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BOOKS BY
WILLIAM EMERSON HITTER

THE HIGHER USEFULNESS OF SCIENCE.

THE PROBABLE INFINITY OF NATURE

AND LIFE.

THE UNITY OF THE ORGANISM, OR

THE ORGANISMAL CONCEPTION OF

LIFE. Illustrated.

THE UNITY OF THE ORGANIC SPECIES,

WITH SPECIAL REFERENCE TO THE

HUMAN SPECIES.

WAR, SCIENCE AND CIVILIZATION.

AN ORGANISMAL CONCEPTION OF

CONSCIOUSNESS.

RICHARD G. BADGER, PUBLISHER, BOSTON

FIGURE 56. SKELETON OF PYTHON.

THE UNITY OF THE
ORGANISM

OR
THE ORGANISMAL CONCEPTION OF LIFE

BY

WILLIAM EMERSON HITTER

Director of the Scripps Institution for

Jiioloyical Research of the University

of California, La Jolla

California

TWO VOLUMES
VOLUME TWO

ILLUSTRATED

BOSTON

RICHARD G. BADGER

THE GORHAM PRESS

COPYRIGHT, 1919, BY RICHARD G. BADGER

All Rights Reserved

This work contains the text of the book:
"An Organismal Theory of Consciousness.'

Made in the United States of America

The Gorham Press, Boston, U. S. A.

CONTENTS

PART I

CRITIQUE OF THE ELEMENTALIST CONCEPTION OF THE

ORGANISM

B. The Production of Individuals by Other Individuals (Concluded)

CHAPTER PAGE

XIV. EVIDENCE FROM METAZOAN GERM-CELLS THAT SUBSTANCES

OTHER THAN CHROMATIN ARE THE PHYSICAL BASES OF
HEREDITY 1

Evidence from spermatozoa, 1. Spermatozoa subject to
heredity as well as "bearers of heredity," 1: (a) Illus-
trated by the ontogeny of mammalian sperm, 4; (b) Illus-
trated by the ontogeny of an insect sperm, 9. Evidence from
the ovum, 15: (a) eggs of ascidians — the facts, 16; (6)
Conklin's interpretation, 19. Critical examination of Conk-
Hn's interpretation, S3.

XV. EVIDENCE FROM SOMATIC HISTOGENESIS IN THE MULTICELLULAR

ORGANISMS ... 32

The mitochondrial theory of heredity, 32. The mitochon-
drial theory tested by the ontogeny of spermatozoa, 36. The
mitochondrial theory tested by histogenesis, 37. The un-
tenable hypothesis that cytoplasm of the ovum is inheritance
material for general but not for special characters, 40. Spe-
cies attributes in single cells of adult orgtvwsms, 43. The
spinules of the ascidian genus styela, 44- The spicules of
sponges and other invertebrates, 50. The "hairs" of higher
plants, 55. Cell-wall structures in higher plants, 57. The
morphology of striated muscle fibers, 60. The physiology of
muscle fibers, 61. Summary of positive information about
the physical basis of heredity, 64.

XVI. THE INHERITANCE MATERIALS OF GERM-CELLS INITIATORS

RATHER THAN DETERMINERS 66

Antecedents of the cytoplasmic and nuclear theories of in-

ix

x Contents

i r. \PTER PAGE

heritance -material. (!6. Function of chromosomes in heredity
aryuin-d and nc<-<>ii<l<iry, 67. The two-fold character of the
j>n>b!i'in of hereditary substance, 70. The probability that
inheritance material becomes such in each ontogeny, 73.
Germ-cells subject to metabolism like all other cells, 74.
Chemical changes in germ-cells during parent's ontogeny, 75.
The possibility of changing sex by influences on the germ,
76. The determiner conception contrary to ordinary chem-
ical principles, 79. Endorsement of E. B. Wilson's proposal
to drop "determiner" from the vocabulary of genetics, 8%.
Advantages in conceiving germ-cell chromosomes as initia-
tors in hereditary development, S3. Inconclusiveness of the
cytological evidence usually appealed to in support of the
chromosome theory, 84. Summing up of the findings
against the chromosome theory, 87. Brief reference to the
untoward implications of the germplasm conception of
heredity, 89.

PART II. THE CONSTRUCTIVE SIDE OF THE
ORGANISMAL CONCEPTION

XVII. GROWTH INTEGRATION 93

The field to be covered by the constructive discussion, 93.
Four types of bio-integration to be treated, 94- Graded
repetitive series as integrative phenomena, 95. Illustrations
from animals, 95. Illustrations from plants, 99. Justifica-
tion for bringing all these phenomena under one head, 103.
Attempted casual explanation of these series, 104- Axial
metabolic gradients as integrative phenomena,, 107. Meristic
gradients and metabolic gradients both phenomena of growth
integration, 111.

XVIII. CHEMICO-FUNCTIONAL INTEGRATION 113

Functional as contrasted with growth integration, 113. The
conception of "internal secretions," 114- Effects of removing

the human thyroid for curative purposes, 115. Experimental
thyroid excision in normal lower animals, 117. The internal
secretion of the duodenal mucous membrane, 119. The na-
ture of the active substances in internal secretions, 121. The
close resemblances and interrelations of the different in-
ternal secretions, 124- Relation between the internal secre-

Contents xi

CHAPTER PAGE

lory and nerrons systems, 128. Composition and nature
of the auloiioinic xi/xtem, 129. Experimental evidence of
connection between the adrenal glands and the nervous sys-
tem, 131. Clinical eridence of adrenal-nervous connection,
133. Summary of present state of knowledge in this field,
137.

XIX. THE ORGAN ISM AT. SIGNIFICANCE OF THE INTERNAL SECRETORY

SYSTEM 141

General inability of elementalism to Interpret Ihe phenom-
ena, l^l. Critique of the view that infernal secretions are
"format ire stuffs," 1/fi. The metaphysical and logical weak-
ness of the view, 1^8. The form of metaphysical absolutism
involved, 151. Confusion of theory of organisms and
theory of the knowledge of organisms, 152. An illustra-
tion of neglect of fact by elementalist theory, 157. A peculiar
elementalist objection to the organic whole, 15X.

XX. NEURAL INTEGRATION 161

Distinction between developmental and functional integra-
tion, 161. Neural and not psychical phenomena the subject
of this chapter, 101. The author's indebtedness to Sher-
rington's work, 162. The fundament alii y of cellular inte-
gration in the reflex-arc, 163. The integration of reflex-
arcs, 168. The spreading and compounding of reflexes, 171.
Antagonistic refl.exes in skeletal muscle groups finally in-
tegratire, 17//. The antagonisms within the autonomic sys-
Icni finally inlegrative, 178. Concluding remarks on the sig-
nificance of neural integration for the organismal standpoint,

XXI. IMPLICATIONS OF THE TROPISTIC AND SEGMENTAL THEORIES OF

NERVE ACTION 18.1

Neglect of the works of Sherrington and Cannon by Jacques
Loeb, 185. The real importance of Loeb's conception of the
nf.rvous system, 187. The organismal character of tropisms
partly recognized by Loeb, 188. The organismal character
of the segmentai theory of the nervous system, 190. Critique
of the element a lixl attempt to interpret tropistic and seg-
mentai theories of the function of the nervous system, 198.
The bearing of this critique on "analysis" in Biological

xii Contents

CHAPTER PAGE

reasoning, 206. Theories of animal behavior in relation to the
"science" of zoology, 208.

XXII. PSYCHICAL INTEGRATION 214

Preliminary remarks: (a) Absolute discrimination between
reflex and psychical phenomena not necessary, 214; (b) The
organism an original datum in all problems of psychic life,
215; (c) Provisional classification of psychical facts, 217.
Likeness between tropistic and higher psychic activity, 220.
First move toward showing the organismal character of the
higher psychic life, 221. Associatwnist psychology a special
case of elementalist biology, 228. Preliminary examination

of objective and subjective, 231. The essence of Wundtian
apperception, 2S2. Remarks on analysis and synthesis, 236.

XXIII. ORGANIC CONNECTION BETWEEN PHYSICAL AND PSYCHICAL 239

A still closer look at the organismal nature of tropisms,
239. The automatic and anticipatory character of tropisms
and other reflexes, 2^1. A still closer look at the likeness
between higher rational life and tropisms, 2I/.2. A still
closer description of the subrational moiety of psychic life,
246. Remarks on the classes of subrational life, 2J^6. Four
certainties about the adaptiveness of subrational psychic
activities, 250. Generally useful to individual and to species,
250. Many useful to species primarily, 250. Variability of
subrational activities, 251. Tendency of subrational activ-
ities to excessiveness, 256. Summary of organismal charac-
ter of all subrational psychic life, 21 '4- Specificity of subra-
tional psychic life, 276.

XXIV. SKETCH OF AN ORGANISMAL THEORY OP CONSCIOUSNESS . 282
Remarks on the hypothetical character of this chapter, 282.

The natural history method and the study of one's self, 282.
Formulation of the central hypothesis, 286. Preliminary
justification of the hypothesis as such, 287. More systematic
justification of the hypothesis, 291. The nature of "outer"
or objective and "inner" or subjective, 292. As to the lowest
terms of self -consciousness, 308. Instinct and physical or-
ganization, 310: Emotion and physical organization, 316.
Glance at the' equilibrative interaction between "body" and
"soul," 323. Support of the hypothesis by the physico-chem-

Contents xiii

CHAPTER PAGE

ical conception of the organism, 32!>. Personality and ele-
mentary chemical substances, 327. On the psychology of
subjective and objective personality, 331. Personality and the
"breath of life" viewed in the light of physical chemistry
of the organism, 336.

POSTSCRIPT 351

BIBLIOGRAPHY 359

GLOSSARY 377

INDEX • 39J

LIST OF ILLUSTRATIONS

FIGURE PAGE

56. Skeleton of Python Frontispiece

36. Sperm of Fur Seal (After Oliver) 4

3T. "Young" Spermatid (After Ballowitz) 5

38. Spermatid of Fur Seal (After Oliver) 5

39. Spermatid of Fur Seal (After Oliver) 6

40. Spermatid of Fur Seal (After Oliver) 8

41. Spermatid of Fur Seal (After Oliver) 8

42. Development of Sperm of Argas Miniatus (After Casteel) 10, 11

43. Development of an Ascidian Egg (After Conklin) ... 17

44. Spinule Cell of Styela Yakutatensis (After Huntsman) . 44

45. Spinule Cell of Styela Plicata (After Huntsman) ... 45

46. Spinule Cell of Styela Greeleyi (After Huntsman) . . 46

47. Spinule Cell of Styela Montereyensis (After Ritter and For-

syth) 47

48. Spicules of Sponges (After Lankester) 51

49. 50, 51. Development of a Spicule (After Lankester) . 52, 53

52. Trichomes of Papaver Orientale (After Cannon) ... 55

53. Trichomes of P. Pilosum (After Cannon) .... 55

54. Trichomes of P. Somniferum (After Cannon) .... 55

55. Side View of Amphioxus (After Parker & Haswell) . . 95

57. Tentacle of Halocynthia Johnsoni (Schematic; After Ritter) 98

58. Acacia Elata 99

59. Vicia Gigantea 100

60. Cassia Sp 100

61. Sequoia Sempervirens 101

PART I

CRITIQUE OF THE ELEMENTALIST CONCEPTION OF
THE ORGANISM

B. The Production of Individuals by Other Individuals

THE UNITY OF THE ORGANISM

Chapter XIV

EVIDENCE FROM METAZOAN GERM-CELLS THAT
SUBSTANCES OTHER THAN CHROMATIN
ARE THE PHYSICAL BASES OF
HEREDITY

Evidence from Spermatozoa

JN our discussion we will be guided by the principle laid
•*• down earlier and followed throughout our treatment
of heredity in the Protozoa, namely, that descriptive onto-
genesis brought to bear on the actual transformations which
result in the production of specific organs and parts, is the
final tribunal for the determination of what substances are
hereditarily formative. The first inquiry will be whether
there exist among the metazoa single-cell organs or parts
which when fully formed exhibit species characters in the
sense of taxonomic biology. If such elements do exist, ob-
servation on the constituents of the undifferentiated cells
which take part in the transformations, obviously may be
expected to give us the information sought as to what
substances are formative.

Spermatozoa Subject to Heredity as WeU as "Bearers of

Heredity"

The comparative anatomy and comparative ontogenesis
of the male germ-cells among animals, which have been pur-
sued with great assiduity and skill during recent decades,

1

2 The Unity of the Organism

furnish perhaps the largest mass of relevant facts we pos-
sess from any one field.

Innumerable researches on fully formed spermatozoa, the
greatest single research being that of Retzius, give us ex-
tensive knowledge of the variety of structure of the sperma-
tozoa in the larger and smaller taxonomic subdivisions of
the animal world. It would be going beyond the evidence
to say that every well-characterized animal species may be
identified by its spermatozoa; but unquestionably the trend
of investigation has been toward such a conclusion. I
believe, for example, it would be impossible to assert on the
basis of evidence that any two species of animals belonging
to different genera, no matter how much alike if their dis-
tinctiveness is not questioned, have indistinguishable sper-
matozoa. "One may say," writes Ballowitz, "that each ani-
mal species has its own sperm-form of definite size." An
attempt to illustrate fully this variety of form and size by
specific examples is out of the question here. We will refer
only to the specificity of the sperm of man himself. Ret-
zius was able to compare in detail sperms of the Chimpanzee,
Orang-Utan, Gibbon, and Homo, and found that while their
resemblance is rather close, each possesses clear differential
marks. For example, the spiral structure of the envelop of
the central piece is considerably more distinct in the Chim-
panzee than in Homo. Worthy of mention is the fact that,
according to Retzius, the sperm of the Chimpanzee resembles
that of Homo more closely than does that of the Orang,
thus falling in with the fact that in several particulars of
adult structure the resemblance of the Chimpanzee to man
is closer than that of the Orang.

The spermatozoa of a given animal group having a closer
resemblance to one another than to those of other groups ;
in other words, having a resemblance due to descent, are
themselves subject to heredity and are not alone concerned
in the transmission of hereditary attributes from parent to

Evidence from Metazoan Germ-Cells 3

offspring1.

Especially important for us is it to notice that to a great
extent the diversity of structure among spermatozoa is in
the locomotor organ, the tail; that is, the organ chiefly
concerned with the unique life of the sperm as such, and
very little if at all concerned directly with fertilization and
hence with hereditary marks of offspring. This fact de-
serves attentive consideration. "In its more usual form the
animal spermatozoon resembles a minute, elongated tad-
pole, which swims very actively about by the vibration of a
long, slender tail." 2 In some respects comparison of the
spermatozoon of the type here indicated with an Appen-
dicularian, a minute Tunicate which possesses a tail through-
out its life, is more instructive. Any one who has had op-
portunity to observe both sperm cell and appendicularian
when alive and active will not have failed to remark the
general resemblance, not only as to form but as to kind
of movement in the two cases. Is the development of the
Appendicularian's tail a manifestation of heredity? Surely
no one would think of giving any but an affirmative answer.
How, then, deny that the development of the spermatozoon's
tail is also a manifestation of heredity? I cannot see that
it would be less inconsistent to affirm that the wriggling ap-
pendicularian is alive but that the wriggling spermatozoon
is not, than to affirm that the ontogeny of the first is guided
by heredity while that of the second is not. To bring the
point onto somewhat more familiar ground, let us revert
to Wilson's comparison of the spermatozoon to the tadpole
stage in the life of the frog. Our contention is that the tail
of the frog's spermatozoon is as indisputably modeled by
heredity as is the tail of the frog's tadpole, and consequently
that we are bound to search for the physical basis of hered-
ity in the former as well as in the latter. Our inquiry is,
then, What observations have we as to the substances con-
cerned in producing the spermatozoon tail?

4 The Unity of the Organism

(a) Illustrated by the Ontogeny of Mammalian Sperm

Without exception, so far as I know, positive description
of spermatogenesis affirms that the tail is produced from

h.c.

ra.p.--

FIGURE 36. SPERM OF FUR SEAL (AFTER OLIVER).

h.c., head cap. n'k., neck, c.p., connecting piece, c.r., cytoplasmic
remnant, g.a., anterior granules, g.p., posterior granules, an.,
annulus. m.p., main piece, e.p., end piece.

other parts of the spermatid than the nucleus. For the lit-
tle we can do in the way of giving objectivity to this general
statement we will first examine the nearly mature, typical

Evidence from Metazoan Germ-Cells 5

mammalian spermatozoon (figure 36) of the Fur Seal.

For the origin of the various parts we will make use of
the summary given by Ballowitz (figure 37). Concerning

FIGURE 37. "YOUNG" SPEHMATID (AFTER BALLOWITZ).

1, nucleus. 2, centrioles. 3, idiosome. 4, mitochondrial body. 5,
chromatoid body. 6, spindle remnant.

the nucleus, ./, "it is certain that in all animals the chroma-
tin-containing part of the sperm proceeds from the
nucleus." 3 On this point there is agreement among observ-
ers ; so whatever may be the truth about the sperm-chromatin

FIGURE 38. SPERMATID OF FUR SEAL (AFTER OLIVER).

n., nucleus, c., centrioles. a.f., axial filament.

as the physical basis of heredity for the adult animal, there
is no question about its being such for the head of the
spermatozoon itself.

6

The Unity of the Organism

The centrioles (2, figure 37) are "almost always double and
occupy a place in the spermatid close beneath the surface of the
cell." ' The pair is typically so placed that the axis joining them
is perpendicular to the surface of the spermatid, the member

a.f.

FIGURE 39. SPERMATID OF FUR SEAL (AFTER OLIVER).

h.c., head cap. n., nucleus, p.c., proximate centriole. d.c., distal
centriole. a.f., axial filament, s., remnant of sphere.

toward the surface being known as the distal centriole and the
one toward the center of the spermatid the proximal centriole
(figure 39 d.c. and p.c.).

As to the part played by each of these in the development of

FIGURE 40. SPERMATID OF FUR SEAL (AFTER OLIVER).

n., nucleus, c.h., chromatin granules, c.t.f., caudal tube filaments.

the sperm we learn that after migrating inward until they
come to lie very near the nucleus, if not in actual contact with
it, the centrioles begin their development with the following out-
come: "The proximal centriole . . . divides into two portions,
closely adherent to the nuclear wall, each connected by a fila-
ment to one distal group. The distal centriole divides into an

Evidence from, Metazoan Germ-Cells 7

anterior and a posterior portion. The -posterior portion becomes
the annulus (an., figure 36) while the anterior one divides again,
forming the Noduli posteriores" 4 (g.p., figure 36). Throughout
their career these bodies or granules are highly stainable with
certain dye-stuffs.

Out of the idiosome (3, figure 37) which as a rule is a body
"fiir sich" — in itself alone — develops the perforatorium, or head
cap of the adult sperm (h.c., figure 36). It is agreed that the
mitochondria (4, figure 37) in the spermatids of many animals,
particularly of many vertebrates "furnish the material" 3 for the
spiral found in the connecting piece (c.p., figure 36) of the
sperm. Although no spiral is present in the Seal sperm it may
be represented, according to Oliver, by numerous granules sur-
rounding the axial filament in the connecting piece. But while
the connecting piece of the Seal sperm seems not to be typical
as regards the spiral, it presents another structure, the caudal
tube, or "manchette" of some authors, in a form which is specially
instructive from our standpoint. In the adult sperm this struc-
ture is a thin sheath enveloping the cytoplasmic part of the con-
necting piece and lying in close contact with the persisting cell
membrane. The point of special interest about it is that its
persistence in the completed sperm of the Seal appears to be
exceptional, for it is known to disappear entirely in the course
of development of the sperm of several other mammals. It is a
transitory or embryonic organ in some species of sperm, but a
permanent one in other species, just as gills, for example, are
transitory organs in the ontogeny of some species, as a frog, but
are permanent in others, as fish,

The development of the tube in the Seal sperm is especially
favorable for observation. "It may be readily followed," writes
Oliver, "from its first appearance up to its final incorporation
in the connecting piece as a peripheral layer, or sheath." * Here
then is a structure having all the essential marks of devel-
opment due to heredity and likewise one the "physical basis" of
which has been carefully observed. Mentioning a long list of
investigators who believe in the "derivation of the caudal tube
by a process of cytoplasmic differentiation alone" Miss Oliver
tells us that her study of the development of the fur seal sperm
is a complete confirmation of this view. As to the very begin-
ning of the tube we read: "Shortly after the centrosomes and
their tail filament have reached the nuclear membrane there
appears in the cytoplasm surrounding the axial thread a series

8

The Unity of the Organism

of delicate filaments attached to the nuclear membrane. The
proximal ends of these arise in a circle around the basal end
of the nucleus with the centrosomes as a center, while their
distal ends project freely into the cytoplasm." 5 (figure 40 c.t.f.)
These filaments "are at first very short and thin, but they in-
crease in length and thickness rapidly. By the progressive dif-
ferentiation of the cytoplasm between them they soon fuse into
a hyaline tube, surrounding the axial thread and open at its lower
extremity." (figure 41, c.f.) The capital point is that we have
here a well-defined structure the development of which is in-

-an.

FIGURE 41. SPEEMATID OF FUR SEAL (AFTER OLIVER).

h.c., head cap. c.t., caudal tube, an., annulus.

dubitably proved to depend primarily on parts of the cell other
than the chromatin. Indeed no one, apparently, has pretended
that the chromatin takes a part in its production, for even those
investigators who have not believed that it arises from the cyto-
plasm alone have held that it originates from the membrane of
the nucleus.

About the chromatoid body (5) and the "spindle remnant" (#)
(figure 37), little need be said in this connection as they seem to
be inconstant structures the significance of which is in much
doubt. Finally, mention should be made of the fact that a con-
siderable portion of the cytoplasm is cast off entirely in the
ontogeny of the sperm of many animals, as for example the seal,

Evidence from Metazoan Germ-Cells 9

the body (c. r., figure 36) being all that is left of the substance
at the late stage represented. So mucli by way of illustration of
the portions of the spermatid, or germ of the spermatozoon, in
the vertebrated animals, for despite the great variety in struct-
ural details presented by the sperm of this part of the animal
kingdom I think all will agree that so far as concerns the chief
point being made in this discussion, what we have presented is
true of the whole phylum.

(b) Illustrated by the Ontogeny of an Insect Sperm

We will now examine the ontogeny of a very different
type of sperm, from another portion of the animal kingdom,
the insecta. The particular species chosen is the fowl-tick
(Argas miniatiis). The investigation made use of is by
Doctor D. B. Casteel.

The series of figures (42 a, b, c, d, e, f, g,} will help to
an understanding of the remarkable, almost unique sperm and
spermogenesis in this animal. Figure 42a shows the nearly ma-
ture primary spermatocyte. Especially to be noted are the
mitochondria, mi., scattered uniformly through the cytoplasm,
and the striated layer, *. I., on the outer surface of the cell.
This layer is sharply demarked from the underlying cytoplasm.
The striae, disposed perpendicular to the surface of the cell, are
excessively fine, and when looked at in situ end on "suggest the
appearance of a faceted compound eye or of honey-comb." 8
Concerning the genesis of this layer Casteel says, in a personal
letter, that the layer begins to appear at the surface and grad-
ually increases in depth until the completed state shown in figure
42a is reached. "The striae," he says, "appear to be forming
from the undifferentiated cytoplasm sheath."

All the figures from 42 b to g have to do with the transforma-
tion of the spermatid (figure 42b) into the spermatozoon. From
figure 42b one sees that the striated layer has disappeared on one
side of the cell and thinned out greatly in a smaller area on the
opposite side; that the nucleus, n, has moved to the surface of
the cell in the middle of the area of disappearance of the striated
layer; that the large plasmosomes, pi. 42a have almost entirely
disappeared from the nucleus ; that the vesicular bodies v.b., as-
sembled for the most part in the vicinity of the nucleus, are

•i.t

fp.

FIGURE 42.
FIGURE 42. DEVELOPMENT OF SPERM OF ARGAS MIXIATUS (AFTER CASTEEL).

a., rupture point of outer tube, c.p., cilia-like processes, fl., flagel-
lum. f.p., finger-form process, g.e., gelatinous envelop, i.e., in-

10

n.— -/

rru\- -

fi-

rm.

FIGURE 42.

vagination cavity, i.t., inner tube, mi., mitochondria, m.r., mito-
chondrial ring, n., nucleus, o.d., oil droplet, o.t., outer tube, pi.,
plasmosome. s.l., striated layer. V.b., vesicular bodies, b.p., begin-
ning of inner tube.

11

12 The Unity of the Organism

in processes of degeneration and disappearance; and that the
mitochondria, mi, look as though they had collected into two
well-defined spherical masses m.r., on the side of the cell oppo-
site the nucleus and near the small area of thinned-out striated
layer. As a matter of fact these two apparent mitochondrial
masses are the opposite sides of a ring seen in optical section.

Transfoimiation of the general form of the spermatid now be-
gins by the indentation of the side opposite the nucleus, this
going on to produce first the quarter-moon shape shown at i.e.,
figure 42c. By the still further growth and narrowing the edges
of the cup finally come together to produce the elongated cavity
shown in figure 42d, o.t. This is the beginning of the outer tube
which becomes long and relatively narrow as development con-
tinues, (figure 42e, ».£.). At an early period in the growth of
this tube the striated layer which naturally becomes shut into the
tube breaks up over most of the circumference of the tube into
what resembles a dense layer of long cilia. However, since the
processes are not motile and later dissolve and produce a gela-
tinous mass within the tube, their resemblance to cilia is only
superficial. This account of the fate of the striated layer applies,
as previously intimated, to most of the circumference of the outer
tube. But the small thin area of the layer opposite the nucleus,
shown in figures 42d and 42c, retains the cilia-like processes.
This persistent basal patch (b.p., figure 42d) is the starting
point for an important part of the future spermatozoon, the
"inner tube" so called by Doctor Casteel (i.t., figure 42e).

While these profound changes are going on in the portion
of the spermatid opposite the nucleus, a stout, somewhat finger-
like process, (f.p., figure 42d) is formed on the nuclear side of
the spermatid, into the base of which the nucleus migrates. In
the meantime the vesicular bodies have entirely disappeared, and
the mitochondria, no longer disposed in the ring of earlier stages,
have assembled into an irregular, rather diffuse mass toward the
basal patch {mi., figure 42d).

At the time when the outer tube has reached its maximum
length and is somewhat coiled, the inner tube, starting at the
basal patch previously described, begins to grow into the cavity
of the outer tube. This growth continues until the inner tube
is approximately as long as the outer tube. Figure 42e presents
an advanced but not completed stage of growth of the inner tube
i.t. In reality, according to Doctor Casteel's interpretation, the
inner tube grows at the expense of the outer tube, for when the

Evidence from Metazoan Germ-Cells 13

two are of nearly equal length the entire spermatozoon is only
about half as long as it was before the inner tube developed.
All of the mitochondria of the cell are drawn into the inner tube
as it grows, and finally form a deeply staining mass at the distal
end of the tube {mi., figures 42e and 42g).

During these transformatory operations the nucleus, greatly
reduced in size proportionally to the spermatozoon as a whole,
has left its former place at the base of the finger-like process,
and been making its way along the wall of the outer tube, bur-
rowing through the gelatinous layer on the outer part of this
tube (n, figure 42e). This migration continues until, when the
distal end of the inner tube reaches nearly the end of the outer
tube, the nucleus lies in the wall of the outer tube opposite the
end of the inner tube (n, figure 42f.)

The final act of transformation takes place after the sperm
has left the' male tick and lies in a spermatophore sac within
the genital ducts of the female. This act begins with the per-
foration of the end (a. figure 42 f) of the outer tube by the
inner tube. Through the opening thus made the whole inner tube
finally passes, really by a slipping back of the outer tube, so
that, the eversion of this latter being completed, the two tubes
constitute one continuous tube. By this act of turning inside
out, the finger-like process in which the nucleus formerly lay
(figure 42d, f.p.) isv brought into close proximity again with the
nucleus.

Some of the details of the final steps in the transformation
Doctor Casteel has not yet been able to make out; but these
are of little consequence for our discussion. Nor has the
act of fertilization been observed. Going on the usual cri-
teria, the end of the sperm containing the nucleus would be
regarded as the head. But surprisingly enough, in moving,
the opposite end, the end containing the mitochondria, goes
foremost.

This sperm and its development are so unique as con-
trasted with those occurring in most animal groups, that
one might be almost inclined to question whether there may
not be something wrong here — whether the case may not be
one of diseased growth, or the result of manipulative mal-

14 The Unity of the Organism

formation, or something else. Any such suspicion is, how-
ever, completely done away with by the fact that much the
same type of spermogenesis is known to occur in other ticks.
Casteel cites particularly the observations of Katharine
Samson on the development of the sperm of Ixodes ricinus
and Ornithodes moubata as furnishing cases to which that
of Argas is "in many respects parallel."

After all that has been said in the previous pages, it is
almost needless to point out the significance for our general
contention of this remarkable case of spermogenesis. Were
we living in that comfortable era of life-philosophy wherein
theologians studied nature for the purpose of proving that
everything in it was made to meet some human need, we
could easily recognize this case as one designed expressly
to assist man in refuting the false dogma of Chromatinic
Omnipotence in heredity.

It is hard to imagine a developmental process in which
denial of form-determining power to non-chromatinic, even
non-nuclear parts of the cell would be a greater folly than
would be the denial of cytoplasmic "form-determination"
in the production of the striated layer of the spermatid, or
of the growth of the outer tube, or of the inner tube, or
of the turning inside-out of the outer tube. In fact, by far
the greater part of the astonishing transformations here
gone through are cytoplasmic, the nuclear changes being
relatively slight.

It may be worth while to remind the reader again, so
boldly does the real truth about hereditary substance stand
out in this case, that the cardinal evil in the chromatin
dogma is that it implies the denial of great masses of the
most direct observational evidence we have as to what the
physical bases of heredity may be, and so tends to detract
attention from them. We may predict that the important
research which has made known this unique case of organic
genesis will pass almost unnoticed by the geneticists of our

Evidence from Metazoan Germ-Cells 15

day. Were their attention called to it they would probably
frankly say that they have little interest in genesis in this
sense. That the sperm here described is not peculiar in
every respect to the species Argas miniatus is certain from
the meager comparative information we possess, as Casteel
has shown. Nevertheless not merely general analogy, but
strong indications contained in even the little comparative
knowledge we have in this particular case, warrant the sup-
position that in some respects the sperm of the species would
be peculiar to the species, to say nothing of the genus,
family and so forth. The development is, consequently, due
to heredity, and the cytoplasm is "inheritance material"
as ascertained by direct observation.

Evidence from the Ovum

We now turn to the ovum to see what can be learned con-
cerning hereditary substance in the development of the
ovum iself. Attention should be called at the outset to the
important difference between the sperm and the ovum in
the kind of specialization in each. The sperm, it will be
noticed, is far more specialized for its own particular life
than is the ovum, this "particular life" of the sperm con-
sisting in its great power of locomotion. As a consequence
of this difference, the ovum as an entity has no such sharp
distinction from the ovum as a germinal element as has the
spermatozoon. This difference is expressed in one way
by the assertion that the fertilized ovum Is the individual
organism in the one-celled stage of its life. No such state-
ment is ever heard about the spermatozoon for the obvious
reason that the sperm does not transform directly into the
embryo as does the egg. From the absence of so distinctive
a character of the ovum as such, it happens that the hered-
ity of the ovum is not so distinguishable from the heredity
of the organism of whose life it is a stage, as is the case

16 The Unity of the Organism

with the sperm. Nevertheless we are bound to recognize
that the egg no less than the sperm has hereditary attri-
butes of its own, and that other substances than chromatin
play a demonstrable part in the production of these. In-
deed the main discoveries concerning what in an earlier
chapter was called the promorphology of the egg are of this
sort. There is one kind of promorphology that is of special
importance to the present stage of this discussion. I refer
to the kind known sometimes as "germinal localization" and
sometimes as "organ forming substances" in the ovum. The
idea, expressed in a sentence, is that in the eggs of some
animals, portions of the egg destined to give rise to par-
ticular parts of the future embryo are visibly different from
other portions before cell division begins, in some cases even
before maturation and fertilization occur. According to
our understanding of heredity, these distinguishable por-
tions of such eggs are themselves hereditary attributes not
only of the animal species to which the eggs belong, but of
the eggs, no less than are distinctive morphological features
of the adult animals or of any developmental stages. The
study of these attributes of eggs is peculiarly interesting
since, belonging to germ-cells par excellence, if we can get
observational evidence on both their origin and destination,
we shall have direct evidence that one and the same substance
is determined on the one hand by heredity, and on the other
is a determiner in the strict genetic sense of hereditary at-
tributes yet to be developed.

(o) Eggs of Ascidianj — The Facts

Because of the great importance of the observations of
E. G. Conklin in this field, and of his general views con-
cerning the bearing of his observations on heredity, we shall
make his work the center of our examination. One of the
most important of Conklin's investigations was on the eggs

Evidence from Metazoan Germ-Cells

17

of several species of Ascidians. Among the great merits of
this investigation are the facts that the normal living eggs
were studied with great care, and that the comparative
method was employed to a considerable extent.

•t.c.

CIX.

C IX.

O.V.

t.C.

Cr.

FKirilK 43. DEVELOPMENT OF AX ASCIDIAX EGG (AFTER COXKMX).

cn., chorion. c.p., clear protoplasm, c.r., crescent of mesodermal
substance, g.v., germinal vesicle, p.b., polar body, p.l., peripheral
layer of protoplasm, t.c., test cells, y.h., yellow hemisphere of

egg- yk-» y°lk-

Three species of simple Ascidians, Ciona intestinalis, Mol-
gula manhattensis and Styela partita furnished the eggs
for the investigations.

We will begin the examination by quoting from the sum-

18 . The Unity of the Organism

mary of results, and then use these as the basis of a closer
scrutiny of the conclusions. Under the head "Organization
of the Egg" we find : "14. In the ovocyte of Cynthia partita
there is a peripheral layer of yellow protoplasm, (p. Z.) a
central mass of gray yolk, («/&.) and a large clear germinal
vesicle (g. v.) which is eccentric toward the animal pole
(figure 43a). These same parts are present in the eggs
of other ascidians, but are differently colored.

"15. When the wall of the germinal vesicle dissolves at
the beginning of maturation divisions a large amount of
clear protoplasm, containing dissolved oxychromatin, is
liberated into the cell body. This clear protoplasm is ec-
centric toward the animal pole and is distinct from the yolk
and peripheral layer.

"16. Immediately after the entrance of the spermatozoon
the yellow and clear protoplasm flow rapidly to the lower
pole, where the yellow protoplasm collects around the point
of entrance ; the clear protoplasm lies at a deeper level. The
yellow protoplasm then spreads out until it covers the sur-
face of the lower hemisphere. This flowing of protoplasm
to the point of entrance of the sperm is comparable with
what takes place in many animals, though much more ex-
tensive and rapid here than elsewhere (figure 43b).

"18. The sperm nucleus moves from the point of en-
trance toward the equator in a path which is apparently
predetermined. This path lies in the plane of first cleavage
and the point, just below the equator, at which the sperm
nucleus stops in its upward movement, becomes the posterior
pole of the embryo. The median plane and the posterior
pole are probably not determined by the path of the sperma-
tozoon but by the structure of the egg. All the axes of the
future animal are now clearly established antero-posterior,
right-left, dorso-ventral.

19. The yellow protoplasm "collects into a yellow cres-
cent with its middle at the posterior pole and its horns ex-

Evidence from Metazoan Germ-Cells 19

tending about half way around the egg just below the
equator (figure 43c. y.h.) This position it retains through-
out the whole development, giving rise to the muscle and
mesenchyme cells (figure 43 d. cr.)

"20. ... At the close of the first cleavage (figure 43d)
the nuclei and clear protoplasm move (c.p.) into the upper
hemisphere and thereafter, throughout development, this
hemisphere contains most of the clear protoplasm and gives
rise to the ectoderm.

"21. . . . When the clear protoplasm moves into the
upper hemisphere the yolk is largely collected in the lower
hemisphere. This yolk-rich area gives rise to the endoderm.

"24. The chief factor of localization is protoplasmic
flowing; cell division is a factor of subordinate value."7

From the numbers given in this quotation it will be recog-
nized that only a portion of the summing up of results is
here presented; and it will be understood that each item in
the summary represents a lengthy, detailed description in
the body of the memoir. This summary we may again sum-
marize as follows: By the time the first division of the egg-
cell is completed the portions of the egg which are to give
rise to three great groups of tissues of the future animal
are distinguishable from one another by definitely visible
attributes. The clear protoplasm situated at the upper
pole of the cell will give rise to the external epithelium or
skin ; the yolk-laden protoplasm in the lower hemisphere will
produce the epithelial lining of the digestive tract; and
from the yellow protoplasm gathered on the surface at the
lower pole will come most of the muscle and connective tis-
sue of the animal.

(6) Conklin's Interpretation

Passing now to a consideration of these facts in relation
to heredity we must not neglect to notice the two-fold aspect

20 The Unity of the Organism

of the question, namely that we have to do on the one hand
with attributes of the egg itself which are results or termini
of the kind that gave rise to the conception of heredity;
and on the other hand with these same attributes not as
results but as causes or forerunners of later-appearing at-
tributes, also by the same criteria due to heredity. That the
egg attributes with which we are dealing are due to heredity
is obvious from the fortunate circumstance that, as already
indicated, the observations under review were comparative
to considerable extent. Thus we have this general com-
parison of the unripe eggs of the three species: "In the liv-
ing eggs of Cynthia this peripheral layer is clear and trans-
parent and contains uniformly but sparsely distributed
yellow pigment, which seems to be associated with these small
refractive spherules. ... In dona and Molgula also these
three areas are distinguishable in the living egg before ma-
turation, but not so clearly as in Cynthia. In Ciona the
peripheral layer is nearly transparent, the yolk is a brown-
ish red. ... In Molgula both the peripheral layer and the
germinal vesicle are transparent, while the yolk is gray,
with a faint lilac tinge." As an example of a very definite
statement of specific difference between the eggs we read,
"In Ciona, the same type of protoplasmic movement occurs
as in Cynthia, but with certain minor differences."

The evidence brought forward by Conklin of species differ-
ences in eggs is not restricted by any means to Ascidians.
In some of the gastropod mollusca, for example, similar re-
sults were reached by a very notable research on the quanti-
tative relations of various egg organs of several species of
the genus Crepidula. To be explicit in a single case only,
the relative volume of macromeres 3A-3D to 3a-3d was
found to be 12.1 :1 in Crepidula plana and 59.3 :1 in C.
convex. From a long series of determinations of difference,
Conklin remarks: "One cannot study the eggs of different
animals without being much impressed with the fact that

Evidence from Metazoan Germ-Cells 21

the distribution of yolk to the four macromeres is highly
characteristic of different species and orders." Yet, as
we shall have occasion to point out later, it seems certain
that facts of this kind have deeper meanings than Conklin
has appreciated.

After all that has been said in previous pages about
specific differences, the point aimed at here will be obvious.
The "decidedly thicker" layer of peripheral protoplasm at
the lower pole in Ciona than in Styela, and the brownish
red yolk in Ciona as contrasted with the white yolk tinged
with lilac in Molgula, are indeed "minor differences." But
species differences in adults and in all other stages are minor
as a rule. The difference between the transparent-white
color of the full-grown Ciona intestmalis and the pale, green-
ish yellow of the adult Molgula manhattensis is also minor
as contrasted with the "same type" of organization of the
two animals. But it is just this being true to type as re-
gards minor differences in living beings that has given rise
to the conception not only of organic species, but to a large
extent, of heredity as well; and how can any one recognize
the differential color of the grown-up Ciona as compared
with the grown-up Molgula as due to heredity, but refuse
to recognize the differential color of the eggs of the two
animals as due to the same cause?

We are, then, bound to accept the attributes of the eggs
as the results of heredity, and so inquire about the "physi-
cal basis" or "bearers" of these attributes. And this brings
us again to the main issue of this section: Have we or have
we not observational evidence that any other substance or
substances than chromatin contribute to the production of
the egg-phenomena under consideration?

Conklin's conclusions and mode of reasoning as to the
bearings of his observations on heredity are so crucial for
the point now engaging us that we must examine them in
considerable detail. Returning to the summary of results

22 The Unity of the Organism

upon which we have already drawn we read: "26. The
organization of the ovocyte is not the initial organization.
The yellow protoplasm (mesoplasm) of the Cynthia egg is
probably derived, at least in part, from sphere material
(archoplasm) which arose from the nucleus at the last
ovogenic division. The yolk (endoplasm) is formed by the
activity of the 'yolk matrix' (Crampton) which also is
probably sphere material. The clear protoplasm (ecto-
plasm) is derived from the germinal vesicle at the first
maturation division. Thus many important regions of the
egg come, at least in part, from the nucleus, and a method
is thereby suggested of harmonizing the facts of cytoplasmic
localization with the nuclear inheritance theory."

Again, we find : "This truly remarkable condition in which
considerable portions of the cytoplasm are traceable to the
nucleus is of the utmost theoretical importance. From all
sides the evidence has been accumulating that the chromo-
somes are the seat of the inheritance material, until now this
theory practically amounts to a demonstration. On the
other hand, all students of the early history of the egg have
observed that the earliest visible differentiations occur in
the cytoplasm, and that the position, size and quality of
the cleavage cells and of various organ bases are controlled
by the cytoplasm. However, in the escape of large quanti-
ties of nuclear material into the cell body, and the forma-
tion there of specific protoplasmic substances we have a
possible mechanism for the nuclear control of the cytoplasm,
and when, as in the case of the ascidians and fresh water
gastropods, these substances are definitely localized in the
egg, and can be traced throughout the development until
they enter into the formation of the particular portions of
the embryo, a specific mechanism for the nuclear control of
development is at hand, and the manner of harmonizing the
facts of cytoplasmic organization with the nuclear inher-
itance theory is clearly indicated."

Evidence from. Metazoan Germ-Cells 23

(c) Critical Examination of Conklin's Interpretation

One is struck at the outset of a critical examination of
these sentences by the fact that what is claimed is that ob-
servations have been made whereby a method is "suggested
of harmonizing the facts of cytoplasmic localization with
the nuclear inheritance theory" ; and that this suggested
harmonization of such localization with the "nuclear in-
heritance theory" (not "chromatinic inheritance theory"
be it noticed) is the final count in a mass of evidence which
"practically amounts to a demonstration" that the "chro-
mosomes are the seat of the inheritance material."

In other words objective facts which are only suggestive
of a conclusion as touching the inheritance role of the whole
nucleus rise to the demonstrational level as touching the
same role of very small parts of the nucleus.

The method by which this particular piece of logical
sleight-of-hand is performed is easy to see, for though va-
ried to meet the exigencies of the special case, it follows the
general scheme of elementalistic interpretation with which
we have become familiar. In the first place, the fact that
the observations furnish very little if any direct evidence
that the chromosomes cause the cytoplasmic flowing and
localization is made innocuous as evidence against the chro-
mosome dogma by assuming that the flowing and localization
do not themselves come under heredity. That such an as-
sumption is implied in the argument seems certain from
the fact that Conklin did not consider "that the general
theory of chromosomes as the bearers of heredity which he
had espoused made it incumbent upon him to take cognizance
of the fact that he himself had in reality testified that
portions of the nucleus other than chromosomes are the
seat of inheritance material. We have here another and a
very notable case of shielding a prevalent theory by defini-
tion; that is, of shutting relevant but inimical facts away

21< The Unity of the Organism

frcin it by definition. That Conklin should have been un-
wittingly led into this is the more surprising in that his
c\vn studies, particularly those on comparative morphology
and physiology of the eggs of several groups of animals,
have added largely to the proof that many of the gross
attributes of the eggs themselves are subject to heredity.
Ir.dccd no biologist has expressed more positively than he
the conception that the egg is the individual animal in one
of its stages of development ". . . from its earliest to its
latest stage an individual is one and the same organism;
the egg of a frog is a frog in an early stage of development
and the characteristics of the adult frog develop out of the
egg, but are not transmitted through it by some 'bearer of
heredity.'"12

One wonders if Conklin really would maintain that only
the attributes of an animal in the adult stage of its life are
subject to heredity; or even that the attributes of any of
its developmental stages are not so subject, were he con-
fronted with the question in this form. Although I have
been unable to find statements in his writings from which
one may positively infer what his answer would be, yet
several passages can be brought together which can not, I
believe, be harmonized with one another, but reveal real
contradiction. Thus in The Mechanism of Heredity, al-
ready cited, we find: "Differentiation, and hence heredity,
consists in the main in the appearance of unlike substances
in protoplasm and their localization in definite regions or
cells. Such a definition is as applicable to the latest stages
of differentiation, such as the formation of muscle fibers,
as it is to the earliest differentiations of the germ cells,
and the one is as truly a case of inheritance as is the other.
In short, different substances appear at an earlier or later
stage in the development of all animals, and these substances
are then sorted out and localized; this is differentiation
[= heredity: see above]. Physiological division of labor in-

.

Evidence from Meia::can Ccr:n-Cdls 23

volves morphological division of substance; sorting out of
functions implies sorting out of the material substratum
of functions." 13

If cytoplasmic sorting out and localization as well in the
earliest as the latest stages of development constitute dif-
ferentiation "and hence heredity," how, one must ask, can
the hypothesis that "the chromosomes arc the scat of the
inheritance material" ever "practically amount to a demon-
stration"? The only way, so far as I can see, to reconcile
these two statements is to say that in so far as the expres-
sion "seat of inheritance material" means anything, both
chromosomes and cytoplasm are such seats, and hence that
neither is the seat of it.

But it is not enough to point out that there is contradic-
tion here. We must try to discover just how so careful a
reasoner as Conklin should fail to detect it, for we may
feel certain that the failure is due not to mere oversight or
carelessness but to some defectiveness in standpoint or gen-
eral mode of procedure. Conklin fully realizes, as our quo-
tations show, that movements of the cytoplasm go far to-
ward determining the attributes of the eggs of Ascidians
and many other animals. But the following makes the
recognition still more definite: "Undoubtedly the most im-
portant of all the localizing factors so far recognized are
cytoplasmiq movements." 14

Assuming that our contention is valid that these localiz-
ing factors of the cytoplasm are inheritance factors (and
the virtual admission of this by Conklin in one of the two
statements which we hold to be contradictory should be
noted) we have still to see by what facts and reasoning Conk-
lin reaches the view that his observations support the
theory that "chromosomes are the seat of inheritance mate-
rial." The observations in this case which support the
chromosome theory are that the three kinds of cytoplasm
of the egg: the yellow protoplasm (mesoplasm), the sphere

26 The Unity of the Organism

material (archoplasm), and the clear protoplasm (ecto-
plasm) come, at least in part, and at one time or another,
from the nucleus. It is not claimed that the chromosomes
or even chromatin can be observed to produce the cytoplas-
mic localization or any of the other distinctive features of
the egg, as for example the different colors characteristic
of different animal species ; or the different kinds of proto-
plasm in the same species. Just what can be observed as
to the relation between the chromosomes and the several
kinds of material which pass out of the nucleus into the
cytoplasm is not much dwelt upon by Conklin in this investi-
gation. The clear protoplasm (ectoplasm) he describes and
figures very definitely as being the major clear mass of the
germinal vesicle set free in the cytoplasm as the nuclear
membrane dissolves at the beginning of maturation. The
chromosomes are said to be distinguishable at this time as
numerous small deeply staining bodies. They can be ob-
served to collect together in the "center of the nuclear
area" and certain things can be made out about the shapes,
and arrangement relative to one another of the individual
chromosomes ; but nothing is recorded to indicate that they
take any part in the movements of the ectoplasm, much less
in the production of it.

Concerning the relation of the other two kinds of cyto-
plasm, the yellow or mesoplasm and the sphere, material or
archoplasm, to the chromosomes, Conklin's description is
still more meager. An investigation by Crampton, however,
has given particular attention to the relation of the sphere
material, or as he calls it the yolk-matrix, to the chromatin.
Crampton's results are the more significant for this discus-
sion in that Conklin was undoubtedly acquainted with them.
As the result of several searching tests of the chemical
character of the yolk-matrix, Crampton says : "In Molgula,
certainly, these granular masses are not chromatin in the
proper sense." 15 Although Crampton believes the substance

Evidence from Metazoan Germ-Cells 27

called by him yolk-matrix and considered by Conklin to be
the same as what he calls sphere material, arises from the
nucleus, this belief rests not on direct evidence of the pass-
age of the substance from the nucleus into the cytoplasm,
but on the facts that when the substance is first seen in
the oocyte it is a small mass situated in the cell-body or
cytoplasm close in contact with the nuclear wall, and that it
reacts to stains and digestive fluids in the same way that
certain granular contents of the nucleus react. But Cramp-
ton is very explicit in pointing out that this substance in the
nucleus is not chromatin, at least of the ordinary kind.
Upon treating the cells with digestive fluids it disappears
from both inside and outside the nucleus, the true chromatin
being then left in clear view as fine granules in the nuclear
reticulum.

So far as concerns the endoplasm, then, even though it
be derived from sphere material and this in turn from the nu-
cleus, the most trustworthy evidence we possess is to the
effect that its primal nuclear source is not chromatin. In
view of such facts as these, made known by Crampton and
others, through what reasoning would Conklin, to whom the
facts are familiar, still hold that taken all-in-all they sup-
port the chromatin dogma of heredity? Readers whose
minds have become sensitized to the general type of reason-
ing which pervades nearly all elementalistic theorizing and
makes it to some extent fallacious, will readily anticipate
about how the argument will run in this case. But it will
be profitable to see it in actuality. After saying that "some
of the important cytoplasmic substances can be actually
seen to come from the nucleus" but that "this does not indi-
cate that these substances exist from the beginning in the
nucleus ; on the contrary there is direct and visible evidence
that they arise epigenetically," Conklin continues: "Such
epigenesis, however, does not signify lack of primary organ-
ization ; on the other hand all the evidence favors the view

28 The Unity cf the Organism

that 7;«c7i of the organization of the cytoplasm is the or-
ganization of the cl.romosomcs, which is definite, determinate
and primary" 10 (Italics mine).

There yen have it again ! Although it is freely granted
that you can see the cytoplasm in the very act of arising
epigenetically and moving about to become definitely located,
that is, to become organized, still what you see is no part of
the real essence of the business. "Back" of this, in the
chromosomes, which, be it specially noticed, can not be seen
to take any active part in the operations, we must conceive
is the "organization" which is "definite, determinate and
primary" — in other words which is The Ultimate Cause, so
far as heredity is concerned. Again I repeat, wearisome
as the iteration has become, that the fallacy in this sort
of reasoning is not in holding that there is some causal
power "back" of the phenomena to be explained, but that
all such power is located there. That is, stating the general
point in its application to the special case, the fallacy lies
not in holding that the chromosomes contribute something
to the hereditary attributes of the ascidians and other ani-
mal groups whose development Conklin investigated, but in
the implied denial that the cytoplasm contributes anything
to it.

Conklin probably would not admit that there is real
contradiction between the observations by himself and oth-
ers, on the part played by cytoplasm in the early stages of
development, and his contention that the evidence now "prac-
tically amounts to a demonstration" of the correctness of
the theory that the "chromosomes are the seat of the in-
heritance material." What he would probably contend is
that the observations are opposed merely to the extreme
form of the chromosome theory. Thus, speaking from an
angle of the general subject a little different from that of
cytoplasmic localization, he writes : "This conclusion is not
a refutation of the nuclear inheritance theory, but it is a

Evidence from Mctazcan Ccrxi-Cdls 29

profound modification of it." 17 And still more recently
he has said, "Many biologists maintain that the nucleus and
more particularly the chromosomes are the exclusive seat of
the 'inheritance material' and that all the 'determiners' of
adult characters are located in them. Against the extreme
form of this theory many general and specific objections
may be urged. General objections are based upon the con-
sideration that the entire cell, cytoplasm as well as nu-
cleus, is concerned in differentiation and that neither is
capable of embryonic development in the absence of the
other. Differentiation is indeed the result of the interaction
of nucleus and cytoplasm, and how then can it be said that
the nucleus is the only seat of the inheritance material?"

An elaborate discussion of whether the language here used
can be harmonized with the statement quoted above about
the demonstration of the correctness of the chromosome
theory, by saying that the views expressed in the last quota-
tion merely involve a "profound modification of the nuclear
inheritance theory" would smack too much of pure dialectics
to deserve a place in this volume. Our sole concern is with
the truth about the thing itself.

Conklin's position would be so far satisfactory if he
would permit us to understand his statements "the entire
cell, cytoplasm as well as nucleus, is concerned in differentia-
tion," and the one about modification of the nuclear inheri-
ance theory, to mean that cytoplasm is "inheritance mate-
rial" and contains "determiners." Evidence that cytoplasm
contains "determiners" is even more positive than is that
for the theory that chromosomes are the seat of such things,
for the simple reason that we can observe abundantly cyto-
plasm in the act of producing hereditary structures, whereas
we rarely observe chromosomes operating directly in this
way. But such permission would not, I fear, be forthcoming.
If it would be, one is at a loss to understand why the terms
"hereditary substance," "physical basis of heredity," "de-

30 The Unity of the Organism

terminers," "factors" and the like, constantly used in con-
nection with the chromosomes are never used in connection
with cytoplasm. Indeed, so well does Conklin present the
general argument for the participation of cytoplasm in
the development of hereditary structures that it is surpris-
ing, not to say disappointing, to find him neglect to present
the most specific argument we have to the same effect, name-
ly that the genesis of a vast range of such structures can
be directly observed to be largely due to cytoplasmic trans-
formations.

One other passage in Conklin's general argument is so
significant that we must reproduce it. "Differentiation is
indeed the result of the interaction of nucleus and cyto-
plasm, and how then can it be said that the nucleus is the
only seat of the inheritance material? If held rigidly, this
theory involves the assumption that the cytoplasm and all
other parts of the cell are the products of the chromosomes,
and that therefore the chromosome and not the cell is the
ultimate independent unit of structure and function; an
assumption which is contrary to fact. Furthermore, since
heredity includes a series of fundamental vital processes
such as assimilation, growth, division and differentiation,
there is something primitive and nai've in the view that this
most general process can be localized in one specific part of
the cell, something which recalls the long-past doctrine that
the life was located in the heart or in the blood, or the
ancient attempts to find the seat of the soul in the pineal
gland or in the ventricles of the brain." 19

This passage contains several well-sent shafts not only
against chromosomal elementalism, but against the elemen-
talist standpoint generally. And I must recur again in con-
nection with it, while the facts of egg organization as pre-
sented by Conklin are fresh in mind, to the perception,
indicated in previous chapters, that the physical-chemistry
conception of the cell must be extended to the organism. If

Evidence from Metazoan Germ-Cells

31

Conklin once sees the full force of this contention, he will,
we may hope, be ready to let go entirely of the idea that
the facts of cytoplasmic organization must be "harmonized
with the nuclear inheritance theory." u He will then see
that there is no more necessity for harmonizing the facts of
cytoplasmic organization with the nuclear inheritance the-
ory than there is for harmonizing the facts of nuclear or-
ganization with the theory of cytoplasmic inheritance.

REFERENCE INDEX

1. Ballowitz 255

2. Wilson, E. B. ('00) 135

3. Ballowitz 27T

4. Oliver 489

5. Oliver 479

6. Casteel 646

7. Conklin ('05) Ill

8. Conklin ('05) 11

9. Conklin ('05) 21

10. Conklin ('05) 114

11. Conklin ('05) 101

12. Conklin ('08) 90

13. Conklin ('08) 92

14. Conklin ('05) 102

15. Crampton ('99) 48

16. Conklin ('05) 101

17. Conklin ('08) 98

18. Conklin ('15) 162

19. Conklin ('15) 163

Chapter XV

EVIDENCE FROM SOMATIC HISTOGENESIS IN
MULTICELLULAR ORGANISMS

WE must now give the greatest possible concreteness
to the general truths stated by Conklin that growth
is the result of the interaction between nucleus and cyto-
plasm and that heredity includes such fundamental vital
processes as assimilation, growth, division, and differentia-
tion. But the one and only way, I again insist, to attain
concreteness and certainty in the matter is through a maxi-
mum of observation coupled with a minimum of inference.
That is, the goal must be reached mainly by direct study of
the anatomical, histological and physiological transforma-
tions through which hereditary attributes are produced. The
issue can be met squarely only by a still further considera-
tion of what we actually know about the participation of
all sorts of elements of relatively undifferentiated cells in
the production of obviously hereditary parts. The study
of interactions between nucleus and cytoplasm and of as-
similation growth, etc., in germ-cells is not enough. What
we have seen in preceding pages about the development of
organs in the protozoa and in spermatozoa is that much
toward the end sought. Our task now is to consider the
local transformations by which structures are produced in
multicellular organisms, especially in those which develop
from eggs.

The Mitochondria! Theory of Heredity

This task may well begin by examining the recent efforts
to locate the "hereditary units" in the mitochondria of the

32

Evidence from Somatic Histogenesis 33

cytoplasm instead of in the chromosomes. This effort is
just as misdirected as is the effort to make chromatin the
sole hereditary substance. According to the organismal
conception, all life phenomena, including those of inheri-
tance, consist in the activities and interactivities of an
enormous number of substances and units and forces, all of
which, in exhaustive analysis, are dependent upon the organ-
ism as a living whole. It is, therefore, as futile to hunt in
one corner as another for the physical basis of heredity
in an exclusive and more or less metaphysical sense. Any
one who has grasped this idea will know beforehand that
the proposal to make mitochondria fully explain heredity is
doomed to failure no less certainly than was the proposal to
make chromosomes or any one kind of cell element play such
a role. But hypotheses, the falsity of which might have been
seen before they were tested, may still be useful. If those
who propose them can be convinced of their fallacy in no
other way than by testing them then it is better that they
should be tested even though much time and labor be given
to the task. Again, the evidence brought out which dis-
proves an hypothesis may be highly useful in establishing
some alternative general view. This result has been espe-
cially striking in the case of the mitochondrial hypothesis of
hereditary substance. By bringing the cell-body back into
the field of interest, from which it had been largely excluded
so far as heredity was concerned, by the nuclear inheritance
theory, the mitochondrial hypothesis has resulted not merely
in proving that mitochondria are not bearers of heredity
in the elementalist sense, but that in a rational sense they
are sometimes rather closely concerned in the production
of hereditary structures ; and, of even greater importance,
that still other cytoplasmic material is likewise so concerned.
The only reason why the mitochondrial theory of hered-
ity is less interesting than the chromatin theory is that
there is so much less observational evidence in support of it.

34 The Unity of the Organism

In fundamental principle the one is no more and no less
acceptable than the other.

Any half plausible suggestion that a particular minute,
obscure part of the germ-cell may be a "bearer of heredity"
seems to endow that part with a peculiar fascination to biol-
ogists who have the elementalist habit of thinking, and this
secures to it an inordinate attention until the untenability
of the hypothesis is so overwhelmingly proved that even the
most credulous are forced to abandon it.

Several biologists have recognized something of the state
of mind here indicated without, however, perceiving its real
meaning. Thus in a review of the work of Meves, which
will be examined presently, we read : "the new interpretation
which Meves gives at this time indicates that many are
still dissatisfied with the all-sufficiency of the [nuclear]
theory, and are eagerly seeking and grasping, as it were,
the first visible sign of any other substance which may serve
to carry the hereditary qualities." The remark to be
made about any statement of this kind is that the real
though usually unperceived ground of dissatisfaction is not
with the all-sufficiency of the nuclear theory of heredity, but
with the all-sufficiency of any theory that attempts to local-
ize the function of carrying heredity in some small, specific
fraction of the germ-cells; and that the attitude which
Doctor Payne well characterizes as "eagerly seeking and
grasping" which has marked so much of recent search after
the physical basis of heredity, has a large measure of genu-
ine illusion in it. Inspired by the ages-old, alluring belief
that an imperceptible final cause and explanation is hidden
somewhere within or behind whatever is grossly sensed, the
pursuit becomes "eager and grasping," which is another
way of saying that it becomes more or less irrational and
fitful.

The truth of these remarks is rather strikingly exempli-
fied in the short, somewhat feverish history of the mitochon-

Evidence from Somatic Histogenesis 35

drial theory of heredity. The meager observational support
for the hypothesis that these particular cell members are
generally bearers of heredity relieves us from the necessity
of examining it in any such detail as we examined the chro-
mosome theory. But there are several things about it of
so much importance that we must look into it somewhat.
The name mitochondria was first used by Benda for a "new
cell organ, perhaps serving a specific function." Benda's
original view was that the function served is that of the
motility of the cell. But in a later publication he presented
observations which seemed to him conclusive proof that the
mitochondria of the sperm are situated in parts of it which
enter the egg at fertilization. This last suggestion gave
an added impulse to the study of the bodies occurring in the
cytoplasm and soon many new names were applied to them,
for they were soon found to present differences in size,
shape, and reaction to chemicals. It seems, however, that
the present state of knowledge justifies us in applying to
them all the one term, mitochondria, though without imply-
ing that they are all exactly alike. They may be held to
be generically alike but specifically different.

The first investigator to set up definitely the hypothesis
that cytoplasmic elements of this class are bearers of hered-
itary qualities seems to have been Meves.

Only a few of the very many investigations since devoted
to the subject can be noticed, these being selected for their
bearing on particular aspects of the problem. In the first
place, the position of Meves himself is important. Accept-
ing the assumptions formulated by O. Hertwig in 1875 of a
"substance which carries over the beginnings (Anlagen) of
the parents to the child," 2 and that "this substance exists
(in the germ-cells) in an original, histologically undifferen-
tiated condition" ; 3 and adding his own reflection that not
all the cytoplasmic parts of the spermatozoon (for example,
the axial fiber of the tail) possess inheritance potencies, he

36 The Unity of the Organism

advanced the hypothesis that the mitochondria answer the
conditions of Hertwig's theory for the cytoplasmic part of
the male, though probably not of the female germ-cells.
Thus Meves reached the rather attractive conception that
"heredity is accomplished through protoplasm and nucleus
together." Were we to know no more than this about his
theory we might suppose his position to be that of a genuine
integrationist. However, his very next sentence does not
permit us to question any further the orthodoxy of his ele-
mentalism. "The qualities of the nucleus," he says, "are
carried over by the chromosomes, those of the plasma by
the chondriosomes." In other words the working together
of nucleus and cytoplasm which he conceives is not of the
sort which makes the part played by each contribute to all
the results, but that one set of elements produces its partic-
ular part of the total effects, while another set produces
another part of the effects.

The Mitochondria], Theory Tested by the Ontogeny of the
Spermatozoa

The utter inadequacy of this hypothesis is shown by some
of the same evidence which revealed to us the inadequacy of
the chromatin theory, namely that when we come to study
the histogenetic processes by which innumerable hereditary
attributes are produced, we find portions of the cytoplasm
other than the mitochondria taking the leading part in the
production. Perhaps no single one of the many instances
examined by us of cytoplasmic participation in the produc-
tion of attributes refutes Meves' hypothesis more complete-
ly and instructively than that of the developing sperm of the
fowl tick already described.

We have already examined this case as one of special weight
in proving that the cell-body, in contradistinction to the nucleus,
is hereditary substance. Now we must see the conclusiveness

Evidence from Somatic Histogenesis 37

with which this particular ontogeny disposes of the mitochondrial
hypothesis of heredity so far as this case is concerned.

Recurrence to the description and figures (42 a, b, c, d, e, f ) will
recall that the mitochondria, widely distributed through the cyto-
plasm in the early spermatocyte (nn.) assemble into a rather
sharply defined ring-shaped mass (m.r., figure 42b) as the cell
transformation proceeds, and finally take a position in the devel-
oping sperm about as remote as they can get from some of the
most actively and fundamentally changing parts of the organism.
(mi., figures 42 e, f, g).

There is no more possibility of explaining the development of
many of the parts of this sperm, the outer sheath, for example
(o.t., figures 36e and f ) as due to the influence of the mitochondria
than as due to the influence of the nucleus. And I point out
again for the hundredth time what the real issue is here. The
observations certainly do not exclude the possibility that the
mitochondria exercise some invisible influence on the development
of, say, the outer sheath. There may be or there may not be such
an influence. But the observations do show conclusively that
cytoplasmic portions of the cell other than mitochondria are
operative in producing the outer sheath.

It should be said in concluding this reference to the sperm of
the chicken tick that Casteel finds the mitochondria located finally
in the end of the sperm opposite that which contains the nucleus ;
and that this end goes ahead in locomotion, the motion being pro-
duced by a circlet of mobile processes at this end. From this he
believes that the contractile elements are mitochondrial in origin.
The conjecture that the mitochondria of the spermatocyte take
part in producing the motor elements of the sperm tail is perhaps
strengthened by the observations of other students, notably Lewis
and Robertson. These investigators were able to follow the mito-
chondria in the ontogeny of the living sperm directly into the
tail, where they transform into two equal threads situated along-
side the axial filament.

T~he Mitochondrial Theory Tested by Histogenesis

If this hypothesis that mitochondria in developing sperm
cells give rise to the motion-producing structures of the
sperm tail, then the mitochondria would be genuine "inheri-
tance material" for these particular elements, the hereditari-

38 The Unity of the Organism

ness of the motile elements being especially striking in the
case of the chicken tick sperm from the fact that the mode
of locomotion in this spermatozoon is almost unique. But
while the case of mitochondria and other non-nuclear parts
of the cell in the development of the spermatozoa, ought to
be conclusive that although mitochondria can not be "hered-
itary units" in any general sense, they, as well as other
cytoplasmic parts, may contribute to the production of
hereditary structures, yet it would not be so accepted, prob-
ably, by the most exacting theorists because such biologists
would not allow that a spermatozoon, being unicellular, can
have organs and parts which are subject to heredity "in
the strict sense" (i.e. in the sense of the definition of hered-
ity set up by these persons). We must, consequently, pro-
ceed to the specific task of this section; namely that of
considering what is known about the part played by mito-
chondria in the histogenesis of hereditary structures in
multi-cellular organisms.

Nearly all the studies centered upon the question of
whether the mitochondria are bearers of heredity have gone
on the assumption, quite inadequate according to my view,
that the problem is to be solved by ascertaining whether or
not the bodies are persistent cell organs, take a definite part
in fertilization, and are contributed in equal quantity by
the female and male germ-cells. In other words the assump-
tion has been that the same criteria which have been relied
upon to prove that chromosomes are bearers of heredity*
must also be applied for deciding whether or not mitochon-
dria have the same office. But numerous studies have also
aimed to follow the mitochondria in the genesis of tissues,
and herein lies the chief importance of investigations in
this domain. Not only have they greatly increased our
knowledge about the role played by various parts of the
cell in histogenesis, notably of the cytoplasmic parts, but
they have put us in possession of much precise information

E-vidence from Somatic Hist agenesis 39

as to what "hereditary substance" is.

The results and conclusions thus far reached are on the
whole so diverse, often so conflicting, that any attempt at
a general review of them would be useless for this discussion.
However, certain of the results, actual or claimed, are im-
portant. For example, at one extreme it is contended that
the mitochondria are the immediate precursors of the most
distinctive elements of all classes of adult tissues. Thus
Meves: "All these differentiations (of embryonic cells into
tissue cells) however heterogeneous they may be, arise
through the metamorphosis of one and the same elementary
constituent of the plasma, the chondriosomes. The chondri-
osomes are the material substratum lying at the basis of the
processes of differentiation, which become the specific sub-
stances of the different tissues." As an extension of this
view we learn from Lewis and Lewis and other reviewers
that the following tissues are reported on the authority of
a long list of workers to be produced by the mitochondria:
fibrillse, myofibrillae, fibrillae of epithelial cells, corneous sub-
stance, secretory granules, fat, leuco-, chloro- and chromo-
plasts, the test substance in foraminifera, and various other
tissue elements.

But several investigators, notably E. V. Cowdry, have
shown the inconclusiveness of the evidence on which the con-
tention is based that neurofibrils originate from mitochon-
dria. "There is no evidence," Cowdry says, "that mito-
chondria are transformed into neurofibrils. . . . The mi-
tochondria do not show, either by a variation in their
morphology, staining reactions, or in any other fashion,
. . . indications of being transformed into material of dif-
ferent chemical composition." 7 Furthermore, he shows that
the mitochondria do not diminish in quantity in any way
commensurate with the increase of neurofibrils, as the neu-
roblasts transform into ganglionic cells. Eminently worthy
of note, as bearing on our contention made some pages

40 The Unity of the Organism

back, that cytoplasm itself is hereditary substance, is Cow-
dry's detailed description of neurofibrils as a "differentia-
tion of the ground substance" of the neuroblasts. And in
a later paper the same author makes a strong case of the
view that while mitochondria are "associated in some way
with the formation of many substances," 8 it is highly im-
probable that they transform into them.

On the whole the tendency of the latest investigations
appears to be to deny that the bodies produce, in a strict
sense, any tissue elements. Thus as a result of their quite
remarkable studies on mitochondria of living cells Lewis and
Lewis say, following the enumeration given above: "The
above theories seem impossible to correlate. It seems evi-
dent that the mitochondria are too universal in all kinds
of cells to have the function of forming any one of the
above structures of differentiated tissue, and in the light of
what cytological chemistry is known, it appears practically
impossible for the mitochondria to form all the cell struc-
tures mentioned above. In view of the fact that the mito-
chondria are .found not only in almost all animal cells but
in plant cells as well it seems more probable that they play
a role in the more general physiology of the cell." The
idea that the mitochondria are primarily concerned with the
metabolism of the cell appears to be gaining ground under
the present comprehensive and critical methods of investi-
gation that are being applied to them.

The Untenable Hypothesis that the Cytoplasm of the Ovum

is Inheritance Material for General but not for

Special Characters

A number of biologists have recently put forward the
hypothesis that while the cytoplasm of the egg-cell may be
"hereditary material" for certain of the general attributes
of the organisms, chromosomes "carry" the hereditary,

Evidence from Somatic Histogenesis 41

of the more specific attributes. This conception has arisen
from a considerable range of observations to the effect that
for quite a time in the early ontogeny of many animals some
of the attributes of the embryo can be seen to come directly
from the cytoplasm of the egg. Thus both Driesch and
Loeb have taken special notice of the fact that, as expressed
by Loeb, "when the protoplasm of the egg possesses a strik-
ing pigment the larva will possess the same for some time
at least"; and that "if such an egg is hybridized with the
sperm of a form whose egg is unpigmented, the larva will,
of course, possess a 'maternal' quality which is due solely
to the protoplasm (Driesch)".10 And in the same connec-
tion, Loeb continues : "It is obvious, then, that during the
first stages of development an influence of the protoplasm
upon heredity may make itself felt, which will disappear as
soon as the protoplasm of the egg has been transformed into
the tissues of the embryo." One of the cardinal questions we
have to consider may be formulated in connection with this
last quotation: Have we a right to assume that because
an obvious influence of the protoplasm upon heredity dis-
appears on the transformation of the protoplasm into tis-
sue, therefore all such influence of the protoplasm ceases?
To answer this question through observations upon the
protoplasm of the cells concerned just before, during, and
just after the transformations is exactly the central aim
of this section. I can not refrain from making use of
another sentence from Loeb to aid in defining the problem
more clearly. "It does not seem to me," he writes, "that a
discussion as to the relative influence of protoplasm and
nucleus upon heredity will prove very fertile, but that it is
necessary to transfer this problem as soon as possible from
the field of histology to that of chemistry or physical chem-
istry." 10 I quite agree that "discussion as to the relative
influence of protoplasm and nucleus upon heredity" can not
be very fruitful. But the grounds of my skepticism are

42 The Unity of the Organism

widely different from those of Loeb. According to my view,
the question is not one to be settled by discussion at all,
but by observation coupled with a measure of consistent
reasoning. Assuming that I am right, to "transfer" the
problem "from the field of histology" if this really means,
as it seems to, that the problem should be taken away from
histology, no matter whether to the field of chemistry or
any other, would be to remove it all the further from ob-
servation and plunge it so much the deeper into discussion.
I have not the slightest doubt that chemistry, especially
biochemistry pursued on the principles of physical chem-
istry, will have to be made large use of before the fullest
possible understanding of the mechanism of heredity is
reached. But this use will have to go hand in hand not
only with morphological studies on germ-cells, but as well
on hosts of cells during the whole ontogeny. Chemical in-
vestigation will have to supplement, it cannot supplant, it
cannot even lead, histogenic investigation. If there is one
thing made more positive than any other about heredity by
modern study of the subject, it is that heredity is some-
thing which pertains to the smaller taxonomic grades of
organisms, races, varieties, species and so forth. It would
seem, accordingly, that hardly any suggestion for the study
of heredity could be wider of the mark than one to trans-
fer it from the only field which makes any pretense of in-
vestigating the details of development, and taking it into
a field like that of physico-chemical activity, which is no-
toriously devoid of the very attributes without which there
would be no such thing as heredity. Having once ascer-
tained by observation as much as possible about how hered-
itary attributes are actually produced, it will then be in
order to learn as much as possible about the chemistry of
the processes. Chemistry can do its share in solving the
problems of heredity after and not before histogenesis has
done its share.

Evidence from Somatic Histogenesis 43

Conklin has expressed more definitely than any other biol-
ogist with whose writings I am acquainted, the idea men-
tioned above, that cytoplasm "influences" heredity in early
ontogenetic stages, and also influences adult attributes of
the major taxonomic groups, but becomes inoperative in
the later stages of development, the heredity of these being
transferred to the nucleus. He says, "In slwrt, the egg
cytoplasm fixes the general type of development and tJie
sperm and egg nuclei supply only the details." And fur-
ther: "We are vertebrates because our mothers were
vertebrates and produced eggs of the vertebrate pattern;
but the color of our skin and hair and eyes, our sex, stature,
and mental peculiarities were determined by the sperm as
well as by the egg from which we came. There is evidence
that the chromosomes of the egg and sperm are the seat of
the differential factors or determiners for Mendelian char-
acters, while the general polarity, symmetry, and pattern'
of the embryo are determined by the cytoplasm of the
egg." n If two points in this last quotation be viewed in
the light of a large mass of relevant evidence not usually
taken into account in recent discussions on heredity, and
if strict consistency in the use of terms be maintained, the
general conclusion will be quite different from that stated
by Conklin. These two points, the conception of "differen-
tial" and of "determiner," must now receive attention ; but
first I will illustrate my position by a case presenting the
kind of evidence to which I have referred.

Species Attributes in Single Cells of Adult Organisms

In general, this evidence comes from the field of histology,
or more strictly histogenesis. The most convincing, because
the most direct, evidence from this source is that pertaining
to the development of hereditary structures in adult meta-
zoa and metaphyta. The structures in such organisms

44 The Unity of the Organism

which are the most indubitably hereditary are those which
distinguish the smaller but yet definite taxonomic groups.
A little consideration will convince one that about the most
crucial cases would be those in which the development of
differential attributes could be traced directly to cell struc-
ture and development. It so happens that vast as is our
knowledge of histogenesis, the part of it which answers
directly to the requirements here laid down is by no means
large.

The S pinnies of the Ascidian Genus Styela

The best instances I have been able to find are super-
ficial appendages in some animals and plants, so small that
they consist of a few cells or even of a single cell. One strik-
ing instance of this sort has come to light in my own studies.

FIGURE 44. SPINULE CELL OF STYELA YAKUTATENSTS (AFTER HUNTSMAN).

n., nucleus.

It is that of the minute spines which cover the inner surface
of the branchial siphon in some species of the ascidian group
of Styalids. Huntsman first called attention to the fact
that each spinule is a single cell, and that at least in some
cases the structures furnish differentiating attributes for
species. Miss Forsyth and I have reexamined the point for
Styela montereyensis and S. yakutatensis, and are able to

Evidence -from Somatic Histogenesis

45

confirm Huntsman's results. As Huntsman had opportun-
ity to study the matter in a larger number of species than
Miss Forsyth and I have had, the following description is
taken largely from his paper. Figures 44a, b, 45, 46 are
from Huntsman and figure 47 is from Ritter and Forsyth.
By comparing figures of what may be called the dorsal view
(figures 44a, 45, 46) with the side view (figure 44b) it is
seen that the distinctive feature about the cell which con-
stitutes the spinule is a shield-like plate on one side of the
somewhat elongated cell, the distal end of which projects
more or less beyond the cell body, the whole resembling to

FIGURE 45. SPINULE CELL OF STYELA PLICATA (AFTER HUNTSMAN).

some extent the end of a finger with its nail. The shield
is harder than the rest of the cell, and is probably com-
posed of the same material as the general "test" of Ascid-
ians, animal cellulose. The spinules are so placed that the
basal portion is embedded in the surface layer of the test
on the inside of the siphon, the shield being on the free side
of the cell with its free edge pointed toward the lumen of
the siphon. The specific attributes furnished by the
spinules depend upon the shape and structure of the shields.
The free edges may be truncate (figure 44, S. yakuta-
tensis), or long-pointed (figure 46, S. greeleyi), or low-
conical (figures 45, 47, S. plicata, S. montereyensis) .
Again the edge may be smooth (figures 46, 47, 5. greeleyi

46 The Unity of the Organism

and S. montereyensis) , or it. may be serrate (figures 44, 45
S. yakutatensis, S. plicata).

Viewing this case in the light of considerations put for-
ward on the preceding pages, the pertinent queries about
the heredity of the shields almost ask themselves : What is

FIGURE 46. SPINULE CELL OF STTELA GREELEYI (AFTEE HUNTSMAN).

the "inheritance material" that causes the shield to be short
and truncate in S. yakutatensis and long-pointed in S.
greeleyi; or that explains the serrated edge in S. yakuta-
tensis and S. plicata as contrasted with the smooth edge in
S. montereyensis and S. greeleyi? Is the "seat" of that ma-
terial in the cytoplasm or the nucleus of the shield-produc-
ing cell? Unfortunately we have no direct observational
information about the genesis of the spinules. But the

Evidence from Somatic Histogenesis 47

indirect evidence which bears on the point favors over-
whelmingly the view that the cytoplasm is chiefly responsible
for the shield. Huntsman supposes the spinules to be de-
rived from the cells of the cellulose tunic characteristic of
ascidians. He may be right in this ; but it may be, too, that
they are derived from the epithelial lining of the siphon.
The matrix of the cellulose tunic is undoubtedly largely if
not wholly produced by the ectodermal cells. It is usually
held to be secreted by these cells; but in some cases, in

FIGURE 47. SPINULE CELL OF STYELA MONTEREYENSIS (AFTEE RITTER AND

FOHSYTH).

Perophora for example, a portion of the cytoplasm of the
cells seems to become transformed into the cellulose. The
process of transformation can be particularly well seen in
the cells which line the branchial siphon of developing blas-
tozooids as shown in figure 32, plate III of my memoir.12
The parts of the cell-bodies turned toward the cellulose are
here long drawn out and the protoplasm gradually becomes
indistinguishable from the surrounding cellulose substance
in which it is imbedded. If now we imagine this protoplasm
to transform not into the characteristic cartilage-like cellu-
lose mass spread over nearly the whole surface of the body,
that from innumerable cells fusing into a common mass, but
each cell to retain its individuality, its protoplasm becoming
the shield of a spinule, we should have what these styelids

48 The Unity of the Organism

actually present.

But whatever be the cells which transform into the spin-
ules, all the available evidence indicates that the cytoplasm
is the chief source of the shield part of the spinule. The
possibility is not excluded that the nuclear chromatin also
plays some part in the development. Indeed investigations,
notably those by Duesberg on the ascidian egg made since
"cytoplasmic inclusions" have come into prominence are
distinctly favorable to such an hypothesis. But this is very
far from proving that such chromatinic influence, assuming
it to exist, reduces the cytoplasm to pure passivity.

In the light of what is here set forth let us examine the
view expressed by Conklin and quoted above that the egg
cytoplasm fixes the general type of development, while the
nuclei of egg and sperm together "supply only the details."
The examination should be the more cogent from the fact
that Conklin's idea was based largely on his investigation
of ascidian eggs, some of which pertained to the very same
genus, Styela, as do the spinules just described. The reader
should not fail to notice that both Conklin and myself are
dealing with single cells, he having to do with egg-cells, while
I am concerned with spinule cells. Nor should the fact be
lost sight of that these two categories of cells stand at
about opposite extremes in the life career of an individual
Styela, the egg being at. the very beginning, while the spinule
is produced at or near the completion of adulthood. Since,
however, both cells possess attributes distinctive of the
species, there can be no more "filling in of details" at one
end of the series than at the other, so far as concerns the
differential attributes under consideration.

Now comes the main point. The differential attributes
of the egg-cell and early embryo, their "polarity, symmetry,
and pattern," are "determined by the cytoplasm," to use
Conklin's own words. What is the evidence that these attri-
butes are thus determined? That of direct observation,

Evidence from Somatic Histogenesis 49

as the examination of Conklin's work has shown. But if on
the basis of such observation it can be asserted that attri-
butes of egg and early embryo are determined by the cyto-
plasm, how escape asserting that the same sort of evidence
touching the production of the adult attributes, those per-
taining to spinule cells, are likewise determined by the cyto-
plasm ?

It was remarked above that if all known relevant facts
were taken into account and consistency in terminology be
maintained, Conklin's statement to the effect that we are
vertebrates because our mothers were vertebrates and pro-
duced eggs of the vertebrate type, but that our species and
racial characters, color of skin and hair, and so forth, are
determined by the chromatin of both parental germ-cells,
would have to be greatly modified. We are now in position
to see what modification is necessary. Although the state-
ment is undoubtedly true that we are vertebrates because
we develop from vertebrate eggs, the implication that the
attributes which identify us with the human species and the
Caucasian race are explained, so far as heredity is con-
cerned, by the chromatin of the germ-cells, whether male or
female or both, is not in accordance with all the observed
facts bearing on the problem. The same kind of evidence
on which the assertion is based that the embryonic charac-
ters are determined by the egg cytoplasm, requires the
assertion that skin and other adult characters are deter-
mined by the same means.

This leads to the remarks we have to make about con-
sistency in the use of terms. The critical reader will hardly
have failed to notice the difference in application of the
word "determined" as used by Conklin in the quotation
we are examining. When it is said that the color of skin,
hair, stature and so on are determined by the germ-cells,
the determining act or condition is far removed from that
which is determined, and no direct causal connection be-

50 The Unity of the Organism

tween the two is established. On the contrary in the state-
ment that the polarity, symmetry and pattern of the egg
are determined by the cytoplasm, the determination is im-
mediate and observed. Manifestly in a literal sense "de-
termined" is properly used in the second connection but not
in the first. The cytoplasm is operative on the spot, so to
speak, in the second case. It is concerned with an immedi-
ate result. The germ-cells on the other hand are not really
determiners. They are not concerned with an end result,
but are if anything instigators of a long developmental
series at the far end of which appear the attributes in
question: skin, color and the rest.

If this case of spinule production stood alone as an
instance of specific characters in adult animals traceable
to cytoplasmic activity of individual cells, it would be a
rather small base on which to erect a general argument in
favor of cytoplasm as inheritance material. But it does
not stand alone. Indeed, the company to which it belongs
will almost certainly be found to be legion when systematic
investigation of the subject shall have been made.

The Spicules of Sponges and Other Invertebrates

I will cite a few more cases. In several widely separated
groups of animals, spicules, usually either calcareous or
silicious, are present in some of the tissues. These are often
produced by one or a very few cells, and often, too, their
shape, size, and probably other attributes differ from
species to species even of the same genus.

The spicular system reaches its greatest development and
has been most studied in sponges. "The spicules of
sponges," writes Sedgwick, "in the diversity, symmetry, and
intricacy of their form, in the perfection and finish of their
architecture, constitute some of the most astonishing ob-
jects in natural history." 13 Figure 48 gives a hint of the

Evidence from Somatic Histogenesis

51

facts on which this statement is based. Although the prob-
lem of what all these spicules are for does not directly con-
cern us, indirectly it does, as the following further remarks
of Sedgwick will indicate. "While it is pretty clear," he
says, "that the main function of the skeletal structures is
the support and protection of the sponge body, it is by no
means easy to give explanations of the diversity and com-

FIGURE 48. SPICtJLES OF SPONGES (AFTER LAXKESTER).

plexity of form which they present. The form of the megas-
cleres is probably connected with the form of the canal sys-
tem with which they are in relation (F. E. Schulze) ; but the
form and even the existence of the microscleres defies any
reasonable explanation." And then comes this statement,
highly significant for almost any discussion of heredity:
"By some spongologists the small spicules are regarded as
functionless, and as having on that account a greater value
for classificatory purposes." 13

If any one wishes to be convinced of the extent to which
the spicule forms differ with different species, he should

52 The Unity of the Organism

consult such systematic monographs as those of Schulze
and Sollas in the reports of the Challenger Expedition. A
picture of such a group of spicules as that shown in figure
48 reminds one of pictures of ice crystals he has seen; and
the question may well be raised, Are not these spicules in
reality crystallization forms, and hence as devoid of hered-
itary significance as are snow flakes? The fact that the
form they have depends on the particular group of sponges
to which they belong, i.e., that they follow the rules of
biological taxonomy, is very strong evidence that they are

FIGURE 49. DEVELOPMENT OF ,A SPICULE (AFTER LANKESTER).

genuine organic productions, and not mere crystallizations.
And the further fact that they follow this rule even though
many of them appear to have no functional significance in-
dicates that their particular forms are due to heredity and
not to modeling by extraneous influences in each individual
sponge. But we are not left to such general evidence for
support of the supposition that they are true organic
products and subject to heredity. Their development has
been studied by several zoologists and the results leave no
doubt about their nature so far as this point is concerned.

A single instance will be enough for our purpose, but it should
be remarked that many others could be given. This is taken from
the excellent summary of what is known about sponges written
by Minchin for Lankester's Treatise on Zoology. "To form a
triradiate spicule three cells migrate into the parenchyma from

Evidence from Somatic Histogenesis 53

the dermal epithelium and become arranged in a trefoil-like figure
(Figure 49). The nucleus of each cell then divides into two, in such
a way that one nucleus is placed more deeply and one more super-
ficially. Between each pair of sister nuclei a minute spicule ray
appears, the three rays being at first distinct from each other but

FIGURE 50 — SEE

soon becoming united at the center of the system (Figure 50 tr.
syst.*). As the rays grow in length the protoplasm of each actino-
blast becomes aggregated around each of the two contained nuclei
and finally more or less completely segmented off to form two
formative cells, of which the one placed more internally travels
to the tip of the spicule ray, while the other remains at the base

FIGURE 51 SEE 49.

b.f.c., basal formative cell, tr.syst., triradiate system.

(Figure 51, b.f.c.}. The apical formative cell sooner or later dis-
appears, returning, apparently, to the epithelium. The basal
formative cell remains at the base of the ray (Figure 51), until
this portion is secreted to its full thickness. It then migrates
slowly outwards along the ray, and in the fully formed spicule is
found adherent to the extreme tip."

54 The Unity of the Organism

That such a mode of development is entirely foreign to crystal
production hardly needs to be remarked. But if further proof
to the same effect were demanded, one other strong piece of evi-
dence is the fact that most of the spicules are not composed of
inorganic substance alone but have a core of organic matter.

Although the technique for the examination of cytoplasm de-
veloped during these last years has not, so far as I know, been
applied to the spicule-producing cells of sponges, we can be
reasonably sure from the study of other secretory cells what the
general results will be when such application shall have been
made. They will bring out numerous details not now known of
how both the nucleus and the cytoplasm act during spicule pro-
duction.

Surely it is not necessary for me to dwell again on the
main point of the evidence here presented. The nuclei of
the spicule-forming cells may take an active part in pro-
ducing the spicules. Indeed from our general knowledge of
nuclear activities, illustrations of which were given in an
earlier chapter, it is probable that such will some day be
demonstrated to be the case. But the proof of nuclear
activity in spicule production will not be disproof of the
already observed cytoplasmic activity in spicule production.

Other animals that may be mentioned in which spicules
are produced in much the same way and have the same taxo-
nomic diversity and constancy are the alcyonaria among
coelenterates, the holothurians among echinoderms, and
some of the compound ascidians, particularly of the family
didemnidae. Relative to the specificity of the structures in
holothurians, we have this piece of significant information :
"These calcareous bodies are of great value to the system-
atist in classifying the smaller groups, such as genera and
species. Although their general characteristics are fairly
similar within the several families, the different shapes of
spicules are not sufficiently constant to be used as diagnos-
tic characters of such large divisions."15 In other words,
so far as these animals are concerned, should it be found,

Evidence from Somatic Histogenesis

55

as it almost certainly will be, that the cytoplasm is "hered-
itary substance" for the production of spicules, the reverse
of Conklin's generalization that the cytoplasm determines
the larger taxonomic features of animals while chromatin is
the seat of the inheritance factors for "filling in details"
turns out to be true. Species-marking details are just
what we are able to see the cytoplasm fill in.

The "Hairs" of Higher Plants

For a few more instances of species characters in multi-
cellular organisms brought down to single cells, we turn to
the plant world.

FIGURE 52. FIGURE 53. FIGURE 54.

FIGURE 52. TRICHOMES OF PAPAVER ORIENTALS (AFTER CANNON).

FIGURE 53. TRICHOMES OF P. PILOSUM (AFTER CANNON).

FIGURE 54. THICHOMES OF P. SOMNIFEHUM (AFTER CANNON).

The "hairs" or trichomes borne on the leaves, flowers
and smaller stems of innumerable flowering plants are
usually composed of only a few cells, so that the characters
they have are often referable to the individual cells. Can-
non has lately investigated these structures in several groups
of plants, and while he was not aiming at the particular
question now occupying us, some of his results are quite to
the point for this discussion. An example of particularly
distinct specificity of the hairs is presented in the following:
"The trichomes of the three species [of poppy] are similar

56 The Unity of the Organism

in form and size, but they are unlike in quality of rough-
ness. In Papaver orientate (52 a, b) and Papaver pilosum
(53 a, b,) the distal ends of the superficial cells project be-
yond the general surface of the trichome and turn out at
a rather acute angle. In Papaver somniferum, however,
these cells did not extend beyond the general surface, with
the effect that the trichomes of this species are smooth (fig-
ure 54 a, b)." 16

This is especially instructive because the attribute in
question pertains to the shape and position of cells, and not
to differentiation within the cells. There, consequently, is
no room for even a reasonable surmise that the attribute is
explained by the chromatin instead of the cytoplasm. Even
though the account gives no information about the position
and behavior of the nuclei of the cells, it is hardly con-
ceivable that any one would maintain that the substance
itself of the cell-tips is not the main factor in the out-
turning of these tips characteristic of Papaver orientate
(figure 52 a, b).

Dealing with quite another type of trichomes, those of
the walnut, Cannon writes: "A character which easily dis-
tinguishes the short secreting trichomes of Juglans cali-
fornica from those of Juglans regia or Juglans nigra
is the length of the head-cells." 17 Both drawings and tables
of measurements of the heads showing lengths and diameters
are given to bring out the positiveness of the distinction.
Furthermore, details of cell division and cell structure dur-
ing the development of the trichomes are furnished; so the
visible evidence that the cytoplasm is "inheritance material"
in this case is beyond question. What the invisible evidence
may be remains for further investigation to discover, but of
one thing we may be sure: no matter how many facts of
development now invisible may later become visible they will
not destroy the validity of the present visible evidence.

How far it would be possible to go on pointing out specific

Evidence from Somatic Histogenesis 57

characters of plants that are referable to individual cells I
do not know; but judging from the instances that come to
view even in my limited knowledge of the subject it might
be carried to an almost indefinite extent. Another instance
which I recall from the experience of my student days is
the case of mosses. The serration of the leaves, I remem-
ber, was one of the features relied upon for generic and
specific characteristics, and I also remember that the indi-
vidual teeth often if not always consisted of one or a very
few cells.

Cell-wall Structures In Higher Plants

That the cell-wall is a structure of great importance in
plants is known to everybody; and the veriest tyro in plant
histology has learned something of the enormous variety
and definiteness of character in different tissues and dif-
ferent kinds of plants presented by this part of the cell.

A very brief reference to two plant structures, pollen-
grains and wood tissue, will be, perhaps, a sufficient re-
minder of what there is for us in this domain. A typically
formed pollen-grain is a minute spheroidal body containing
two cells, one known as the antheridial or germinative cell
and the other as the sterile or vegetative cell. The wall of
the grain consists of an outer coat, the exine, and an inner,
the intine. The elaborateness of structure which these coats
may reach is astonishing if regarded in the crepuscular light
of the theory that cells are "simple" things. The most dis-
tinctive thing about the pollen-grain is the pollen tube
which is produced on one side of the grain and through
which the antheridial cell reaches the ovule in fertilization.
The taxonomic variety which is our main interest just here
pertains largely to the sculpturing of the surface of the
exine and to the structure of the exine at the point where
the pollen tube will break through. It is well known to
botanists that the "spikes, warts, ridges, combs, etc." of

58 The Unity of the Organism

the surface of the grains are in general definitive of taxo-
nomic groups of plants. And concerning the places of
emergence of the pollen tube we read: "The number of
these peculiarly organized points of exit is a fixed one in
each species, and often in whole genera and families."
As to the way these various structures are produced we have
this very definite statement: "The sculpturing upon the
outer surfaces of the spores of mosses and ferns and the
corresponding pollen grains of the Phanerogams can in
most cases be attributed to the activity of the protoplasm
surrounding the developing spores." 19

That the main tissues of plants present taxonomic char-
acters is amply illustrated in the wood-tissues. The facts
concerning these tissues have been almost forced into
prominence by the needs of fossil botany, though they have
also been much studied as a part of ordinary plant mor-
phology. The section on fossil wood in Zittel's Handbook of
Palaeontology is a considerable resume of knowledge in this
field, and contains numerous statements and figures which
bring out impressively the general truth of the specificity
of the tissues of trees. After necessary allowance is made
for the strictly botanical unsatisfactoriness of many of the
species and genera recognized by palaeobotanists, it is not
doubted, so far as I know, that on the whole the kinds of
tissue they describe do in reality represent different kinds
of trees. A single illustrative quotation will serve to make
concrete what is here dealt with in general terms : "The
phloem segments, like those of the xylem, are divided by
few-seriate pith rays into rather regular two- to four-
seriate rows of cells made up of thin-walled, small-celled
elements in the trunk of Zamia ftoridana, etc., and Stan-
geria. But ... in the trunks of Cycas, Dion, Encepha-
lartos, Macrozamia and doubtless most cycads, as likewise
in the Cycadeoideae, sclerenchymatous elements are more
or less numerously and regularly interspersed among the

Evidence from Somatic Histogenesis 59

row cells, thus giving much more strength to the stem." 20
And the author goes into considerable detail in discussing
the presence and the absence of "wood tracheids," "scalari-
form pittings," "border pits," "spirally thickened walls"
and "sieve tubes" in the genera Zamia, Cordaites, Stangeria
and Cycas.

That this well-nigh endless variety of character of the cell-
wall in adult plant tissues is due to the activity of the cell
protoplasm appears never to be questioned by botanists
so long as they are dealing with the actual structure and
development of the wall. "The cell-membrane is produced
by the protoplasm," we read in the section on morphology
in the Lehrbuch der Botanik by Strasburger, Jost, Schenck
and Karsten (llth German edition) this section being from
the pen of Strasburger himself. This simple, unqualified
statement of fact by Strasburger is the more noteworthy
because, as we saw in another connection, he has been one
of the extremists on the chromosome dogma of heredity.
Reading his statement that the cell-membrane is produced
by the protoplasm, with the indubitable fact in mind that
this membrane presents innumerable characters which are
taxonomically definitive, and hence are hereditary according
to the best criteria we have of hereditary characters, it
seems impossible to avoid seeing that it implies an irrec-
oncilable contradiction of Strasburger's often-repeated
view that the chromosomes are the sole bearers of heredity.

To round out the primarily factual part of this discussion
two questions remain to be considered : first, that of heredity
in the main classes of tissues of multicellular organisms ;
and second, that of the results being reached by the latest
methods of cytoplasmic study on the behavior of different
portions of the cells in the histogenesis of these tissues.
What is implied in these two questions can be made clear by
a special case. Is the minute structure of striated muscle
tissue, for example, subject to heredity? If so, are the

60 The Unity of the Organism

hereditary attributes determined by the chromatin or by
the cytoplasm of the myogenic cells? Applying our usual
test for hereditary structures, we ask whether or not this
tissue presents attributes characteristic of the taxonomic
groups, species, genera, families and so on. Here again,
while we have a vast store of knowledge about the structure
and genesis of muscle tissue in many kinds of animals, only
incidentally, as a rule, have the studies been made from the
taxonomic standpoint.

The Morphology of Striated Muscle Fibers

For one series of very recent studies that comes near this
standpoint we are indebted to H. E. Jordan. On the taxo-
nomic aspect of the matter Jordan writes: "A quite gen-
eral consensus of opinion considers them more or less closely
related, and ranks them both between Crustaceans and
Arachnoids. Lvmukis muscle, however, is in appearance very
much more like vertebrate than like insect muscle; while the
muscle of the marine arthropod Anoplodactylus is of the
typical insect type." 21 Of the numerous differences between
the fibers of the two animals compared, reference to two will
suffice for our purpose. They concern the so-called M
and Z lines found in the light bands of most striated muscle
tissue. What Jordan regards as one of the important re-
sults of his work on Limulus muscle was the evidence secured
that the Z line represents a membrane, as some observers
have believed, the specially convincing evidence being the
fact that the "line" is attached to the sarcolemma periph-
erally and to the nuclear wall centrally. These rela-
tions are lacking, he says, in the sea-spider's muscle. The
M line, he tells us, is especially well developed and hence
easily demonstrated in the sea-spider muscle in some states
of contraction, while he failed, as have other students, to
detect it at all in the Lvrmdus muscle.22

Evidence from Somatic Histogenesis 61

So we come again to the real issue. Assuming the Z
membrane to be a cytoplasmic structure, as it has practi-
cally always been held to be, are we going to deny that the
cytoplasm itself causes the peculiarity of the Z membrane
in the sea-spider as compared with that in Limtdus, that
denial being necessitated by the dogma that the real "seat"
of the difference is the chromatin of the nucleus operating
by some invisible "factor" perhaps of the nature of an
enzyme?

The extent of variety in striated muscle tissue is brought
impressively to view in such a comparative study as that
by Marceau. His main object is not to find differences but
to discover whether in spite of structural differences they
have similar traits, as if they might all be derived from a
single primitive form which has undergone more or less
profound modification.

Of the many differences which the investigation sought to re-
duce to orderliness on the basis indicated, only two will be men-
tioned. From an elaborate table of measurements of the diameter
of fibers, we find the following results for the sheep and pig : 23

Maximum Minimum Average

Sheep 25/i 5/t 15p,

Pig 45 5 20

The other point selected concerns "striated scleriform trans-
verse bands" characteristic of the muscle fibers of the vertebrate
heart. This time the animals we choose are the horse and cow.
The thickness of the band is given as exactly the same in these
two, but the distance between the bands is 140ju for the horse and
120/i for the cow.24

The Physiology of Muscle Fibers

We might go on almost endlessly, pointing out slight but con-
stant specific differences that involve differences in muscle struc-
ture, questioning in each case whether the hereditary cause of
this difference lies in the cytoplasm or chromatin of the muscle

62 The Unity of the Organism

cells. But such repetition would be useless for the present dis-
cussion. Striated muscle tissue is specially favorable for testing
hypotheses about inheritance material from the functional side
as well as from the structural s.ide. For example, there are innu-
merable differences, larger and smaller, in the limb movements
of animals belonging to different species and genera. I know of
no observations which precisely connect activities of this sort
with muscle tissue; but information concerning the electromotor
force in various animals is available. The following on the au-
thority of Englemann, taken from Winterstein will serve our
purpose. The values are those of the "demarcation current" of
galvanic electricity of heart muscle, this current being generated
by making a cut surface at the base of the heart and the natural
surface at the apex act as a galvanic pile: 25

Animal species Electromotor force in D.

Anguilla fluviatilis 0.0265

Rana esculenta 0.031 1

Triton cristatus 0.0124

Tropidonotus natrix 0.036

Testudo graeca 0.022

Columba livia 0.0458

Cygnus oler 0.0168

Mus musculus v. albino 0.040

Mus rattus v. albino 0.0446

Lepus cuniculus 0.0363

These investigations appear to have been made from the stand-
point of general physiology, and therefore not to have been car-
ried out with the systematic exactness and exhaustiveness de-
manded for taxonomic discrimination. We may consequently
presume that more searching examination of the same series
would considerably modify these results, but we have no reason
to suppose that they would eliminate altogether the differences
due to the animal species^

After due allowance is made for the purely physiological
and environic causes which undoubtedly explain a great
many of these differences, probably no biologist would hesi-
tate to grant that many of them have an hereditary basis.
Turning again to the question of the seat of the hereditary

Evidencr from Somatic Histogenesis 63

factors, we are now especially attracted by the functional
aspect of the subject. To an unsophisticated physiologist
studying the phenomena involved in this question, it would
probably never occur that more than one answer is possible.
Well-informed as such a physiologist may be supposed to be
on the important part known to be played by the nucleus in
the life of the cell, he would undoubtedly take it for granted
that the whole nucleus, its chromatin with the rest, con-
tributes in some fundamental way to the result. But unless
well indoctrinated beforehand with the chromosome dogma
of heredity, he would almost certainly be amazed were
some one to contend seriously that the cytoplasm is not the
material basis of the hereditary peculiarities exhibited. He
would reply, "Why, you are virtually denying that the
substance of the muscle fiber is the real seat of muscular
activity, thus implying a contradiction of the 'universally
accepted principle that the potential chemical energy of the
muscle substance is the primary source of muscular energy
in all its manifestations' 2e for surely muscular energy 'in
all its manifestations' would'include those elements of mus-
cular activity which are hereditarily distinctive of different
kinds of animals."

That the cytoplasm is at least the main source of the
muscle substance furnishing this energy would not be ques-
tioned, probably, by any histologist, but the definiteness of
view held at the present time on this subject is worth re-
calling and is indicated by such statements as the following:
"The energy of contraction is the transformed surface-
energy of the ultimate structural elements or colloidal par-
ticles (submicrons) composing the fibrils."27

Presumably there would be much difference of view among
physiologists as to the validity of the chemico-physical part
of Lillie's theory of muscular contraction ; but apparently
there would be very little dissent from that part of his view
which locates the processes, whatever their exact nature, in

64 The Unity of the Organism

the muscle fibers. So it would be merely a matter of suf-
ficient patience to go over all the tissue systems, epithelial,
glandular, bony, nervous, and the rest, and point out nu-
merous certain, and innumerable probable instances of dif-
ferences for different taxonomic groups of animals, and to
show that these hereditary differences are expressed pri-
marily in the cytoplasm of the cells.

Summary of Positive Information about the Physical Basis

of Heredity

We have explored a vast region of fact and theory con-
cerning propagation and development in organisms, for the
purpose of ascertaining what is actually known about the
organs and substances by which hereditary attributes are
produced. Expressing the matter in terminology familiar
to current discussion on heredity, we have been trying to
find what is actually known about the physical basis of
heredity. If clear-cut, unequivocal information of the
kind sought is contained in all»we have seen, it ought to be
statable in a few simple sentences.

What has been accomplished may be epitomized in two
such sentences:

First. Overwhelming observational evidence has been se-
cured that the cytoplasm of cells participates directly m
the formation of organic parts which have hereditary at-
tributes.

Second. A great mass of evidence, partly of observation
and partly of legitimate inference from the principles of
organic integration, has been secured, that the chromosomes
of the germ-cells in plants and animals which propagate by
means of such cells, participate in the production of or-
ganic parts having hereditary attributes.

Any substance which plays such parts in development
may be named a physical basis of heredity; and these two

Evidence from Somatic Histogenesis

65

groups, or categories of knowledge must, it seems, serve as
the foundation of all legitimate reasoning about such "basis
of heredity," or "inheritance material," or "hereditary fac-
tors," or "bearers of hereditary qualities," or whatever ex-
pression for the idea be employed.

REFERENCE INDEX

1. Payne 190

2. Meves 816

3. Meves 81T

4. Meves 820

5. Meves 850

6. Meves 845

7. Cowdry ('14) 416

8. Cowdry ('16) 436

9. Lewis and Lewis 393

10. Loeb ('06) 181

11. Conklin, ('15) 176

12. Ritter ('93)

13. Sedgwick 82

14. Minchin ('00) 107

15. Goodrich 224

16. Cannon 13

17. Cannon 23

18. Goebel 367

19. Campbell 51

20. Wieland 197

21. Jordan ('16) 493

22. Jordan ('16) fig. 7 496

23. Marceau 273

24. Marceau 280

25. Winterstein Ill

26. Luciani Ill, 85

27. Lillie, R. S. ('16) 255

Chapter XVI

THE INHERITANCE MATERIALS OF GERM-CELLS
INITIATORS RATHER THAN DETERMINERS

Antecedents of the Cytoplasmic and Nuclear Theories of
Inheritance Material

FOR the purpose of calling vividly to mind the character
of the evidence on which the two propositions rest
with which the last chapter ended, it will be profitable to
cast a glance back on the course along which biology has
come down to us, with a view to finding a sharply out-
standing spot in the early growth of knowledge which led to
each of them. On the botanical side such a spot in the
knowledge of cytoplasm as hereditary substance, is the
work of Schleiden on the microscopic structure of adult
and developing plant tissues. The publication of his Ueber
Phylogenesis, 1838, may be taken as the starting point of
our knowledge of cellular transformation in the production
of tissues. It should be remembered that the observers of
that period had very hazy notions about the distinction
between nucleus and cell-body, or cytoplasm. On the
zoological side the publication by Ehrenberg, in 1836, of
Die Infusionsthierchen als volkommene Organismen may, I
think, be looked upon as the first milestone in the progress
of knowledge of cytoplasmic transformation into tissue sub-
stance.

The ever-broadening stream of knowledge of the chromo-
somes in relation to heredity is usually held to have orig-
inated in the discovery forty years ago, by O. Hertwig,

66

•Inheritance Materials of Germ-Cetts 67

that the most essential fact in fertilization is the union of
the nuclei of the male and female germ-cells.

That cytoplasm is a physical basis of heredity is proved
by a great body of direct observational knowledge. That
the chromatin of chromosomes is a physical basis of heredity
is proved by much observational knowledge when this knowl-
edge is supplemented by reasoning involving the principles
of biological comparison and correlation. These two masses
of knowledge constitute, as already indicated, the founda-
tion of all legitimate reasoning about inheritance material.
Whether chromatin and cytoplasm are the only substances
which participate in the formation of hereditary structures
can not now be stated with certainty, though there are both
observational and general grounds for believing that they
are not. But into this question we need not enter in this
discussion. Nor is it necessary for our purpose to inquire
very particularly whether the conceptions of chromatiin
and cytoplasm really imply just two substances or two great
classes of substances, though it is best to have in mind the
undoubted fact that the latter alternative is the true one.
Beyond a doubt "chromatin" and "cytoplasm" ought al-
ways to be understood to mean "kinds of chromatin" and
"kinds of cytoplasm."

Function, of Chromosomes in Heredity Acquired and
Secondary

The question which specially concerns us here is that of
what the relation is between chromatin and cytoplasm in
virtue of which they play the particular roles they are found
to play in producing hereditary structures. Perhaps the
most important aspect of this general question is that which
the theory of phyletic evolution naturally brings up: does
the evidence in hand suggest any answer to the question
whether chromatin or cytoplasm is the more primitive and

68 The Unitt) of the Organism

fundamental as hereditary substance? Surveying as we
have the whole field of organic propagation, including the
process in unicellular organisms as well as in multicellular,
and sexless as well as sexual methods of reproduction, and
taking the facts as they actually present themselves, it
seems as though but one answer to this inquiry is possible:
The substances included under tht generic term cytoplasm
are the more fundamental and primitizre.

The only possible way of escaping this conclusion is by
excluding from the conception of heredity the vast majority
of developmental phenomena presented by unicellular organ-
isms and by monogenic propagation in multicellular organ-
isms. As our examination of these provinces revealed, such
exclusion is exactly what the chromatin dogma of heredity
has undertaken, implicitly or explicitly, to fix upon biology.
The utter unwarrantableness of this undertaking was made
sufficiently obvious, we may assume, by the examination;
so we need spend no time on that now. All that is necessary
is to remind ourselves vividly of the main positive outcome
of the examination: Heredity is a universal phenomenon of
the living world. It is coextensive with organic propagation
and development, while "carrying heredity" by chromo-
somes is, according to the evidence, very far from a uni-
versal phenomenon. It is a long way from being coex-
tensive with organic propagation and development. We
may, consequently, proceed in our quest of a more rational,
more consistent, more satisfactory conception of the purely
operative side of producing hereditary structures. Pur-
suing the quest, we remind ourselves of having found that
only the most meager observational evidence is afforded by
the protophyta and protozoa, and by monogenic metaphyta
and metazoa, that chromatin is hereditary substance, while
these organisms afford an overwhelming mass of such evi-
dence that cytoplasm is hereditary substance.

But if in the lower, more primitive moiety of the organic

Inheritance Materials of Germ-Cells 69

realm, chromatin is a physical basis of heredity to only a
limited extent and in a partial way but has this office widely
and positively fixed in the higher moiety, the moiety, that is,
in which bisexual propagation is fully established, what
other conclusion can be drawn consistently with the modes
of reasoning universally sanctioned by evolutionists, than
that the function of "carrying hereditary qualities" by the
chromosomes in higher organisms is a secondary or ac-
quired, or better a delegated or assigned function? Hered-
ity is far older, phylogenetically, and far broader tax-
onomically than is the chromatin mechanism by which it
now in part manifests itself. Under this interpretation the
acquisition by chromosomes of the function of carrying
heredity would belong to the same evolutional type as for
example the acquisition by certain cells of the function of
muscular contraction, or by certain other cells of con-
ducting nervous stimuli. The advantage and satisfaction
of a conception of the role of chromosomes in heredity
which ranges them naturally and easily with all other or-
gans and tissues of the plant and animal body will be
quickly seen by every one to whom the seemingly endless
chance of discovering new interrelationships and consis-
tencies in living nature is one of the most rewarding things
about biological investigation.

While we are duly impressed with the importance of per-
ceiving that chromosomes fall into the great class of other
organs and tissues when considered from the standpoint
of phyletic differentiation of structure and function, we
should not fail to notice that within the class they hold on
a number of counts a very distinct place. Probably the
most distinctive of these counts, at any rate the one most
important for this discussion, is the fact that while the vast
majority of tissues, taking the term in its usual meaning,
stand at or near the termination of the ontogenic series —
are, in other words, the final stage in the series — the chro-

70 The Unity of the Organism

mosomes in their function as bearers of heredity stand at
the beginning. They represent the initial stage in the series.
Furthermore, their function in this respect is unique as
contrasted with the function of other tissues, in that while
other tissues have, typically, each a single function which
they perform immediately, the chromatin of a given germ-
cell has a great complex of functions, namely, that of ini-
tiating the development through which all the attributes of
the individual during its whole life-career are developed.
In a word, chromosomes of germ-cells are not determiners or
carriers of determiners ; they are initiators or carriers of
initiators. They may, then, be called bearers, or carriers
of heredity in a very literal sense, namely in the sense that
they are made use of by one individual, the parent, to carry
across or transfer from itself to another individual, the
child, the hereditary attributes of the species in a latent or
potential state. By virtue of their being thus used, they
are members of a developmental series, in which series their
place is at the beginning and not at the end, the nature of
the series depending on the phylogenic history of the par-
ticular organism to which the particular chromosomes be-
long.

The Two-fold Character of the Problem of Hereditary

Substance

At this point it becomes a matter of the greatest im-
portance for theories about hereditary substance to dis-
tinguish between the problem of the operation of such sub-
stance in the developing individual, and that of how such
substance ever came to be hereditary substance; stated
otherwise, between the problems of how any substance par-
ticipates either directly or as an agent in the building up of
a structure having hereditary attributes, and that of how
the substance itself became impressed with the attributes, in

Inheritance Materials of Germ-Cells 71

a latent state, of the progenitors of the developing in-
dividual.

Investigation of the first of these problems is to a large
extent a matter of observation, as we have seen in the pre-
ceding pages. The sub-science of histogenesis consists
largely in tracing out the processes by which completed
tissues arise by the transformation of less differentiated or
undifferentiated cells. And when such newly arisen tissues
and structures are proved to be hereditary by such evi-
dence as we have called attention to, then the study of
histogenesis comes to be so far a study of hereditary sub-
stance.

When, however, we turn to the other problem, that of
how hereditary substance comes to be such, we are in a
different, a much more difficult case, for so far science has
succeeded in getting almost no observational hold upon it;
and despite the vast discussion it has received the darkness
that envelops it is hardly an iota less black than it was the
day of its original formulation. But stygian as the darkness
is, here, especially as to details, we yet are able to see,
probably, the quarter from which light will come if ever it
does come. That quarter is the physical-chemistry con-
ception of the organism as a system of phases the whole of
which, as a species entity, is essential to its equilibrated
activities. This nature of the organism, together with some-
thing akin to its internal secretory and enzymic productiv-
ity, enables it, we may conjecture, by some means now
wholly unknown, to reflect its totality of transferable at-
tributes upon the germinal cells and to transform them into a
latent state. Ungrudging acknowledgment of the complete-
ness of our ignorance of how any part of a cell or any other
portion of an organism becomes endowed with the capacity to
develop or causally to affect the development of an organism
similar to that from which it came, should be an important
item in the preparation to accept any and all indubitable

72 The rtiiti/ of the Organism

facts connected with heredity, even though no causal ex-
planation of them is forthcoming.

The childlikeness, as Conklin well characterized it, of
the belief that chromosomes are a simple and complete ex-
planation of inheritance would not be so bad in itself. If
it stopped there, as genuine childlikeness would, no posi-
tive harm could be done. It is the making of this belief
the starting point of a grand speculation which blinds the
eyes and closes the mind to a vast number of facts and
legitimate inferences about heredity, that plays havoc with
thinking on genetics. It would be a great gain if gen-
etic theory would recognize wholeheartedly that all or-
ganic development, as contrasted with mere enlargement,
consists more fundamentally and obviously in transforma-
tion of substances than it does in unchanged continuity of
substances. For under such recognition the futility of
attempting to explain the transformation of one lot of
substance by referring it to another lot which does not
transform, or in other words to explain development by
something that does not itself develop, would be manifest.

That the chromosome theory of heredity in reality deep-
ens rather than illumines the darkness which surrounds the
problem is seen when one reflects that not only does it
throw no light on the question of how the chromosomes come
to be bearers of heredity, but that it creates the new and
equally difficult question of how the chromosomes (which ac-
cording to the theory maintain unchanged their individuality
not only from generation to generation but throughout each
ontogeny) are yet able to be causally operative in the cell-
bodies undergoing the transformations which they actually
do undergo in the developing organism. That the germ-
plasm-chromosome theory of heredity could have led its
devotees to sidestep the details of ontogeny, especially those
of histogenesis, to the extent which our review has shown
it to have done, would be unbelievable but for what is

Inheritance Materials of Germ-Cells 73

actually before us in the recent history of biology.

Viewing heredity as being definitively a kind of organic
transformation — transformation, that is, in accordance with
a pre-existing or ancestral pattern more than it is a kind
of continuity— it becomes obvious that even were the dem-
onstration to become complete that the chromosomes are
the only portions of the germ-cells * essential to fertiliza-
tion, they still would not be proved bearers of heredity in
such sense as the germ-plasm theory holds them to be. They
would not because the problem of the transformations
which constitute ontogeny would still be untouched. The
theory would be established only when the demonstration
should be produced that the chromosomes cause immedi-
ately all the particular ontogenic transformations known
to be hereditary. All that would be proved about heredity
by demonstration that the chromosomes alone participate
in fertilization would be that the chromosomes alone con-
stitute the first ontogenic stage of the hereditary parts of
the particular organism to which the fertilized egg gives
rise.

The Probability That Inheritance Material Becomes Such
In Each Ontogeny

But because thus far failure has attended all efforts to
get knowledge of how hereditary substance is produced,
are we obliged to own that we know nothing at all, even
inferentially, about its production? And is the search for
such knowledge to be given up as hopeless? My answer is
an energetic negative to both these questions. In the first
place, there is much evidence to support the hypothesis,
very general to be sure but yet by no means devoid of use-
fulness, that hereditary substance becomes such in some

* The utter unwarrantableness of the common assumption that as
regards the male germ-cell such a demonstration is "practically com-
plete" will be noticed presently.

74 The Unity of the Organism

way through being subject to the metabolic processes com-
mon to the whole organism. Undoubtedly the germ-plasm
dogma itself has tended strongly to divert attention from
this aspect of the problem of germinal material— indeed,
has tended to minimize the importance of the metabolism
of such material even if it has not tended to deny that the
material is subject to this process.

So important is it from the organismal standpoint to
conceive the material basis of heredity as part and parcel
of the organism generally, especially as regards the basal
growth and sustentative processes, that we must examine in
some fullness the evidence favorable to such a conception.
In its most brazenly evidence-ignoring form, the germ-plasm
dogma asserts that the female parent does not really pro-
duce the eggs or the male parent the sperm, as they seem
to, but that these are produced by previous germs ad MI-
finitum. There are, to be sure, quite a number of observ-
able facts, as those of the early formation of germ-cells
in several animals, that can be forced into a seeming sup-
port of such a conception. But the familiar and all but uni-
versal fact that multicellular organisms, plants and animals
alike, are sexually immature for a shorter or longer part of
their lives, the very essence of the immaturity being the un-
developed state of the reproductive system, would be a
sufficient refutation of the view for any mind not made
impervious to facts by long and faithful sophistication.

Germ-Cells Subject to Metabolism Like All Other Cells

The biological commonplace that all germ-cells, like all
other cells, undergo a process of growing and maturing
before they can perform their distinctive office, and that
this process depends upon the retention of the germs by the
parent organism, ought, as already indicated, to be a suf-
ficient antidote againjst the germinal continuity fallacy,

Inheritance Materials of Germ-Cetts 75

even though nothing were known as to exactly what goes
on in the germ while it is growing and ripening. But we
are by no means without positive knowledge under this
head. In fact the last few cell divisions immediately pre-
ceding the ripening of both ova and spermatozoa, and the
ripening processes themselves, have received searching ex-
amination during the last few decades, with the result that
hardly any cytological phenomena are better known than
are the profound morphological changes which accompany
these processes. That these changes are particularly mani-
fest in the chromosomes, the assumed seat of the determiners
of heredity, is one of the very things that has aroused so
much interest in the processes. Nor are we wholly unin-
formed about the chemical changes taking place in the
growing germ-cells. Unfortunately knowledge in this field
has hardly passed the stage of early infancy, but at least
enough is known to warrant the assertion that the young
germ-cells are subject, as are all the other cells, to the
general metabolism of the organism.

Chemical Changes in Germ-Cetts During Parent's Ontogeny

About the most striking information we have in this
field is what has come from such investigations as those
on the chemical changes which occur in the sex glands and
other body parts during reproduction in some fishes.* Mie-
scher's work was ground-breaking in this domain for it
was the first to show that the "sexual organs in the salmon
develop at the expense of the muscular system, and that the
salmine deposited in the testis during the breeding season
must be derived from the proteins of the muscle, since the

* Notable among these studies are: Histochemische und physiologische
Arbeiten, gesammelt und JieraMsgegeben von seinen Freunden, by Mie-
cher; and Changes in the Chemical Composition of the Herring during
the Reproduction Period, by Milroy. Biochemical Journ., v. in, 1908, p.
366.

76 The Unity of the Organism

fish does not take any food during the period." l The work
of Riddle and his collaborators is producing evidence to the
same effect.

Such researches do not, to be sure, prove that the chem-
ico-physiological changes extend to the chromosomes of the
germ-cells, much less to the imaginary determiners of hered-
ity in the chromosomes. But viewing the results in the
light of the well-grounded general belief that the mo»st
fundamental test of living substance is metabolic change,
it is seen that any hypothesis which assumes the existence
in the germ-cells of something virtually not subject to the
general metabolism of the cells, assumes at the same time the
burden of furnishing objective evidence that such a something
does exist.

As a matter of fact the prime offense of the germ-plasm-
determiner hypothesis is that its very essence places it
beyond the reach of scientific observation. Such truth as it
may contain cannot be made really effective because it can
not be proved, and such error as it may contain can not
be robbed of its power for evil because it cannot be dis-
proved. In a word, the hypothesis is one that belongs to
the realm of dialectics primarily, and has no just claim to
a place in inductive science.

The Possibility of Changmg Sex By Influences on the Germ

But perhaps the most conclusive evidence of the funda-
mental dependence of true germinal material upon the or-
ganism, should somewhat fuller verification of the obser-
vations be obtained, are results like those reached by Whit-
man, King, Whitney, Riddle, and by R. Hertwig and his
students, according to which sex may be reversed in several
animal species by various conditions extraneous to the
germ itself, acting on the germ-cells from which the animal
is to develop. The instance of this usually regarded as best
established is afforded by certain species of frogs and toads.

Inheritance Materials of Germ-Cetts 77

The widely known result reached by Hertwig and verified
and extended by Kuschakewitsch expressed in a single sen-
tence, is that the number of males and of females produced
by the eggs of a given female depends upon whether the
eggs are fertilized when newly ripe or when over-ripe, a great
predominance of males coming from the latter class. In
a tabulation of the results of four sets of experiments by
Hertwig 2 almost every case shows the number of males
increased when the time elapsing between deposition of the
eggs and fertilization was increased, the highest percentage
of males in any one lot being 759, fertilization in this in-
stance having been twenty-two hours after the fertilization
of the last preceding lot.

In a species of toad, Bufo lentiginosus, results have been
obtained just the reverse of those on the frog, that is, the
proportional number of females has been experimentally
increased. This was accomplished by fertilizing the eggs
in water made slightly alkaline. Since frog eggs are known
to absorb water when they remain long in it, as in the case
of those which gave rise to a preponderance of males in
the Hertwig method of experimenting; and since alkaline
solutions extract water from eggs, and likewise cause them
to produce a preponderance of females, Miss King drew
the obvious conclusion that the quantity of water contained
in eggs of these animals may be a factor in determining the
sex of the animals developed from the eggs. Although
Miss King recognizes that her experiments do not furnish
final proof of the conclusion she draws, she believes, rightly
it would seem, that they weigh heavily in that direction.
"As they stand," she writes, "the results strongly suggest
that sex in Bufo is determined at or near the time of fer-
tilization, and that external factors acting during this pe-
riod may influence the sex-determining mechanism in such
a way as to cause it to produce one sex or the other. The
results also seem to indicate that in Bufo sex is determined in

78 The Unity of the Organism

the egg, and that it may depend in some way on the rela-
tive amount of water in the egg at the time of fertiliza-
tion." 3

Riddle, perhaps the most outspoken opponent of sex pre-
destination now writing, strongly espouses the hypothesis
that the sex to which a particular egg will give rise is de-
pendent partly on the quantity of water which that egg
contains. But whether water is a factor in determining sex
or not, the evidence presented by Riddle, coming partly from
researches by C. O. Whitman and partly from his own,
constitutes, when taken with the evidence to the same effect
presented by other investigators, almost if not quite com-
plete proof that sex is not the hard-and-fast thing which
most present-day genetic speculation would make it.

Furthermore the evidence produced by these two inves-
tigators seems to connect the decision as to which sex a
particular egg shall give rise, with some condition of the
parents. It is well known to all zoologists, in the United
States at least, that at the time of his death Professor Whit-
man had accumulated a vast store of data on the habits,
particularly the breeding habits, of pigeons. To Doctor
Riddle, who had worked with Whitman considerably, fell
the task of carrying on to some extent Whitman's experi-
ments and of preparing for publication the results which
Whitman left in the rough. The following quotation from
Riddle's paper referred to above, summarizes Whitman's
results that are especially important for us now: "Whit-
man found that if certain very distantly related pigeons
[i.e., two individuals from different families] are mated
that only male offspring resulted. If the matings were
made of individuals not quite so distantly related — different
genera usually — and if to this situation be added the ele-
ment of overwork at reproduction [i.e., the birds not being
permitted to- nest their own eggs, but forced to keep laying
eggs in rapid succession] then the first several pairs of

Inheritance Materials of Germ-Cells 79

eggs produced in the spring will produce all or nearly all
males. The last several pairs of eggs laid in autumn will
produce all, or nearly all, females. At the transition period
in the summer he found that some pairs, or clutches, of eggs
produced both a male and a female. In these cases it was
usually the first egg that produced the male ; and the sec-
ond egg — laid forty hours after the first — that gave rise
to a female." 4

Into Riddle's interesting discussion of Whitman's results
and his own chemical studies on the eggs of pigeons and
hens we need not go. Suffice it to say that it seems to me
Riddle is justified by the evidence now in our possession,
in his contention that "sex rests upon a quantitative and
reversible basis" and that in this sense it has been controlled
by conditions extraneous to the germ-cells themselves. This
does not imply, as I understand, that such control would
necessarily be practicable or even possible in all organisms,
nor does it preclude the possibility that in some species
there may be dimorphic or partially dimorphic spermatozoa
or ova as regards sex production. Neither does it preclude
the possibility that in some cases where a preponderance
of one sex has been observed, this is due to selective mor-
tality or some process other than the actual shifting of the
sex tendency in the particular eggs.

These several concordant bodies of testimony must, it
would appear, open the eyes of biologists sooner or later to
the ludicrousness of a theory that would make the parent
organism hardly more than a combined culture medium and
incubating oven for its germ-cells.

The Determiner Conception Contrary to Ordinary Chem-
ical Principles

If, on the basis of such facts as we have, we try to come
still closer to the questions of how the assimilative and
morphogenic processes of the organism occur, whether in

80 The Unity of the Organism

the production of hereditary substance or in the transform-
ation of such substance into actual hereditary structures
and activities, we find ourselves hedged about on every side
by partial knowledge, by dubious knowledge, and by com-
plete ignorance on many fundamental points.

However, chemical considerations seem to point the way
to future discovery. In the first place, it seems necessary
to recognize that the whole germ-plasm conception as orig-
inally promulgated, with its interminable system of "bear-
ers" and consummators, was contrary to what is well known
about chemical processes generally. Thus the continuity
presented by a complex chemical operation does not consist
in an unchanged series of individual entities of some sort,
such as determinants and determiners are, or originally
were conceived to be, but rather in a regular succession of
transformations. For example, when chromic hydroxide,
which is grayish-green, is dissolved in acid, a green solu-
tion results, which turns to greenish violet or pure violet
if allowed to stand a long time. Exactly what the chemical
changes are that correspond to these color changes I do
not know, and probably the information which chemists have
on the subject is not exhaustive. At any rate modern
chemistry conceives the phenomenon to consist in a succes-
sion of reactions and transformations, the various colors
and shades being attributes of the compounds that exist
in the various stages along the way, and not as due to in-
dividual bodies carried by the preceding stages for the
express and exclusive purpose of producing the particular
colors that are observed, as would be implied in such a
metaphysical scheme as was the germ-plasm theory elab-
orated by Weismann.

Enzymic chemical action presents perhaps a still better
starting point for imagining what the fundamental hered-
itary processes may be than does ordinary chemical activity
like that just instanced. The essence of this kind 0f activ-

Inheritance Materials of Germ-Cells 81

ity is, as everybody knows, that in some way enzymes cause
or at least facilitate transformation in other substances.
Thus the attribute of solubility of the sugar into which
starch is transformed, through the action of the salivary
enzyme ptyalin, is not held to be due to a determiner for
solubility carried by the ptyalin and passed on into the
sugar, but rather it is recognized that solubility is one of
the attributes possessed by the kind of sugar into which
starch is converted by the ptyalin. The solubility is thought
of rather as an attribute of the sugar and not as something
once latent in the ptyalin which produced the sugar. A
few details of the action of the enzyme in this case illustrate
the point still better. Maltose, which is the chief if not
the only sugar resulting from the action of ptyalin, is not
reached by a single bound, as one might say, but through a
series of bodies known as dextrins, at least three of which
have been recognized. These are amylo- erythro- and
achroo-dextrin, named from the color they display when
treated with iodine, the first mentioned turning blue, the
second red, and the third remaining colorless. What mod-
ern chemist would think of explaining the blue of the amylo-
dextrin by a "determiner" for that color in the ptyalin
or even in the starch, the red of the erythrodextrin by an-
other determiner for red, and so on? That is the sort of
explaining chemists of a century ago did, but they have
long since learned not merely the futility but the scientific
evil of such explanation.

If it were germane to our present task we might go on
and show that the gene conception in modern genetics is
really a revival in biology to-day of the gene conception
which passed muster in chemistry a hundred years ago,
when oxygen and hydrogen were named. Such an exposi-
tion would be appropriate to a history of scientific theory
or to a treatise on the theory of natural knowledge, but
hardly to the present work.

82 The Unity of the Organism

Endorsement of E. B. Wilson's Proposal to Drop "De-
terminer" From the Vocabulary of Genetics

In his Croonian Lecture having the title The Bearing of
Cytological Research on Heredity, E. B. Wilson said, "In
the meantime it would be well to drop the term 'determiner'
or 'determining factor' from the vocabulary of both cytol-
ogy and genetics." 5 If the facts and arguments set forth
in the preceding pages are valid, they constitute a demon-
stration that not only would it "be well to drop the term
'determiner,' " but that it must be dropped, at least in its
present application, before thought and investigation on the
mechanism of heredity can be free and in very deed truth-
seeking. "What we really mean to say," Wilson continues,
"is 'differential' or 'differential factor,' for it has become
entirely clear that every so-called unit character is pro-
duced by the cooperation of a multitude of determining
causes." So far as these statements go they are in strict
accord with the organismal standpoint maintained in this
volume, and we may also say, with the physical-chemistry
standpoint.

Where attributes of adult organisms have been so defin-
itely correlated with particular chromosomes and possibly
parts of chromosomes of the germ-cells as seems to be the
case in the fruit flies, such chromosomes are unquestionably
differential, and since they stand at the very beginning of a
long and complex transforming and developing series, they
may very properly be called differential factors even though
they do not themselves participate substantively in the
transformation. The general similarity of their mode of
action to that of enzymes is certainly considerable : a minute
quantity of the substance is capable of inducing or facili-
tating the transformation of a large amount of other sub-
stance in a perfectly definite manner, and the inducing agent
is not itself consumed.

Inheritance Materials of Germ-Cells 83

Advantages of Conceiving Germ-Cell Chromosomes as Initia-
tors m Hereditary Development

This chemical way of viewing chromatin and chromosomes
sanctions the idea that these are to be regarded as initiators
of developmental processes which lead to hereditary attri-
butes, rather than determiners of those attributes. If one
wants to know in what way this conception would have
an advantage over the determiner conception as a working
hypothesis, my reply is that the advantage is two-fold.
First, it would surely correct the tendency of genetic re-
search under the guidance of the determiner hypothesis, to
restrict its attention to attributes of adults at one end of
the ontogenic series and to the chromosomes of the germ-
cells at the other end, and to ignore or touch only in the
lightest way all the intervening parts of the series. This
correction would result because the new standpoint would
bring the whole series of continuities and transformations
alike into proper perspective, revealing thus that the mem-
bers of the series intervening between germ and adult must
be investigated in exactly the same way and with the same
objects in view as the end members, if complete understand-
ing of the hereditary process be the goal of research. It
could not then happen that the egg-cell would be repre-
sented, as it now so commonly is, as a relatively large sac
containing nothing significant for heredity except the rela-
tively small chromosomes. Nor could nearly the whole
mass of ontogenic phenomena, especially those of histo-
genesis, be treated so lightly in speculation and so largely
neglected in investigation as they have been under the
domination of the determiner theory.

The second advantage in the initiator conception is that
since it would recognize the "differential factor" of the
chromosomes to be in reality due to the fact that the whole
ontogenic series to which the chromosomes belong is differ-

84 The Unity of the Organism

ential, that is, that it pertains to a particular species or
kind of organism, it would put an end to the notion by
which recent genetic science has been so largely dominated,
that the problem of how the series came to be thus specific
or differential may be solved by speculation, and it would
incite geneticists to efforts to solve the problem by obser-
vation aided by experiment. It is impossible to refute the
charge that genetics is to-day more interested in an elab-
orate system of conceptions — of speculation, in other words
— than it is in observed or possibly observable phenomena.
We cannot keep too constantly before our minds the fact
of our almost complete ignorance of how any substance
becomes hereditary substance whether through the "inheri-
tance of acquired characters" or in any other way. Hence
mere speculation on the subject after the manner of the pan-
gens idea is much worse than nothing if permitted to run into
a bewildering and enslaving system like that of the germ-
plasm theory as it came from Weismann's mind. Neverthe-
less it is quite germane to the present discussion to point
out that whatever might be the nature of the chemical ac-
tion, whether enzymic or some other, through which the
series of ontogenic transformations should be accomplished,
the character and subtlety of these processes seem to make
them, more than any others we know, competent with some
modification to serve as the go-between for impressing the
germinal material with the latent attributes of the species.

Inconclusiveness of the Cytological Evidence Usually Ap-
pealed to in Support of the Chromosome TJieory

And this leads to the concluding statements of this dis-
cussion. The three categories of cytological fact which
have been weightiest in the formation and maintenance of
the chromosome theory of heredity are the individuality and
continuity, chiefly numerical, of the chromosomes from par-

Inheritance Materials of Germ-Cells 85

ent to offspring; the apparent equality (it should never be
forgotten that the dogma of equality does not rest on rigor-
ous quantitative investigation) of the chromosomes in the
male and female germ-cells ; and the assumption that in
the male germ-cells the chromosomes alone are concerned
in fertilization. Even if these groups of assumed fact were
established with absolute certainty they would fah1 far short
of being direct and final proof that chromatin is the only
hereditary substance. That this is true must be apparent
to all well-informed, carefully thinking biologists. The
most important grounds for this inconclusiveness are in-
volved in the facts and arguments set forth in the preced-
ing pages, but they may be summarized here, and in addi-
tion two other grounds may be pointed out.

First and foremost, in my opinion, is the general truth
that chromosomes or even chromatic substance can not pos-
sibly be recognized as the sole bearers of hereditary sub-
stance, because the evidence is enormous in quantity and
direct and indisputable in quality that other substances par-
ticipate actively in the production of hereditary attributes.
There is no way of escaping this conclusion except by nar-
rowing the definition of heredity for the very purpose of
bringing it within the scope of the chromosome theory of
hereditary substance. The scope and fundamentality of
this aspect of the problem is sufficiently dwelt upon, we
may hope, in what has gone before.

The two additional grounds for skepticism as to the con-
clusiveness of the cytological evidence will now be pointed
out. First, as to the evidence from the individuality and
continuity of the chromosomes. All that any careful
thinker claims or can claim for this evidence is that the
individuality and continuity of chromosomes as observed
are what might be expected if they were the germinal de-
pository of the organism's hereditary attributes. And the
question is constantly and naturally afeked, what other

86 The Unity of the Organism

meaning can the whole remarkable series of phenomena of
maturation and fertilization have than that attributed to
them by the chromosome theory? My rejoinder to this
argument may begin with a reply to the question. As long
as knowledge of the chemistry and physiology of cells —
germ-cells with the rest — is as fragmentary and inconclusive
as it now is, certainty as to the meaning of the phenomena
mentioned is out of the question. However, it would seem
quite probable that they are concerned primarily with the
nutritive and assimilative processes of the cell and only
derivatively with heredity. Furthermore, the question asked
may well be paired off with another of similar character,
namely, what is the meaning of the almost if not quite com-
plete breaking up and disappearance of the chromosomes in
the so-called resting stages of the immature germ-cells, this
being accompanied by the dissolution of the nuclear mem-
brane so as to allow the freest possible commingling of the
whole nuclear contents with the cytoplasm of the cell? Have
we not at least as much factual right to suppose that during
this mixing of substances the chromatin, and so later the
chromosomes, are influenced by the cytoplasm, as that the
reverse influence takes place? Is it not entirely possible
that this process is one of the very means or occasions of
impressing the chromosomes with the attributes of the or-
ganism which, as we have seen, apparently must take place
whether acquired characters are ever inherited or not?

And now as to the argument from the assumed exclusive
participation of the chromosomes of the male germ-cell in
fertilization. First of all, it can not be admitted for a
moment that the chromosomes are proved to be as ex-
clusively the fertilizing agents as they are generally as-
sumed to be. Even in such extreme cases of seeming ex-
clusiveness of chromosomal participation as that claimed
by Strasburger for the pollen cells of some plants, neither
Strasburger nor any one else has claimed, so far as I know,

Inheritance Materials of Germ-Cells 87

that all other male substances than chromatin are excluded
as rigidly as would be required by experiments in a chemical
laboratory designed to ascertain the action of a particular
chemical element or substance in its purity. It is certain,
for example, that in almost if not quite all male germinal
elements in animals, a thin outer layer from the cytoplasmic
part of the spermatid is present on the head of the sperma-
tozoon. Furthermore, it is well known that at least the
"intermediate piece" of the sperm tail, which is not usually
regarded as chromosomal in origin, remains in the egg at
fertilization. Nor is there any good ground for supposing
that the non-chromatinic portions of the nucleus are ab-
solutely excluded. The almost certain presence in the egg
at fertilization of at least these male substances other than
chromatin can by no means be regarded as insignificant for
heredity, especially if the initiator conception of germinal
material is held. It seems to follow of necessity that if the
fertilizing substances, whatever their source, be conceived
to act in an organic system of the physical chemistry sort
after the manner of enzymes, no such quantitative relation
subsists between these fertilizing substances and the prod-
ucts of organic growth as the chromosome theory implies ;
nor can their action be so narrowly localized in the egg.
Their action would be conceived to involve the entire ovum
ab initio, and not the chromosomes alone.

Summing Up of the Findings Against the Chromosome

Theory

The general result of our critique is that the whole at-
tempt to interpret the physical basis of heredity in accord-
ance with elementalist conceptions has failed and must con-
tinue to fail, so fat as its main aim is concerned. We are
led to see that the germ-plasm dogma, no matter how often
or how completely it changes its nomenclatural habiliments,

88 The Unity of the Organism

as in the shifting from determinants to determiners, or from
determiners to gens, or from gens to factors, involves a
rejection of the conception that the germinal elements of
organisms after being discharged are literally detached
parts of those organisms. This conception was well on the
road to incorporation into the great body of established
biological truth when it was headed off by Weismann's
diametrically opposed hypothesis of germinal isolation.

I would insist that the defense of the organismal concep-
tion in this volume is really a carrying out of such a con-
ception of the organism and its germinal products as is
implied by the old view that the germ is a part of the parent
organism. It would hardly be possible to express more sat-
isfactorily in a single sentence the most inclusive theo-
rem, as it might be called, the demonstration of which is the
aim of the part of this volume devoted to the means by which
organisms propagate their kind, than the following from
E. B. Wilson: "To the modern student the germ is, in
Huxley's words, simply a detached living portion of the
substance of a preexisting living body carrying with it a
definite structural organization characteristic of the
species." 6

Coupling this statement by Wilson with another from
one of his latest writings,5 to the effect that we ought to
drop the term determiner because in reality what it means
is differential, I call attention to the fact that the "dif-
ferential factor" of the later statement and the "definite
structural organization characteristic of the species" of
the earlier statement are in essence one and the same. The
only difference is that in the earlier statement it is the
whole germ-cell that is recognized to be a detached part of
the organism, while the later statement can be brought
down to the chromosomes because of the greater refinement
of knowledge attained since the earlier one was made. The
point I wish to make stand forth with the greatest possible

Inheritance Materials of Germ-Cells 89

boldness is that the germ-cell chromosomes may properly
enough be said to be differential, if only one never loses
sight of the fact that they are differential in no other sense
than are any other particles or substances of the germ-
cells or any other cells which participate in the production
of species-attributes.

Brief Reference to the Untoward Implications of the Germ-
plasm Conception of Heredity

The somewhat laborious task of exhibiting the difference
between conceiving the physical basis of heredity from the
elementalist and from the organismalist standpoints may
well be brought to a close by calling attention to the impli-
cation of the two conceptions as applied to heredity in man
himself. Looked at from this direction the germ-plasm
dogma is seen to be chargable with the grave offence of
having added its weight to a conception of human life, the
overcoming of which has been consciously or unconsciously
man's aim throughout the whole vast drama of his hard,
slow progress from lower to higher levels of civilization —
the conception that his life is the result of forces against
which his aspirations and efforts are impotent. As ap-
plied to man this form of fatalism is no less sure and no less
dire in its tendencies than have been any of the innumerable
theistic forms of fatalism that have prevailed through the
centuries. It is almost certain that the ardor with which
Eugenics has been espoused by several biologists is due
in considerable measure to the fact that they have felt
more or less definitely this sinister implication of the theory,
and have turned to Eugenics as the only weapon against
its evil forebodings. The germ-plasmic eugenist virtually
says, "Yes, indeed is man a reasoning, willing, aspiring
animal, but all his activities in these ways are futile so far
as the race as a whole is concerned, except as they are

90 The Unity of the Organism

brought to bear, extrinsically and operatively rather than
organically, on the Germ-Plasm." This form of the Eu-
genic idea corresponds in spirit to the propitiative offerings
of primitive religion. It aims to mollify by human agency
powers that act upon men's lives, but which are in them-
selves largely extraneous, largely evil, and wholly irre-
sponsible.

What eugenists of this school have failed to see, evidently,
is that even were unit-factors as differentiate from one
another in heredity as the extremest Mendelist conceives
them to be, and that even were the germ-plasm improved up
to the level of his highest hopes, his results in terms of ac-
tual human lives and social conditions would be distressingly
meager. They would be so, because whether unit-factors
exist independently in heredity or not, they certainly do
not exist thus independently in development and function.
In these ways they interact upon one another in the most
vital manner, as physiology, especially of the internal se-
cretions and the nervous system, and as physiological and
social psychology are rapidly and conclusively demon-
strating.

We thus end our examination of the means by which or-
ganisms produce others of their kind with the conclusion
that the material through which reproduction is accom-
plished is in the most vital way part and parcel of the
organism, that is, that the germ-cells are somehow stamped
through and through, potentially, with the characteristics
of the kind, or race, or species to which the producing or-
ganism belongs. And with this we are ready to pass to the
examination of those integrative phenomena of the organ-
ism generally, one manifestation of which is this very nature
of the germ-cells.

REFERENCE INDEX

1. Marshall 292 4. Riddle 10

2. Hertwig, R. ('12) 75 5. Wilson, E. B. ('14) 351

3. King 232 6. Wilson^ E. B. ('00) 7

PART II

THE CONSTRUCTIVE SIDE OF THE ORGANISMAL
CONCEPTION

Chapter XV 11
GROWTH INTEGRATION

The Field to be Covered by the Constructive Discussion

ACCEPTING the inevitable destructive result of our cri-
tique of the elementalist standpoint, that the attempt
to interpret living beings in the terms of their constituent
parts alone always leads to partial failure and disappoint-
ment, or to the worse result of illusionment as to the trust-
worthiness of the explanations proposed; and accepting the
constructive result that everything in the critical study tends
to show that no part of any organism can be rightly inter-
preted except as part of an individual organism, this indi-
vidual being in turn interpreted as a member of a taxonomic
group, it is revealed that we are only on the threshold of the
positive, the constructive side of our general enterprise. Even
though the conclusion be unescapable that the living organ-
ism somehow acts causally on its parts, the problem still
remains as to the modus operandi of that acting. The "some-
how" which came to us as an incident of our critical study
has yet to be inquired into.

Stated more specifically the task now before us is that of
examining closely and systematically the interdependences
among the parts of the individual organism. Although these
interdependences are among the most obvious and general
of all organic phenomena such an examination of them biol-
ogy has not yet made systematically. Indeed — and here is
one of the most vital things for us to see — a cardinal charge
against the elementalist standpoint is that in its very nature

93

94 The Unity of the Organism

it not only does not encourage, it actually stands against
such examination. Its opposition to comprehensiveness and
systematization is profound and essential.

Our examination will begin with a single brief, two-parted
definition : The structural and functional interdependence
found to exist among the parts of an organism we call bio-
integratedness; and the process of moving on from grade
to grade of interdependence among the differentiating parts
which constitutes ontogenesis in the individual we call bio-
integration.

Four Types of Bio-integration to Be Treated

In the present state of knowledge and for the discussion
now before us four types or kinds of bio-integratedness and
bio-integration may be recognized as pertaining to the in-
dividual organism :

1. Growth integration, most obvious in graded me'ristic
series, but also expressed in the "axial gradients" of Child.

2. Chemico- functional integration, known so far chiefly
in connection with internal secretions.

3. Neural integration, comprising the interdependences
among the parts of the nervous system, and the involvement
with this of the muscular, glandular and other organs.

4. Psychic integration, very closely connected with neural
integration, but approached from the side of the totality of
activities of living beings rather than from the side of nerve-
organ activity, and so taking cognizance of a vast number
of phenomena not yet definitely correlatable with neural
phenomena.

The full presentation of facts and arguments under these
four heads would reach far beyond the limits set for the
present work. We are, consequently, obliged to restrict
ourselves to a small portion of the best established and most
compelling evidence under each head.

Growth Integration 95

Graded Repetitive Series as Integrative Phenomena

This, perhaps the simplest form of integrational phenom-
ena known to biology, is seen almost everywhere, but shows
itself most typically and strikingly in plants and in many
lower animals. Reference is made to the gradation in the
repetitive or meristic parts appearing in so many organisms.
The most obvious criterion of such gradation is the rela-
tive size of the parts, but, as we shall see later, there is con-
siderable reason for supposing the gradation is not re-
stricted to size. The few examples to which space can be
given are selected to represent as wide a range as possible
of the phenomena under consideration.

Illustrations from Anvmals

The lancelets, fish-like animals of the genus Amptiioxus,
may be noticed first (Figure 55). It will be observed that

FIGURE 55 — SIDE VIEW OF AMPHIOXUS (AFTER PARKER & HASWELL). nch.,
notochord. cir., cirri, or.hd., oral hood, myom., myomeres. dors.fr.,
dorsal fin rays, cd.f., caudal fin. gon., gonads

the creature tapers off toward both ends and that the series
of metameres, myom, usually called myomeres because they
compose the main body-musculature, diminish not only in
dorso-ventral measurement from near the middle each way,
but are also thickest in the mid-region and become thinner
as they progress toward each end.

Something of this size scheme of body-parts is very com-

96 The Unity of the Organism

mon in the animal kingdom. Figure 56, a photograph of
the skeleton of a python (see frontispiece), shows in a gen-
eral way the size-relations of metameric skeletal parts in a
higher vertebrate. Something of the extent to which the
proportionality of parts of the individual metameres is car-
ried out in this skeleton is shown by tabulating a series of
measurements of the parts :

Position of

vertebra

L.V. T.V. T.ex.zyg. H.D.S. W.D.S.

L.R.

5

6 mm.

5 mm.

11 mm.

15 mm.

4 mm.

25 mm.

50

11

16

23

11

8

48

100

13

17.5

28.5

9

7

79

150

13

16.5

26.5

7.5

7

82.5

200

11.5

15

22.5

5.5

7.5

69-5

250

10

10.5

16.5

5

5.5

52.5

300

6

5

7

4

2

0

327

4

3

4

2

2

0

"Position of vertebra" refers to the serial number, beginning
with the head-end, of the vertebra measured. Legend: L.V.,
Length of vertebra, measured from the posterior edge of one
dorsal spine to the anterior edge of the one next behind it. T.V.,
thickness of vertebral centrum in its thinnest part, i.e., near the
middle. T. ex. zyg., thickness of vertebra at extreme of posterior
zygopophyses (articulating processes) . H.D.S., height of dorsal
spine. W.D.S., width of dorsal spine. L.R., length of rib.

The starfishes are another class of animals which exhibit
beautifully this size gradation of repeated parts, both their
"tube-feet" and the calcareous skeletal supports being
graded proportionately to the tapering arms of the animal.
The following table presents a series of measurements of the
two organ systems just mentioned, from a single arm of
Astrospecten calvfornicus. The dimensions are in millimet-
ers, and the series proceed from the proximal to the distal
end of the arm.

Growth Integration 97

TABLE

Series Tube- feet Ambulacral plates Adambulacral plates

number Length Length Width Thickness Length Width Thickness

5

3.9

6.4

3.1

1.2

3.8

3.6

1.0

10

3.7

3.9

2.7

1.3

3.5

2.4

1.2

15

3.5

3.5

2.4

1.3

2.8

2.2

1.4

20

3.1

2.6

2.0

1.2

2.4

2.0

1.2

25

2.2

2.3

2.0

1.0

2.1

1.9

1.1

30

2.1

2.1

1.9

0.9

1.9

1.7

1.2

35

2.3

1.8

1.6

1.0

1.7

1.3

1.1

40

1.7

1.2

l.l

0.9

1.4

1.0

1.0

45

1.8

0.8

1.0

0.6

1.0

0.8

0.9

50

1.3

0.7

0.7

0.5

0.7

0.7

0.6

55

0.9

60

0.8

-

65

0.7

68

0.5

Not only do these graded meristic series appear in the
individual makeup of a great range of animal species, but
they occur in the colonies of many species in which aggrega-
tions are produced by budding. Sometimes, as in many al-
cyonaria, the size gradations are very obvious, while in other
groups the distinctions are so small as to be discoverable
only by close quantitative study. An example of this latter is
furnished by the plumularian hydroids. A typical colony of
the genus here studied, Torrey writes, "closely resembles a
feather, of which the shaft is represented by the stem and the
veins by the two ranks of alternating branchlets, or hydro-
cladia, corresponding to barbs. Each hydrocladium is
divided by more or less definite nodes into internodes and
bears on one aspect — the same in all hydrocladia — a compact
series of hydranths, one to each internode."

Without entering into the tabular and graphic details
contained in this study, the author's summarized statement
concerning one of the tables will suffice : "It will be seen from
the table that, as the tip of the colony is approached, not

98

The Unity of the Organism

only do the hydrocladia possess fewer and fewer hydroth-
ecae, but the dimensions of the latter through the mesial
nematophore reaches its minimum more and more rapidly.
Since the hydrothecae, once formed, do not enlarge with age,
it is clear that for such colonies as this, there is a limit of
growth and a specific form." 1

This correlation and proportionality among repetitive
parts is frequently observed within the bounds of particular

a.

FIGURE 57 — TENTACLE OF HALOCYNTHIA JOHNSONI (SCHEMATIC; AFTER
RITTEH).
'a, axis, a', axis of branch, b', b", primary and secondary branches.

organs, as for example, in the branching tentacles occurring
in various groups. A detailed study of one of these cases
was made by me some years ago on the tentacles of an as-
cidian. Figure (pi. 12, figure 13) of this study, sup-
plemented by the following statement, illustrates the
point. "Although this figure is diagrammatic in a way, it is
accurate as to numbers of branches. The positions, too, of
all the branches and length of the primaries were deter-
mined by micrometer measurements, and the secondaries were
drawn as accurately as possible." 2 For the rest, the figure
(Figure 57) tells its own story.

Growth Integration
Illustrations from Plants

99

But it is in the plant world that these graded series of
homonymous parts in individual organisms arc most strik-
ingly seen. It occurs in what is perhaps its most typical,
least modified expression in the arrangement and size rela-
tion of parts in the leaves of many ferns and palms. But
the compound leaves of innumerable flowering plants illus-
trate it very beautifully. Figures 58, 59, and 60 (Acacia,
Vicia and Cassia) show three types of compound leaves
based on the mode of gradation of the leaflets. These might

FIGURE 58. ACACIA ELATA.

100

The Unity of the Organism

be described as the bi-gradient, the direct gradient and the
reverse gradient types, depending on whether the gradation
is from the mid-region of the axis both ways (figure 58), from
the proximal toward the distal end (figure 59), or from the
distal toward the proximal end (figure 60).

FIGURE 59. VICIA OIOAKTEA.

FIGURE 60. CASSIA SP.

Almost all simple leaves of seed plants show something of
the same scheme. As examples, typical elliptical-entire
leaves of the elm and poplar and such typical lobed leaves as
those of most oaks may be pointed to.

Nearly every twig of a tree which represents a single
annual growth impulse, in cases where the growing period is
restricted to a small part of each year, presents a size gra-
dient in the leaves distributed along the axis. A particularly
striking illustration of this is furnished by the California

Growth Integration 101

coast redwood, Sequoia sempervirens (figure 61), where the
new segment is short, is added end on to the one before it
until a considerable succession of segments is produced, and
where the leaves are retained for several years. That each
segment in these leaves is an annual production is not cer-
tain, probably several segments being sometimes formed in a

FIGURE 61. SEQUOIA SEMPERVIREXS.

single season ; but however that may be, that growth occurs
in a series of impulses, each of which is sharply recorded
in the size gradations of the repeated parts, is obvious
enough.

It is a familiar fact, too, that in many plants a similar
quantitative gradation of the reproductive parts along an
axis occurs, but the extent to which this scheme pervades
the constituents of the members of the series, even to the
seeds, appears not to have attracted much interest on the
part of botanists. To illustrate this point I present a single
set of measurements, one of many which I have collected,

102 The Unity of the Organism

of parts of the fructiferous organs of plants. These meas-
urements are of Frasera perryl, an abundant annual in
southern California and rather specially favorable for such
a study in that the fruit stalk is single in each plant, stands
up intact and rigid after it is fully ripe and dry, and is al-
most mathematically regular in the disposition of its parts.
The table was compiled from measurements of a single plant,
and three measurements pertaining to each seed vessel are
given, namely, the length of the interval on the main axis
between each two vessels, the length of the pedicels which
bear the vessels, and the length of the vessels themselves.
The measurements are all in millimeters. Several other di-
mensions might have been taken, which would almost cer-
tainly have produced similar results.

Series
Number

Length of
Internode

Length of
Pedicel

Length of
Capsule

1

27

36

21

2

19

29

18

3

19

32

18

4

22

34

16

5

23

29

16

6

19-5

28

17

7

18.5

25

17.5

8

17

25.5

17.5

9

19

25

18

10

18

25

16

11

18

21

16.5

12

15

21

16

13

15

22

13.5

14

15

22

14

15 15 21.5 13

That these gradations hold, at least in some plants, even
to the seeds is certain as the following tabulation of the
weight of seeds from different parts of the seeding axis of a
wild mustard plant (Brassica nigra) shows. The figures
were compiled from the weights of seeds taken from groups

Growth Integration 103

of twenty capsules from the bases, middle portions, and dis-
tal ends, respectively, of six such stalks. The weights given
are in grams.

Total weight Number of Seeds Av. Wt. per Seed
Base .903 950 .00095

Middle .694 733 .00088

Tip .330 525 .00063

This mere glance at an exceedingly common phenomenon
in living nature must suffice for the present.

Justification for Bringing All These Phenomena Under One

Head

Probably about the first question that most persons would
raise concerning what we have presented would be as to how
far the series dealt with have anything to do with one an-
other. Especially, we may apprehend, would most biologists
question the justifiability of bringing together the meristie
phenomena in animals and the repetition of parts in plants.
If such a collocation of phenomena must be justified on the
basis of known causal factors, then undoubtedly is justifica-
tion impossible in the present state of knowledge. But justi-
fication of this sort is not called for by the point now oc-
cupying us. What concerns us at present is the quite for-
mal fact that when any lot of homonymous objects fall into
a quantitatively graded series the members of that series
have a fixed relation to the scries as a whole. They are not
interchangeable with one another. Each is a function,
mathematically speaking, of its set or series. Vertebra TO
of the python's skeleton, myotome m of the amphioxus body,
tube-foot in of the starfish arm, branchlet m of the ascidian
tentacle, leaflet m of the vetch leaf or of the redwood shoot,
seed-vessel m of Frasera, seed-lot m of the mustard plant,
and m, or any other member you choose from any other

104 The Unity of the Organism

series whatsoever, is a determinate thing; it is what it is
partly because of its position in the series regardless of
whether the physical or other producing agent of the dif-
ferent series be the same or wholly different.

Even this purely structural formal basis establishes
the fact of a measure of integratedness for all individual
organisms in which the phenomenon appears. But to leave
the subject at that would be superficial and unsatisfactory
indeed. However, reflection makes it almost certain that
there is some sort of causal basis for the phenomena. This
conclusion follows first from the fact that the series result
from the growth of the organism, and second from the cer-
tainty, at least in many cases, that the continuance of life
of the individual involves the maintenance of the series, this
in turn involving some measure of metabolic interdepen-
dence among the members of the series.

Attempted Causal Explanation of These Series

For establishing the general truth of this type of inte-
gration we need not, in strictness, go any further than we
have gone. Nevertheless, the importance of the subject
justifies a few remarks on attempts that have been made to
explain the series causally. The best known of these comes
from botanists, and conceives that the diminishing series of
leaves and other structures, seen with more or less distinct-
ness almost universally among plants, is due to the increas-
ing remoteness of the successive parts from the roots of the
plant, that is, from the main source of the plant's food. It
is obvious, however, that this explanation is not of general
application, since in animals the food does not come from a
root system which anchors the organism to its food-yielding
medium. Nor is it possible to bring the series in all animals
into correlation with a blood circulatory system, as their
existence in many coelenterates, hydroids and alcyonarians

Growth Integration 105

for example, where no circulation exists, shows. It is al-
most certain, too, that the series occur in many plants that
have no sap system such as is assumed by the physiological
explanation above indicated. Many of the marine algae
come under this head, a striking example of which is the kelp
Macrocystis pt/rifera of the western coast of both Americas.
That the lamina; of this plant fall into a beautiful direct
gradient series is a fact which can not escape the- notice
of any one who sees them. The question of whether each
streamer of laminae reaches finally and necessarily a limit of
growth in which the size series is present is not so certain,
but from considerable attention to the question I am almost
sure this is the case, although the point needs more study.
Another interesting and probably useful course of reason-
ing about organic growth attempts to connect the results of
growth with autocatalytic chemical action. Although these
attempts have not, so far as I am aware, taken special cog-
nizance of the natural size series which are occupying us,
but have been concerned with the weights or volumes of or-
ganisms at various stages of growth, there is little doubt
that the phenomena we have been considering are closely
connected with those dealt with in these attempts. T. Brails-
ford Robertson seems to have devoted more thought to this
matter than any one else. The following, taken from the
summary of conclusions found in his original paper, pre-
sents the most essential parts of his theory: "(1) In any
particular cycle of growth of an organism or of a particular
tissue or organ of an organism the maximum increase in vol-
ume or in weight in a unit of time occurs when the total
growth due to the cycle is half completed. (£) Any particu-

x

lar cycle of growth obeys the formula log - - K(t-tx),

A. X

where a: is the amount (in weight or volume) of growth
which has been attained in time t, A is the total amount of
growth attained during the cycle, K is a constant, and £1 is

106 The Unity of the Organism

the time at which growth is half completed." 3

Assuming that the growth of an amphioxus, lot us say, to
adulthood represents a growth cycle of this statement, that
the production of somites begins with the most anterior one
and proceeds toward the tail, and that the successive growth-
increments (corresponding to x in the formula) arc regis-
tered in the somites as we find them, then the animal's body
as exhibited by its myomeres would correspond fairly well
to Robertson's statement under (1), as would several of the
other growth series we have glanced at, and as would also
great numbers of series presented by ordinary plants and
animals.

The formula for growth contained in (2) is, according
to Robertson, "such as would be expected to hold good were
growth the expression of an autocatalytic chemical reac-
tion." Assuming the general correctness of these state-
ments, no one interested in the larger problems of organic
growth could hesitate to believe that they must be impor-
tant in some way.

However, that the relations shown do not prove that au-
tocatalytic chemical activity is a cause of growth in any-
thing more than a subordinate, contributory way, is obvious
on reflection". In the first place, Robertson himself has
pointed out, in substance, that such action says nothing
about the particular shape which the mass of transformed
substance takes, but since some characteristic configuration
or shape is fundamental to all organic growth, the entities
for which A and x stand in the formula are really only ab-
stractions. Although the formula may apply approximately
to a great many organisms, it will apply to none exactly,
except by chance to a very occasional one. This is the gen-
eral form of criticism, illustrations of which are seen in the
fact that in the series of direct and inverse gradients shown
in the vetch (figure 59) and Cassia (figure 60), respectively,
the formula appears not to apply at all. The general pur-

Growth Integration 107

port of the strictures here placed upon the value of this
explanation of growth is well brought out by Moeser, who
says, probably with literal truthfulness, "One will not find
two germinating plants (Keimpflanzen) which would have
exactly the same growth curve, even though they proceed
from seed of absolutely the same weight and grow under
exactly the came conditions."

The kernel of this criticism is that even though it should
be established, as very likely it will be, that autocatalytic
action is an essential factor in all growth, it can be a causal
explanation in only a partial and subordinate sense. This is
so because although K is a constant for a particular indi-
vidual as observed, it assumes different values for different
groups, partly at least because no account is taken of size
or configuration. Moreover, even if these factors were con-
sidered, there would be nothing corresponding to a physical
constant (depending only on autocatalytic action), since
one of the most distinctive things about organic growth is
that it is differential, the differentials corresponding to the
taxonomic group to which the individuals belong.

Even though we are still much in the dark as to what the
causal nexus is between the growth processes and the quan-
titatively graded series so widely seen in nature, it seems
certain that some such nexus exists, and that its operation
constitutes a true integrational factor in the individual. It
appears, too, that this type of integration is about the
simplest and that it accompanies the simplest type of dif-
ferentiation, the two together constituting the simplest type
of organization above the organization of the cell. But into
this interesting subject we can not now go.

Axial Metabolic Gradients as Integrative Phenomena

For the most thorough and sustained experimental inves-
tigation of the primary intcgrative processes in growing

108 The Unity of the Organism

organisms that has been made, biology is indebted to C. M.
Child. In two recent volumes he has summed up and sys-
tematized the elaborate researches prosecuted by him in this
field almost exclusively for fifteen years, and has presented
his conclusions more fully than in any of his previous writ-
ings.

The limitation set for the present to the constructive part
of our enterprise makes it impossible to do more than touch
at a few points the great mass of experimental evidence on
which Doctor Child bases his conclusions. Fortunately,
however, the kernel of the conclusions can be stated rather
clearly in a small space.

Although (surprisingly, it seems to me) Child refers
hardly at all to the graded meristic series occurring in na-
ture, to which the preceding pages have been devoted, it
can hardly be doubted that the phenomena with which he
deals, and calls "axial gradients," come under the same head
as do those which have been occupying us. The phenomena
which in the first instance Child has been concerned with,
have been brought to light mainly through studies on re-
generation in many lower animals. But the general con-
clusions reached are far broader than this ; indeed they ex-
tend to well-nigh the whole scope of organic growth, but
especially to growth which involves elongation either of the
whole organism or of certain parts of organisms. Thus the
head-tail type of individual, whether the body be segmented
as in arthropods and in many worms, or unsegmented as in
other worms and in molluscs, is perhaps the most striking
exemplification of the axial gradations with which Child
deals. The following quotation shows the generality with
which he views the matter from the ontogenic side: "Gra-
dients in rate of cell division, size of cells, condition or
amount of protoplasm in the cells, rate of growth, and rate
and sequence of differentiation are very characteristic fea-
tures of both animal and plant development. Such gra-

Growth Integration 109

dients are definitely related to the axes of the individual or
its parts, and are evidently expressions of axial metabolic
gradients. While the existence of such gradients indicates
the existence of gradients in activity of some sort, the
various kinds of gradients are not all necessarily pres-
ent where metabolic gradients exist. In some cases the vis-
ible gradient may be a gradient in rate of growth or in
protoplasmic constitution ; in still others a gradient in
sequence of differentiation, etc., and sometimes metabolic
gradients exist without any structural indications of their
presence. At best these various kinds of gradients are
merely general indications of differences in metabolic rate
and undoubtedly in many cases the visible^ differences along
an axis represent something more than differences in meta-
bolic rate. The important point is that visible indications
of graded differences in metabolic rate occur so generally
in definite relations to the chief axes of the body.5

One phase of this general statement is the developmental
correlation that various regions of the body in many lower
animals have with the head or anterior end, these regions
being developmentally dominated, in Child's expression, by
the anterior end proportionally to the distance of the re-
gion from the end.

A typical case is furnished by flat-worms of the genus
Planaria, animals especially favorable for experiments in
regeneration, since they are very hardy to laboratory con-
ditions and have great powers of reconstituting themselves
from pieces of various sizes, shapes and positions cut from
them. "Any piece of the planarian body," says Child, "is
capable of giving rise to all parts posterior to its own level,
whether a head is present or not, but no piece is capable of
producing any part characteristic of more anterior levels
than itself, unless a head begins to form first." 6

From a great mass of experimental evidence produced b^
Child and others we have the following: "These facts force

110 The Unity of the Organism

us to the conclusion that in such experimental reproductions
there is a relation of dominance and subordination of parts.
The apical or head region develops independently of other
parts but controls or dominates their development, and in
general any level of the body dominates more posterior or
basal levels and is dominated by more anterior or apical
levels." 7

A really unique merit in Child's work is the fact that he
has given special attention to the connection of these axial
gradients manifesting themselves in various structural and
functional ways, with the fundamental metabolism of the'
organism. Several methods of experimenting have been em-
ployed to this end, the one most frequently used being what
he calls the susceptibility or survival-time method. The es-
sence of this depends upon the fact, determined by many ob-
servers, "that a relation exists between the general meta-
bolic condition of organisms, or their parts, and their sus-
ceptibility to a very large number of substances which act
as poisons, i.e., which in one way or another make meta-
bolism impossible, and that difference in susceptibility may
be used with certain precautions and within certain limits as
a means of distinguishing differences in metabolic condition,
and more specifically, differences in metabolic rate." 8

The demonstration of metabolic gradients by this method
depends upon the fact that "death and disintegration' of dif-
ferent parts of the body usuallv follow a regular sequence,"
this making it possible "to determine the time, not merely of
disintegration of the whole animal, but of the various re-
gions of the body." 9

Another way of showing difference in rate of metabolism
in different parts of the organism is by the use of the biom-
eter, an apparatus for estimating minute quantities of carbon
dioxide, recently devised by S. Tashiro in connection with
his important researches on carbon dioxide production in
nerves. By these methods it is shown, pointing to a single

Growth Integration 111

instance, that in pieces of a flat-worm isolated by cutting
"the rate of metabolism is higher in long anterior pieces
than in posterior pieces of the same length."

Starting from this low but seemingly universal level of
integrative phenomena in the individual, Child formulates
views of the nature of organisms that agree very well with
the organismal standpoint upheld in this volume. He
writes : "The organic individual appears to be a unity of
some sort. Its individuality consists primarily in this unity,
and the process of individuation is the progress of integra-
tion of a mere aggregation into such a unity, for this unity
is not simply the unity of a chance aggregation, but one of
a very particular kind and highly constant character for
each kind of individual. In all except the simplest individ-
uals it determines a remarkable degree of uniformity and
consistency, both in the special relations of parts and the
order of their appearance in time, and also in coordination
or harmony of functional relation to these parts after their
development." ll

Meristic Gradients and Metabolic Gradients Both
Phenomena, of Growth Integration

In view, then, of the exceedingly wide prevalence in living
nature of axially disposed meristic series quantitively
graded, and of the equally wide or even wider prevalence of
axial gradients on the basis of metabolic activity, the
gradients of both sorts arising as fundamental growth phe-
nomena, it appears impossible to avoid recognizing our first
category of integration, namely, growth integration, as
about the most simple and primal and universal of all these
categories, at least for multicellular organisms. It seems
as though the other kinds of differentiation and integration
are superposed, as one might express it, upon this primordial
kind. To a consideration of the other, superimposed inte-

112 The Unity of the Organism

grations we now pass, taking them again in their seeming
order of obviousness.

REFERENCE INDEX

1. Torrey 139 7. Child (1) 215

2. Ritter ('09) 71 8. Child (1) 66

3. Robertson 612 9. Child (1) 77

4. Moeser 373 10. Child (1) 73

5. Child (2) 65 11. Child (2) 2

6. Child (1) 213

Chapter XVIII
CHEMICO-FUNCTIONAL INTEGRATION

Functional as Contrasted with Growth Integration

A SSUMING Child's theory of metabolic gradients to be
•*T^well grounded, we are furnished thereby with one im-
portant insight into the chemical processes involved in the
unity of the individual organism. But that process is con-
cerned primarily with the growth, with the production of the
individual. The question now is, are there chemical proc-
esses the object of which is to maintain the functional unity
of the complete or nearly complete individual? Are there
chemical operations the office of which is to preserve a
proper interrelation among the parts of the organism as
these perform their special offices?

That such is to some extent the significance of most if
not all internal secretions as usually understood is indicated
by the fact that the functional disturbances attending re-
moval of the thyroids or other glands from various animals ;
and by the further fact that where internal secretions play
a part in development, their action is rather that of stimula-
tor, or at least modifier, than of true producer.

The conception of internal secretions as being at least as
much regulators of physiological function as of growth is
illustrated by cases of hypopituitarism of the post-
adolescent type, like those described by Gushing, for ex-
ample. In the series of cases of disease due to "pituitary
deficiency" the first symptoms appeared when the subjects
were from thirty to forty years old.

113

114 The Unity of the Organism

The Conception of "Internal Secretions"

The nature of the phenomena now to be considered, and
their significance for our discussion make it desirable to
think about these secretions from the broad standpoint first
stated, according to Bayliss, by Brown-Sequard and d'Arson-
val, namely, as materials produced by any living cells or
tissues which are discharged into the blood or lymph and
have specific effects on other parts or functions of the or-
ganism. Regarded thus it is now known that many cells of
the organism produce internal secretions. Although we are
more concerned with function than with structure in this
discussion, our purpose will be best served by beginning
with a morphological classification of the secretion-pro-
ducing cells. They may be divided into two categories,
those which are disposed into definite organs, or glands, the
ductless glands of long standing in anatomy; and those
which are not assembled in such organs. Knowledge of this
second class of cells is of recent date, and is fuller from the
functional than from the structural standpoint. The chief
glands, to which the name Endocrine has lately been given
by Schafer, are now so well known as hardly to need men-
tion. They are the thyroid apparatus, including the thy-
roids and the parathyroids, the suprarenal body, the pitui-
tary body, and probably the thymus and pineal bodies. Cells
now known to produce internal secretions but which are not
arranged in glands are certain cells of the pancreas scat-
tered among the pancreatic cells proper; certain cells of
the alimentary mucous membrane ; the interstitial cells of
the ovary and of the testis, and probably certain cells of
the placenta, of the mammary gland, and of the uterus.

Following our usual course of making the treatment
merely illustrative rather than aiming at exhaustiveness,
our selection will include one example from each of these
groups. From the glandular category we take the thyroid

Chemico-Functional Integration 115

apparatus, and from the non-glandular a portion of the
alimentary mucous membrane, namely, that of the duo-
denum.

Effects of Removing the Human Thyroid for Curative

Purposes

As definite knowledge of the great physiological impor-
tance of internal secretions begins with human surgery —
with operations on the thyroid apparatus — we may well
begin our study here. This is the better starting point
in that there is no more striking illustration of how great
a part of the whole organism may be implicated in the
action of internal secretions than is afforded by the prod-
ucts of the thyroids and parathyroids.

The subject first came into clear light in the early eighties
of the last century through the experiences of Swiss sur-
geons, Theodor Kocher and J. L. Reverdin especially, who
removed the thyroids to cure goitre, this disease being spe-
cially prevalent in some parts of Switzerland. The patients
operated on were found to improve rapidly for a time after
the operation, but later untoward symptoms began to mani-
fest themselves. Because the variety and pervasiveness of
these symptoms in a typical case are highly instructive for
us we present them in detail, selecting a description from
Human Physiology by Luciani : "Patients who have under-
gone total thyroidectomy . . . experience the initial symp-
toms of glandular deficiency either at once or at latest some
weeks after the operation. They feel weak, complain of
heaviness of the limbs, and more or less diffuse dull pains,
particularly in the legs, which may become acute and assume
the character of pains in the bones.

"Other more serious symptoms are gradually associated
with the preceding. After four or five months the face
and the extremities swell and become cold, the muscles are

116 The Unity of the Organism

torpid, sometimes rigid, often exhibiting muscular tremors,
and are incapable of carrying out any delicate manual acts
of precision. At first the swelling is variable ; it is more
pronounced in the morning than in the evening, but steadily
increases until it becomes permanent. It is not ordinary
oedema, in which percussion with the fingers leaves a depres-
sion; it is a hard and elastic swelling. It is specially local-
ized in the hands, feet and face, where it produces a char-
acteristic alteration of the countenance. The lower eyelids
are the first to present a sacculated, semi-transparent swell-
ing, which is hard to the touch ; then the infiltration spreads
to the folds of the face, which become smoothed out ; to the
nose, which gets rounded; to the lips which swell, and
bulge outward, saliva dribbling from them. The features
are coarsened and expressionless like those of a cretm.

"The mental functions accord with this appearance, since
they are blunted, so that the patients lose their memory,
become deaf, taciturn, melancholy, self-absorbed, and reply
extremely slowly to questions. They further complain of
slight but perpetual headache; feel an almost constant sen-
sation of cold, which is most acute at the extremities ; at
times they are seized with vertigo, and may even lose con-
sciousness.

"All the symptoms become still further aggravated. The
whole body may grow more bulky from the extension of the
swelling. The skin loses its elasticity, can only be picked
up in large folds, and becomes dry owing to defective
capacity for sweating. The epidermis desquamates in more
or less extensive lamellae, particularly on the hands and feet ;
the hair turns grey, falls out, and gets constantly thinner.

"The heart functions weakly, but with ordinary rhythm ;
the pulse is small and thready. Examination of the blood
shows nothing constant; but there is often a more or less
pronounced and progressive oligocyth<Tmia, which undoubt-
edly contributes to the characteristic pallor of the skin,

Chemico-Functional Integration 117

this being of the earthy, yellow-spotted hue peculiar to
cretins.

"The respiratory rhythm is almost always normal; the
digestive apparatus functions well, as also the urinary
system. The spleen is not enlarged."

This complex (syndrome in medical terminology) of
manifestations is known technically as cachexia thyreo-
priva.

Later experience by other surgeons with the same opera-
tion discovered that in some cases the effects are much more
acute and rapid, and may be replaced by what has been
called "tetany" (though having little in common with ordi-
nary tetanus), ending in death more often than otherwise.

Experimental Thyroid Excision in Normal Lower Animals

No sooner had the far-reaching influence of the thyroid
for the human organism begun to be recognized in this way
than experimentation on inferior mammals was invoked for
further light on the subject. Moritz Schiff, the ground-
breaker in this field, published in 1884 the results of the
removal of the thyroid from a large number of dogs. In
all cases where the whole thyroid apparatus was excised
the dogs soon died after a run of such symptoms as tremor,
spasms, and convulsions. Nor did Schiff rest content with
merely ascertaining the effects of removal, complete and
partial, of the thyroid apparatus. He found that these
effects could be entirely prevented by grafting a portion of
the gland under the skin or into the body cavity of the
animal before the thyroid operation, or by injecting thyroid
juice into the blood or lymphatic systems, or by feeding raw
thyroid to the dogs. The story of how these experiments
led to the now widely practiced treatment of myxoedema with
thyroid or thyroid extract would be out of place here,
though it should not be passed wholly unnoticed.

118 The Unity of the Organism

So an enormous mass of evidence, experimental, surgical
and clinical, is now in court demonstrating that for some
animals at least, among them being the human and the
canine species, products of the thyroid apparatus are in-
dispensable to the normal life, the symmetrical growth and
balanced physiological activities of the organism. That
the apparatus is essential to the "Hormonic Equilibrium"
of the organism in some animals is beyond question.

While no pretense can be made at an exhaustive exami-
nation of this evidence two phases of our discussion make
it desirable to cary the examination on the manifestational
side a little farther. One of these is the importance of
making as objective and emphatic as possible the extent of
the manifestations in the individual; the other is the ques-
tion of the generality, taxonomically speaking, of the thy-
roid apparatus.

In the interest of the first of these I present, verbatim, the
report of a single case of complete thyroidectomy, the animal in
this instance being a fox. The individual concerned was a female
less than one year old.

"Oct. 28. Glands removed; good recovery.

"Oct. 2Q. Normal, but does not eat.

"Oct. 30. Salivation, rapid breathing, strong tremors and tet-
any from 7 A. M. to 2 P. M. ; quiescent but weak during the rest
of the afternoon.

"Nov. 1. „ Normal, but rather weak; eats; no sign of tremors
or salivation during the day.

"Nov. 2. Restless ; slight tremors ; dyspnoea ; does not eat.

"Nov. 3. Some depression, but no tremors or salivation until
4 P. M. ; does not eat. At 4 P. M. spasms appeared and con-
tinued unabated as long as observed (7 P. M.).

"Nov. 4. Found dead at 4 A. M. Post-mortem examination
revealed no parathyroids nor accessory thyroids." :

As to taxonomic range and character of manifestation of thy-
roid influence, much diversity might have been anticipated on
general natural history grounds. As far as investigations have
gone they realize these anticipations. A summary of results will
serve our purpose, and this is at hand in Schafer's volume already

Chendco-Functional Integration 119

cited.

Concerning the effects of removal of the thyroid appa-
ratus he says :

"The most acute symptoms are exhibited by carnivora such as
dogs, cats, foxes, and wolves (Vincent), and the young of herbi-
vora (v. Eiselsberg, Sutherland Simpson) and are of a nervous
nature. . . . Some species exhibit no symptoms whatever — at
least when the operation is performed on the adult. Horsley
states that this is the case witli birds and rabbits; but according
to Gley, the latter are affected if care is taken to find and re-
move all four parathyroids, and Doyon and Jouty obtained typical
tetany in hens which had been parathyroidectomized. . . . Ac-
cording to Vincent and Jolly badgers are totally unaffected by
complete removal of both thyroids and parathyroids." 3

From the anatomical characteristics of the organs,
and from the known effectiveness of minute portions of
them, such statements as the last must be taken with re-
serve. Although these results show by their diversity that
an enormous amount of study remains to be done on the
comparative side, they leave no question that the secretion
of the thyroid apparatus is important for the general
health and equilibrium of most animals in which it occurs.
The measure of this importance in the eyes of some authori-
ties is seen in such a statement as, "No cell anywhere in the
body can reach morphological perfection without thyroid
stimulus." 4

The Internal Secretion of the Duodenal Mucous Membrane

We now pass to an examination of the effects of the
internal secretion of the duodenal mucous membrane. This
particular secretion is selected for the reasons that it is,
according to Bayliss, one of its discoverers, "the most
typical of all the chemical messengers"; that it was one of
the first to be investigated ; and that it is one of the few
which have been isolated as definite substances.

120 The Unity of the Organism

The mode of operation of this secretion is tersely stated
by Bayliss: "Food entering the duodenum causes the pro-
duction of a special substance which enters the blood and
excites the pancreas to pour into the duodenum a digestive
juice." 5

That the presence of various substances, especially acids,
in the duodenum, induces a flow of pancreatic juice was
known when Bayliss and Starling began their work in this
field; but up to that time the excitation of the pancreas to
such action was supposed to be through a nerve reflex.
These investigators had reported in 1902 6 that acid in the
duodenum is able to cause the pancreas to secrete after
nervous communication between the intestinal wall and the
pancreas is excluded. They went further and obtained an
extract from the duodenal mucous membrane which, being
injected into a vein, induced a copious flow of pancreatic
juice. The substance, whatever it is, which acts thus they
call secretm. It has been surmised by a few physiologists
that the effects are not due to the direct action of the secre-
tion on the pancreatic gland-cells, but that the influence
is exerted through the vaso-dilator mechanism. But this
surmise is negatived by the demonstration that the secretin
will induce the flow of pancreatic juice while it does not
alter the blood pressure. The case seems, then, fully estab-
lished, and is so clear-cut and relatively simple an instance
of the coordinated functioning of two wholly distinct parts
of the body through chemical means, that it is desirable
to get sharply before us the known steps in the process.
In the course of normal digestion, food acidulated in the
stomach passes into the duodenum. Here, probably in
virtue of its acidity, it acts upon the cells of the mucous
membrane in such a way as to induce them to produce a
substance which is discharged, not into the intestines, there
to take its part in digestion, but into the blood. By the
blood stream the substance is carried through its whole

Chemico-Functional Integration

circuit, hence through the lungs and so on, around to the
pancreas, the typical gland-cells of which it excites into
activity, so that the pancreatic juice, an "external" instead
of an "internal" secretion, is poured into the duodenum to
exercise its digestive office on the same food which started
the cycle of activities.

It was with this substance particularly before their minds
that the authors adopted the name hormone to designate
substances which act thus. "The group of substances re-
ferred to," says Bayliss, "which includes adrenaline and the
various internal secretions, is characterized by the prop-
erty of serving as chemical messengers, by which the activity
of certain organs is coordinated with that of others. They
enable a chemical correlation of the functions of the organ-
ism to be brought about through the blood, side by side
with that which is the function of the nervous system." 7

This reference to the side-by-side activities of chemical
messengers and nervous system in integrating the organism
touches a subject of the utmost importance. Considera-
tion of it must, however, be deferred until we have looked
a little more into the nature of hormones.

The Nature of the Active Substances in Internal Secretions

That the peculiar iodine-rich albuminous substance ob-
tained from the thyroid by Baumann in 1895 and since
observed by other investigators, contributes in some essen-
tial way to the action of the secretion of this gland is the
belief of apparently a large majority of authors (Bayliss,
Eppinger, Howell, etc.), but not of all (Luciani). In view
of the uncertainty on the point Schafer's proposal "to ex-
press our ignorance by a term which implies no theory"
may well be accepted, with the proviso that the term pro-
posed be really taken as evidence that something though
not everything is known about the substance. A part of

The Unity of the Organism

the proposal is worth quoting. "I propose therefore pro-
visionally to apply the word thyrine to denote the active
principle, whether it be identical with or contained in the
iodothyrin of Baumann or not." 8

Thyrine then becomes the name of a substance the source
and some of the activities of which are known, but whose
main physical and chemical attributes are unknown.

Concerning the mode of action of thyrine there are sev-
eral divergent views, all based on some evidence and so
perhaps not entirely antagonistic. Is the antitoxic theory
of Luciani 9 partly right, right as regards the parathyroid
secretion (Moussu, Vassale and Generali), and partly
wrong, wrong as to the secretion of the thyroid proper, this
being trophic rather than antitoxic? 10 May there not be
more in the enzymic theory suggested some years ago, than
later writings have been inclined to favor? Does the fact
that internal secretions seem to be simpler than enzymes,
as indicated by their greater resistance to heat, preclude the
possibility that their normal mode of action is of the enzyme
type after all? That is, may it not be necessary to extend
the conception of enzymic action (which is surely generic
anyway) to include the various sorts of activity presented
by hormones, understood in the sense given it by its origi-
nators ?

But neither can the resemblance of internal secretions
to drugs, so far as their action is concerned, be overlooked.
This has been dwelt upon by Schafer " and has important
bearings on the problems of the origin as well as on the
chemical nature of the substances.

Another aspect of the mode of action of internal secre-
tions is that of whether the effects are to stimulate or inhibit
the activity of the organ or tissue on which they operate.
Schafer and others make a special point of this, directing
the attention to the fact, by'way of illustration, that the
adrenaline of the suprarenal medulla causes contraction of

Chemico-FiMictional Integration 123

the plain muscle of the blood vessels and inhibition of that
of the intestines.12 The distinction has an undoubted natu-
ral grounding, and so is in the interest of accurate descrip-
tion and clear conception.

As to the actual chemical composition of internal secre-
tions, knowledge is exceedingly meager. More is known
about adrenaline, the active principle of the suprarenal
gland, than about that of the secretion of any other gland
or tissue. This was isolated by the Japanese chemist
Jokichi Takamine in 1901, and has since been more fully
examined by several investigators, notably by T. B. Aldrich.
It is described as a micro-crystalline substance occurring in
at least five crystal forms. Aldrich assigns to it the em-
pirical formula CgHjoNO.^, this structure placing it not far
from tyrosin in the benzene or aromatic series. Of special
interest is the astonishingly minute quantites which produce
physiological effects. According to Aldrich 0.000001 gram
of an aqueous solution of the chloride per kilo of body
weight injected into the blood system raises the blood pres-
sure 14 mm. of mercury.13

The chemical nature of Tethelin, lately isolated from the
anterior lobe of the pituitary of the ox, has been studied
by its discoverer, Robertson. It is described as white or
pale cream colored, readily powdered, highlv deliquescent,
and having a greasy odor and slightly acid reaction in
aqueous solutions. It contains 1.4 per cent of phosphorus
and four atoms of nitrogen for every atom of phosphorus.
The phosphorus-nitrogen content of the substance is con-
sidered by Robertson as specially significant, since this
seems to ally it chemically with "phytin," a substance found
in the rapidly growing parts of plants, and in milk. The
natural suggestion is that the growth-promoting substances
in plants, milk, and the pituitary secretion are chemically
related.

124 The Unity of the Organism

The Close Resemblances and Interrelations of the Different
Internal Secretions

Even the meager, merely illustrative examination of in-
ternal secretions we have been able to make brings out the
close resemblance there is between the several endocrine
glands, and also between the physiological effects of the
various secretions. These resemblances suggest an intimate
organic interrelationship among all the internal secretion-
producing parts of the body.

All investigators in this field, no matter to how restricted
a section of it their efforts are primarily directed, seem to
come upon the interdependence of the sources and activities
of hormones. To illustrate, Gushing, whose central interest
has been the hyphophysis, is led to conclude that experi-
mentally induced hyphophyseal deficiency works histologi-
cal changes in many if not all the other ductless glands.
It is not surprising, consequently that far-reaching theories
have been elaborated on the basis of these relationships.

Bayliss refers with approval to Elliott for the conserva-
tism with which he sums up the present state of knowledge on
this aspect of the general subject. But even so, features are
pointed out "which suggest a common bond":

"(1) Carbohydrate metabolism is influenced, not only by
the pancreas, but also by the thyroid in super- activity, in
acromegaly, and by the injection of adrenaline.

"(2) Growth is affected by the testis and the cortex of
the suprarenals, arrested by the absence of the thyroid.

"(3) Nervous implications.

"(4) The pituitary becomes hypertrophied when the thy-
roid is removed. Acromegaly may lead to enlargement of
the thyroid." 14

At the other extreme of what may be regarded as legiti-
mate scientific theorizing, we have the views of Sajous, who
believes research will finally demonstrate a relationship be-

Chemico-Functwnal Integration 125

tween all the ductless glands the combined functioning of
which dominates most of the activities, normal and patho-
logical, of the organism.

Sajous' elaborately worked-out theories of internal se-
cretions, especially in their relation to disease and medical
practice, are opposed at many points to prevailing opinion
based on present day research. Nevertheless regarded from
the standpoint of general biology there would seem to be
much merit in his effort on the one hand to find a common
ground in the metabolic processes for all the phenomena at-
tributed to endocrinal activity; and on the other hand to
find a more consistent morphological and physiological basis
of definition and classification of internal secretions and the
structures which produce them than has yet been recognized.
For example, whether Sajous is right or not in contending
that the pituitary body does not produce an internal secre-
tion, certain it is that the non-glandular structure of its
posterior part, extracts of which alone have slowing effects
on the heart, is strongly suggestive to the critical naturalist
that the inclusion without qualification of this part at least
of the organ among the endocrine glands is an instance of
what is known to taxonomists as "lumping" in classification
— a kind of practice that advance in knowledge always finds
to be inadequate for purposes that are critical. Sajous'
late summary of his views is highly suggestive to the general
biologist, even though it is excessively theoretical in some
parts.

As far as a much interested outsider can judge, the pres-
ent state of understanding of the relationships among the
internal secretions is set forth with exceptional judiciousness
by Waller. "There can be little doubt," the author opens
his discussion by remarking, "that the various internal secre-
tions are most closely correlated, yet perhaps the most dif-
ficult, and also the most fascinating problem of present day
medicine, is to assign to each its proper and right share of

126 The Unity of the Organism

importance." 15 This statement, coupled with the fact that
one of the main objects of the discussion is to display the
many contradictions which the author's large experience as
a practitioner has found in the action of the internal se-
cretions, is about the most striking, and from our stand-
point most significant thing about this paper.

As one illustration of the agreement of action of the se-
cretions, or at least of the influence of the endocrine glands,
it is pointed out that changes in calcium metabolism have
been observed after removal of the thymus ; in disease of the
pituitary and of the pineal bodies ; after castration ; after
ovariotomy; after removal of the suprarenals, and after re-
moval of the thyroids and parathyroids.

Of the numerous instances of contradiction which he
brings out, we mention only that concerning tetany. This
he shows may result from either removal of the thyroid or
from an overdose of thyroid extract. The explanation of
the contradictions in the action of a given secretion favored
by the author is that of the "varying influence of the other
internal secretions." But the descriptions given seem to
leave no doubt that difference in type of individuals also
comes into the explanation. Thus among children afflicted
with enlarged tonsils and adenoids, the two distinct types
dependent upon the character of the symptoms, is a case in
point. One type is dull and stupid, stunted in growth, has
dry coarse skin, and may display symptoms of rickets. The
other type is vivacious physically and mentally, given to
peevishness, irritability and quick fatigue, and always want-
ing a change of activity. This type is over-tall for its age,
perspires readily and is fine-skinned. Both types of cases
are benefited at least for a time, Waller says, by treatment
with thyroid extract.

Concerning the general nature of the interdependence
among internal secretions, this author's views seem to me so
eminently sound that I cannot refrain from quoting them in

Chemico-Functional Integration 127

some fullness : "Considerable stress has been laid upon the
antagonism of different internal secretions by various au-
thors. I believe -we should gain a truer insight into their
working if we dwelt rather upon their harmony. It does not
strike me as a very high conception of the human organism
that health should consist in the balance of dissentient or
antagonistic forces. It would seem far more ideal that all
the internal secretions should work together for the common
good of the organism, and that when some special demand is
made upon a particular gland the others will work in har-
mony with it. Every gland is probably necessary for the
perfect activity of the rest, and the harmony between the
glands is demonstrated by physiological experiments. . . .
When it is found that the removal of an organ constantly
induces either atrophy or hypertrophy of some other organ,
we can reasonably deduce that in the first case the organ
removed is essential to the welfare of the one that atrophies
in its absence, and in the second case that the hypertrophying
organ is endeavoring to replace the lost one, in some de-
gree, and that therefore the two organs have a kindred
function."16

As an example of the first case, the fact is cited that the
removal of the thyroid or of the anterior part of the pitui-
tary induces the atrophy of the testicles or the ovaries. The
second case is illustrated by the hypertrophy of the supra-
renal from the . removal of the thyroids, and also by the
hypertrophy of either the thyroid or the pituitary on re-
moval of the other. "The demonstrated facts of hyper-
trophy," we read, "clearly point to an entente or even a
triple alliance between thyroid, hypophysis and suprarenals.
And the genital system is absolutely dependent upon the
integrity of these three." 17

Stating now, in a single paragraph, the results of inves-
tigations in this field, we have: The different parts and ac-
tivities of the organism are maintained in their normal state,

128 The Unity of the Organism

both as to the essential nature of each, considered individ-
ually, and as to their relation with one another, by a number
of exceedingly powerful and subtle chemical substances (in-
ternal secretions) which are produced by certain parts, are
passed into the blood, and by it are carried about over the
whole organism to exert their appropriate influences on
other parts and functions. Because of the peculiar way these
substances do their work, they have been called chemical
messengers, or to have a distinctive name, hormones.

Relation Between the Internal Secretory and Nervous
Systems

But no physiological truth is better known than that one
of the main offices of the nervous system is to correlate the
organs and parts of the body with one another. It is but
natural to suppose, therefore, that if there is a chemical
scheme for accomplishing the same end, the two are in some
way closely related.

That the relations which exist between the cerebro-spinal
nervous system, the autonomic nervous system (including the
sympathetic), and the internal secretions, constitute one of
the most important subjects in the whole physiological do-
main, at the same time that it is one of the most recondite
and difficult to investigate, has come gradually to view
through the work of the last few decades. We will try to
extract enough from the mass that has been written on the
subject, to illustrate the principles involved. The modern
period of knowledge of what was formerly but rather in-
definitely included under the term sympathetic nervous sys-
tem, has revealed that we have to do with a portion of the
general nervous mechanism which in reality is a subdivision
of a larger category.

Chemico-Functional Integration 129

•
Composition and Nature of the Autonomic System

The name autonomic was given to this category by Lang-
ley. "The autonomic nervous system," he says, "means the
nervous system of the glands and of the involuntary muscles ;
it governs the 'organic' functions of the body"; and further:
"The word implies a certain degree of independent action,
but exercised under the control of a higher power." 18 Bay-
liss adds : "It is necessary to be quite clear that the au-
tonomic system includes the sympathetic, since some writers
abroad use the name as applying to all the visceral nervous
system other than the sympathetic, speaking of sympathetic
and autonomic." 19

Perhaps the most important single fact which differen-
tiates the autonomic from the cerebro-spinal system is the
intercalation, everywhere in the autonomic system, of an
extra neurone between the cerebro-spinal nerve and the part
innervated. Cannon states this distinction very clearly:
"The skeletal muscles receive their nerve supply direct from
the central nervous system, i. e., the nerve fibers distributed
to these muscles are parts of the neurones whose cell-bodies
lie within the brain or spinal cord. The glands and smooth
muscles of the viscera, on the contrary, are, so far as is now
known, never innervated directly from the central nervous
system. The neurones reaching out from the brain or spinal
cord never come into immediate relation with the gland or
smooth muscle cells ; there are always interposed between the
cerebrospinal neurones and the visceral extra neurones
whose bodies lie wholly outside the central nervous
system." 20

Cannon's suggestion that these interposed neurones may
function as transformers for impulses received from the cere-
brospinal system should be noted here.

Three sharp subdivisions of the autonomic nervous sys-
tem are recognizable. One is known as the vagal or cranial

130 The Unity of the Organism

autonomic, because it is largely made up of fibers from the
vagus, or tenth pair of cranial nerves. Another is the sym-
pathetic, or better, the thoracico-lumbar autonomic, be-
cause its fibers originate from the great visceral sympathetic
ganglia. This is by far the most extensive of the three sub-
divisions, and is the only one that is distributed to all parts
of the body. The third is the sacral autonomic. As its name
implies, it is quite restricted in extent, its fibers being dis-
tributed to the extreme distal end of the intestine, the urin-
ary bladder, and some of the external genital organs. But
the differences between the three which are most important
for us are physiological, a particularly important difference
being that the thoracico-lumbar division acts antagonistic-
ally to both the end divisions. Stimulation of the fibers of
.the sympathetic has just the opposite effect to the same
stimulus applied to the fibers of the others. "The sympa-
thetic fibers check, the vagal autonomic fibers excite, the
movements of the intestines ; the sympathetic dilates, the
vagal autonomic contracts, the pupil; the sympathetic
hastens, the vagal autonomic slows, the heart." 21 The
sacral contracts the lower part of the large intestine and re-
laxes the outlet of the bladder, while the sympathetic relaxes
the same part of the intestine and contracts the same part
of the bladder. Cannon states the general principle thus :
"When the mid-part meets either end part in any viscus
their effects are antagonistic" 22

While the incompleteness of knowledge in this field needs
emphasizing, yet that knowledge is sufficient to put some of
the main features beyond question, and to make clear the
great importance of the subject and of fuller knowledge on
it. Touching these general aspects Professor L. F. Barker
writes : "While we do not yet understand the exact mechan-
isms of association among the activities of the cerebrum,
the endocrine glands, and the reciprocally antagonistic au-
tonomic domains and their end-organs, we can begin to see

Chemico-Functional Integration 131

the paths which must be followed in order that more exact
knowledge may be gained."

Experimental Evidence of Connection Between the Adrenal
Glands and the Nervous System

Some of the most important information we have in this
field is furnished by Cannon and his collaborators concern-
ing the secretion of the adrenals and its relation to the
autonomic and central nervous systems. It had been proved
before Cannon began his investigations that adrenin injected
into the blood has exactly the same effect on certain parts
of the organism as does the sympathetic autonomic nerves
with which the same parts are supplied, and that the effect
of the secretion is direct and not through the nerves. In
other words, it had been proved that the organism has two
methods by which the same activity of certain of its parts
can be induced, one nervous, the other chemical. Thus the
dilation of the pupils, the erection of hairs, the inhibition
of activities of the alimentary canal, and the liberation of
sugar from the liver can be induced either through sym-
pathetic autonomic centers or by the secretion of the adrenal
bodies. This in itself was important evidence of interrela-
tion between the nervous system and internal secretory sys-
tem. But the experimental researches prosecuted in Can-
non's laboratory have proved that a connection exists be-
tween the autonomic-adrenal phenomena and the cerebro-
spinal system through the sensory nerves, and with the
psychic life of the animal ; and have shown the probable sig-
nificance of the entire scheme for the life of the organism as
a whole.

To be a little more specific, they have proved:
(1) That strong excitation of sensory nerves stimulates
reflexly the adrenal glands and causes them to pour an
increased amount of adrenin into the blood.

132 The Unity of the Organism

(2) That emotional excitement (as the fright of a cat by
a dog) similarly increases the flow of adrenin.

(3) That this increase of adrenin in the blood may in-
crease the liberation of sugar from the liver into the blood
to such an extent as to make sugar appear in the urine, thus
demonstrating a true "emotional glycosuria."

(4) That the increased adrenin of the blood thus pro-
duced is probably advantageous to the organism in that it
enhances its ability to meet special stresses that naturally
accompany special excitement, as of fear, anger, or pain,
this advantage consisting partly in augmentation of the
working energy of the muscles, probably through the sugar
delivered to them, and in increasing the coagulability of the
blood, thereby reducing the danger from bleeding wounds.

Summing up his conclusions as to utility, Cannon writes :
"These changes in the body are, each one of them, directly
serviceable in making the organism more efficient in the
struggle which fear or rage or pain may involve; for fear
and rage are organic preparations for action, and pain is
the most powerful known stimulus to supreme exertion. The
organism which with the aid of increased adrenal secretion
can best muster its energies, can best call forth sugar to
supply the laboring muscles, can best lessen fatigue, and can
best send blood to the parts essential in the run or the fight
for life, is most likely to survive." 24

But fear and rage are, one hardly need be reminded, in
part psychic phenomena, and hence inseparably connected
with the higher centers of the cerebrospinal nervous system.
Though in the main reflex and automatic, they are neverthe-
less to some extent subject in man to intelligent control.
Thus the way is open for a measure of rational understand-
ing of the structural-functional means by which human be-
ings "tap," as William James would say, and bring under
direction those remarkable "reservoirs" of ordinarily unused
energy about which everybody knows something from his

Chemico-Functional Integration 133

own experience, and upon which nobody has written more
intelligently than James.

An excellent beginning has been made, then, in the ex-
perimental demonstration of the integration of the endo-
crinal and common glandular systems, the blood circulatory
system, the autonomic and cerebrospinal nervous systems,
and the emotional-psychic life of animals.

Clinical Evidence of Adrenal-Nervous Connection

But important knowledge and general views in this field
are also coming from clinical medicine, and pharmacology.
A general presentation of the results reached down to
1913 is contained in Innere Sekretion und Nervensystem,
by H. Eppinger and others. A particularly significant body
of evidence coming from this source concerns the relation
between the sympathetic or middle autonomic nervous ap-
paratus and the two end autonomic systems, the cranial and
sacral. These two groups act, it will be recalled, antagon-
istically to each other. Eppinger and others have shown
that the thoracico-lumbar, or sympathetic, and cranial, or
vagal systems differ in susceptibility to stimuli in different
individuals, and perhaps in the same individual at different
times, thus making the two groups what is called sympa-
theticotonic and vagotonic with reference to each other, de-
pending on whether the sympathetic or the vagal is the more
susceptible to stimuli. This difference can be demonstrated
by the administration of various drugs, as adrenin and pilo-
carpin. But it is known, according to Eppinger, that the
thyroid toxin stimulates both the sympathetic and the vagal.
From this it results that over-stimulus of either may occur
through this source, and go to the extent of producing the
characteristic symptoms of Basedow's or Graves's disease
(rapid heart beat, exophthalmia, diarrhea, etc.). These
symptoms may occur in varying degree, depending on

134 The Unity of the Organism

whether the patient is sympathetico- or vagotonic, the sym-
patheticotonic type of the disease being characterized by
marked protrusion of the eyeballs, especially rapid heart
beat, absence of sweats, diarrhea, and disturbance of the
respiration; while the vagotonic type is characterized by
slight protrusion of the eyes and increase of heart action,
by outbreaks of sweat, diarrhea, and by faultiness in the
respiratory rhythm.

While some observers, like Falta, do not believe the facts
now known can be definitely classed in this manner, the ef-
fort, justified by some positive knowledge, has at least the
merit of specifying to some extent the intricate reciprocal
action between the thyroid apparatus and the nervous sys-
tem, and also between the different portions of the auto-
nomic system; and to this extent all students of the subject
bear witness. Thus Falta : "In my opinion everything speaks
for the fact that in Basedow's disease the entire nervous
system is in a condition of over-excitement and that the
pictures presented by the vegetative nervous system are as
uncommonly manifold and always changing."

The indication of prime importance in this is that in these
antagonistic divisions of the autonomic nervous and endo-
crinal glandular systems, operating together with the other
portions of the organism, there is a balancing-off or equil-
ibrating apparatus through which the whole complex of vege-
tative functions is carried on, all of which in turn are con-
nected with the psychic functions. Probably no better il-
lustration can be found of the conception of the organism as
fundamentally dynamic. According to this conception nor-
mality, both in function and in structure, consists not in
rigid, invariable activities and organs, but in a ceaseless play
of constitutively antagonistic forces and structures. By this
conception the whole life of the organism, physical and psy-
chical, may be crudely likened to the performance of the
tight-rope walker, which depends on numberless balancing

Chemico-Functional Integration 135

activities. Let the performer be really motionless in every
part for one instant, and he falls.

The treatment of tetany and its relation to internal se-
cretions, especially to that of the thyroid, by Eduard Phleps
in the work now under consideration, is another excellent il-
lustration of how interpretation may run in accordance with
this conception of the animal organism. The essence of the
section, as touching this question, is contained in the fol-
lowing:

"On the ground of clinical symptoms authoritative clin-
icians like Eulenburg, Kahler and Nothnagel explained tet-
anus as a disease of the entire nervous system. Later it was
proved that this disease was not due to primary organic
changes of the nervous system, but to secondary functional
disorders. . . . The view of those authors, who refer the
disease to the simple effect of a substance of the epithelial
granules acting normally and continuously on the whole
nervous system, finds here a further development of that old
theory because for them the clinical picture of a regular
grouping of nervous stimulus- and response-phenomena
arises from an impairment of the close functional relation
between the nervous system and the epithelial granules ( Mac-
Callum, Chvostek jun., Biedl, Eppinger, Falta, Rudinger,
Jonas, et al.). In agreement with these authors we con-
ceive the action of the epithelio-secretive substance as that of
a hormone in the sense of Starling and Bayliss, which must
have its essential point of attack on certain reflex stations
of the central nervous system." 26

Taking cognizance, now, of the fact that most if not all
the cells known to produce internal secretions arise embry-
onically from epithelium, as does also almost all nervous
tissue, we have the suggestion of a deep-seated combination
scheme, chemical-and-nervous, for integrating the organism.
The best investigated example of what is here referred to is
the suprarenal bodies. It is now fully established that the

136 The Unity of the Organism

inner or medullary portion of the organ, the part which pro-
duces the adrenin, is developed from the same neuroblastic
mass out of which the sympathetic autonomic ganglia arise.
Furthermore, it seems beyond question that the so-called
chromophil, or sometimes the adrenin granules in the chief
cells of the functioning gland play a fundamental role. Com-
bining these facts with the equally well ascertained facts
that the cortex of the gland is derived from the same em-
bryonic mesoblastic mass which gives rise to the genital
glands ; and that in adults cortical changes in the supra-
renals are intimately correlated with reproductive changes,
and a general view of the factual basis on which the sugges-
tion rests is before us. We may, I think, regard the sugges-
tion not only as justified but as revolutionary in comparison
with any theory that was scientifically justifiable until re-
cently. The following further quotation from Phleps brings
the idea into still clearer view :

"The unqualified dependence of the nervous system on the
epithelial bodies (this last used in the sense of general physi-
ological considerations, and only by way of illustration),
and the absolutely vital significance of this enables us to see
many things in a new light. We learn that many motor,
sensory and vasomotor-trophic functions of the central nerv-
ous system even up to the highest reflex stations having the
most complicated cortical functions, are in constant func-
tional cooperation with organs which heretofore have not re-
ceived sufficient consideration from this standpoint. The
results compel a change of our views concerning the con-
stantly dominating position of the nervous system. We may
see in many of its activities only the most manifold inter-
mediary roles between glandular functions which are in the
closest relation to metabolism, and the sum total of all re-
actions which follow external stimuli." 27

Chemico-Functional Integration 137

Summary of Present State of Knowledge In This Field

Speaking generally, we may say that the trend of all re-
sults, experimental and clinical, is unquestionably toward a
demonstration of the closest interaction between the entire
internal secretory system and both the autonomic and cere-
brospinal nervous systems, this interaction affecting the
whole of both the growth and the functioning of the animal
organism.

But we must remind ourselves again how fragmentary
knowledge is in this great realm. Unanswered questions meet
one on the very threshold of any portion he enters. By
way of illustration, take the phenomenon of abnormal growth
known as acromegaly. This malady is characterized, as the
name indicates, by a "peculiar non-congenital hypertrophy
of the upper and lower extremities and of the head." 5 Such,
according to Schafer, is the definition given by Pierre Marie,
who first fully described the disease. The main visible symp-
toms consist in the enlargement of the bones of the head,
hands, feet, chest, etc., especially in their terminal portions.
Through such growth the nose and lower jaw, especially the
chin, become protrudent. But the whole skeleton is more or
less affected, and there is a corresponding over-development
of the muscles, the affected person becoming abnormally
strong. That acromegaly is constantly associated with an
abnormal condition of the hypophysis is recognized by ap-
parently all authorities. Whether the abnormality of the
gland is a cause or only an accompaniment of the disease is
an open question in the minds of some. However, the view
of a large majority is that such a causal relation does exist.
"That the acromegalic skeletal growth," says Schafer, "is
produced by hypertrophy and oversecretion (or perverted
secretiooi) of the anterior lobe is highly probable, both as the
result of partial extirpation in animals and from the effect of
operative removal of the pituitary tumours in man." 29 The

138 The Unity of the Organism

particular issue here, it will be noticed, is the vitally im-
portant one of what might be called the functional as
distinguished from the hereditary cause or at least incitement
of growth.

In illustration of the importance of understanding the
unification among these complex systems, the manifestation
of which is in turn dependent upon the organism as a whole,
the following from the address by L. F. Barker already re-
ferred to, is impressive.

"In how far these sudden and violent excitations of the
autonomic nervous system which accompany strong emo-
tions are due to the intervention of the glands of internal
secretion, and in how far they depend upon direct neural
conduction from the brain, we are as yet but ill-informed. I
need only remind you of the vasodilation of the face in the
blush of shame, of the palpitation of the heart in joy, of the
stimulation of the sudoriparous glands which precedes the
sweat of anxiety, of the stimulation of the vasoconstrictors,
the pupil dilators and the pilomotors in the pallor, mydriasis
and goose-skin of fright, to illustrate some of these violent
autonomic excitations." 30

The references here, it will be noted, are primarily mani-
festations pertaining to the surface, the integumentary
parts of the body, and their scope is what especially inter-
ests us. Very nearly the whole list of these parts is in-
volved, and probably a complete inventory would be still
more inclusive.

Now notice the range of manifestations at a deeper level
that are involved. "The balance maintained normally be-
tween the two antagonistic systems the vagal and sympa-
thetic autonomic is one of the most interesting of physio-
logical phenomena. Think, for example, of the rate of the
heart-beat — how constantly it is maintained at a given level
in each individual when the body is at rest ; the impulses ar-
riving through the vagal system just balance those arriving

Chemico-Functional Integration 139

through the sympathetic system, so as to maintain a rate of
approximately seventy-two beats per minute. And a similar
balance is maintained in other autonomic domains (e.g., the
pupils, bronchial musculature, gastric glands, gastro-intes-
tinal muscle, sweat glands, bladder muscles, etc.)."

And Barker then calls attention to the extent to which the
normal processes of the body depend upon temporary upsets
of these equilibria, examples of which are watering of the
mouth at the smell or sight of food which appeals to the
appetite through these senses, the flow of gastric and pan-
creatic juices at the proper time, through indirect stimula-
tion ; the sudden relaxation of the sphincter and contraction
of the detrusor of the bladder in micturition; the violent
contractions of all the muscles concerned in parturition in
the female, and so on.

We may summarize the results of this chapter thus:

(1) The researches of recent years on the internal secre-
tory system and its connection with the great subdivisions
of the nervous system, and with the blood, muscular, and
reproductive systems, have laid a solid foundation for an un-
derstanding of the chemico-functional basis of the animal
organism's unity.

(2) The emotional phase of the psychic life of the ani-
mal is proved to be in direct organic connection with this
basis.

From these results there naturally springs the important
question: What relation has human consciousness to this
same basis? An attempt to answer this question will be an
unavoidable part of our treatment later of the psychic in-
tegration of the organism.

REFERENCE INDEX

1. Luciani II, 13 5. Bayliss 707

2. Carlson and Woelfel 49 6. Bayliss and Starling 325

3. Schafer (1914) 18 7. Bayliss 706

4. Hertoghe 194 8. Schafer 28

140

The Unity of the Organism

9. Luciani II, 22

10. Luciani II, 35

11. Schafer 9

12. Schafer 11

13. Aldrich 457

14. Bayliss 721

15. Waller 277

16. Waller 280

17. Waller 281

18. Langley 240

19. Bayliss 484

20. Cannon, W. B.('16) .... 22

21. Garrison 144

22. Cannon, W. B. ('16) ... 34

23. Garrison 146

24. Cannon, W. B. ('13) ... 372

25. Falta 59

26. Eppinger, et al 214

27. Eppinger, et al 217

28. Schafer 106

29. Schafer 69

30. Garrison 146

Chapter XIX

THE ORGANISMAL SIGNIFICANCE OF THE
INTERNAL SECRETORY SYSTEM

TT remains now to consider the real purpose for which
•*• the presentation of facts and views on internal secre-
tions has been made, namely that of showing critically the
significance of this secretory system for the organismal
conception.

General Inability of Element alism to Interpret the
Phenomena,

There is perhaps no better way of approaching this part
of our task than by noticing the inability of elementalistic
biology to deal in a really intelligent and consistent manner
with the phenomena in this field. The breakdown of bio-ele-
mentalism when confronted with the phenomena of "chemical
messengers" nowhere finds more cogent illustration than in
the effort to identify internal secretions with the organ-
forming substances hypothesized by Sachs and others.

Although in what follows the exposure of inconsistency
and fallacy will have to be drastic and may seem personal,
the truth is it is wholly impersonal in spirit and is directed
at a system of bad reasoning born of what might be called
a juvenile metaphysics of the living world.

The objective achievements of Jacques Loeb and others
of the school he represents, in experimental biology, merit
the admiration of all lovers of observational truth. One
may be, too, more tolerant of their faults as reasoners

141

The Unity of the Organism

than he could be but for his recognition of their high service
against all the traditional forms of supernaturalism. The
real case against the school is, as I see it, two-fold. First, in
their zeal to substitute naturalism for supernaturalism they
fail to notice that supernaturalism is in its very essence
finalistic, and they are led to imagine they have attained,
or can attain, natural explanations that fully supplant the
old supernatural explanations. This results in the con-
version of their supposed naturalism into something which
is essentially another kind of supernaturalism. The second
part of the case against the school is its abuse of the most
common principles of the knowledge-getting processes in ob-
jective biology. For the general good of the biological
sciences the urgent need of reformation touching both as-
pects of the case has led me to examine the particular in-
stance of Loeb's treatment of internal secretions at greater
length than would otherwise be justifiable.

Critique of the View That Internal Secretions are "Formative

Stuffs"

Although Loeb is the only author, so far as I know, who
has expressly contended for the identity of internal secre-
tions with Sachs' organ-forming substances, the assumption
is so accordant with the spirit of elementalism, and Loeb is
so typical and eminent a protagonist of this philosophy,
that his proposal will probably find many adherents. It is
consequently desirable to see what there is in the effort to
bring hormones into such a historical setting.

The statement of Loeb's views is contained in "The Or-
ganism as a Whole from a Physico-Chemical Viewpoint,"
1916. Referring to his espousal twenty-five years ago of
Sachs' hypothesis to explain heteromorphosis, he writes : "At
that time the idea of the existence of such organ-forming
substances was received with some scepticism, but since then

Significance of the Internal Secretory System 143

so many proofs for their existence have been obtained that
the idea is no longer questioned. Such substances are known
now under the name of 'internal secretions' or 'hormones';
their connection with the theory of Sachs was forgotten with
the introduction of the new nomenclature." l

A case to which Loeb makes special reference as proof
that internal secretions are the same as Sachs' organ-form-
ing substances is that of the effect of thyroid on the meta-
morphosis of the tadpoles of frogs and toads, demonstrated
by Gudernatsch. The author's mode of using this case in
illustration of his contention is highly instructive. He re-
fers to the legless condition of the tadpoles and calls atten-
tion to so much of Gudernatsch's results as pertain to these
members. Gudernatsch found, Loeb points out, that where-
as in the usual course of things the tadpoles live from four
months to a year before the legs grow out, by feeding them
on thyroid gland these members can be induced to appear
at any time. "We must, therefore, draw the conclusion,"
Loeb says, "that the normal outgrowth of legs in a tadpole
is due to the presence in the body of substances similar to
the thyroid in their action (it may possibly be thyroid sub-
stance) which are either formed in the body or taken up in
the food." When the case is presented in this way and
nothing more said about it, it certainly looks considerably
as if the thyroid substance or something like it is leg-form-
ing substance; and such an interpretation would be enticing
were it really true, as Loeb says, that no other substance
seems to have such an effect.

When, however, one consults the account given by Guder-
natsch himself as to what his experiments were and what
they established, the whole matter stands in a quite different
light. First of all, the fact that in these experiments mam-
malian thyroid was largely used as food for the tadpoles, thus
bringing it to pass that if thyroid substance was specific
organ-forming substance, then frogs' legs were produced by

144 The Unity of the Organism

mammalian substance, shows at once that there is something
badly askew in the theory. So we are incited to examine
the facts, and particularly the reasonings, carefully.

The following is taken from one of Gudernatsch's papers :
"The most striking and at the same time unquestionable re-
sults were attained by thyroid feeding. . . . The influence
of the thyroid food was such that it stopped any further
growth but on the contrary led to an abnormal diminution
of the size in the animals treated, while simultaneously it ac-
celerated the differentiation of the body immensely and
brought it to a premature end." 3 In other words, the effect
of thyroid food was to stop the increase in size of the frog's
larva and start, almost at once, its transformation into the
adult. Now this transformation does not consist merely in
the production of legs, but in a whole series of changes,
some of which, like leg transformation, are progressive,
while others are regressive. For example, one of the re-
gressive changes to which Gudernatsch gives particular
attention is the resorption of the tail. "Reduction of the
body mass (resorption of the tail, loss of water, therefore
an increasing compactness of the body, etc.)"4 more than
in normal development, the author says, goes hand in hand
with the progressive changes. That is, when the entire series
of results of thyroid feeding are considered, and not merely
one result picked out arbitrarily, then in case we choose to
say the thyroid substance is organ-forming as regards legs,
we should have to say it is organ-destroying as regards
tail. Furthermore, if we call the thyroid substance organ-
forming, consistency would compel us to recognize that one
and the same substance is not only formative of legs but
of numerous other organs and parts, as of the skin, mouth,
respiratory and blood systems, all of which undergo, as is
well known, progressive changes during metamorphosis. Nor
is this complex series of morphological changes the while
story. Striking and characteristic changes in the habits

Significance of the Internal Secretory System 145

of the tadpoles resulted. "Towards the end of metamor-
phosis the animals hardly moved about in the water. They
were always lying quietly, generally on their backs. When
disturbed they would move for a few seconds in a somewhat
convulsive manner and then drop again to the bottom of
the dish, while tadpoles fed on other material would swim
about for a long time." 5 So mammalian thyroid substance
is not only organ-forming for a whole series of frog organs
but it is habit-forming for a variety of frog habits !

But we must not let the ludicrousness of this veer us
away from the reasoning in the case. Taking the facts
actually brought out by Gudernatsch, what becomes of the
specificity of the substance, which according to Loeb's state-
ments was what Sachs hypothesized? "Sachs suggested that
there must be in each organism as many specific organ-form-
ing substances as there are organs in the body." 6 The truth
appears to be that thyroid substance in this case is organ-
forming in much the same sense that water is organ-forming
for the leaves, flowers, and fruit in a squash vine, which could
not develop without water. Indeed the analogy suggested
goes further than appears at first sight. As everybody
knows, the effect on young plants of a scant water supply
is to stunt the plant as to size and to hasten its blossoming
and fruiting. That is, an under-supply of water has an
effect on immature plants similar to that of an over-supply
of thyroid substance on immature frogs, namely that of re-
tarding growth and hastening metamorphosis. The total
effect in each case is systemic. In other words, the real
significance of the instance used by Loeb is just the opposite
of his interpretation of it. Thyroid substance is organ-
formmg only through being organism-transforming. Full
justification of this way of interpreting the part in
development played by thyroid substance is furnished by the
recent studies of B. M. Allen. This investigator has, like
Gudernatsch, experimented with frog larvae. He has, how-

146 The Unity of the Organism

ever, supplemented Gudernatsch's work by depriving the
larvae of thyroid altogether instead of giving them an extra
allowance through feeding. Allen extirpates the entire em-
bryonic thyroid from tadpoles long before any indications of
metamorphosis appear. What he finds of special impor-
tance for the present discussion is contained in the follow-
ing:

"While Gudernatsch showed that thyroid feeding acceler-
ates development, this work shows that the total absence of
the th}7roid gland produces complete cessation of somatic
differentiation at a certain stage but does not hinder con-
tinued growth in size." 7

O

The first part of this statement taken alone might be
looked upon as confirmatory of the "formative stuff" theory
of thyroid substance. But the phrase "at a certain stage"
implies, as Allen's paper as a whole conclusively shows, that
the exact opposite of such a conception is alone tenable.
What actually happens, Allen brings to light, is that in
spite of the complete absence of thyroid substance trans-
formation of the larva is begun but is not carried through.
That is, the organs and parts of the adult frog are laid
down but (with the exception of the reproductive glands)
are not completed. "It is evident," Allen says, "that the
thyroid gland is in no wise essential to the earlier phases of
development, but that at a certain definite stage, further
development of the soma is dependent upon it." 7 It should
be mentioned that Allen makes a rather special point of the
accordance of his results with those of Gudernatsch.

If the greatly hastened and modified metamorphosis of the
frog tadpole observed by Gudernatsch is inducible by no
other means than by thyroid substance (which while pos-
sible is not at all certain), then is the substance causal in
the important sense of being not only competent but in-
dispensable. But even so it falls far short of being a com-
plete causal explanation of the phenomena under contempla-

Significance of the Internal Secretory System 147

tion, as Allen's results directly prove.

A defect in Loeb's reasoning is his ignoring the truth that
thyroid substance lacks in the case cited the prime attribute
of a sufficient cause, namely full competency. The actual
substance which enters into the new legs, and as far as that
goes, into the other new parts, is probably provided very
little, if at all, by the specific substance or hormone of the
thyroid. The causal role played is the relatively humble one
of excitor or stimulator to an activity not essentially new
but only exceptional as to time. Of course Loeb does not
need to be told that hormones incite growth rather than
provide the substance itself out of which the organs and
parts are built. Indeed portions of his account of this very
case show positively that he is aware of this. We read :
"Thus we see that the mesenchyme cells giving rise to legs
may lie dormant for months or a year but will grow out
when a certain type of substances, e.g., thyroid, circulates in
the blood. There may exist an analogy between the activ-
ating effect of the thyroid substance and the activating
effect of the spermatozoon or butyric acid (or other par-
thenogenetic agencies) upon the egg."8

This suggestion of analogy between the action of thyroid
secretion and "butyric acid (or other parthenogenetic agen-
cies)" is well taken. The resemblance between the two
agencies, as judged by their effects on development, is cer-
tainly rather close. Very well; would Loeb, then, call bu-
tyric acid organ-forming substance, and identify it with the
"formative stuff" of Sachs ? Certainly any substance which
will rouse the latent developmental capacities of an egg into
activity is in a minor sense formative, especially if these
capacities are wholly unable to start without some such
agency. But since butyric acid, or some one of the other
dozens or scores of parthenogenetic agencies, may activate
the eggs of many, many species of animals ; and since the
eggs of many, many species may be activated by any one of

148 The Unity of the Organism

scores of such agencies, what about the specificity of the
formative substances which Loeb himself expressly says was
part of Sachs' conceptions?

Again, how reconcile the contention that thyroid sub-
stance or something like it is organ-forming for the legs
of a tadpole with the statement quoted a few paragraphs
back about mesenchyme cells which give rise to legs?

The Metaphysical and Logical Weakness of the View

Obviously Loeb's treatment of this subject contains irre-
conciliable contradictions. Is it then worthless? My answer
is, no, not by any means. But how comes it that a scien-
tist of his great experience and merited distinction can run
into such self-destroying speculations and statements, seem-
ingly without rational discomfort to either himself or others
of the school which he represents?

The answer takes us back to some of the most funda-
mental issues between the elementalist and organismal stand-
points, and though not requiring us to palliate in any degree
such offenses against scientific reasoning, it partly explains
how tolerance for such offenses is begotten, and discovers
a nucleus of genuine merit in Loeb's position. Bringing the
matter to as basal a statement as possible, what we find is
that this whole book on the Organism as a Whole is written
on the theory that the only alternative to the assumption
of supernaturalism is materialism. Instead of supernatural
forces of some sort (Platonic Ideas, entelechies, psychoids,
"supergenes") for explaining the organism when regarded
as alive and whole, his assumption is that material elements
as known to us in inorganic nature are the sufficient causal
explanation of organic phenomena.

Were this theory correct — were it true that the "vital
principle" must either be conceived as supernatural or that
the inorganic elements taken by themselves are competent

Significance of the Internal Secretory System 149

to produce organisms, then it would be impossible for biol-
ogy to do much better in its reasoning and general attitude
than Loeb and other elementalists do when they undertake
to construct a philosophy of organisms. I agree whole-
heartedly that all supernaturalism, no matter what nomen-
clatorial garb it takes on, must be repudiated by the sciences
of organic beings. Ideas, or psychoids, or entelechies, or
"principles" of any kind conceived as independent of, or
even separable from, sensible objects are quite as repugnant
to me, an organismalist, as they are to any elementalist. The
essence of my contention is that the natural substitute for
these imponderable things are the living, individual organ^
isms themselves, and not the particles of which they are com-
posed. Each and every individual organism is a natural
reality by exactly the same criteria that the atoms, mole-
cules, cells and tissues of which it is composed are natural
realities. And since each individual is to some extent differ-
ent from every other, and maintains its individuality in full
possession of these differences, by its power of transforming
foreign substance into its own substance, it is ultimate both
as to structure and as to causal power in as deep and literal
a sense as the material particles of which it is composed are
ultimate.

Loeb's considerable attention to the views of Claude Ber-
nard is fortunate for us, since it affords a chance to show
from still another angle the inevitable breakdown of ele-
mentalist reasoning when it is brought face to 'face with or-
ganic phenomena as actual nature presents them to the
modern student. Loeb calls attention, properly, to the fact
that one of the things on which Bernard placed special em-
phasis, as Bichat before him had done, is the organizing syn-
theses which go on in the living being. The real advance, it
seems to me, which Bernard made over any of his predeces-
sors, was the positiveness of his rejection of a vital force
as something supernatural — as something, using his own

150 The Unity of the Organism

words, "under the government of a special principle, a pecu-
liar power what name soever be given it, whether soul, or
archeon, or psyche, or plastic intermediary, or guiding
spirit, or vital force, or vital properties";9 and his rejec-
tion of these conceptions because of his recognizing that the
unitariness of the organism removes the necessity for as-
suming any such extraneous principle. "When we say,"
Bernard writes, "that life is a guiding idea, or the evolu-
tive force of the being, we merely express the thought of a
unity in the succession of all the morphological and chemical
changes effected by the germ, from the beginning to the
end of life. Our mind grasps that unity as a conception it
cannot escape. . . ." 10

Up to this point the position held by Bernard is entirely
satisfactory for the organismal conception as I am trying
to develop it, and is, according to my view, unassailable.
But the unquoted part of the last sentence contains a state-
ment which reveals Bernard on a by-road leading away from
the promised land toward which he was headed as long as he
was speaking in terms of biology proper. The rest of the
sentence follows : "and explains it as 'a force' ; but the mis-
take is in supposing that this metaphysical force acts after
the manner of a physiological force." 10 Stated in a nut-
shell, the by-road which Bernard is entering here is that of a
kind of separatedness, but inevitable concomitance, or paral-
lelism between the phenomenal and neumonal worlds which,
according to the views upheld in this volume, does not exist.
"We need here," Bernard says, "to draw a distinction be-
tween the metaphysical world and the phenomenal physical
world, which serves as its basis, but which can borrow noth-
ing from it." This Leibnitzian theory, according to which
"everything takes place in the soul as though there were no
body, and in the body everything takes place as though
there were no soul," Bernard says science "recognizes and
adopts in our day." But in our day, this year nineteen

Significance of the Internal Secretory System 151

hundred and eighteen, science, at least so much of it as
speaks through this volume, though understanding fully
what this dualist theory is, rejects it. It denies that things
take place in the soul as though there were no body ; and
that things take place in the body as though there were no
soul. On the contrary it affirms that what in rather un-
critical language we call "the body" and "the soul" are in
the most intimate and indissoluble connection with and de-
pendence upon each other, and together constitute "the
Organism."

The Form of Metaphysical Absolutism Involved

A full, systematic justification of this position is beyond
the province of this volume; and in this chapter we are con-
cerned solely with the "body," the strictly morphological
and physiological aspect of the subject. Nevertheless, this
much of contact with the "soul" aspect was unavoidable for
the reason that Bernard and also Loeb have run into it in
such fashion as to color deeply their discussions and out-
look. This coloring is the more unfortunate and the more
insistent in requiring attention from the fact that the au-
thors, especially Loeb, are apparently unaware of such
coloring. For example, Loeb writes on the first page of his
book, after saying that the atomistic theory of matter and
electricity are now in all probability on a "permanent
basis" : "This permits us to state as an ultimate aim of the
physical sciences the visualization of all phenomena in terms
of groupings and displacements of ultimate, particles, and
since there is no discontinuity between the matter consti-
tuting the living and non-living world the goal of biology
can be expressed in the same way."

Statements like this, many of which can easily be quoted
from Loeb's writings, leave no question about his meta-
physical affinities. The conception of "ultimate particles"

152 The Unity of the Organism

as explanation of all phenomena, is exactly what I mean by
elementalist absolutism.

Confusion of Theory of Organisms and Theory of the
Knowledge of Organisms

Although the thoroughgoing metaphysical character of
these statements is evidenced by the finalism which crops
out at several points, this is not the aspect of the matter
which chiefly interests us here. Rather what we are con-
cerned with is the fact that affirmations about the "aim of
the physical sciences" and the "goal of biology" do not
belong, properly speaking, to the provinces of physical
science and biology at all, but to quite a different science,
namely that which deals with the nature of knowledge itself.
The "physical sciences" are the vast accumulation of man's
positive knowledge, theories, hypotheses, and so forth, about
physical nature; they certainly are not physical nature it-
self. Consequently a statement of the character and aims
of that knowledge is not a statement about the phenomena
to which the knowledge pertains. And the same reasoning
applies to the affirmation about the goal of biology.

All this is only another way of saying what Loeb him-
self virtually tells us, namely that his entire discussion of
the organism as a whole is made from the standpoint of one
particular theory of the ultimate nature of living beings,
that theory being the mechanistic. Recall the complete title
of the book The Organism as a Whole From the Physico-
Chemical Viewpoint. To treat the subject from this view-
point is of course perfectly legitimate. When, however, the
assumption is made that such a treatment is the only really
legitimate one because it rests on ultimate truth, then sound
science is bound to protest, chiefly because of the obvious
biological inadequacy, and at some points, perversion and
contradiction displayed in the treatment.

Significance of the Internal Secretory System 153

This brings us back to the way internal secretions are
dealt with by Loeb. His failure to distinguish between the
two very distinct fields of theory above indicated, namely
theories about the phenomena of living beings, and theories
about knowledge of these phenomena, largely explains the
defects of the theoretical parts of his work. And we are
now in position to give our criticism greater explicitness.
Consider for example in the light of what has just been said
about confusion relative to kinds of theory, the irreconcil-
able statements, cited on an earlier page, that the mesen-
chyme cells give rise to the legs of the tadpole, and that
thyroid substance is organ-forming substance for the legs.
Stated briefly the case seems to be thus : Loeb recognizes, as
every one must, that internal secretions constitute a physico-
chemical agency for bringing about that harmonious de-
velopment and functioning so characteristic of the organ-
ism. But this harmony is one of the very things which has
seemed to some biologists inexplicable without the assump-
tion of supernatural influences of some sort ; hence Driesch's
attempt to modernize the ancient entelechy. But since in-
ternal secretions play the role that entelechies are supposed
to play, namely that of establishing and maintaining the
unity and equilibrium of the organism, the need for ente-
lechies no longer exists ; at least, this would be so for all
persons who do not contend that "ultimate explanation" is
the "goal of science."

To those who hold these absolutist beliefs as to the power
and aims of science three favorite courses are open and are
followed by different representatives of the school, depend-
ing on the taste, training and outlook of the person. One
course consists in pointing out, taking an illustrative case,
that internal secretions, being only contributing causes, do
not constitute an ultimate explanation, so that entelechies
or something similar are as necessary as before. This would
be the course followed by the vitalistic wing of the absolu-

154 The Unity of the Organism

tist school. For them internal secretions would be, in so far
as they contribute to the harmony of the organism, merely
agencies produced and used by supernatural causes.

Another course, and perhaps the one most frequented by
element alists, would be to contend that internal secretions
are sufficient as a causal explanation of organic unity to
make the entelechy or any similar notion quite superfluous,
even though these substances are far from a complete ex-
planation. The reasoning of this group of elementalists as
to this situation is substantially as follows: although in-
ternal secretions fall far short of fully explaining organic
unity and harmony, the action of these being merely that
of incitors and inhibitors, they are yet genuinely causal,
genuinely physico-chemical and so are on the road toward
complete explanation of the phenomena. All that is nec-
essary consequently, is to believe that still further advance
in the same direction will reach finally a full elementalistic
explanation ; that is, an explanation which will have no need
of either supernatural elements or the organism as a whole.
The attitude of this large class is one primarily of faith
rather than of reliance on positive knowledge; they are
inspired more by what they believe they will do in the future
than by what they actually have done. They are preemi-
nently men of promises. Although their achievements in
experimental science are indeed large, the results reached
by them are prized more on account of what they are believed
to augur for the future than for their present meaning and
worth.

Then, finally, there is the group of elementalistic absolu-
tists, of whom the author of The Organism as a Whole from
the Physico-Chemical Viewpoint is one of the most eminent
in our day, who, as we have pointed out, by confusing theo-
ries about objective phenomena with theories about the
knowledge of such phenomena, are led to affirm that such
phenomena as the unity of the organism are fully explained

Significance of the Internal Secretory System 155

by internal secretions, when as a matter of fact they are
only partly explained thus.

The surprising thing about the confusion of this group
is that wholly irreconcilable positions are held with impun-
ity, such for instance as those according to which thyroid
substance is held to be the organ-forming substance of frogs'
legs in one part of a discussion, and mesenchyme cells are
acknowledged to be of this nature in another part. The
contradiction is, to be sure, often of such character as
easily to escape the uncritical reader; but as to the au-
thors of such contradictions no other explanation seems
possible than that of wrong habits of scientific thought be-
gotten of untenable a priori conceptions. For example, a
Uasty reading of the discussion under review might lead one
to suppose that thyroid substance is not, after all, regarded
by Loeb as anything more than one contributing cause of
frogs' legs, mesenchyme cells being recognized as another
cause. Close attention to the text does not, however, war-
rant this generous interpretation of the author's position.
Going back to his espousal of the theory of Sachs and other
botanists as to organ-forming substances, we read : "Specific
shoot-producing substances are carried to the apex, while
specific root-producing substances are carried to the base of
a plant. When a piece is cut from a branch of willow the
root-forming substances must continue to flow to the basal
end of the piece, and since their further progress is blocked
there they induce the formation of roots at the basal
end." u

If this means anything it means that the shoots and roots
are actually built up by material carried about in the wil-
low branch. There is nothing in the language that can be
interpreted as meaning that shoot-and-root-forming sub-
stances are mere stimulators of some other substances which
become the actual shoots and roots. Yet it is with forma-
tive substances of this sort that Loeb in some parts of

156 The Unity of the Organism

his discussion explicitly identifies internal secretions. "At
that time the idea of the existence of such specific organ-
forming substances was received with some skepticism. . . .
Such substances are known now under the name 'internal
secretions' or 'hormones'; their connection with the theory
of Sachs was forgotten with new nomenclature." Then
follows the reference to the tadpole legs ; so that were consis-
tency really a jewel to the author, he could not escape mean-
ing that the substance of the legs was actually derived di-
rectly from the particular internal secretion in question, in
this case thyroid substance; and the later statement about
the formation of legs from dormant mesenchyme cells
through the mere activating effect of thyroid substance is by
implication contradicted.

This brings up again a matter about the interpretation
of development which we have dwelt upon in several other
connections, that of protest against rejecting the indubi-
table evidence of the senses in favor of a priori conceptions.
The crucial question in the present case is this : which is the
more fundamentally organ-forming substance for frogs' legs,
the mesenchyme cells which "though giving rise to legs may
lie dormant for months," or the thyroid substance which
may stimulate these cells into premature activity? While
Loeb's discussion does not raise this question definitely, the
implication is unescapable that thyroid substance is for him
the more fundamental. What else is the meaning of the
contention that this substance is organ-forming while no-
where do we find the mesenchyme cells so designated? Yet
the observational evidence is that the production of legs is
accomplished through the transformation of mesenchyme
and other cells which in the larva are not leg-substance, but
in the adult are leg-substance. Hence it follows that so far
as actual observation is concerned the mesenchymatous and
other larval substances are more entitled to be called organ-
forming than is the thyroid substance.

Significance of the Internal Secretory System 157

An Illustration of Neglect of Fact By Element alist Theory

These reflections lead to a still deeper level of the inherent
faultiness of elementalistic absolutism in biology, the toler-
ance which it engenders for ignoring relevant facts ; or
stated otherwise, of arbitrarily selecting from a great com-
plex of facts just those which suit the argument, and dis-
regarding all the others. For example, recall Loeb's ref-
erence to precocious leg-production in frog tadpoles as
though the effect of thyroid feeding stood alone rather than
as one among a great concatenation of effects, some con-
structive and some destructive, this complex of phenomena
constituting the metamorphosis of the young into the adult.
Loeb's use of Gudernatsch's results amounts to a positive
obscuration for the reader of what these important experi-
ments really teach. Only by the culling of facts to suit the
argument and the use of certain words and phrases, as
"influence," "responsible for," and so on, with equivocal
meanings, can these results be made to support the conten-
tion that thyroid substance is specific organ-forming sub-
stance for frogs' legs. The patent fact is that certain
mesenchyme and other cells of the larva are organ-forming
for legs, and there is no straightforward way of talking
about the causes of the transformation of a given group of
more or less undifferentiated tadpole cells into the much
enlarged and highly differentiated group called a leg, with-
out recognizing the whole organism as causal of the par-
ticular transformation. Probably no set of discoveries con-
cerning the development of the individual has ever been
made which so objectifies the means employed by the whole
in producing and correlating its constituent parts as those
on internal secretions ; and not the least significant fact is
that these substances are themselves produced by the or-
ganism.

Even were Loeb's contention valid that thyroid substance

158 The Unity of the Organism

is specific organ-forming substance, the indubitable fact that
in normal development this substance is itself a product of
the organism's activities, throws it into a very subordinate
place as a cause of development. The truth is, the main
upshot of the effort to explain ontogeny on elementalistic
principles amounts to an effort to avoid recognizing the most
positive and definite entity in the whole situation, namely
the organism taken alive, normal and untampered with.

A Peculiar Element alls t Objection to the Organic Whole

We now come to the last point to be noticed in connec-
tion with the elementalist attempt to deal with internal se-
cretions as related to the organism as a whole. Instancing
the familiar way in which a particular part of a flat-worm
will give rise to a new head after being cut away from the
original animal, when no head would have been formed at this
place had not the animal been cut, Loeb writes : "How does
the 'whole' suppress all this formative power in the part be-
fore the latter is isolated? It almost seems as if the isola-
tion itself were the emancipation of the part from the tyr-
anny of the whole. The explanation of this tyranny or of
the correlation of the parts in the whole is to be found, how-
ever, in a different influence." 12

Then follows the statement previously quoted about the
specific organ-forming substances of Sachs and other bot-
anists, and the assumed identity of these with internal se-
cretions.

Without raising the question concerning the evidence for
the assumption that the production of a flat-worm's head
as indicated is dependent upon internal secretions, let us
consider a moment the interesting conception thrown into
the treatment that the whole flat-worm tyrannizes over its
parts. Why this? Is it "mere rhetoric"? We are not per-
mitted to judge it thus, for no one has pronounced against

Significance of the Internal Secretory System 159

this sort of thing in science more frequently than Loeb. The
conception that the "whole" exercises a "tyranny" over the
parts we must accept as being seriously scientific with Loeb.
Well then, since tyranny is "absolute power arbitrarily or
unjustly administered," according to the dictionary, it is
certainly interesting to an organismalist to find so eminent
an elementalist acknowledging the organism as a whole to be
truly causal relative to its parts — for it is hardly conceiv-
able that even the extreme pliability of elementalist practice
as regards the definition of words would venture to hold the
absolute power which constitutes tyranny to be without
causal efficiency. Power thus potent but which could cause
nothing, not even the destruction of the parts (for surely
the tyranny of the whole does not destroy the parts), would
be too queer a conception for anybody to father deliber-
ately.

But the most interesting thing about this idea of the
"tyranny" of the whole over the parts remains to be no-
ticed. Tyranny is not merely absolute power; it is such
power exercised unjustly or "in a manner contrary to law
or justice."

Here we come, I think, to about the last ditch of the ele-
mentalist position. On what ground does one conceive the
power exercised by the whole organism over its parts, to be
contrary to law or justice. According to what legisla-
ture or court is there a law of the parts of an organism more
just than the law of the whole? None whatever in nature,
it must I think, be admitted. The only ground for the ele-
mentalist's pronouncement, that the whole acts tyrannously
toward the parts, that it acts in a manner "contrary to law
or justice" is in the mind of the person who makes the pro-
nouncement.

The truth is — and it is of great importance since its
influence reaches far beyond the confines of scientific tech-
nicalities— any scientist, especially any biologist, who is

160 The Unity of the Organism

through and through an elementalist, is necessarily a protes-
tant against all law except the law of elements.

The scientific elementalist is inevitably anarchistic toward
all the most common, most objective, structures and laws of
nature.

His faith is in the laws of the obscure or invisible world
and against those of the everywhere visible world. Atoms
are more real to his mind than are lands and waters, plants
and animals.

REFERENCE INDEX

1. Loeb ('16) 155 7. Allen ('18) 515

2. Loeb ('16) 156 8. Loeb ('16) 156

3. Gudernatsch 475 9. Bernard 516

4. Gudernatsch 476 10. Bernard 519

5. Gudernatsch 476 11. Loeb ('16) 154

6. Loeb ('16) 159 12. Loeb ('16) 154

Chapter XX
NEURAL INTEGRATION

Neural and Not Psychical Phenomena the Subject of This

Chapter

THE fact should be firmly fixed in mind at the outset
that in this chapter we have nothing to do with the
organism's consciousness and volition. We are to deal with
the nervous system purely on the basis of its physical activi-
ties. Whether or not consciousness or something of the es-
sential nature of it appertains to part or all of the activities
. to be treated we do not have to decide, so far as this discus-
sion is concerned. Our task here is to be strictly anatomical
and physiological, and not at all psychological.

Distinction Between Developmental and Functional
Integration

The discussion may be opened by calling attention again
to a matter noticed incidentally in the last chapter, namely
the relative parts played by internal secretions and the ner-
vous mechanism in integrating the developing organism on
the one hand, and the functioning organism on the other.
We saw that the role of internal secretions in preserving
developmental equilibrium in the individual is so conspicu-
ous as to throw the strictly physio-logical role of the sub-
stances somewhat into the background. Nevertheless as evi-
dence, particularly clinical evidence, has made clear, their
part in functional equilibrium is far-reaching.

161

162 The Unity of the Organism

We must now point out specifically what appeared only
incidentally in the previous discussion, namely, that the
integrative action of hormones and of the nervous system
are to a considerable extent the reverse of each other as
regards their relation to development and to adult func-
tion. That is, while harmones are, perhaps, more impor-
tant in development and become relatively less significant
with the advancing age of the individual, the nervous sys-
tem plays a minor part in integrating the developmental
processes but becomes of supreme importance in this way
for the functioning of the adult. Or, stating the generali-
zation in another way, integration of the organism is ac-
complished by chemical means chiefly, and by neural means
little or not at all, during early life, and by neural means
chiefly and by chemical means secondarily in later life.

One can hardly fail to see, in a general way, the bearing
of this on the familiar truth that the life of the individual
among the higher animals, man especially, is successively
vegetal, animal, emotional, and finally rational and intel-
lectual in its dominant characteristics, with the successive
stages of earlier and later childhood, youth, and earlier
and later maturity.

The Author's Indebtedness to Sherrington's Work

My dependence upon Sherrington's work in this field will
be so great as to make it impossible to acknowledge it at
every point. I therefore make at the outset the general
statement that a large part of my discussion consists of a
re-wording and rearranging of facts and ideas contained in
this physiologist's important book, The Integrative Action
of the Nervous System. But while my chief reliance here
is on Sherrington's work, the writings of Cannon and his
collaborators have been the most important source of what
I have to say about the autonomic nervous system. Can-

Neural Integration 163

non's recent book, Bodily Changes in Pain, Hunger, Fear
and Rage, has been especially drawn upon.

The Fundamentality of Cellular Integration in the Reflex

Arc

In no department of physiology do cells keep themselves
more persistently in the attention of the student than in the
physiology of the nervous system. But likewise nowhere
is the fundamental dependence of cells on other cells more
clearly seen ; for while, speaking from the standpoint of
general functions, cells may be looked upon as individual
units, when it comes to the study of nerve cells as such,
that is, as constituents of the functioning nervous system,
the individual cell is found to be no longer the basal unit.
Viewed thus the reflex-arc and not the individual cell is the
unit.

The distinction thus indicated is important from the or-
ganismal standpoint and must be considered a little more
fully. The general functions of cells to which reference i*
made above are those common to all cells, even those of the
tissues of fully differentiated multicellular animals. No
matter what tissue be under consideration, muscle, gland,
epithelial or what not, so long as it is truly living, each and
every cell assimilates, breathes, excretes, and carries on all
the metabolic processes. Thus far each cell is an indepen-
dent unit in a high degree. Now while all tissue cells, using
the term tissue in its common histological sense, have a rela-
tional or integrational function in addition to these indi-
vidual functions, it is in the nervous system that this inte-
grative aspect of cell life is most positive and definable.
The reflex-arc, as the unit of the nervous system, is itself a
combination of three indispensable parts or elements : the
receptor, the conductor and the effector. Typically these
structures contain at least four cells, one for the receptor,

164 The Unity of the Organism

two for the conductor, and ortc for the effector. An illus-
tration would be a reflex-arc consisting of a tactile cell of
a touch organ, the two cells constituting the conducting
path, and an effector cell in a muscle. A point of special
interest in connection with such an arc is that the dependence
of the parts on one another is such, in the highest develop-
ment of the arc, that the specific action of each part is
dependent on the specific and connected action of the other
parts. "The optic nerve itself," to quote Sherrington, "is
unable to enter into a heightened phase of its own specific
activity on the application of light. Initiation of nervous
activity by light is the exclusive (in this instance) function
of cells in the retina, retinal receptors." And of course
without brain cells as effectors for vision the specific activi-
ties of receptors and conductors would be impossible. Not
only would an optic nerve fiber's conducting ability be useless
without a retinal cell on the one hand and a brain cell on the
other, but the very conductive act itself would not be fully
performed.

This specific dependence, as it might be called, of the
parts of a reflex-arc is so significant that another example
may be profitably noticed. A striking one is afforded by
the effects of the passage of gall stones through the gall
ducts, instanced by Sherrington, partly on the basis of his
own studies. The excruciating pains associated with this
malady are due to the distention of the wall of the gall duct
by the passage through it of the mineralized organic con-
cretions which constitute the gall stones. The point in
this for us is the fact that though the stimulus which pro-
duces the pain is mechanical and acts upon the wall of the
duct, this stimulus is so peculiar that other sorts of me-
chanical stimuli of the same tissue, even to cutting and
wounding, give no sense of pain. Though the duct may
be cut without causing pain, pain may be produced by
injecting the duct to distention with neutral fluid. "Marked

Neural Integration 165

reflex effects can be excited [by fluid] from the very organs
the cutting and wounding of which remains without effect." 2
The reflex mechanism involved is adjusted to respond only
to a stimulus of a special kind. In this it is comparable to
the optical mechanism referred to above.

These illustrations show something of the general nature
of the reflex-arc as an integrated structure. But we want
to know something about the part contributed by the
different constituents of the arc to this nature. The spe-
cific office of sense organs viewed as the receptor members of
the arcs will first engage attention. "The main function of
the receptor is therefore to lower the threshold of excit-
ability of the arc for one kmd of stimulus, and to heighten
it for all others."3 This principle is so important that we
must allow no vagueness as to its meaning. It means that
while more than one sort of stimulus may put a particular
sense apparatus into operation, there is one and only one
kind, namely, that to which the sense organ is naturally
subject, that elicits the reflex in its normal or type expres-
sion. A sense organ may be looked upon as a group of
receptors attuned to a special stimulus, as contrasted with
that of the general stimuli to which an organism is always
subject by being always in contact with its environment.

In illustration Sherrington instances the fact that the
threshold for the touch-sensation is lower for a mechanical
stimulus than for an electrical stimulus. Having regard to
the whole lot of reflex-arcs of the body, we may say that
the different sense organs constitute mechanisms of selective
.excitability for the different stimuli, each organ being so
adapted to its natural stimulus that it responds to this
better than to any other. This reference to the sense or-
gans as adaptive makes it desirable to notice the fact that,
according to Sherrington, electricity is never an adequate
stimulus of any sense organ because it is not a natural
stimulus for any, since it does not enter into the natural

166 The Unity of the Organism

environment of the organism. This is important in view
of the large use made of electricity in experimental work
on nerve physiology. To what extent may accepted gen-
eralizations concerning various reactions be influenced by
the inadequacy of the stimuli applied?

We glance next at the physiological character of the con-
ducting member of the arc. It is hardly necessary to say
that typically this part is found in the nerve cords every-
where present in the body. The most general truth of sig-
nificance for us is the fact that having regard to conduction
through the arc as a whole, the process is different in many
important respects from conduction in the nerve cord alone.
Conduction in the arc as an integrated whole is quite dif-
ferent from conduction in the part of the arc specially
devoted to that office. To illustrate by a quotation : "An-
other remarkable difference between reflex-arc conduction
and nerve-trunk conduction is the irreversibility of direc-
tion in the former and the reversibility of the latter."
This is only another way of expressing the well-established
fact that nerve impulses are incapable, in the higher organ-
isms especially, of running in more than one direction.
Sensory excitations can go only centripetally and motor ex-
citations only peripherally. As Sherrington points out, this
is part of the "law of forward direction" of the neural system.
It is fully established that nerve impulses may run in both
directions when a stimulus is applied at any point or a
nerve trunk, whether the trunk be motor or sensory. Taken
by itself this fact might encourage a somewhat careless ob-
server to think of the nervous system as more or less hit-or-
miss in its structure and action, it being able to work one
way as well as another, the final result being determined by
where the stimuli happen to be applied. As a matter of
fact, though, when we come to consider the real unit of
neural organization, the reflex-arc, instead of any of the
constituents of that arc taken separately, all suggestion of

Neural Integration 167

haphazardness disappears. Nerve conduction in a nerve
trunk as an isolated phenomenon has neither existence nor
meaning for the actual organism. The conducting element
par excellence of the real unit of nerve organization is
differentiated with reference to the other elements of that
unit, and normally acts only in such relation. The circum-
stance that this element is found to have the ability to act
somewhat differently under different but abnormal rela-
tions signifies little or nothing, so far as one can see, for
the normal workings of the nervous system. What it does
show is something of the diversity and plasticity of the
organism because of its latent ability to act otherwise than
in nature it does act when conditions are imposed upon it
which are wholly new to it.

Although the explanation of this irreversibility of direc-
tion appears not to be known with certainty, the suggestion
that it may be connected with a difference in permeability
of the synaptic membrane between cells of the reflex-arc to
certain ions, depending upon which side of the membrane is
presented to these ions, is plausible and indicates the re-
sourcefulness, as one might say, of structural and functional
method by which the organization of the living being is ac-
complished. Should this suggestion prove to be correct,
the question would arise, how comes it that the synaptic
membrane is thus differential in its action? And no ade-
quate answer would be forthcoming that did not take cogni-
zance of the fact that the membrane assumed this differential
mode of acting as part and parcel of the differentiation and
integration as the reflex-arc as a whole.

Sherrington deals with a whole series of other differences
between nerve trunk conduction and reflex-arc conduction,
such as the phenomena of summation of stimuli, that is,
the adding up of excitations too slight taken singly to pro-
duce reaction, until the aggregate brings response. Such
are rhythmic activity in response to stimuli which are not

168 The Unity of the Organism

rhythmic, or at least not of the same time-intervals as the
excitations, gradation in intensity of reaction, and so on.
To these he devotes more than a hundred pages, all under
the common heading, "Coordination in the Simple Reflex."
It is unnecessary to follow the matter further, sufficient
having been given to show something of the variety of ends,
all definitely and specifically contributory to the needs of
the organism as such, and all accomplished through differ-
entiation and integration of one relatively simple organic
structure, the reflex-arc.

The Integration of Reflex-Arcs

Our next task is to learn something of the combinations
among the myriads of reflex-arcs of which the higher nerv-
ous system is composed. The definition of the reflex-arc
given on a preceding page presents this mechanism in its
simplest terms, and so far as the definition indicates it would
seem possible for it actually to exist and be operative in
this simplicity. In fact, physiologists are accustomed to
assume such an entity and to call it the simple reflex-arc,
and its activity the simple reflex. But while such a concep-
tion is convenient and helpful for didactic purposes, espe-
cially as an aid to visualizing the earliest stage of cell inte-
gration in the evolution of the nervous system, as a matter
of fact, according to Sherrington, it probably never exists
in nature. "A simple reflex is probably a pure abstract
conception, because all , parts of the nervous system are
connected together and no part of it is probably ever
capable of reaction without affecting and being affected by
various other parts, and it is a system certainly never abso-
lutely at rest." 5

The factual basis for the conception that one of the most
essential distinctions between the nervous system and other
systems of the body is the integratedness of the former, is

Neural Integration 169

well expressed by Donaldson: "Isolated groups of nerve-
cells do not occur. Indeed, a group of nerve-cells discon-
nected from the other nerve-tissues of the body, as muscles
or glands are disconnected, would be without physiological
significance. It is desirable, therefore, to emphasize the
fact that by dissection the nervous system is found to be
connected throughout its entire extent." 6

I would ask the reader to consider these statements in the
light of the cellular conception of the organism maintained
in this volume and especially dwelt upon in the chapters on
the cell-theory, namely, that the true way of viewing the
organism is not as being built up of cells in the sense of
having been constructed by the bringing together of pre-
viously isolated cells, as a brick house is built up of bricks ;
but rather as being composed of cells through resolving
itself into these as it increases in size and differentiates
itself into its organs and functions. With such, a concep-
tion Sherrington's formulation, based on the findings of
vast observational and experimental research on the struc-
ture and action of the nervous system, is in perfect accord.
If we see in the completed nervous system a complex
mechanism developing as a unit subservient at all stages to
the needs of the organism as a whole, the myriads of reflex-
arcs then present themselves as final, as end-stages in the
differentiation and not as initial states; and the universal
organic and functional connection with one another, affirmed
in the quotation, would be just what we might expect.

If, on the other hand, the integral nervous system
were built up in a literal sense, that is, by the actual coming
into connection with one another of previously isolated
simple reflex-arcs, such arcs ought to be demonstrable both
in the ontogeny of higher animals and in the adults of the
lower metazoa.

No one should be beguiled into the notion that the readily
observed facts of ontogeny of the nervous system, the vari-

170 The Unity of the Organism

ous processes, dendrites and axones, do actually grow out
on nerve cells and bring cells into connection with one
another and with receptor and effector cells, and that a
functional coordination is thus finally reached does not
exist in any way or degree in the early stages. Proof that
the completed stages of neural integration cannot be ac-
complished without the production of cell-growths which
put cells into connection with one another is a very different
thing from proof that these cells were once wholly isolated
from one another, and that the outgrowths which establish
the final connections were initiated by impulses which origi-
nated wholly within the cells ; as, by way of illustration, the
branches of two young trees standing not far apart might
come into contact as the trees increased in size. It is a
matter of elementary knowledge of animal development that
in this sense the cells of the nervous system are never iso-
lated either from other nerve cells or cells of certain other
parts of the organism. Until we know vastly more than
we do know of the chemical nature of intercellular substances
and of the chemical and physical activities which go on at
the planes of contact between cells, nothing could be more
gratuitous and unscientific than to assume that nerve cells
differentiate as they would were they not in some measure
vitally associated with one another from the very begin-
ning. Indeed, such positive knowledge as we have tends
strongly against such an assumption.

Recall, for example, the fundamental part chemical mes-
sengers play in development. And Sherrington's insistence
on the role of intercellular substance and "surfaces of sepa-
ration" between cells in the functioning of the adult nervous
system is much to the point for this contention. Intercellu-
lar as well as mtracellular conduction must be, he main-
tains, expected in the reflex-arc on the basis of the cell
theory.7

Neural Integration 171

The Spreading and Compounding of Reflexes

As a practical matter study of the integrative action of
the nervous systems never gets away from integration among
reflex-arcs and reflexes any more than it does from the
cellular intergration fundamental to the reflex-arc itself.
The general nature of the study always involves the "spread
of reflexes over a wide range of nervous arcs." The idea
is more fully stated in the following: "This compounding of
reflexes with orderliness of coadjustment and of sequence
constitutes coordination, and want of it inco-ordination.
We may therefore in regard to coordination distinguish
coordination of reflexes simultaneously proceeding, and co-
ordination of reflexes successively proceeding. The main
secret of nervous coordination lies evidently in the com-
pounding of reflexes." For the rest, all we can do or need
do to meet the requirements of this discussion is to get at
the main principles as illustrated by examples of this com-
pounding of reflexes.

As a starting point for the discussion of this larger
aspect of neural integration, Sherrington takes what he has
called "the principle of the common path." Basal to this
conception is the familiar fact that a stimulus applied at a
single point, on the surface of the body, for example, gives
rise to a nerve impulse which may reach a great number
of muscles or glands. A single receptor with its neurone
must be in communication with a great number of effectors,
some of which are very different in kind. Looking at the
arcs from the effector ends, as one may say, it is obvious
that the impulses reaching innumerable effectors must come
over a single conducting course.

On the other hand, since a given muscle or gland may be
reached by impulses from a great number of reflex-arcs,
the effector and its neurone must be the "common path"
for all these impulses, often very different in quality and

172 The Umty of the Organism

source. "While the receptive neurone forms a private path
exclusively serving impulses of one source only, the final
or efferent neurone isy so to say, a public path, common
to impulses arising at any of many sources of reception." 9

An example which illustrates the general principle of the
common path, and several phenomena incidental to this,
and one which has been much investigated by Sherrington,
is that of the scratching reflex in dogs. It has long been
known that in several land vertebrates which have the
habit of scratching the side and back of the forward part
of the body with the hind foot, the scratching movement
may be elicited by appropriate stimuli applied to the area
reached by the foot as a pure reflex; that is, in the absence
of any chance for impulses from the brain to reach the
parts involved in the activities. Since the scratching move-
ment consists in bringing the hind leg forward and upward,
and for scratching, a rhythmic movement of the foot, the
muscles, both flexor and extensor, of the thigh, leg and
foot must be involved. And since the reflex can be induced
by a stimulus applied at any point within the large recep-
tive field (i. e., nearly the whole side and back of the body),
impulses started from various parts of the field must pass
through one and the same neurone in the muscles concerned.

And here comes in a fact showing another aspect of the
integration of reflexes in this case. A stimulus at a given
point in the field too weak by itself to elicit the reflex may
bring it on when acting in combination with weak stimuli
at other points in the field. Sherrington calls reflexes
which act together in this way allied reflexes.

Still another kind of combination of reflexes involving
the common path principle, even more significant than allied
reflexes, are what are known as proprio-ceptive reflexes.
The kernel of this class of reactions is the existence of re-
ceptors in the deep tissues of the body, that is, not belonging
to the surface and hence not subject to stimuli from the

Neural Integration 173

external world, but because of their situation subject only
to stimuli "given by the organism itself." Specifically the
sources of the stimuli are the muscles, joints, blood vessels
and so on, which by the regular activities are always in
more or less movable contact with one another and with
other parts and organs. Stimuli from the environment
acting upon the skin receptors give rise to reflexes which
put muscles into activity, and these activities serve in turn
as stimuli to receptors in the deep parts ; and the impulses
arising from these deep receptors may pass to still other
muscles over the same effector neurons used by impulses
coming from skin receptors. The proprio-ceptive field is
a sort of relay and supplement and extension of the field
of contact with the environment, or extero-ceptive field.

A simile used by Sherrington helps us to understand the
import of this compounding of reflexes : "The receptor
system bears, therefore, to the efferent paths the relation
of the wide ingress of a funnel to the narrow egress. Fur-
ther, each receptor stands in connection not with one efferent
only but with many — perhaps with all — though as to some
of these only through synapses of high resistance. The
simile to a funnel will therefore be bettered by supposing
that within the general systemic funnel, of which the base
is five times wider than the egress, the conducting paths
from each receptor may be represented as a funnel inverted
so that its wider end is more or less coextensive with the
whole plane of emergence of the final common paths. This
gives some idea of the enormous formation of common
paths from tributary paths which must take place." 10

And Sherrington forces home the truth of the scope of
the combinations by calling attention to the fact that under
strychnine poisoning "a muscle can be excited from prac-
tically any afferent nerve in the body." This is equiva-
lent to saying, he remarks, "that each final common path is
in connection with practically each one of all the receptors

174 • The Unity of the Organism

of the body."11

Even though this statement should prove to be too strong,
it certainly contains truth enough to show "the profusion
in which common paths exist."

The functional side of the total reflex system, the struc-
tural side of which was visualized by the funnel simile stated
above, may be regarded as pretty well presented by certain
phenomena known as irradiation. What is meant by this
is quite clear, in its general outlines, from the following:

"The more intense the spinal reflex . . . the wider, as a
general rule, the extent to which the motor discharge spreads
around its focal area. Thus, as stimulation of the planta
causing the flexion-reflex is increased there is added to the
flexion of the homonymous hind limb extension of the crossed
hind limb, then in the homonymous fore limb extension at
elbow and retraction at shoulder, then at the crossed fore
limb flexion at elbow, extension at the wrist, and some
protraction at shoulder; also turning of the head toward
the homonymous side, and often opening of the mouth,
also lateral deviation of the tail. According to circum-
stance, especially according to intensity of stimulation, the
field of end-effect of the flexion-reflex may vary from a
minute field occupying part of a flexor muscle of the knee
to a field including musculature in all four limbs and neck
and head and tail." 12

Antagonistic Reflexes in, Skeletal Muscle Groups Finally

Integrative

So far what has been said about the interaction between
reflexes has dealt only with interactions 'which are har-
monious with one another in various modes of combination.
But there are antagonistic or competitive as well as har-
monious interactions. These must now receive attention.
As an illustration we make use again of the scratching re-

Neural Integration 175

flex. "If," says Sherrington, "while stimulation of the skin
of the shoulder is evoking the scratch-reflex, the skin of
the hind foot of the same side is stimulated, the scratching
may be arrested." 13

Then the author proceeds to show, by description and
diagrams, how the two excitations here involved, one from
the skin of the side of the body, the other from the hind
foot of the same side, have the "same final common path"
to the muscles of the hip and leg, which, however, they "use
to different effect"; that is, the one to excite, the other to
inhibit, contraction of the muscles concerned.

At once there arises the important question: What is the
meaning of such antagonisms, such seeming want of har-
mony, as this? How deep-seated is the competition thus
shown? Does it amount to an "ultimate truth" in the na-
ture of the organism, thereby furnishing an argument in
favor of the "struggle of the parts" as an explanation of
the organism? Or is it possible that the antagonism is
secondary to the wider needs and activities of the organism
as a whole?

A partial answer to this inquiry is found in the character
of the stimuli operative in the two opposing fields. "Stimu-
lation of the skin of the hind foot by any of the various
stimuli that have the character of threatening the part with
damage causes the leg to be flexed, drawing the foot up
by steady maintained contraction of the flexors of the ankle,
knee and hip." 14

In other connections Sherrington dwells on the peculiari-
ties of effect and importance from the standpoint of adap-
tation, of stimuli of this class, as for example pricking,
strong squeezing, injurious heat, and so on.

From the descriptions we notice that the actions of the
hind leg involved in the scratching-reflex are considerably
different from those involved in withdrawing the hind foot
from stimuli of harmful portent. But since the same mus-

176 The Unity of the Organism

cles are necessarily involved in the two sorts of action, and
since the stimuli reaching these muscles from the two
sources must use the same final paths to those muscles, back
of the antagonism between the two sets of reflexes is the
question, which, in a given case, is more important to the
organism, the scratching action or the withdrawal-from-
danger action?

Since the scratch-reflex in the dog is probably connected
primarily with flea and other insect bites, and since on the
whole it may be assumed that these are rather annoyances
than real dangers to life and limb as the "nocuous" excita-
tions are by fundamental nature, it would be fair to infer
that though either reflex might under certain circumstances
inhibit the other, the threshold for the injury-escaping re-
flex would be lower. I do not know that there is anything
in the evidence which bears directly on this point, but the
question is one that would surely arise were the whole" sub-
ject of antagonistic reflexes to be looked at from the stand-
point of the needs and adaptations of the normally living
organism.

The way certain other reflexes, antagonistic in a sense,
are yet correlated in a larger sense, is more obvious than in
the case just given. Thus the reflex appertaining to two
limb muscles which oppose each other does not merely ac-
tivate the muscle which contracts ; it simultaneously causes
depression of the opposing muscle. Cooperative antagon-
ism, as it might be called, of this general sort is widespread
among higher animals, and applies to glandular, circulatory
and various other mechanisms as well as to the muscular.
Indeed, it being undoubtedly true, as Sherrington repeatedly
points out, that the "outward behavior" of animals involves
a great variety of movements which proceed in an orderly
sequence, if they are normal, it seems almost necessary to
suppose that really all normal reflexes must be cooperative
and harmonious with reference to the organism as a whole,

Neural Integration 177

even though when viewed one by one or in secondary groups
they are antagonistic.

Such a conception of the real nature of antagonistic re-
flexes is favored by the seemingly general fact that these
reflexes are seldom if ever really destructive of one another,
since they do not act upon one another simultaneously,
Their antagonism consists in a successional opposition to
one another. As they follow one another one acts in the
opposite direction to the other, and the antagonism is the
more real in that frequently they overlap to some extent.
But as already said this overlapping probably never amounts
to complete coincidence. Such overlappings and other
forms of partial opposition constitute the phenomena of in-
hibition which play a great role in the sum total of reflexes
of the organism. This is part of the method by which
transition is accomplished from one reflex to another, where
the same muscles, for example, execute both. But the fact
that the transition is accomplished normally "without con-
fusion," to use Sherrington's phrase, shows the subordina-
tion of the inhibitions to the organism as a whole.

Another important fact to which Sherrington calls at-
tention is that inhibitions which reflexes produce upon one
another never, so far as is known, result in injury to the
tissues involved. Genuine opposition of reflexes, as of any
other sorts of physical or chemical action, would, according
to all our conceptions of natural bodies, have deleterious
effects on the opposing bodies. As a matter of fact, in-
hibiting reflexes not only do not injure the mechanisms in-
volved, but actually prepare them for greater functional
activity later on.14

This beneficent effect, as it might be called, of inhibition
is perhaps illustrated by certain forms of compensatory
reflexes. Thus, stimulation of the central end of the nerve
to the extensor muscles of the dog's knee results in contrac-
tion of the flexors of the hip and knee. But on removal of

178 The Unity of the Organism

this stimulus contraction of the extensor muscles imme-
diately succeeds, this "rebound" being especially marked
in the vasto-crwretus muscle, the main knee extensor.15

TJie Antagonisms within the Autonomic System Finally

Integrative

It was mentioned above that "cooperative antagonism" in
nervous action is widespread in the animal mechanism. The
illustrations given pertained to the cerebrospinal system
and the innervation of skeletal muscles. We must now fol-
low this subject farther and deal with the somewhat similar
phenomena presented by the autonomic nervous system.
This subject was treated to some extent in the chapter on
internal secretions, to the extent, that is, that it implicated
the endocrinal glandular system. The presentation here
will involve some repetition of what was said in the previous
discussion, but the nature and importance of the subject
will justify this.

Cannon appears to have been the first to make clear the
similarity between the opposing action of the subdivisions of
the autonomic nervous system and what Sherrington calls
the reciprocal innervation of antagonistic skeletal muscles.
"As the above description has shown," Cannon writes, "there
are peripheral oppositions in the viscera corresponding to
the oppositions between flexor and extensor muscles." 16

The description referred to is summed up in the state-
ment that many of the viscera and other parts of the body
are innervated by either the cranial or sacral, i.e., the ter-
minal autonomies, and also by the thoracico-lumbar or mid-
dle autonomic, this double innervation being such as to be
statable thus: "When the mid-part meets either end-part in
any viscus their effects are antagonistic" 17 The heart-
beat is slowed by the cranial autonomic and quickened by the
thoracico-lumbar. Contraction of the smooth muscular

Neural Integration 179

layer of the stomach and small intestine to produce "tone" is
increased on the whole by the cranial autonomic, while gas-
tric peristalsis and secretion are inhibited and the arterioles
of these organs are contracted by the thoracico-lumbar.18
The pupil of the eye is contracted by the cranial autonomic
and dilated by the thoracico-lumbar. The lower part of the
large intestine is contracted by the sacral autonomic and is
relaxed by the thoracico-lumbar. The discharge tube of
the urinary bladder is relaxed by the sacral and contracted
by the thoracico-lumbar. The blood vessels of the erectile
tissue of the external genitals are dilated by the sacral
autonomic and contracted by the thoracico-lumbar; and
so on.

Now it is especially important, as Cannon says, to notice
the kind of service these subdivisions of the autonomic sys-
tem perform for the organism. On considering the func-
tions of the cranial division, one recognizes that they have
to do with bodily conservation. "By narrowing the pupil
of the eye they shield the retina from excessive light. By
slowing the heart rate, they give the cardiac muscle longer
periods for rest and invigo ration. And by providing for the
flow of gastric juice and by supplying the muscular tone
necessary for contraction of the alimentary canal, they
prove fundamentally essential to the processes of proper
digestion and absorption by which energy-yielding material
is taken into the body and stored. To the cranial division
of the visceral nerves, therefore, belongs the quiet service
of building up reserves and fortifying the body against times
of need or stress." 19

Pasing to the sacral division, one sees that as concerns
its distribution to the digestive and urinary viscera, its of-
fice is that of accomplishing the discharge of refuse material
from the body. Hence, "like the cranial division, the sacral
is engaged in internal service to the body, in the perform-
ance of acts leading immediately to greater comfort." 20

180 The Unity of the Organism

So much in illustration of the service of the two end di-
visions of the autonomic nervous system.

What, exactly, we now inquire, is the nature of the service
performed by the thoracico-lumbar division? Since this
part is, as already seen, antagonistic in its action to both
the cranial and sacral parts wherever it innervates an organ
or part also innervated by these end-parts, the inference is
readily drawn that its service to the organism as a whole
would also be in a sense opposed to the services of the
cranial and sacral divisions. Only a moment's reflection
is necessary to recognize that at some points at least this
is so. For instance, quickening of the heart beat through
innervation by the thoracic autonomic nerves, thereby send-
ing the blood stream more rapidly and strongly through the
whole body, is clearly opposed, so far as these acts in them-
selves are concerned, to the slowing of the heart and hence
of the blood stream by the vagus nerve, i.e., by cranial au-
tonomic innervation. Now this can mean nothing else than
that whereas the vagal (cranial) autonomic action is in
the interest of upbuilding and conserving the organism in
its whole normal, wonted life, the speeded-up action through
the thoracic autonomic is in the interest of some special,
more or less temporary need or condition. But obvious and
important as is this opposition between the two subdivisions
as concerns heart innervation, of not less importance, though
less easily observable in their full scope, are the oppositions
and reciprocations brought about in the peripheral blood
vessels of the whole body through the innervation of the
muscles of the arterial walls by the thoracico-lumbar au-
tonomic in connection with the heart action. These in-
nervations, coupled with those of the smooth muscles of the
gastro-intestinal canal, of the sweat and probably other
peripheral glands, of the smooth muscles of the hairs in
mammals, of the adrenal medulla for the secretion of
adrenin, and of the liver for releasing stored carbohydrates

Neural Integration 181

into the blood, by the thoracico-lumbar autonomic, produce
an equilibrating mechanism of well-nigh inconceivable com-
.plexity and delicacy, the workings of which are, however,
beginning to be revealed to us, both as to details and as to
rationale for the life of the organism as a whole.

The whole scheme is adjusted, we may say, for these two
necessities of the organism: (1) to serve the organism in its
uninterrupted and uninterruptible, and therefore normal,
growth and existence; and (2) to secure the perpetuity of
the organism through the recurring times of special demand
and stress which are inevitable from the external conditions
of life under which all living beings exist.

The supreme significance of all this from our standpoint
is that we get a glimpse of the means by which an extensive
and fundamental number of parts of the organism are sub-
ordinated in their special activities to the special needs of
the whole. The following from Cannon's chapter, Fatigue
and Blood Pressure, gives concreteness and some particu-
larity to this statement : "In connection with the foregoing
considerations [of facts bearing on the value of increased
arterial pressure in pain and strong emotion] the action of
adrenin on the distribution of the blood in the body is highly
interesting. By measuring alternations in the volume of
various viscera and limbs, Oliver and Schafer proved that
the viscera of the splanchnic area — e.g., the spleen, the
kidneys, and the intestines — suffer a considerable decrease
of volume when adrenin is administered, whereas the limbs
into which the blood is forced from the splanchnic region
actually increase in size. The action of the adrenin indicates
the relative degrees of sympathetic innervation." 21

This last sentence is, of course, what specially concerns
us in this discussion. Continuing, and having in view his
own researches into the beneficial effects on fatigued striated
muscles of adrenin in the blood, Cannon says : "At times of
pain and excitement sympathetic discharges, probably aided

182 The Unity of the Organism

by the adrenal secretion simultaneously liberated, will drive
the blood out of the vegetative organs of the interior, which
serve the routine needs of the body, into the skeletal muscles
which have to meet by extra action the urgent demands of
struggle or escape. But there are exceptions to the general
statement that by adrenin the viscera are emptied of their
blood. It is well known that adrenin has a vasodilator, not
a vasoconstrictor, action on the arteries of the heart; it is
well known also that adrenin affects the vessels of the brain
and the lungs only slightly if at all. From this evidence
we may infer that sympathetic impulses, though causing
constriction of the arteries of the abdominal viscera, have
no effective influence on those of the pulmonary and intra-
cranial areas and actually increase the blood supply to the
heart. Thus the absolutely and immediately essential or-
gans— those the ancients called the 'tripod of life' — the
heart, the lungs, the brain (as well as its instruments, the
skeletal muscles) — are in times of excitement abundantly
supplied with blood taken from the organs of less importance
in critical moments." And Cannon concludes with the very
pertinent remark : "This shifting of the blood so that there
is an assured adequate supply to structures essential for the
preservation of the individual may reasonably be inter-
preted as a fact of prime biological significance." Indeed
so ! Even more significant, I believe, than Cannon has shown
us — as we shall see in a later chapter. But the limitations
set at the outset of this chapter, namely that of dealing
only with neural processes strictly, especially as these mani-
fest themselves in reflexes, permits us to go no farther than
to call attention to the fact that while Cannon finds "the
most significant feature" of the reactions he has studied to
be "that they are of the nature of reflexes — they are not
willed movements, indeed they are often distressingly be-
yond the control of the will," 23 he yet has coupled them in
the most positive way with the emotions, especially with

Neural Integration 183

those of fear and rage, and with pain. Otherwise stated, he
has coupled them, more definitely than this has been done
before, with the conscious psychic life of the organism.

Concluding Remarks on the Significance of Neural Integra-
tion for the Organismal Standpoint

Perhaps enough illustration and general discussion have
been presented to convince the reader not only that "the
nervous system functions as a whole" 24 but that this func-
tioning is strictly subservient to the needs of the organism
as a whole, whether the normal individual, living normally,
or the normal individual living under special stress, be con-
sidered. It is hoped the reader will not have failed, despite
the brevity and inadequacy of the presentation, to perceive
the fundamental truth that the organism's totality of activi-
ties, executed to so large an extent through the agency of the
neural mechanism, are in turn subordinate to the needs of
the organism as related to its natural environment. This is
only another way of expressing the truth, which has become
almost trite since the idea of evolution has been an essen-
tial part of biological philosophy, that all, or at least by
far the major part, of the activities of organisms are
adaptive.

In connection with no other organ-system than the nerv-
ous does the truth come out so patently that the unity and
wholeness of the particular system, the unity and wholeness
of the organism, and the adaptiveness of the organism to its
environment are bound together inseparably; indeed, may
almost be called different aspects of one and the same truth
of living beings. The very conception of adaptation, at
least as touching neural activity, seems dependent upon the
correlatedness, the unitedness, of the differentiated parts.
The conception of adaptation, so far at least as reflexes are
concerned, depends essentially on the mode of relation of

184

The Unity of the Organism

parts. It is non-existent except through a particular form
of integration.

Every specific act of every part of the nervous system
is primarily in the interest of some other part and function
of the organism than itself.

This is particularly true for all adaptive acts of the
system.

Considerations of this sort seem not only to justify, but
to render necessary some such view of purpose in the or-
ganism as the following: "The infinite fertility of the or-
ganism as a field for adapted reactions has become more
apparent. The purpose of a reflex seems as legitimate and
urgent an object for natural inquiry as the purpose of the
colouring of an insect or a blossom. And the importance to
physiology is, that the reflex reaction cannot be really in-
telligible to the physiologist until he knows its aim." 25 No
biologist whose mind is both open and reasonably pene-
trating will hesitate to accept these views as correct.

What I wish to call particular attention to is that the
last sentence quoted calls for an additional clause: "the re-
flex reaction cannot be really intelligible to the physiologist
until he knows its aim," and he can know its aim only by
considering it in the light of the organism's entire complex
of normal activities; i.e., in accordance with the conception
of the organism as a whole.

REFERENCE INDEX

1. Sherrington 6

2. Sherrington 12

3. Sherrington 12

4. Sherrington :. 38

5. Sherrington 8

6. Donaldson 605

7. Sherrington 16

8. Sherrington 8

9. Sherrington 116

10. Sherrington 145

11. Sherrington 146

12. Sherrington 151

13. Sherrington 136

14. Sherrington 136

15. Sherrington 205

16. Cannon, W. B. ('16) 35

17. Cannon, W. B. ('16) 34

18. Cannon, W. B. ('14) 262

19. Cannon, W. B. ('16) 31

20. Cannon, W. B. ('16) 34

21. Cannon, W. B. ('16) 107

22. Cannon, W. B. ('16) 108

23. Cannon, W. B. ('16) 185

24. Sherrington 114

25. Sherrington 236

Chapter XXI

IMPLICATIONS OF THE TROPISTIC AND SEG-
MENTAL THEORIES OF NERVE ACTION

HAVING now carried our study of the nervous sys-
tem far enough to enforce the conception that the
very essence of this system, even the wholly reflex aspect of
it, is its unifying, its integrating office for the organism, it
remains to see what the elementalist mode of reasoning can
do with the system in this aspect of it.

As in the case of internal secretions, we can get at this
subject in no way better than through the writings of
Jacques Loeb. This way of approach will be seen to be the
more advantageous when Loeb's long-continued and dis-
tinguished experimental studies on some of the activities of
several kinds of lower animals are borne in mind.

Neglect of the Works of Sherrmgton and Cannon by
Jacques Loeb

Despite Loeb's express statement in the preface to The
Organism as a Whole that the volume is intended to be a
companion to The Comparative Physiology of the Brain, on
which account a discussion of the central nervous system is
omitted from The Organism as a Whole, the omission seems
very unfortunate. Much important work on the physiology
of the nervous system, especially on its integrating function,
has been done since The Comparative Physiology of the

185

186 The Unity of the Organism

Brain was published,* Sherrington's really epochal book
being especially notable among these later productions.
Some of the most fundamental views and arguments pre-
sented by Sherrington and others are irreconcilable with
conceptions which are at the very center of the elementalist
philosophy, and which find repeated expression in Loeb's
writings on the nervous system. It is, consequently, a mat-
ter of surprise and disappointment that a volume devoted
expressly to the organism as a whole should leave almost
unmentioned the very organ-system which is most distinctive
of the organism thus viewed, especially since this leaves un-
touched clearly formulated theoretical issues of cardinal
importance.

Likewise the disposal of Cannon's work, upon which we
have drawn so largely in the preceding chapter, by five
lines in The Organism as a Whole 1 is truly astonishing !

Indeed, so striking is the defectiveness of The Organism
as a Whole in this respect that one can not help recognizing
that despite its author's statement in the preface as to his
intentions, the omission amounts to an evasion even though
not so intended.

However, the expressed statement by Loeb of the place he
wishes the Comparative Physiology of the Brain to hold
in his interpretation of the organism has great usefulness
to us : it is tantamount to an assurance that the views set
forth in the Comparative Physiology have undergone no
essential change because of the discoveries and arguments
produced since the publication of that work. So our exam-
ination of Loeb's position on this important matter must
concern itself largely with facts and views contained in
the Comparative Physiology of the Brain.

* The copyright of Comparative Physiology of the Brain bears the
date 1900, while The Integrative Action of the Nervous System, by
Sherrington, than which few more important books on the system have
been written, was copyrighted in 1906.

Implications of the Theories of Nerve Action 187

The Real Importance of Loeb's Conception of the Nervous

System

First of all, it is with genuine satisfaction that I recognize
the eminent service rendered physiology and general biology
by this author's experimental investigations on animal ac-
tivities. I have long considered that one of physiology's
foremost achievements is the clarification of the conception
that nerve phenomena, the most characteristic features of
which are response to stimuli and the conduction of the
effects of stimulation, are not wholly unique attributes of
nerve tissue, but are elaborations of attributes inherent in
all protoplasm. Perhaps no single physiologist has con-
tributed more to this clarification than has Loeb. Par-
ticularly important has been his demonstration that nerve
centers in the sense of ganglionic masses which determine
reflexes are not fundamental to the coordinated and adapted
movements of animals ; that reflexes take place in many
animals normally and in many others experimentally, where
no such centers exist, as for example in the ascidian after
the single ganglionic mass is cut out.

The pertinacity and technical skill with which Loeb has
followed up these ideas are highly commendable. By focus-
sing attention on the fact that since plants have no nervous
system, they are, in so far as they exhibit movement induced
by stimuli, illustrations of the principle of protoplasmic
response in actually differentiated organic beings ; and by
carrying the same conception over into the animal world and
demonstrating that many activities here also are dependent
upon direct protoplasmic response and not necessarily on
a specially differentiated ganglionic mass, he was led to
formulate the tropism theory. This theory, as we shall
show presently, has a significance which Loeb himself seems
not to have fully comprehended, his general standpoint be-
ing unconductive to such comprehension. The theory must,

188 The Unity of the Organism

consequently, hold a large place in our effort to see what Loeb
as a representative of the elementalist school in biology is
able to do with the integrative action of the nervous system.
Let it be understood that with the general controversy
which has gone on so long and warmly concerning the value
of the tropism theory for explaining animal behavior, we are
little if at all concerned in this discussion. The point of
cardinal importance for us, and the thing about the theory
which, as mentioned above, Loeb and his followers seem not
to have perceived clearly, is the fact that the theory is
really dependent for so much of validity as it has on the
conception of the organism as a whole. In other words, the
theory is in essence an organismal and not an elementalistic
theory. Hence it results that since Loeb is the author and
chief proponent of the tropism theory, but is at the same
time an extreme elementalist, his general position as an in-
terpreter of animal activities and the nervous system con-
tains fundamental contradictions. Indicating these con-
tradictions and exploring, though incompletely, the course
along which the truth lies, will form a natural conclusion to
our study of the integrative office of the nervous system.

The Organismal Character of Tropisms Partly Recognized

By Loeb

I first call attention to the fact that Loeb himself has seen,
though "as in a glass darkly," the organismal character of
the tropism theory. Speaking of the lack of a sharp boun-
dary between reflexes and instincts, he says some authors
are wont to speak of reflexes when the responses to stimuli
pertain to single parts or organs, but they "speak of in-
stincts where the reactions of the animal as a whole are
involved (as is the case in tropisms)."

And the same idea comes out still more definitely in a
proposal to discard the term reflex as applied to such a re-

Implications of the Theories of Nerve Action 189

sponse as the clasping act by the arms of the decerebrated
male, frog, and substituting the term tropism. "It is better,''
he says, "to call them tropisms since the organism as a whole
is involved." 3

But Loeb's most illuminating recognition of the organis-
mal character of tropisms is found not in any positive
acknowledgements but in his statements of what a tropism
consists of essentially. This he has told us many times and
in varied phraseology during the last twenty-five years. The
following from The Organism as a Whole will serve well as
the basis of our consideration : "Animals possess photo-
sensitive elements on the surface of their bodies, in the eyes,
or occasionally also in epithelial cells of their skin. These
photosensitive elements are arranged symmetrically in the
body and through nerves are connected with symmetrical
groups of muscles. The light causes chemical changes in
the eyes (or the photosensitive elements of the skin). The
mass of photochemical reaction products formed in the
retina (or its homologues) influences the central nervous sys-
tem and through this the tension or energy production of
the muscles. If the rate of photochemical reaction is equal
in both eyes this effect on the symmetrical muscles is equal,
and the muscles of both sides of the body work with equal
energy; as a consequence the animal will not be deviated
from the direction in which it was moving. This happens
when the axis or plane of symmetry of the animal goes
through the source of light, provided only one source of
light be present." 4

Reduced to its lowest terms, the tropistic mechanism de-
scribed here and in many other places consists of: first,
portions of the body surface having specific irritability, as
receptor organs, these being usually paired and usually
symmetrical, though often unsymmetrical ; second, muscles
as effectors, almost always paired and usually symmetrical;
third, nerves, variously constituted, as conducting paths

190 The Unity of the Organism

connecting more or less directly the sensory areas with the
muscles. In a word, tropistic movements are determined by
a definite number of definite kinds of parts or organs, defi-
nitely arranged with reference to one another and to the
whole; that is, upon an organized body called a living being,
or briefly an organism. A morphological entity is funda-
mental to the tropism conception. Tropisms are explained,
partly, by these organizations. Loeb does not avoid, in fact
at times appears not to try to avoid, recognizing this : "The
irritable structures at the surface of the body and the ar-
rangement of the muscles determine the character of the
reflex act." "The explanation of them [tropisms] depends
first upon the specific irritability of certain elements of the
body-surface, and, second, upon the relations of symmetry
of the body." 5

The Organismal Character of the Segmental Theory of the
Nervous System

Another province in which Loeb has done distinguished
work and into which the organism as such constantly ob-
trudes itself and will not accept relegation to a secondary
place, is that of the coordinated action of the central nerv-
ous system in those animals in which that system consists
of an axial series of ganglia. Here again, as in the tropism
theory, the basal perception underlying the segmental theory
of nerve physiology is the truth that nerve "centers" as the
"seats" of particular activities and functions in the old sense
(in the sense, that is, of being the real source of these
activities) do not exist. According to the segmental theory
the real unit of activity of an animal made up of metameres
or segments is the segment itself, the nerve ganglion of the
segment being only one element in the complex. The idea
is set forth with fullness and perspicacity in the chapters
of the Physiology of the Brain dealing with worms and

Implications of the Theories of Nerve Action 191

arthropods, and especially in the one on the segmental the-
ory in vertebrates.

The starting-point chosen by Loeb for the treatment is
the earthworm. It is generally known that this animal is
jointed, or segmented, that every segment has a pair of nerve
ganglia situated underneath the intestine on the floor of
the body wall, and that the pairs of ganglia are all connected
with one another fore-and-aft, so that the whole constitutes
a chain of ganglia extending the entire length of the body.
In addition to this series of ventral ganglia, there is a pair
of large ganglia above the gullet at the extreme front end
of the creature, these being connected with the ventral nerve
chain by commissures. These dorsal nerve masses are usu-
ally called the supra-esophageal ganglia, but are often
spoken of as the brain.

On the basis of the "center" theory of the nervous sys-
tem, it was very natural to raise the query: Is the earth-
worm's brain the "seat", the primal source of the coordi-
nated activities, the crawling and burrowing which make
up so large a part of the life activities of these crea-
tures? Loeb's own formulation of the question is good.
"Does coordinated progressive movement in which all the
segments of the body participate, depend upon the brain?"

The first attempt to answer this question experimentally
seems to have been made by Friedlander. Having made a
small opening in the side of the body of a living worm and
cut out a piece of the nerve cord so that a complete hiatus
in the cord was produced, the author fully expected, he tells
us, that after the healing of the wound in the body wall,
which took place very quickly and completely, the parts of
the body in front of and behind the section of the cord would
either behave like two separated individuals, or that the
front part would draw the rear part passively about. But
neither result followed. "Animals from the middle of which
a piece of the central nervous system is wholly wanting,

192 The Unity of the Organism

crawl exactly like normal animals except as to a small dif-
ference to be noticed later." 6 The wave of muscle con-
traction and other movements, starting at the anterior end,
passed over the point at which the ganglionic chain was in-
terrupted and on to the hind end of the body just as though
there were no such interruption. "The movements were
coordinated exactly in the same way as in the normal ani-
mal."

These experiments, together with others that have since
been performed by other investigators, overthrow, as Loeb
says, "the idea that coordination in these animals is deter-
mined by a special centre of coordination which is located
in the brain." 7

But if the brain is not the coordinating center and does
not contain it, what and where is that center? The answer is
there is no such center. Coordination is not a central matter
at all. Rather it is a matter of the working together of
many parts, each duly balanced off with many others, and
properly subordinated to the whole animal.

The details of the truth thus stated in general terms can
be illustrated more advantageously by referring to some of
Loeb's own observations on a group of annelids considerably
higher in the scale of life than the earthworm. I refer to
the pile worms familiar to many persons accumstomed to
marine shore fishing. This group of worms, known collec-
tively as nereid annelids, are far more "heady" than earth-
worms in that the useful body member named head is more
definitely set off from the rest of the body than it is in the
earthworm, and is outfitted with eye-spots, touch appen-
dages, and so on, wholly lacking in the earthworm. Nereis
is much more highly "cephalized" than is the earthworm, so
brain centralization would be expected to present features
here not occurring in the more lowly worm. When nereids
are deprived of the supra-esophageal ganglia their behavior
is such as to suggest at first sight support for the concep-

Implications of the Theories of Nerve Action 193

tion that the brain is a fundamentally originative and coor-
diriative member. For example, an individual thus depleted
will crawl about unceasingly on the sand of the aquarium
bottom, without, however, showing any signs of burrowing
into the sand, though this operation is very characteristic
of the normal animal. This looks as if the brain were indeed
the seat of the burrowing instinct. But not so. The bur-
rowing activity can be induced not only in animals deprived
of the brain, but even in parts of animals from which the
heads have been cut away altogether. Loeb placed a brain-
less piece of a Nereis on the sand, and as usual it remained
quiet. "I then gradually covered the forward end with
sand. The rest of the animal immediately began to make
the typical movements which the animal makes in forcing its
way into the sand. At the same time the glands began to
secrete the sticky substance which cements the particles
of sand together, forming the wall of the burrow-hole."
"But, why," Loeb asks, "does the Nereis not burrow when
deprived of its brain? For the simple reason that it makes
use of the organs of the mouth in burrowing, and these are
amputated with the head. Hence it is the loss of the pe-
ripheral head-organs which keeps the decapitated Nereis from
burrowing, and not the loss of the brain."

Since the coordinated activities involved in burrowing can
be accomplished through the acting together of various su-
perficial and deeper body parts — sense organs, muscles, and
so on — without the intermediation of the brain, these pe-
ripheral members are the real "seat" of such activities. The
author then goes on to inquire what part the brain does play
in the creature's normal life. "If we compare," he says,
"the conduct of a Nereis whose brain has been amputated
with that of a normal worm, the difference seems to be of the
same nature as that between an insane and a rational human
being. . . . The peculiar irritability by means of which the
Nereis draws its head back and moves backward out of the

194< The Unity of the Organism

tube depends upon organs which are located in the forward
end of the body and whose sensory nerves go to the supra-
esophageal ganglion." 9

One office of the Nereis brain is, therefore, to make a
great complex of activities normal, or "of the same nature"
as sane activities in "rational human beings." Not the ac-
tivities themselves but their normality are dependent on the
brain. This distinction is undoubtedly general, but it is
definite so far as it goes. Beyond question to the student
who should compare the movements of the normal and the
brainless Nereis with sufficient care, the distinction between
the two would be as positive, as indubitable, and as definable
as that between almost any two complexes of organic phe-
nomena. The unsatisfactoriness of the distinction is not its
vagueness but its generalness. It is sufficiently definite
though not sufficiently analytic.

Just what the brain does to normalize the activities ; in
other words, how the change of character of the worm's
behavior — its "restlessness" and general insane-like conduct
• — results from the absence of the brain, is not known. Loeb
makes a suggestion toward an explanation, but recognizes
what he offers as nothing more than a suggestion ; so to go
into it is not worth our while.

But there is an obvious usefulness of the ganglia and
whole neural chain in such animals aside from the general
one of normalizing behavior. It serves as a common con-
ducting and relay system between the peripheral sense or-
gans and locomotor muscles. Loeb's statement of the matter
is concise and may be adequate: "The central nervous sys-
tem does not control response to stimulation; it merely
serves as a conductor from the point of stimulation to the
muscle through which weaker stimuli may pass, and pass
more rapidly than would be possible if the muscle were stim-
ulated directly.

"In the Annelids each ganglion is a relay station for

Implications of the Theories of Nerve Action 195

the sensory and motor nerves of the corresponding segment.
If the head exercises a stronger influence upon the behavior
of the animal than any other segment, as in Nereis, for in-
stance, I believe it is due to the fact that in the oral end
more kinds of irritability are present and more peripheral
organs are differentiated (sense organs, mouth, etc.) than in
the other segments." 10

Loeb has done good service in bringing together the evi-
dence for, and emphasizing the truth that, the segmental
make-up of the vertebrate head, so long and so ardently
studied by anatomists and embryologists, is as important a
matter for neural physiology as it is for morphology, and
that as a matter of fact the vertebrate spinal cord works
more or less on the segmental principle, and in this respect
falls into line with the central nerve axis of annelids and
arthropods.

The first, or one of the first, investigators to recognize
this truth was Schrader, in his studies on the activities of
frogs whose brains had been wholly or partly removed. The
frequently quoted and later fully confirmed statement by
this investigator is of prime significance for the general
truth of what we might call fore-and-aft neural integration
in vertebrates. "The se*ries of experiments we have given
teaches us that the central nervous system of the frog can be
divided into a series of sections, each of which is capable of
performing an independent function. It brings the central
nervous system of the frog into closer relation with the cen-
tral nervous system of the lower forms, which consist of a
series of distinct ganglia that are connected by commissures.
It speaks against the absolute monarchy of a single central
apparatus and against the existence of different kinds of
centers, and invites us to seek for the centralization in the
many-sided coupling of relatively independent stations."

As statements much at variance from this concerning the
"central" function and power of different parts of the

196 The Unity of the Organism

frog's brain are not uncommon, and as the facts are spe-
cially important for us, it is desirable to consider the evi-
dence a little more in detail. To this end the following from
Holmes may be taken as an authoritative summing up of
results so far rea'ched in this field. "A frog with the brain
removed as far back as the medulla is still capable of per-
forming regular leaps and swimming movements of the
limbs. When thrown on its back it rights itself, and still
performs compensatory motions when tilted or rotated.
Breathing is normal, and the animal swallows pieces of food
that are placed in its mouth." 12

If the removed portion of the brain is made yet larger,
by including the anterior part of the medulla, the animal is
still capable of performing a long series of regular activi-
ties. "Locomotion is effected mainly by creeping, but the
frog is nevertheless capable of springing in the ordinary
manner. In the water it swims by alternate movements of
the limbs. . . . The breathing and swallowing reflexes are
still normal, but the croaking reflex is no longer performed.
The reflex of snapping at food is not destroyed. ... If a
piece of meat is rubbed against the frog's nose, the animal
snaps at it and uses the fore legs to stuff it into the mouth.
The same reaction may be brought about by using the finger,
but after the finger is seized and it is found that the object
is too large to be stuffed into the mouth, the frog begins to
reject the morsel, and uses the fore legs to push it away.
Truly a remarkable combination of reflexes!" 12

And further: "There is no center for coordinated locomo-
tion in the medulla. Disturbances of locomotion begin with
the fore limbs. If the medulla is cut across at the tip of
the calamus scriptorius, the animal sinks on its breast, and
the fore limbs are for a considerable time helpless, although
the hind limbs are capable of performing vigorous coor-
dinated movements. The reason for this is that the injury
lies so near the region from which the nerves of the fore

Implications of tlie Theories of Nerve Action 197

limbs arise that the movements of these members are very nat-
urally affected." 13 It is the interdependence fore-and-aft
of reflex activities like those here indicated, that is, of the
fore limbs acting with the mouth, and the hind limbs with
the fore limbs, that suggested to Loeb the idea of "chain
reflexes," a conception of special interest from our stand-
point.

That the segmental mode of functioning of the vertebrate
nerve axis is not limited to the lower portion of the great
vertebrate phylum is, as Loeb points out, shown by such
investigations as those of Goltz, and of Goltz and Ewald on
the dog. These are given so fully in the larger text-books
of physiology and are thereby so readily accessible to all
specially interested in the subject, that a very brief presen-
tation will suffice.

Two main groups of reflex centers in the nerve cord are
made out, the cervico-thoracic and the lumbo-sacral. The
first group contains the center for movements of the an-
terior limbs, for respiratory movements, for acceleration
of heart action, for dilation of the pupil of the eye, and for
several other activities at the anterior end of the body. The
lumbo-sacral group contains the center of movements for
the posterior limbs, for tontrol of the anal sphincter, the
activities of the generative organs, and several other parts.

The existence of such centers in the dog's cord was proved
by section of the cord at different levels under the most
careful operative conditions. Had the investigation stopped
with experiments of this sort the facts brought out might
have been interpreted as confirmatory of the "seat" notion
of a nerve center. When, however, it was discovered that a
dog from which most of the cord is removed may live for
many months in a good state of health, all the vegetative
functions being carried on almost typically, it became clear
that the various centers of the cord are not the seats of life
activities in any such fundamental sense as was earlier

198 The Unity of the Organism

supposed.

For one thing the results seemed to show that the sym-
pathetic nervous system shares with the central system
more profoundly than had been known before, in determin-
ing life processes.

The conclusions which flow with apparent certainty from
the observations are well summed up in the following: "Un-
doubtedly all such activities [of visceral life] may subsist
and function in a comparatively normal fashion after re-
moval of all spinal influence. The office of the spinal sys-
tem in regard to the visceral life seems to consist in en-
dowing these functions with greater energy, and in con-
ferring greater stability and more solid equilibrium on the
general constitution of the animal." 14

The reader will hardly fail to recall, on reading this state-
ment about the function of the mammalian spinal cord,
Loeb's statement, quoted some pages back, about the nor-
malizing function of the Nereis brain. "Stabilizing" the
dog's function means much the same as "normalizing" the
annelid's functions. Nor should the reader neglect to notice
that he might substitute the word "integratedness" for
"equilibrium" in the quotation without change in the es-
sential meaning of the sentence.

Critique of the Element alist Attempt to Interpret Tropistic

and Segmental Theories of the Function of the

Nervous System

We now return to the central point of the present sub-
topic, namely that of what a genuine bio-elementalist is able
to do when confronted by the facts in possession of modern
physiology bearing on the tropistic and segmental theories
of the nervous system.

First we are compelled by the evidence to recognize the
general soundness of the doctrines. Second, we recognize

Implications of the Theories of Nerve Action 199

that probably no biologist has dealt with the conception
more comprehensively and illuminatingly than Loeb. Third,
we find that his espousal and able treatment of the theories
has led him into positions so thoroughly organismal in both
essence and expression as to be quite irreconcilable with his
own more general elementalistic philosophy.

It remains to point out more specifically than we have
wherein the organismal implications of his teachings as ex-
emplified by his writings constitute a refutation of the
elementalistic implications of his teachings as exemplified
by his strivings after an "ultimate explanation" of organic
phenomena in the terms of physics and chemistry.

The essence of the irreconcilability of the two positions
may be put into a general form thus : in innumerable state-
ments and definitions found in his discussions of these the-
ories, Loeb is compelled to introduce the organism either as
a whole or in considerable portions, as a causal explanation
of particular phenomena with which he deals ; and the com-
pulsion to such introduction makes it impossible for him or
any one else to dispense with the causes thus introduced by
resolving them into ultimate elements of any sort, whether
organic or inorganic.

Illustration and justification of this general statement
must be given. The following typical sentence may intro-
duce the discussion: "The irritable structures at the sur-
face of the body, and the arrangement of the muscles, de-
termine the character of the reflex act."15

Notice what it is that "determines" the character of the
act. "Structures" and "muscles" do it, these being "ar-
ranged" so-and-so, the irritable structures definitely on the
surface of the body and the muscles within the body. The
point needing special attention is that not chemical com-
pounds or even living substances but structures — organs —
and these arranged; that is, entities which neither exist nor
can exist except through the agency of an organism, enter

200 The Unity of the Organism

into the definition.

That the structures referred to and all others of the or-
ganism are composed of chemical substances and nothing
else, is beyond question. Why, then, does the author not
bring into the definition this truth about the ultimate make-
up of the structures? The central aim of his researches
on animal activities being what they are— "a physico-
chemical analysis of behavior," is it not surprising that he
should be satisfied with even a working description or defini-
tion of a reflex act which takes no notice of the results of
such analysis? If the understanding of complicated life
phenomena consists in resolving them into their simple ele-
mentary components (see the first sentence of the Physi-
ology of the Bram), then a definition of a tropism that
would contribute largely to an understanding of it should
not be content with such proximate constituents as "irri-
table structures," "muscles," and so on, but should go right
back to the ultimate physico-chemical elements.

The typical elementalist answer to these restrictive criti-
cisms is well known. It is, substantially, that the understand-
ing and hence the definition of tropism, or for that matter
of any other life phenomenon, is final or ultimate only when
expressed in physico-chemical terms. The criticism sug-
gested concerning the mere proximateness and hence inade-
quacy of a definition that uses such terms as "structures,"
"muscles" and "body surface" is allowed to have a large
measure of validity, the usage bein'g justified mainly, it is
claimed, on practical grounds. The morphological concep-
tions involved, it is held, are so strongly intrenched in bio-
logical terminology, indeed are so necessary in a historic
and subsidiary sense, that it is very inconvenient, if indeed it
is not impossible, to dispense with them. In a word, the
contention is that while morphology and general physiology
are necessary, their necessity is secondary or subsidiary to
physics and chemistry. But what should be seen in this

Implications of the Theories of Nerve Action 201

with the greatest possible clarity is the error, the fallacy, not
of objective fact but of reasoning. Stated in general terms,
the fallacy consists in the tacit assumption that the indis-
pensability and adequacy of physico-chemical elements to
explain organic behavior are of a sort that exclude the in-
dispensability and adequacy of "structures," as sense or-
gans and muscles, from the explanation of the same phe-
nomena.

To go into the epistemological and logical necessities
involved in the situation with which we are here confronted
is entirely beyond the scope of this volume. However, in
the interest not so much of Truth in the abstract as of
healthy, wholesome, useful science, biologists will have to
cease employing such deprecatory epithets as "anthropo-
morphism," "metaphysics," "rhetoric" and the like to gain
for themselves a sense of security in the use of language and
reasoning which can not endure for an hour the searchlight
of really careful thinking and expression.

To focus the general statement of the fallacy on the par-
ticular matter in hand, one must see that both factually
and epistemologically the organs and other morphological
and general functional elements, or factors found by the
analysis of the organism, are "fundamental" or "ultimate"
for the phenomena to be explained by exactly the same cri-
teria that the physico-chemical elements are fundamental
or ultimate.

A reflex act or a trophism can no more be intelligibly
expressed or understood or conceived as an objective fact
without sense organs, muscles, etc., than without physico-
chemical substances. If one questions the truth of this af-
firmation let him test the matter by trying to express a re-
flex act in the terms of the physico-chemical elements known
from analysis to be "behind^,' such an act. To begin with, he
finds it necessary to fix upon some particular reflex act,
scratch-reflex, perhaps. Such particular act must be taken

202 The Unity of the Organism

as the starting point, otherwise the experimenter will be com-
mitted to the Idea of the act rather than the act itself.
And I take it for granted that no truly present-day biologist
is willing, even though he may not be able to justify fully his
unwillingness, to commit himself to that ancient position.
Very well then, the scratch-reflex is going to be expressed in
terms of the oxygen, nitrogen, carbon, phosphorus, and so
on, which we are sure are "behind" it. Having recognized
the necessity of starting with some particular reflex act, and
having settled upon the scratch-reflex, true to his recogni-
tion of the necessity for particularity he sees he must go
still further in the same direction. If his analysis is to be
exhaustive and his expression adequate and final, it will
have to particularize still further and concern itself with
the reflex of some particular animal species, very likely even
with some single individual or small group of individuals.
A vast mass of evidence makes it almost certain that a
dog's scratch-reflex is different from a cat's, and both are
different from an ox's, a frog's, and so on. Suppose, then,
the dog's reflex settled upon, and defined so fully that it
is distinguished from every other reflex whatever. The stu-
dent is now ready for his main undertaking, that of ex-
pressing the dog's scratch-reflex in terms of the physico-
chemical elements to which the reflex is reducible.

The next step is to examine the chemical elements con-
cerned, as these are treated in inorganic chemistry, for the
purpose of seeing what their "terms" are; that is, what
their attributes or properties are, the special purpose of this
examination being to ascertain what terms, i.e., what at-
tributes, answer to, or correspond to, and so can be used to
express, the dog's scratch- reflex. The outcome of this
examination is unequivocal. It finds no terms, no attri-
butes whatever, which by themselves suggest even remotely
the reflex under consideration.

But since the examiner has satisfied himself that no other

Implications of the Theories of Nerve Action 203

simple elements, material or immaterial, are "behind" the
reflex, and since the reflex is an indubitable reality, there is
no escape from the conclusion that something other than the
original inorganic simples must have intervened between
these simples and the reflex. And what is that something?
Does common experience and common sense hesitate with its
answer? If it does its hesitancy is probably from surprise
that so obvious a matter should be made the subject of
serious questioning. "The dog is what has intervened be-
tween the chemical simples and the reflex." That is what
common experience must answer and will unhesitatingly
answer once it recovers from its surprise at being ques-
tioned on a subject so open and daylight clear.

But then science comes forward with its criticism of this
common-sense answer. There is no gainsaying, it admits,
the truth of the nai've answer thus given. But this answer,
science says, is a mere truism. It leaves the case just where
it was before science began its analysis, so is worthless for
scientific purposes, however useful it may be for ordinary
purposes.

This rejoinder by science must be looked into carefully;
otherwise its weakness will be missed. It must be exacted
of science that she show more explicitly than she has what
she means by explaining the dog's scratch-reflex by refer-
ring it to the physico-chemical elements at the basis of the
act. Let us, we must insist, hear you express a dog's
scratch-reflex in the terms of oxygen, carbon, et cetera.

That such expression is possible is freely granted. But
how can it be done? That is a key question. It can be
done in one and only one way, namely by adding to the
attributes, that is, to the "terms" which inorganic chemistry
recognizes in the chemical elements concerned, just those
attributes and terms which the dog's scratch-reflex requires
in order that the elements may explain the reflex. We can
say that besides the specific gravity, combining weight, and

204 The Unity of tlie Organism

other well-known attributes of the elements, they possess
dog's-scratch-reflex attributes. But, the name thus sug-
gested for the newly discovered attributes being cumber-
some, we may devise for them some term more simple and
convenient — for example, do-sca-re-x, doscarex.

In virtue, then, of the doscarecious powers of oxygen,
carbon, and so forth (the fact that these powers can only
be assigned in the lump and not distributively to the several
elements should not be lost sight of though it may be neg-
lected for the present argument), a complete physico-chem-
ical explanation of the phenomenon under consideration is
reached.

Does this discussion advance the interpretation, the under-
standing of biological phenomena beyond the familiar sar-
casm about explaining drug-induced sleep as due to a dor-
mitive principle of the drug? Yes, I maintain, it does,
because in place of a "principle" attached to no particular
drug but to any sleep-producing substance, the doscarecious
powers recognized by us are definitely assigned to the chem-
ical elements known to be the sole constituents of organ-
isms. The new powers take their place perfectly, definitely
and positively among the other attributes of the elements,
the assignment being based on the solid ground of analyses,
laboratory and other, made through years of scientific re-
search. But an exceedingly important point which comes
in sight here is that though these doscarecious powers are
proved to be real ones, they are latent and wholly unknown
to inorganic chemistry for the reason that they never mani-
fest themselves under any other combination of things than
just that which in its totality Zoology has named dog. A dog,
and a dog only, is able to cause oxygen, carbon and the
other elements to reveal these particular scratch-reflex
powers. The dog comes in as a sine qua non to the pro-
duction of, and hence to the causal explanation of, the
particular group of activities under consideration. We

Implications of the Theories of Nerve Action 205

have no evidence that the chemical elements operating them-
selves can actualize their own latent doscarecious powers.

The reader will hardly fail to connect what we are say-
ing with the familiar phenomena of the assimilation by
organisms of nutrient substances. All our argument really
does to the usual conception of this phenomenon is to focus
attention upon the fundamental importance of the individual
organism as a factor, as a cause, of the chemical transfor-
mations wrought in the nutrient substances. The insuf-
ficiency of statement of the assimilative, or anabolic, or
synthetic aspect of the metabolic cycle in the organism,
lies in its failure to bring out clearly enough the indubitable
fact that the final results are innumerable activities and sub-
stances which pertain solely to the living, normal individual
— which are strictly personal and private, as one may say.

The current mode of expression according to which the
assimilative syntheses take place in the organism is quite
misleading in that it permits or even encourages a concep-
tion that the syntheses have a measure of detachment from,
and independence of, the life of the organism as a working
unit, which as a matter of fact they do not have. Again, the
usual statement that the* syntheses result in organic sub-
stances of more complex, higher make-up is inadequate in
that it diverts attention from the fact that these new sub-
stances belong to, are fundamentally part and parcel of the
organism. They are not "any old organic substances" but
are exactly those substances necessary to maintain the nor-
mal life of the particular individual organism. Hence de-
spite the indubitable fact that the final results are reached
by way of innumerable purely physical and chemical opera-
ations, the organism itself, acting as an integer no matter
how complex, is always to the fore as a controlling, domi-
nating factor.

206 The Unity of the Organism

The Bearing of This Critique on "Analysis" in Biological

Reasoning

The considerations thus briefly set forth lead to certain
still more general ideas of the utmost importance. The
natural entities to which we apply the descriptive terms
living and organic are specially distinguished by the chem-
ical and physical syntheses which they accomplish by virtue
of their inherent constitution. So as concerns the most
characteristic of these syntheses, especially the chemical
ones, they are to a very considerable and fundamental ex-
tent definitive of the organic individual, species, genus, and
so on, of taxonomic biology. This is equivalent to saying
that the synthetic operations regarded each by itself ter-
minate in results which are in large measure unique and so
unforeseeable from anything we know about the original
elements as such; that is, before they have actually been
subjected to the particular synthetic transformations under
consideration.

And this again is equivalent to saying that synthesis —
transformatory synthetic processes and products — is more
distinctive of living beings than are analytic products and
processes.

Finally, it follows from these facts about the synthetic
nature of organisms, and from the established principles of
thought, that analysis alone is incapable of interpreting, of
understanding organic beings. No natural object which in
its nature is more distinctively synthetic than analytic can
be understood by knowledge-processes which are more ana-
lytic than synthetic.

This conclusion goes to the very heart of the elementalistic
position, and, as stated in the discussion on internal secre-
tions, is really as much an epistemological as a biological
problem.

Reverting again to Loeb's writings, the conclusion to

Implications of the Theories of Nerve Action 207

which we are forced joins issue with the very opening sen-
tence of the Physiology of the Brain, as we have already in-
dicated. "The understanding of complicated phenomena,"
particularly as presented by organisms, can not be accom-
plished through "an analysis by which they are resolved into
their simple elementary components." Unquestionably an-
alysis is essential to, but it is not adequate for, full under-
standing of the phenomena.

The very nature of organic synthesis and of knowledge-
getting precludes ilie possibility of attaining the kmd and
degree of understanding of organisms which element alist
biology claims to have attained and promises to attain.

"A complete explanation of life in terms of physics and
chemistry" is impossible for the sufficient reason that physics
and chemistry as such do not contain any of the really dis-
tinctive terms of life. Those terms can only be brought
into physics and chemistry after and not before the phe-
nomena of life have been searchingly scrutinized ; that is, an-
alyzed and found to involve physics and chemistry. The
terms of life are in the original data on the phenomena of
life, and no sort of analysis can possibly make this other-
wise.

Our position, it should be noticed, touches the hackneyed
controversy over vitalism and materialism only in so far as
the course of reasoning we have pursued involves the recog-
nition that each organism (the dog, for instance) is a
natural object possessed of certain causal powers, by ex-
actly the same logical and epistemological criteria that any
simple chemical element or chemical compound is a natural
object.

Stated in a brief and common-sense way, our contention
is that the attributes which make a dog a living body are no
less natural than are the attributes which make carbon a
chemical body. The tiresome and meager-fruited contro-
versy between Materialists and Vitalists may be character-

208 The Unity of the Organism

ized as due to the fact that neither party has taken the
trouble to establish clearly, even in their own minds, the
meaning of the word natural. As a consequence of this
slipshodness the two groups agree tacitly, in treating the
inorganic and the organic worlds as though natural does
not mean the same thing in the two realms. The implica-
tion is that if the inorganic world, for instance, be held to
be natural by both parties, for the Vitalists the living realm
is largely super-natural, while for the Materialists the same
realm is largely iw/ra-natural.

Theories of Animal Behavior in Relation to the Science of

Zoology

This somewhat protracted though wofully insufficient
treatment of neural integration may close with a brief
section on some of the still larger biological and methodo-
logical implications of the conclusions reached. Special at-
tention is called to the fact that the culminating part of
our argument has involved data and conceptions which are
as unequivocally zoological, morphological, and physiologi-
cal, as any of the data and conceptions are unequivocally
physical and chemicaL Physical chemistry, or any other
aspect of inorganic chemistry, is utterly powerless, so far
as we can see, to discover such facts, as for example, that
oxygen, carbon, nitrogen, etc., possess latent "doscarecious"
powers. This final section is in the direction, consequently,
of establishing the parity, to claim the least, of zoology,
botany, morphology, and general physiology, with chemistry
and physics, in the great complex group of biological
science.

We may first allude to a favorite mode of expression of
materialistic elementalists. Whenever fuller analysis has
proved some group of animal phenomena not hitherto con-
nected directly with physico-chemical substances and forces,

Implications of the Theories of Nerve Action 209

to be in reality dependent on such agencies, this school is
wont to remark in substance that investigation has finally
"transferred" the phenomena from the provinces of zoology,
morphology, general physiology and the other sciences of
animal life, to physics and chemistry. Our argument puts
beyond question the logical inadequacy of such a statement.
Analysis does not by any means transfer the phenomena
from zoology, etc., into physics and chemistry. Neither
analysis nor any other agency can any more take the study
of animal phenomena away from zoology and put it into
physics and chemistry than it can take bread-making away
from the baker's art and put it into physics and chemistry.
The chemist may undoubtedly take to bread-making and find
that his new employment has much in common with his old ;
but in so far as he really succeeds at the new, he is more a
baker than a chemist. He has not transferred bread-making
to chemistry, but if anything has done just the reverse.
What analysis actually does in these cases is to extend the
bounds of physico-chemical forces and laws into zoology,
morphology, etc., and to prove that if zoological, morpholog-
ical and physiological undertakings are to move into ever
greater fullness, aid from physics and chemistry is indis-
pensable.

Thus critical examination of the reasoning of elemen-
talist biology reveals the logical fallacy in any sort of state-
ment which involves the assumption that the older sciences
of organic beings, like taxonomic botany and zoology, geo-
graphical distribution, morphology, general physiology and
so on, are not and never can be relegated to places of minor
or secondary importance in biology. But it is the practical
harmfidness of such assumptions rather than the logical
fallacies underlying them which chiefly concern us in this
volume, and no part of our whole subject is more vitally
affected by such harmfulness than this of the behavior, even
the purely tropistic behavior, of animals. The whole round

210 The Unity of the Organism

of animal biology — attitude toward research problems and
undertakings, valuations and interest in different fields of
knowledge, educational aims and method — all are affected.

In the light, for instance, of such a complex of animal
behavior as that presented by the northern fur seal, its
mating, breeding, migrating and other habits, the conten-
tion that the myriads of complicated phenomena which go
to make up animal life can be understood by converting
zoology into a laboratory and experimental science, to the
end of analyzing the phenomena into their "simple elemen-
tary components," is so ludicrous as hardly to need argu-
mentative refutation. Indeed, it seems as though persons
obsessed by a theory to the extent of being impervious to
the ludicrousness of the contention, are likely to be also
impervious to the true reasoning involved in it. It may, I
think, be assumed that so much of zoology as has formed
this remarkable conception of itself will before long drop
into the background by scientific gravitation despite its
present great vogue.

One of the leading motives, consequently, of this con-
structive part of my enterprise is to establish the essen-
tiality of general zoology and its time-sanctioned depart-
ments on so solid a basis of philosophic reasoning that the
necessary methodology of the regenerated science of the
future will be clearly seen in broad outline.

If the considerations inadequately presented in these last
pages and in other parts of this volume once get secure
lodgment in biological thought it will become manifest that
the "behavior" of any animal species (as of the fur seal,
to take at random any one of thousands of species that
would illustrate the point quite as well) can mean nothing
less to a really scientific biology than the whole series of
activities of at least one individual animal from its birth to
its natural death. Consequently an "understanding of the
complicated phenomena" thus presented can not be secured

Implications of the Theories of Nerve Action

by any amount whatever of analysis, but only through an
endless series of never-divorced analyses and syntheses, this
series running on through years of effort by scores of
investigators in the field (on the Pribilof Islands, on the
Behring Sea and far down into the North Pacific Ocean)
and in the laboratory, by general zoologists, mammalogists,
anatomists, embryologists, physiologists, comparative psy-
chologists, bio-chemists and physical chemists.

How, let one ask himself, would the resolving of a fur
seal's behavior into reflexes as the simple elementary com-
ponents of that behavior, contribute to an understanding of
the annual oceanic migration of the animal or the fighting
of the males for the females, if that analysis accomplished
nothing beyond proving a certain measure of identity be-
tween the reflexes of a fur seal and, for instance, those of
an earthworm? Or how would the understanding sought
be enhanced by carrying the analysis to a still deeper level
— to the physico-chemical level — and discovering just how
oxygen, or some particular proteid substance, participates
in the reflex? We must never lose sight of what the biolo-
gist's task is as regards understanding. It is to investigate
for the purpose of understanding the facts presented by
living beings in nature. The behavior of the fur seal as the
animal lives its normal life is what is to be studied in the
case chosen, and one of the most characteristic things in
this particular behavior is a yearly journey of several thou-
sand miles through the Pacific Ocean. That is one of the
phenomena to be studied and understood, and no amount of
analysis resulting in discoveries which do not apply par-
ticularly to that particular phenomenon can be admitted
as an explanation of that phenomenon. As compared with
any of the other natural sciences, biology is preeminently
the science of individuals — or natural objects which though
alike in innumerable attributes are unlike in innumerable
other attributes, and in no aspect of organisms do the dif-

212 The Unity of the Organism

ferentials which make individuals, species, genera and so
on come out so importantly as in behavior.

The evidence being now overwhelming that all organic
phenomena, including behavior, are dependent upon physico-
chemical substances and forces, one of the most pressing ques-
tions of procedure in biological research is that of bringing
the older and, generally, less exact natural history aspects of
the science into closer, more vital cooperation with its newer
experimental and more quantitatively exact aspects. Specif-
ically stated, work of the type long prosecuted by explor-
ing expeditions, botanical and zoological gardens, museums,
botanical, zoological and biological societies, and govern-
ment biological surveys; and that of laboratories in the
strict modern sense, the morphological, physiological, and
bio-chemical laboratories, must join hands more closely and
effectively than they have heretofore to' insure continued
progress in the organic sciences. Several movements of the
day in biology could be mentioned whose meaning, viewed
from our standpoint, can hardly be mistaken. Perhaps the
most conspicuous of these is that congeries of research ac-
tivities known as ecology. In spite of frequent deprecia-
tive comments about ecology, especially because of its in-
definiteness as to both content and delimitation, it has the
merit — from our standpoint the very great merit — of facing
organic nature as it actually is, that is, of having for its
subject matter the modes of life of organisms as nature
presents them, and hence of recognizing the laboratory as
an agency, but only as one among other agencies, for deal-
ing with its subject. As to method, while ecology recognizes
the indispensability of the laboratory and experimentation
in the narrow sense, it refuses to let such experimentation
usurp the whole of its interest and effort.

So our study of the organism's integratedness as exem-
plified by its activities, that is, by its behavior, and by the
mechanism through which these activities are carried on,

Implications of the Theories of Nerve Action 213

leads to the somewhat unexpected though entirely natural
result which may be summarily stated thus: To gain un-
derstanding of the behavior of living beings is admitted by
everybody to be the chief reason for investigating such ac-
tivities. Due consideration of the nature of the activities
and of the nature of understanding makes it certain that the
phenomena themselves are highly integrative and integrated,
or synthetic, and that understanding of them depends as
much on synthetic knowledge-getting as on analytic knowl-
edge-getting. Perception of this last truth necessitates,
again, a sort of synthesis, or integration, of the numerous
research agencies.

REFERENCE INDEX

1. Loeb ('16) 285 9. Loeb ('02) 94

2. Loeb ('12) 70 10. Loeb ('02) 101

3. Loeb ('16) 284 11. Schrader 177

4. Loeb ('16> 257 12. Holmes ('06) 313

5. Loeb ('02) 7 13. Holmes ('06) 315

6. Friedlander 363 14. Luciani iii, 355

7. Loeb ('02) 85 15. Loeb ('02) 5

8. Loeb ('02) 91

Chapter XXII

PSYCHIC INTEGRATION

Preliminary Remarks

(a) Absolute Discrimination Between Reflex and Psychical
Phenomena Not Necessary

I T will be recalled that in our discussion of neural inte-
•*• gration we limited ourselves strictly to those manifesta-
tions and activities of organisms which are, so far as ob-
servation can determine, strictly reflex, that is, show no
evidence of intelligence and volition, or even necessarily of
instinct. What we have to do next is to consider the unity,
the integratedness of the animal organism as manifested in
the vast array of its activities which by universal consent
are designated by such terms as instinctive, emotional, vo-
litional, conscious and intelligent.

Be it noted that this aim will not exact of us, any more
than did the last, a sharp delimitation between reflex or
purely mechanical acts and psychical or conscious acts.
Just as in the former discussion we were concerned with the
integrative character of those acts which are indubitably
reflex, so here our object is to study the integrative or syn-
thetic character of those acts which are indubitably psychi-
cal. We shall now be dealing with acts which have unques-
tionable psychical attributes, that is, show something of
individual plasticity and something of correspondence to
individual needs; which are, in other words, to some degree
individually determined to meet individual requirements
either of external or internal imposition and intention.

214

Psychic Integration 215

(b) The Organism an Original Datum in All Problems of
Psychic Life

Another preliminary remark of high importance concerns
the question of what, precisely, it is with which we have to
do — of what we start from and what is ever in sight, in the
discussion. Our fidelity to the organism, living in its natural
setting, as the foremost objective reality in this treatise,
prevents us ab initio from being satisfied with a Body, and
a Mind or Soul, as these have figured from time immemorial
in discourse about the higher animals, particularly about
man.

If in all the world there is such a thing as objective
truth, what we start with and have ever to deal with in
studying psychic phenomena, just as in studying all other
phenomena of animals, are individual objects or bodies of
very particular construction and activity. And by no pos-
sibility can consistent thought and statement avoid acknowl-
edging that that vast assemblage of acts and other manifes-
tations which are called psychical are yet only part and
parcel of the still vaster assemblage of acts and manifesta-
tions presented by the very same living objects, that is, by or-
ganisms. Our occupation will be basally with an object, some
particular organism, having innumerable attributes, which
being classified fall rather roughly into two great groups,
one of which we name physical or material and the other
psychical or spiritual. For short, the physical or material
•group is called the Body, while the psychical or spiritual
group is called the Soul or Mind.

Our discussion, then, will never lose sight of the fact that
the acts with which we deal are acts of the organism and not
of any of its parts merely, whether these be conceived as
material or psychical. No matter how far particular acts
may be dependent upon, and so explicable by, particular
parts, this dependence can not in reality be the whole story,

216 The Unity of the Organism

for the sufficient reason that the parts are, finally, non-
existent except as derivatives of and dependencies upon the
organism, as our whole treatise has abundantly shown.

Discussion of the Nervous System, the Brain, the Cerebral
Cortex, Neurones, Reflexes, the Senses, Responses, Emo-
tions, Consciousness, Will, Reason, and so on, as though any
of these are now or ever have been or ever will be independent
entities, or things to which the organism is subordinate, is
from our standpoint one of the deep inadequacies and mis-
fortunes of much biological and psychological thinking. To
the whole attitude of the zoological naturalist, who by na-
tive endowment and by training is imbued with the spirit
of his motto "neglect nothing" in the study of animals, this
habit of the special sciences of animal life is intolerable.

The ancient problem of the relation of "Mind" to "Body"
is one of those problems which run on endlessly in discussion
simply because the partisans of one theory or another never
know exactly what they are discussing — never know just
where they start from, in what direction they are going, or
what the end would be like if they reached it.

Lest this statement be taken as foreshadowing both a
right statement and a "final solution" of the problem, I
affirm very positively that it foreshadows nothing of the sort.
All I hope to do is to add considerably to a clear statement
of the problem, to add a little to our comprehension of
whither we are going, and to contribute a bit to the "final
solution," whatever that may mean.

The real problem of psychic integration formulates itself,
for us, in a two-parted way: Given any particular act or
action-system* which is unquestionably psychical, (1) how
many and what parts of the organism are essentially in-
volved in it? and (2) does the act or action-system bear such

* This phrase I borrow from Jennings (Behavior of the Lower Organ-
isms, p. 107) and mention incidentally here that we shall find it ex-
tremely useful later on.

Psychic Integration 217

relation to other acts or action-systems and to material
parts of the organism as to warrant the ascription to these
acts of causal influence on other acts and on the material
parts ?

(c) Provisional Classification of Psychical Facts

A third and final introductory remark touches on the
question of what shall be recognized as contained in the
organism's system of psychic attributes. Following our
regular custom of beginning with the phenomenon under con-
templation at its fullest, most indubitable expression, we
shall not go far amiss if we accept the time-honored trium-
virate of feeling, will, and intellect as the most obvious sub-
groups of highest psychic attributes ; for only a hopelessly
sophisticated philosophy and psychology can hesitate to
acknowledge that every full-grown, normal, civilized human
organism, at least, is at once a sort of reservoir of feeling,
sentiment, and emotion ; a dynamo of resolution and exe-
cution ; and a granary of intelligence and reason. ( See, for
example, Thinking, Feeling, Doing, by E. W. Scripture.)

Perhaps the only thing that needs saying about these
sub-systems of mind is that our general standpoint aligns
us squarely with the tendency in present-day psychology to
accept them for what they actually are, striving to become
acquainted with them and to assess their importance on this
basis. To ascertain first of all the facts on the psychic side
of the living animal, then next to interpret, to correlate, to
explain these facts, are cardinal principles of procedure in
our enterprise. For one thing, as an evolutional zoologist
of many years' practice in speculating on how animal parts
originated (even those of almost infinite simplicity as com-
pared with the mind of man), I am too familiar with the
limitations and pitfalls of the genetic method to be be-
guiled into making some one theory of the origin of mind

The Unity of the Organism

the corner-stone of my interpretation.*

Furthermore, as a naturalist faithful to the mandate "neg-
lect nothing," I am in full accord with psychology's aban-
donment of the earlier supposition that a leading aim of
psychology must be to prove that some one psychic province
is all-dominant, the others being merely secondary and trib-
utary.

Thus all forms of the theory that the psychic empire is
at bottom Intellect, the heaven-ordained monarch of which
is Unconditioned Ideation, are incompatible with our stand-
point. Something of the weight and variety of authorita-
tive sanction which are pitted against us here is indicated
by such names as Descartes, Locke, Leibnitz, Hume, Kant,
and Herbart of the proximate past, and Wundt, Royce,
Howison and Bradley of the immediate past.

And theories like those of Fichte, Schopenhauer, Hart-
mann, Nietzsche, and divers pale, fitful present-day lights,
which would accord to Will hegemony over the entire psychic
realm, are still less tolerable to us.

Feeling has won its rightful place in psychology so recently
that there seems little danger of its pushing its claims to
position and power beyond reason. Except perhaps in the
form of sensationalism, or the theory that all knowledge ac-
tually does originate in sensations, psychology of the west-
ern world appears never to have attempted seriously the
deification f of Feeling as it has of Reason and Will. When

* I count it as one of the pieces of good luck in my scientific career
that, through no merit of my own, a technical memoir of mine containing
an elaborate theory of the origin of the vertebral column has lain in
editorial keeping unpublished for a decade and a half.

t It is possible, as my friend Professor G. M. Stratton suggests to me,
that the German philosopher Fr. H. Jacobi, came nearer doing this than
any one else. His teachings gave, however, a definite place to positive
knowledge, so that, according to Hoffding, his faith and his knowledge
constituted two distinct philosophies. What he wrote probably does not,
consequently, contradict the statement in the text. Jacobi seems not to
have exerted much influence on the main current of German philosophy
and life.

Psychic Integration 219

Thomas Hobbes identified imagination with fancy, "orig-
inal fancy" with sen'se, and "decaying sense" with memory,
and held sense to be caused by "so many several motions of
the matter, by which it presseth our organs diversely," and
when he defended the general doctrine that "there is no
conception in a man's mind which hath not at first, totally
or by parts, been begotten upon the organs of sense," 1 he
did indeed blaze a trail which might easily lead to an over-
exaltation of the sensuous and emotional side of life. But
the eminently practical character of Hobbes' undertaking
being remembered (he was writing not for the love of specu-
lation but to save his country from political chaos and mis-
ery begotten as he believed from false theories and impossible
desires and ambitions) one may expect to find in him ele-
ments of steadiness and restraint which would make for safety
in speculation.

One such element is clearly seen in his opinion that he who
is "born a man" and lives "with the use of his five senses"
has all the native equipment necessary to realize the best in
him both for himself and for his country.2 The necessity
of being "born a man" is the point to be especially noticed.
That, with all it may imply, is on a par with the necessity
of using the five senses. So whatever of scientific hobby-
riding under such captions as "sensationalism," "empiric-
ism," "associationalism," may have followed in the wake of
Hobbes' writings, Hobbes himself, I am quite sure, was at
heart a genuine organismalist and is entitled to high es-
teem as one of the very first moderns to speak strongly for
the importance of the body generally to the psychic life of
man. Listen to this:

"Natural sense and imagination are not subject to ab-
surdity. Nature itself cannot err; and as men abound in
copiousness of language, so they become more wise, or more
mad than ordinary." "Between true science and erroneous
doctrines, ignorance is in the middle."

220 The Unity of the Organism

Having completed a reconnoissance of the field in which
we are to work, of its expanse, its contents, and its main
subdivisions, we are prepared to. take up the task proper.
Approaching it from our standpoint, one naturally sur-
mises that between the organism's neural unity as manifested
by its reflexes studied in the last chapter, and its psychical
unity known to psychology and to be considered presently,
a vital unity of still higher order exists. By unity I mean a
unity so intimate, so reciprocating, so mutually constitutive
that the term parallelism, with the meaning given it in much
of recent psychology, is wofully inadequate for it. Such a
unity, if it exists, must be sought by inspecting the entire
gamut of psychic life, from the simplest responses to stim-
uli on up through simple reflex responses, the tropisms, the
primal affective and emotive responses, through the perceiv-
ing, the imagining, the conceiving, the reasoning operations,
to the very highest constructive human mental achievings.

Likeness Between Tropistic and Higher Psychic Activity

An important move, starting from the highest phase of
rational mind, has been made toward recognizing the nature
of this unity. This move is the more significant for our
enterprise in that the investigator who has made it is neither
an elementalist nor an organismalist, but an eminent sub-
jective idealist. Josiah Royce is the student who has per-
formed this service. Stated in a single sentence, the ad-
vance he has made toward discovering the union consists
in the recognition of certain fundamental resemblances be-
tween some of the very highest operations of man's mind
and the pure tropistic operations of lower animals.

Royce's contribution to this subject is contained in his
Outlines of Psychology, and nowhere else so far as I know.
From the preface of this book I gather the following, partly
by way of quotation and partly from obvious inference.

Psychic Integration 221

"To my mind," says Royce, "an interesting side-light has
been shed upon the well-known controversies between the
associationists on the one hand, and the school of Wundt
and the partisans of 'mental activities' generally, on the
other, by the stress that Professor Loeb has recently laid
upon the part that what he calls 'tropisms' play in the life
of animals of all grades." 4

Then after telling in a few sentences what tropisms are,
Royce continues : "Now it is especially notable that the 'tro-
pisms' of Loeb are not, like the 'reflex actions' of the theories,
modes of activity primarily determined by the functions of
specific nerve-centres. Furthermore, they are more general
and elemental in their character than are any of the acquired
habits of an organism." (At this point Royce takes up, for
a moment, a matter to one side of my main purpose, namely
the problem of "self-activity" and "spontaneity"; so I ven-
ture to change somewhat the order and emphasis of his ar-
gument.) "Now it has occurred to me to maintain, in sub-
stance, that the factor in mental life which Wundt's school
defines as 'Apperception' . , . may well be treated, from the
purely psychological point of view, as the conscious aspect
or accompaniment of a collection of tendencies of the type
which Loeb has called 'tropisms.' " °

Then we have: "Wundt has insisted that his 'Appercep-
tion' is no disembodied spiritual entity. I conceive that Loeb
has indicated to us, in the concept of the 'tropism,' how
a power more or less directive of the course of our asso-
ciations, and more general than is any of the tendencies
that are due, in us, to habit, or to specific experience, can
find its embodiment in the most elemental activities of our
organism." (

What, now, is the bearing of this idea of Royce's on the
main theme of this chapter, the organism's unity as mani-
fested in and influenced by its psychic life? As an initial
step toward answering this question, the reader is asked to

222 The Unity of the Organism

recur to the chapter on tropistic activities and their an-
atomical groundwork, recalling that it was the special aim
of our discussion to show the inevitable organismal trend of
the whole doctrine of tropisms. It should be remembered
also that tropisms are, all of them, probably, beyond ques-
tion adaptive in fundamental nature; i.e., they work in the
interest of either the individual as a whole or of the species
to which the individual belongs. The circumstance that
under occasional more or less artificial conditions the ac-
tivities of an animal may result in injury to it or even in its
death, is not proof that on the whole those activities are not
advantageous. A horse or man may make himself sick now
and then by taking the wrong kind of food or too much
food, but this does not prove eating to be useless on the
whole, or non-adaptive.

Another important thing to bear in mind about tropisms
is their automaticity, or preferably their intrinsicality.
They are rooted in and partake of the very essence of the or-
ganism— so much so that they manifest themselves inevitably
when the right external and internal conditions are present,
whether the general ends which they normally serve are at-
tained in the particular instance or not. The flight of the
moth toward the flame, even at the sacrifice of its life some-
times, is a manifestation of a tendency that wqrks, on the
whole, for the good of the animal. That the moth follows
the impulse even to death merely shows how tremendously
deep-seated this type of reaction is. That the activity may
result in injury or death in a special case is just because the
case is special, i.e., it is a departure from the regular con-
ditions under which the reaction-type became incorporated
in the organization of the creature. Being always poised
for a particular kind of action, and having a supply of en-
ergy to execute the action, are unquestionably among the
most distinctive attributes of animal organisms. Such or-
ganisms are distinguished from plant organisms not only

Psychic Integration 223

by the present fact of inherent activity of the animal, but
by their inherent preparedness for acting to meet new and
more or less unusual situations. This action and action-
readiness are the real meaning of the neuro-muscular system.
All biotic organization is anticipatory in various ways, but
animals are almost exclusively anticipatory in action.

It is just these attributes that Royce recognizes as com-
mon ground between certain of the highest psychic activi-
ties of man and tropistic activities. With this overplus,
and in some cases useless or even injurious activity (in-
stanced by the flight of the moth toward and around the
flame), let us now pass to the upper end of the gamut of
animal activity for illustrations. A very few must suffice.
The first chosen is one of exalted creativeness in art.

From the vast domain of art a more instructive illustra-
tion of over-wealth of self-activity can hardly be found than
is afforded by William Shakespeare. A recent investigation
of his works undertaken with a view to finding what they
tell about the "native endowments of the author" and prose-
cuted with that love for accurate, exhaustive knowledge
which is the very soul of "modern science, leads to the result
that of these endowments "the most outstanding perhaps is
his exuberant vitality." This characteristic of the man is
exhibited in the "reckless volubility of almost every char-
acter, the piling up of fancy upon fancy, of jest upon jest,
the long embellishment of humor and foolery and horseplay
for no other reason than the delight they afford." 7 And
incidentally, the strict individualism of this sort of thing is
exemplified by one of these same Shakespearian characters:
"Come, come," says Mercutio to Benvolio, "thou art as hot
a Jack in thy mood as any in Italy. . . . Nay, an there
were two such, we should have none shortly, for one would
kill the other." "What has Queen Mab to do with the ac-
tion of the play of Romeo and Juliet? Nothing; but Mer-
cutio mentions her, and before any one can stop him he has

The Unity of the Organism

poured forth fifty lines of purest fantasy. . . . Whole
scenes," this student declares, "exist for no other reason than
that the author's brain is teeming with situations and humor
and infinite jest." 8

Now I hear in imagination expressions of astonishment,
rising to protest, even to ridicule, on the part of some biol-
ogists, and to horror on the part of some literateurs, at the
idea of suggesting that there is anything really in common
between the two groups of phenomena here placed side by
side — the activities of a moth and of Shakespeare ! For the
moment I do no more in reply than ask the reader to take
cognizance of the fact that the whole training and occupa-
tion of the naturalist consist largely in comparing all sorts
of things, inorganic as well as organic, which to cursory
observation seem unlike, for the purpose of finding whether
closer and broader examination can not discover resem-
blances and affinities which may throw light on the ever-
insistent problem of origin and causal relationship. From
that procedure, and that alone, initially, came the theory of
organic evolution. It is the quintessence of the organic
method. To him who is so instructed and disciplined that
the recognition of likeness and kinship between, for example,
the prothallus of the fern and the flowering plant, or between
a horse's fore-foot and a man's hand, will receive no shock
from the comparison. The intrinsic justification of the
comparison will be deferred until we have a few other illus-
trations before us. Another illustration will be taken from
an author, J. J. Rousseau, whose activities stand about mid-
way between art proper and science proper.

"I felt," Rousseau says in his Confessions, "that writing
for bread would soon have exhausted my genius, etc.,"
Again: "Nothing vigorous or great can come from a pen
totally venal." And finally: "In a severe winter, in the
month of February, and in the situation I have described, I
went every day, morning, and evening, to pass a couple of

Psychic Integration 225

hours in an open alcove which was at the bottom of the
garden. ... It was in this place, then, exposed to freezing
cold that without being sheltered from the wind, I composed,
in the space pf three weeks, my letter to D'Alembert on
theatres."

If this sort of thing, one may note in passing, is a case
of "struggle for existence," the existence struggled for is
on the highest plane of psychic life and not on the plane
of mere brute continuance.

The only other example will be one of activity in science
proper, i.e., "pure intellect" as far as there is such a thing.

The case of some man devoting the best of his life to the
working out of a great germal idea — an Aristotle, a Coper-
nicus, a Galileo, a Kant, a Darwin will serve our purpose
best. Of these we choose the case of Darwin. Consider first
the youth and the young man keenly alive to the flood of
sense impressions pouring in upon him from external nature,
and mentally — "internally" — "restless," as Royce would
say, from an undefined though strong dissatisfaction with
the stereotyped school and university curricula and modes
of dealing with subjects. Later comes the set of environ-
mental influences (chiefly through the naturalist Henley),
quite incidental to his regular, prescribed environment, to
which he responds with eagerness and effectiveness — an al-
most automatic choosing of fields of intellectual activity.
Out of ajl these fragmentary and by-the-way experiences —
"contents of consciousness" — there is organized a body of
natural knowledge, and such definiteness and promise of ten-
dency as to justify an appointment to a post of consider-
able responsibility and unique opportunity, that of natural-
ist to H. H. Exploring Ship Beagle.

During the voyage and from the new and strange con-
tacts with nature afforded by it, there arises another state
of "restlessness," this time concerning the origin of organic
species, the "mystery of mysteries," as Darwin himself put

226 The Unity of the Organism

it. A matter deserving special notice is that the truly for-
ward, the creative step came after, and was conditioned
upon, a period of dissatisfaction with the prevalent teaching
on the subject. Then the considerable time of semi-con-
structive observation and thinking and feeling under guid-
ance of the general surmise that species arise naturally and
not supernaturally, as all his earlier experiences — "contents
of consciousness" — had taken for granted. And at last the
final, for him, great conception, the hypothesis of the "strug-
gle for existence" and "survival of the fittest" as a cause of
the transformation of species. The suddenness and spon-
taneousness with which this idea emerged into consciousness
should be specially noticed. Once the merest suggestion of
it hove in sight, the whole hypothesis formed itself, organ-
ized itself, rapidly and completely.

The sense in which the process may justly be called spon-
taneous is important. Although we well know that the
famous hypothesis was suggested by the reading of Malthus'
work on population, we know equally well that the most es-
sential features of the hypothesis were not contained in the
teachings of Malthus. There was something genuinely new
in the hypothesis. Out of the former total of experiences
came that which did not actually exist in those experiences.
Although the hypothesis was clearly a product of something
which went before, it was a synthetic product in the strictest
sense, in essentially the sense that a chemical compound is a
synthetic product of its interacting elements, the sense that
the most distinctive attributes of the compound can not be
found in the elements taken separately, but only after the
interaction has actually occurred.

We must not fail to consider the long period of Darwin's
strict "self-activity" in collecting evidence, pro and con,
bearing on his hypothesis ; and the activity designed, notice,
to ascertain whether or not there is a process going on in
the outer world of plants and animals corresponding to the

Psychic Integration 227

process he had conceived, i.e., had pictured in his "inner
world" of consciousness. The genuineness of the individual,
the personal, the unique character of mental life and mental
creation can hardly be more strikingly illustrated than by
such cases as this of Darwin's when the conception, the hy-
pothesis, is kept to one's self so long in order "to prove"
whether it is "true" or not.

Now I want to call particular attention to the indubitable
fact that these illustrations are only extreme manifesta-
tions of attributes which are universal in the human animal
at least. There is no normal human known to anthropology
which has not some measure, no matter how small, of creative
impulse in art and in science.

As a conclusion to this presentation of instances I must
again insist upon one of my cardinal points: that the in-
dividually active and creative power of the human organism
on its psychical side is not a whit less real, less objective, less
a natural phenomenon to the natural historian than is the
individually creative power of physical growth and variation,
and reflex and tropistic action. Indeed, the thorough-going,
consistent zoological naturalist, the substance of whose
science is largely animal behavior in all its aspects, can not
possibly approve the effort to separate completely the two
sorts of creation.

First Move Toward Showing the Organismal Character of
the Higher Psychic Life

Now for the further scrutiny of such psychical facts as
those typified by the examples presented, for the pur-
pose of seeing what has been done and may yet be done to-
ward bringing them into accord with the organismal con-
ception, the pole star of all our previous discussions. This
examination will begin, as others have begun, by showing
how elementalistic attempts to interpret organic phenom-

228 The Unity of the Organism

ena soon reveal their inadequacy and finally break down as
the efforts come to face the increasing complexity which
progress of objective research always finds in such phenom-
ena.

Associationist Psychology a Special Case of Elementalist

Biology

In the particular psychical realm we are now to examine,
elementalist theory has appeared most prominently as what
is called Associationism. This flourished first in England as
the school of English Associationists, David Hartley, near
the middle of the eighteenth century, being usually consid-
ered its founder. Psychologists of this group hold that
ideas, which for them appear to be identical with sensations,
are the "ultimate elements" of psychic phenomena. "Ac-
cording to this theory, rigidly carried out, all genesis of new
products is due to the combination of pre-existing ele-
ments." Even the passions, according to Hartley "must
be aggregates of the ideas, or traces of the sensible pleas-
ures and pains ; which ideas make up, by their number and
mutual influence upon one another, for the faintness and
transitory nature of each singly taken." 9 The "piling up
of fancy upon fancy, of jest upon jest, the long embel-
lishment of humor and foolery and horseplay" which Pro-
fessor Manly shows characterize many of the Shakespearian
plays, would be explained, according to this kind of psy-
chology, not really by the author Shakespeare but by the
"aggregation," in some way, within him of ideas.

And, similarly, the works which in popular language are
said to be by a Darwin, a Humboldt, a Copernicus, an Aris-
totle, are in reality not by but merely m these men. The men
were only the places of aggregation of the elements — the
ultimates — by which the teaching on the .origin of species,
on the general character of the earth, on the solar system,

Psychic Integration

on the deeper meaning of external nature, were produced.
Again the old story with which we have become familiar: not
the organism, but elements of, or perhaps merely in it, are
the causal explanation of whatever occurrences are associ-
ated with the organism. It is, I think, safe to assume that
both the merits and the demerits of associationist psychol-
ogy have been made patent enough, at least to English-
speaking students, by the writings of James and others.

If only the doctrine of "association of ideas" can be satis-
fied to do what it is really able to do, and not insist upon
trying to do what it can not do, its usefulness is great and
its permanence in psychology assured.

As indicated above, the "huge error," as James expresses
it, by which the "whole historic doctrine of psychological
association is tainted" 10 is only another miscarriage of the
elementalist mode of reasoning, and so is subject to the
general type of criticism which the reader has met in every
chapter in this book.

In order to divest the criticism as much as possible of
personal flavor I shall make large use of James' language.
"All these writers," says James, referring to Hobbes, Hume,
Priestley, Hodgson and the later English associationists,
"hold more or less explicitly to the notion of atomistic 'ideas'
which occur. In Germany, the same mythological suppo-
sition has been more radically grasped, and carried out to
a still more logical, if more repulsive, extreme, by Herbart
and his followers, who until recently may be said to have
reigned supreme in their native country."

Now the objection to the doctrine of "atomistic ideas"
does not so much concern the conception of ideas as atoms
as the nature attributed to these atoms, namely in assum-
ing them to be immutable, and sufficient in their isolate
capacities to account for the thought and other products
arising from their "association." The following two quota-
tions illustrate the form this criticism, the essence of which

230 Tlie Unity of the Organism

is now very familiar to us as biologists, takes when it ap-
pears in garments of a psychologist's making. The "huge
error" of the association doctrine, mentioned above, James
explains, is "that of the construction of our thoughts out
of the compounding of themselves together of immutable
and incessantly recurring 'simple ideas.' " 10

If there be any doubt as to the meaning of this surpris-
ingly un-James-like wording, there certainly can not be as
to the following: "For Herbart each idea is a permanently
existing entity, the entrance whereof into consciousness is
but an accidental determination of its being. So far as it
succeeds in occupying the theatre of consciousness, it crowds
out another idea previously there. . . . The ingenuity with
which most special cases of association are formulated in
this mechanical language of struggle and inhibition, is great,
and surpasses in analytic thoroughness anything that has
been done by the British school. This, however, is a doubt-
ful merit, in a case where the elements dealt with are arti-
ficial ; and I must confess that to my mind there is something
almost hideous in the glib Herbartian jargon about Vorstel-
lungsmassen and their Hemmungen and Hemmungssummen,
and sinken and erheben and schwehen, and Verschmelzungen
and C ample xionen." ll

The long and short of the "huge error" of associationist
psychology is that ideas are no such independent, immutable,
simple entities as the doctrine supposes ; that in their origin
and in all they are, and all they do, and all that comes forth
from their association, they are in some sort and measure
dependent upon — what? Something. Search after this
something has been a large motive of more recent psycho-
logical inquiry.

One way of supplementing and rectifying associationist
doctrines is to epitomize the shortcoming of these doctrines
in the statement that they recognize only the objective side
of the association process, whereas the subjective side is

Psychic Integration 231

equally important. Thus Pillsbury: "It was a neglect of
the subjective conditions and the insistence upon the objec-
tive side of the problem that has led the English Associa-
tional School into disrepute. The explanations that they
gave were true as far as they went, but their incompleteness
vitiated the conclusions as soon as they laid claim to uni-
versality." 12 And the author then shows, convincingly
enough, how two sets of subjective factors, attention and
choice, play a large and important role in determining the
"associative train." And further, "A complete explanation
of association demands that both sets of factors [objective
and subjective] be taken into account; to omit either is to
fail in the solution of the problem." 13

Preliminary Examination of Objective and Subjective

There is undoubtedly a real gain in having proved, as
Pillsbury and others surely have, that a "side" other than
the objective in association does exist. It is highly ad-
vantageous, also, to have learned enough about this other
"side" to make the term subjective an appropriate name
for it. But any one coming to a study of the associational
activities of the organism's psychic life as we have, namely
as naturalists, can not avoid, if true to his traditions and
methods, wanting to know how these two "sides," the objec-
tive and the subjective, go together — what the nature is of
their relation. For the very fact that they are two sides
of one thing is to the naturalist prima facie evidence that
they are in vital organic connection. Even the two sides of
an inanimate thing, like the earth with its two hemispheres,
have a relation to each other too important for the earth
sciences to ignore or even to put off as merely "parallel."
But when it comes to an entity like a live animal, the quin-
tessence of which is organization, the question of how two
of its "sides" so important as its objective and its subjec-

232 The Unity of the Organism

tive are related, becomes most fundamental, especially when
a 'subject like that of psychical association is up for con-
sideration.

It is, then, fundamental to our enterprise to find out all
we can about the connection between the objective and sub-
jective aspects or sides of the organism. It would be folly
to expect results of any value from an effort of this sort
without having first given attention to the nature of each
of the sides. Now the objective "side" comes to much the
same thing as the physical or material organism as we are
conceiving the "sides." But since this has been the sub-
ject of our whole treatise up to the last two chapters, and
even of the greater part of these, we are already possessed
of enough understanding of this "side" for our present
purpose.

As to the subjective "side" the case is different. Into
its nature, its makeup and activities, we have looked very
little — only in a bird's-eye-view fashion thus far in the
present chapter, and into its marginal or transitional zone
in the preceding two chapters. We are, consequently,
obliged to penetrate further into the subjective realm itself
before the main problem, that of the relation between the
sides, is attacked.

The Essence of Wundtian Apperception

This carries us back to the point at which we brought
Royce's suggestion of a relation between Loeb's tropism
theory and Wundt's apperception theory into the discus-
sion, the return to this point being for the purpose of using
Wundt's conception to induct us further into the nature of
mental life. The importance of examining Wundt's con-
ception is two-fold. In the first place, we want to know
whether or not it is genuinely descriptive of man's highest
psychical life. If it is, nothing can stand in the way of its

Psychic Integration 233

acceptance, so far, by the anthropological zoologist. But in
the second place we must know whether or not it carries, a's
some critics believe it does, transcendental or supernatural-
istic implications. If this charge be true it is of course to
this extent unsanctionable from our standpoint.

Wundt's most concise characterization of apperception
which I have found is, "The process by which any content
of consciousness is brought to clear comprehension we call
apperception."

A content of consciousness is any definable experience we
may have. All consciousness whatever is consciousness of
something or other. This "something or other," no matter
what, nor whether regarded as a whole or in part, is a con-
tent of consciousness, according to my understanding.

What is most distinctive about Wundt's characterization
may be regarded as centering around the word clear. When
a particular content gets itself into the lime-light of con-
sciousness— when it becomes the center of attention — the
process by which it does so is apperception. On the other
hand, the process by which* contents, though brought into
consciousness, come only into its outer zones or edges, and
do not monopolize attention, is perception. Though this
getting of a content into clearness in consciousness, this
monopolizing of attention, may take place passively or
actively so far as the mind as a whole is concerned, the ac-
tive way seems to be the more distinctive and important, at
least for mental life as a whole.

It is apparently this positive activity of apperception,
directed toward making particular contents of conscious-
ness clear, which has brought criticism upon Wundt's con-
ception. "Wundt talks," says Pillsbury, "almost as if
there were a faculty or force of apperception, something
behind and superior to consciousness, which brings about
the change in clearness of the impressions. There is in
the brain a definite centre of apperception, and in conscious-

234 The Unity of the Organism

ness a force very closely related to will, that in and of itself
chooses certain ideas for elevation to the high places of
consciousness, and equally arbitrarily rejects others." And
then follows this in Pillsbury's criticism, which brings out
unmistakably its real purport: "It is very much like the
self-conscious unity of apperception of Kant, which gives
the final form and order to the various disconnected elements
of the mind, and is in so far something inexplicable, a factor
in experience that must be assumed without any further
discussion of its nature, origin, or laws of action." 13 A
suspicion, obviously, that the transcendentalism of Kant
broods over Wundt's theory. As to the justification of this
suspicion we need not be concerned here. Enough for
us at this point to recognize that from the standpoint of
description as natural history practices the art, or aims to
practice it, Wundt's account of the way the mind works in
a vast range of its activities seems true, and as far as it
goes is satisfactory.

Not only the matter of clearness of the contents of con-
sciousness, but their makeup as well is important. Al-
though "psychical elements" figure largely in Wundt's sys-
tem, one finds no intimation that the whole mind and its
contents can be "explained" by reducing them to "ultimate
elements" after the familiar manner of elementalist explana-
tion. "All the contents of psychical experience," Wundt
says, "are of a composite character." And it follows from
this that "psychical elements, or the absolutely simple and
irreducible components of psychical phenomena, are the
products of analysis and abstraction." 16

The two words, "analysis" and "abstraction," need par-
ticular consideration. The psychical elements found by
analysis do not exist, as such, in nature. Analysis, in this
case, is logical or thought-analysis, and not objective analy-
sis. We should do well to recall what was said in the dis-
cussion of reflexes, namely that the "simple reflex," though

Psychic Integration 235

legitimate and useful as an aid to interpreting the phenom-
ena of reflex nerve action, has no actual existence in nature.
It, like the "psychical element," is an abstraction. This is
only another way of saying that psychical elements are
what they are because they are parts of the mind as a whole,
just as we have seen over and over again physical elements
of the body are what they are because they are parts of the
body. "The specific character of a given psychical process
depends for the most part not on the nature of its elements
so much as on their union into a composite psychical com-
pound. Thus, the idea of an extended body or a rhythm, are
all specific forms of psychical experience. But their charac-
ter as such is as little determined by their sensational and af-
fective elements as are the chemical properties of a compound
body by the properties of its chemical elements. Specific char-
acter and elementary nature of psychical processes are, ac-
cordingly, two entirely different concepts." 17

We must not miss an essential point in this, that since
with psychical elements just as with chemical elements we
never know exactly what OR how much each particular ele-
ment contributes to the compound, we are obliged to con-
ceive the attributes of the compounds as pertaining to the
elements collectively even before the compounding has been
done.

So it comes about that because all contents of conscious-
ness are fundamentally composite, but are also resolvable into
components of various grades of complexity, synthesis and
analysis have a prominent place in the Wundtian system.
Of these, synthesis is the more definitive and fundamental,
since it enters into the very nature of consciousness itself.
Consciousness is, according to Wundt, the "inter-connec-
tion of psychical compounds." "It is the name for the gen-
eral synthesis of psychical processes, in which synthesis the
single compounds are marked off as more intimate combina-
tions." 18 The meaning of this is made clearer by the state-

236 The Unity of the Organism

ment that unconscious states like deep sleep, faint, and so
on, are the interruption of these interconnections.

Remarks On Analysis and Synthesis

This brings us to where we can see the important distinc-
tion between an aggregation and a synthesis — in a psychical
sense particularly — and likewise between a fragmentation
and an analysis. No mere aggregation,* as of ideas or emo-
tions, would make consciousness. Only a synthesis of con-
stituents can do that. And, contrariwise, while the mere
severance of the synthesized components produces uncon-
sciousness, an analysis of them results, not in unconscious-
ness, but in a consciousness of the constituent parts of the
contents of consciousness. The essence of consciousness is
unitariness — integratedness, in our general terminology — as
regards the contents of an individual organism's psychical
nature, so th.at whatever analytical processes the mind per-
forms must move within the bounds of its own unitariness
or integratedness. Were we to conceive the analytic opera-
tions of the mind to exceed or even quite to equal its synthetic
operations, we should have to conceive it as utterly negating
consciousness, i.e., as destroying itself. A man could ana-
lyze his own mind in an elementalist sense only by suiciding.
In other words, he could never do it, simply because he
would have killed himself by the very process of analyzing
before he had completed his job.

These remarks on the distinction between synthesis and
aggregation, and between analysis and fragmentation, are
not quite what Wundt says. They go somewhat beyond his
actual expression, but are legitimate inferences, I am quite
sure, from his discussion as a whole. And they help us toward
what we want to accomplish, namely to discover still more

* Recall our previous remarks on this subject, e.g., pp. 182 and 268,
and also the quotation from Hartley, p. 228.

Psychic Integration 237

than Royce discovered about the relation between apperccp-
tive processes, as Wundt conceives them, and the processes
known as tropisms.

Anticipating our results, and stating them in the most
general terms possible, we may say that the "apperceptive
synthesis" of Wundt, and what may be called tropistic syn-
thesis, have a common ground in the kind of synthesis which
is the very essence of that kind of organization to which the
term life is applied. To be alive is to be an organic in-
dividual; to be an organic individual is to be an indi-
vidual that perpetually synthesizes itself from substances
extraneous to itself (food, in the narrower sense, and
oxygen) ; and to be a psychically endowed individual
is to be an individual which in addition to synthesizing
a physical nature from the substances mentioned, synthesizes
a psychical nature from physical and chemical contacts and
interactions between the individual and the external world,
the physical contacts being called stimuli. Viewing the
matter thus, it is seen to be -highly probable that in its ulti-
mate essences the dependence of the psychical nature of the
organism on stimuli is connected, directly and inseparably,
with the dependence of its material nature on material nu-
triment.

We should, I think, be surprised were a demonstration
to be produced that psychic life has as little connection with
metabolic processes as the text-books of psychology would
lead one to suppose. Every modern psychologist, like every
modern biologist, accepts, unquestioningly I presume, the
conception that in some way the psychic life is no less de-
pendent on the nutritive substances and processes than is
the physical life. Yet that "some way" appears to be gen-
erally regarded as so remote and obscure as to be beyond the
reach of profitable treatment by psychological science, judg-
ing from the considerable number of standard text-books
which I have consulted on the point. In only one of these

238 The Unity of the Organism

(Elements of Physiological Psychology, by Ladd and Wood-
ward) do I find the word "metabolism" in the index.

Our task may then be restated as that of making out
more fully and clearly than Royce did the connection be-
tween apperception and tropisms, which is involved presum-
ably in the whole problem of organic synthesis from its
highest manifestations in psychic synthesis to its lowest
manifestation as metabolic synthesis ; of bringing to more
specificity the general statement made above. But such
statements are altogether too sweeping and abstract to satis-
fy scientific description and explanation in our day. A
chapter must now consequently be devoted to making them
more definite.

REFERENCE INDEX

1. Hobbes 15 10. James ('90) 1,553

2. Hobbes 21 11. James ('90) I, 603

3. Hobbes 25 12. Pillsbury 106

4. Royce ix 13. Pillsbury 112

, 5. Royce x 14. Wundt 229

6. Royce xi 15. Pillsbury 270

7. Manly 3 16. Wundt 32

8. Manly 4 17. Wundt 33

9. Baldwin I, 80 18. Wundt 223

Chapter XXIII

A Still Closer Look at the Organismal Nature of Tropisms

HAVING selected tropisms as a strategic point in our
program of search for the vital connection, if such
exists, between the physical and the psychical, we must turn
again to this subject. Our previous treatment of the tro-
pism theory brought out the essential organismal character
of the type of activity to which the term tropism has. been
applied. The result of tha"t treatment might be epitomized
by saying that in so far as the theory rests on accurate
and adequate description, it is genuinely organismal and
genuinely sound, but in so far as it rests on causal ex-
planation that is elementalistic in spirit and expression it is
genuinely unsound. Our present aim will be furthered by
illustrating this epitomized stricture on the theory in a little
different way from which we objectified our criticism in the
earlier treatment.

Every one familiar with current explanatory discussion
of tropisms must have noticed the large and free manner in
which the word substances is made use of in the explanations.
Thus, to illustrate: The larvae of certain butterflies emerge
from their winter nests under the influence of the warm
spring sunshine, crawl to the tips of the branches of some
shrub or tree, eat the buds and tender leaves there; then,
after feeding to satiety "turn tail" and crawl down the

239

240 The Unity of the Organism

branches. This rather complex and, to the insects, highly
useful performance Loeb and others have proved to consist
of a series of reflexes so interconnected as to come under
the tropistic type of activity. And Loeb, e.g., in the chap-
ter, On the Theory of Animal Instincts (Physiology of the
Brain) uses the case to good effect in support of his
contention that the traditional instinct-and-nerve-center ex-
planation of such phenomena is utterly inadequate.

So far, good. As to the straightforward presentation of
facts, Loeb's position seems unassailable. But what about
the causal explanation of the facts? What, exactly, is it
that sends the larvae up the branches? What causes the
eating activities? What makes the creatures then turn
about and finally sends them down the branches? That
several environmental factors, the warm weather, the
sunlight, the character of the plant buds and leaves, and so
on, are involved is brought out clearly enough. But what
about the factors pertaining to the larvae themselves ? The
body-shape, the skin, the sense drgans, the muscles are, as
was emphasized in the previous discussion of tropisms, freely
recognized after a fashion by the tropism theory. But
deeper still than these — what? Chemical substances "ac-
cording to requirement," in the language of the cook books.
Until the caterpillars have taken food they are positively
"heliotropic," that is, literally, are induced by sunlight to
move toward the sun, after the higher spring temperature
has caused chemical changes in their bodies essential to
such movement. But by eating to satiety the chemical
changes essential to the positive heliotropism are inhibited
and a negatively heliotropic state comes on. "We can im-
agine," writes Loeb, "that the taking up of food leads to
the destruction of the substances in the skin of the animal
which are sensitive to light, upon which substances the helio-
tropism depends, or that through the consumption of food
the action of these substances is indirectly prevented." 1 In

Organic Connection Between Physical and Psychical

the total scheme, then, actual and imagined, various "sub-
stances" are indispensable.

That the imaginary constituents constitute a very im-
portant part of the explanation is obvious. This fact is,
however, not specially objectionable. It is not if its true
character is never forgotten. But here comes the point I
wish to make focal just now. If imagination is to be given
a place at all in the argument it must have a larger place
than Loeb has accorded it. Otherwise the teachings of
evolution, i.e., the genetic continuity in biology, are tacitly
repudiated. Attention has previously been called, especially
in the chapter on the organism and its chemistry, to the deep
current of virtual anti-geneticism which runs through physi-
ology generally, and particularly through bio-chemistry.
Undoubtedly we can imagine "substances" produced and de-
stroyed in such a complex of activities as that described, to
meet exactly the needs of the larva; but can we legitimately
imagine them to be so produced and so destroyed by any
other means than just by the particular animals in question,
that is, by the organisms? Various of our discussions, but
particularly those in which the specificity of protoplasm
have been dwelt upon, constitute a decisive negative answer
to this question. No causal explanation of the requisite
"substances" imagined can stop short of the organism, alive
and normal, as an essential and "causal factor" in the phe-
nomena presented. Causal explanation of tropisms which
aims to reach a physico-chemical basis is really organismal
as well as are tropisms seen through the medium of pure
description.

The Automatic and Anticipatory Character of Tropisms and
Other Reflexes

Nor should the reader fail to note the intrinsicality, the
adaptiveness, and the anticipatoriness, of tropisms as illus-

The Unity of the Organism

trated in this example. Conscious will and choice seemingly
do not come into the operation at all. Given the right con-
ditions, internal and external, the caterpillar goes through
the concatenation of operations necessary for its existence,
willy-nilly. Moreover, the actions initiated by the warm
weather, the larvae being yet down at the base of the shrubs
or branches, have in organic prospect, as one might say, a
supply of food peculiar to the species. And this supply,
be it remarked, is several inches at least, and several min-
utes at least, away from the larva at the beginning of its
round of activities. Its future, even more obviously than
its present, existence is involved in the acts. Anticipatori-
ness is perhaps the most conspicuous attribute of the adap-
tiveness of such activities. C. Lloyd Morgan has well ex-
pressed the truth that one of the most important lessons
to be learned from the study of animal behavior "in its or-
ganic aspect" is the fact that "living cells may react to
stimuli in a manner which we perceive to be subservient to a
biological end, and yet react without conscious purpose —
that is, automatically." 2

But from our examination of the cell-theory we conclude
that "living cells" in this statement ought to read "living
organisms."

So much by way of further preparation, in the reflex and
tropistic phases of animal life, for our search after a vital
connection between the physical and the psychical. It will
now be advantageous to return to that supremely important
aspect of human psychic life already examined somewhat,
namely that of Wundtian apperception.

A Still Closer Look at the Likeness Between Higher Ra-
tional Life and Tropisms

As Royce's statement of the objections to the concep-
tion of Wundt contains several points that will be useful to

Organic Connection Between Physical and Psychical 243

us, we reproduce more of his sentences. "It has been ob-
jected to the partisans of Wundt that the term 'appercep-
tion,' as thus used, seems to signify a factor in mental life
which can be explained neither in terms of what we have
called sensitiveness, nor in terms of the law of habit. It
has also been objected that the conception of a conscious
process, engaged in influencing its own states, is a concep-
tion which confuses together metaphysical and psycholog-
ical motives. The psychologist, engaged as he is, not in
studying how Reason forms the world, but in observing and
reducing to rule the mere phenomena of human mental life as
they occur, is not interested, it has been asserted, in a power
whose influence upon mental phenomena seems to be of so
ambiguous a character as is that which the Wundtian 'ap-
perception' possesses." 3

Again: "This is the place," Royce writes, "neither to
expound nor estimate Wundt's theory. But it does here
concern us to point out that what occurs m mind whenever
we are actively attentive is attended with a feeling of rest-
lessness, which makes its dissatisfied with all those associa-
tive processes that do not tend to further our current in-
tellectual interests. On the other hand, the cerebral proc-
esses that accompany active attention are certainly such as
tend to inhibit many associative processes that would, tf
free, hmder our current intellectual interests." Mean-
while, "our active attention itself is always the expression
of interests which possess the same elemental character that
we have all along been illustrating in the foregoing para-
graphs. The attentive inventor is eager about the beau-
tiful things that he thinks of while he is trying to invent.
The attentive hostess is eager about social success. The
attentive caged animal is eager about whatever suggests a
way of escape." 4

The discussion from which these sentences are taken is
contained in a chapter near the end of the book, entitled

244 The Unity of the Organism

The Conditions of Mental Initiative, and in order that the
reader may get the full force of what Royce is talking
about, he is earnestly recommended to read the entire chap-
ter. Only thus can the "foregoing paragraphs" mentioned
be adequately appraised. But we must try, in our own
way, to get the essence of the matter. Royce's presenta-
tion is his way of insisting upon the facts of psychical life
and activity, high and low, which have given rise to the
Wundtian conception of apperception, these facts being
the indubitably initiatory, directive and selective qualities
of mind in all its grades. Furthermore, Royce dwells on
the homogeneity, as one may express it, of this intrinsicality
of mental life — its initiative, its persistence, and its selectiv-
ity— with the individual or fluctuating variations which
have played so large a part in theorizing about organic
evolution and heredity during the Darwinian era of biology.
And he goes back still further in good modern biological
fashion, and connects these variations with organic growth
itself, thus calling attention to the fact that variations of
this particular sort can not be referred to environmental
influence.

At this point we may stop, as biologists, to supplement
Royce's argument by pointing out that variations of the
sort indicated, are referable to environmental influence only
in the sense that growth is so referable. An organism's
securing and taking in of its nutritive substances are un-
doubtedly a kind of response to contact with its environ-
ment, and in that broad sense growth may be said to be
due to environmental influence. If the organism had no
nourishment, if it received no environmental influence of this
kind, it certainly would not grow. At the same time, since
the organism manages somehow to build a great variety of
tissues and organs out of one and the same supply of nour-
ishment; that is in response to one and the same "environ-
mental influence" (as we are agreeing to use the phrase

Organic Connection Between Physical and Psychical 245

here) there is no course open but to recognize that the or-
ganism is a very important, because indispensable, factor in
its own growth and differentiation. "Self differentiation,"
so far as the whole organism is concerned, is a fact than
which no other in the whole domain of biology is better es-
tablished. Indeed, self differentiation is really a special
form of self growth and surely no one would contend that
environmental influence is more than an essential factor in
the growth of an organism. To hold it to be a complete
explanation of the phenomenon would be too manifestly ab-
surd to receive serious consideration. It would be to con-
tend, in effect, that one of the processes of the organism
(its growth) is more than all the processes of the whole or-
ganism. But since most if not all Variation depends, either
directly or indirectly, upon growth, what is more natural
than that the living, growing organism should display much
self variation?

That such variations are among the most common phe-
nomena presented by organic beings, there is no shadow of
doubt to any one who views the problem broadly and crit-
ically, and with no domineering preconceptions as to what
ought to be and ought not to be ; who, in other words, views
the organic world as a natural historian, guided by the
mandate "neglect nothing," instead of as a physicist in the
mathematico-laboratory sense, guided by the mandate "neg-
lect everything which can not be made to conform to gen-
eral mathematically statable law."

These remarks about the relation of mental activity to
growth, differentiation and variation of the organism, and
to environmental influence would apply throughout, muta-
tis mutandis, to tropistic activity.

246 The Unity of the Organism

A Stitt Closer Description of tlie Subrational Moiety of
Psychic Life

And this brings us to where our final return may be made
for purely descriptive and comparative purposes, to the
subrational moiety of psychic life, the purpose of the re-
turn this time being to characterize this moiety as faith-
fully but as briefly as possible on the basis of the total re-
sults of researches in the field up to the present time. So
bulky and varied are these results that to examine them
exhaustively is hardly possible for any one person even
though he be a specialist in the field. Much less possible is
it, then, for a general zoologist to make such an examina-
tion. Nevertheless it is, I believe, possible to give an
epitome of the present state of knowledge that shall be true
in all fundamental respects and highly significant for our
enterprise.

Remarks on the Classes of Subrational Life

In giving this epitome we shall not try to maintain a sharp
distinction between reflexes, whether of the tropistic or any
other type, and instincts. To begin with, as always, when
a large and intensively cultivated domain of science is en-
tered for the purpose of extracting from it its most certain
major results, we may take it for granted that the ex-
tremists touching any portion of the field over which di-
vergence is wide and warm, are unsafe guides for the gen-
eral student. Thus, the student who enters the realm of
animal behavior for such a purpose as that for which we
are now entering it soon sees that those specialists who find
nothing but tropisms, and these of the most uncompro-
mising sort, in the activities of much of the animal king-
dom, are not the ones to whose guidance he can entrust

Organic Connection Between Physical and Psychical 247

himself, no matter how voluminous, and perhaps excellent
in quality, their experimental researches may be.

For instance, such a view as that of Bohn's, according to
which the word instinct ought to be eliminated from the
terminology of science "as a legacy of the past, the middle
ages, the theologians and the metaphysicians," 5 is so obvi-
ously unjustifiable to any well-informed zoologist as to
make him suspicious of such a writer all along the line,
especially wherever his judgment and scientific poise are im-
plicated.

This question of the reality of instincts I use to illus-
trate the peril to the general student of the unpoised spe-
cialist, because it is germane to the present discussion. In
general zoology the type of animal behavior to which the
term instinctive is applied is not less conspicuous and real,
to say the least, than is the type described as tropistic.
For an experimentalist to come out of his laboratory and
tell a broadly experienced entomologist or ornithologist, for
example, that the familiar achievements of young insects
of many species, and of numerous young birds should not
be called instinctive because (as the experimenter asserts)
they are reducible to the tropistic or perchance the simple
reflex type of reaction, may justly be characterized as sci-
entific impertinence. It is as though an embryologist, hav-
ing discovered that a bird's wing is the genetic counterpart
of a salamander's forelimb, should instruct the ornitholo-
gist that it is wrong for him to call the bird's wing a wing,
because the member may be reduced to a lower type of limb.
Unquestionably the experimental specialist often pro-
duces results which necessitate changes in the general zo-
ologist's conceptions and nomenclature. But it is not his
province to take into his own hands the revision of the
fundamental terms of zoology. Any one moderately in-
structed in the history of zoology knows that "instinct" is
a hardly less well-grounded zoological term than "birth" or

248 The Unity of the Organism

"intelligence," or many another indispensable term.

Our inquiry is not as to whether there are such things as
instincts, but how they operate and what they signify for
the animals possessing them. Perhaps of highest interest
to us is the fact that innumerable instincts, if indeed not all,
are as indubitably hereditary as are any animal endowments
whatever. This comes out especially convincingly in those
numberless cases where the instinctive operations develop
strictly pari passu with the anatomical development of the
young, there being absolutely no opportunity for them to
learn, even subconsciously.

Take as an example the crustacean Amphithoe longimana,
in which Holmes compared in detail the activities of the
newly hatched young with those of the adult. "Amphithoe
lives in tubular nests which are usually lodged among sea
weed. The nests are somewhat longer than the animal,
and are spun of a web-like material into which bits of sea
weed are often incorporated which help to conceal the oc-
cupant. In its nest Amphithoe lies in wait for prey, ready
to dart out upon any small creature which touches the ends
of its long antennae.

"The activities of the adult Amphithoe, with the excep-
tion of those concerned in reproduction, are almost ex-
actly paralleled by those of the young. I have taken the
eggs from the maternal brood pouch shortly before hatch-
ing and kept them isolated in individual dishes. For some
time after emerging from the egg the young were weak and
had imperfect control of their movements, which were jerky
and irregular. Soon the minute creatures would crawl and
swim much like the adults, and the next day they began
constructing nests which were the same shape as those formed
by their parents." Then comes a part of the description
to which the reader's special attention is called because it
brings out, partly by implication, a richness of detail in be-
havior which defies full expression, and which every care-

Organic Connection Between Physical and Psychical 249

fully observing zoologist knows to be characteristic of the
activities of nearly all animals. Especial attention is in-
vited to this, because this elusive wealth of behavior is
usually overlooked by the cursory observer on the one hand
and by the experimentalist on the other. "The attitudes in
the nest," Holmes writes, "the waving of the antennae, the
beating of the swimmerets, the restless movements of the
legs and mouth-parts, springing after food, belligerency
toward passers by, the little unobtrusive signs of timidity,
the reversal of position in the nest on the approach of
danger and the general behavior outside of the nest, were,
on the next day after hatching, almost exactly the same as
in older individuals. The only differences in behavior were
due to the feebleness of the young and their imperfect con-
trol of their movements." One never reads a description
like this by a typical experimentalist, especially if he be a
pure tropist, or by a meagerly trained zoologist ! Then the
final statement: "The young are hatched with all the in-
stincts necessary fully to equip them for the business of life.
No experience is necessary to teach them what is advan-
tageous for them to do."

The impossibility should be noticed of drawing a sharp
line in this description between instinctive and purely reflex
acts. "Reversal of position in the nest on approach of
danger" is clearly instinctive. But "beating of swimmer-
ets," and especially the "restless movements of the legs" —
are these instinctive or wholly reflex? Probably they arc
reflex, though the leg movements may well be partly in-
stinctive. A whole volume of examples as unquestionable
as this could be compiled, and all groups of animals from
mammals down to worms at least would be represented.

250 The Unity of the Organism

Four Certainties About the Adaptiveness of Subrational
Psychic Activities

Concerning the purposefulness or adaptiveness of activi-
ties of this general type, I think four things may be re-
garded as absolutely certain.

Generally Useful to Individual and to Species

First, a vast majority of them are recognizably contribu-
tory to the perpetuity of both the individual in its normal
life, and of the species. But for them neither individual
nor species would continue to exist. This is so obvious that
further remark upon it is unnecessary.

Many Useful to Species Primarily

Second, in a large number of instances particular acts
by particular individuals are in the interest of the species
primarily and of the individuals only secondarily or not at
all. This is shown most conclusively in cases like that of
several species of salmon, where the individual normally goes
through activities which secure the continuance of the spe-
cies but which end in the death of the individual. A large
and varied number of cases of this type occur, especially
among insects. But the supremacy of species over individ-
ual needs appears under various other forms. Thus almost
certainly such tropistic activities as that of the moth going
to its death or injury in the flame is of this sort. This
case may be stated in general terms thus: Owing to lack
of any ability on the part of an individual to modify its in-
herited mode of action to meet a special situation, it acts in
the old way even though the new situation, while in gen-
eral like the old, yet differs from it enough to make it peril-
ous to the individual if it acts unmodifiedly in the old racial

Organic Connection Between Physical and Psychical 251

way. The preeminently racial utility and hereditary char-
acter of instincts is certainly one of the most interesting
things about them for the present discussion.

Variability of Subrational Activities

The third certainty about reflex and instinctive activities
is that they are by no means so stereotyped and invariable
as older cursory observation or as much theorizing, espe-
cially about tropisms and instincts, has held them to be.
Darwin, in the notable chapter on Instinct in The Origm of
Species, was the first to attack seriously the notion of such
invariability in dealing with instincts. He undertook to
show that the instinctive type of activity is subject to vari-
ation just as are all other aspects of animal life.

A telling set of recent investigations under this head is
by the Peckhams. That on the solitary wasp, Ammophila
urnaria, is particularly to the point because the earlier
writers had used its habits of paralyzing caterpillars by
stinging them and storing them up as food for its young to
illustrate the undeviating and unerring character of in-.
stincts. But the extensive studies of these entomologists
led them to write: "The one preeminent, unmistakable and
ever-present fact is variability. Variability in every par-
ticular— in shape of the nest and the manner of digging
it, in the condition of the nest (whether closed or open)
when left temporarily, in the method of stinging the prey,
in the degree of malaxation, in the manner of carrying the
victim, in the way of closing the nest, and last, and most im-
portant of all, in the condition produced in the victims by
stinging." 7

No present-day authority so far as I know contends that
instincts operate in a hard-and-fast manner comparable to
the workings of any man-made machine. They are now
universally recognized to be subject to the same general

The Unity of the Organism

principles of variation to which all organic phenomena are
subject. Furthermore, under the searching investigation
and criticism of numerous workers, notably H. S. Jennings
and Ills followers, the tropism theory has been deprived, for
most biologists, of its inorganically mechanistic character.
The principles of "random movements," "avoiding reac-
tions," "trial and error," and others, are thoroughly estab-
lished and the recognition of them may be said to have so
modified the doctrine of tropisms as to make it one of or-
ganic mechanism rather than of inorganic mechanism — as it
virtually would be according to the thoroughgoing elemen-
talistic conception of it. The "mechanistic conception of
life," one may remark, has very much to commend it if only
the machines conceived are recognized to be alive. My re-
marks under this head * may be consulted by the reader
who wishes to follow this point.

What is meant by random movements is made clear by
the following: "In the earthworm and the larvae of blow-
flies which are negatively phototactic it has been shown by
the writer that movements which bring the animal toward
the light are checked or reversed and only those which hap-
pen to direct the animal away from the light are followed
up. Whatever immediate orienting tendency the light may
have in these cases is relatively unimportant as compared
with the clement of selection of favorable movements in di-
recting the animal away from the light."

Here it will be noticed that the end, beneficial to the ani-
mal, is reached through a combination of orienting reac-
tions of the rigidly tropistic type, -i.e., the type dependent
on the movement of the animal directly toward or away
from the source of light by the symmetrical plan of the
body, and a sort of reaction in which the particular body-
form and the direction of light rays are of only secondary

* See "Machines, living," in the index of The Probable Infinity of Na-
ture and Life.

Organic Connection Between Physical and Psychical 253

significance. But this latter type of activity, wholly di-
vorced from a direct-orienting reaction, and even from a
bilateral body symmetry, is of wide application among the
lower animals. It was first brought clearly to the atten-
tion of biologists by Jennings in his now well-known inves-
tigations on Paramecium and other protozoans. These in-
vestigations formed the bases of the "avoiding reaction" and
the "trial and error" conceptions now generally recognized
to be of much importance in the behavior of all animals, es-
pecially of those in which a high measure of bodily activity
occurs but in which there is little or no intelligence. Jen-
nings' lucid account of his results in the chapter Be-
havior of the Infusoria; Paramecium (Behavior of Lower
Organisms) is strongly commended to the reader.

The following paragraph must suffice for our reference
to this work. After describing the behavior of Parame-
cium, Jennings writes : "This method of behaving is per-
haps as effective a plan for meeting all sorts of conditions as
could be devised for so simple a creature. On getting into
difficulties the animal retraces its course for a distance, then
tries going ahead in various directions, till it finds one in
which there is no further ^ obstacle to its progress. In this
direction it continues. Through systematically testing the
surroundings, by swinging the anterior end in a circle, and
through performing the entire reaction repeatedly, the in-
fusorian is bound in time to find any existing egress from
the difficulties, even though it be but a narrow and tortuous
passageway." And this complex and highly useful be-
havior is performed by an organism which, so far as the
best anatomical researches have been able to determine, is
entirely devoid of a nervous system, and consists of a single
cell!

But the "trial and error" scheme here exemplified is by
no means confined to unicellular, non-nervous animals, nor
to experimentally produced conditions. That it is opera-

254 The Unity of the Organism

tive in nature, and among animals with rather highly de-
veloped nervous systems I shall illustrate by describing
briefly a performance witnessed by me some years ago.
This was the capture and engulfment of food by a nemertean
worm.*

These marine worms are of considerable size, some reach-
ing a length of many inches, even a few feet, and ranging
in thickness from less than an eighth of an inch to nearly an
inch. Externally they give the impression of being very
lowly in organization, the body being devoid of limbs or
other appendages, and without segmentation. However,
when they are examined internally a surprisingly high grade
of organization is found, the muscular, digestive, blood and
nervous systems being on a par, probably, with those of
any invertebrates below the crustaceans and insects. The
nervous system, particularly the brairi^ is relatively large,
though not differentiated into diverse ganglionic masses and
connecting strands to the extent found in jointed worms.
The creatures are poorly equipped with external sense or-
gans, there being no tentacles nor any certain olfactory or
auditory organs. And eyes, when present, are so minute
and simple as to be without power of sight in the ordinary
sense; almost certainly they are mere light-perceiving or-
gans.

The most distinctive anatomical feature of the nemerte-
ans is a very long and thin though muscular and flexible hol-
low tube situated at the anterior end of the animal, which
is usually carried stowed away in a pouch within the body.
While thus retracted the tube has some such relation to
the rest of the animal that a glove-finger would have to the

* Greatly to my regret I am unable to say what the species or even the
genus was of either the nemertean or the annelid here referred to. The
observation was made at the Shumagin Islands, Alaska, and under cir-
cumstances that rendered it quite impossible to "look up" the species.
And my knowledge of the taxonomy of these groups of worms is alto-
gether too meager to enable me to identify genera even, offhand.

Organic Connection Between Physical and Psychical 255

glove were it completely inverted into the hand of the
glove. This tube is used in the capture of prey, the ani-
mals being carnivorous and highly voracious. The mode of
employing the apparatus consists essentially in thrusting
the tube out with almost the speed of lightning, the object
being to bring the organ into contact with the prey at many
points. The lash is not used as a lasso for catching or as
a spear for piercing the prey, but for paralyzing it, prob-
ably by a toxic secretion spread over the whole surface.
The more effectually to accomplish this, the lash is shot out
at a victim again and again.

Now for the aspect of the whole operation of food-taking
which specially concerns us. Being quite sightless and
touchless in the usual sense, the lash must be used as an ex-
ploring or finding as well as a paralyzing or killing organ;
and since its great length and limberness preclude it from
being used as an ordinary tentacle is used, the finding op-
eration is accomplished by repeated out-thrustings of the
tube. In the instance witnessed the prey was an annelid
worm, a creature well provided with locomotor organs, and
a good crawler. On this account the victim-to-be was able,
in the early stages of the onset, to move out of contact with
the nemertean now and then. At such times the prey could
be relocated only by darting out the lash at random, except
as to general direction. So it resulted that many of the
thrusts missed the mark; but they were instantly repeated
with a little variation of direction, till the victim was lo-
cated again. The whole performance reminded one of the
game of blind-man's-buff, a game in which the seeker paws
around in the general vicinity, as he believes, where the one
sought was last touched.

The effectiveness of the try, try again method was at-
tested in this instance by the fact that the annelid was hit
and the paralyzing dose administered times enough to put
the annelid into so helpless a state that the nemertean was

256 The Unity of the Organism

finally able to get its mouth into contact with its prey.
Then the victim, itself but little smaller than the nemertean,
disappeared down the latter's "throat" with almost the
rapidity with which the lash was retracted into and thrust
out of its pouch. How much of this highly complex per-
formance, so eminently useful to the nemertean, was purely
reflex, how much chemotactic, and how much instinctive.''
And who will assert positively that there was no trace of
consciousness, even of intelligence, in it?

An extremely interesting line of inquiry is suggested by
cases of "trial and error" like this where at one extreme the
"errors" are not much less numerous than the successes, and,
at the other extreme, are cases in which the errors are re-
duced almost to nil. A type case of this last would be the
poise-and-spring of a cat after its prey. With little doubt
a closely graded series could be made out running through
from one extreme to the other. A cardinal interest in the
inquiry would be as to the extent to which the simple reflex,
tropistic reflex, instinct, and intelligence figure in the dif-
ferent grades. Would it not turn out that the gradual
diminution of error through the series would be, generally
speaking, concomitant with the increase of intelligence? I
suspect so.

Tendency of Subrational Activities to Excessiveness

The fourth and last certainty about reflex and instinc-
tive activities to receive attention is their tendency to ex-
cessiveness — their way of going beyond what is necessary or
even really safe for the welfare of the organism. Although
from several points of view this is one of the most impor-
tant aspects of the whole subject, it has received surpris-
ingly little attention, especially by the modern school of ex-
perimental zoology.

Probably every one who has observed animals widely and

Organic Connection Between Physical and Psychical 257

thoughtfully has been impressed with the exuberance of their
performances. That they are ever wont to overdo things,
even operations which are when done in measured fashion
absolutely essential to their existence, is matter of common
knowledge. Holmes has some comments under this head
which may fitly introduce our presentation. "With all their
wonderful adaptiveness instincts are far from ideally per-
fect. Much of Mark Twain's remarks on the futility and
imbecility, the wasted effort and labor at cross purposes
shown in the behavior of ants may easily be verified by any
observer."

A common form taken by excessiveness of action is repe-
tition. Very many, perhaps all, animals are notorious re-
peaters. A few out of the many available instances will
suffice to fix the phenomenon in mind. Some time ago a
small whale (probably a half-grown Humpback, Megaptera
versabttis) came near shore at La Jolla, California, and re-
mained in the same small area for days. While there it
went through a particular set of movements known to whal-
ers as "breaching" scores of times, each set being exactly,
so far as one could see from the shore, like every other set.
The performance consisted, in this case, of a leap out of
the water, which carried the body clear of the surface of
the sea, the direction of emergence being probably thirty
degrees from the perpendicular. During the ascent the ani-
mal turned with a characteristic twist to the left and came
down on its head and left side with a great splash. Once
back in the ocean the creature reversed the course it was
going when making the leap, returned to some distance from
where it had emerged, reversed its course again, and re-
peated the leap identically, to all appearances, even as to
the spot of emergence and direction of travel. Why so
many times the same performance in the same spot? That
is the problem which concerns us here. Even though we
conceive it to be somehow adaptive — connected in some in-

258 The Unity of the Organism

direct way possibly with feeding or reproduction or migra-
tion or some other vital function — the question still remains,
why so much of it? And to this no probable or even ra-
tional answer is forthcoming from the standpoint of adapta-
tion and utility, taking these terms in their usual meaning.

Here is another case from the mammalia, the possible
adaptive significance of which is still more remote, if any-
thing, than that of the behavior of the whale. Many indi-
vidual mice of the genus Peromyscus being used by Doctor
Sumner and Mr. Collins in their researches on heredity and
environmental influence at the Scripps Institution take to
throwing back summersaults in their cages. The more com-
mon performance consists in a run along the floor of the
wooden cage and up its side to near the top, then a quick,
strong jump backward clear across the cage, the feet being
uppermost during the first part of the leap but coming to
rights again by the time the landing is made. Here again
the question of why the mice do this seemingly useless thing
is not so interesting for the present discussion as that of
why they do it so much.

The high flight of some species of birds, the great eleva-
tions being reached by long, regular upward spirals, would
appear to come under the head of non-adaptive, superfluous
action. The sand-hill crane, Grus mexicana, may be taken
as an instance of a bird given to this habit. Surely such
flights by this species can have nothing to do with food-
getting, since in the excursions the bird is going directly
away from, instead of into, the region where its food
abounds. It eats snakes, frogs and other creeping animals,
and various seeds and roots. Nor is there any evidence that
the flights are concerned with the mating function, nor yet
with migration, though one might possibly imagine that
while on the excursions the birds learn, after a fashion, the
topography of the surrounding regions.

The high-diving and booming of the night-hawk, Cordefas

Organic Connection Between Physical and Psychical 259

lirginianus, repeated time after time in the early evening and
occasionally in midday when an approaching storm cools
the air, would seem to be another performance of the non-
adaptive sort. The suggestion that this is a courtship af-
fair can hardly stand, in view of the fact that at least as
often as otherwise the birds which do it are entirely alone.
Nor can one see how so extensive and swift a dive, with so
much noise, can be advantageous for the capture of flying
insects.

And reflect on the quantity of movement of many ani-
mals. Can any one believe that mammals and lizards run,
birds and insects fly, and fishes swim just exactly so much as
and no more than, they must in order to survive? Would
it be contended that the Golden Plover, to take a well known
case of extensive migration, would certainly succumb in the
struggle for existence on arty thing less than a journey from
the high latitudes of the northern hemisphere well into the
southern hemisphere and back, each year? There is a vast
difference between a necessity for migration to some ex-
tent and a necessity for migration of a particular quantity.
One of the great weaknesses of the natural selection theory
has been, I am very sure, its slight regard for quantity ;
quantity of need, quantity of performance', quantity of
benefit.

These examples serve to illustrate the fact that among
the higher animals at least, much muscular activity occurs
which is not at all, or only partly, adaptive. But by far the
more common occurrence of excessive activity is in connec-
tion with behavior which is more or less obviously adap-
tive. "A good thing carried to excess," in the familiar
phrase, expresses well what is in mind here.

This excessiveness of adaptive activity is naturally more
easily recognized in animals which are most easily observed
and most active generally. Thus it is from birds and in-
sects that examples can be most readily drawn.

260 The Unity of the Organism

•

Let the current view be accepted that the song of pas-
serine birds is associated adaptively with the mating func-
tion. Even so, no one who has given careful attention to
the matter can have failed to recognize that with many
species much more singing is done than actual pairing and
breeding call for. I have kept almost daily notes for sev-
eral years on the singing of the Western Meadow Lark,
Sternella magnu neglect a, in the vicinity of La Jolla. The
birds are resident the whole year through, and as they come
familiarly around my home and laboratory, the observations
can be quite full. Although the breeding time is restricted
to late February, March, April, and sometimes May, there
is not a month in the year when songs may not be heard,
most of the time in full volume. Significantly, I believe, the
song is at its ebb during some weeks just before the nesting
period begins. Nor does the singing of the males seem to
be connected in any close way with mating. The birds do
not pair off closely and permanently, even for the breeding
season. Most of the singing, which occurs chiefly in the
morning and early forenoon and again toward evening, is
done while the singer is, more commonly than otherwise,
quite alone on some telephone pole or wire. And the mode
of singing does not change at all when mating begins. An-
other interesting fact about the singing of this species is
the considerable range of temperature and light conditions
over which the song is invariable, so far as these factors are
concerned. The song may be as full and frequent on
cloudy, misty mornings as on sunny ones; and over a con-
siderable range of temperature the song is quite independent
of the particular degree marked by the thermometer.

While the song habits of this bird are undoubtedly some-
what exceptional in their looseness of correlation with mat-
ing and with environmental conditions, certain it is that
much this sort of thing is observable with several resident
species which I have observed. The house finch, Carpodacus

Organic Connection Between Physical and Psychical 261

Mexicamis, and the California towhee, Pipilo fuscus, may be
specially mentioned in this connection. The fact that do-
mesticated song birds, like the canary, may be brought to
sing almost perpetually is only an extreme manifestation
of tendency among song birds to sing in excess of any strict
utility of song.

Think of the monotonous repetition in the croaking of
frogs, the chirping of crickets, the stridulations of cicadas,
and so on! I have counted more than five hundred con-
secutive chirps of a cricket in about half an hour, with
only a little variation as to notes or intervals. And this is
surely a very moderate example of what actually occurs —
as any one can easily convince himself by listening and count-
ing almost any still night, almost anywhere where crickets
live. Probably the chirping pf crickets is employed in mat-
ing. Very well. But are the thousands of chirps uttered
by a given individual each night for many nights, the small-
est number upon which the species can survive? Even ask-
ing of the question reveals the monstrosity of a theory that
would necessitate an affirmative answer to it — as strict ad-
herence to the natural selectionist meaning of utility un-
doubtedly would.

In place of bringing forward additional instances, which
could easily be done, to show that vocal sounds and bodily
performances of various sorts more or less obviously con-
nected with mating among higher animals are produced in
excess of what the strict application of the rule of physio-
logical economy would dictate, I shall do no more than util-
ize the conclusions of two investigators who seem specially
qualified to speak on the subject, and assume that these con-
clusions would receive the sanction of all zoologists who have
given serious attention to the matter and have formed their
judgments unbiased in favor of any explanatory theory.

The first of these investigators is W. H. Hudson, who
represents a period a little antecedent to the present spe-

262 The Unity of the Organism

cially critical experimental era. I quote from his well-
known The Naturalist in La Plata, published in 1892: "I
wish now to put this question : What relation that we can
see or imagine to the passion of love and the business of
courtship have these dancing and vocal performances in
nine cases out of ten? In such cases, for instance, as that
of the scissor-tail tyrant-bird, and its pyrotechnic evening
displays, when a number of couples leave their nests, con-
taining eggs and young, to join in a wild aerial dance; the
mad exhibitions of ypecahas and ibises, and the jacanas'
beautiful display of grouped wings ; the triplet dances of
the spur-winged lapwing, to perform which two birds already
mated are compelled to call in a third to complete the set;
the harmonious duets of the oven-birds, and the duets and
choruses of nearly all the wood-hewers, and the wing-slap-
ping aerial displays of the whistling widgeons ; will it be
seriously contended that the female of this species makes
choice of the male able to administer the most vigorous and
artistic slaps? . . . There are many species in which the
male, singly or with others, practises antics or sings during
the love-season before the female; and when all such cases,
or rather those which are most striking and bizarre, are
brought together, and when it is gratuitously asserted that
the females do choose the males that show off in the best
manner or that sing best, a case for sexual selection seems
to be made out. How unfair the argument is, based on these
carefully selected cases gathered from all regions of the
globe, and often not properly reported, is seen when we turn
from the book to Nature, and closely consider the habits and
actions of all the species inhabiting any one district. We
see then that such cases as those described and made so much
of in the 'Descent of Man,' and cases like those mentioned
in this chapter, are not essentially different in character,
but are manifestations of one instinct, which appears to be
almost universal among the higher animals. The explana-

Organic Connection Between Physical and Psychical 263

tion I have to offer lies very much on the surface. . . . We
see that the inferior animals, when the conditions of life are
favorable, are subject to periodical fits of gladness, affecting
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 occasionally
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 him-
self in some extravagant way." X1

The reader is asked to note what Hudson says about pick-
ing out such evidence as will help the case for sexual selec-
tion, and saying nothing about evidence which will not help
it. Beyond question the dogma of natural selection, espe-
cially the Weismannian perversion of it, has flourished
largely on this sort of thing. Nor has natural selection alone
among biological theories had the benefit of assorted evi-
dence. Indeed the whole elementalistic mode of interpret-
ing living nature may be characterized as one whose doc-
trines depend largely upon "special privilege," to adopt a
phrase lately much used in the economic world, as to evi-
dence for their support.

The other investigator upon whom we draw is Prof. Julian
S. Huxley, whose work is that of a field zoologist imbued
with the exacting spirit of the present day. Huxley's stud-
ies are devoted to the mating habits of birds, so there can
be no question that the activities he describes are intimately
connected with reproduction. Of the numerous species
dealt with in the paper now before us, we notice first the
Great Crested Grebe. It is highly significant that in this
species mating takes place before the so-called courtship
performances begin, so this latter process can not be es-
sential to securing a mate. The female is "courted" after
she is got possession of. The courtship activities begin soon

264 The Unity of the Organism

after pairing, two entirely different sets of ceremonies be-
ing involved in the activities. One of these Huxley calls
ceremonies of mutual display, the other, ceremonies of coi-
tion. The highly elaborate mutual display performances
are fully described but can not be reproduced here. They
consist in a variety of body attitudes, head and wing and
feather movements, swimmings and divings, and call-notes,
the whole lasting some minutes. Concerning this prelim-
inary operation, Huxley writes :

"The most noticeable thing about all these ceremonies is
that they are 'self-exhausting' — they do not lead on to any-
thing further. Looked at from the physiological point of
view, they seem to me to be nothing but 'expressions of emo-
tion' : the birds act thus because they are impelled to do
so, because they enjoy it. Looked at, on the other hand,
from the evolutionary point of view, they seem to have been
developed as a bond to keep the pair together." 12

Following these preliminaries, the ceremonies of coition
take place, these being less striking, though characteristic.

Speaking of his studies on the mating habits of some of
the warblers, and referring to differences of interpretation
between himself and W. P. Pycraft, another observer in the
same field, Huxley writes : "In this, Mr. Pycraft and myself
are, I think, agreed; to both of us the 'display' of the male
Warbler is nothing but a direct expression of sexual excite-
ment, scarcely, if at all, modified by Darwinian Sexual Se-
lection— nothing but the way in which nervous disturbance
caused by sexual excitement happens to liberate itself. Gen-
eral nervous discharge will cause general muscular contrac-
tion; and something approaching this is here seen — rapid
hopping, extension and fluttering of the wings, spreading
of the tail, bristling up of the feathers on head and
throat, and utterance of a series of quick sounds. This ex-
presses a condition of readiness to pair, and doubtless to
the female comes to be a symbol of the act of pairing.

Organic Connection Between Physical and Psychical 265

Hence, as far as the female is concerned, the act of pairing
has come to depend upon this stimulus (acting of course on
a suitable internal physiological state). This is no more
strange in the bird than it is that in ourselves thoughts and
emotions of love well up at the sight of some tangible ob-
ject connected with the beloved." 13

But it is in the sex function itself that the tendency to
overdo manifests itself with greatest force. In fact, the fa-
miliar and ominous expression "sexual excesses" as applied
to the human animal, indicates very truthfully what is be-
fore us. The whole phenomenon of competing and fighting
among the males of all higher animals for possession of the
females, with its momentous consequences in dozens of ways,
may truly be said to rest back on the excessiveness of the
sex impulse and instinct. Since as a general rule the males
and females of animal species are approximately equal in
numbers, pairing off two by two, after the manner of the
population of Noah's ark, might occasion but little and mild
competition could each male and each female be satisfied
with one mate, in accordance with the allotment which the
numerical equality would make. And the pertinent question
may be raised in passing, would not such a mode of pairing
secure the perpetuation of the species quite as well as, pos-
sibly better than, the method which is so largely in vogue?

Highly suggestive seems to me in this connection, observa-
tions I have recently been able to make on the mating hab-
its of one of the California "surf perches" (Cymatogaster
aggregatus}. This is one of the numerous viviparous bony
fishes peculiar to our coast. The species under attention
lives quite normally, as far as one can see, in the aquaria of
the Scripps Institution; so what may be assumed to be its
typical habits can be observed continuously.

Strict monogamy appears to prevail in the species. At
least this is true with the specimens — three males and four
females under observation, and so far as a particular breed-

266 The Unity of the Organism

ing period is concerned. Each male begins his attentions
while his fiancee, so to speak, is heavily gravid from the pre-
vious mating (when and how accomplished we unfortunately
know nothing about beyond the fact that it must have been
before the individuals under observation were brought to
the aquarium from the sea, about six weeks before the mat-
ing began.)

In the case of one pair, the amours of the male continued
more than two weeks, the first few days of which were
before the family of young began to be born, the period of
parturition extending over three days. Although there
was no indication on the part of the other males of inten-
tions or even desires toward the spouse (as she may now
be called) of this male, he was quite pugnacious, directing his
seemingly unnecessary operations against the other females
as well as against the other males. It should be said, how-
ever, that his antipathies were considerably greater against
the males than against the other females. The other two
males took partners after much the same fashion ; but since
both of these were somewhat smaller, and fully acknowl-
edged the over-lordship of the one singled out in our account,
their performances were less clear cut.

Specially noteworthy is the character of the amours of
the male, which alone or almost alone, seems to take an in-
terest in the performance. No contact, or at least only the
slightest, of the male with the female was seen though the
fish were under observation much of the time. A peculiar
downward darting of the male first on one side then on the
other of the female, close to her but not quite touching her,
was one of the favorite manoeuvers. But various rapid
circlings about, up and down, head-on and tail-on, over and
under, and in nearly all possible ways, may be witnessed.

The full meaning of this monogamic (temporarily so, at
least), largely non-tactual type of mating we do not know
partly because we have not yet all the facts ; but I suspect

Organic Connection Between Physical and Psychical 267

it to be important. But this much is clear as to its bear-
ing upon the point uppermost in this discussion: There is
an excessiveness of activity in a variety of ways, particularly
in the driving of other females, the presence of which in the
vicinity of the mate is merely incidental and utterly harm-
less.

Obviously it is the demand, instinctive or organic or both,
for more sexual gratification than the natural numerical
scheme of the two sexes provides, and the actual necessities
of race perpetuation demand, which is largely responsible
for the contests to secure mates, so characteristic of all
higher animals. The bull fur seal must have forty or fifty
mates, instead of the one which the numerical equality of
the two sexes would naturally give him ; hence the fierce com-
bats among the males, with the result that a great majority
of the whole male population at any one time is forced to
remain outside the "harems" during the mating season. And
some such eliminative process must occur in all species where
the sexes are about equal in numbers, and where promiscuity
in pairing is practised.

Nor are the injuries and disasters which may result from
the driving power of the sex-impulse restricted to compet-
ing individuals of the same sex. The mates sought after not
infrequently suffer seriously from the excesses of the seek-
ing males, the females being usually more passive and hence
the more liable to injury in this way. Thus, J. S. Huxley
has lately told of the exhaustion and actual death of the
female mallard duck from being repeatedly "tread" by the
males, the same and different individual males participating
in the strangely destructive performance.

Finally, the individual itself is not safe from self-injury
through its own sex impulses. Some of the forms which this
sort of thing may take in the human species are too famil-
iar, too disastrous and too repugnant to need illustration
in proof of their reality. That they occur also more or less

268 The Unity of the Organism

among animals is well known to all who have had consid-
erable experience with domestic animals.

Excessive activity in connection with the alimentary func-
tion must now be glanced at. That there is no nice quanti-
tative balance between the food necessities of the animal and
the food gathering instincts and impulses and efforts on the
basis of the principle of natural economy and parsimony,
is shown conclusively it would seem by many animals which
have the storing habit. The honey bee is an example of
this among insects. Given a sufficient supply of flowers to
work on, in the wild state these bees seem always to store
away more food material than they consume.

The extent of their honey-making is limited rather by
the raw material available and by their own restricted phys-
ical powers than by their nutritional needs. This is the im-
pression I have from my observations on wild and tame bees
and I find it to coincide with that of other naturalists whose
opportunity for observing wild bees has been much greater
than mine. For example my esteemed naturalist friend, Mr.
Frank Stephens of San Diego, California, reminds me that
the view is confirmed by the fact that in "bee trees" a por-
tion of the comb containing honey is not infrequently black
and shows signs of being old.

Darwin made quite a point, it may be recalled, of the
economy in some aspects of the. bee's work. "The comb of
the hive-bee," he says, "as far as we can see, is absolutely
perfect in economising labour and wax." (Cell-Making In-
stinct of the Hive-Bee, in The Origin of Species.) 14 But a
thoroughly economic adjustment between different parts of
a given complicated operation, and economy of the opera-
tion as a whole, are very different.

As an instance of excessive repetition in the food-getting
activities among the insects, the following from Fabre may
be taken as fairly typical. A solitary wasp of the genus
Sphex captures and slays a locust, but instead of using it

Organic Connection Between Physical and Psychical 269

at once for food, or of taking it directly into her home, she
sometimes leaves it on the road, and runs to her home, even
though this is threatened by no danger. Then after a time
she returns to the game. This going-and-coming may be
performed repeatedly before the carcass is finally taken into
the dwelling. If by chance the game is removed during the
absence of the wasp, the wasp returns to the spot where her
load was left, but, not finding it, she, nevertheless, keeps up
the going-and-coming for some time. The first back-and-
forth journey from game to dwelling is explicable, Fabre
shows. "But what is the use of the other visits, repeated
so speedily one after another?" Fabre inquires.15 Something
like this almost every one must have seen, who has watched in-
sects at all.

I am quite certain that the acorn storing habit of the
California woodpecker, Melanerpes formicivorus bairdi, is
quite beyond any use the bird makes of the acorns. In the
first place, despite much discussion of the question whether
the acorns are used at all, and if so how, the case is by no
means clear. But the point I particularly wish to make is
that whatever use, if any, the birds make of the acorns,
whether as food directly or as culture media for worms or
insects, these in turn to be eaten by the birds, they store up
many more than they utilize. This seems to me highly prob-
able from the fact, which I have ascertained by numerous
examinations at different places and times, that many holes
contain dried up and wasted acorns which show no signs
of having been picked at or otherwise moved after they were
inserted into the holes. Furthermore, the great extent of
the hole-drilling and filling in itself seems to exceed the
bounds of necessity, especially in view of the certainty that
the bird's chief food supply is from quite another source. A
pine log fifty feet long and one hundred thirty-six inches in
girth at the middle, which I found in the San Jacinto Moun-
tains, contained on a fairly careful estimate 31,800 holes,

270 The Unity of the Organism

many of them containing acorns.

But even were it certain that the acorns are utilized in any
manner and to some extent in connection with the feeding
function, there are still other evidences than that just ad-
duced of the imperfect and excessive operation of the acorn-
storing instinct. As is well known, the bird sometimes ex-
tends its drilling operations to wooden buildings to the ex-
tent of making itself a great nuisance. I have seen a case
where the birds had pierced the rustic of an uninhabited
house, so that when the acorns were inserted, instead of
filling the puncture as they would fill holes in a tree, they
would drop down into the space between the rustic and the
inner wall. Apparently the failure to stop the hole, and
failure also to perceive why, or to recognize that the hole
could not thus be stopped, "fooled" the birds into putting
one acorn after another into the same hole, endlessly almost,
judging by the great quantity of nuts piled up at the bot-
tom of the space.

While the storing habit of the California woodpecker is
undoubtedly exceptional as to extent, it is by no means
wholly unique. At least one species of blue- jay (Cyanocitta
cristata) has much the same habit, in the opinion of most
ornithologists who have studied the habits of the bird. An
experienced naturalist, E. H. Forbush, has recently said
concerning Mark Twain's "Baker's Blue Jay Yarn," in A
Tramp Abroad, "All of this is not merely amusing; it is good
ornithology in so far as it reports the way a Jay acts."
This story, it may be said for the benefit of any reader so
unfortunate as not to know it, turns upon the performance
of a jay similar to that narrated above about the California
woodpecker, the acorns, and the old house.

The habit of the shrikes (genus Lanius) of impaling their
victims and leaving them, almost certainly operates more or
less independently of, and often in excess of, the food re-
quirements of the birds. "My observations," says Forbush,

Organic Connection Between Physical and Psychical 271

"have led me to believe that it rarely returns to eat what it
has thus cached, unless driven to do so by hunger resulting
from adverse fortunes of the chase." 17

Nor is there much if any question that something of the
same sort occurs among mammals which have the food stor-
ing habit. E. T. Seton quotes the following from Dr. John
Wright concerning the big eastern chipmunk (Tanuas stria-
tus griseus) : "It is a most provident little creature, con-
tinuing to add to its winter store, if food is abundant, until
driven in by the severity of the frost. Indeed, it seems not
to know when it has enough, jf we may judge by the surplus
left in the spring, being sometimes a peck of corn or nuts
for a single squirrel." 18 There are many other statements
by the best authorities, especially concerning numerous spe-
cies of mice, which strongly suggest a like superabundance
of storing activities. But for the rest I will mention a case
that has come to my own notice.

I am indebted to Mr. Frank Stephens for information
about and the opportunity to witness to some extent for
myself the operations of the storing instinct and feeding
habits of the Antelope Ground-Squirrel (Ammospervnophi-
lus leucurus). This chipmunk-like little squirrel proves to
be so readily domesticable that it becomes almost as familiar
a household member, at least for Mr. Stephens' household,
as a domestic cat. Although an account of the habits of the
single individual in Mr. Stephens' possession can not yet be
told fully by a long ways, a few points of much interest
for the present discussion are positive enough.

In the first place the genuinely instinctive character of
the storing habit is established by the fact that although the
specimen under observation was taken soon after birth, and
has lived all its life in complete isolation from parents and
all its kind and has been furnished artificially with an
abundance of food, its storing operations are carried on
constantly and almost as perfectly, so far as one can judge.

272 The Unity of the Organism

as though it were living in the natural state. This fact 'in
itself is evidence that the instinct is not determined solely
by immediate needs of the individual. But much more
convincing evidence furnished by this case to this effect is in
the particular way the instinct works. For example, this
species possesses cheek pouches for carrying food as do so
many rodents which have the storing habit. When nuts,
grain, etc., are presented to the animal she very rarely eats
them immediately even though manifestly hungry, but carries
them away to some distance, one at a time; going back and
forth and placing the articles in her two pouches till these
are quite full. And these little pre-storage journeys, as
they may be called, are often definite in character. At any
given time they end at nearly the same spot, and the animal
takes nearly the same position while the article is being pre-
pared for and inserted into the pouches. This is clearly
the typical procedure in filling the pouches, though it is
varied considerably from time to time.

As to what follows the pouch-filling there is considerable
variation — normally so it appears. In case the animal is
hungry she may quietly extract the nuts from the pouches
and eat them. Or she may run about for some time with
her cheeks bulging full. Or she may take her load off some-
where and lay it away either in some cache previously es-
tablished or in a new one. The cache may be in a bed of sand
if this is at hand ; or it may be in or under some old garment
or piece of cloth or paper which the surroundings may
present.

An especially interesting fact noticed by Mr. Stephens
is the tendency shown on the part of this squirrel to carry
the articles to as distant a place from where it gets them
as can well be reached.

On the whole there is no doubt that we have here a var-
iedly illustrative example of activity over and above need
in the operation of an instinct.

Organic Connection Between Physical and Psychical 273

This bare touch, so far as instances are concerned, of
overactivity in connection with reflexes, and especially with
instincts which are on the whole useful, leads naturally to
the great field of animal play. Space limitations prohibit
us from taking more than a bird's eye view of this field.
Fortunately, however, even such a view can be quite effective
for our purpose because of the well-known work of Karl
Groos, The Play of Animals. Our sole purpose here, as
in the rest of this discussion, is to answer the question
whether animals do or do not carry their activities which on
the whole are fundamental -to their existence beyond what
is necessary for their own individual requirements. With
Groos's explanatory theory of play we are concerned only
so far as it involves the question of fact upon which our
present interest centers. That most if not all animal activ-
ity which can rightfully be called play, and which is not
intelligent, is instinctive, we believe Groos has conclusively
shown. The explanation adopted by Spencer and others
that play is the useless imitation by young animals of useful
activities performed by their seniors, the imitative acts be-
ing useless because merely the overflow of "surplus energy,"
is certainly inadequate, as Groos has insisted. That animals
constantly go through performances playfully which they
have had no chance to see or to have otherwise impressed
upon them from without, is as certain as that they constantly
perform useful acts in this way.

It consequently results that a source of energy for play,
that is, for actions which are not immediately essential to
the existence of the organism, must be an endowment of the
organism no less certainly than that a source of energy must
exist for actions which are essential to its existence. So
Groos's statement: "A condition of surplus energy still ap-
pears as the conditio sme qua non that permits the force
of the instincts to be so augmented that finally, when a real
occasion for their use is wanting, they form their own mo-

274 The Unity of the Organism

tive, and so permit indulgence in merely sportive acts,"
becomes a statement of fact if by "surplus energy" we
understand energy available for, and upon occasion used for,
acts which are not indispensable to the existence of the in-
dividual.

The quantity and generality of play performed by animals
may be taken as one important measure of the extent of the
energy possessed over and above what is essential for their
normal individual existences, and this without reference to
whether or not the play may be useful as a preparation for
future essential activities, or for recreation only. The fact
can hardly be too much insisted upon that ulterior useful-
ness of the organism's acts, whether to the species generally,
to offspring, or to the individual's own future, cannot pos-
sibly be a sufficient explanation of the energy immediately re-
quired for the act itself. Even though an animal does noth-
ing whatever except by reason of its hereditary endowments,
or in the interest of its offspring; and though the real pur-
pose of much that it does looks to its own future, it must
nevertheless continue to eat, digest and assimilate, and
breathe. The subdivision of biology which has come to be
known as physiology has for its distinctive task exactly
that of studying the present activities of the organism. With
the organism's past, whether individual or racial, and with
its future, whether individual or racial, physiology can be
concerned only indirectly.

Summary of Organismal Character of All Subrational

Psychic Life

Having now examined broadly though far from exhaus-
tively the psychic life of the animal in each of its most ob-
vious phases, the highest rational phase, the emotional phase,
the instinctive phase, and the reflex phase (in which tropisms
are included) for purely descriptive and classificatory pur-

Organic Connection Between Physical and Psychical 275

poses, let us briefly summarize what we have learned.

In each phase we have found the organism, living, whole
and normal, indispensable to a comprehension of the phe-
nomena examined. Or, expressed in a different way, we have
found it possible in each phase to reach only a very imperfect
understanding of the phenomena by referring them to the
elements which can be discovered in them. For example, the
theory of association of ideas is inadequate to explain ra-
tional life, in such manifestations as apperception and mental
initiative and creativeness.

In the emotional phase, in such emotions as fear, rage and
sex passion, not only does cursory observation recognize
the involvement of a large part of the organism, but physi-
ological investigation is able greatly to extend our recogni-
tion of this involvement by showing how the nervous system
in its cerebro-spinal and its autonomic divisions, the cir-
culatory, the alimentary, and the internal-secretory sys-
tems, are essentially and reciprocally involved.

As to the organismal character of psychic life in the
phase of instinct, it suffices to recall that one of the most
widely accepted criteria for differentiating instinctive from
reflex activities is that the former involve the organism as a
unity, a whole, while reflexes, according to this criterion,
pertain only to limited portions of the organism. "An
instinct is a more or less complicated activity manifested by
an organism which is acting, first, as a whole rather than as
a part." 20 To this statement of the matter may be added
that when the instinctive act is in the interest of the indi-
vidual performing it, the act is not only by but for. the in-
dividual as a whole.

As to the reflex phase (if that is to be reckoned as psy-
chic) the organismal nature of tropisms has had so large
a place in our discussion that surely no more need be done in
this summary than to remind the reader of our discussion of
tropisms. And even reflexes of a simpler fern than the

276 The Unity of the Organism

tropisms — indeed the abstract conception of the "simple
reflex," though not, perhaps, involving the conception of the
organism as a whole, yet is not comprehensible on elemen-
talistic principles, as our examination of Sherrington's in-
vestigations revealed. And such phenomena as those of the
spreading and compounding of reflexes are quite incompre-
hensible except on the organismal principle, even though the
whole organism may not be involved, observably at least, in
particular reflex acts.

Specificity of Subrational Psychic Life

The concluding section of this descriptive chapter on
psychic integratedness must be devoted to the specificity,
not to say individuality, of animal behavior in all its phases.
The vast body of trustworthy detailed knowledge now in our
possession justifies, I am quite sure, the following general-
ized statement under this head: It is exactly on the psychic
side of animal life, psychic being taken in the broadest sense,
that animals are most differentiated from one another, both
as to individuals and as to species.

Taxonomic zoology is based almost entirely on structural
attributes of animals. This results from reasons that are
obvious, speaking generally, and constitutes a justification
of the fact from a practical standpoint. Nevertheless the
purely practical advantages of the classificatory systems as
they have been built up have been, and are, gained at the
expense of several rather serious disadvantages. One of
these is* as advance of knowledge leads us to realize, that our
well-nigh exclusive attention to structural differences and
likenesses has tended strongly to divert attention from func-
tional differences and likenesses. It is of fundamental impor-
tance for a truly comprehensive science of organic beings,
that is, for a philosophical biology, to regard our synoptic
classifications not as a final result of knowledge-getting, but

Organic Connection Between Physical and Psychical 277

rather as a life-sized sketch, as one may say, of the whole
living world, to facilitate the gigantic task of completing
the picture through the cooperation of numberless artists,
the completion to be accomplished by filling in the sketch
with the entire round of attributes, structural and func-
tional, presented by the natural lives of organisms. I have
dwelt somewhat at length on this matter elsewhere,* and can
refer to it here only as a background for what I wish to say
about psychical specificity.

Two extracts must suffice. "No biological phenomenon
is adequately interpreted or dealt with experimentally, until
it has been considered with reference to the place that the
organism to which it pertains holds in the system of classi-
fication." And further: "What I affirm is that the inductive
evidence has now gone so far toward proving every sharply
differentiated species to contain some differentia in all the
main provinces of their structure and function, that to as-
sume the absence of such differentia in any given case is
unwarranted." 21

I want to utilize these earlier general statements about
organic specificity, as a basis on which to rest a generaliza-
tion concerning the specificity of psychic attributes. So
enormous is the observational data available for illustration
here, that in lieu of presenting any of them I am going to
state in a wholly dogmatic fashion the generalization toward
which we are certainly being led by modern crucial researches
on animal behavior. Let us imagine ourselves possessed of an
approximately exhaustive descriptive knowledge of the be-
havior of the whole animal world, this knowledge being, how-
ever, unaccompanied by any knowledge whatever of the cor-
poreal nature of the animals. This behavior-knowledge
would fall naturally into categories larger and smaller, the

* The Place of Description, Definition and Classification in Philoso-
phical Biology, in The Higher Usefulness of Science (Boston, 1918) ;
also The Scientific Monthly, November, 1916.

278 TJie Unity of the Organism

smaller ones being for the most part subdivisions under the
larger. Then let us imagine this system of behavior-knowl-
edge compared exhaustively with a later-acquired, equally
exhaustive knowledge of the corporeal nature of all animals.
The two systems would be found to match each other very
nearly as closely as though the two had been worked out to-
gether, much as they are being actually elaborated by struc-
tural and functional zoology. In other words, the species,
genera, orders, and so on, of animals are differentiated from
one another and coordinated with one another by their "be-
havior," that is, by their whole round of psychical and reflex
attributes, much as they are by their corporeal attributes.

The inductive evidence for such a generalization is being
produced at the present time by three quite definitely marked-
off kinds of research. These may be designated as ( 1 ) quali-
tative field researches, (2) laboratory experimental re-
searches, and (3) quantitative field researches, the definitely
quantitative method being statistical.

The first-mentioned class of investigations is typified by
the earlier field zoologists, whose aim was to learn, as ex-
haustively as possible in a purely qualitative way, the habits
of animals in nature. Workers of this class are the typical
zoological naturalists of the history of animal biology.
Aristotle, Conrad Gesner, John Ray, Charles Linne, P. S.
Pallas, Gilbert White, J. J. Audubon, J. H. Fabre, A. R.
Wallace and A. Forel may be named as conspicuous examples
of pre-modern members of this class ; and Charles Darwin
stands out sharply as a representative of it, but as a transi-
tion to the modern period, the transitional character of
Darwin being seen not only in the doctrines he proposed but
as well in his intimate combination of the experimental
method with the older method of observation.

By the modern period of research in field zoology I mean
the period during which, while the natural history point of
view and attitude are retained, the critical rigor of modern

Organic Connection Between Physical and Psychical 279

science generally is practiced, and experimentation in one
form or another is employed as a supplementary agency
wherever and whenever possible. One of the best examples
of this type of zoological research and writing is Ants, by
W. M. Wheeler. But a considerable portion, and fortunate-
ly an increasing portion, of experimental research in animal
behavior is being done quite in the spirit of field zoology.
The work of R. M. Yerkes deserves mention as perhaps the
most definitely purposed and executed combination of the
field and experimental methods for investigating the behavior
of mammals and birds, that has yet been made. But much
of the research recently named animal ecology tends strong-
ly toward rigor in field investigation. This kind of study is
specially adapted to bring out the specific nature of behav-
ior, since the group of organisms, species, etc., as a whole,
occupies a central place in the student's interest, so that if
behavior is attended to at all its differential features are
likely to receive attention along with the differential struc-
tural features.

(2) Laboratory experimentation on animal behavior has,
as previously indicated, been prosecuted more intensely and
widely in the modern period than either of the other classes
of investigation. In fact, it may be said to be distinctive of
the period, and to have set the standard as regards rigor for
the other types of investigation. From its very nature, how-
ever, it is not calculated to bring the specificity of behavior
to a central place in the student's interest. Singling out
as it does one or a few attributes at a time as they are ex-
hibited by one or a few individuals of one or a few species,
breadth and penetration of comparison are liable to be sacri-
ficed. This kind of research tends to be extremely particu-
laristic in every way. Nevertheless, painstaking and judi-
cious workers, like Englemann, Forel, Binet, Wasmann,
Romanes, C. Lloyd Morgan, Verworn, Jennings, Loeb,
Holmes, and Parker, generally state what species their in-

280 The Unity of the Organism

vestigations have dealt with, so a reader interested in the
question of specificity can usually detect evidences of dif-
ferences in the behavior of different species, even though the
investigator himself was obviously little interested in the
subject, and so took no pains to bring out such evidence.

Indeed, the fact that species-differences in behavior can so
frequently be recognized in descriptions even though the
writer's general attitude may, if anything, militate against
the disclosure of the differences, is rather strong evidence
of the general prevalence of such differences.

(3) Although statistical investigation of animal behavior
has been much less prosecuted than has either of the other
types it is, nevertheless, within the limits of its availability,
a very valuable method for revealing species differentia, its
efficacy consisting in the fact that species may be compared
with reference to different behavior traits taken one by one,
and on the basis of quantitative data covering considerable
samples of whole populations. The method is specially ap-
plicable to the minute floating life of the seas and lakes,
known as plankton, and is being much employed to this end
at the Scripps Institution for Biological Research. It can-
not be described in detail here, but consists essentially, as
employed at this Institution, in collecting great quantities
of organisms by agencies as nearly quantitatively constant
and accurate as possible, in counting the organisms thus
secured, and in correlating the biological values thus ob-
tained with quantitative studies on the physical environment
of the organisms, these environmental determinations being
made simultaneously with the collection of the organisms.
By this means one element in the behavior, that namely of
the up-and-down journeys in the sea, long known to be per-
formed by many oceanic species, has been studied with a fair
degree of quantitative accuracy as to the extent of travel,
time required for each journey, and environmental influence.
A considerable series of species have been compared on this
basis.

Organic Connection Between Physical and Psychical 281

Two quotations are all that space will permit us to give
for showing what these investigations are bringing out on the
subject of specificity in behavior. The animals referred to
in the first quotation constitute a group of small, arrow-
shaped worms known as the Chaetognatha — bristle-jaws.
"Each species occurring in the San Diego region has its own
definite and specific manner of vertical distribution, just as
truly as it has its own morphological characteristics." 22

Similar results have been obtained by the same methods
applied to a very different group of animals, minute crus-
taceans of the ubiquitous order Copepoda. The investiga-
tions on this group have been made by Dr. C. O. Esterly,
and the results are specially interesting in this case, because
Doctor Esterly has applied laboratory experimentation, to
some extent, to the same animals, and has found a good de-
gree of concordance in the results of the statistical and the
experimental investigations. "A heterogeneous assortment of
forms may be obtained in the same collection but each has its
own way of reacting toward the elements of its environ-
ment." 23

It is the indubitable trend in one direction of the vast
evidence from these three quite different classes of research
on animal behavior that to my mind justifies such a con-
ception of specificity of psychical and reactive animal life
as that formulated above. Something of the probable mean-
ing of this specificity we shall see in the next chapter.

REFERENCE INDEX

1. Loeb ('02) 189 13. Huxley, J. S. 156

2. Morgan, C. L 10 14. Darwin, C I, 353

3. Royce 328 15. Kellogg, V. L 648

4. Royce 329 16. Nature Lovers' Library II, 218

5. Holmes ('11) 92 17. Nature Lovers' Library

6. Holmes ('11) 93 III 100

7. Peckham, G. W. and E. S. 53 18. Seton I, 360

8. Holmes ('11) 21 19. Groos 15

9. Jennings ('06) 49 20. Wheeler ('10) 518

10. Holmes ('11) 95 21. Ritter ('18) 119

11. Wallace, A. R 555 22. Michael 46

12. Huxley, J. S 151 23. Esterly 11

Chapter XXIV

SKETCH OF AN ORGANISMAL THEORY OF
CONSCIOUSNESS

Remarks on the Hypothetical Character of This Chapter

HYPOTHESIS and theory will dominate in the task
upon which we now enter and in this respect the
present chapter will differ sharply from the preceding chap-
ters. Fact, description, classification, and restrained gen-
eralization have been the leading motives up to this point.
One main and several subsidiary hypotheses will be central
in the discussion. Into the presence of these will be sum-
moned many of the facts and generalizations previously set
forth. The purpose in this summoning will be on the one
hand to test the hypotheses by the facts and generalizations
and on the other hand to see how the facts will look in the
light of the hypotheses.

This announcement of the hypothetical and theoretical
character of the task now before us, will give us two advan-
tages: It will justify a dogmatic form of expression at times
which we should not otherwise feel privileged to use; and
will justify a brevity of treatment which would not be pos-
sible were we aiming at thorough generalization and demon-
stration. Hence the justification of undertaking to deal
with so vast and vital a subject in the limits of a sketch.

The Natural History Method and the Study of One's Self

Insistent as I have been on the importance of the natural
history way of approaching the phenomena of the living

282

Sketch of an Organlsmal Theory of Consciousness 283

world, in entering upon the present discussion I must em-
phasize this more than ever and must call attention to the
particular character of this importance in our present un-
dertaking.

The natural history method of viewing organic beings is
per se the comprehensive method, one of its best mottos
being, as we have repeatedly seen, "neglect nothing." That
knowledge of organisms separates itself sharply into de-
partments is no deterrent to the naturalist against utilizing
any knowledge he may come upon that will contribute to his
main aim — that of understanding organisms. Who or what
shall restrain me from observing and carefully thinking
about any fact of my own being which promises to help me
on my road to such understanding? The foremost zoolo-
gists, of modern times especially, have amply recognized and
freely used this principle so far as all physical and some of
the lower psychical attributes are concerned. But when it
comes to man's higher psychical attributes, zoologists have
usually said, sometimes expressly, sometimes tacitly, that
these belong to a wholly different realm, a realm with which
we have little or nothing to do. And their position of
"hands off" as touching man's higher psychic life, has re-
ceived the readier, fuller sanction in that it has accorded
well with the prevalent views and practices of those students,
anthropologists, economists, sociologists, and ethicists who
have made these higher reaches of human life their special
fields of inquiry. But the course of nature can not be per-
manently thwarted. Such an attempt to wrench human life
asunder is bound to fail finally. In the several subdivisions
of biology, normal advance has tended to stay the wrenching
process, comparative psychology being notable in this ten-
dency.

The opposition to such organic disunion consistently
maintained throughout this book reaches its culmination in
these chapters on psychic integration. In what follows we

284 The Unity of the Organism

shall pass more freely than ever from one phase or aspect
to another, over the entire gamut of psychic life both in the
individual and in the animal kingdom. If facts of my own
subjective life will serve my purpose, I shall be as free to
requisition them as to requisition facts of any phase or as-
pect of my objective life. If the ethical or esthetic or social
attributes of the human animal will best illuminate a point,
these shall be brought in with as little misgiving as will be
anatomical or embryological or physiological or instinctive
attributes.

So great store do I lay on this catholicity of attitude
toward psychic life, that I shall show by a single instance
that at least a few other present-day zoologists have some-
what similar feelings about the zoological character of psy-
chical phenomena. Referring to the controversies which
have inevitably arisen over the problem of instinct, W. M.
Wheeler says that such controversy "is pardonable, at least
to some extent, since the subject itself presents no less than
four aspects, according as it is studied from the ethological,
physiological, psychological or metaphysical points of view."
"From the first two of these," the author continues, "in-
stinct is open to objective biological study in the form of
the 'instinct actions.' These may be studied by the physiol-
ogist merely as a regularly coordinated series of movements
depending on changes in the tissues and organs, and by the
ethologist to the extent that they tend to bring the organism
into effective relationship with its living and inorganic en-
vironment. But that these movements have a deeper origin
in psychological changes may be inferred on the basis of
analogy from our own subjective experience which shows us
our instincts arising as impulses and cravings, the so-called
'instinct-feelings'; and these in turn yield abundant material
for metaphysical and ethical speculation." From the
context of these sentences we may infer that Wheeler rec-
ognizes that the four aspects mentioned under which the

Sketch of an Organismal Theory of Consciousness 285

subject of instincts presents itself, represent the same num-
ber of valid departments of man's mental life. The point
I wish to make is that although a zoologist may recognize
without cavil that speculation on psychological, ethical,
and metaphysical problems which arise in connection with
instincts, are legitimate activities of man, and might prop-
erly deny that it is incumbent upon him to do much specu-
lating of this sort, yet it would be incumbent on him to take
due cognizance of these speculative attributes of the human
animal. A truly scientific zoology can not justify itself in
issuing a manifesto to the effect that certain attributes pre-
sented by some animals do not fall within its province. It
may more or less constantly neglect or refuse on practical
grounds, to deal with certain attributes ; but that is a very
different matter from a formal declaration such as many
present-day zoologists make, that with these attributes
zoology has nothing to do. Such a declaration is self-stunt-
ing, if not self-stultifying, in that it is a virtual self-inhi-
bition by zoology of its own growth.

These reflections may be terminated by defining the mo-
tives and the mental attitude with which I approach the
great problem of consciousness. I come to it not as a meta-
physician, not as a psychologist, not as a physiologist, not
even as an anthropologist, but as an anthropological zoolo-
gist ; as a zoologist who in course of his regular professional
work takes up the animal group of which he himself is a mem-
ber, chancing as he does to possess among other attributes
that of knowing his own life directly, that is, through sub-
jective or self-conscious experience, as well as indirectly
through objective experience.

Approaching the problem of consciousness in such an at-
titude and for such a motive, it is impossible to view it other-
wise than as one aspect of the larger problem of life gener-
ally. For while the psychologically and metaphysically im-
portant question of whether consciousness is coextensive

286 The Unity of the Organism

with life need not be raised by the naturalist, the indubitable
fact that at least a large sector of life is conscious ; in other
words, the fact that consciousness is a part of life, he can
not ignore if he is to deal with consciousness at all. For the
naturalist, then, no hypothesis or theory of consciousness
can be satisfactory which is not clearly and expressly em-
bedded in and an essential part of an hypothesis or theory
of life generally. Our central hypothesis, drafted in ac-
cordance with these principles, may now be given.

Formulation of the Central Hypothesis

All the manifestations which in the aggregate we call
Life, from those presented by the simplest plants to those
of a consciously psychical nature presented by man and
numerous other animals, result from the chemical reaction
between the organism and the respiratory gases they take,
oxygen being almost certainly the effective gas for nearly
all animals. An essential implication of this proposition
is that every living individual organism has the value,
chemically speaking, of an elementary chemical substance.

Let us be promptly explicit in recognizing the character
of the two propositions contained in this hypothesis. They
are manifestly chemical in large part, and a complete demon-
stration of their truth is impossible without the aid of chem-
ical research focussed directly upon them. But though
clearly chemical, equally clearly they go beyond — far beyond
—present chemical knowledge. To speak of a whole organ-
ism as equivalent to a chemical element seems at first sight
not only unwarranted by positive chemical knowledge, but
opposed by such knowledge. Furthermore, the term "re-
action" as used in the first proposition undoubtedly seems
quite foreign to the technical meaning which chemistry has
attached to the word. Indeed so remote to say the least,
are these fundamental propositions of the hypothesis from

Sketch of an Organismal Theory of Consciousness 287

definite chemical knowledge, that if they are entitled to rank
as constituting a legitimate scientific hypothesis, this must
be on grounds other than those of present-day technical
chemistry quite as much as on those of such chemistry. In
attempting, consequently, to establish the propositions on a
true and useful hypothetical basis, it will be permissible to
notice these other grounds first.

»
Preliminary Justification of the Hypothesis as Such

The proposition that each living individual has the chem-
ical value of an elementary substance, will receive attention
first, and the initial step will be to inquire what, in general,
the criterion is of an elementary chemical substance. Here,
for instance, is a lump of phosphorus. In virtue of what is
it declared to be such a substance? Not primarily, let us
specially notice, because the phosphorus is simple, that is
to say, is an element in the sense of not being reducible to
still simpler substances. Rather the basal criterion of its
being a chemical substance is that upon its being brought
into contact under certain conditions with certain other
chemical substances, oxygen for instance, there is produced
a third substance having very different attributes from either
of the original substances. Transformation of substances
chiefly through interaction upon one another is the founda-
tion fact which has brought it to pass that substances are
described as chemical. That is the fact upon which the
science of chemistry primarily rests. Facts and problems
of simplicity and complexity, relative and absolute, are
later and secondary. The task of chemistry "consists in the
investigation of substances and those of their processes by
which the physical attributes of the substances undergo
permanent changes." (Handworterbuch der Naturwissen-
schaft.)

Every adequate definition of chemistry and chemical sub-

288 The Unity of the Organism

stance and chemical action contains the idea of transforma-
tion in one form or another. Clearness on this point is in-
dispensable to our purpose. Chemistry is too often defined,
even in elementary text books and in dictionaries, as though
the "composition of matter" were its initial and most es-
sential function. But this conception is surely contrary to
the history and most essential nature of the science. There
is, it seems, entire agreement among competent writers that
scientific chemistry is a direct descendant of Alchemy, and
a very imperfect knowledge of the history of Alchemy re-
veals the fact that the every-where present, normal trans-
formations in nature, particularly in inorganic nature, were
the foundation phenomena of this old art. One has only to
recall the place held by the idea of the transmutation of
metals, this idea having usually the practical aim of chang-
ing the "base metals" into "noble metals." The "phil-
osopher's stone" and the "great elixir" were magical some-
things by which the transmutations could be accomplished.

Greatly significant from our standpoint is the fact that
one of the objectives of Robert Boyle (middle of the seven-
teenth century), who, perhaps as much as any one man, is
entitled to be called the father of experimental chemistry,
was to rectify the false and mystical notions prevalent in his
time about "Elements," "Principles," "Essences," etc. "Tell
me what you mean by your Principles and your Elements,"
Boyle demanded, "then I can discuss them with you as work-
ing instruments for advancing knowledge."

What is "behind" the transformations — forces, elements,
principles, essences, spirits or what not — is indeed an impor-
tant and, properly asked, a legitimate question. But — and
here is the most vital fact of all — it is a question which can
not be raised even, until after the transformations have been
observed, nor can an answer of objective value be given un-
less the whole round of observed phenomena, the substances
previous to transformation, the transformatory processes,

Sketch of an Organismal Theory of Consciousness 289

and the new substances, be accepted at their face value, that
is to say, at a value which is as near to ultimate truth as any
truth whatever, connected with the phenomena.

The elemental constitution of bodies is an inference, al-
ways and solely, drawn from their observed corporeal attri-
butes. And chemistry is the science which assumes the task
of drawing, elaborating, and systematizing these inferences
on the basis of the transformation of the attributes. The
meaning of the statement that chemistry is one of the natural
sciences is that chemistry is the science which uses its natural
history observations to penetrate still more deeply into the
constitution of bodies. Natura a natura vmcitur, nature is
surrounded by, is contained in nature, is as fundamental a
truth for chemistry as for any other natural science. A
living being is as much a natural body as is a piece of phos-
phorus, and its obvious attributes, its outer-layer attributes,
are as essential to its nature as are its inner, its hidden
attributes. So any genuinely transformatory changes, and
genuinely new products arising through the reaction between
the living body and some other body is so far chemical in
nature, and the reacting bodies are so far chemical.

A long step toward justifying the proposition that each
individual living organism has the value, chemically, of an
elementary substance, will be taken if it can be shown that
any qualitatively new product whatever results from the
interaction between the organism acting as a unit, as one,
as an element, and some other element. Having regard to
the entire world of living beings, the chances for finding new
products which may have arisen in this manner are prac-
tically if not theoretically infinite. Manifestly, then, only a
very small sector of the entire range of such possible produc-
tions can be searched. It must, consequently, be our aim, as
always in handling inductive natural history evidence, to
choose for examination evidence which shall be most clear-
cut, most illustrative, and most convincing.

290 The Unity of the Organism

The sector of organic phenomena best capable of yielding
such evidence is, I believe, exactly this of psychic life. And
within the great range of this life, the higher conscious life
of man is most replete with the evidence we seek. Again
within the range of man's higher life, each individual's own
private life, even his subjective life, his consciousness, is the
evidence most certain and convincing. Translating this last
statement into familiar language, one sees that it is only
another way — the scientific way — of affirming the truth, that
the greatest of all certainties of which man is capable is that
of his own existence. I am saying, virtually, that when we
analyze, after the manner of objective science, this old fa-
miliar affirmation about certainty, and carry the analysis
as far as we are at present able to, we find that the sense, or
better, the feeling of certainty of self-existence and self-
identity is in last analysis one of the effects of a transforma-
tory interaction between ourselves and some substance (oxy-
gen?) in our breath, as stated in the first of our two propo-
sitions.

That proposition seems then to be hardly more than a
recognition that psychic phenomena containing at least the
germ of consciousness is a kind of chemical product which
has not heretofore been clearly recognized as such, the lack
of recognition being due to the strangeness of the product as
compared with any chemical products with which experimen-
tal chemistry has hitherto occupied itself. But looked at in
a really broad and deep way, is it any more difficult for me
to interpret a state of consciousness in myself to be a result
of chemical action between me and the air (oxygen?) I
breathe, than for me to interpret the dim greenish-white
luminosity of a piece of phosphorus to be a result of the
chemical action between the phosphorus and the air essential
to the glowing? From a purely chemical standpoint I do
not believe we have any ground for holding that some prod-
ucts of chemical reaction are more comprehensible or less

Sketch of an Organismal Theory of Consciousness 291

comprehensible than are others.

Chemically viewed the problem now on our hands is en-
tirely one of fact — fact as determined by observation alone,
and by observation with the aid of experimentation. If it
can be shown that each individual conscious being really does
behave like a chemical substance in the process of reacting;
and if the result of such reaction (jan be shown to have even
one of the essential marks of a -chemical product, both propo-
sitions of my two-parted hypothesis are warrantable and the
hypothesis becomes genuinely scientific — a genuine "working
hypothesis" — one, that is, for bio-chemistry to take seri-
ously.

More Systematic Justification of the Hypothesis

That the propositions are demonstrable to the extent of
the demand just indicated is my contention. This conten-
tion I will now try to make good and will begin with a few
remarks on a question concerning the hypothesis which
ought to arise instinctively in the mind of every one. That
question is : Does such a conception of psychic life and con-
sciousness as that contained in our hypothesis imply any real
infringement upon or derogation from me, in the deepest
sense a real entity properly designated by the terms person
and personality?

On saying that this query ought to arise mstmcti-vely, I
do not mean ought in the ethical sense, but in the organismal
sense. That is, in a sense which implies that the very nature
of the conscious organism is that it is not only self-existent
in a measure like every natural object, but that it is self-iden-
tifiable, and within certain bounds, self-determinative of its
own acts. Now recognizing it to be thus by its "very na-
ture" is only another way of recognizing that it is so in its
instincts as well as in its physical organization. But since
instinct is more fundamental, more deep-rooted in the or-

292 The Unity of the Organism

ganism than is intellect, as phylogenic and ontogenic psy-
chology make clear, if a pronouncement implying a de-
rogation from the reality and natural prerogatives of the
individual be issued from the intellect, a response of protest
and antagonism would be expected from instinct. This would
be expected as an ordinary organic impulse to self-defense
and self-preservation.

The Nature of "Outer" or Objective and "Inner" or
Subjective

What we have to do consequently is to scrutinize the con-
scious individual in order to see if it presents any uniqueness
of attributes and of transformatory power in reacting with
other bodies that is on a par with the uniqueness of an ordi-
nary chemical substance in the same respects. Now it is,
'as suggested some pages back, exactly in the conscious, the
subjective life, that such uniqueness is most easily demon-
strable. There are several ways in which the conscious indi-
vidual manifests this uniqueness. A particularly convincing
way, I think, is in the relation between what are commonly
known as the objective, or "outer," and the subjective or
"inner" sides of mental life. This, consequently, will be the
approach to the subject chosen by us and we will enter upon
it by returning to Royce, first to his "Outlines of Psychol-
ogy," then a little later to some of his specifically philosophi-
cal writings.

In the first chapter of the Outlines, devoted to initial defi-
nitions and explanations, Royce states, simply and clearly,
a distinction "between our physical and mental life," which
elsewhere he has worked out with great elaboration. Thus:
"Physical facts are usually conceived as 'public property,'
patent to all properly equipped observers. All such observ-
ers, according to our customary view, see the same physical
facts. But psychical facts are essentially 'private property,'

Sketch of an Organismal Theory of Consciousness 293

existent for one alone. This constitutes the very conception
of the difference between 'inner' psychical or mental, and
physical or 'outer' facts."4

Ever-present, and obvious as is the comparison here made,
it nevertheless is of so great importance that we must stop
and reflect upon it, for we shall surely fail to grasp the full
measure of what is to follow if we are lukewarm toward one
of the elements of it. The element I refer to is the unique-
ness, the essentially personal character of inner as contrast-
ed with outer facts. Every normal person is ready enough
to insist that his thoughts, his feelings, his emotions and all
the rest of his higher psychical experiences are his and his
alone. The tremendous reality and force of the rights of
"private opinion," of "personal conscience" and so forth,
among civilized men, hardly need to be expatiated on.

The character of the uniqueness of these experiences, how-
ever, concerns practical living less vitally, so we give it less
attention. The whole vast range of my mental life, from
the lowest, simplest, vaguest sensations to the highest, most
bewildering complex emotions, passions, imaginings and
thoughts, are my own, absolutely, so far as other persons
are concerned. I cannot share them to the least extent with
another person. Of course I can let others, especially my
most intimate associates, my dearest friends, know a good
deal about these experiences of mine. But after all, gladly
as I would share many of them with these friends, it is utter-
ly impossible for me to do so. My experiences must remain
wholly outside of their consciousness. No two persons can
have the same experience any more than they can have the
same hands or stomachs. Nor is this all. If mental life is
subject to the general biological laws of variation into which
we have latterly gained much insight, I am obliged to sup-
pose that these experiences of mine, the whole retinue of sen-
sations, feelings, emotions and thoughts, differ somewhat
from the corresponding experiences of other persons. And

294 The Unity of the Organism

all observation confirms this supposition— much of it strong-
ly. Inferential evidence could hardly be stronger than that
my particular emotional response to opera singing, for
example, is quite different from that of many other persons.
Obviously we are here skirting the edge of what modern
realism in formal philosophy calls pluralism, and deals with
in part as the question of whether percepts are strictly indi-
vidual and personal. No philosopher with whose views I
have become acquainted, has discussed this question so fully,
and in my opinion, so illuminatingly as Sellars. The follow-
ing sentences taken from his chapter, The Advance of the
Personal, show clearly, it seems to me, that the conclusions
he has reached, working from the purely philosophical side,
are essentially the same as those arrived at by me, advancing
from the biological side: "What may be called the sensory
content of our percepts is important, — I do not wish to
be understood to belittle it, — but so are the meanings which
arise in conection with our bodily activities and motor ad-
justments to stimuli. Here again, we are face to face with
individual factors in perception which even the idealist must
recognise and somehow explain. Evidently, perception is not
a mere passive presentation, but a construction whose gene-
tic elements can be partially traced. Finally, let us call to
mind that percepts are continuous with feelings and with the
so-called organic sensations. . . . Once vaguely objective,
feeling is now considered subjective or personal."1 Many
other sentences and paragraphs of like purport could be
quoted from this author. I have selected this for the two-
fold reason that it indicates the measure of my agreement
with his view as to the personal character of percepts and
the rest of conscious life; and at the same time indicates
wherein I shall have to out-do him in the matter of validat-
ing the individual. A part of our task, to be reached a little
later, will be to show that although feeling and all the rest
of psychic life is indeed subjectively personal, it is also

Sketch of an Organismal Theory of Consciousness 295

objectively personal. In other words, it will be my task to
remote, or at least to show the way to remove, the vagueness
which Sellars asserts, rightly, has hitherto clouded this
side of personality. To do this thing is, indeed, one of my
most important chances to contribute to a "better philosophy
of life."

But since our psychical life, especially our conscious life,
is a vast — incalculably vast- — complex of experiences, of
"contents," sounds, sights, memories, feelings, ideas, many
of which are set off very sharply from the rest, are clearly
characterizable, and are wonderfully persistent; and since
innumerable of these are coming along all the while which
have much of genuine newness about them ; and since further,
these contents of consciousness are intertwined with and are
determinative of a vast complex of other contents called voli-
tions which in turn are linked up with and are more or less
directive of bodily activities of many kinds, some purely re-
flex and some instinctive, it seems impossible to escape recog-
nizing, even if one wanted to, that if the verb "to create" has
any definite meaning at all the normal, self-conscious animal
organism is about the most creative thing we know or can
conceive. Indeed it is altogether likely that the very notion
of creation, whether natural or supernatural, came initially
from the creative activity and the impulse to such activity,
of man himself.

We may justly say, I think, that we know all creativeness,
chemical creativeness with the rest, through being in our own
deepest natures creative, that is, transformative and trans-
formative in the way which we call chemical. We learn
about the processes of life and call some of the most essen-
tial of them chemical just by performing those processes as
some of our most essential attributes. A portion of the pro-
cess which goes on within us, together with the corresponding
product, constitutes what we call the science of bio-chemis-
try. This means that according to our hypothesis "objec-

The Unity of the Organism

tive" and "subjective," or "outer" and "inner" as applied to
life, are something quite different from what they have been
either in traditional philosophies, or in most, at least, of
recent psychology. "When we speak," Royce writes, "of
our physiological processes as internal, the word 'internal,'
although it here generally implies 'hidden, in whole or in part,
from actual outer observation', does not imply 'directly felt
by us ourselves.' " 5 My hypothesis implies a denial of the
correctness of this statement. I say that in the sum total
of the "contents of consciousness," a nether segment, as one
might call it, of physiological processes is "directly felt by
ourselves." There is no content of consciousness which does
not contain an element that is internal or subjective in what-
ever sense any other content of consciousness is internal or
subjective. And per contra, there is no content of conscious-
ness which is not objective to some extent, in whatever sense
any other content of consciousness is objective. The mind,
according to this conception, is not something which uses
the brain or any other part of the organism merely as a tool
with which to make thoughts and other contents of con-
sciousness. Nor on the other hand is consciousness of the
nature of a secretion, the gland for which is the brain, though
unquestionably the brain has an essential part in the pro-
duction of thought and the higher contents of consciousness.
Among the consequences of the reaction between the or-
ganism and the air we breathe are consciousness with its
marvellously rich and varied contents.

But at this point I must specially request the reader to
notice that I am not pretending to describe and explain all
the contents of consciousness. In other words it is not a
theory of knowledge, but a theory of consciousness that I
am sketching; and knowledge in the strict sense, and con-
sciousness are very different. They differ, according to my
understanding, much as the fully developed, physical organ-
ism differs from the living substance, or protoplasm, of

Sketch of an Organismal Theory of Consciousness 297

which the organism is composed. Consequently I am not
even concerned primarily with sensation in so far as this im-
plies sense organs or even nerves and nerve terminals of the
simplest kind. Rather I am dealing with the stages and con-
ditions antecedent to consciousness and in which it is latent,
in much such way as the cytologist when he studies the living
substance of all sorts of tissue-cells is not dealing with organs
and the organism in the full sense, but only with their sub-
strata. But although it is not knowledge, properly speaking,
either in its conceptual or perceptual aspect that I am dis-
cussing, since my enterprise does take me across the border
line and a short distance into the realm of knowledge, I must,
in the interest of historical continuity and setting, say a
little more than I have said about the general nature of
knowledge.

My assertion should be taken literally that there is no
content of consciousness which is purely either subjective or
objective, inner or outer, conceptive or perceptive, ideational
or impressional, or whatever form of expression be given the
antithesis here implied. That every content of consciousness
which exists or can be conceived has an essential element of
both members of the antithesis is exactly what I mean. To
illustrate, even the axioms, postulates, or whatever else may
be counted as most ultimate in mathematics contain an ele-
ment of the outer, or objective, as well as of the inner, or
subjective. These mathematical contents of consciousness I
single out to illustrate my meaning because they have been
clung to by philosophers and scientists more tenaciously than
any others as purely subjective or mental. And further
there is a strategic gain in this reference to mathematics in
that it brings into the open the fundamental opposition of
my hypothesis to one main root of Cartesian philosophy;
the philosophy, that is, from which the modern doctrine of
psycho-physical parallelism has grown. Our thinking, which
Descartes held proves our existence, really proves it only in

298 The Unity of the Organism

so far as it shows that among the activities essential to the
human organism thinking is one. In other words the "there-
fore" in "I think, therefore I am," is true only because "I
am, therefore I think," the reverse proposition, is also true
and includes the other truth. The lesser truth is true be-
cause it is an essential part of the larger truth, much in
the same way that the cells of a multicellular organism are
alive because they are essential parts of the organism.

We need not inquire how, from this serious shortcoming of
Descartes' description of psychic life Descartes went on to
the conclusion that "there is nothing really existing apart
from our thought" and that "neither extension, nor figure,
nor local motion, nor anything similar that can be attrib-
uted to body, pertains to our nature, and nothing save
thought alone ; and, consequently, that the notion we have of
our mind precedes that of any corporeal thing, and is more
certain, seeing we still doubt whether there is any body in
existence, while we readily perceive that we think."18 Nor
need we concern ourselves with the voluminous and tedious
reasonings by which a considerable number of moderns, fol-
lowing Descartes's lead, have convinced themselves that they
have "reduced" all reality or at least all reality that really
amounts to anything, to quantity. Enough now to remark
that every modern biologist who really accepts the basal data
of his science, must agree that "Psycho-physical paralellism
. . . stands to-day as the scandalous but irrefutable conse-
quence of postulating a material world without qualities and
a world of minds that lack spatiality and exists — nowhere"
One way of characterizing my hypothesis would be to say
that it is an effort to remove this scandal by showing where-
in the postulation noted by Dr. Montague is not true.

The genetic relationships of my hypothesis can be still
farther indicated by coming on down from Descartes to
Hume then from Hume to Huxley and finally to G. F. Stout
and John Dewey as philosophers of to-day. Hume's nom-

Sketch of an Organismal Theory of Consciousness 299

enclature for the subjective and objective sides of man's
psychic life is "Relations of Ideas" for the first, and "Mat-
ters of Fact" for the second. Of the first kind says Hume,
"Are the sciences of Geometry, Algebra and Arithmetic ; and
in short, every affirmation which is intuitively or demonstra-
tively certain." . . . "That three times five is equal to half
of thirty" is a simple illustration of the relation of
ideas. And, "Propositions of this kind are dis-
coverable by the mere operation of thought, with-
out dependence on what is anywhere existent in the uni-
verse."20 And further on, Part 2, same section, we read:
"It must certainly be allowed, that nature has kept us at a
great distance from all her secrets, and has afforded us only
the knowledge of a few superficial qualities of objects ; while
she conceals from us those powers and principles on which
the influence of those objects entirely depends." Then Hume
goes into a discussion of the operations and relations of the
"superficial qualities" and "secret" powers of objects which
is so similar to my treatment of the relation of the organism
to the attributes of certain objects (chapters 20 and 21 this
book, and, more particularly, my essay Is Nature Infinite?21)
that it seems as though his words must have been in my mind
when I thought out what I have there written, though I cer-
tainly was not conscious of Hume's views. And this sub-
conscious influence appears the more probable in that I have
almost conclusive proof of having read his argument not long
before my own was written. I am certain, however, that if
his statements were in my mind they were only in its pro-
conscious part and were not nor ever had been in its full-
conscious part. In other words, if I had read his words I
had not. grasped their full significance. This probable in-
stance of the "sub-" or "pro"-conscious I refer to not so
much because of its interest in this instance, as because of its
bearing on my conception of the nature of consciousness.
The discussion by Hume to which I refer is that in which he

300 The Unity of the Organism

talks about the sensible qualities and the "secret powers"
of the bread we eat. "Our senses inform us of the color,
weight, and consistence of the bread," he says, "but neither
sense nor reason can ever inform us of those qualities which
fit it for the nourishment and support of a human body."
The particular puzzle upon which Hume comes in this
matter is the fact that although the examination here and
now of a natural object gives us absolutely no clue as to what
latent attributes ("secret powers," he calls them) the ob-
ject may possess, when we examine a second object of the
same kind we assume that the same secret powers are pos-
sessed by the second object. "If a body of like colour and
consistence with that bread, which we have formerly eat,
be presented to us, we make no scruple of repeating the ex-
periment, and foresee, with certainty, like nourishment and
support. Now this is a process of the mind, of thought,"
Hume goes on to say, "of which I would willingly know the
foundation." "The bread," he says, a little farther on,
"which formerly I eat, nourished me; that is, a body of such
sensible qualities was, at that time, endued with such secret
powers : but does it follow that other bread must also nour-
ish me at another time, and that like sensible qualities must
always be attended with like secret powers? The conse-
quences seem nowise necessary. At least, it must be acknowl- _
edged that there is here a consequence drawn by the mind;
that there is a certain step taken; a process of thought,
and an inference, which wants to be explained." Then after
a little further argument to show the necessity of recog-
nizing such a process we find this to me exceedingly interest-
ing passage: "There is required a medium, which may en-
able the mind to draw such an inference, if indeed it be drawn
by reasoning and argument. What that medium is, I must
confess, passes my comprehension; and it is incumbent on
those to produce it, who assert that it really exists, and is
the origin of all our conclusions concerning matter of fact."

Sketch of an Organismal Theory of Consciousness 301

The great merit here shown by Hume is his ability to push
the analysis of his problem to the very limit of the positive
information he had to go on, recognise exactly wherein his
information was lacking, and then stop without running off
into a purely speculative substitute for his deficient knowl-
edge. According to my hypothesis the unknown "medium"
which he saw must exist, the* researches of a century and a
half since he wrote, in chemistry, physiology, general zoology
and botany, and psychology, have enabled us to see is the
individual animal organism reaching with the respiratory
substance (oxygen?) it takes in. In this one particular and,
from the standpoint to which we have been accustomed, very
peculiar case, the reaction is at one and the same time part
of the essence of both ideas and impressions in the Humean
sense, the reaction being the "medium" or the "certain step"
by which the inference is drawn, this inferring being possible
because of the continuity of the organism as a person, or
self, and the persistence of the respiratory substance as the
same identical thing from the past through the present into
the future.

We will now notice how Huxley, because of his much more
extensive knowledge of the structure and function of animals
than Hume possessed, was able to draw still closer than Hume
could to the heart of the old Mind-Body puzzle. The gist
of Huxley's position on, and contribution to, the problem
can conveniently be presented through his remarks on the
question of innateness of various aspects of psychic life,
these remarks occurring in his essay on Hume. After point-
ing out that neither Locke nor Hume seemed to know exact-
ly what Descartes, the originator of the modern conception
of innate ideas, meant by his phrase "idees naturelles,1" Hux-
ley quotes Descartes as follows: "I have used this term in
the same sense as when we say that generosity is innate in
certain families ; or that certain maladies such as gout or
gravel, are innate in others ; not that children born in these

302 The Unity of the Organism

families are troubled with such diseases in their mother's
womb; but because they are born with the disposition or
faculty of contracting them."22 Then after further quota-
tions to the same effect Huxley writes : "Whoever denies
what is, in fact, an inconceivable proposition, that sensations
pass, as such, from the external world into the mind, must
admit the conclusion here laid down by Descartes, that,
strictly speaking, sensations, and a fortiori, all the other
contents of the mind, are innate. Or, to state the matter in
accordance with views previously expounded, that they are
products of the inherent properties of the thinking organ, in
which they lie potentially, before they are called into exist-
ence by their appropriate causes."

The upshot of this clearly is that innate for Descartes and
Huxley means hardly anything else than hereditary, as ap-
plied to the psychical as well as to the physical attributes of
animals. The ample justification in our day of the view
that psychical attributes are hereditary should, it would
seem, restore to full standing in biology, the conception of
innate ideas — only, of course, in a very different sense from
that into which later Idealists have perverted it.

It is in this discussion that Huxley makes one of the most
direct and unanswerable arguments against materialism that
can be made : "The more completely the materialistic posi-
tion is admitted, the easier it is to show that the idealistic
position is unassailable, if the idealist confines himself with-
in the limits of positive knowledge."23 That is to say, if the
materialist insists that all traces of innateness of ideas and
other contents of the mind must be repudiated, he virtually
contends that heredity of whatever sort, whether of physical
or psychical attributes, must be repudiated. With this con-
ception of innateness in the entire psychic aspect of the
organism before him Huxley asks: "What is meant by ex-
perience?"

"It is the conversion," he replies, "by unknown causes, of

Sketch of an Organismal Theory of Consciousness 303

these innate potentialities into actual experiences."24 Now
these "unknown causes" are, according to my view, essential-
ly the same as the "medium" which Hume recognized must
exist for making the "step" possible from the "superficial
qualities" to the "secret powers" of natural objects and from
the "secret powers" of one object to those of another. They
are, to repeat, the reaction of the organism in its latently
psychical aspect, with "the breath of life," that is, with the
oxygen, or whatever be the gaseous constituent of the air
which is active in respiration. And I believe we can see to
a considerable extent why Huxley considered these causes as
wholly unknown. It was because physiology and bio-chemis-
try in his day were not yet able to view the organism from
the standpoint of physical chemistry. Because of this ina-
bility Huxley nor any other physiologist of his period had
an adequate structural ground-work for thinking organis-
mally about living things. They were consequently obliged,
really, to think of all psychic phenomena, and consciousness
with the rest, as being restricted to the nervous system.
That such was Huxley's view at any rate, we know from
his own words : "No one who is cognisant of the facts pf the
case nowadays doubts," he writes, "that the roots of psychol-
ogy lie in the physiology of the nervous system." The im-
portant revision of this statement which our hypothesis calls
for is that while the roots of psychology are indeed in the
nervous system they are by no means in that system alone.
They pass through it to a much deeper level, so to speak, and
in passing draw great nutriment from it.

In a brief but important paper starting off with the prop-
osition that a philosopher can not legitimately question the
existence of the external world — that all he can rightly do is
to inquire what that world is and how we can know it at all,
G. F. Stout comes to the kernel of the problem in considera-
bly the same way that Hume and Huxley came to it. "For
primitive consciousness and for our own unreflective con-

304 The Unity of the Organism

sciousness," he says, "sense experience and the correlative
agency which conditions it coalesce in one unanalysed total
object. They coalesce in such a way that the sense-presenta-
tion appears as possessing the independence of the not-self,
and the independent not-self seems to be given with the same
immediacy as the sense-presentation." And, "this complex
but unanalysed cognition," Stout continues, "is the germ
from which our detailed knowledge of matter develops."39
If proved true my hypothesis would be a considerable for-
ward step, I believe, in analysing this "unanalysed cogni-
tion." For although Stout's assertion "the independent not-
self is not matter" seems at first sight to exclude oxygen or
any other constituent of our breath from such a place in
the external world of his conception as that which it has in
that world according to my conception this exclusion is, I
think, only seemingly so, for a sentence farther on the author
says matter "essentially includes the qualification of the in-
dependent not-self by the content of sense-experience." The
seeming discrepancy is probably due to the generality of the
term matter. I too would say that the "independent not-
self" is not matter were I to mean by matter the total sub-
stance of the external world. But in the sense that the effec-
tive respiratory gas (oxygen supposedly) is matter, my
hypothesis would require me to hold that the not-self has an
essential material component, which component is really the
attribute of the gas in virtue of which it reacts with the
organism in the peculiar way it does to produce conscio.us-
ness. It seems to me that what Stout seeks in the "quali-
fication of the independent not-self by the content of sense-
experience" is the immediately consciousness-producing attri-
bute of the respiratory gas. We might state the point this
way: Oxygen (or the effective respiratory gas) has a double
status in human consciousness. First and most fundamental-
ly, it has the status of an immediate and essential participant
in producing all consciousness whatever; and second it has

Sketch of an Organismal Theory of Consciousness 305

the status of an indirect participant in producing the par-
ticular consciousness which we call observational knowledge
of the gas. Our knowledge of this one gas is due to two
tilings, (1) to our reaction to it through our sense organs
in the usual psychological meaning of react; and (2) to our
reaction with it through the protoplasmic basis of all con-
sciousness, reaction in this case having the meaning which
chemistry has given the word. What the relation is be-
tween the attributes of the gas in virtue of which it reacts
with the organism in these two ways, and also what the rela-
tion is between the attributes of the organism in virtue of
which it reacts with the gas in these two ways, are questions
with which a theory of knowledge would deal but which lies
outside of the scope of this sketch, which, as has already
been said, restricts itself to a theory of consciousness. I
may, however, refer in passing to the fact that chemistry
appears to be all at sea on the problem of the relation be-
tween the chemical and the physical attributes of all sub-
stances whatever; so the difficulties about oxygen in this one
particular are not an unshared difficulty.

Finally, to bring this exposition of the historical setting
of my hypothesis down to the present hour, I call attention
to the way the hypothesis connects with the best that formal
philosophy in our own day has done, or as I suspect is
competent to do, towards making out what "experience" is.
No philosopher with whom I have met has gone farther in
this direction than John Dewey. In his recent essay, A Re-
covery of Philosophy, we read: "Dialectic developments of
the notion of self-preservation, of the conatus essendi, often
ignore all the important facts of the actual process. They
argue as if self-control, self-development, went on directly as
a sort of unrolling push from within. But life endures only
in virtue of the support of the environment."26 The italics
are mine and mark the most vital part of the quotation for
us. And a page farther on: "Experience is no slipping

306 The Unity of the Organism

along in a path fixed by inner consciousness. Private con-
sciousness is an incidental outcome of experience of a vitally
objective sort; it is not its source. Undergoing, however,
is never mere passivity. The most patient patient is more
than a receptor. He is also an agent — a reactor." . . .
Again the italics are mine. I take the liberty to end the quo-
tation at "reactor" though the remaining part of the sen-
tence is important for Dewey's particular purpose. But my
aim is different. I want to fix attention on the two state-
ments italicised for the purpose of showing how my hypo-
thesis connects with Dewey's general conception of experi-
ence. When Dewey says life endures only as supported by
the environment, he is speaking in very general terms, having
reference, I imagine, more to social and other bulk aspects
of environment. My hypothesis, on the contrary, makes the
dependence of life on environment exceedingly specific in that
it undertakes to show the particular thing in the environ-
ment, namely, the respiratory part of the atmosphere, which
is physiologically basal to self-development and self-pre-
servation. The Self which traditional philosophy has strug-
gled so hard to understand is literally, the human organism,
according to my hypothesis. And when in this discussion I
speak of it as reacting with the respiratory air to produce
consciousness, I am using the verb to react in a very specific,
physico-chemico-biological sense, while Dewey is using it in
a general sense, and explicitly at least, with only a psy-
chological implication.

The "self" which I am suggesting does indeed imply
"another" no less unequivocally than does the "self" of ad-
vanced social psychology. But the "self" and the "other"
implied by my hypothesis differ from those of current philo-
sophical theory in that the roots of both are not only in
the social relationships of the human species, but extend
right on through these into sub-human relationships, even
down into the very constitution of inorganic nature. The

Sketch of an Organismal Theory of Consciousness 307

"self" and the "other" of my conception are more personally
objective, and more cosmic in their affinities, than are the
"self" and the "other" of social psychology.

Continuing now with our examination of the foundation
of my hypothesis I find it convenient, especially because of
my reference a few pages back, to Huxley's unanswerable
contention for an essence of truth in both materialism and
idealism, to call attention to a natural history fact in the
higher mental life of man which I take to be a strong con-
firmation of the contention. This fact concerns the general
difference between what are commonly known as the mate-
rialistic and the idealistic attitudes of mind. This difference
comes, I believe, to the same thing finally, as the difference
between the objective and subjective attitudes, and is also
the difference, at bottom, between what in rather loose though
prevalent expression, is called the difference between the
scientific and the philosophic attitudes. It would seem that
the philosopher who declares himself to be an Absolute Ideal-
ist, as Royce does, is under heavy obligation, especially if
he enters the field of psychology, to explain the fact that the
originators of great interpretative ideas of nature have in-
variably recognized that their hypotheses must be "proved" ;
that is, that the subjective experience which constitutes the
hypothesis must be found to have its counterpart in the ex-
ternal world of sense. If "Reason creates the world," even
in the recondite meaning of Royce's philosophy, how hap-
pened it that Newton should have been so "restless" for evi-
dence of an objective, an external counterpart to the subjec-
tive result he had reached by mathematical reasoning, that
he held back his reasoned creation for sixteen years, waiting
for the proof, the sense-perceptual or at least the sense-
percepfibfc experience, that should round out his reasoned
truth? May not, I ask, the very kernel of the difference
between science at its best and philosophy at its best be in
this, that the typical scientist is somewhat deficient in "rest-

308 The Unity of the Organism

lessness," adopting Royce's terminology, for internal or sub-
jective reality ; while the philosopher of the schools is some-
what deficient in restlessness for external or objective reality?
We could say with almost literal chemical accuracy that the
curiosity and eagerness of the naturalist for yet unobserved
objective truth is due to an unsatisfied affinity which is weak,
or in some instances, wholly lacking, in the subjective idealist.
The facts which seem to justify our chemico-organismal
hypothesis of conscious psychic life, seem also to imply a
complete interpenetration of objective science and idealistic
philosophy.

As to the Lowest Terms of Self -Consciousness

Let us now veer our course in examining self-conscious life,
and see what can be made out about its roots and rootlets
instead of about its fruitage.

We are often reminded that our knowledge about our in-
ternal organs, our heart, liver, lungs, et cetera, comes only
through observations by the anatomist and physiologist;
that we are quite unconscious of these organs in our own
bodies, especially if they are working normally. Now I
point out that to be perceptually conscious of a liver, let us
say, as a specialized morphological entity performing its
appropriate functions, is a very different matter from being
conscious of those primal, undifferentiated processes which
are basal to life itself, and so are common to all the tissues
whether liver, muscle, brain, or what not, so long as they are
actually living. That that which is truly organic, in the
sense of pertaining to the fully constituted organism, must
be regarded from this standpoint as well as from the stand-
point of their final state of differentiation, is one of the
common-places of modern biology. Let a person in as near-
ly perfect health as he ever experiences, do his best to elimi-
nate all external and internal stimuli of his specialized

Sketch of an Organismal Theory of Consciousness 309

sensory parts ; also all remembering, all feeling of the usual
kind, all imagining, and all thinking. Then let him answer
the question: How do I know I am alive? An undertaking
of this sort is wholly introspective in the sense of being
such that each person must engage in it for himself alone.
He can not show his results to anybody else. A good bit ot
ingenuity may be exercised on it and the outcome will be
found to be rather surprising if not very conclusive as to the
purpose for which the experiment was tried. But the results
as reported may be of some value. Personally, I believe I
can follow my consciousness down to where I can recognize
its most basal remaining "content" to be an awareness of
what I may call extension without definite limitations. It
seems to me I can detect something to which I could not,
from its nature alone, apply the terms "I" or "me" as some-
thing differentiated from everything else. Possibly what I
note is wholly fanciful, but I seem to feel myself in about
the condition of psychical life which I imagine a star fish is in.

Of course I realize how far such a statement is from being
purified of all thought and other ordinary mental elements.
Nevertheless, I believe it to be of some value as evidence
that consciousness is an attribute of the organism as a
whole, and can neither be held to contain an element which
can exist separately from the organism, nor be restricted
to any particular part of the organism as the brain or the
nervous system. There seems to be some evidence "directly
felt by us ourselves," and that evidence points to this con-
clusion as to the nature and "seat" of consciousness. The
point is susceptible, I am quite sure, of rather rigid experi-
mental examination. However, the further experiments
which have suggested themselves to me involve difficulties
more formidable than I have thus far been in position to
attempt.

The reader acquainted with James's notable Chapter X,
"The Consciousness of Self" (The Principles of Psychology,

310 The Unity of the Organism

Vol. 1) will recognize the difference between such introspec-
tive experimentation as that here indicated, and that so il-
luminatingly described by James as tried on himself. While
James's undertaking was to give an account of the thought
and other processes in consciousness as he could observe them
in himself, what I want to accomplish requires me to get rid
of, to ignore as far as possible, the very things which James
was studying. I want to find whether any "content of con-
sciousness" remains after thought and the other usual men-
tal contents are out of the reckoning. I believe, however,
that James opens the way to such an hypothesis as mine.
Thus in a footnote we read, "The sense of my bodily exist-
ence, however obscurely recognized as such, mat/ then be the
absolute original of my conscious selfhood, the fundamental
perception that / am. All appropriations may be made to
it by a Thought not at the moment immediately cognized by
itself. Whether these are not only logical possibilities but
actual facts is something not y«t dogmatically decided in
the text."6

Except for a little misgiving arising from uncertainty as
to the exact meaning of "Thought" in this quotation, I be-
lieve my hypothesis does what James says his text leaves un-
decided.

This foot-note of James's may serve as a switch key to
shift the current of our discussion from the psycho-con-
scious phase of life through the psycho-physical to the purely
physico-chemical phase. The course along which this shifting
will run can be designated thus: full-fledged intellect (al-
ready examined), instinct, emotion, bio-physico-chemical or-
ganization.

Instinct and Physical Organization

The discussion from which we have just turned of the
relation between "inner" and "outer," between "subjective"

Sketch of an Organismal Theory of Consciousness 311

and "objective," must be regarded as meeting the require-
ments of this sketch so far as the first member of the series
is concerned; and the relation between instinct and physical
organization will now receive attention. The evidence of
vital connection here is so abundant and clear-cut, and the
views of competent observers are so unanimous that the sub-
ject can be disposed of quite summarily. Probably the most
indubitable single block of evidence comes from nest-building
and cocoon-spinning insects. Many of the facts from this
field have been so much exploited for the very purposes to
which we now invoke them that a few quotations from and
remarks upon the writings of naturalists generally acknowl-
edged for learning and judicious thinking will suffice.

We turn first to W. M. Wheeler, and take to begin with,
words which he in turn quotes from Bergson: "As Bergson
says," we read, " 'It has often been remarked that most in-
stincts are the prolongation, or better, the achievement, of
the work of organization itself. Where does the activity of
instinct begin? Where does that of nature end? It is im-
possible to say. In the metamorphoses of the larva into the
nymph and into the perfect insect, metamorphoses which
often require appropriate adaptations and a kind of initia-
tive on the part of the larva, there is no sharp line of de-
marcation between the instinct of the animal and the organiz-
ing work of the living matter. It is immaterial whether we
say that instinct organizes the instruments which it is going
to use, or that the organization prolongs itself into the in-
stinct by which it is to be used.' ' And Wheeler continues :
"The spinning of the cocoon by the larval ant is a good
example of the kind of instinct to which Bergson refers.
From one point of view this is merely an act of development,
and the cocoon, or result of the secretive activity of the seric-
teries and of the spinning movements of the larva, is a pro-
tective envelope. But an envelope with the same protective
function may be produced by other insect larvae simply as a

The Unity of the Organism

thick, chitinous secretion from the whole outer surface of the
hypodermis. Here, too, we have an activity which, though
manifested in a very different way, is even more clearly one
of growth and development. And when the workers of
(Ecophytta or Polyrhachis use their larvae for weaving the
silken envelope of the nest, as described in Chapter XIII,
we have a further extension and modification of the cocoon-
spinning activities. In this case the spinning powers of the
larva are utilized for the purpose of producing an envelope,
not for its individual self, but for the whole colony. In
conventional works this latter activity would be assigned a
prominent place as a typical instinct, the spinning of the
cocoon might also be included under this head, but the form-
ation of the puparium, or pupal skin, would be excluded
as a purely physiological or developmental process, yet this
last, no less than the two other cases, has all the fundamental
characteristics of an instinct."7

Then immediately follows this statement, especially signi-
ficant for the proposition of our hypothesis which assigns to
the individual organism the chemical value of an elementary
substance: "Viewed in this light there is nothing surprising
about the complexity and relative fixity of an instinct, for it
is inseparably correlated with the structural organization,
and in this we have long been familiar, both with the de-
pendence of the complexity and fixity of parts on heredity
and the modifiability of these parts during the life-cycle
of the individual. Fixed or instinctive behavior has its
counterpart in inherited morphological structure as does
modifiable, or plastic, behavior in well-known ontogenetic
and functional changes."

The statement that surprise is largely taken away from
such elaborate manifestations of instinct as those here de-
picted, by recognizing that the instincts are "inseparably
correlated with structural organization" and have their
"counterpart in inherited morphological structure," will, no

Sketch of an Organismal Theory of Consciousness 313

doubt, receive the assent of most zoologists, as will also the
statement that our long familiarity with structural organi-
zation and morphological inheritance is what makes us re-
gard these without surprise, and, by inference, as compre-
hensible. It is not that the corporeal form and structure of
the worker ants and of the larvae which they manipulate as
spinning instruments and shuttles for making the nest, are
necessarily simpler and, on that account, more comprehen-
sible than are the instinctive acts of the workers, but that
during our whole lives we have been familiar with structure,
and ourselves exist as "structural organizations." This is
equivalent to saying that we have always been not only learn-
ing but directly experiencing interdependences and correla-
tions among the common body-parts and body-acts, and so
regard them as comprehensible, as explicable. To compre-
hend really an external complex of structures and activities
is to live the counterpart of it. To understand such a com-
plex scientifically is to understand it through a course of
observation and reasoning; that is, rationally. To explain
such a complex is to bring in, or recognize consciously one by
one the constituent elements of the complex, and recognize
all these as parts of the ensemble. It is to recognize the
elements in both their isolate and integrate capacities.

So much for the evidence of integration between instinct
and physical organization as presented by one carefully phil-
osophical naturalist. Several other naturalists have gone
nearly as far, but this single instance is so typical and conclu-
sive as to the objective facts that it will suffice. In com-
menting on the significance of being surprised at such rarely
witnessed performances as those furnished by these ants,
while we are not surprised at common structures and acts
of equal or greater complexity furnished by more familiar
animals and by ourselves, I go beyond, though only a little
beyond Wheeler.

The only other zoologist to whom I turn for evidence of

314 The Unity of the Organism

vital relation between instinct and structure is C. O. Whit-
man. His testimony supplements Wheeler's in that it is
more exclusively and radically objective than is Wheeler's;
that is, it verges less toward the subjective- type of presenta-
tion and draws nearer to the bio-chemical ground work. Al-
though Whitman wrote relatively little on animal behavior,
that little seems to me to contain some of the most important
observations and conclusions which have been produced in
this branch of zoology. What I utilize is taken from his
address Animal Behavior. The animals upon which Whit-
man's chief studies were made were leeches of the genus Clep-
sine; a salamander (Necturus) ; and pigeons of several spe-
cies. Our purpose will be best served by quoting a few sen-
tences which go direct to the heart of the question in hand,
that namely of the vital connection of instinct and basal
physical structure. "The view here taken," Whitman writes,
"places the primary roots of instinct in the constitutional ac-
tivities of protoplasm and regards instinct in every stage of
its evolution as action depending essentially upon organiza-
tion".8 Then, apparently to clarify and emphasize the last
clause about the dependence of instinct or organization, he
adds a footnote thus : "Professor Loeb refers instinct back
to '(1) polar differences in the chemical constitution in the
egg substance, and (2) the presence of such substances in the
egg as determine heliotropic, chemotropic, stereotropic, and
similar phenomena of irritability.' According to this view,
the power to respond to stimuli lies in unorganized chemical
substances, and the same powers exist in the adult as in the
egg, because the same chemical substances are present. Or-
ganization serves at all stages merely as a mechanical means
of giving definite directions to responses.

"The view I have taken regards instinctive action as
organic action, whatever be the stage of manifestation. The
egg differs from the adult in having an organization of a
very simple primary order, and correspondingly simple pow-

Sketch of an Organismal Theory of Consciousness 315

ers of response. Instinct and organization are, to me, two
aspects of one and the same thing, hence both have onto-
genetic and phylogenetic development."

These statements show, as do those given in our discussion
of the cell-theory, how far Whitman went away from full-
fledged elementalism and toward organismalism. But his
treatment of instinct and animal behavior reveals what his
treatment of the cell-theory does not, at least so clearly ;
namely, how far he also went on the way to the natural his-
tory mode as contrasted with the mechanistic mode of phil-
osophizing on biological phenomena. And this gives me a
pleasant opportunity to testify to the genuinely naturalist
current that ran through his life and work. An unforgettable
visit which I had with him among his pigeons not long before
he died, permitted me to see something of the character and
depth of his interest in those animals. His whole attitude
toward them — his wonderfully broad information about, and
understanding of their general ways of life and personal
idiosyncrasies, his solicitude for them, and his measured af-
fection for them — was such as is never displayed by any
one who has not very much of the real naturalist about him,
in his personality as well as in his knowledge. The individual
pigeons, many of them at any rate, appeared to be realities
to him in a deep sense and not merely "mechanical means for
giving definite directions to responses" of chemical sub-
stances. But after all this is said, it must also be said that
there is no evidence that Whitman ever grasped fully the con-
ception that the "constitutional activities of protoplasm" in
which he believed instincts to be rooted, must be the consti-
tutional activities of protoplasms (protoplasm in the plural
number), because no individual pigeon is either any other in-
dividual nor even exactly like any other; and also that the
existence of protoplasms is dependent upon the organisms
to which they belong as well as upon the chemical substances
of which they are composed. Whitman went so far on the

316 The Unity of the Organism

road toward organismalism as to believe genuinely in the
organic and organisation, but not far enough to make him
accept unreservedly individual organisms.

We are able to state definitely wherein lies the great and
rather unique merit of Whitman's investigations on animal
behavior. (1) By a judicious combination of pure observa-
tion and observation aided by experiment and conception, he
pushed psychic phenomena in the form of instinct down al-
most to the physico-chemical level; that is, to the proto-
plasmic level. (2) He at the same time remained positively
within the organic, the living realm. His merit is that of
restraint as well as of positive achievement. He did not per-
mit his enthusiasm for physical explanation to betray
him into adopting a phraseology which, while sounding like
an explanation of instinct, amounts in reality to a denial or
a repudiation of it.

So much for the evidence of vital connection between in-
stinct and organization. According to the schedule indi-
cated a few pages back for reviewing systematically this con-
nection through the entire range of psychic life, we have
next to glance at the connection between the emotions and
organization.

Emotion and Physical Organization

Approaching this subject as we now are from the direction
of psychology proper, the well-known James-Lange interpre-
tation of emotion comes immediately to mind. It will be
advantageous for our sketch not to focus attention too close-
ly on any theory or discussion but to take in as much as we
can of the entire field, keeping in the foreground our own
personal experiences and observations as contrasted with the
descriptions and views of authorities. What I mean is that
the reader shall take himself in hand for serious study as to
his emotional life, watching himself from hour to hour, day

Sketch of an Organismal Theory of Consciousness 317

to day, and year to year under all the varied conditions,
happenings, purposes, and impulses to which he is subject.
In doing this a special point should be made of looking back
scrutinizingly at experiences of particular satisfaction, ela-
tion, joy, sorrow, irritation, anger,' fear, dread, humiliation,
and shame, as soon after their occurrence as possible that
they may be fresh in memory. But incidents and episodes
of one's remoter past which stand out with special vividness
from the intensity of the particular emotions when they were
experienced, or because of results which flowed from them,
will be found illuminating.

To what extent and in what particular fashion was our
bodily organization implicated in the feelings and emotions
we experienced, is our problem. Fortunately one can "live
over again" as we say; can "work himself into" rather pro-
nounced emotional states, through a combination of memory
and imagination. That is, he can be much of a genuine dram-
atist when all alone, as touching events and scenes of his
own past experience. What happens to your body when you
do that sort of thing? is the central question before us. The
very criterion by which you answer this question you will
find will be that of how far the body-manifestations appro-
priate to the particular emotions are elicited through your
efforts. If your hands do not clinch somewhat, if many of
your arm, leg, and abdominal muscles do not contract some-
what, if your respiration does not quicken somewhat, and
other manifestations, various corporeal indices of anger, do
not appear quite independently of direct intention on your
part, you will be sure you have not "worked up" a genuine
state of anger. The only real knowledge of an emotion is a
lived knowledge of that emotion. In order to be a true actor
your body parts must act, directly, automatically, spon-
taneously, so far as any rational purpose is concerned. And
what is true of anger is clearly true of all other emotions.
Our emotional activities may be described as instinctive

318 The Unity of the Organism

and reflex activities, the feeling-impulse of which comes
through intelligence, but is not of intelligence — is not under
the direct guidance and control of intelligence. According
to this interpretation no animal, no matter how highly con-
stituted as to instincts and reflexes, could have emotion un-
less it had intelligence. Emotional activity is instinctive and
reflex activity of an intelligent organism, with, however, the
element of intellect eliminated or in abeyance for the time
being as regards these particular acts. This is what I would
call the natural history description of emotion. And I be-
lieve it is in essential accord with James's conception of emo-
tion, but his description is a psycho-physiological rather
than a natural history description. I am quite sure that
what I have just said means virtually the same as the follow-
ing: "// we fancy some strong emotion, and then try to ab-
stract from our consciousness of it all the feelings of its
bodily symptoms, we find we have nothing left behind, no
'mind-stuff' out of which the emotion can be constituted,
and that a cold and neutral state of intellectual perception
is all that remains."9

I will now point out wherein I believe the natural history
description and interpretation of emotion are somewhat truer
and better than those given by James and other physiologi-
cal psychologists — and, I may add — very much truer and
better than those given by certain writers who approach the
subject from the physiological side pure and simple. James's
epigrammatic statements about being afraid because we
tremble when we meet a bear in the woods ; about being sorry
because we cry ; about being angry because we strike, do his
own position some injustice, I think. This is an instance in
which his gift for piquant writing succeeded too well. But
the fact ought to be noticed that what he actually says is
that as between the usual statement, namely, that we tremble
because we are afraid, cry because we are sorry, strike be-
cause we are angry, and his way of stating the case, his way is

Sketch of an Orgamsmal Theory of Consciousness 319

"more rational." It is only relative, not absolute truth, he
is aiming at in these statements. Nevertheless, after due al-
lowance is made for an expressional miscue to some extent,
there is yet substantial defect in his presentation. Speaking
in general terms, the defectiveness is not so much in the
antithesis set up as in the restrictedness implied. Or, bring-
ing the criticism around toward our particular standpoint,
the statement falls short of being organismal.

Cannon has, I believe, indicated the direction in which the
adequate statement lies. He writes : "We do not 'feel sorry
because we cry,' as James contended, but we cry because
when we are sorry or overjoyed or violently angry or full of
tender affection — when any one of these diverse emotional
states is present — there are nervous discharges by sympathe-
tic channels to various viscera, including the lachrymal
glands. In terror and rage and intense elation, for example,
the responses in the viscera seem too uniform to offer a satis-
factory means of distinguishing states which, in man at least,
are very different in subjective quality. For this reason I
am inclined to urge that the visceral changes merely contri-
bute to an emotional complex more or less indefinite, but
still pertinent, feelings of disturbance in organs of which
we are not usually conscious." What Cannon's criticism
amounts to, expressed in other language is: while freely
granting that organs and functions in the usual physiologi-
cal sense play an essential part in emotion, neither the vis-
ceral nor any other single set of organs is sufficient to account
for the whole of any emotion. Visceral changes contribute
to the "emotional complex," but the real source of the feel-
ings involved is embedded elsewhere and more broadly in the
organization. Cannon suggests : "the natural response is a
pattern reaction, like inborn reflexes of low order." 1J "The
typical facial and bodily expressions," he writes, "automati-
cally assumed in different emotions, indicate discharge of pe-
culiar groupings of neurones in the several effective states."

320 The Unity of the Organism

Without stopping to examine this language in detail, our
aim will be achieved by pointing out that the more closely the
various emotions are scrutinized, and the more effort there
is made to refer them to their causes, the more varied are they
found to be, and the more widely are we led to search in the
organization for causal factors. The mental attitude of per-
fect openness toward any and all facts, both of effect and
cause, which may occur in a given organic situation, is one
of the leading characterizations of the organismal conception.
The assertion that the organism as a whole is the causal ex-
planation of an emotion or an "emotion complex" is justified
by two considerations: (1) Except for the organism viewed
alive and whole and under both its ontogenic and phylogenic
aspects, the emotion would not exist; and (2) so wide-spread
and subtle does common observation recognize the parts of
the organism involved to be in many of its emotional activi-
ties that for practical purposes, it is better to work on the
hypothesis that all parts of the organism are implicated than
to adopt the alternative hypothesis that certain parts only
are involved ; that is, that some parts are not involved.

As a matter of fact, I believe that in spirit James' hypo-
thesis is organismal even though, probably from his training
and career in formal anatomy, physiology, and psychology,
he never became entirely free from the Body-Soul antithesis
and the dogmatisms of "nerve physiology," which have so
dominated modern physiology and psychology. This opinion
I base on the general tenor of his discussions particularly
of the emotions, rather than on his direct formulation of his
theory of emotion. I will quote a few passages that seem
particularly to trend in this direction. "No reader of the
last two chapters {The Production of Movement, and In-
stinct] will be inclined to doubt the fact that objects do
excite bodily changes by a preorganized mechanism, or the
farther fact that the changes are so indefinitely numerous
and subtle that the entire organism may be called a sound-

Sketch of an Organismal Theory of Consciousness

mg-board, which every change of consciousness, however
slight, may make reverberate. The various premutations and
combinations of which these organic activities are susceptible
make it abstractly possible that no shade of emotion, how-
ever slight, should be without a bodily reverberation as
unique, when taken in its totality, as is the mental mood it-
self. The immense number of parts modified in each emotion
is what makes it so difficult for us to reproduce in cold blood
the total and integral expression of any one of them. We
may catch the trick with the voluntary muscles, but fail with
the skin, glands, heart, and other viscera."12 I ask the read-
er to make special note of the part of the quotation be-
ginning, "The various permutations" as we shall have more
to say about it a few pages farther on.

Again we read: "Our whole cubic capacity is sensibly
alive ; and each morsel of it contributes its pulsations of feel-
ing, dim or sharp, pleasant, painful, or dubious, to that sense
of personality that every one of us unfamiliarly carries with
him. It is surprising what little items give accent to these
complexes of sensibility."1 I hope the reader will notice
how easy it would be for me to contend that these state-
ments come near to my statement about "inner" and "outer,"
or subjective and objective; and also to my formal hypo-
thesis as to the nature of consciousness. However, I do not
wish to make too much of such a contention, though I shall
bring up the point again presently. All I want to do just
here is to make still clearer the meaning of my view that
James was organismal in spirit, though not wholly so in for-
mal statement. To me one of the strongest evidences of
this was his obvious effort, as indicated by these and many
other passages in many other writings than his Psychology,
to describe fully the phenomena with which he chanced to
deal. As I have remarked in substance so many times in this
book, one of the most unmistakable signs of the elementalist
attitude in biology is incomplete and more or less perverted

322 The Unity of the Organism

description. And nowhere, perhaps, in the whole biological
realm is there a better chance for description of the genuine-
ly natural history, organismal kind — the kind a cardinal
motto of which is "neglect nothing," than in this very field
of human emotions, especially of one's 'own emotions. Nor
can I refrain from reminding the reader that one of the
master works in this field is Darwin's The Expression of the
Emotion* m Man and Animals,27 and that while a leading
motive of its author was to interpret the emotions in ac-
cordance with the theory of descent and the natural selec-
tion hypothesis, probably the most lasting value of the work
is from its fullness and excellence as a natural history de-
scription of the emotions and their objective expression.

As to the fact of vital interdependence between psychic
life and physical life through the emotions, personal experi-
ence and observation, backed up and supplemented by many
authoritative writings, among which those of Darwin and
James stand out strongly, there seems no longer any room
for question. The role of the emotions as between "Body"
and "Soul" may be crudely likened to the splice which a skill-
ful sailor weaves into two pieces of rope in joining them so
that there shall be no knot and as great strength as in any
other part of the rope. In the recent period of psychology
— of so-called physiological psychology — we have frequently
heard about psychology "without a Soul;" and such an idea
has seemed repugnant to many persons. But if we could
show that this modern psychology is "without a Body" by
the same token that it is "without a Soul," the legitimate mis-
givings about the soullessness of the psychology ought to be
allayed. And really the organismal conception of psychic
life is seen, especially when we examine it in the phase of the
emotions, to amount to such a composition of the Body-Soul
antithesis. "Body" we can see, as it figured in the old psy-
chology, virtually signified what we usually mean by corpse,
or cadaver. "The Body," in that sense was not alive at all.

Sketch of an Organismal Theory of Consciousness 323

It was not alive because all the life was taken out of it (by
the theoretical antithesis) and put into "The Soul."

Glance at the Equiltbrative Interaction Between "Body" and

"Soul"

Going forward from such predominantly observational
descriptions of psychic life in its emotional phase as those of
Darwin and James, to such experimental descriptions as
those being produced by the investigations of Pawlow, of
Crile, and especially of Cannon, we are getting considerable
insight into the rationale of how "Body" and "Soul" vitalize
each other. Modern researches on the physiology or the
psychology (which one calls it depends entirely on the direc-
tion of his approach) of psychic life is revealing something
of the why and how of the poet's instinctive perception, "Soul
needs Body as much as Body needs Soul." Only one aspect
of this "why and how" need be noticed in the present discus-
sion. That is the fact of the balancing off of antagonistic
emotions to make the normal emotional life just as reflex-
actions and instinctive actions are largely phenomena of
equilibration, or balancing-off.

It should be recalled that we have found this antagonistic-
equilibrative principle to run through the entire neuro-psy-
chic life. In the strictly reflex phase the mode of operation
of the opposing muscles, the flexors and extensors of the
limbs, as brought out by Sherrington, was cited as a good
illustration of the principle. A manifestation of the prin-
ciple in a broader way, as measured by the extent of organic
parts involved, was seen in the relation of the vagal (cranial)
and splanchnic (thoracico-lumbar) autonomies, as empha-
sized by Cannon (Chap. 19, this book) this illustration being
chiefly in the reflex phase. In a yet higher phase we saw,
again from Cannon's work, the principle in operation
through the emotions (Chap. 23) thus bringing it up to the

324 The Unity of the Organism

phase of lower conscious life.

The reader should not forget the insistence throughout
our presentation of these antagonistic phenomena, that al-
ways the oppositions and antagonisms and competitions are
fundamentally constitutive as to the normal organism. Even
the most pronounced of them are yet in the interest of the
organism as a whole. They are always partial phenomena
relative to the whole organism. They have evolved in strict
accordance with and sub-ordination to the fundamental na-
ture of the organism in its totality. The opposing muscles
of our limbs can not break or tear one another under normal
conditions. Even antagonisms among the parts of the or-
ganism are possible because the parts belong to the organism.
The antagonisms of the parts do not produce the organism,
primarily, but are themselves produced by the organism, or
at least, are a portion of the means or methods by which the
organism lives and enlarges, develops and functions. All this,
be it noticed, holds not merely as touching purely physical
organization * but as to the entire gamut of psychic life,
at least up to and including instinctive and emotional life.

Support of the Hypothesis by the Physico-Chemical Con-
ception of the Organism

This prepares us for the final step of switching the discus-
sion from the psycho-conscious aspect of life to the bio-
physico-chemical aspect. The place in our discussion to
which this return naturally takes us is that wherein we con-
sidered the organism's chemical nature as interpreted by phy-
sical chemistry. That interpretation has been presented by
several physiologists but with special insight and cogency by
F. G. Hopkins. For example, our citation in Chapter 4 of
the statement that the conception of the organism as a

* Recall the discussions of growth and chemico- functional integration,
chapters 17, 18, and 19.

Sketch of an Organismal Theory of Consciousness 325

chemical laboratory "is rapidly gaining ground," should be
recalled, as should also the opinion of Hopkins : "the chem-
ical response of the tissues to the chemical stimulus of foreign
substances of simple constitution is of profound biological
significance," and that further study of the phenomena "must
throw vivid light on the potentialities of the tissue labora-
tories."1 So far this chemical laboratory conception of the
tissues may be said to be strictly chemical; but let us recall
what the interpretation is when it passes from chemistry in
the exclusive sense to physical chemistry and becomes more
specific as to the laboratory apparatus, as one may say,
through which the "tissues" work. In other words, recall
the conception of the cell and its mode of operating, as
viewed by physical chemistry. The quotations given in
Chapter 4 may well be repeated in part: ". . . the living
cell as we now know it is not a mass of matter composed of a
congregation of like molecules, but a highly differentiated
system ; the cell in the modern phraseology of physical chem-
istry, is a system of coexisting phases of different consti-
tutions." 15 Then from this review our own contention, set
forth especially in Chapter 7, that wherever in such state-
ments as those just quoted from Hopkins "the term cell oc-
curs the term organism really ought to be used."

It is important for our cause generally that the full
weight of our argument in support of the view that on the
strictly physical plane, the organism rather than the cell
is really the equilibration system toward which physico-
chemical knowledge is tending, should be in the reader's con-
sciousness. At this point if, consequently, this is not so,
he is urged to read what is said on the point in Chapters
4 and 7 especially.

Our central purpose now is to show that the organismal
hypothesis of consciousness articulates directly and natur-
ally with the same conception of the organism. Undoubtedly
it is in the emotional phase of psychic life that this articu-

326 The Unity of the Organism

lation is most open to common observation. Compare, for
example, James' "Our whole cubic capacity is sensibly alive;
and each morsel of it contributes its pulsations of feeling,
dim or sharp, pleasant, painful, or dubious, to that sense of
personality that every one of us unfamiliarly carries with
him," with Hopkins' "On ultimate analysis we can scarcely
speak at all of living matter in the cell; at any rate, we
cannot, without gross misuse of terms, speak of the cell-
life as being associated with any one particular type of mole-
cule. Its life is the expression of a particular dynamic equil-
ibrium which obtains in a polyphasic system . . . 'life' as we
instinctively define it, is a property of the cell as a whole,
because it depends upon the organization of processes, upon
the equilibrium displayed by the totality of the coexisting
phases." 16 Also compare Hopkins' statement that among
the different "phases" of the cell in which its life inheres,
"are to be reckoned not only the differentiated parts of the
bio-plasm strictly defined (if we can define it strictly), the
macro-and-micro-nuclei, nerve fibers, muscle fibers, etc., but
the materials which support the cell structure, and which
have been termed metaplastic constituents of the cell," with
James' "each morsel" of our cubic capacity "contributes its
pulsations of feeling, etc."

The congruity of these statements is apparent even when
taken as here exhibited; that is, each as standing by itself
at about the two extremes of the scale of life. When, how-
ever, they are viewed in connection with my general argument
that "cell" in Hopkins' statement ought to be replaced by
"organism"; and in connection with what we have learned
from Cannon and others about the mechanism by means of
which the organism operates in the phase of conscious emo-
tion, it seems as though our organismal hypothesis of con-
sciousness comes near to a demonstration. And so far as
ordinary descriptive natural history is concerned, I believe
this to be true. However, I recognize, keenly enough, that

Sketch of an Organismal Theory of Consciousness 327

from the standpoint of bio-chemistry, and physiology, and
also from that of philosophy in the traditional sense, that
demonstration is not only far away, but is attainable, if at
all, only by surmounting very formidable difficulties. So I
reassure the dubious reader that all I am claiming is that
my two propositions about the nature of consciousness to-
gether constitute a legitimate scientific hypothesis.

Personality and Elementary Chemical Substances

With both the physico-chemical aspect and the psychical
aspect of our hypothesis now before us more fully and
sharply than they have been hitherto we will examine an ob-
jection to it which I apprehend will be the most serious the
hypothesis will meet; namely that to the proposition that
each individual organism has the value in a chemical sense
of an elementary substance. And since this objection will
probably be more intolerant and stubborn from the side of
physics and chemistry than from that of natural history and
psychology I will adjust my remarks with reference to tke
opposition as thus anticipated.

The considerations I am going to present might have been,
in strict expository coherence, presented as a part of my
discussion of the uniqueness of the individual consciousness
as marked by its necessary privacy and its difference from
all other individual consciousness. What we are now to
emphasize is the fundamentally of objective as contrasted
with subjective personality of such highly developed animals
as song birds, domesticable animals, and civilized man.

A complete definition of "personality" is not obligatory
for our purpose. Only this much need be said about the
meaning we shall give the word: First, we deny the right
claimed by some authors to make personality purely psy-
chical, or spiritual — a thing of the "inner," or "deeper"
self; "Self" that is, in a thorough-going subjectivistic sense.

328 The Unity of the Organism

It is on this ground, as I understand, that some psycholo-
gists, as G. F. Stout, and apparently C. Lloyd Morgan,29
deny personality to animals. All I will say on this question
here is that I am quite sure that every close observer of the
higher animals will recognize that if he undertakes to .give
truly full report of his observations on their behavior he wil
have to speak of the personality of some at least of ther
just as he would of the personality of observed human beings,
or he will be obliged to call the same thing by some other
name — a kind of procedure against which we have spoken
strongly throughout this volume. For us, whatever person-
ality may be, we must conceive it to be founded upon, and
conformable to, the organism. "Organism" must be the more
inclusive term. "Person" must stand to "Organism" in the
logical relation of species to genus.

Another meaning of personality in this particular dis-
cussion will concern the uniqueness of each organism as to its
psychical attributes regarded in their totality. By unique-
ness I mean not merely the fact that each organism is itself,
perceptually regarded, but that it is not a replica, a dupli-
cate of any other. It is not only another organism but it is
in some measure a different other organism. For the benefit
of those physical- and metaphysical-minded readers who have
never informed themselves much about the facts of natural
history and have never tried seriously to think in the nat-
ural history manner I would remark that what I have just
said concerning the uniqueness of the individual organism
is only re-asserting in a more refined way what botany and
zoology have recognized more or less definitely since Dar-
win's time at least, and have partially expressed in the terms
"individual difference" and "individual variation."

With this we come to the cardinal point: // individual
animal organisms, especially individual humans under civi-
lization, be contemplated with due heed to the motto "neglect
nothing" the conviction wul be reached that each and every

Sketch of an Organismal Theory of Consciousness 329

one has literally as much of uniqueness about it as has an
elementary chemical substance.

In order to bring out the truth" of this statement we must
exhibit, in the regular natural history manner, the resem-
blances and differences between chemical elements on the one
hand and the resemblances and differences between human
beings on the other, and then pool the results of these com-
parisons.

To the carrying out of this enterprise the so-called peri-
odic law in chemistry is of very great importance. The
essence of this law, stated from the natural history stand-
point, is that the chemical elements range themselves into
natural species and genera after much the fashion that plants
and animals do ; and that the classification is based mostly on
the chemical attributes of the substances, but partly on their
physical attributes also. Thus the "halogen group," that to
which lithium, sodium, and potassium belong, is a genus in
the sense of descriptive natural history, its species being the
substances mentioned with others not enumerated. Also the
group often spoken of in chemical laboratories as "the iron
group" — the genus containing the species iron, cobalt, nickel,
platinum, etc., illustrates the point. Two species of the last
genus, iron and nickel, will be used in our study. Let us
compare some household utensil made of iron with a similar
one made of nickel. For the ordinary uses to which these
implements would be put the difference between the sub-
stances of which they are made would hardly be noticed.
The higher specific gravity of nickel (8.5 plus) is so slight
as compared with that of iron (7.8) that the greater weight
of the nickel implement would probably not be noticed. Nor
would the slightly lower melting point of nickel nor its much
lower magnetic capacity be recognized. The most avail-
able distinguishing difference is in color, the ordinary house-
keeper answering you, if you ask how she knows a nickel
from an iron implement, that the nickel piece is silvery bright

330 The Unity of the Organism

while the iron piece is black.

See now what this means. Actually, as is well known to
every beginning student in analytical chemistry, these two
metals are very similar in color as well as in other physical
attributes — so much so, in fact, that some authors apply
the same term "silver white" to both. What a housekeeper
really means when she says she knows one implement to be
of nickel because it is bright and the other to be of iron
because it is black, is that she is depending on a chemical
rather than a physical attribute for a distinguishing mark;
the attribute, that is, in virtue of which iron is acted upon
much more readily by oxygen in the presence of moisture
than is nickel. The much greater liability of iron than
nickel to tarnish and rust is a chemical rather than a phy-
sical difference between them. This fact, namely that of
the dependence of distinguishing differences between sub-
stances more upon chemical than upon physical attributes is
of very wide applicability in nature, and is greatly impor-
tant both scientifically and philosophically.

Now turn from comparing these two elementary chemical
substances to a comparison of any two human organisms, or ;
persons who might be members of a household to which
the implements might belong. And make the comparison first
on the basis of the physical attributes just as we began
comparing the implements of nickel and iron. Does any
reader doubt that he would find it much easier to distinguish
the persons than the metals? As to purely morphological,
that is, physical differences between almost any two persons
(with the possible exception of certain rare instances of
"identical" twins), there is no room for question. General
shape of head, face and features, and the size and propor-
tions of the various parts of the body furnish many unmis-
takable distinguishing attributes.

Sketch of an Organismal Theory of Consciousness 331
On the Psychology of Subjective and Objective Personality

•

But unerring as are the differentiating marks on the
physical side, such marks are few as compared with those
on the psychical side. Noting first certain merely physico-
psychical differences think of the manners of speech and of
hand writing, to mention only two items ! Undoubtedly these
differences are to a considerable extent physical but no one
would seriously question that psychical factors come in all
along the line. This is perhaps most obvious in speech as
evidenced by voice modulations, intonations, gesticulations,
and facial and bodily expressions. Again, differentials are
everywhere recognizable in responses to sensory stimuli,
especially in the matter of reaction-time. There are the
quick and accurate persons, and the quick and inaccurate
ones ; and there are the slow and accurate and the slow and
inaccurate types, to go only a step in description and classi-
fication on this basis.

Then we proceed to compare the unequivocal psychical
phases of life: the feeling, the emotional, the esthetic, the
religious, and the intellectual phases. Here we pass into
a realm of what might properly be called objective privacy
in psychology, individuals for the study of which would be
largely the student's most intimate and most enduring friends
and associates, human and animal. Such a psychology would
be undeniably so particular and intimate that much of it
would be unpublishable even if it had an interest beyond
the few persons concerned. At the same time there are some
portions of it of great public importance, one such por-
tion being exactly what we are in need of in the present dis-
cussion. I refer to the exceedingly familiar but scientifically
much neglected definite and sustained psychical differences
of individuals who by reason of being members of the same
household or same small community are subject to nearly
identical influence so far as concerns such fundamental en-

332 The Unity of the Organism

vironic factors as food in the narrow sense, drink, air, light
and temperature. The duty before us is that of testifying to,
of viseing, the objectively psychical individual as we did the
subjectively psychical individual earlier in this sketch.
"What is needed," writes Sellars, "is not vague statements to
the effect that individuals cannot be separated or that they
are aspects of one another, but definitions and analyses." 17
Sellars is here raising his voice against the tendency in
present-day social psychology to make the individual a kind
of incident in the social order, a by-product of Society. It
is a satisfaction that the regular course of my psychological
argument has brought me to where I also may contribute
something to the definition and analyses essential to check-
ing the tendency indicated by Sellars. If it can be shown
biologically and psychologically all in one that personality
is indubitably objective, both substantively and kinetically,
not only the Individual but Society will be the gainer, I am
very sure. For my contribution we will examine in outline
what may appropriately be called the action-system (adopt-
ing and expanding Jennings' term) as it manifests itself in
a small homogeneous group of human beings. Our study will
be, in other words, one in domestic and neighborhood psy-
chology.

The "material" in this instance must be nay own household
and the handful of persons constituting the colony of the
Scripps Institution for Biological Research. This group
is rather specially favorable for such a study in that its
geographic severance from other groups, and its strictly
rural habitat give it an exceptionally natural, simple, and
uniform environment. The analysis might run along any
one or all of several axes ; but our purpose will be accom-
plished by following one only. That one shall be the reac-
tion, the behavior, of individual members of the group in
response to the stimulus of the world war. Were complete-
ness to be aimed at in the analysis, every individual in the

Sketch of an Organismal Theory of Consciousness 333

group would have to be considered. Such a treatment would
be highly instructive but space limitations forbid us going to
such length. We must restrict ourselves to a few of the
more pronouncedly individualistic behaviors and must treat
even these in a very sketchy fashion. To be remarked at
the outset is the fact that every member of the group is
deeply loyal to America and to the cause of the Allies. On
the very door-sill of the examination we recognize two well-
differentiated aspects to each person's action-system, namely
an aspect of commonality for nearly all members of the
group ; and an aspect of very pronounced differentiality for
many of them.

Behaviors-in-common will receive attention first. In the
uniform growth, from the very beginning of the struggle in
August, 1914, of belief in the general rightness of the cause
of the Entente ; of realization of the meaning of the struggle ;
and of sentiments and resolutions of devotion to the foreign
nations with which our nation is finally joined, these experi-
ences have been very much at one. To be sure this common-
ness has fallen far short of identity. But as to essentials
resemblance has been far greater than difference. For ex-
ample every adult has accepted unhesitatingly his and her
obligations to the Red Cross ; to the appeals for aid from
IBelgium, France, and the other despoiled countries; to the
increasing cost of living; to the buying of Government
Bonds ; and to the appeals and regulations of the Food Ad-
ministration. Naturally there has been difference in the
particular way and extent of response of each in these mat-
ters ; but in essence there has been nothing differential.

We turn now to behavior-not-in-common ; behavior, that
iis, which has differentiated the members personally with
great sharpness. This examination is much more important
for the subject in hand. The reference here is to each one's
"bit" as the common phrase had it when our country was
first entering the conflict. The "war work" (as the expres-

334 The Unity of the Organism

sion has gradually become with the advance toward the cli-
max of the gigantic struggle) into which each has gravitated
has much the appearance of the naturalness and inevitability
presented by the falling of a stone or the flowing of water.
The case grows so significant at this point that I must par-
ticularize somewhat more than I have heretofore. A becomes
an acknowledged leader in "drives" for Red Cross funds,
Liberty Bond sales, etc. B becomes a regular consultant
on the knitting of Red Cross articles. C is a highly skilled
deviser and maker of dishes from "substitute" foods. D
is appointed an official of the National Food Administra-
tion. E becomes an official teacher of girls and women as to
the peculiar duties and obligations of their sex in war times.
F concentrates nearly the whole of his physical energy upon
an elaboration of the view that a victory over Germany and
her allies cannot be really complete without being spiritual
as well as material — that the philosophy or theory of life
being fought for by Germany must be overthrown as well as
her armed forces. Of the forty adult members of the group
fully one-half have been incited in a special degree to some
activity that has a distinct personal character, some of these,
as above indicated, being very pronouncedly so. The per-
sonality of these reactions comes to view most distinctly in
the fact, absolutely certain to an observer whose acquain-
tance with the persons has been intimate and has extended
over some years, that no one of those who has settled into
one of the special, definite, and important pieces of work
could wholly replace any of the others in their special tasks.
Probably each could do something at the "job" of any of
the others were conditions such as to force him to try; but
success under such conditions would surely be partial, very
much so in some of the cases.

This automatic definition and classification of persons sub-
ject to a common major stimulus, with nearly the same gen-
eral environic conditions, and with almost complete freedom

Sketch of an Organismal Theory of Consciousness 335

of action so far as concerns the particular stimulus, seems
to me a phenomenon of very great importance since it de-
pends upon principles of organic beings, especially upon
principles of civilized man's "being," which are well-nigh if
not entirely universal, I am sure. Undoubtedly the phenom-
enon is often much obscured through counteracting ele-
ments in the environment, especially in social customs, eco-
nomic conditions and general education among civilized men.
But in spite of all these, attentive observation will nearly
always be able to recognize it. Highly significant is it as
bearing on this particular aspect of the matter, that the
niches finally found by most of the persons were obviously
determined to some extent by long continued previous activi-
ties and unmistakable natural "gifts."

Another noteworthy fact is the clear indication of not
mere acceptance, but positive satisfaction on the part of
most if not all the persons, once they are "settled" to their
"jobs," this satisfaction prevailing despite the strenuousness,
perplexity, and wear-and-tear entailed. During the first
weeks of America's plunge into the maelstrom the anxious
psychical casting about in our little group, as throughout
the whole land, presents to the anthropological biologist as
he looks back upon it a case of trial and error on a gigantic
scale, the scene being replete with jumbled elements of noble
zeal, splendid efficiency, mis-expenditure of strength and
funds, and ludicrous proposals. But out of this, as out of
this unprecedented instance of world-wide "struggle for ex-
istence," there is quite sure to come, indeed is coming, as one
of its first fruits, personality more real and powerful and
fuller of grandeur than ever.

While personalities come forth with special distinctness
of outline and forcefulness of expression during occasional
events of vast import to the race like the present war in-
volving literally the whole civilized portion of the human
species, yet I would insist that the difference between the

336 The Unity of the Organism

manifestations at such times and at ordinary times is al-
most entirely one of degree, rather than of essential nature.
The attentive observer will not fail to find personalities as
here understood always and everywhere, no matter how sim-
ple and lowly the lives, and monochrome the external condi-
tions. In little details of intelligent, but still more of reflex,
instinctive, and emotional life, all of which compounded to-
gether makes what we often call temperament, the keen and
sympathetic observer will always see persons in the deep
sense here indicated. Not the transcendent genuises merely,
the Aristotles, the Shakespeares, the Napoleons, have the
right to be called personalities, because of the unique powers
with which they are endowed ; but each and every one of civi-
lization's humblest-ranked myriads, and each and every
nature-tutored denizen of the virgin forest, of the untilled
plain, and of the unregenerate desert, have the same right-
in-kind.

Personality and the "Breath of Life" Viewed in the Light
of Physical Chemistry of the Organism

Swinging the discussion back now on the physico-chemical
aspect of the organism, I recall first the truth alluded to a
little while ago, namely, that it is preeminently the chemical
rather the physical attributes of elementary inorganic sub-
stances which furnish the distinguishing marks of these sub-
stances. Even in the inorganic world we saw that substances
are most readily and decisively differentiated from one an-
other by the transformation-products resulting from the
reaction of the substances upon one another. "Transforma-
tion of energy," using a form of expression favored by the
disembodying tendencies in recent chemical theory, is the
most distinctive thing about all chemistry, inorganic as well
as organic. The oxidation and other chemically reactive
changes and products of nickel and iron, we noticed, are the

Sketch of an Organismal Theory of Consciousness 337

most differentiative things about these metals. Let us push
the application of this criterion of difference a little farther
in comparing human persons. We give energy-transforma-
tion and work performed a leading place here also. And
being naturalistically chemical rather than chemically chem-
ical we are forced to touch the "high spots" only at first
regardless of what may be in between them. We are free to
seize upon the end or completed products of the reactions
and transformations. What reaction-products, I ask, of
nickel and iron towards any other substance or set of condi-
tions are more unlike than the reaction-products of an effi-
cient Department-of-Justice official, let us say and an ef-
ficient food conserving house-keeper, in this time of common
national danger? Yet these diverse products may come from
not only the same danger stimulus, but likewise from as nearly
identical physico-chemical environic stimuli as it is possible
to secure. Were official and house-keeper to eat of the same
food, drink of the same fluids, breathe of the same air, and
be subject to the same temperatures month in and month
out the difference in product would not be a whit less.

So stands the case when viewed in its "high places" only.
But the high places are as real places as any whatever. No
realities, it matters not how obscure or subtle, pertaining to
the intermediate places, can make the high places other
than what they are. Judging human beings by what they do,
by work done through the transformation of the substances
and energies which they take from the external world, their
personalities °xe surely not less well-attested than are the
individualities ^.' elementary chemical substances.* But it
will not do to be satisfied with touching the high places in
this rather jaunty fashion. Some attention must be given to

* A rather full discussion of the point here touched may be found in
my essay, The Higher Usefulness of Science, where I raise and try to
answer the query, "What is nature because man is a part of it?" Per-
haps a less ambiguous way of asking the question would be, "What must
nature be in order that it may produce such an animal as man?"

338 TJie Unity of the Organism

the subtler aspects of the problem. The little we shall do
in this way may be introduced by the query, what reason
is there for including in our hypothesis the supposition that
it is "some substance in the air, almost certainly oxygen,"
with which th'e organism reacts chemically, to produce con-
sciousness and all other phenomena of life? Why single out
this substance from the other elementary substances essential
to life, as for instance carbon or nitrogen ? * My reply be-
gins by recalling the immemorial recognition of the "breath
of life" the "life giving air" and so on, of universal experi-
ence. It is well to recall likewise such semi-philosophic con-
ceptions as that of the pneuma or "psychical breath of life"
of later Greco-Roman philosophy. The inextricable en-
tanglement, historically, of breath and air with spirits is
also worth remembering, especially the continuance of this
into the modern period of scientific analysis, unmistakeable
traces of which are seen in the writings of William Harvey
and the foremost physiologists of the era to which he be-
longed. For example, the spiritus nitro-aereus of John
Mayow which, we now know, was his term for oxygen as
glimpsed first in the history of science, may be mentioned.

More important than any of these reminders from the his-
tory of knowledge is that of the familiar fact that the most
crucial evidences of truly independent or autonomous life of
the individual higher animal are respiratory. That the new
born human babe's first breathing-act is its first genuine in-
dependent life-act is one of the most commonplace of truths.
And recall how the "return of life" as we say of the nearly
drowned person, and of one who has "fainted dead away"
is marked by the resumption of respiratory activities. Cer-
tain reflexes, as those from stimulating the eyelids, and pos-

* The argument in answer to this query should be taken as an exten-
sion of, and in important respects a replacement of, that contained in
my essay, Is nature infinite?" wherein I discuss the specificity of in-
dividual organisms as indicated by how they use their nutrient sub-
stances.

Sketch of an Organismal Theory of Consciousness 339

sibly certain heart flutterings, may be more persistent move-
ments than those connected with breathing. But these are
less certain signs of individual life. It is only to philosophy
of the elementalist sort that the mere twitch of a hand or an
eyelid or a trace of heart action would be a satisfactory
proof of life. Nor would it be to a philosopher of this school
should the "living substance" under observation happen to
pertain to a loved relation or friend. Satisfactory evidence
of life in this case would come only with the nearly simul-
taneous return of breathing and consciousness. A right
interesting section could be written at this point on the
importance of nutriment in the ordinary sense, and of drink,
as compared with air at the very beginning and ending stages
of the individual life. For instance such questions would
have to be considered as that of the independence of the new
individual for a while at the outset on food-yolk in many
animals below the mammals, and on placental connections in
mammals; that is on material metabolically elaborated by
the older or parent individual. But such a discussion not
being indispensable to this sketch, must be foregone. Enough
here to emphasize the fact that while it may be entirely jus-
tifiable to regard oxygen as a food as some good modern
physiologists do the two important facts should never be lost
sight of that (1) oxygen (air) is the one and only ever-
present and never varying constituent of the dietary. In
other words that it is the one constituent which nature sup-
plies as by "free grace" to use a good old theological ex-
pression; and that (2) oxygen is the one and only food that
needs no digesting and so no digestive organs or tissues set
apart for its metabolic elaboration.*

Oxygen is the only food which passes directly as such to

* Were the view held by some physiologists, that the alveolar epithe-
lium of the lungs transmits atmospheric oxygen to the blood by an active
process spoken of as a secreting, this statement would need modifying
somewhat. However, the view does not seem to be accepted by most
authorities.

340 The Unity of the Organism

every part of the organism. In oxygen the organism finds
one of its most fundamental food materials for which it does
not normally have to go in search or to compete with other
organisms. The familiar fact and its significance appear not
to have attracted the attention of biologists much. Even
L. J. Henderson 30 who has written so illuminatingly on
many aspects of organic adaptiveness says nothing definite
on this point. These two facts are weighty reasons for my
proposal to look upon oxygen as one chemically elementary
substance and the organism as another, the reaction be-
tween which is basal in the production of consciousness and
all life phenomena. Consequently the problem of how, ex-
actly, the organism endowed with full-fledged consciousness
reacts toward oxygen is certainly one of the most important
of all problems on the purely physico-chemical side of life.
And, as said early in this sketch, it is just here that my the-
ory is most avowedly hypothetical. It would be quite out of
the question to present in the remaining pages of this book,
even had I the requisite knowledge for doing so, all that
might profitably be said on the subject. Consequently only
two or three of what seem to me the most crucial matters will
be mentioned.

In the first place I ask the reader to recall what has been
said in various of the preceding chapters which have brought
out the indubitable trend of the interpretation of life phe-
nomena according to the principles of physical chemistry,
away from the elementalistic conception of the organism.
The interpretation of the organic cell as a system of phases
in dynamic equilibrium, so strongly set forth by Hopkins
and Bayless will be remembered. And this will call to mind
the sharp way in which the new conception, with its appeal
to the role of surface-layers, membranes, and areas of con-
tact between all sorts of constituent substances, sets itself
over against such pseudo-objective conceptions as that of
biogens, not to mention the horde of out and out subjectivis-

Sketch of an Organismal Theory of Consciousness 341

tic "elements" of which pangens <xnd determinants have per-
haps had the greatest vogue. The importance of the anti-
elementalistic tendency of physical chemistry when it comes
to be applied to biological problems is greatly enhanced, it
appears to me, by the circumstance that J. Willard Gibbs,
who was one of the very first to appreciate in a full scientific
sense the importance of massive as contrasted with minute-
particle phenomena in inorganic nature, and so was one of
the "fathers" of physical-chemistry, made no assumptions
about the invisible composition of substances in his treat-
ment of "Heterogeneous Equilibrium" and allied topics.
"Certainly," writes Gibbs, "one is building on an insecure
foundation who rests his work on hypotheses concerning the
constitution of matter." 31 If this is true as touching the
relatively simple structures and movements in the lifeless
world how much more obviously true is it as touching the liv-
ing world, and especially such life phenomena as human con-
sciousness !

So we are able to requisition one of the admittedly most
important advances of modern times in inorganic science as
support for the supposition that the air we breathe, and
presumably its oxygen, contributes in some direct and funda-
mental way to the production of consciousness even though
this substance, if its "ultimate nature" is what inorganic
chemistry and physics have hitherto attributed to it, has lit-
tle or nothing to suggest that it possesses such a unique
latent attribute. The reader should not fail to recall here
Hume's recognition of the "secret powers" of substances.

But is it not possible that physico-chemical and physi-
ological knowledge of oxygen and air, the "breath of life,"
do contain somewhat more to justify the supposition than is
usually recognized? In this connection I relate that one of
the most mentally adhesive statements I ever heard from a
bio-chemist, its adhesiveness depending largely on the fact
that the chemist was one of great experience as a laboratory

The Unity of the Organism

investigator, was to the effect that chemical analyses make
known what they find and absolutely no more. In other
words such analyses never exclude the possibility of sub-
stances other than those found. And this chemist asserted
furthermore that all organic analyses leave residues to some
extent. No manipulative methods are known, it appears,
capable of effecting a really complete analysis of any or-
ganic substance. Whether these restrictions on analyses
still hold I am not sure, though I have seen or heard nothing
which leads me to suppose they do not.

It is this general shadow of manipulative imperfection
which overhangs all formal physics and chemistry, coupled
with the advances being made from time to time in our
knowledge of oxygen and air which has led me to put into my
hypothesis a shade of doubt as to whether oxygen is the con-
stituent of the air the reaction of which with the organism
produces consciousness. The demonstration of helium and
argon, and probably neon, crypton, and xenon in atmos-
pheric air, all within a little more than two decades, has
influenced my thinking in the same direction. Besides, the
idea, become a commonplace of physics and chemistry in a
single night, figuratively speaking, that the "atom is as com-
plex as the solar system" has had its part in shaping my
conceptions ; as have also such well-credentialed conceptions
from the inorganic sciences as that "Uranium II" is "a long-
lived element" which is the "parent of the actinium series of
elements, but has no genetic connection with the uranium
series"; and that "in the lead pleiad there are seven ele-
ments having quite different atomic weights."

The extent to which, as exemplified by this case, the inor-
ganic sciences have found themselves driven into the organic
realm for terms with which to express their new conceptions
must impress every thoughtful person. Earlier, what we
might describe as purely contemporaneous physical dynamics
had to borrow such terms as energy, power, force, work,

Sketch of an Organismal Theory of Consciotisness 343

from the nomenclature of living" beings. Later, with the per-
meation of all knowledge by the conception of the natural
or derivative origin of everything (a genuinely organic con-
ception, notice), has come even for elementary chemical sub-
stances, the induction into physics and chemistry of such
ideas as genetic relations, parenthood, and length of life.
So my suggestion that the air we breathe must be recognized
to possess latent attributes which by reacting with the or-
ganism produce consciousness, falls into a genetic series in
the history of the interpretation of nature.

The very important question, as already indicated, of ex-
actly how atmospheric or molecular oxygen operates in the
living being generally and the conscious being particularly,
is largely for the future to answer. One should never fail,
however, to couple this question with the same question as to
the behavior of oxygen, and for that matter of any other
chemical substance, in any reaction whatever. Exactly how,
for example, does oxygen operate with hydrogen to produce
the attribute of refrangibility of water; or with phosphorus
to produce the peculiar glow which that substance may ex-
hibit under some conditions?

Concerning the positive knowledge and the views as to
details of the action of oxygen in connection with the or-
ganism, only a little can be said here though that little may
be very important. Looked at from the standpoint of the
old, the orthodoxly atomistic chemistry, probably the most
anomalous thing about my hypothesis is that the organism
conceived as equivalent, chemically speaking, to an elemen-
tary substance, is the unquestioned fact that the organism is
not only composed of several chemical substances, but that
one of -these is oxygen itself. Stated baldly, the anomaly is
that two chemical substances are supposed to react upon each
other, one of which (the organism) is known not only not
to be simple, but to contain the other substance. But even
the old chemistry with its "compound radicals," of which

The Unity of the Organism

cyanogen (CN)2 is said to have been the first discovered, and
of which the unitedly-acting combinations of carbon and
hydrogen as methyl, CH3, affords some slight support for
our conception so far as the mere matter of chemically uni-
tary compoundedness is concerned. In so far, however, as
technical chemistry can be drawn upon for supporting our
hypothesis, it is the new, or physical chemistry, as has
been repeatedly stated, that is our main reliance. Unless I
am greatly deceived, the real inwardness of that great move-
ment in inorganic science is against the age-old conception of
the ultimate adequacy of atoms to explain inorganic na-
ture, almost as positively as the organismal conception is
against the ultimate adequacy of any constituent element
whatever, to explain organic nature. The surface energies,
for example, developed at contact faces and giving rise to
the phenomena of adsorption * appear to be not a whit less
real and ultimate energies than are any that can be attrib-
uted to atoms and molecules taken as such. And, be it no-
ticed, one of the most distinctive things about these areal and
massive energies is that they dominate atomic and molecular
energies to a certain extent. This is just what the now uni-
versally recognized principle of "mass action" is in so far as
such action has been studied enough to make possible its for-
mulation into law; that is enough to learn how it influences
velocity and quantity of chemical change. But would any
careful physicist or chemist pretend to know to a certainty
that such action is restricted to influence of that sort? Surely
not. Are we certain for instance that it can not under any

* Adsorption is the loading of the surface of a solid body immersed in
a solution, with the dissolved substance. Thus it is by adsorption that
charcoal takes the coloring matter out of a colored solution. The action
results from the facts that there is surface tension at the inter-faces be-
tween the charcoal and the liquid, and that this tension is lessened by
the presence of the dissolved color-substance in the liquid. The sub-
stance then moves to the place of lessened tension and concentrates on
the surface of the solid.32 The principle has very wide application in
nature, particularly in organic nature, where colloidal substances and
water are in contact so extensively.

Sketch of an Organismal Theory of Consciousness 345

circumstances influence qualitative as well as quantitative
change? Surely we are not. This of course is far from
contending that mass action actually does influence quali-
ties. My sole point is that so long as there is lack of
certainty that it does not or may not exert such influence
any assumption which implies such certainty is unwarranted
and unscientific.

Putting together, then, the physically massive concep-
tions of inorganic chemistry and the organismal conceptions
of bio-chemistry what seems to follow touching the chemico-
substantive composition of organisms is that a portion of
all the substances essential to life, carbon, oxygen and others,
have been combined from all eternity (whatever be the mean-
ing of the phrase) in the peculiar way called organic, while
other portions have remained in the state called inorganic.
This leads me to remark, quite incidentally so far as this
discussion is concerned, that according to this view the as-
sumption would be that organisms have always existed, or at
least that they have existed as long as "matter" or anything
else of which we have any information or clear conception,
has existed. The warrantableness of this assumption I am re-
lieved from arguing here from having treated the problem
at some length in another place. (Are we obliged to suppose
the spontaneous generation of life ever occurred?)53 All
that need be said now about the outcome of that discussion is
that the warrantableness lies in the absence of any ground
for assuming the contrary. I take my position squarely on
the direct evidence in the case. All the evidence of that sort
we have — and in that discussion I emphasize the fact of its
vast quantity — is to the effect that organisms are produced
by other organisms known as parents and in no other way.*

* To the stock and rather vapid rejoinder that such a solution of the
problem of the origin of life is no solution at all, but only a putting off of
the difficulty, the obvious reply from my standpoint is that I am making
no pretense of "solving the problem," as "solution" would be meant in
the anticipated rejoinder. From my standpoint, however, the everlast-
ingly-from-parents hypothesis would be a solution of the problem if the
hypothesis were proved true.

346 The Unity of the Organism

We can now state briefly as much more of the bio-chemical
aspect of the problem as seems indispensable to our present
argument. A few remarks on what the physiology of our day
often calls tissue respiration will compass what is in mind.
The key fact in this is of two-fold character: (1) The tissues
of the organism, not its blood or any other fluids, contain the
substance which is in the strictest sense living. (2) This
substance is called living because chemical changes of a very
distinctive sort are going on in it. These changes are of a
fundamentally double nature as regards atmospheric or
molecular oxygen; namely, combinative and incorporative
change, and separative and expulsive change. The last-men-
tioned, the separative and expulsive change, is known as oxi-
dation and manifests itself to ordinary experience in the dis-
charge of oxygen combined with carbon as carbon dioxide,
and in the setting free of energy in the form of muscular and
other work, and of heat. The first-mentioned, or incorpora-
tive change, consists in taking in and storing up oxygen,
"somehow," as the more carefully worded physiologies put
it. This statement may be taken as a very brief natural
history description of the most fundamental steps in what
formal physiology calls metabolism with its two aspects, the
constructive, or anabolic, and the destructive, or katabolic.
Probably no one will question that this conception of the
foundations of the life process for nearly, if not quite, all
animal life is that held by the best physiologists since the
time of C. Bernard at least. No physiologist whom I have
consulted has stated the nature of the process more definitely
than has Sir Michael Foster. "The Respiration," he writes,
"of the muscle then does not consist in throwing into the
blood oxidizable substances, there to be oxidized into car-
bonic acid and other matters; but it does consist in the as-
sumption and storing up of oxygen somehow or other in its
substance, in the building up by help of that oxygen of
explosive decomposable substances, and in the carrying out

Sketch of an Orgtwismal Theory of Consciousness 347

of decompositions whereby carbonic acid and other matters
are discharged first into the substance of the muscle and
subsequently, into the blood." 34 And he points out in other
connections that what is true of muscle in this regard is es-
sentially true of all other tissue systems. In another still
more recent text book we read: "Nothing definite is known,
however, as to the nature of the probable combinations
formed by oxygen with the different materials for building
up muscles and other tissues, or of the intermediate anabolic
and katabolic forms through which it passes in combining
with carbon into carbonic acid." 35 And this author then
expresses what are, apparently, his own views, by quoting
from Foster as follows : "The whole mystery of life lies hid-
den in the story of that progress [that of construction and
destruction in the tissues] and for the present we must be
content with simply knowing the beginning and the end."

The kernel of my suggestion so far as metabolism is Con-
cerned, is that the anabolic, or the assimilative, the truly
synthetic aspect of the complete operation, is the continual
renewal, or keeping up of the oxygen constituent of the
organism which comes to it by heredity, that is which has
always been in the "line of descent." It is the maintenance
of what might be spoken of as the original oxygen constitu-
ent of the organism. There would always then be operating
in the organism oxygen of two sources, that from the one
source designated, employing our well-established evolutional
terminology, phylogenic or hereditary oxygen ; and the other
ontogenic or individual oxygen. In general the same kind
of reasoning would hold for the other chemical simples, car-
bon, nitrogen, and so on; but these are in quite a different
status from oxygen owing to the fact that they are not
normally taken by the animal organism in the pure or uncom-
bined state, but only in some other organic combination,
as food in the ordinary sense.

Metabolically expressed, then, we may say in short that

348 The Unity of the Organism

the warrantableness for considering the individual organism
as a chemical element, is the fact that it maintains its identity
as regards all its elementary constituents except one, oxygen,
be wrenching these, so to speak, from other organic com-
pounds (by digesting these) and then by synthesizing the
elements into its own particular substance. Another way of
expressing the same conception is to say that the organism
is an element, chemically speaking, because it reacts directly
in a chemical sense with another element.

Did this chapter pretend to be anything more than a
sketch of a theory of consciousness a considerable discus-
sion of the "activation" of oxygen would naturally come in
somewhere, perhaps at this point. The essense of activation
is the fact that when oxygen passes into the organism by the
respiratory process it is somehow changed into a condition
which enables it to oxidize living tissue-substances as it
can not to any degree, seemingly, when brought into con-
tact with the same substances outside the organism. This
discussion would involve the various theories which have
been put forward to account for this phenomenon, as those
which make use of the principle of enzymes, of peroxides or
of some other. All that our aims here require us to notice
is that nothing conclusive as touching the nature of activa-
tion would come from the discussion. How unsatisfactory
a state this whole subject is in may be seen from the follow-
ing words of a foremost American biochemist: "It has been
a popular practice to appeal to hypothetical enzymes to
explain some of the obscure chemical transformations in the
organism. Thus we have been wandering through the mazes
of the oxidases, oxygenases, peroxidases, reductases, cata-
lases and other products of perplexing nomenclature in the
hope of escaping the uncertainties of intermediary meta-
bolism." 36

Sketch of an Organismal Theory of Consciousness 34*9

Summed-up Statement of Justification of the Hypothesis

The final gathering-up-and-putting-together may now be
made of all that has been said about the physico-chemical
aspect of the organism on the one hand, and about its psy-
chical aspect on the other. That is to say, we are now ready
to epitomize the results of our examination of the ancient
and honorable but withal unsolved problem of how Body and
Soul go together. As regards "body" or "the physical" we
have been led to the physico-chemical conception of the or-
ganism as a well-nigh inconceivably complex mass of sub-
stances, mostly in the colloidal state, operating as a system
of phases in dynamic or constantly changing equilibrium.
As regards "soul" or "the psychical," we have found also a
series of phases of activities, namely the phases of intellect
and reason, those of instinct, those of feeling and emotion,
those of the will, those of the tropisms and the "simple re-
flexes," and finally those of simple protoplasmic response.
According to my hypothesis, the phases of the bio-chemico-
physical sort and the phases of the psychical sort have com-
mon ground in the organism as a whole, the phases of in-
tellect and reason corresponding to the cerebro-spinal nerv-
ous system ; the phase of instinct corresponding probably to
the autonomic nervous system ; the phases of feeling and emo-
tion corresponding mainly to the glandular and visceral sys-
tems; those of the will to the body-muscular system; those
of the tropisms and simple reflexes to the receptor-conductor-
effector systems; and finally those of simple protoplasmic
response to the fundamental protoplasmic mechanism of
response, whatever its structure.

According to the scheme presented in the sketch and
summed up here, just as physical functioning and physical
form reach back to the very dawning of animal life, both in
the individual and in the race or type, so consciousness with
its nether limits in what, following the terminology of em-

350 The Unity of the Organism

bryology (see section on the pro-morphology of the egg-
cell, Chap. 8), might be called pro-consciousness, is an
attribute of all animal organisms. As comparative anatomy
and physiology have made us familiar with the physical as-
pect of the animal organism existing as the fully realized
or developed adult at one end of the ontogenic series, and as
the unrealized adult or germ at the other end of the same
series, exactly so its psychology gradually familiarizing us
with the realized, or adult mind at one end of the ontogenic
series, and as the unrealized or germinal mind at the other
end of the same series. When we affirm that the completed
individual organism is latent in the germ, we must under-
stand that the psychical aspect no less than the physical
aspect is so latent. With very little doubt, it seems to me,
the real meaning of the so-called sub-conscious, and of
psycho-analysis as a method of investigating it, is that the
ontogenic stages of the psychic life of the human organism
are being discovered and that a method of investigating these
stages is being worked out. Freud and his followers have
been and still are somewhat in the dark, I think, as to just
what they are doing, albeit their discoveries and methods are
of the utmost importance.

REFERENCE INDEX
1. Wheeler, W. M 519 21. Ritter, W. E. (19182) Essay 2

3. Sedgwick & Taylor 262-3 22. Huxley, T. H 98

4. Royce, J 2 23. Huxley, T. H 96

5. Royce, J 3 24. Huxley, T. H 100

6. James, Wm 1-341 25. Huxley, T. H 94

7. Wheeler, W. M 521 26. Dewey, John 9

8. Whitman, C. 0 310 27. Darwin, Chas

9. James, Wm 11-451 * 28. Crile, G. W

10. Cannon, W. B 280 29. Morgan, C. Lloyd 256

11. Cannon, W. B 282 30. Henderson, L. J

12. James, Wm 11-450 31. Gibbs, J. W Pref. x

13. James, Wm 11-451 32. Lewis, Wm. C. McC.-.II, 303

14. Hopkins, F. G 217 33. Ritter, Wm. E. (1918=) Essay 1

15. Hopkins, F. G 220 34. Foster, M 469

16. Hopkins, F. G 220 35. Luciani, L I, 395

17. Sellars, R. W 75 36. Mendel, L. B 21

18. Descartes, Rene..Pt. 1, Sec. 8 37. Ritter, Wm. E. (19182)

19. Montague, W. P 120 38. Wendt, G. L 442

20. Hume, David. . .Sec. IV, Ft. 1 39. Stout, G. F 680

POSTSCRIPT

THE argument in favor of the organismal way of viewing
living nature has now run what appears to me its natural
course, to its inevitable end. Yet I cannot bring myself to
write "Finis" without making a few remarks which though
connected vitally with the argument, do not seem an essential
part of it.

These remarks concern the general effect of the organismal
standpoint on those who may grasp it firmly and adopt it
unreservedly. Since, as pointed out in the "Historic Back-
ground" with which this book opens, the standpoint has been
recognized by biologists with varying degrees of fullness
from the time of Aristotle at least, there can be no doubt
that the human mind is naturally attuned, as one might say,
to this general type of response to organic phenomena. It
seems therefore fitting that a presentation like that which I
have made should be accompanied by a few words on the
probable influence of a wide prevalence of the organismal
view. The pertinent question will be asked, how could it
have come to pass that if the standpoint has been so long
in the world it should have missed full recognition and have
failed to exert its due influence? The reply is obvious to an
attentive reader of this book: At no time until the present
in the long historical growth of knowledge of the living world
has information been sufficient to make possible a rounded-
out statement of the conception. To illustrate, it is only in
the very last years that enough has been known of the
physical chemistry of the cell to engender such an interpre-
tation of this exceedingly important biological entity as that
which biochemists are just now reaching. Yet this interpre-

351

352 Postscript

tation is indispensable to anything even approaching a full
development of the organismal view.

But nothing stands out more boldly from the pages of
this book than the insufficiency even yet, of actual knowledge
for making the standpoint complete. If therefore, I append
to my presentation a brief reference to the larger effect the
view has had on myself, and on this basis forecast what the
effect would be on thinking people generally were they to
make it their own, such a forecast will surely be in harmony
with the larger purpose of the book, even though the antici-
patory remarks have no place in the presentation itself.

The long and laborious gathering and arranging of facts,
and weighing of natural evidence and formal arguments
which has constituted the development of the standpoint in
my own mind, has compelled me to re-examine and re-assess
the whole frame and fabric of my spiritual life. Nothing, so
far as I can tell, has escaped. Not my scientific knowledge
alone — my professional stock-in-trade — but all my ideas and
beliefs touching religion, art, society, politics, industry, per-
sonal relations, and private living, have come in for their
share of scrutiny and renovation.

An exceedingly brief "synoptic" classification and char-
acterization * of the entire range of these effects can be
given in the terms of formal science and philosophy.

As to classification, the effects fall into a two-fold group-
ing. One of the groups appertains to the great province of
the nature of knowledge; the other to the equally great
province of the nature of morals.

The characterization of effects on the nature of knowl-
edge which seems to me most inclusive and most practically
significant, may be stated thus: By the validation of ob-
jective knowledge, largely through the principle of what I
have called standardization of reality, but partly through

* See my essay, The Place of Description, Definition and Classifica-
tion (Ritter, 1918).

Postscript 353

the organismal hypothesis of consciousness, such knowledge
is elevated to the rank of strict equality with "pure thought,"
often so-called; that is, with subjective, or intuitive knowl-
edge. In this way mathematico-mechanistic science is de-
prived of the regal place it has claimed for itself since the
era of Descartes and Leibnitz, and is brought to the plane
of absolute equality as to importance and dignity, with
sense-experiential science. By thus adjusting the claims of
these two great realisms of science, an attitude toward the
infinite totality of nature, and a methodology for interpret-
ing it, which have hitherto borne the stamp of subjection
and inferiority assume their rightful places in the great
hierarchy of philosophical science. This leveling-down of
mathematical mechanics and the deductive method and level-
ing-up of observational knowledge and the inductive method,
implies the complete overthrow of psycho-physical dualism
in psychology, and the rescue of personality from bondage
to a theoretically infinite monotony of "Matter and Energy."

The characterization of the effects of the organismal view
on morals centers around the perception that in the establish-
ment of human personality the persons are organically in-
terdependent upon one another; that is, interdependent
through their "attributes of relation," this resulting in the
incorporation of men into a, pluralistic universe far more
real and vital than philosophic pluralism has hitherto been
in position to grasp. Through a type of human conduct
guided by knowledge of these principles of personality and
the interdependence of personalities, and through supple-
menting mathematico-mechanistic methods of study by a
rigid application of observational and statistical methods, a
genuine science of morals, both theoretical and practical,
is made attainable.

That my enterprise of developing the organismal view is
only part and parcel of the general current of interpretation
of living nature which has flowed through the centuries seems

354 Postscript

clear even from my meager acquaintance with the history of
philosophic thought. Thus we read in Windelband (A His-
tory of Philosophy, Eng. by Tufts,) : "For the decisive fac-
tor in the philosophical movement of the nineteenth century
is doubtless the question as to the degree of importance
which the natural-science conception of phenomena may
claim for our view of the world and life as a whole." (624).
Then after speaking of the sharp antithesis between the
Weltanschauung elaborated by the "Highly strained idealism
of the German Philosophy" of the early nineteenth century,
and the "materialistic Weltanschauung" of the later decades
of the same century, the author writes : "If we are to bring
out from the philosophical literature of this century and
emphasize those movements in which the above characteristic
antithesis has found its most important manifestation, we
have to do primarily with the question, in what sense the
psychical life can be subjected to the natural-science mode
of cognition." (p. 625).

That Part II of this book of mine, especially Chaps. 20
to 24, go a long way toward answering the cardinal question
formulated by Windelband appears to me certain. And, I
may add, it also seems quite clear to me that the gigantic
struggle at arms which that philosopher's nation has now
brought upon the world, is one of the strongest proofs that
philosophic thought and, following this, social and political
leadership in Germany have failed miserably to discover the
Via Media between the Weltanschauung of the "highly
strained idealism of the German Philosophy" and the mate-
rialistic Weltanschauung which has finally reached its nat-
ural climax in militaristic brutism, and is almost certainly
(Sept., 1918) approaching its overthrow.

Nothing could more fittingly end this book, devoted as it
is to demonstrating the operative nature of organic unity
in one of its great segments, than a reference to the fact
that the philosophy of life now determining German morals,

Postscript 355

and which has drawn its inspiration largely from the hypo-
thesis of natural selection, has failed — pathetically beyond
the power of words to express if done unintentionally; and
criminally in equal measure if done intentionally — to under-
stand the real meaning of Darwin's teaching as a whole.

Certain it is that had the German philosophers of Macht-
politik recognized the place of unqualified supremacy as-
cribed by Darwin to the mental and moral endowments of
man, it would have been impossible for them to make the
dogma of survival of the fittest serve their ends in any such
way as they have made it, and done so honestly. Attentive
reading of The Descent of Man makes it perfectly plain
that Darwin simply accepted all the higher human attri-
butes— moral, esthetic, and religious, no less than those of
the intellect — as fundamental data in his reasoning about
man's evolution. His sole effort as touching these was merely
to see in how far they could be regarded either as helped
forward in their development by natural selection, or at least
as not inconsistent with it. Apparently it never even oc-
curred to him to regard his hypothesis as supreme-over-all,
so that all attributes whatever, the noblest ones of man with
the rest, must either be forced into conformity with it, or
their reality and power virtually denied. "I fully subscribe
to the judgment," runs the opening sentence of the chapter
on "The Moral Sense," etc., "of those writers who maintain
that, of all the differences between man and the lower ani-
mals, the moral sense or conscience is by far the most im-
portant." And, especially significant at this time, Darwin
quotes with obvious approval, an apostrophe to Duty by
Kant, in which this "Wondrous thought" is represented as
"holding up its naked law" in the soul, and demanding
reverence. Darwin's entire discussion in this part of the
Descent makes it clear that what he had in mind was to
discover as far as possible the germs of "conscience," of
"feeling of right and wrong," of an "inward monitor," of

356 Postscript

"sympathy," of "parental and filial affection," of "social
affection," of the "instinct of self-sacrifice" and so on, in the
lower animals so as to have a starting point for these attri-
butes as they occur in civilized man. It was not at all his
purpose to show, as the German perversion of the struggle-
and-survival hypothesis holds, that the evolution of man
has consisted largely in a farther differentiation and intensi-
fication of the dominantly brute attributes, with an infusion
as a kind of by-product from the struggle for existence, of
certain "humanistic sentimentalities," which in reality are
signs of weakness and must be suppressed.*

And this perversion by German science and philosophy of
Darwin's teaching is rooted very deep in German culture and
character. The straightforward, common-sense descriptions
and inductions of the practical-minded, country-dwelling,
country-loving, unacademic English naturalist were alto-
gether too simple and unsophisticated to satisfy a Kultur
permeated through and through with the "highly strained
idealism" of Kant, Fichte, Hegel, and Schopenhauer. The
two worst errors committed by Darwin were his over-em-
phasis on the natural selection hypothesis, and his pro-
pounding of the gemmule-pangenesis hypothesis; and it is
highly characteristic that it was in just these two "strained"
speculations that German biology and practical philosophy
should have taken up Darwinism the most ardently and over-
worked it the most absurdly and disastrously.

My examination of the germplasm-determinant theory of
Weismann in Part I of this book has revealed something of
the scope and nature which the gemmule fallacy was destined
to assume when it fell subject to German speculation. The
more subtle and far-reaching and humanly practical conse-
quences of the adoption and elaboration of the struggle-and-

* The effort which Dr. George Nasmyth has made in his book Social
Progress and the Darwinian Theory to set right Darwin's position in
this matter, ought to bear fruit after a while.

Postscript 357

survival hypothesis by German speculation has not yet been
subjected to thorough-going biological criticism, though sev-
eral moves in this direction have been made.

Even the realism of recent German political and economic
theory and practice is a "highly strained" speculative real-
ism. This philosophical monstrosity is largely attributable,
demonstrably so I believe, to a cultural and governmental
system in which the principle of universal organic personality
is grossly violated. And what a price in misery and blood
and treasure the whole world, but old Europe particularly,
is paying for a consummation which a truer philosophy of
life would have foreseen and forestalled !

Can the leaders of German Kultur be convinced of the
fundamental fallacy of their theory of human and national
life, only by discovering that their military establishment,
built up through many decades of patient, costly organiza-
tion and discipline, but under guidance of a philosophy of
mechanism and brutism, is yet incapable of overpowering a
military establishment, a large portion of which may be im-
provised in the course of a few months, if such improvision
be under guidance of a philosophy of personality and hu-
manism ?

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GLOSSARY

ACROGEMALY. A disease charac-
terized by hypertrophy of the
terminal parts of the body, as
of the face and extremities ;. an
outgrowth involving bony and
soft parts.

ADRENALS. A pair of small glands
situated in front of the kidneys.
They are glands of internal se-
cretion, their secretion exercising
in particular a regulating effect
on the nerves of the heart and
blood vessels.

ADRENIN. The "active principle"
in the secretion of the adrenal
glands.

ALCYONARIA, include many corals
and other coelenterate animals
with eight mesenteries and eight
tentacles.

ALVEOLAR. In anatomy, a numer-
ously-pocketed, or sacculated,
structure, typified by the ter-
minal cavities of the lungs, but
occurring in various tissues; be-
lieved to constitute also one kind
of protoplasmic structure.

AMOJBA. A unicellular animal, a
genus of rhizopodous Protozoa.

AMPHIBIAN. An animal living
both in water and on land.
Properly a class of vertebrates
whose young are typically aquat-
ic and respire by gills; examples,
frogs, toads and salamanders.

.\MPHIOXUS, literally pointed, or
sharp at both ends. The cur-
rent name for one of the very
simplest and lowest vertebrate
animals occurring in the sand
and mud of the seashore in
many parts of the world.

ANABOLIC, the chemical up-build-

ing of the living body; construc-
tive metabolism.

ANTIBODIES. "The products of a
reaction of the body towards a
natural or artificial introduction
into it of certain foreign sub-
stances, bacteria and their poi-
sons, vegetable poisons of other
kinds, and various albuminoids."
The name antibodies has refer-
ence to the antagonism thess
products have for the introduced
substances.

ANTIGENS. Substances the reac-
tion of which with the living
body produce antibodies.

ASCIDIANS. Marine animals hav-
ing a gelatinous or leathery en-
velope containing cellulose. In
the larval stage a notochord or
forerunner of the vertebral col-
rmn is present. Some free liv-
ing species retain the notochord
all through their lives.

AXON. The long, slender, sparce-
ly-branched, fibrillar process of
a ganglion cell; contrasted with
the shorter, more branched, more
irregular dendron.

AXOSTYLE. A slender, flexible rod
of organic substance forming a
supporting axis for the body in
some Protozoa.

BIOGEN. Literally life producer.
Imaginary ultimate units of life.
Such special significance as the
"biogen theory" has over other
theories which make imaginary
vital or physiological units a
goal of the ultimate explanation
of life, is found in the fact that
the biogen theory aims to be
more definitely chemical than

377

378

Glossary

the others. The German physi-
ologist Max Verworm has elab-
orated this speculation more
fully than has any one else.

BIOPHOR. Literally life carrier.
Biophors, the imaginary ultimate
vital units of the Weismannian
system of speculative biology,
differ from biogens in the fact
that Weismann, not being a
chemist or even a physiologist,
but a zoologist interested in re-
production and heredity, rather
than in function generally, did
not undertake to put his specu-
lation on a chemical basis.

BIOPLASM. Formative living mat-
ter; not differing in any but a
speculative way from proto-
plasm.

Bionc. Pertaining to living
beings.

BLASTOGENESIS. Reproduction by
budding, as used in this book;
in general, propagation from an
undifferentiated germinal mass.

BLASTOMEHES. The first segments
or cells formed by the division
of the ovum.

BLASTOZOOIDS. The united individ-
uals produced by budding, and
constituting the colony, or cor-
mus in the compound ascidians.

BLASTTJLA. The stage of develop-
ment of the embryo from the
ovum in many animals, in which
the organism consists of a hol-
low sphere the wall of which is
composed of a single layer of
cells.

BRYOZOA. Literally "moss ani-
mals," from the resemblance,
fancied more than real, of some,
of the species to mosses; also
called Polyzoa. Marine animals
occurring abundantly on all
shores. Most of the species
propagate by budding as well as
by eggs and sperm, the bud-pro-
duced individuals remaining at-
tached to one another to form
colonies, as in many hydroids and

ascidians. Each individual con-
sists of a body proper bearing
a circle of tentacles, and an en-
veloping case often calcareous,
into which the body may be
quickly and completely re-
tracted.

CALYMMA. The much-vacuolated
portion of the body of radio-
laria, situated outside the central
capsule, the vacuoles containing
fluid impregnated with gas. The
main office of the structure
seems to be in connection with
the flotation of the animals.

CAMBIUM. The layer in woody
plants between the outer dead
layer, or bark, and the inner
dead mass, or wood proper, from
which new tissue is formed; the
true growing part of plants
which live several years and at-
tain a large size.

CARTESIAN PHILOSOPHY. The mode
of viewing man and nature in-
augurated in modern times by
Rene Descartes. The most dis-
tinctive thing about it is the
sharpness with which the dual-
ism, or antithesis, between mind
and matter stands out in it. Its
great practical importance for
the present era lies in its genetic
relationship to psycho-physical
parallelism in psychology, and to
all forms of idealism in philos-
ophy.

CENTROLECITHAL EGGS. Eggs in
which the protoplasmic portion
and nucleus constitute a surface
layer, the inner mass being
chiefly yolk, that is food ma-
terial for the future embryo.
The eggs of most insects and
crustaceans are of this type.

CEPHALIZE. The tendency among
animals for a head to become
differentiated from the rest of
the body.

CHEMICAL MESSENGERS. Substances
produced by the organism,
either in special glands, the

Glossary

379

glands of internal secretion, or
in general tissues, and carried
by the blood and lymph over the
body generally, to influence the
growth or functioning of other
tissues. Hormones is another
name given to these substances.

CHEMOTROPIC. Pertaining to the
reaction of organisms to chem-
ical stimuli.

CHROMATIN. The finely granular
substance most distinctive of the
cell-nucleus. Its name comes
from the readiness with which
it is colored by many dye-stuffs.

CHROMATOPHORES. Pigment-bear-
ing sacs, often single cells, in
plants and animals. It is by
means of these that the rapid
color changes in the skin of
many animals are accomplished.

CHROMIDOSOMES. One of the many
names given to minute specially
stainable bodies in the cytoplasm
of many cells.

CHROMOSOMES. The more or less
definite bodies of the cell-nu-
cleus into which the chromatin
granules are grouped. Their
constancy of structure and rela-
tion to hereditary characters
have given them great promi-
nence in much of recent biolog-
ical theory.

CHROMOPHIL. Any body in the liv-
ing organism that has an avidity
for staining reagents.

Coccus. In the classification of
bacteria according to their
shapes, those which are spherical
are called cocci.

COELENTERATA. A group of ani-
mals that have a digestive cav-
ity, but nothing corresponding
to the abdominal cavity; also
called radiata from the radial
arrangement of the body.

CORTICAL. Pertaining to the cor-
tex, or outer layer of an organ,
as of the brain, bark of the tree,
&c.

CRETIN. An individual aifected

with cretinous disease, a disease
characterized by certain bodily
deformities and mental impair-
ments. The malady frequently
accompanies goitre, and is now
considered due to deficiency in
secretion of the thyroid gland.
CRUSTACEA. A class of inverte-
brate animals belonging to the
great phylum arthropoda, briefly
characterized by their exoskele-
ton and paired jointed appen-

CYTOLOGY. Science of the cell.

CYTOPLASM. Substance of the cell-
body, as opposed to the cell-nu-
cleus.

DETERMINANTS. That particular
class of imaginary ultimate vital
units by which the development
of hereditary attributes is deter-
mined. They were invented by
Weismann, and were conceived
to constitute the germ-plasm,
and to be located primarily in
the chromosomes of the egg and
sperm. In later speculation de-
terminer is used more frequently
than determinant — on some ac-
count that is not clear.

DIATOMS. An immense group of
aquatic, unicellular algae espe-
cially characterized by their
firm, box-like, regularly-shaped,
chitinous shell.

DINOFLAGELLATES. Literally or-
ganisms which are two-lashed,
. owing to the two flagella pos-
sessed by most of the species.
A group of aquatic unicellular
organisms almost as numerous as
the diatoms. The photo-synthe-
sizing power of living substance
possessed by these two groups,
and their enormous abundance
at and near the surface of the
bodies of water in which they
live, make them fundamentally
important for all the life of the
waters of the earth.

DISTAL. A common anatomical
term signifying away from a

380

Glossary

given point, usually some defi-
nite feature, as the attachment
of a muscle, taken as a point of
reference.

DORSO- VENTRAL. A term much
used in the anatomy of the
higher animals to signify a di-
rection from-back-to-belly of a
creature.

DUODENUM. The first portion of
the small intestine between the
stomach and the jejunum.

DYNAMIC CENTER. A phrase used
rather frequently in recent biol-
ogy, especially in the biology of
the cell, to express the concep-
tion that certain structures, as
the centrosome, are in someway
not clearly specifiable the "seat"
of various vital activities. The
phrase has some such implica-
tion for general physiology as
"nerve center" had and with
many still has, for nerve phys-
iology.

ECOLOGICAL. Pertaining to Ecol-
ogy, the science of organisms in
relation to their natural environ-
ments. This old but newly ap-
preciated and named branch of
the science of living nature, may
properly be regarded as the nat-
ural history of plants and ani-
mals modified to meet the mod-
ern demands of comprehensive-
ness and exactness in dealing
with a great province of nat-
ural phenomena.

ECTOPLASM. The outermost, some-
what denser layer of protop'asm
in many cells, especially in nrmy
unicellular animals. Opposed to
endoplafm, the inner, more fluid
mass. The ectoplasm in proto-
zoans corresponds to the skin of
higher animals. The presence of
a more or less sharply set-off
outer layer or membrane or skin
in all organisms whatever is
coming to be recognized as hav-
ing a more fundamental physio-
logical meaning than that of a

protection for the delicate parts
underneath, now that so much is
being learned by physical chem-
istry about surface phenomena.

ENERGY. Work and capacity to do
work. It is important to note
that work and capacity to work
necessarily imply some object,
organic or inorganic, to do the
work, and hence that when the
energy of a horse, or of a stream
of water, is spoken of the word
energy has a very different
meaning from what it has in
such a phrase as the "energy
conception" of nature or of the
organism, the implication in
these cases usually being that
energy is the real essence of
nature and of the organism, the
shape and other so-called static
attributes which all bodies pre-
sent, being only incidental and
mere appearances.

ENZYME. A chemical substance
produced by an organism, plant
or animal, to the end of bring-
ing about chemical transforma-
tion in other substances, but
without itself being transformed.
The ptyalin of saliva by which
starch is changed into sugar, is
typical. Enzymes play a very
great part, especially in diges-
tion and nutrition, in the physio-
logical processes of all organ-
isms.

EPIGENESIS. That theory of devel-
opment of the individual organ-
ism which holds the organs and
parts to be actually new produc-
tions, and not merely enlarge-
ments or actualizations of what
already existed, this latter con-
ception of development consti-
tuting the theory of preforma-
tion. Although these opposing
theories were debated with fury,
almost, some years ago, little is
heard about them now though
none of the particular problems
around which the discussions

Glossary

381

centered can be said to have
been solved.

EPIMORPHOSIS. The mode of re-
generation of organisms in
which a multiplication of cells
on the surface of injury is first
produced, then from this "em-
bryonal tissue," the new organ
or part is formed; contrasting
with morphalaxis, a mode of
new formation which consists in
a direct transformation of an
already existing part into the
new part.

FACTOR (in Genetics). A hypo-
thetical unit of structure or of
chemical composition, contained
in the germ-cell, which in some
way is held to condition the de-
velopment of a particular char-
acter in the adult, or of a com-
plex of characters which are
transmitted in constant associa-
tion with one another. Factors
are believed to interact with one
another in development, and at
times to be so "linked" that they
are only partially independent in
transmission.

FLAGELLTJM. A lash-like appen-
dage or large cilium serving as
an organ of locomotion in some
Protozoa and some bacteria.

FORAMIXIFERA. A class of rhizo-
podous marine Protozoans, usu-
ally having a porous shell.

FORMATIVE STUFFS. Hypothetical
substances which are supposed
to be formed in one part of an
organism and transported to an-
other part, there to produce, or
to influence the production of
new organs. For example, sev-
eral botanists have supposed that
the flower substance of some
plants is actually produced in
the leaves.

GAMETE. A reproductive cell
which unites with another repro-
ductive cell to form a zygote.

GASTRULA. That stage of embry-
onic development in many ani-

mals which consists of two germ-
layers inclosing a central cavity.
It is produced from the blastula
(which see) by the in-sinking of
one-half of this into the other.

GEMMIPAROUS. Producing gemmae,
or buds (reproducing by bud-
ding), applicable to both plants
and many animals.

GEMMULE. In the original and
proper sense a small aggregation
of cells set apart in the tissues
of some plants and animals,
notably in many sponges, for the
purpose of reproduction. In ori-
gin and structure gemmules are
more like buds than eggs,
though the end served is very
similar to that served by seeds.
In a secondary and wholly hypo-
thetical sense, gemmules are
imaginary, minute bodies given
off by all the tissue cells of an
organism and assembled in the
germ cells, there to cause the
development of the next genera-
tion. This taking of a very con-
crete name from botany and
zoology, and using it in a wholly
imaginary way to explain hered-
itary development was due orig-
inally to Charles Darwin, but
with more or less unimportant
variations of meaning has since
been resorted to by many of the
best known biologists.
This example indicates the great
importance for biology, espe-
cially for the biology of repro-
duction and development, of dis-
tinguishing between the same
terms used in a strictly objective
and descriptive sense on the one
hand, and in hypothetical, or
purely imaginary sense on the
other.

GENE. A term much used in pres-
ent-day genetical science, but ap-
parently not differing in any sig-
nificant particular from factor
(which see).

GEXETIC. Pertaining to genetics,

382

Glossary

evolutionary science dealing with
natural propagation and devel-
opment, the interest centering at
present in that portion of devel-
opment which is hereditary and
involves sex cells.

GERM-PLASM. Actually all the pro-
toplasm of the germ-cells which
participates in development; the-
oretically merely the small por-
tion of the germ-cells supposed
to be "hereditary substance."

GONAD. A mass of undifferen-
tiated, generative tissue from
which the male and female re-
productive glands originate.

GOXOPHORE. The ultimate gener-
ative zooid of a hydrozoan, giv-
ing origin directly to the genera-
tive elements.

HECTOCOTYLIZED. Applied to the
remarkably altered condition as-
sumed by one of the arms of the
male cephalopod to make it an
organ for impregnating the fe-
male.

HELIOTHOPIC. Responding to the
stimulus of sunlight.

HETEROMORPHOSIS. A kind of re-
generation in which the part pro-
duced is different from that
which was lost, as, for example,
when an antenna-like structure
grows in the place of an eye-
stalk, in some crustaceans when
the eye stalk is cut off.

HISTOGENESIS. The process of tis-
sue genesis, or production, from
undifferentiated cell masses, in
plants and animals.

HISTOLOGY. The science of tissues,
plant or animal; microscopical
anatomy.

HOHMOXE. Literally something
which excites or stirs up. Orig-
inally and strictly applied to
those internal secretions (which
see), the office of which is to
incite the parts on which they
act to greater activity. But in-
ternal secretions are also known
now which retard or inhibit the

action of the part they affect;
and to these it has been proposed
to apply the term chalone, that
which slackens. But some phys-
iologists use hormones as syn-
onymous with "internal secre-
tions."

HOMONYMOUS. As used in this
book, an anatomical term refer-
ring to the different members of
a series which differ more or
less, but still all have the same
general name. Thus all the
pairs of appendages of a lobster
are homonymous, or ambulatory
appendages originally, although
used for a variety of purposes
now.

HYDRANTH. One of the bud-pro-
duced polyps of a hydroid col-
ony.

IDIOPLASM. Literally plasm which
is very specially one's own. First
used to designate the hypothet-
ical part of the germ-cells which
is supposed to be alone respon-
sible for hereditary transmission.
Idioplasm may be regarded as
the historical antecedent of
'germ-pla#m (which see).

INTERSTITIAL. Pertaining to or sit-
uated in an intervening space; a
term much used in anatomy to
signify within an organ.

IXTEHXAL SECRETION. The term
has long been used in the phys-
iology of the higher animals, in
contradistinction to "external se-
cretion," to designate the prod-
ucts of glands, like the thyroid,
which discharge their products
into the blood or lymph, instead
of upon the surface of the body
or into the digestive or some
other cavity of the body. The
existence of internal secretions
was known long before anything
was known about their use;
hence this non-committal name,
so far as function is concerned.
The recent discovery of their
office has suggested the name

Glossary

383

hormone (which see) for them,
and has revealed their great im-
portance not only for physiology,
but for philosophical biology.

INVOLUTION. Literally inrolling, or
inwrapping. In descriptive biol-
ogy used to signify the return of
an organ to its original or nor-
mal condition after some violent
or pronounced deformation of it.
Sometimes, but apparently un-
justifiably, used as a synonym of
degeneration. Since the doctrine
of evolution has become prom-
inent in biology, a process the
opposite of evolution has been
thought by some to be necessary,
and to this involution has been
applied.

JELLY-FISH. In the interest of dis-
criminative knowledge, the habit,
rather common among people
who' have the opportunity to see
the transparent, somewhat gela-
tinous-appearing animals of the
ocean, of calling them all "jelly-
fishes" should be abandoned. The
name should be restricted, to the
regularly disc- or dome-shaped,
tentaculated animals belonging
to the coelenterate phylum, thus
enlarging the bounds of definite,
popular zoological information,
by recognizing that marine ani-
mals of several large and very
distinct classes have this general
consistency and appearance.

KARYOPLASM. A cytological name
referring to the substance or
plasm distinctive of the cell-nu-
cleus.

KARYOSOME. A small, discrete,
rather constant body which
stains readily, contained in the
cell-nucleus; frequently synony-
mous with nucleolus.

KATABOLISM. The down-breaking,
or descensive phase of metabol-
ism; the opposite of anabolism
(which see).

KINETO-NXTCLEUS. One of the nu-
clei in the two-nuclear protozoa

supposed to be concerned in
some special way with the move-
ment of the flagella or cilia of
these animals.

LAMELLAE, singular lamella. A
term much used in anatomy to
designate the thin plates, scales,
etc., that are so numerous and
varied in form and size in nearly
all organisms.

LARVA. Properly applied only to
stages in the lives of individual
animals which pass into succeed-
ing stages through a deep-sealed
metamorphosis, as for example
the grub or maggot of a fly, and
its transformation into the adult.
Larval stages and profound
metamorphoses are very common
and widespread in the animal
kingdom.

LIMULUS. The technical genus
name for the horse-shoe crab, an
animal of special interest to gen-
eral zoology in several ways.

MACRONUCLEUS. In the infusoria, a
group of protozoans, there is one
large nucleus and one or several
much smaller nuclei. The first is
called, from its relatively large
size, the macro-nucleus; the
others micro-nuclei. From the
behavior of the two kinds of nu-
clei at conjugation and division,
the micronuclei are known to be
intimately connected with these
processes, while the macronucleus
seems to be more concerned with
the nutritive functions of the
animal.

MANUBHIUM. In morphology a
part or organ which resembles a
handle; specially the clapper-
like, or handle-like portion of a
medusa which is found within
the "bell." The animal's mouth
is at the end of the manubrium,
and most of its digestive cavity
within the stalk of the manu-
brium.

MATRIX. In biology the ground
substance in which cells are em-

384

Glossary

bedded in some tissues, and
which is produced as a secretion
by the cells. It is one kind of
intercellular substance. The
opalescent, almost homogeneous
chief mass of ordinary cartilage
is a typical matrix.

MELANIN. A rather general term
in biology, especially in zoology,
applied to dark brown to black
pigments.

MEROTOMY. The automatic cutting
off of parts or segments in liv-
ing organisms.

MESENCHYME. Undifferentiated
mesoderm that produces con-
nective tissues, some muscles,
and certain other structures in
the animal body.

METABOLIC. Pertaining to metab-
olism, the process of chemical
building up and breaking down
in the living organism.

METAMERIC. Pertaining to the
longitudinal series of parts or
joints into which the bodies of
many higher animals, such as
earthworms, lobsters and fishes,
are divided.

METAPLASTIC. Pertaining to meta-
plasm — applied to changes which
cells sometimes undergo from
one plasmic type to another; also
applied to certain supposedly
lifeless inclusions in the proto-
plasm of cells.

METAZOA. Multicellular animals.

MICRONUCLEUS. See macronucleus.

MICHOPHYLE. In botany and zool-
ogy the aperture in the coats of
the ovule and ovum through
which the male fertilizing cell
penetrates.

MONERA. Hypothetical simple
structureless masses of proto-
plasm (without any nucleus).
Assumed by Haeckel as the low-
est members of the evolutionary
series. Advance of knowledge
has found no evidence of such
organisms.

MORPHALLAXIS. A kind of regen-

eration in which part of an or-
ganism transforms directly into
a new and different part.

MORPHOLOGICAL. Pertaining to
morphology, the science of form
and structure.

MORULA. A stage in the embryonic
development of many animals, in
which the ovum has completely
segmented, but the segmentation
cavity has not yet been formed.

MYONEME. A thread-like contrac-
tile structure in the cytoplasm of
certain higher protozoa.

NEMATOPHORE. A body of defense
and offense developed in certain
hydroids, consisting of a chitin-
ous receptacle in which thread-
cells are immersed; the nettling
organs on the tentacles of large
jelly-fishes.

NEURAL. Pertaining to nerves.

NETTHOBLASTS. Undeveloped' nerve
cells.

NUCLEO-PLASM. Nuclear substance,
including the different nuclear
ingredients.

NucLEb-pROTEiN. One of the com-
pounds of nucleins and paranu-
cleins.

(EDEMA. Dropsy, a vasomotor
neurosis characterized by non-
inflammatory swellings on vari-
ous parts of the body.

ONTOGENY. The development of an
individual organism from germ
to completed or adult stage.

ORGANELLE. A little organ, and
organoid, organ-like, are terms
applied to the organs of unicel-
lular plants and animals, not so
much because of their small size
and indenniteness of form and
structure as on account of the
theory that a true organ must
be composed of cells, and cannot
be a part of a cell. These terms
are among the sequelae of the
cell-theory.

ORIENTING. Finding or fixing the
positions or directions.

PANGEN, and PANGENESIS. These

Glossary

385

terms, basal in Darwin's famous
hypothesis of heredity, mean all-
generator and all-generative only
in the sense that all parts of the
body of the organism give off
gemmules (which see), which
assemble in the germ-cells to en-
able these to reproduce the or-
ganism. Thus the pan, or all-
generative power was conceived
as having its original "seat" in
the organism all-in-all. In other
words Darwin's speculation was
almost diametrically opposed to
the transformation it has under-
gone latterly, especially in the
prolific mind of Weismann, the
germ cells alone, or rather the
germ-plasm being the all-genera-
tor, according to these specula-
tions.

PARATHYROIDS. Small glands lying
near the thyroid but not func-
tionally connected with the lat-
ter.

PARTHENOGENESIS. Reproduction
by means of unfertilized eggs.

PELLICULA. The cuticle or outer-
most body membrane in some
unicellular and other low organ-
isms.

PHASE. This old and familiar word
has taken on new and greater
importance, both scientific and
philosophic, with the recent ad-
vance of knowledge in the region
of over-lap between physical and
chemical phenomena, this ad-
vance making what is generally
called physical chemistry. A
phase in pure physics, as it may
be called, has reference to the
position of the particles of a
body when the particles are un-
dergoing change. For example,
corresponding particles in two
succeeding waves of water or air
are in the same phase. In phys-
ical chemistry phase has refer-
ence not to position but to state
or condition of the constituent
particles of a heterogeneous, or

unlike system. Thus, a combina-
tion of liquid water (in common
language just water) and solid
water, or ice, is a two-phase sys-
tem of water. Philosophically
viewed, the great significance of
phases is that the positions and
states of the particles are possible,
even conceivable, only in relation
to the larger, containing part or
whole. Something of the bear-
ing of this on the theory of plu-
ralism (which see), when this
theory is approached from the
strictly objective side, will be
easily seen.

PHLOGISTON. An imaginary sub-
stance formerly supposed to ex-
ist in all combustible bodies, and
to be the cause of fire and flame.
For nearly a century before the
discovery of oxydation as the
true cause of fire, by Lavoisier,
the phlogistic theory dominated
much of chemical science. The
chief interest in the theory now
is in its relation to the observa-
tional and logical processes in-
volved in interpreting the gen-
erative processes of nature
everywhere. The phlogistic the-
ory may be taken as a type of
elementalistic causal explanation
of natural production.

PHYLOGENIC. Pertaining to phy-
logeny, the development of the
race; concerning ancestral or-
ganisms, real and hypothetical.

PITUITARY GLAND. A gland of in-
ternal secretion, situated at the
base of the brain, and connected
in the embryo with the roof of
the mouth.

PLURALISM, philosophical (so used
in this book). The conception
that in its deepest nature the
universe is multiform and com-
plex; the opposite of Monism,
the conception that some single
Essence or Substance, more or
less known or unknown, is the
foundation of all things.

386

Glossary

PLUTEUS. Name given the charac-
teristic process-bearing larva of
sea-urchins and their near rela-
tives. These larvae are of con-
siderable general interest because
of the extensive use made of sea-
urchin eggs in experimental em-
bryology, the eggs being easily
obtained and easily kept in the
laboratory.

PROTEINS. Nitrogenous substances
found in the bodies of plants
and animals. These substances
are usually considered to be the
most fundamental, from the
chemical standpoint, in organic
beings.

PROTISTA. A group name intended
to include all unicellular organ-
isms; i.e., both protophyta, one-
celled plants, and protozoa, or
one-celled animals.

PSETJDOPODIA. Literally false feet.
They are temporary protrusions
of the protoplasm of some pro-
tozoa, especially of the rhizopo-
dous class, typified by the amoe-
ba, the name having reference to
the locomotor office of the proc-
esses. But their food-taking and
digesting office should be noted
also.

PTYALIN. The unorganized fer-
ment, or enzyme of saliva, chiefly
instrumental in the conversion of
starch into sugar.

RADIOLARIA. One of the main sub-
divisions of the protozoa, espe-
cially characterized by their gen-
erally spherical outline, and ra-
diating structures, some soft and
extensile, others stiff and per-
manent. The radiolaria are al-
most all marine.

REGULATION. Much used in studies
in the regeneration of organisms,
to express the power many
plants and animals have of un-
dergoing structural or func-
tional readjustments in order to
retain, or to regain, their typical
form ; a significant adaptation of

a general term to a technical
end.

RETICULAK. Net-like, a term much
used in anatomy, as many por-
tions in both plants and animals
of many grades, present this
type of structure, though the
netting never has the regularity
of manufactured netting.

RHLZOPODA. The great subdivision
of the protozoa especially char-
acterized by sending out pseudo-
podia (which see). Amoeba is
usually mentioned as the type of
this subdivision, but the larger
number, probably, of rhizopods
possess shells of one sort and an-
other, while . amoeba is entirely
naked during all its active life.

SAHCODE. Literally like flesh. The
name originally applied to what,
under microscopic examination,
seemed to be the fundamental
living-substance of animals. La-
ter discovered to correspond to
what was known as protoplasm
in the cells of plants.

SABCODYCTIUM. A protoplasmic
network of the surface of the
calymma of a radiolarian.

SELF-DIFFERENTIATION, SELF-REGU-
LATION, &c. It is not without
philosophical significance that
the term self has forced its way
into technical biology, something
as it has into technical philoso-
phy. In biology the term is par-
ticularly common in connection
with developmental phenomena
and has reference to operations
wliich depend primarily on the
organism itself, and can be re-
ferred to "external factors" only
remotely and in a round-about
way.

SERICTERIES. Glands by which silk
and silk-like substances are se-
creted in many insects.

SOMA, SOMATIC. The body and
pertaining to the body. Much
used in later discussions of
heredity in a strongly hypothet-

Glossary

387

ical sense, to indicate the com-
plete independence, so far as
development is concerned, of the
body from the germ. The an-
tithesis is often made stronger by
speaking of the substance of the
body and the substance of the
germ, using the terms somatic-
plasm and germplasm. From
the philosophical standpoint it is
instructive to compare the the-
oretically complete separation of
body and germ in modern gen-
etics, with the theoretically com-
plete separation of body and
soul in philosophy and psychol-
ogy-

SPECIFICITY. The state of being
specific, that is, of being mani-
fested as phenomena distinguish-
able from all other phenomena.
The group of terms kindred to
specific and species, long import-
ant in systematic and taxonomic
biology, are becoming increas-
ingly so with the advance of
knowledge, especially in the do-
mains of the chemistry of differ-
ent kinds of organisms, and of
comparative behavior and psy-
chology.

SPORAZOA. One of the main sub-
divisions of the protozoa a lead-
ing characteristic of which is in-
dicated by the name, that char-
acteristic being the commonness
with which propagation occurs in
the group by means of spores
produced within the body of the
animal. By far the greater num-
ber of the species of the group
are parasitic, many of them dis-
ease producing.

SPORULATION. The process of con-
verting into spores, as in the
sporazoa, in some other animals,
and in many plants. Spores dif-
fer from eggs, on the one hand,
and seeds on the other, only in
the fact that spores are not sex
cells, that is, do not need to unite
with other cells in order to de-

velop, as is the case with most
eggs and seeds.

SPONTANEOUS GENERATION. The
difference between "spontaneous"
in this phrase and in the phrase
"spontaneous action," as of an
animal, should not be thissed. In
the latter connection the word
has nearly if not quite its orig-
inal meaning, that is, of one's
own accord, or initiative; acting
by and through one's self alone.
The adjective pronouns mea, tua
and sud are said to have been
used always with sponte in good
Latin prose. Strictly, then, if
life really originated from some-
thing which was not living, that
is by a "fortuitous" concourse or
interaction among chemical ele-
ments of different sort, spon-
taneous would not be the proper
term .to describe the operation,
simply because it would have in-
volved fundamentally several
selves, even if the different ele-
ments could each be called a
self. It would not have been an
operation identifiable by my,
your, or his or its, but by their.
Plurality rather than singularity
of action would be the essence
of the conception.

STEREOTROPIC. Reacting to stimuli
of contact with solid objects.

STOLON. A prolongation of the
body of some plants and animals
that gives rise to new individuals
by budding.

StTDORiPARous. Sweat-producing.

SYMBIOTIC. Pertaining to a state
of living together of two dissim-
ilar organisms to the advantage
of both.

SYNAPTIC. In cytology pertaining
to synapsis, the conjugation of
chromosomes in sex cells preced-
ing the reduction divisions con-
nected with the maturation of
germ cells.

SYNCYTIUM. As used in this book,
a cytological term applied to a

388

Glossary

protoplasmic mass containing
many nuclei, but not set off into
distinct cells. The entire em-
bryo is of this character in some
animals. In fact a few embry-
ologists have contended that dur-
ing the embryonal stages of
most, if not all animals, the cells
are connected by protoplasmic
strands and bridges, making
them syncytia. The undoubted
wide prevalence of syncytial
structure among animals espe-
cially, has been used as an argu-
ment against the cell-theory.
SYNTHESIS. From the organismal
standpoint not many terms used
in biology are more important
than this. The etymological
meaning, placed or put together,
expresses only a part of the to-
getherness of an organism; the
part, namely, which pertains to
the assimilative activity per-
formed by the organism on its
nutritive substances. This proc-
ess may be regarded as a syn-
thesizing one in nearly the lit-
eral sense (though even here the
process is more one of self-ac-
tivity and less one of external
agency than seems to be implied
in the original word). But it is
when we come to consider the
original nature and power of the
organism by virtue of which it
assimilates food, that the inad-
equacy of synthesis, except in a
much modified sense, comes to
light, for the organism's ability
to assimilate, that is to put or
place together, its nutritive sub-
stances is wholly dependent, so
far as we have evidence, on the
fact of its being already and
originally a together entity. An
organism is able to put together,
or synthesize, its food just be-
cause it itself is together, or syn-
thesized. A synthesized state is
a prior condition to synthesizing.
To be an organism at all is to be

synthesized.

SYSTEMATIC. Pertaining to a sys-
tem; literally a standing or being
together. It is unfortunate that
"systematic" has come to be re-
stricted in its application in re-
cent biology to the formal classi-
fication of plant and animal
species. As a matter of fact a
necessary consequence of the
unity of all phenomena of the
living world is that all these
phenomena "stand" in some nat-
ural and ascertainable relation
with all other phenomena, so
that all biological knowledge
whatever must of necessity be
systematic if it really corre-
sponds to nature.

TAXONOMY. Mode of arrangement,
the branch of biology which
deals with the classification of
the species of plants and ani-
mals.

TEST. As used in zoology and bot-
any, an external covering or
tunic, usually nearly lifeless,
tough and resistant. Its office is
mostly protective.

THYMUS. A gland of internal se-
cretion found in the neck region
in all vertebrates, and connected
originally with the gill system.

THYROID. One of the most impor-
tant glands of internal secretion,
located, as is the thymus, in the
neck region, but connected em-
bryonically with the pharynx
rather than with the gills proper.

TROCHOPHORE. A larval stage in
the lives of many marine worms
and molluscs, characterized by
being well organized for swim-
ming by means of cilia variously
disposed on the surface of the
body.

TRYPSIX. One of the chief "active
principles," or enzymes of pan-
creatic juice. It splits proteids
into simpler compounds. It is
produced by some plants as well
as many other animals than man

Glossary

389

and vertebrates.

TUNIC. In botany and zoology,
any well differentiated membran-
ous covering of an organ or an
entire organism; much the same
as a test.

TUNICATE. Name of a group of
marine animals, most sharply
characterized by the cellulose-
containing tunic, or test, which
envelopes the body; by the pe-
culiar basket-like respiratory sys-
tem; and by the notochord or
precursor of the vertebral col-
umn, possessed by all the species
in the embryonal life, and by a
few during the whole life, fre-
quently used synonymously with
Ascidian, which see.

VAGAI,. Pertaining to the vagus
nerves, one of the tenth pair of
cranial nerves in all true verte-
brates.

VASO-CONSTRTCTOR. Applied to the
nerves which cause contraction
of the walls of blood vessels.

VASO-DILATOH. Applied to nerves
which cause, or more exactly,
permit a widening of the blood

vessels by diminishing the tonus
of the muscles of the vessel
walls. Since the smaller blood
vessels are all supplied with both
constrictor and dilator nerves
the constant balancing between
these antagonistic influences,
both kinds of impulse being in
response to the general needs of
the organism, this scheme illus-
trates well a principle of equili-
bration widely operative in the
animal kingdom.

VISCERAL. A term used in zoology
to indicate not only the totality
of internal organs, but also the
side of .the animal on which
these are situated.

ZYGOTE. A body formed by the
conjugation of two reproductive
cells, called gametes. Gametes
and zygotes may be either uni-
cellular organisms, or the repro-
ductive cells of multicellular or-
ganisms.

ZYMOGEN. The enzyme-producing
substance in the secretory cells
of glands the secretions of which
contain enzymes.

INDEX

Abderhalden, E., i, 102

Absolutism, metaphysical, ii, 151

Acacia, ii, 99

Accidental products of chemical
change, i, 111

Acorns, storing of, by woodpeck-
ers, ii, 269

Acromegaly, ii, 124, 137

Action, autocatalytic, ii, 107

Action-system, ii, 216

Activity, adaptive, excessiveness
of, ii, 257

Activities, instinctive, tendency to
excessiveness of, ii, 256, 268

Adaptation, of neural activity, ii,
183

Adaptive parallelism, i, 335

Adaptiveness, ii, 241 ; of subra-
tional psychic activities, 250

Adrenal glands, and nervous sys-
tem, ii, 131

Adrenaline, chemical composition
of, ii, 123

Adrenin, in blood, ii, 131; effects
on fatigued muscles, 181

Adsorption, meaning of, ii, 344

Adventitious buds, i, 38

Aggregation, principle of, i, 182;
of protozoa, 268; and synthesis,
ii, 236

Air, in relation to consciousness, ii,
290; breath, and spirits, 338

Albumen, living and dead, distinc-
tion between, i, 78

Alchemy, ii, 288

Alcyonaria, ii, 97

Aldrich, T. B., ii, 123

Alga, i, 41

Allen, B. M., on origin of sex-
cells, i, 73; on removal of thy-
roid, ii, 145

Alternation of generations, i, 316

American biologists and organis-
mal theory, i, 11

Amoeba, complexity of, i, 289; to
man, 291 ; as organism and as
cell, 297

Amours, of fishes, ii, 266

Amphibia, i, 204

Amphioxus, on isolated blasto-
meres of, i, 204; on graded
series in, ii, 106

Analysis, in biological reasoning,
ii, 206; and abstraction, 234; and
synthesis, 235; remarks on, 236;
of organic substance, 341

Anatomist, of protozoa, i, 286

Ancestral, i, 290

Anger, ii, 132, 317

Animal, odors of, i, 84; behavior,
ii, 208; human, 284; kingdom,
284

Animals, as analytical chemists of
one another, i, 88

Annelids, ii, 192

Antagonisms, ii, 175; within au-
tonomic nervous system, 178; co-
operative, 178

Anthropologist, ii, 285

Anthropology, ii, 227

Anthropomorphism, ii, 201

Anti-bodies, i, 100

Anticipatoriness, ii, 242

Antigens, i, 100

Ants, behavior of, ii, 257, 279;
larval, spinning of cocoon by,
311

Aphids, i, 353

Appendicularia, ii, 3

Apperception, relation to tropisms,
ii, 221; definition by Wundt,
233; real nature of, 243

Apples, odor of, i, 87

Argon, ii, 341

Aristotle, i, 2; ii, 278

Arrhenius, S., i, 101

Art, creativeness in, ii, 223; crea-
tive impulse in, 227

391

892

Index

Artificial parthenogenesis, i, 345

Ascidian, bud propagation in, i,
50, 309; egg development in, ii,
17; tentacles of, 98; ganglion
of, 187

Assimilation, ii, 205

Association in psychology, objec-
tive and subjective sides of, ii,
230

Associationism, ii, 228

Associationists, ii, 221 ; and ele-
mentalists, 228

Atomistic theory, ii, 151

Atoms, ii, 149, 160

Attention and choice, ii, 231

Attitudes, difference between sci-
entific and philosophic, ii, 307;
elementalist, and emotions, 321

Attributes, correlation of, i, 215;
ii, 202; physical or material, and
psychical or spiritual, 215; struc-
tural and functional, 277; eth-
ical, 284; observed corporeal,
289; physical and chemical, 305;
latent of oxygen, 341 ; latent,
343

Autocatalytic action, factor in
growth, ii, 107

Automaticity, ii, 221

Autonomic nervous system, ii, 128;
vagal or cranial, 129; sympa-
thetic or thoracico-lumbar, 130;
sacral, 130; antagonisms within,
178

Avoiding reactions, ii, 252

Axioms, ii, 297

Axones, ii, 170

Baboon, i, 98

Bacillus biitschlii, i, 262

Bacteria, i, 310; membrane and
surface structure, 257; un-
doubted organisms, 263; classifi-
cation of, 266

Balance of organs, i, 7

Balanoglossus, i, 223

Balfour, F. M., i, 268

Ballowitz, E., ii, 2

Barker, L. F., on interrelation be-
tween internal secretions and
nervous system, ii, 130, 138

Basedow's disease, ii, 138

Bateson, W., i, 22

Bayliss, on accidental chemical
products, i, 112; and Starling,
ii, 120; on meaning of autonomic
nervous system, 129

Bearers, i, 306; of heredity, 338

Bees, honey, ii, 268

Behavior, complexity of, i, 289;
animal, ii, 208, 210, 227; be-
havior-knowledge, 277

Behring Sea, ii, 211

Benda, C., ii, 35

Benecke, W., i, 258

Bernard, Claude, ii, 149

Bichat, i, 4

Bio-chemical substances, phylog-
eny of, i, 110

Bio-chemistry, and taxonomist, i,
94

Biococcus, i, 22; 319

Biogen conception, i, 194

Biogenesis, theory of, i, 27; versus
spontaneous generation, 316

Bio-integration, types of, ii, 94

Biologist, anthropological, ii, 335

Biology, real, i, 5; goal of, ii,
152; elementalist, and associa-
tionist psychology, 228; subdivi-
sions of, 283

Birds, high flight of, ii, 258; song
habits of, 260; mating habits of,
263

Blastomeres, i, 203; position in the
whole, 206

Blastula, i, 203

Blepharoplast, i, 255, 329

Blood, and bloods, i, 91 ; cor-
puscles, white, 297; adrenin in,
ii, 131

Blue-jay, storing habits of, ii, 270

Body, i, 321; ii, 150, 215; relation
to mind, 216; constitution of,
289; vs. corpse or cadaver, 322;
and soul, 323

Born, G., i, 207

Botanical, diagnosis, i, 265

Botany, elementary instruction in,
i, 236

Boyle, Robert, ii, 288

Brain, not coordinating center, ii,
191; normality dependent on,
194; as element of organism, 216

Index

393

Brandt, Percy, i, 39

Bread, "secret powers" of, ii, 300

"Breath of Life," ii, 303, 336,

338

Brown, A. P., i, 95
Briicke, E., conception of the cell,

i, 129
Bud, adventitious in plants, i, 38;

propagation in compound ascid-

ian, 50, 309; in bryozoa, 53
Burrows, M. T., on tissue cultures,

i, 173; on organ formation in

such cultures, 176
Butterflies, larvae of, ii, 239

Calkins, G. N., i, 240

Cambium, buds from, i, 39

Canary birds, ii, 261

Canidae, i, 96

Cannon, W. A., on chromosomes
and Mendelism, i, 356; on
heredity in plant hairs, ii, 55

Cannon, W. B., on autonomic nerve
action and adrenin, ii, 129, 131,
162, 178, 185, 319, 323

Carbon, ii, 338

Carrel, Alexis, on tissue cultures,
i, 168, 174; isolated tissues and
"morphological plan" of organ-
ism, 177

Carrying, characters of adult, i,
224; hereditary qualities, second-
ary and acquired, ii, 67

Cassia, ii, 99, 106

Casteel, D. B., ii, 9

Castle, W. E., definition of hered-
ity, i, 315

Caterpillar, ii, 239

Caudal tube of spermatozoan, ii,
7

Causal factors, ii, 103

Cause, in heredity, i, 313; sufficient,
ii, 147; contributing, 153; un-
known, of experience, 303

Cell, as chemical laboratory, i, 82;
physical chemical conception of,
116; as an organism, Briicke,
129; as key to ultimate biologi-
cal problems, 163, 181 ; in logical
and factual aspects, 228; as
elementary organism, 228; ag-
gregations, 295; colonies, 295;

evolution of, Minchin, E. A.,
307

Cell-membrane, produced by pro-
toplasm, ii, 59

Cell-nucleus, and protoplasm, i,
126

Cell-state, i, 295

Cell-system, i, 216

Cell-theory, what it is, i, 150 et
seq.; inadequacy of, 11 and 158;
attempt to subordinate protista
to, 280 et seq.; and integration
of nervous system, ii, 169

Cell-wall, in higher plants, ii, 51

Ceils, subordinate to living beings,
i, 293; used by living beings,
294; isolated, 294

Cellular centers, i, 332

Centers, dynamic in cells, i, 333;
of apperception, ii, 233

Centrioles, i, 333

Centrosome, i, 330, 331

Cerebral cortex, ii, 216

Cerebrum, ii, 130

Ceremonies, self-exhausting, of
mating, ii, 264

Cha?tognatha, ii, 281

Chcetopterus, i, 12

Chain reflexes, ii, 197

Challenger Expedition, ii, 7

Characters, special and general
in heredity, ii, 40

Chemical action and interaction, i,
215; autocatalytic in organic
growth, ii, 105

Chemical basis of genus and spe-
cies, i, 107

Chemical, criterion of, ii, 289

Chemical messengers, i, 23; ii, 119,
121, 128, 170

Chemico-functional integration, ii,
94

Chemico-naturalist inquiries, i, 105,
109

Chemist, and naturalist, i, 107

Chemistry, and organism, i, 75; of
organisms, 91; comparative, 105;
and variation, 115; in solving
problems of heredity, ii, 42;
definition of, 287; physical, 303,
336; periodic law of, 329; atom-
istic, 343

Index

Chicken pox, i, 264

Child, C. M., and physiological cor-
relation, i, 17; and metabolic
gradients, ii, 108

Chipmunk, storing habit of, ii, 271

Chondriosome, ii, 36; as material
substratum of different tissues,
39

Chromatin, theory of, i, 314; rela-
tively undifferentiated, 318; evi-
dence of, as hereditary sub-
stance, 326; physicial basis of
heredity, 328; supposed omnipo-
tence in heredity, ii, 14; kinds
of, 67

Chromatinists, i, 319

Chromosomal elementalism, i, 320

Chromosomal hypothesis of hered-
ity, evidence ' for, i, 324, 326 et
seq.

Chromosome dogma, ii, 59

Chromosomes, i, 21, 306, 324; as
immediate ancestors, 319; in
fertilization, 342; accessory, 347;
X and Y, 350; seat of inheri-
tance material, ii, 22; in rela-
tion to heredity, 66; initiators in
heredity, 83

Chun, C., i, 301

Cilia, i, 330

Ciliary tuft of spirillum, i, 259

Classification, i, 99, 296; of
physical facts, ii, 217; synoptic,
276

Chemical evidence of adrenal-
nervous connection, ii, 133

Cohn, F., on classification of bac-
teria, 266

Collins, H. H., ii, 258

Common-paths in nerve physiol-
ogy, ii, 171

Common-sense, i, 32

Comparison, i, 99; of shells of
rhizopod and nautilus, 237;
sacrifice of, in experimental
method, ii, 279.

Competition, ii, 175

Condition, molecular appeals to,
i, 276

Conjugation, i, 269

Conklin, E. G., on egg as stage in
life of organism, i, 193; on

development of Ascidian egg, ii,
17; on hereditary characters de-
termined by cytoplasm, and by
chromatin, 42, et seq.

Coordination, neural, not a "cen-
tral" process, ii, 192

Corycella, i, 270

Consciousness, ii, 161 ; contents of,
233; organismal theory of, 282;
and chemical action, 290; theory
of, and theory of knowledge,
296; an attribute of the organ-
ism as a whole, 309; and
physico-chemical conception of
organism, 324; and pro-con- ;
sciousness, 350

Contents of consciousness, ii, 225,
233

Courtship of animals, 262

Cowdry, E. V., i, 437

Crampton, H. E., ii, 26

Crane, sand-hill, ii, 258

Crepidula, ii, 20

Cretin, ii, 116

Crickets, chirping of, ii, 261

Crithidia, i, 334

Ctenophore, i, 201

Cushing, H., ii, 113, 124

Cuvier, i, 5

Cycads, ii, 58

Cytoplasm, and Karyoplasm, i,
135; Kinds of, ii, 67; funda-
mental and primitive as heredi-
tary substance, 68

Cytoplasmic activity, in spicule
production, ii, 52

Cytoplasmists, i, 319

Cytostome, i, 248

Dances, of lapwing, ii, 262

Darwin, Chas., as naturalist, i, 75; ;
as example of creativeness in j
science, ii, 225; on comb of hive*
bee, 168

Davidson, H. C., on plant as sym-
biotic colony, i, 35; on "planto-
gens," 36

De Bary, and cell theory, i, 162

Definition, i, 296

Dendrites, ii, 170

Descartes, Rene, ii, 298

Descent, i, 315

Index

395

Description, i, 99

Determinants, Weismannian, i, 21,
225; changed to determiners, 348

"Determined," different meanings
of, ii, 49

Determiner, fascination of for
some minds, i, 306; meaning in
true objective sense, ii, 16; and
dialectics, 76; theory of, con-
trary to chemical principles,
79; Wilson's proposal to drop,
82

Development, in protozoa, i, 267;
cause of, ii, 158

Developmental mechanics, i, 18

Dewey, John, ii, 298; on "Self"
and environment, 305

Diagnosis, medical, i, 265

Dialectics, and determiner hy-
pothesis, ii, 76

Diatoms, i, 310

Difference, chemical, between or-
ganisms, i, 83; between germ-
cells, importance of, 214; re-
semblances and, 317; in func-
tion and behavior, ii, 276

Differential factor, ii, 82

Differentiation and integration, ii,
168

Dinoflagellates, i, 310

Directing activity, of develop-
mental process, i, 70

Display, mutual, ii, 264

Distribution, vertical, ii, 281

Division, of labor, i, 205; deter-
mined by growth, 220; physio-
logical, ii, 24

Dobell, C. C., on nuclei in bac-
teria, i, 262; on Ehrenberg's
conception of protozoa, 284; on
protozoa as non-cellular, 290

Dog, as causal explanation, ii,
203, 204

Donaldson, H. H., ii, 169

Doncaster, L., i, 352

Dormitive principle, ii, 204

Dramatist, ii, 317

Driesch, Hans, on cell theory, i,
153; totipotence theory of, 202,
et seq.

Ductless glands, ii, 114
Dujardin, Felix, interpretation of

protozoa, i, 280; and plasmic

elementalism, 320
Duodenal mucous membrane, ii,

119
Dynamic center, of cell, i, 333

Earthworm, ii, 191

K chinas, i, 202

Ecology, ii, 212, 279

Economy, physiological, ii, 261

Ectoderm, i, 46

Effect, i, 313

Ege, of chick, studied chemically,
i, 79; of frog, 199; floating, 213;
hereditary attributes of, 214; as
stage in development of indi-
vidual, ii, 24

Ehrenberg, C. G., interpretation
of protozoa, i, 280, and ii, 66

Elements, photosensitive, ii, 189;
physical and chemical, 235;
psychical, 235; chemical, cri-
terion of, 286

Elementalism, i, 2; narrowing in-
fluence of, 230; cellular, 286;
and internal secretion, ii, 141

Elementalist, conception, i, 280;
speculation, 319 ; and organismal
standpoints, ii, 148; theory, and
neglect of fact, 157; anarchistic,
160; attempt to interpret trop-
istic and segmental theories,
198; biology and associationist
psychology, 228

Elementalistic interpretation, ii,
23

Elementary organism, i, 227

Embryogeny, i, 277

Embryology, methods of, i, 224;
of protozoa, 268; and genetics,
311, 324

Embryo, i, 204, 272

Emotions, and physical organiza-
tion, ii, 216; natural history de-
scription of, 318, 322; elemen-
talist description of, 321

Emotional, attitude, i, 322; Glyco-
suria, ii, 132; psychic life of
animals, 133

Empedocles, i, 3, 40

Endoderm, i, 46

Endocrine glands, ii, 114, 130

396

Index

Endoplasm, i, 277

Energy, formative and regenera-
tive, i, 11; of modern physics,
76; and matter, 141; conception
of, 320; of contraction, ii, 63;
and substance, 337; and power,
force, work, 342; surface, 344

Entelechy, ii, 149

Environmental influence, ii, 244

Enzymes, digestive, comparative
chemistry of, i, 104; facilitate
transformation, ii, 81

Epicurus, i, 3

Epithelium, and internal secre-
tions, ii, 135

Eppinger, H., ii, 133

Equilibrium, general notion of, i,
17; physical chemistry concep-
tion of, 216

Essences, ii, 288

Esterly, C. O., ii, 208

Evidence, direct and indirect, of
mechanism of heredity, i, 325;
favorable to chromatin as hered-
itary substance, 326; visible and
invisible, ii, 56; picking out of,
ii, 263

Evolution, i, 291, ii, 241

Evolutions, four simultaneous, i,
321

Eudendrium, i, 67

Eugenics, i, 305; and fatalism, ii,
89

Excessiveness, of instinctive activ-
ity, ii, 259; of sex impulse, 265

Excitability, threshold of, ii, 165;
selective, 165

Excitor, ii, 147

Experience, ii, 305; subjective and
objective, 285; causes of, un-
known, 303

Experimentation, limitations of, ii,
212; laboratory, 279

Explanation, spurious type of, ii,
81; causal, 146, 240; of trop-
isms, 190

Expression of emotion, ii, 264

Factorial hypothesis in heredity, i,

21

Factors, in heredity, i, 42, 306
Facts, neglect of, by elementalists,

ii, 157; matters of, 299

Falta, W., ii, 134

Fatalism, ii, 89

Fatigue, and sugar in blood, ii,
132; and blood pressure, 181

Fear, and adrenin in blood, ii,
132; with physical and psvchical
life, 183

Feeling, intellect, will, ii, 217

Felidas, i, 96

Ferns, ii, 99

Fertilization, hybrid, 5, 344

Fielde, A., on odors in ants, i, 89

Finalism, ii, 152

Finch, house, ii, 260

Fishes, sperm of different species,
i, 102; floating eggs and young
of, 213; mating habits of, ii, 265

Flagella, of bacteria, 259; rela-
tion to nucleus, 328, 330; struc-
ture and origin of, 370

Flat-worms, ii, 109

Flavors, of animals and plants, i,
84

Fluid, as phase of system, i, 216

Foetus, i, 272

Forbush, E. H., ii, 270

Forces, antagonistic, ii, 127; con-
stitutively antagonistic, 134;
abuse of the term, 298

Fore-foot, horse's, and man's hand,
ii, 224

"Formative stuffs," i, 16; and in-
ternal secretions, ii, 142; of
Sachs, 147

Formi-determination, cytoplasmic,
ii, 14

Forsyth, Ruth, ii, 44

Fossil wood, ii, 58

Foster, Sir Michael, ii, 346

Fragmentation, and analysis, ii,
236

Freud, S., ii, 350

Friedlander, B., ii, 103

Frog, eggs of, i, 199; pigment
cells of, 339; sex determination
in, ii, 76; larva and thyroid of,
143; croaking of, 261

Fruit flies, supposed connection
between mutations and chromo-
somes in, i, 354

Fundamental, criterion of, ii, 201

Index

397

Fungus, i, 41

Fur-seal, development of sperm

of, ii, 4; migrations of, ii, 210;

mating habits of, 267
Fusibility of tissues, as test of

relationship, i, 143

Gall-stones, ii, 164

Gametes, i, 269

Ganglia, supra-oesophageal, ii,
191; as relay stations, 194

Gastrula, i, 203

Gates, R. R., i, 353

Gemmule, as method of asexual
propagation, i, 309; as specula-
tive entities (see term in glos-
sary)

Gene, a hypothetical entity in
modern genetics, i, 20; static
nature of the conception, 42;
historic antecedent of, ii, 84

Geneticists, modern school of, i,
324

Genetics, tendency to ignore em-
bryology, i, 311; inattention to
developmental facts, ii, 14;
thinking on, 72

Germ, nature of, i, 223; imaginary
structureless, 268

German Deep-Sea Expedition, i,
278

Germ-cells, in hydroids, i, 60;
promorphology of, 211; '^hrow
of" the soma, 319; subject to
metabolism, ii, 74

Germ-disc, i, 208

Germinal, continuity, i, 310; ma-
terial, 209; localization, ii, 16

Germ-layers, i, 45; subservient to
organism, 48; and the germ-
plasm theory of Weismann, 58

Germ-plasm theory, type effect of,
on observation, i, 66; implica-
tions of, 318; extreme form of
contrary to inductive science, ii,
76

Germ-regions, organ forming, i,
209

Giardia, i, 254

Gibbs, J. Willard, ii, 341

"Gifts," natural, ii, 235

Ginkgo, i, 330

Gladness, fits of, ii, 263

Glands, ductless, endocrine, ii, 114

Glycogen, i, 215

Goal of biology, ii, 152

Goette, Alexander, review and
criticism by Weismann on sex-
cells in hydroids, i, 62, et seq.;
organismal trend of results by,
68

Goitre, ii, 115

Gradients, direct and inverse, ii,
106; axial metabolic, 107

Grafts, i, 143

Granules, epithelial, ii, 135

Grasshopper, i, 357

Graves' disease, ii, 133

Grebe, great crested, ii, 263

Grennell, Joseph, i, 85

Groos, Karl, ii, 273

Growth, determines division, i,
220; integration, ii, 93, 94; cycle
of, 105; of an organism, 105;
explanation of, 107

Gudernatsch, J. F., ii, 143

Habits, mating, of birds, ii, 263;
of viviparous fishes, 265; stor-
ing, of honey bee, 268, of wood-
pecker, 269, of mammals, 271;
in nature, 278

Haeckel, E., on radiolaria, i, 235;
moneron theory, 256, 320

Haecker, V., on radiolaria, i, 236;
on ontogeny of radiolaria, 278

Hairs of higher plants, ii, 55

Half-embryo, i, 199, 201

Hallez, P., i, 218

Hardens, A., i, 104

Hargitt, C. W., i, 67

Harmer, S., i, 54

Harmonic, equipotential system,
i, 205; equilibrium, ii, 118

Harmony, i, 3; in health, ii, 127

Harrison, R. G., on tissue cul-
tures, i, 168, et seq.

Heidenhain, H., i, 332

Heinke, Fr., i, 213

Heliotropic, ii, 240

Helium, ii, 342

Hemoglobin, i, 95

Henderson, L. J., ii, 339

Herbart, as extremist in associa-

398

Index

tionist psychology, ii, 229

Herbst, C., i, 344

"Hereditary substance," contrary
to facts in hydromedusae, i, 68

Heredity, and elementalistic phil-
osophy, i, 20; nature of, 305;
stronghold of biological elemen-
talism, 305; effort to restrict to
sexual propagation, 308; denned
by E. G. Conklin, 308; com-
plex of causes, 313; definition,
314; chromatin in, 320, 321; and
sex, 348; cytological basis of,
349; spermatozoon's tail mani-
festation of, ii, 3; mitochondrial
theory of, 33; and sponge spic-
ules, 53; summary of informa-
tion on physical basis of, 64;
narrowing definition of, 85

Herlitzka, Amedeo, i, 205

Hertwig, O., theory of Biogene-
sis of, i, 27; experiments on
half embryos, 200; on centro-
some, 332

Hertwig, R., on cell-theory as ap-
plied to protozoa, i, 288; on
experimental determination of
sex, ii, 76

Higher Usefulness of Science, ii,
337

His, Wilhelm, i, 208

Histogenesis, and the mechanism
of heredity, i, 325, and ii, 32;
and species characters in adults,
43

Hobbes, Thomas, and sensational-
ism in philosophy, ii, 219

Holmes, S. J., an organism as
symbiotic community, i, 183; on
brainless frogs, ii, 195; on ac-
tivities of Amphithoe, 248; on
behavior of ants, 257

Hooker, D., i, 339

Hopkins, F. G., on the cell as a
chemical laboratory, i, 82; on
the physical chemistry of the
cell, 114 et seq.; on the cell
constituents essential to the cell
as a system of phases, 192; ver-
sus particular types of mole-
cules as an explanation of life,
194

Honey-bee, eggs and chromosomes
of, i, 352; storing habits of, ii,
268

Hormones, importance of, to or-
ganismal conception, i, 23; na-
ture of action of, ii, 121; rela-
tion to nerve action, 128; sup-
posed identification with "form-
ative stuffs," 142; integrative
office of, compared with that of
nervous system, 162

Hudson, W! H., ii, 261

Human being in one-celled stage,
i, 217

Hume, David, ii, 298

Hunter ciliates, i, 235

Huntsman, A. G., ii, 44

Huxley, Julian S., ii, 263

Huxley, T. H., on the physical
basis of life, i, 121 ; on the
cell-theory, 288, 296; on innate
ideas as conceived by Descartes,
ii, 298; against materialism, 302

Hydroids, germ-cells in, 60; sepa-
rated blastomeres in eggs of,
204; graded growth series in, ii,
97

Hypophysis, ii, 124; alliance with
thyroid and adrenals, 127

Hypopituitarism, ii, 113

Hypothesis, ii, 282; of conscious-
ness, 286; "working," 291

Idea, central of this book, i, 24
Ideas, atomistic and association
of, ii, 229; "relations of," in
Hume's system, 299; innate, 301
"Identical stuffs," i, 123
Impertinence, scientific, ii, 247
Individual, man, i, 31 ; tree, 31 ; ex-
altation of, 196; normal, ii, 205
Individuality, in the living world,
i, 30; Huxleyan, 43; of chromo-
somes, 85; of organism, ii, 111
Inheritance, i, 312; nuclear theory,
and cytoplasmic localization, ii,
22; of acquired characters, 24;
material — imitator rather than
determiner, 66; probability that
substance becomes such in each
ontogeny, 73
Inhibition of reflexes, ii, 176

Index

399

Initiative, mental, ii, 244

Initiator hypothesis in heredity,
advantages of, ii, 83

Inner mass, and outer layer, of
body, i, 301

Insane-like conduct from absence
of brain, ii, 194

Insect, eggs of, i, 220; sperm of,
ii, 9

Instinct, involves animal as a
whole, ii, 188; importance of as
zoological term; 247; variability
of, 251; and intelligence, 256;
food-gathering, 268; problem of,
284; "instinct actions," "instinct
feelings," 284; and physical or-
ganization, 310

Integratedness, of Oiardia, i, 255;
and equilibrium, ii, 198

Integration, growth, ii, 93; func-
tional, 113; distinction between
developmental and functional,
161; cellular, in reflex arc, 163;
and differentiation, 168; psy-
chic, 214

Integrity, organismal, i, 26

Intellect, feeling, will, ii, 217; and
instinct, 256

Interaction, chemical, i, 215

Intercellular substance, ii, 170

Interdependence, metabolic, ii, 104

Internal secretions, importance of,
to organismal conception, i, 23;
conception of, ii, 113; interrela-
tions of, 124; and epithelium,
135; and "formative stuffs," 142

Internal secretory systems, ii, 128;
and nervous systems, 128

Investigation of distribution of
sex-cells, i, 73; statistical of
animal behavior, ii, 280

Iron, ii, 339, 337

Irradiation, ii, 174

Isolated cells, i, 167

James, William, on human energy,
ii, 132; on associationist psy-
chology, 229; on consciousness
of self, 309

Jelly-fish, i, 235

Jennings, H. S., on ultra-Mendel-
ism, i, 42; on complexity of

protozoan behavior, 289; on trial

and error, etc., ii, 252
Johnson, H. P., i, 272
Jolly, J., i, 176
Jordan, H. E., ii, 60

Kant, E., self-conscious unity of
apperception, and transcenden-
talism, ii, 233

Karyoplasm, i, 135

Kelp, ii, 105

Key, cell as, to biological phe-
nomena, i, 229

King, Miss H. D., ii, 76

Knowledge, analytic and synthet-
ic, i, 25; -getting, ii, 213; ana-
lytic, 213; theory of, 296; nature
of, 297

Kofoid, C. A., i, 244; on soil amoe-
ba, 328

Kofoid and Christiansen, on neu-
rometer system in flagellates, i,
254

Kolle and Wassermann, i, 258

Laboratory, as an agency, ii, 212
Lamarck, i, 5, 75
Lancelet, ii, 95
Langley, J. M., ii, 129
Lapwing, dance of, ii, 262
Larvae, grafting together, i, 207;

specific characters in, 214; as

"carrier" of adult characters,

224

Law, of elements, ii, 160
Lead, ii, 341
Leaves, compound, ii, 99
Leeches, ii, 311
Leibnitzian theory, ii, 150
Lemon trees, i, 38
Lewis and Lewis, ii, 40
Life, ii, 286; vegetal, emotional,

rational and intellectual, 162;

tripod of, 182; subjective, 284
Like produces like, i, 315
Likenesses, functional, ii, 276
Lillie, F. R., as pre-organismalist,

i, 11; on "properties of whole"

in the embryo, 12 and 193
Lillie, R. S., i, 340
Limulus muscle, ii, 60
Lineus lacteus, i, 189

400

Index

Living beings, in nature, ii, 211

Living substance, i, 115

Living units, hypothetical, i, 19

Localization, by protoplasmic flow-
ing, ii, 19

Locust, i, 218

Locy, Wm. A., i, 280

Loeb, Jacques, i, 23; on relative in-
fluence of nucleus and protoplasm
on heredity, ii, 41 ; on identifica-
tion of internal secretions with
"formative stuffs," 141; on
"ultimate aim" of biology, 151 ;
and "organism as a whole," 185;
neglect by, of work of Sher-
rington and Cannon, 185; on
understanding of natural phe-
nomena, 207; organismal ten-
dency of tropism theoiy of,
240

Logic, pure, i, 22

Loomis, L. M., i, 85

Love, emotions of, ii, 265

Luciani, L., ii, 115

Lucretius, i, 3

Machines, living, ii, 252

Macrocystw pyrifera, ii, 105

Mammals, storing habits of, ii, 271

Manly, J. M., on "exuberant vital-
ity" of Shakespeare, ii, 223

Marceau, F., ii, 61

Marshall, F. H. A., ii, 79

Materialism, author's attitude to-
ward, ii, 207; Huxley against,
302

Mathematics, ii, 297

Mating habits, of birds, ii, 263;
of fishes, 265

Matter, and energy, in modern
physics, i, 76, 141 ; and force,
196; composition of, 288, 341;
generality of the term, 304

Mass action, ii, 344

McClung, C. E., i, 347

Mcllvane, Charles, i, 87

McMurrich, J., Playfair, on germ
layers, i, 47; character of cell
division in embryo, 219

Meadow lark, western, song habit
of, ii, 259

Mechanism of heredity, i, 315,
322 ; organic vs. inorganic, ii, 252

Medussetta, i, 236

Meirowsky, E., i, 339

Melanin, i, 339; formed in cyto-
plasm, 341

Membrane, and surface structure
of bacteria, i, 257; synaptic, be-
tween cells of reflex arc, ii, 167

Mendel, Gregor, i, 305

Mendelian inheritance and chro-
mosomes, i, 356

Mendelism as a creed, i, 324

Mental, sense, i, 3; initiative and
restlessness, ii, 243

Meristic parts, ii, 95; meristic phe-
nomena in plants and in ani-
mals, 103

Merotomy, i, 276

Mesenchyme cells, dormant in tad-
poles, ii, 147; as inheritance
material, 155

Messengers, chemical, secretin as
example of, ii, 119, 121

Mesoderm, i, 46

Metabolic processes, and the or-
ganism's supremacy over its
cells, i, 294; interdependence of,
ii, 104

Metabolism, i, 215; germ cells sub-
ject to, ii, 74; katabolic and
anabolic, 346

Metals, "base" and "noble," ii, 288

Metameres, ii, 95

Metamorphosis, ii, 145

Metaphysician, ii, 285

Metaphysics, chromatin and, i,
321; juvenile, ii, 141; as an epi-
thet, 201

Metaplasy, of differentiated cells,
i, 186 *

Metazoa, i, 268

Metcalf, M. M., i, 289

Method, experimental, ii, 278; sta-
tistical, employed at Scripps In-
stitution, 280; natural history,
in study of self, 282; impor-
tance of, 282

Meves, F., ii, 35

Meyer, Arthur, i, 258

Mice, summersaults of, ii, 258

Michael, E. L., ii, 281

Microbes, specificity of, i, 265

Miescher and Kossel, i, 79, 102

Migration, of sex-cells, i, 61 ; ex-

Index

401

tent of, by birds, ii, 259

Milk, i, 104

Minchin, E. A., Biococcus theory
of, i, 2; on evolution of cell,
307; as chromatinist, 318

Mind, relation to body, ii, 215,
216; "mind stuff," SIS

Miracles, i, 322

Mitochondria, in ontogeny of in-
sect sperm, ii, 9; not trans-
formed into neuro-fibrils, 37

Mitochondrial theory of heredity,
ii, 33

Moeser, W., ii, 107

Molecular condition, appeals to, i,
276, 282

Molecules, ii, 149

Mollusca, i, 221

Moore, V. A., i, 259

Monogamy, in viviparous fishes,

11, 265 '

Moneron theory of Haeckel, i, 256,
320

Morgan, C. Lloyd, ii, 328

Morgan, T. H., and regeneration,
i, 180; on half -embryos of frog,
200; on eggs of phylloxerans,
353; on mutations and heredity
in fruit flies, 354

Morphallaxis, i, 180

Morphological, plan of the organ-
ism, i, 177, 195; entity, funda-
mental to tropism conception, ii,
190

Mortensen, T. H., i, 216

Mosaic, theory of development, i,

12, 198, 205; "mosaic picture,"
organism as, 208

Mosquitoes, i, 213
Mosses, leaves of, ii, 57
Mushrooms, poisonous species of,

i, 87

Musk-deer, i, 86
Mustard plant, weight of seeds of,

ii, 102
Mutation, connections of, with

chromosomes, i, 353
Myomeres, ii, 95, 106
Mystification of protoplasm, i, 121

self, ii, 282; importance of, 283

Natural selection, and selected
evidence, ii, 263; slight regard
for quantity by, 259

Naturalism, ii, 142

Naturalist, and evolution theory,
i, 76, 286; ii, 224; zoological,
227, 278

Nature, of things, i, 34; not sim-
ple, 236; of knowledge, ii, 152

Nautilus, comparison of, with
rhizopod, i, 237

Needs, nutritional, ii, 268

Neglect nothing, naturalist's mot-
to, i, 94; ii, 216, 245, 283

Nemertean, regeneration of, i,
189; capture of prey by, ii, 255

Neoformation, i, 276

Nereis, ii, 192

Neresheimer, E. R., i, 245.

Nerve, centers, ii, 187; physiology
of, 320

Nervous system, in protozoa, i,
243; autonomic, ii, 128; and in-
ternal secretions, 128; integra-
tive action of, 162 et seq.;
Loeb's important conception of,
187

Neural integration, ii, 94, 161

Neurones, ii, 216

Neurophanes, i, 245

Neuro-motor apparatus, i, 243;
system, 255

Nickel, ii, 327, 329

Night-hawk, ii, 258

Nomenclature of germ-plasm the-
ory, ii, 87

Non-cellular, protozoa interpreted
as, i, 290

Nucleus, i, 207, 331 ; in bacteria,
261 ; part in pigment formation,
340; in oxidizing action, 341; of
"utmost theoretical importance,"
ii, 22; control of cytoplasm by,
22

Nusbaum, J., on tissue trans-
formation, i, 190

Nusbaum and Oxner, on regenera-
tion of nemertean, i, 189

Natural history, special ability of,
i, 113; methods of studying the

Objective, side of psychical asso-
ciation, ii, 230; and "outer," 292

402

Index

Observation, method of, ii, 278

Odors of animals and plants, i, 84

(Edema, ii, 116

Oil-drops and globules, in eggs, i,
213, 215

Oliver, J. R., ii, 7

Omnipotence of chromatin in
heredity, ii, 14

One-cell stage of organism's life,
i, 214

Ontogeny, misuse of term, i, 271 ;
of protozoa, i, 277; effort to ex-
plain on elementalistic princi-
ples, ii, 158

Optic nerve, ii, 164

Organ, i, 245

Organ-forming substances, in the
ovum, ii, 16; theory of, 141

Organ-germs, i, 209

Organelles, i, 248

Organic formation, and isolated
fragments, i, 176

Organic matter, i, 114

Organicists, i, 7

Organ-independence, i, 40

Organism, as a whole, familiarity
of expression, i, 1; definition of,
18; distrust of by biologists, 25;
and its chemistry, 75; and its
protoplasm, 120; and its cells,
150; in interpreting the cell, 156;
substitution of, for cell, 194;
consisting of one cell, 227; fic-
tion of structureless, 256; as
cause, 276; "organless," 232,
320; hypothetical primitive, 319;
fundamentally dynamic, ii, 134;
"body" and "soul" combined,
151; as causal explanation, 199;
as cause of chemical transfor-
mation, 205; a natural object,
207; attributes and acts of, 215;
living, 275 ; physico-chemical
conception of, and conscious-
ness, 324

Organism-transforming action of
thyroid substance, i, 145

Organismal theory, i, 2, 280; con-
structive side of discussion of,
ii, 91; and C. M. Child's results,
111; and elementalist stand-
point, 148; in relation to trop-

isms, 188; of consciousness, 282

Organismalism, i, 2; and correla-
tion, 17; remarks on the term,
28

Organismalist, ii, 149

Organization, law of embryonic
development, i, 14; of infusoria,
282; of chromosomes, ii, 28;
physical, and instinct, 310; and
emotion, 316

Organizing power of living beings,
i, 211

"Organless organisms," i, 232, 320

Organogenesis, i, 325

Organoids, i, 248

Organs, fallacious teachings about,
i, 242; rudimentary, 277; "can
belong only to multicellular or-
ganisms," 281

Origin of species, i, 4

Osborn, H. F., i, 320

Outer layer, universality in or-
ganisms, i, 301

Ovum, i, 210; as entity and as
germinal entity, ii, 15

Oxidation, in animal body, i, 340;
ii, 346

Oxygen, "doscarecious" powers of,
ii, 204; in relation to conscious-
ness, 290 et seq.; as respiratory
substance, 301 ; latent attribute
of, 341 ; hereditary, ontogenic
and individual, 347; "activation"
of, 348

Pacific Ocean, journey of fur
seals in, ii, 211

Pain, ii, 183

Pairing, promiscuity in, ii, 267

Paleobotanists, ii, 58

Paleontologists, i, 322

Palms, ii, 299

Pancreas and pancreatic juice, ii,
120

Pangens of Darwin, i, 19

Parallelism, psycho-physical, irre-
solvable inconsistency of, with
organismal standpoint, ii, 150,
220; historical basis of, 297

Paramecium, i, 326; ii, 253

Paratoid gland, of toad, i, 111

Parthenogenetic eggs, i, 351

Index

403

Particles, ultimate, ii, 151

Partisanship in science, i, 338

Payne, F., ii, 34

Pearl, R., i, 311

Peckhams, G. W. and E. S., ii,
251

Percepts, ii, 294

Periodic law, in chemistry, ii, 329

Personal conscience, ii, 292

Personality, ii, 335; and person,
295; and elementary substances,
327

Petrels, i, 85

Phases, fluid, of cell system, i,
216; of the cell, ii, 326

Philosopher's stone, ii, 288

Philosophy, cartesian, ii, 297

Phleps, Ed., ii, 135

Phlogiston, i, 225

Phosphorus, ii, 290; a simple, 287;
glow of, 343

Phylogeny, of biochemical sub-
stances, i, 110

Phylloxerans, eggs of, i, 352

Physical basis of life (Huxley), i,
121; of heredity, ii, 64

Physical chemistry, in biology, i,
114; and protoplasm doctrine,
140; and the organismal stand-
point, 191; conception ' of cell,
215, 333; conception of organ-
ism, ii, 71; limitations of, in
biology, 208; absence of in
earlier biology, 336; anti-ele-
mentalistic tendency of, 341

Physical science, ii, 152

Physio-chemical substances and
forces, in behavior, ii, 312

Physics, and chemistry in hered-
ity, i, 115; province of, 141

Physiologist, ii, 285

Physiology, and heredity of mus-
cle fibers, ii, 61; distinctive task
of, 274

Phytin, ii, 123

Pigeons, control of sex in, ii, 78;
Whitman's study of habits of,
314

Pigment, from chromatin, i, 338;
bearers, 340

Pigmentation, i, 213

Pill bugs, i, 219

Pillsbury, W. B., ii, 231

Pineal body, ii, 114

Pituitary Body, ii, 114, 124

Plankton, ii, 280

Plantagens, i, 36

Planaria, ii, 109

Plasma, i, 134

Plasmic elementalism, i, 311

Play of animals, ii, 273

Pleasure and Pain, criterion of, i,
289

Pleomorphism, i, 266

Plover, Golden, ii, 259

Pluralism, philosophical, ii, 294

Pluteus larva, i, 203

Poisoning, strychnine, ii, 173

Polarity, in plants and animals, i,
181

Pollen grains, in fertilization, i,
343; structure of membrane of,
ii, 57

Porospora gigantea, i, 270

Post-generation, i, 207

Postulate, ii, 297

Powers, secret, of substances, ii,
300

Precipitin reaction, i, 100

Predisposition, i, 206

Primitive, i, 290

Primrose, evening, i, 353

Primun movens, i, 276

Principle, of unity of organiza-
tion, i, 166; of aggregation, 182

Principles, Alchemists', ii, 288

Private opinion, ii, 292

Promiscuity in mating, ii, 267

Promorphology, of germ cells, i,
211; metaphysical, 225

Properties of the whole, i, 13, 148

Prophysiology, i, 212

Protista, i, 230, 280

Protophyta, i, 230

Protoplasm, as goal of biology, i,
5; mystification of, 120; and the
organism, 120; Max Schultze on,
125; latest views as to morphol-
ogy of, 133; specificity of, 143;
as general term must be used in
plural number, 148

Protozoa, theoretical simplicity of,
i, 230, 286; development of, 267;
theoretical aggregation of to

404

Index

produce metazoa, 268

Protozoan colony, theory of, as
nature of metazoan, i, 222

Protozoology, i, 280

Pseudopodia, i, 240; pseudopodial
system, 239

Psychic activities, subrational,
four certainties about, ii, 250

Psychic integration, ii, 94; discus-
sion of, 214 et seq.

Psychic life, subrational moiety
of, ii, 246; phases of, 274; spec-
ificity of, 276; man's higher,
283; catholicity of attitude
toward, 284

Psychical, organic connection be-
tween physical and, ii, 239

Psychical element, an abstraction,
ii, 235

Psycho-analysis, ii, 350

Psycho-physical parallelism (see
parallelism)

Psychoids, ii, 149

Psychologist, ii, 285

Psychology, associationist, ii, 228,
and Wm. James, 229; without a
soul and without a body, 322;
social and domestic, 332

Purpose, of reflex, ii, 184

Python, proportionality of parts
in skeleton of, ii, 96

Radicals, compound, in chemistry,

ii, 343
Radiolarian, compared with jelly

fish, i, 235; swarm spores of, 278
Rage, ii, 183

Random movements, ii, 252
Reaction, chemical and neural, ii,

286
Reason, ii, 216; "forms the

world," 243
Receptors, superficial and deep,

ii, 173
Redwood, adventitious buds in, i,

38; meristic growth series in, ii,

101
Reed, H. S., on adventitious buds

in lemon trees, i, 38
Reflex arc, cellular integration in,

ii, 163
Reflexes, simple, an abstract con-

ception, ii, 168; compounding
and spreading of, 171 ; scratch,
allied, proprio-ceptive, 172; an-
tagonistic, 174, 176; inhibition
of, and compensatory, 177; pur-
pose of, 184; relation to in-
stinct, 188, 246

Regulation, formative, i, 199

Reichert, E. T., i, 95

Relationship, the problem of
causal, ii, 224

Rennet, i, 104

Repetition, in organic growth, ii,
95 et seq.; in instinctive activ-
ity, 257 et seq.

Repetitive parts, ii, 95

Reproduction, asexual, and hered-
ity, i, 309 et seq.

Researches in biology, field, labor-
atory, quantitative, ii, 278

Resemblance, importance in doc-
trines of heredity, i, 312, 315;
and difference, 317

Respiration, life, and conscious-
ness, ii, 286 et seq.; of muscle,
346

Responses, ii, 216

"Restlessness," mental, ii, 225, 243,
307 .

Retzius, G., comparative re-
searches on spermatozoa, i, 216,
ii, 2

Reversibility, of direction in nerve
conduction, ii, 166

Rhizopod, i, 123

Rhus, chemistry of, i, 87

Riddle, O., ii, 76

Ritter and Forsyth, ii, 45

Robertson, T. B., on chemical ac-
tion in growth, ii, 105; on
tethelin, 123

Rousseau, J. J., ii, 224

Roux, W., terminology of "De-
velopmental mechanics," i, 18;
Mosaic theory of, 198

Royce, J., relation between trop-
isms and apperception, i, 23, ii,
220; on mental initiative and
restlessness, 243

Ruzicka, V., i, 262

Sachs, J., i, 16; law of, 219

Index

405

"San Diego region" (oceanic), ii,
281

Saint-Hilaire, J., i, 7

Sajous, E. de M., ii, 124

Salamander, ii, 311

Salmon, ii, 250

Salpa, i, 316

Sarcode, identification with plant
protoplasm, i, 123

Schafer, E., on endocrine organs,
ii, 14; drug-like action of inter-
nal secretion, 122

Schaudinn, F., i, 261

Schleiden, M. J., conception of
plant, i, 34

Schultze, Max, on protoplasm and
cell, i, 125 et seq.

Schwann, Th., theory of cells, i,
150

Science, positive, i, 298; creative
impulse in, ii, 227

Scientific, spirit, i, 338; attitude,
difference between, and philo-
sophic, ii, 307

Scratch-reflex, workings of, ii,
172; possible "ultimate explan-
ation" of, 202

Scripps Institution, statistical
methods in, ii, 280; for Biolog-
ical Research, 332

Sea Urchin, properties of eggs of,
i, 107; experiments on develop-
ing eggs of, 202

"Seat" of heredity, i, 321; of in-
heritance material, ii, 23; brain
supposed seat of coordination,
ii, 191

"Secret powers" of substances, ii,
341

Secretin, ii, 120

Secretions, internal, integrative
office of, ii, 113 et seq.

Secretory systems, internal, ii, 128

Sedgwick, A., ii, 50

Seeds, gradations in, i, 102

Segmental theory of nerve action,
ii, 185; of nervous system, 190

Segregation, in heredity, i, 355

Selection, sexual, ii, 262

Self-preservation and regulation,
i, 18; -differentiation, 199, 208;
ii, 245; -activity, 226; -exhaust-

ing ceremonies, 264; -injury
through sex impulse, 267; nat-
ural history method in study of,
282; -defence and -preservation,
292; or person, 301; -control,
305; -development, 305

Sellars, R. W., individuality in
percepts, ii, 294; the individual,
and social psychology, 332

Sensationalism, ii, 218

Sense organ, ii, 165; senses, 216

Sequoia sempervirens, adventi-
tious buds of, i, 38; growth
series in, ii, 101

Series, graded repetitive, ii, 95

Serum, rabbit, i, 101

Sex, cell production, theory of, i,
61; as unit-character, 348; de-
pendence upon chromosomes,
346, 350; impulse, excessiveness
of, ii, 265; power of, 267

Sexes, numerical proportion of, ii,
267

Shakespeare, Wm., "reckless volu-
bility" of, ii, 223

Sherrington, S. C., fundamental
work on nervous system, i, 22,
ii, 162 et seq.; neglect of, by
J. Loeb, 185

Shrikes, ii, 270

Simple reflex, an abstract concep-
tion, ii, 168

Simplicity, "ultimate," i, 320

Skin, universal presence of, in or-
ganisms, i, 300

Smallpox, i, 264

Smell, i, 84

Smith, J. B., i, 214

Society, and individual, ii, 332

Soma, i, 319

Song habits of birds, ii, 260

Soul, composed of "seminal
atoms," i, 3; as aspe'ct of organ-
ism, ii, 215; and body, interac-
tion between, 323

Specialist, the unpoised, ii, 247

Species, difference in egg of, ii, 20

Specific differences between germ
cells, importance of, i, 214

Specification of organic matter, i,
111

Specificity, chemical, of organ-

406

Index

isms, i, 83; of corresponding
proteins, 95; of protoplasms,
143; of sperm of anthropoids
and man, ii, 2; of animal be-
havior, 276; of psychical and
reactive life, 281

Speculations, i, 293

Spermatozoa, species differences
in, i, 216; structure of tail of,
333; structure of head of, 342,
and ii, 87; two kinds of, i, 347;
subject to heredity, 399; resem-
blance to tadpole, ii, 3; ontog-
eny of mammalian, 4

Spicules of sponges, ii, 50

Spinules of Ascidian, ii, 44

Spirits, historical relation to es-
sences, ii, 288; historical relation
to breath and air, 338

Sponge tissues, i, 144

Spontaneous, generation, ii, 316;
in mental activity, 226; on
meaning as applied to "origin
of life" (see glossary)

Spores, i, 269; of mosses and
ferns, ii, 258

Squid, i, 221

Squirrel, storing habits of, ii, 271

Stage, one celled, of animal life,
i, 214

Star fishes, ii, 96

Starch grains, i, 214

Stein, Fr., i, 286

Stentor, supposed nerves in, i, 245;
ontogeny of, 272

Stephens, Frank, on storing habits
of bees, ii, 268; of antelope
ground squirrel, 271

Stevens, Miss M. M., i, 350

Stimuli, summation of, ii, 167

Storing habits, of honey bee, ii,
268; of woodpecker, 269; of
mammals, 270

Stout, G. F., ii, 298; unanalyzed
cognition in consciousness, 308;
on personality in animals, 328

Strassburger, Ed., i, 342; and
chromosome dogma, ii, 259

Striated muscle tissue and hered-
ity, ii, 59

Structurelessness, i, 285

Structures, ii, 199; constitutively

antagonistic, 134 ; organ-form-
ing, 141

Struggle, of the parts, ii, 175; for
existence, 225

Studies, practical, i, 214

Sub-conscious, ii, 350

Subjective, ii, 231; side of asso-
ciation, 230; and "inner," 292

Substance, of soul, i, 4; organic
production of, 81; simple homo-
geneous, 281; imaginary, 240;
nutritive, 244; criterion of ele-
mentary, 287; abuse of the
term, 2*98; respiratory, 301; and
energies, 337; free and vague
appeals to, ii, 239

Sugar, and enzyme action, ii, 81 ;
from liver into blood, 132

Summation, of stimuli, ii, 167

Sumner, F. B., ii, 258

Supernaturalism, and finalism, ii,
142; and materialism, 148

Suprarenal body, ii, 114, 135; in
alliance with thyroid and pitui-
tary, 127

Surface energy, of muscle fibers,
ii, 63

Surfaces of separation, ii, 170

Surf perches, matings, habits of,
ii, 265

Surgeons, Swiss, ii, 115

Surplus energy, ii, 273

Sutton, W. S., i, 356

Swarm spores, of radiolaria, i, 278

Swezy, olive, i, 291

Symbiotic theory of organism, i,
35, 183

Symmetry of animals, ii, 189

Sympathetic nervous system, ii,
128

Synthesis, organizing, ii, 149; as-
similative, 205; and analysis,
235; apperceptive, 237

System of nature, i, 4

Systematic biochemistry, i, 95

Tadpoles, of frogs and toads, ii,

143

Takamine, J., ii, 123
Taste, i, 84
Taxonomic, in organic grades, ii,

42; discrimination, 62

Index

407

Taxonomist, i, 213

Terms, of oxygen, ii, 202; of life,
207

Tetany, i, 117, 135

Tethelin, ii, 123

Theory, of structive of proto-
plasm, i, 138; wrong better than
none, 292; Leibnitzian, ii, 150;
of organisms, and of knowledge,
152; elementalist, 157; tropistic
and segmental, of nerve action,
185; of tropisms, 232; of apper-
ception, 232; of knowledge, 296;
of consciousness, 296

Thomson, J. A., i, 309

Threshold of excitability, ii, 165

Thymus, ii, 114

Thyroid, ii, 124; apparatus, 114;
effect of removing, 115, 117;
triple alliance with pituitary
and adrenal, 127; substance,
143; organ-transforming sub-
stance from, 145

Tissue, laboratories, i, 83; mix-
tures, 143; cultures, 168; falla-
cious teaching about, 240; of
multicellular organisms, 281 ;
isolated, 294; of trees, specific-
ity of, ii, 58

Toads, ii, 76

Tonniges, C., i, 326

Torrey, H. B., ii, 97

Totipotence, theory of, i, 202

Transformation, essence of evolu-
tion, i, 41; heredity works by, i,
312, 322; of substances, ii, 72,
287

Transmission, in connection with
heredity, i, 312

Transmutation of metals, ii, 288

Transcendentalism, ii, 233

Treviranus, i, 5

Trial and error, ii, 252, 256

Trichocysts, i, 326

Tripylea, i, 278

Tropism theory, essentially an or-
ganismal theory, ii, 188; relation
to apperception, 232

Tropisms, explained by organism,
ii, 190; organismal nature of,
239; automatic and anticipatory
character of, 241; higher ra-

tional life and, 242

Tropistic and higher psychic ac-
tivity, ii, 220

Tropistic mechanism, ii, 189

Tropistic theories of nerve action,
ii, 185

Truth, ultimate, ii, 152, 289

Trypsin, i, 106

Type, i, 7

Tyranny, of the whole, ii, 158

Ultimate, problem, i, 35; criterior
of, ii, 149, 201; particles, ii, 151;
truth, 152, 175, 289; elements.
228

Ultramicroscopic organisms, i, 265

Unicellular, i, 290

Units, representative, i, 306

Unity, the organism the "only
real," i, 12, 26, 205; physiolog-
ical, 14; of the individual, 33;
thought of, ii, 150

Uranium H., ii, 341

Utility, racial, ii, 251; natural se-
lectionist meaning of, 261

Variation, ii, 245

Vertebrates, man a, because moth-
ers were, ii, 43

Verworn, M., ii, 279

Vetch, ii, 106

Virgin propagation, i, 352

Vital force, ii, 149

Vitalism, i, 113; author's attitude
toward, ii, 207

Viviparous bony fishes, mating
habits of, ii, 265

Volition, ii, 161

Wallace, A. R., ii, 278

Waller, H. E., ii, 126

Warblers, ii, 264

Wasps, instincts, variability of, ii,
251; excessive action of, 268

Wassermann, i, 258

Watase, S., i, 221

Webber, H. J., i, 330

Weismann, A., studies of sex-cells
in hydroids, i, 60; Goette's re-
sults contrary to, 68; metaphys-
ics of, 225, 348

Whale, breaching of, ii, 257

Wheeler, W. M., sense of smell,

408

Index

and odors, of ants, i, 87, 89;
early embryology of insects,
218; as field naturalist of mod-
ern type, ii, 278; on problem of
instinct, 284; on instinct and
bodily organization, 311

Whitman, C. O., as pre-organis-
malist, i, 11; on cell-theory, 220;
relation between instinct and
structure, 314

Whole, embryo, i, 204; "tyrannizes
over parts," ii, 159

Will, feeling, and intellect, ii, 216,
217

Wilson, E. B., as pre-organismal-
ist, i, 11; on cell structure, 135;
statement of cell-theory, 151 ; on
"real unity," 192; on early em-
bryology of amphioxus, 204; on
promorphology, 217; on x and y
chromosomes, 350; on connec-
tion between chromosomes and
Mendelian inheritance, 356; pro-
posal to drop "determiner" as
genetic term, ii, 82; on germ as
detached portion of parent, 88

Wilson, H. V., i, 144

Winterstein, H., ii, 62

Woodpecker, California, storing
habits of, ii, 269

Wood-tissues, ii, 58

Work, energy, power, force, ii, 342

World, external, ii, 303

Wundt, W., and apperception, i,
23; definition of apperception,
ii, 233; seeming transcendental
element in the apperception of,
234

Yerkes, R. M., on combined ex-
perimental and field research in
behavior, ii, 279

Ziegler, C., i, 201
Zoja, R., i, 204
Zoological diagnosis, i, 265
Zoologist, anthropological, ii, 285
Zoology, instruction in elementary,
i, 236; and the science of be-
havior, ii, 208; fundamental
terms of, 247; taxonomic, 276
Zygote, i, 269

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