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Reprinted from THE JOURNAL OP EXPEIBMBNTAL ZOOLOGY VOL. 9, No. 1.

WILLIAM KEITH BROOKS

A SKETCH OF HIS LIFE BY SOME OF HIS FORMER PUPILS

AND ASSOCIATES'

•

William Keith Brooks, second of the four sons of Oliver Allen
Brooks and Ellenora Bradbury Kingsley, was born at Cleveland,
Ohio, March 25, 1848. His parents were both descended from
the early settlers of Massachusetts, the first of the name having
come to America from England, before the year 1634. His
father, who was born in Middlebury, Vermont, had removed to
Cleveland in 1835, where he was engaged in business.

As a boy, Brooks was studious and thoughtful. He obtained
his early education in the public schools of Cleveland, and entered
Hobart College at the age of eighteen. Two years later he entered
the junior class at Williams College, from which he received the
degree of Bachelor of Arts in 1870. Although in college he had

1 Biographical sketches of Professor W. K. Brooks have been published as fol-
lows:

1. E. A. Andrews; Sketch of William Keith Brooks. Pop. Sci. Monthly, vol.
55 no. 3, July 1899, pp. 400-409, with portrait.

2. E. A. Andrews; William Keith Brooks. Science, N. S. vol. 28 Dec. 4, 1908,
pp. 777-786, and Jan. 1, 1909, p. 31.

3. Edwin G. Conklin; The Life and Work of Professor Brooks. Anatomical
Record, vol. 3, no. 1, January 1909, pp. 1-13, with portrait.

4. Edwin G. Conklin; William Keith Brooks. Proc. Am. Phil. Soc., no. 190,
1909, pp. 3-10.

5. Edwin G. Conklin; Biographical Memoir of William Keith Brooks, 1848-
1908. National Academy of Sciences, Biographical Memoirs, vol. 7, February
1910.

6. E. A. Andrews; Biography of William Keith Brooks, in "Leading Americans,"
Holt & Co., New York. 1910. "

A meeting commemorative of Prof. Brooks was held in McCoy Hall, Johns
Hopkins University, Nov. 12, 1908. Addresses were made by Professor B. L.
Gildersleeve, Dr. H. M. Kurd, Professor W. H. Howell, Professor E. A. Andrews,
Dr. Caswell Grave and Professor W. H. Browne. See Johns Hopkins Univer-
sity Circular, January 1909.

THE JOURNAL OF EXPERIMENTAL ZOOLOGY, VOL. 9, NO. 1.

2 WILLIAM KEITH BROOKS

shown special interest in philosophy and in studies with the micro-
scope, he was uncertain on graduation whether to devote himself
to natural history, to mathematics or to Greek, in all of which
subjects he excelled. After leaving Williams College he spent
a short time with his father in business, but this occupation was
not to his liking and he gave it up to become a teacher in a school
for boys at Niagara Falls. After holding that position for two
years he became a graduate student at Harvard College under
Louis Agassiz, who was then at the zenith of his career, and at
the seaside laboratory established by this great master in 1873 on
the Island of Penikese, Brooks began a life-long devotion to the
study of marine zoology. In 1875 he was appointed assistant
in the museum of the Boston Society of Natural History and in
the same year received the degree of Doctor of Philosophy from
Harvard. It was during the summer of this year, while at home
on his vacation, that he organized, together with Theodore B.
Comstock and Albert H. Tut tie, a class for laboratory instruction
in zoology and botany for teachers.

With the opening of the Johns Hopkins University in 1876, one
of the twenty fellowships was awarded Brooks, who thus at its
very foundation entered the service of the institution with which
he was to remain connected until his death. He was immediately
advanced to the position of Associate and later was successively
appointed Associate Professor of Comparative Anatomy, Asso-
ciate Professor of Morphology, Professor of Animal Morphology,
Professor of Zoology and Head of the Biological Department.
In 1878 he was made Director of the Chesapeake Zoological
Laboratory of the University, an institution which he organized
and which became a potent adjunct to the Baltimore laboratory
in the training of biologists.

Professor Brooks was the recipient of numerous public honors.
When but thirty-six years of age he was elected a member of
the National Academy. He was chosen a member of the Ameri-
can Philosophical Society in 1886, and of the Academy of Natural
Sciences in 1887. He was Lowell lecturer in 1901 and gave one
of the three general addresses before the International Zoolog-
ical Congress at Boston, in 1907. He received the honorary de-

A SKETCH OF HIS LIFE 6

o-ree of LL.D. from Williams College in 1893, from Hobart College
in 1899, and from the University of Pennsylvania in 1906. For
his discoveries on the life history of the American oyster he was
awarded the medal of the Societe d'Acclimatation of Paris, and
for his work on the Stomatopoda, a Challenger medal. He was
editor of the "Memoirs from the Biological Laboratory" of the
Johns Hopkins University, joint editor of the " Studies from the
Biological Laboratory" of the Johns Hopkins University, and
one of the editors of the Journal of Experimental Zoology. He
was a member of the Boston Society of Natural History, the
American Academy of Arts and Sciences, the American Society
of Zoologists, and of the Maryland Academy of Sciences, and was
a Fellow of the American Association for the Advancement of
Science and of the Royal Microscopical Society.

On June 13, 1878, Professor Brooks married Amelia Katherine
Schultz (deceased 1901), daughter of Edward Thomas Schultz,
and Susan Rebecca (Martin) Schultz of Baltimore. Two chil-
dren were born, Charles Edward Brooks and Mrs. Menetta
White (Brooks) Daniel, both of whom survive him.

A congenital defect of the heart had always caused Professor
Brooks to lead a less active life physically than do most men,
and to this trouble other bodily ills were added as life advanced.
After a continuous prostrating illness of nine months he died
at his home "Brightside," near Baltimore, November 12, 1908.

As a stimulating teacher, an ardent and successful investigator,
and a philosophic naturalist, the influence of Brooks on the de-
velopment of zoology in this country has been very great. His
students are scattered widely in college and university, in muse-
ums, and scientific stations in this country and abroad, and many
have become eminent in their own fields of work. His discover-
ies, numerous and important, have enriched zoology and have
been incorporated into the permanent literature of that science.
Certain of his memoirs are models of completeness and beauty.
His brilliant and thorough work on the oyster fisheries of Mary-
land has made his name familiar to economists and to intelligent
legislators. In an age perhaps over-eager in the pursuit of new
knowledge Brooks has called attention back to the fundamental

4 WILLIAM KEITH BROOKS

nature of knowledge itself in such a way that his helpfulness
has been gladly and gratefully recognized in many circles of
science.

In his personal character Brooks combined gentleness and
strength and a rare wisdom. In university matters and in all the
affairs of life he was a lover of freedom and of justice tempered
with kindliness. Although looked upon from the beginning as a
master mind, he was totally free from formality and never ass amed
the authoritative air of the traditional professor, but met his
students and associates in all simplicity and frankness as fellow
student and inquirer. What he was as a man and a student was
fully revealed, and the singularly deep influence which he exerted
upon those who worked with him constitutes a remarkable
tribute to his genuine ability and worth. The close friend-
ship between him and his students was evidenced in many ways
in the daily life of the laboratory, and at the evening gather-
ings at his home. It was given more definite expression on the
occasion of his promotion to a full professorship^ and again on
his fiftieth birthday, when his pupils came together at Bright-
side to present to him formally the portrait for which he had
sat at their request.

The appreciations, reviews and recollections embodied in the
following pages and coming from former students and associates
record some of the labors and some of the traits, human and pro-
fessional, of a profound thinker and tireless worker.

SOME RECOLLECTIONS OF PROFESSOR BROOKS CHRONOLOG-
ICALLY ARRANGED

1876-79.- Among the company of twenty young men who
came together in Baltimore in 1876 as the first group of "Fellows"
of Johns Hopkins University, were three biologists. One of these
was of less than average stature, wearing a serious face, with
close-set eyes, quiet and unhurried in his movements, speaking
not frequently, and never with haste. This was W. K. Brooks
of Williams and Harvard. The biological department was at
once organized by Professor Martin with Dr. Brooks as an Asso-

3 ProfessorS. F. Clarke, Williams College.

A SKETCH OF HIS LIFE 5

ciate, and it is an illustration of that quiet impressiveness of
Brooks' simple manner that his appointment was immediately
recognized by every one to be most eminently fitting.

He quickly gained our respect and admiration by the con-
stant seriousness of his thoughts, and the simplicity and gen-
uineness of his statements : simple in expression but showing care-
ful and deep reflection. Our affection was won and held by his
genuine, never-failing interest in, and friendship for us.

As I recall his reading to me of the then unfinished manuscript
of his book on Heredity in my room on Centre street in 1876, of
the many long talks on biological subjects, in either his room or
mine at the University, at Brightside, or at Crisfield, Fort Wool
or Beaufort, I become aware again of the constant seriousness,
and power of his thought, which awoke and continually increased
an admiration for his intellectual ability.

Brooks' friendship was even and steadfast. It never found
great expression in words, but it never wavered. I felt this dur-
ing my early years of association with him, and the conviction
was but strengthened with the growing years. This steadfast-
ness of affection and confidence in his friends, his perfect simpli-
city and genuineness, and his serious and profound mind are to
me the sources of Brooks ' great and lasting influence on men.

1883-84.* The first time I saw Professor Brooks was in 1883.
The year before, while I was endeavoring to make out some of the
points in the structure of Balanoglossus, then imperfectly known,
it was announced in the Johns Hopkins Circular that a littoral
species of that animal had been found at Hampton, Va. At
Mr. Adam Sedgwick's suggestion I wrote to Brooks asking if
I might come over to investigate it. Brooks, as his friends will
remember, did not habitually answer letters, but as it happened
he did answer that one and sent me a cordial invitation to come
and try. Such leave was no little thing to give, for Balanoglos-
sus must have been known to be one of the prizes of the station,
but in professional generosity Brooks was royal and lavish.

* Professor William Bateson, Cambridge, England.

6 WILLIAM KEITH BROOKS

From the first moment of meeting in the empty warehouse of
the Normal College, which then served for a laboratory, we be-
came friends. He was of course much my senior, but there is no
other word which so well expresses the happy unconstrained feel-
ing that I felt towards him and that he showed towards me. It
had been settled that I was to live at Mr. Cock's boarding house,
across the creek, where the Brooks family had their quarters, and
we thus spent several weeks in constant intimacy.

He was not the least like any one else I had ever known, and I
find it difficult to express the charm which his personality had
for me then, and has had increasingly since. He was, as I soon
found, on account of superficial eccentricities reputed a reserved
and rather inaccessible man. In general company he would
indeed often remain silent and I think he had moods in which
a morbid shyness would take complete possession of him, but once
at his ease he was another man. At such times he would talk
abundantly, but his speech was always thatof the taciturn observer,
with the special, holding quality that the speech of such men has.
He spoke in short incisive phrases, full of novelty, suggestion,
and humorously inventive thought, sometimes, but not often,
rising to enthusiasm. I see him now, with his short, round fig-
ure, sitting on the piazza at Mr. Cock's or lying flat on his bed—
a posture he often took when in a talking mood — ruminating his
thoughts, which, if the truth must be told, were periodically in-
terrupted by his devotion to tobacco. What a strange combi-
nation it was! The grave, kindly face, the earnest solemnity of
philosophical speculation and the homely quid. Now, I suppose,
no university professor, howrever contemplative, dare use tobacco
in this particular way ; but I wonder if any university professor-
ruminates spacious ideas as Brooks used to do, daily through
long vacant hours of leisure, to the delight and elevation of a
youthful listener. Those are the times of true education

"when lofty thought
Lifts the young heart above its mortal lair/'

Many of Brooks' pupils must look back on similar pleasant
hours of intimate, informal summer laboratory life as critical
moments in their development. For myself I know that it was

A SKETCH OF HIS LIFE 7

through Brooks that I first came to realize the problem which for
years has been my chief interest and concern. At Cambridge
in the eighties morphology held us like a spell. That part of
biology was concrete. The discovery of definite, incontrovert-
ible fact is the best kind of scientific work, and morphological
research was still bringing up new facts in quantity. It scarcely
occurred to us that the supply of that particular class of fact was
exhaustible, still less that facts of other classes might have a wider
significance. In 1883 Brooks was just finishing his book "Her-
edity", and naturally his talk used to turn largely on this subject.
He used especially to recur to his ideas on the nature and causes
of variation, and to the conception which he developed in "Her-
edity," that the functions of the male and female germ cells are
distinct. The leading thought was that which he expresses in
his book (p. 312) that " the obscurity and complexity of the phen-
omena of heredity afford no ground for the belief that the subject
is outside the legitimate province of scientific enquiry." He
deplored the fact that he had no opportunity for the requisite
experiments in breeding, but he saw plainly that such experiments
were the first necessity for progress in biology.

To me the whole province was new. Variation and heredity
with us had stood as axioms. For Brooks they were problems.
As he talked of them the insistence of these problems became
imminent and oppressive. It all sounded rather inchoate and
vaporous at first, intangible as compared with the facts of develop-
ment which we knew well how to pursue, but with the lapse of
time the impression became strong that Brooks was on the right
line. That autumn I went home feeling that though in technique
we were a long way ahead of Johns Hopkins — I had the pleasure
of showing off the Jung microtome, then the latest thing in pro-
gress, to the admiring Baltimore men — yet somehow Brooks had
access to novelties of a more serious description.

In the following summer I was again with Brooks at Beaufort,
N. C., but in that year I soon fell ill and was for a long time too
weak for much talk of any kind. Indeed, but for the devoted
ministrations of Brooks and his students, who for weeks performed
for me the offices of the trained nurse, I might never have left

WILLIAM KEITH BROOKS

Beaufort alive. The "Heredity" had meanwhile appeared and I
am afraid Brooks was disappointed with the reception it met, for
it was noticed with little more than formal sympathy. Looked
at in the light of subsequent knowledge its purpose was indeed
rather, as he says, "to turn the attention of others into this
channel" than to make an independent advance. In the preface
he wrote: "I have little hope that my views will be accepted
in the form in which they are here presented, but I do hope that
they may serve to bind together and to vitalize the mass of facts
which we already possess and that they may thus incite and direct
new experiments." That function he and his book did at length
admirably perform for many, both in England and in America.

1885-89.* In going over my memories of Dr. Brooks I find tnat
rny mind does not separate him from his environment. I con-
tinually see him in the semicommunal life of the laboratory,
whether in Baltimore or Beaufort, Woods Hole or the islands of
the West Indian sea, which so stirred and charmed him. Even
his home life with its restful, satisfying beauty was but a detached
fragment of the other larger existence. I think of him as the cen-
tral figure, wise and kind, of a circle of young men coining from
many quarters, from New England, the Middle States, the West,
and the South, from Canada, England and Japan, a society from
which older members were always going out to honorable careers
and into which new were coming to learn the ways and traditions '
of the school. Very different were we, but knit together from
the start by the strong bond of a common interest, and presently
by growing appreciation of him who made the school. It took
us but a short time to learn that here was no mere work-shop, well
organized and in which we might acquire the requisite degree of
skill in a profession, but that we were in the company of a master
mind, wide ranging in the fields of knowledge and inquiry, pro-
found in contemplative thought, and with the acuteness of the
observer who discovers what has been hidden.

As I dwell on the man and try to single out mental habits and

4 Professor H. V. Wilson, University of North Carolina.

A SKETCH OF HIS LIFE 9

attributes from the whole of his personality, I come to many that
arrest and enchain my attention.

It is interesting to consider his practice and advice to beginners
in the study of Nature. It was to start out, not from a general
principle, but from some phenomenon that had caught the eye
and become a nucleus for thought. Continued, persistent obser-
vation and reflection circling round such a center would yield,
he held, solid results in the shape of new facts and would sooner
or later lead one into living contact with great questions. This
method of work was eminently characteristic of his independent,
individualistic temperament.

The serenity of Dr. Brooks impressed every one. In a mind so
strong, active, and keen, calm temperateness was doubly notice-
able. This peace of mind must have been due in part to the fact
that his critical insight was unobscured by self-seeking. A firm
gaze fixed on the distant goal held the immediately advantageous
in its proper place, and gave him a confidence, a quiet boldness
that we all recognized.

Brooks frequently said that he tried always to be a reasonable
man. And in dealing with men and their ways I am convinced
that reasoning did guide him in remarkable degree. His log-
ical habit of thought came in, however, for more congenial exer-
cise in professional work. Do we not all remember the pleasure
he had in the skillful disengagement of the idea from the mass of
details, and in its portrayal, language and drawing mutually con-
tributing to clearness?

I recall also his strong and helpful faith in the value of labor
spent in searching out the order of the universe, the way things
happen in nature. For, as he often said, such knowledge both
makes the conscious life of man fuller and nobler, and is the
basis on which rests all our control of natural phenomena.

The machinery of Professor Brooks' department, the lectures,
set tasks and routine, was simple. Experience has shown, how-
ever, that it was not inadequate, on the contrary, that it was well
adapted to the purpose in view. Brooks' underlying assumptions
were that graduate students had come to stay some time, would
work as hard as they could, and that they had enough independence

10 WILLIAM KEITH BROOKS

of mind and enough elementary training to handle books and
journals which record the actual state and progress of zoology.
Of lectures there was one now and then from Professor Brooks on
any subject. A round of lectures by older students in the de-
partment was given some years, and this was excellent practice.

The journal club was serious. It met weekly and the arrange-
ment was such that each graduate student reported a number of
times during the year. A reading club met weekly in the even-
ing at Professor Brooks' house. Some pleasant book of general
zoological interest, often one of travel, was read, after which came
tea. In the laboratory again once a week readings of a more
serious nature and with some discussion were held. The "Origin
of Species" was in this way gone through, and "Agassiz's Essay
on Classification."

Professor Brooks had compiled an elaborate list of the litera-
ture, with which it was supposed candidates for the doctor's
degree were to make themselves familiar. It included the
text-books of the period and important memoirs on the various
subdivisions of zoology. The list was long. Perhaps some stu-
dents completed it. But we all read with considerable diligence
and it was the custom to make careful abstracts. On the basis
of this common reading a good deal of informal talk and dis-
cussion was maintained among us.

We lived in the laboratory all day and the younger men learned
much from the older, especially in matters of technique. Brooks
gave excellent suggestions on drawing and would occasionally go
through the form of taking a micro-photograph. A beginner in
my time was usually given some material, referred to a paper or
two on comparative anatomy or embryology, and told to verify
the research. At intervals, frequent enough, Brooks looked at
his figures, notes, and preparations and had something to say
about the matter. Frequently before this first testing and form-
ing exercise was completed, the man would be put at another.
Two or three filled the year. Then came a long season at the
seaside laboratory, in all probability the first for the student and
teeming with experience. There was daily collecting, much study
of live animals, much rearing of embryos and larva?. The pelagic

A SKETCH OF HIS LIFE 11

%

fauna got in the tow net or at times by dipping came in for a good
deal of attention. Numerous quick dissections were made, and
quantities of notes and drawings. Brooks exercised little or no
supervision over such work, but the older men were a great help
to the younger. The larger manuals such as Balfour's Embry-
ology, and later Korschelt and Heider, were fairly thumbed.
The industry and "go" of Brooks' summer laboratories was re-
markable, the lamps lit in the evening, and someone frequently
sitting up all through the night to " follow a development."
Toward the end of the season, when a little perspective had been
acquired and mere mass and variety began to pall, a special form
or two was singled out as promising something in the way of new
results, and the path of research was thus opened up. The ma-
terial so collected was studied in detail during the folio wing winter.
More intensive reading bearing on the problems as they became
defined was undertaken. Informal, short, but helpful talks about
the work were had with Brooks from time to time. He would
examine particular preparations, quickly to be sure, or would
criticise figures. There was never any leading or " nursing"
on his part. By the end of the year, though, some grasp of the
methods of research had been acquired, and the following summer
at the seaside usually found the student able to pursue the line
of inquiry on which he had already started, or to strike off into
an associated field.

1887-96.* No account of Professor Brooks should omit men-
tion of his love for plants. It is true that this interest was appar-
ently, to a large extent, of secondary influence as far as his pub-
lished work was concerned • but it was very real and occupied a
constant place in his thoughts on the broader problems of nature.
The relations of the living organism to the environment were well
exemplified for him in the plant world. He always kept at hand
something of botanical interest, and would invite you into his little
greenhouse, or into his garden, to exhibit with pride some product
of his own skill. Nowhere, perhaps, was his innate sympathy for

'' Professor Henry McE. Knower, University of Cincinnati.

12 WILLIAM KEITH BROOKS

nature more clearly shown than here. His knowledge of botany
was also constantly drawn upon whenever he dwelt on problems
of heredity, variation, adaptation, etc.

Brooks' affection for nature was also expressed in his observ-
ations on the common domestic animals about him. The exact
nature of this interest was unique to my experience, until I heard
him read aloud from the pages of Gilbert White. Then I realized
how the students of Brooks were being kept in contact with, and
inspired by, a spirit which had survived as a legacy, transmitted
to this true naturalist from a former century.

Brooks' influence extended also to undergraduates and he im-
pressed beginners in a manner which carries a lesson to our bio-
logical teachers. The routine of facts to be perceived was left
to the books and the laboratory assistant; Brooks brought out in
graphic lectures, the larger aspects of biology. With him, details
invariably led to some interesting relation or law. He drew well
and deliberately on the blackboard, and was direct, simple, and
clear. The result was to give an impression of nature as a sys-
tem of interesting problems, glimpses of life-histories, adapta-
tions, and action. All was alive and presented by an intimate
friend. The student listened to a master.

Many who may not be recognized as students of Brooks have
thus had awakened in them an interest in nature, and an insight
into her methods, which made a lasting impression upon them.
This is affirmed by a number of those now conspicuous in the fields
of medical science and practice who heard these lectures.

More than one undergraduate was diverted into a life-long de-
votion to biological science by this man who stood so steadfastly
for the highest ideals of research in this field. His talks of
nature were most persuasive, and presented to the minds of his
listeners a vivid picture of the blue sea, the coral reefs, and the
wonderful adjustments of the life with which they teem. This
vision was so real as to supplant all ordinary motives of life, and
inspire the sympathetic listener with the desire to follow the path
of so genuine a leader.

It was not difficult to arouse his interest. Anyone who made
a sane observation of nature could kindle it as readily as an ex-

A SKETCH OF HIS LIFE 13

pert biologist; and if the observation happened to be foolish the
reaction was often equally illuminating. In this breadth of inter-
est we are reminded of Darwin.

It is strange that Brooks never published particularly on ani-
mal behavior. He was constantly alive to the interest and oppor-
tunities presented by this field, and urged several students to
follow such studies; but the trend was, then, too strongly set
toward morphological work to permit of their diversion into this
other channel. I believe that his very special interest in the trans-
formations and life-histories of echinoderms, liver-flukes, and in-
sects, which has not been generally emphasized in accounts of
his life, was greatly affected by the charm of observing the living
larvse and their reactions. His enthusiasm induced several men
to work on these groups, as can be seen in a number of papers by
students whose publications are not enumerated in these ac-
counts.

It seems to me that it was this every-day intimacy with living
things, and his insistent reflections on their adaptive responses,
that attracted and held his students: This combined with a
single-hearted devotion to a high standard of scientific work and
thought.

1 888-98* During my residence of ten years at the Johns Hop-
kins University, as undergraduate, graduate, and assistant, suc-
cessively, I was thrown by fortunate circumstances into relations
with Professor Brooks which ultimately assumed an intimate
and personal character, and it is with deep appreciation and grat-
itude that, as I now look back upon those years, I realize the
influence which he exerted upon me. He was one who set by his
example the ideals which he wished his students to follow in scien-
tific work. Many for the first time learned from him the real
meaning of the " search for truth" as day by day in simplicity
and sincerity he taught us through his own truth-loving nature
veracity of thought and action. Many, too, learned under him-
for the first time to take a philosophic outlook on zoological phe-

0 Professor George Lefevre, University of Missouri.

14 WILLIAM KEITH BROOKS

nomena. And his life was such that it made all around him feel
in some measure the charm of the naturalist's calling.

Although the great and lasting debt, which all who came under
his instruction owe to him, springs from the inspiration that un-
consciously passed from him to us, it was his personal qualities,
his gentleness, his kindliness, his thoughtfulness of others, as well
as the quaint humor that characterized so many of his acts and
sayings, that humanized and endeared him to his students. His
capacity for sympathy was never shown more strikingly than dur-
ing those dark days that followed upon the death of Professor
Humphrey and Dr. Conant from yellow fever after the disastrous
expedition to Jamaica in 1897. All who were there at the time
will recall how deeply moved he was, nor will any of us be likely
to forget the simple sincerity of the man as he stood among us and
talked of the nobility of the sacrifice of a life for the sake of others
and for the cause of science.

I think my earliest definite recollection of Professor Brooks is
of seeing him walk into the lecture room in an undergraduate
class wearing a long rubber overcoat which he proceeded at once to
use on himself for the purpose of illustrating the morphological rela-
tions of the squid's mantle, while holding out the upturned collar
to demonstrate the position of the siphon. I still have my notes
on his undergraduate lectures and in reading them over I am
struck afresh by the recollection of their clearness and beauty,
although the subjects upon which he talked before the class fol-
lowed each other without apparent order or relation. As I later
learned while acting as his assistant, he was apt to lecture upon
anything that he happened to be thinking about at the time, not
infrequently changing the subject at the very last moment, to
the dismay of the assistant who would then have to prepare has-
tily an entirely different set of charts and specimens from those
which he had been previously instructed to have ready.

When I began my graduate work in zoology, I was, like every
one else at the start, cast adrift, to sink or swim ; and for all one
knew at the time, Brooks seemed absolutely indifferent as to the
outcome. He had given me a bottle containing a few shriveled
and collapsed specimens of Doliolum, with instructions to work

A SKETCH OF HIS LIFE 15

on the proliferous stolon. I remember with mortification how I
floundered helplessly through the first few months in what ap-
peared to me a hopeless struggle to reach solid ground, until one
day I happened to find out something new about that stolon.
It was a very trivial point, but in the exuberance of my first dis-
covery I showed it to Professor Brooks, and from that moment
his attitude toward me changed as if by magic. I was forthwith
consecrated to the study of the Tunicata. Brooks had, however,
the habit of suddenly suggesting and urging upon a student a to-
tally different problem from the one upon which he was working,
and this caused the greatest consternation among us during our
earlier years, until we found by experience that he usually for-
got about the matter in a few days. In this connection I cannot
refrain from quoting from a letter which he wrote me from Bal-
timore while I was absorbed in studying the embryology of Ap-
pendicularia at the Beaufort Laboratory in the summer of 1895.
"I have just heard from Bigelow," he wrote, "that the medusa
which I have been studying (Gonionemus) is now abundant in the
Eel Pond at Woods Hole. If you could get the embryology and
metamorphosis, it would make a fine thesis, and I write in the
hope that you may be disposed to go to Woods Hole at once to
try to study it, and to get specimens of the adult for me." The
idea of dropping all of my work and setting out on a journey from
North Carolina to Massachusetts to collect jelly fishes did not
appeal very strongly to me, and I remained in Beaufort, but just
how I escaped from the situation, which was quite embarrassing
at the time, I do not now remember.

The recollections of Professor Brooks that are the most vivid
and interesting ones to me are chiefly associated with our summers
at the marine laboratories, for it was there, away from the routine
and greater restraint of the life in Baltimore, that we came to
know him most intimately and affectionately. In the daily
companionship with him, for he constantly shared with us both
the joys and hardships of the work, the lovable side of his nature
was conspicuously open to us. A thousand incidents associated
with him at Beaufort crowd my memory as I recall him there,
the center of our life, the enthusiastic naturalist, the wise coun-

16 WILLIAM KEITH BROOKS

seller and teacher, the sympathetic friend, his droll humor always
in evidence, but with never a trace of unkindness. I remember
a day when one of the men, a rather puritanical student, who had
been struggling with some refractory material, in a moment of
discouragement told Professor Brooks that he could do nothing
with it. In his characteristic way he made no reply at the time,
but some hours later returned and said quietly, "Did you ever-
try swearing? That helps sometimes."

Most delightful of all is the recollection of long evenings on the
verandah, where, after the day's work was done, we sometimes
sat listening to his talk on nature and philosophy. True it is
that we were not always able to follow him closely in his meta-
physical moods but we learned at least to feel something of the
relation that exists between the study of phenomena and the
philosophic inquiry into underlying causes.

1900-05. 7 To work on aphids, to read Witlaczil, these were my
first instructions. After that he seemed to have lost interest in me.
and he showed none in aphids. Months later he startled me by
suddenly proposing three elaborate dissections, a study of the
lamellibranch gill, and of the brooding habits of Cyclas.

To improve his pedagogy seemed an easy thing at that time;
to-day I am thankful that he left me alone, and neither pushed nor
pulled. Into the sea of work suggested I plunged, bat Brooks
furnished no life-belts. Instead he gave opportunity, and some-
thing more.

In my time the " Foundations" were being read, discussed, and
not wholly understood. The typewriter in the laboratory and
at Brightside, clicked incessantly. The Lowell Lectures were in
the making; bulky translations from Hertwig and from Heider
were completed though never published ; many essays and shorter
papers were written; Berkeley was quoted; and the pile of incom-
ing reprints remained unclassified on the floor.

He seemed to be writing much, and the larger problems,
for the time, triumphed over the microscope. The doom of his
morphological studies was practically sealed by illness that grad-

7 Professor Otto C. Glascr, University of Michigan.

William Keith Brooks

student nt William* College
1868

•
a day

•rial, i.i.

ing

•

•

•

,
To

day I

'

•

.

ore in
Heider

A SKETCH OF HIS LIFE 17

ually became worse, until his system, enfeebled by a weak heart,
scarcely resisted the other difficulties that began to burden him.
It is true that periods of improvement alternated with those of
depression, but the doubt that hovered over him cast its shadow
through the laboratory. It was at this tune that Mrs. Brooks
died.

Such health as he had known, never came back wholly, and
months passed before interest in life and work returned. With
renewed vigor he studied his hydroids, his salpas, and the oyster,
began to complete researches half-forgotten, and to start new
ones. The Sunday evenings at Brightside, too, were resumed,
and amid clouds of smoke, he read Berkeley or his own writings.
This was Indian summer.

At home, much of his most vital teaching was done. Stretched
comfortably in his steamer chair, in full view of the books and pic-
tures that he loved, and surrounded by a family, not in the narrow
sense, but one in which his students, his negro servants, his dogs,
and his flowers had each a place, he was thoroughly at ease. Often,
as he laid his hand affectionately on Jupe's great head, he spoke
with tenderness of the details of his home-life.

If one thing must be singled out to explain the affection he
inspired, it is that he himself was affectionate. The loyalty that
led him to give of his own small income in times of need and
made him speak of former students as though they had been with
him only yesterday, included other things, his science, his duties
as a teacher, and his university. In its period of hardship he
economized, and offers from other institutions did not shake him.

His interests were human, and his science a pathway along
which he walked in humility to view the world and to interpret
it. The great problems were not mere exercises for the mind, but
human difficulties. The teacher and the mnn were inseparable
and it was no less the man than the teacher who inspired others.

1905-08.* It was during Professor Brooks' declining years that
he honored me with his friendship. On these visits of his to the

1 Dr. A. G. Mayer, Carnegie Institution of Washington.

THE JOURNAL OF EXPERIMENTAL ZOOLOGY, VOL. 9, NO. 1.

18 WILLIAM KEITH BROOKS

Carnegie Laboratory at Tortugas I was much impressed with
his broad kindliness and tolerance of spirit and with his interest
in the world. The force and independence of his character also
were obvious and it was clear that he would have been a deep
student of living things under any conditions of life. He was a
thinker even more than an observer. He was the follower of no
school, and few men have been so little dominated by the thoughts
of the world around them.

Still it was not his power and originality alone that made him
great and reverenced among us. It was his spirit that led us
onward in our science. The little boy who studied dragon flies
in the pool of his father's yard had had many years pass over him,
yet in his simple wondering love of nature he remained as in his
childhood days. This deep reverence for the universe of which
he felt he formed so small a part, made him careless of many things
we deem important in our daily life, for his thoughts were not
apon things of the moment but were far beyond in the border-
land between the known and the unknown.

THE CHESAPEAKE ZOOLOGICAL LABORATORY9

Professor Brooks' early experience at Penikese under Louis
Agassiz must have had a great effect upon him. From that time
on his interest in marine zoology was one of the dominant influ-
ences in his life. One of his first important acts at the Johns
Hopkins University was to organize (in 1878) a movable seaside
station under the name of the Chesapeake Zoological Laboratory
and during the following twenty-eight years he was constantly
to be found during the warmer season at some point on the
coast or in the West Indies accompanied by a party of students,
all engaged in the study of marine life.

The importance of this Laboratory in the development of the
biological department of the Johns Hopkins University and in
the general advance of zoology in America may be estimated from
the large number of students who worked at the laboratory and

* Professor E. G. Conklin, Princeton University, in National Academy
Biographical Memoirs, vol. 7.

A SKETCH OF HIS LIFE 19

the large number of papers which they published. Doctor Brooks
expected all of his graduate students to spend a season or more
at this laboratory. He rightly estimated such work as the most
valuable experience a beginner could have, for in this way the
student became acquainted with animals under natural condi-
tions; he had the opportunity of laying a broad foundation for
his future work as a naturalist, of finding for himself some matters
to investigate, and thus early to acquire the mental habit of the
independent investigator.

The Chesapeake Laboratory, as said, was not limited to one
place. For the first few years of its existence it was located at
several different points on Chesapeake Bay; afterwards it was
located at Beaufort, North Carolina; then at different places in
the Bahama Islands, and finally in Jamaica. In the various
expeditions of Brooks and his students to these different places
they made not only a biological sarvey of each region, but they
did work of most fundamental and far-reaching importance
on the various groups of animals found. Out of these expeditions
has grown the beautiful and permanent station of the U. S.
Fisheries Bureau at Beaufort, North Carolina, in which Brooks
took great interest and pride. It was on these expeditions that
his students came to know him most intimately and affectionately.
In the memory of each of them is fixed some scene of his enthu-
siasm over the discovery of a rare form or of an unknown stage in
some life history; his long vigils full of exciting discoveries; his
quiet talks on nature and philosophy.

The Chesapeake Zoological Laboratory occupied so large a
place in the life and work of Professor Brooks that it seems desir-
able to reproduce here, in his own words, a more detailed account
of the aims and history of that laboratory during, its first nine
years. The following is taken from a report by Professor Brooks
on "The Zoological Work of the Johns Hopkins University, 1878-
86," published in the Johns Hopkins University Circulars, vol.
6, No. 54:

In natural science the policy of the University is to promote the study
of life, rather than to accumulate specimens: and since natural laws are
best studied in their simplest manifestations, much attention has been

20 WILLIAM KEITH BROOKS

given to the investigation of the simpler forms of life, with confidence
that this will ultimately contribute to a clearer insight into all vital
phenomena.

The oldest forms of life are marine: every great group of animals is
represented in the ocean, while many important and instructive groups
have no terrestrial representatives ; omitting the insects, more than four-
fifths of the known species of animals are marine, and the total amount
of animal life in the ocean is incomparably greater than upon the land.
In a word, the ocean is now, as it has been at all stages in the earth's
history, the home of life; and it is there, and there only, that we find the
living representatives of the oldest fossils, and are thus enabled to study
the continuous history of life from its simplest to its most complex
manifestations.

On the sand flats at the mouth of the Chesapeake Bay, we find, living
side by side, animals like Lingula/Amphioxus, Limulus and Balanoglos-
sus, which are the representatives of some of the oldest and most primi-
tive types of animal life ; and all attempts to trace out the natural rela-
tionships of any group of animals, lead us at once to forms which are
found only in the ocean.

The animals which have contributed most extensively to the formation
of the earth's crust, the corals and foraminifera and radiolarians, abound
in the ocean to-day, and it is only by studying their life, by observations
at the seashore, that we can understand and interpret their geological
influence.

Nearly every one of the great generalizations of morphology is based
upon the study of marine animals, and most of the problems which are
now awaiting a solution must be answered in the same way.

For these reasons our chief aim in zoology and animal morphology
has been to provide means for research upon the marine animals of the
Atlantic coast, and for nine years, successive parties, composed of instruc-
tors, fellows and students in this department, together with instructors
and advanced students from other institutions have spent at the sea-
shore all the months in which marine work is practicable. Their time and
energy have been devoted to research rather than to the preservation of
collections, and the wisdom of this course can be estimated by examina-
tion of the accompanying list of publications [here omitted] ; all of which
are based, either in part or entirely, upon researches which we have
carried on at the seashore.

The wisdom of our policy is well illustrated by the fact that the leading
naturalist of America, himself the head of one of the largest scientific

A SKETCH OF HIS LIFE 21

collections in the world, says in his annual report for 1884,10 that the
expenses of an immense natural-history collection are so great that it
would be far cheaper, with the present facilities and the cost of travel,
to supply the student with the necessary funds for valuable researches,
than to go on for years spending in salaries of curators and the care of
collections, sums of money which, if spent in a different manner, in
promoting original investigation in the field or in the laboratory and in
providing means for the publication of such original researches, would
do far more towards the promotion of natural history than our past meth-
ods of spending our resources.

This fact has become widely recognized during the last ten years,
as is shown by the establishment of marine laboratories by several of
the European institutions of learning; and in the summers of 1883 and
1884 we had with us at our laboratory a young English naturalist (Wm.
Bateson) who had been provided by the Royal Society of London with
funds for his researches, the results of which have recently been published
in England.

The Johns Hopkins University was among the first to recognize and
act upon this new departure in zoology, and our little marine station
is almost as old as the great Naples laboratory. Briefly stated its history

*

is as follows:

In 1878 a small appropriation was made to enable a party of biologists
from the University to spend a few weeks at the seashore in the study of
marine zoology. Through the influence of Maj. Gen. Q. A. Gillmore,
the Secretary of War permitted us to occupy the vacant building at
Fort Wool. Prof. Spencer F. Baird also exerted his influence with the
Secretary of War in our behalf, and aided us in many other ways; fur-
nishing us with dredging apparatus and with three small row-boats.
The scientific results of our season's work were printed in an illustrated
volume, the cost of publishing which was borne by the following citizens
of Baltimore : Samuel M. Shoemaker, John W. Garrett, John W. McCoy,
Enoch Pratt, P. R. Uhler, T. B. Ferguson, Dr. Geo. Reuling, President
Oilman, Professor Martin and others.

In 1879 the appropriation for the maintenance of the laboratory was
renewed, and in order to present an opportunity for studying the oyster
beds of Maryland, the laboratory was opened in three of the barges of
the Maryland Fish Commission at Crisfield, Maryland, a point which
proved to be very unfavorable. Maj. T. B. Ferguson, the State Fish
Commissioner, not only provided the barges for our accommodation,

10 Report of the Museum of Comparative Zoology, Cambridge, Mass.

WILLIAM KEITH BROOKS

but he also fitted the steam yacht Lookout with dredging apparatus, and
rendered us valuable help in dredging and collecting. Through his influ-
ence a small steam launch was also detailed from the U. S. Navy for
our use.

The next year the Trustees of the University voted to continue the
laboratory for three years more, 1880-1-2, and they provided a liberal
annual appropriation of $1,000 for current expenses, which was renewed
annually in 1883-4-5-6, and was expended in rent, wages, fuel, laboratory
supplies, repairs, etc. They also appropriated the sum of $4,500 for
permanent outfit, and most of this was used in the purchase of two boats;
a Herreshoff steam launch twenty-seven feet long and eight feet beam,
and a center-board sloop forty-seven feet long and fourteen feet beam.

After an examination of all the available localities the town of Beau-
fort, N. C., about four hundred miles south of Baltimore, was selected
as the site for the laboratory, and a vacant house, suitable for the accom-
modation of a small party, was found and rented as a laboratory and lodg-
ings for the party, and it has been occupied during the seasons of 1880-
1-2-4-5, and by two students in 1886. As the director was, in 1883, a
member of the Maryland Oyster Commission, the outfit of the laboratory
was that year moved from Beaufort into the Chesapeake Bay, and we
occupied a building which we rented from the Normal School at Hamp-
ton, Va. As Hampton proved to be a very unfavorable place for our
work we returned to Beaufort the next year, and we have accordingly
spent five seasons at Beaufort.

During the season of 1886 the zoological students of the University
were stationed at three widely separated points of the seacoast. A party
of seven under my direction visited the Bahama Islands, two were at
Beaufort, and one occupied the University table at the station of the
U. S. Fish Commission at Woods Hole.

The party which visited the Bahamas consisted of seven persons, and
our expedition occupied two months, about half of this being coiisumed
by the journey.

The season which is most suitable for our work ends in July, and we had
hoped to reach the Islands in time for ten or twelve weeks of work there,
but the difficulty which I experienced in my attempts to obtain a proper
vessel delayed us in Baltimore, and as we met with many delays after
we started, we were nearly three weeks in reaching our destination.

We stopped at Beaufort to ship our laboratory outfit and furniture,
but the vessel, a schooner of 49 tons, was so small that all the available
space was needed for our accommodation, and we were forced to leave
part of our outfit behind at Beaufort.

A SKETCH OF HIS LIFE 23

We reached our destination, Green Turtle Key, on June 2nd, and n •-
mained there until July 1st. The fauna proved to be so rich and varied
and so easily accessible that we were able to do good work, notwithstand-
ing the shortness of our stay and the very primitive character of our lab-
oratory. This was a small dwelling house which we rented. It was not
very well adapted for our purposes, and we occupied as lodgings the
rooms which we used as work-rooms.

Record of the various sessions

For the following brief records of the various sessions we are
indebted in large part to Prof. E. A. Andrews.

1878: 8 weeks, Ft. Wool, Virginia; 7 members. Brooks studied cm-
bryology of Lingula.

1879: June 25- August 8, Crisfield, Maryland; 11 members. Brooks
studied the oyster. Three barges served as laboratory and
quarters. Swarms of mosquitos led to the abandonment of
this locality early in August, and the removal of the labora-
tory to Ft. Wool, until September 15.

1880: April 23-September 30, Beaufort, North Carolina; 6 members.
Laboratory and quarters were in the Gibbs house. A steam
launch was bought and the laboratory equipped by means of
an appropriation from the University.

1881: May 2-end of August, Beaufort, North Carolina; 12 members.
An "Elementary Seaside School" had been announced, with
lectures by Brooks and S. F. Clarke; fee for the course, $25.

1882: May 1-end of September, Beaufort, North Carolina; 8 members.

1883 : May 1-October 1 , Hampton, Virginia. As a member of the Mary-
land Oyster Commission Brooks was obliged to spend this
summer on the Chesapeake. The new machine shop of
the Hampton Institute was rented as a laboratory, and a
fast sloop was added to the equipment. Wm. Bateson there
joined the party to study the development of Balanoglossus.

1884: June 1-September 19, Beaufort, North Carolina; 10 members.
The illness of Brooks obliged him to return after a month,
leaving the laboratory in charge of H. W. Conn. Bateson,
who was again with the party, was also seriously ill.

1885: May 23-September 15, Beaufort, North Carolina; 11 members.
Brooks became a licensed pilot to take the steam launch in
and out of Beaufort Inlet.

24 WILLIAM KEITH BROOKS

1886: June 2- July 1, Green Turtle Key, Abaco, Bahamas; 7 members.
The party left Baltimore, May 1, in a small Bay schooner,
chartered by the day, with Brooks as pilot. With head winds,
mishaps and a stop at Beaufort to take on laboratory furni-
ture they did not reach their destination until June 2.

1887: March 1-July 1, Nassau, Bahamas; 12 members. After this
session, owing to financial losses on the part of the Uni-
versity, the Chesapeake Zoological Laboratory was tem-
porarily suspended and its outfit dispersed.

1888 and 1889: Brooks, with some of his students, was at Woods Hole,
Massachusetts, as naturalist in charge of the U. S. Fish
Commission Station.

1891: May 26-September 1, Kingston, Jamaica; 15 members. The
Chesapeake Zoological Laboratory was established at Port
Henderson, on the harbor opposite Kingston.

1892: A party of three, in charge of Professor Andrews, was located
at Alice Town, North Bimini, Bahamas. Brooks did not go.

1893: April 20-July 23, Port Henderson, Jamaica; 7 members. Brooks
did not go and Dr. R. P. Bigelow was acting director.

1894: April 7-July 7, Beaufort, North Carolina; 9 members. Brooks
was present.

1895: June 6-August 13, Beaufort, North Carolina; 4 members. Doc-
tor Sigerfoos was acting director; Brooks \vas not pre-
sent.

1896: April 29-July 30, Port Henderson, Jamaica; 4 members. Dr.
F. S. Conant was acting director; Brooks was there for a
while.

1897: June-September, Port Antonio, Jamaica; 12 members. Prof.
James Ellis Humphrey was acting director. Humphrey
died there of yellow fever, August 12; Dr. Franklin Story
Conant contracted the fever there, and died on his return
to Boston in September.

1898: Beaufort, North Carolina; 6 members. Prof. H. V. Wilson was
director. In this and all subsequent years students went,
with little or no aid from the University, to the U. S. Fish
Commission Station at Beaufort.

1901-1906: Brooks was again at Beaufort in 1901 and 1903, and at
the Marine Laboratory of the Carnegie Institution at Dry
Tortugas, Florida, in 1905 and 1906.

A SKETCH OF HIS LIFE 25

PROFESSOR BROOKS AS AN INVESTIGATOR AND WRITER

Professor Brooks' investigations lay mainly in the field of
animal morphology and embryology. In this field he was an
acute observer possessed of great patience and pertinacity. His
philosophic insight and breadth of view, moreover, made him
alert to the significance of what he observed, and his memoirs
are hence notable for their suggestive and broad theoretical dis-
cussions. Fundamental resemblances in the development and
anatomy of forms were the phenomena in which he was especially
interested. Interrelationships between groups and the phyloge-
netic value of embryonic and larval characteristics were the specu-
lative problems on which he brought his discoveries to bear. In
reaching conclusions from facts he showed the caution of the
observer who had seen much, and his soundness of judgment is
widely recognized. Nevertheless he was at times not averse to
bold speculation, as may be seen in his instructive discussion of
the nature of the early pre-cambrian fauna, and the origin of the
existing great groups of animals (The Genus Salpa and The
Foundations of Zoology) . His morphological studies embraced a
number of invertebrate groups, pelagic tunicates, mollusks, mol-
luscoidea, Crustacea, and hydromedusaB.

The illustrations in Brooks' memoirs are striking. It was his
practice to make them himself, and they have the artistic excel-
lence combined with truthfulness of detail found only in the work
of the artist-naturalist. Most of his drawings were in pen and
ink, the shaded parts stippled, and made on a large scale suitable
for reduction. They represented much labor, but Brooks was a
quick worker in this style, which he preferred above all others.
The mechanical process of stippling aided him, he maintained, to
abstract his mind and to follow out lines of thought quite unrelated
to the drawing. With respect to his artistic skill Brooks was with-
out egotism, and when the drawings were once reproduced the
originals were thrown away.

Together with skill in drawing Professor Brooks was unusually
fortunate in possessing literary power in a marked degree. His
subject is presented in an order and manner that makes it easy

26 WILLIAM KEITH BROOKS

for the reader to follow, and his command of language is admir-
ably suited to the needs of the naturalist. His technical papers
always show order and proportion and a fine precision in the use
of words. These qualities appear too in his one. text-book, the
" Handbook of Invertebrate Zoology," a manual so excellent that
it has been a model for many later books in this field. His popular
articles and lectures reveal the same logical habit of mind, but
in these it is the graphic description that especially seizes the mind
of the reader. Particularly pleasing and effective are the descrip-
tions of scenes of nature in which animals dominate, or of the be-
havior of individual animals. In the argumentative portions of
his later writings dealing with heredity and the philosophical
aspects of nature, Brooks is not always easy to follow. He leaves
a good deal of responsibility on the reader. Yet these writings
contain much that is beautiful in style as well as in idea, much
that is very quotable, real " nuggets of wisdom, products of deep
thought as well as of careful observation."11

Researches on the Tunicata.1- Brooks' first contributions on
the SalpidaB appeared in 1875-76. He observed that the eggs,
which are borne by the individuals of the chain, arise really in
the solitary Salpa and are passed into the stolon early in its devel-
opment. Each individual of the chain receives usually one (in
some species more) of these eggs and serves as nurse to the embryo
which comes from it. Salpa, therefore, does not show true alter-
nation of generations, and Chamisso's discovery of an ap-
parent metagenesis in this form must be looked on as a misin-
terpretation of the phenomena. In his later study Brooks found
that the spermatozoa as well as the eggs come from the mass of
germ cells lying in the ventral part of the solitary Salpa, so that
the solitary Salpa is in reality a potential bisexual animal.

Brooks worked out in far greater detail and with greater clear-
ness than any other student the development of the buds upon the
stolon, and showed the fundamental harmony of the process of

11 President D. S. Jordan.

11 Professor M. M. Metcalf, Oberlin College.

A SKETCH OF HIS LIFE 27

budding in Salpa with that in Pyrosoma and in the Clavelinidae
among the ascidians. The stolon is bilaterally symmetrical, its
planes of symmetry coinciding with those of the solitary Salpa
which bears it, and at first the planes of symmetry of every
member of the chain coincide with those of the stolon and the
solitary Salpa. Very soon, however, a twisting of the chain occurs
which leads to the formation of a double row of Salpse, each row
with the dorsal surfaces of its members turned outward while the
ventral surfaces of the two rows are turned toward one another,
and the right sides of the members of one row and the left sides of
those of the other row are turned toward the base of the stolon.

He showed that the placenta of Salpa does not resemble the
mammalian placenta in its method of nourishing the embryo,
but that certain cells in the placenta, taking nourishment from
the blood stream of the nurse (the chain Salpa), grow to very
large size, then lose their connection with the placenta and wander
to different parts of the embryo, where they break down and nour-
ish the growing tissues of the embryo.

Salensky's generally accurate work upon the embryology of
many species of Salpa contained one fundamental error, since
he described the embryos as arising not from true blastomeres
but from follicle cells, the blastomeres degenerating early in the
developmental history. Brooks, recognizing the improbability
of any such conditions, succeeded in tracing the development of
the egg itself until from its blastomeres the organs arise. He found
that the blastomeres develop very slowly; that the follicle cells,
on the other hand, proliferate very rapidly and take on the form
of the rudiments of the several organs, the organs being thus
blocked out in these extra-embryonic cells, while as yet the blas-
tomeres are very few in number. Later the blastomeres multiply
and pass into the different parts of the mold thus formed for them
by the follicle cells, and gradually use as food the degenerating
follicle cells that surround them.

In his latest, unpublished work he traced the cleavage of the
egg; he found a clear gastrula arising by invagination from the group
of blastomeres; he observed the hollow dorsal nerve tube, finding
it at first considerably elongated; he found a postero-dorsal rod

28 WILLIAM KEITH BROOKS

of blastomeres, the notochord, which later for the most part
passed into a postero-ventral protuberance there to degenerate
and share in the formation of the eleoblast which he thus clearly
showed to be a degenerate tail ; at one time he was studying struct-
ures in some of his embryos which seemed to be a pair of true
stigmata, but his final decision in regard to them is unknown.

Brooks' embryological work convinced him that Salpa, though
now perfectly adapted for pelagic life, has not always been pelagic,
but that it is descended from sessile forms like the ascidians, and
that some of the features which so well adapt Salpa for a pelagic
existence arose during this sessile stage in its ancestry, or were
then much improved over the earlier condition illustrated in
Appendicularia. Having found this most typically pelagic of
all pelagic animals to be a migrant from the ocean bottom, he
was led to review the whole pelagic fauna, and as a result of this
review reached the conclusion that nearly all pelagic animals of
considerable size or complex structure have had a similar history
and are descended from forms that once lived on or near the ocean
bottom.

The memoirs upon the Salpidse are of such comprehensive
character and fundamental importance that they must be desig-
nated as monumental. This massive character of his work,
together with the soundness of judgment displayed, has unquest-
ionably made Brooks the foremost student of the group. It
is he, more than all others, who succeeded in showing that beneath
the perplexing maze of secondary phenomena which so obscures
the development of this group, there is a general conformity to the
development of other chordates.

Researches on the Crustacea.13 Professor Brooks' interest in
the Crustacea began early, for as a boy he had collected the fresh-
water shrimp, Palaemonetes exilipes, in the Rocky River near his
Cleveland home, and in the marine laboratory of Alexander
Agassiz he had observed with astonishment "the lively interest
in shells," displayed by the newly hatched hermit crabs. That

11 Professor F. H. Herrick, Western Reserve University.

A SKETCH OF HIS LIFE 29

the impressions thus made were strong may be gathered from the
fact that many years later he urged the writer to study the devel-
opment of this shrimp, if possible, and that after the lapse of
nearly a quarter of a century he wrote the graphic account of the
behavior of the hermit crab which appears in the introductory
lecture of his work on "The Foundations of Zoology."

Altogether there are about fifteen papers on the embryology,
metamorphosis, habits and classification of the higher Crustacea
which singly or jointly bear the name of Brooks, and all were
issued during a period of fourteen years, from 1879 to 1892. More-
over, his "Handbook of Invertebrate Zoology" contains much
original matter pertaining to this class of animals. His first con-
tribution in this field was on the larval stages of the stomatopod,
Squilla empusa, and represented the first "Scientific Results of
the Chesapeake Zoological Laboratory" for 1878, and it was
upon tne adults and the larvaB of this sub-order that some of his
most notable work was later accomplished.

At Beaufort, North Carolina, during the season of 1880, Brooks'
interest in Crustacea deepened, for he saw in their structure and
in the metamorphosis which they so beautifully displayed, a
means of attacking several larger problems, such as "the laws of
larval development," the analysis of secondary adaptations, and
the meaning of metamerism in both the lower and higher animals.
He had pondered over the works of Professor Claus on crustacean
development and morphology, and for upwards of four years, from
1880 to 1883, his own elaborate notes and pen drawings on the
Macrura had grown to such an extent that they filled a large
portfolio. From this source he drew materials from time to time
for publication, as certain subjects happened to engage his special
attention. Without doubt he had contemplated an extended
monograph, which was only partially fulfilled in the work on
'The Embryology and Metamorphosis of the Macrura," pub-
lished in 1892.

The works by which Brooks will be best known to all future
students of crustacean zoology are undoubtedly his monograph
on "Lucifer: A Study in Morphology," published in the Philoso-
phical Transactions of the Royal Society of Great Britain for

30 WILLIAM KEITH BROOKS

1882, and his " Report on the Stomatopoda," which appeared as
part of the sixteenth volume of the Scientific results of the Chal-
lenger Expedition in 1886. In the former work, we are told that
in April 1880, he found at Beaufort "a single Lucifer with two
eggs attached to one of its appendages," and that he was "led
by the great importance and interest of the subject to make every
effort to trace its life-history." Success came only after months
of repeated failure, when at last he could say with evident satis-
faction: "I have seen the eggs of Lucifer pass out of the oviduct.
I have seen the Nauplius embryo escape from the same egg which
I had seen laid, and I have traced every moult from the Nauplius
to the adult in isolated specimens. There is therefore no crusta-
cean with the metamorphosis of which we are more thoroughly
acquainted than we now are with that of this extremely interesting
genus." Not only did he discover that Lucifer emerged from the
egg as a true Nauplius, but what was even more novel, that the
egg underwent a total and regular segmentation, and gave rise to
an egg-gastrula of the invaginate type. After giving an exhaustive
analysis of the developmental stages of Lucifer, and comparing
its successive appendages with those of other representative Mala-
costraca, he concludes that the three-jointed Nauplius larva repre-
sents a true ancestor, that there is essentially but one kind of
homology presented by metameric animals, and that, therefore,
the remote ancestor of the Crustacea does not represent a com-
munity of once independent parts.

The monograph on the Stomatopoda is distinguished by the
great ingenuity shown in classifying all of the known larvae of
this sub-order, and in tracing them to their proper genera, for
he had no living material to work with, excepting the two species
from the southern coast of the United States, Squilla empusa and
Lysiosquilla excavatrix, which he had previously studied, and
which he used for exact comparisons so far as possible. He said
of the collection submitted to him, that while it contained only
fifteen species of adults, eight of which were new, it was very rich
in larvse. In speaking of the eggs, he remarked that since they
were not carried about by the female, attached to her body or
appendages, as is the rule in the higher Crustacea, they quickly

A SKETCH OF HIS LIFE 31

perished when deprived of the constant current thus supplied,
and that it was very difficult to procure them at all, adding
that he knew of no young Stomatopod which had been reared
from an egg outside the burrow1 or in an aquarium.

The sentence just quoted was written in 1885, and we can ap-
preciate the pleasure he must have experienced in being able to do
the very thing to which he alludes, two years later at Nassau, for
on the first or second day after reaching the Bahamas one of his
students, Dr. E. A. Andrews, brought him "a Gonodactylus
and a bunch of yellow eggs," which had been broken out of a
coral rock. Feeling sure that at last he was on the track of a
stomatopod's eggs, he started at once for the beach, and it was
not long, as he tells us, before "the problem was solved, and I
went home and to bed, confident that I should next day get all
the embryological material I needed." The notable paper in
which he has described how a stomatopod crustacean was for
the first time reared from an egg, and followed in all its successive
stages, alive, should not be overlooked, though appearing as a
chapter in another work (The Embryology and Metamorphosis
of the Macrura. Chapter III.).

Researches upon the Ccelenterata.^ Exclusive of preliminary
accounts afterwards published in more amplified form, and of
popular writings, Professor Brooks produced either alone, or in
cooperation with his students, ten papers upon ccelenterates.
All are the results of labors of his maturity, for he was thirty-two
years of age when the first was published. This may account in
some measure for the high standard he maintained throughout
these papers, for next to Agassiz we must rank him as the greatest
student of the ccelenterates of our country.

The excellence of his work depends not upon the number of
species he described as new to science, for of these he names but
eleven during the whole twenty-seven years covered by his writ-
ings on ccelenterates. It is in the fields of embryology and anatomy
that Brooks' work stands preeminent; and his life-histories of

14 Dr. A. G. Mayer, Carnegie Institution.

32 WILLIAM KEITH BROOKS

Liriope, Cunoctantha, Eutima, and Philalidium McCradyi are
classics of science in their thoroughness, wealth of accurate illus-
tration, and that subtle charm in description which was their
author's own. Through patient searching upon many a collecting
trip at Beaufort, he was the first to find and describe the hydroids
of Turritopsis, Nemopsis, Phortis, and Stomotoca, while his
studies along the shores of the Chesapeake led to discovery of the
ephyrse and early growth-stages of the free swimming medusa of
Dactylometra.

His summers in the Bahamas led to the discovery of the re-
markable process of the development of medusa-bearing hydroid
blastostyles upon the gonads in Epenthesis (Philalidium) Mc-
Cradyi. He also sectioned and beautifully figured, the marginal
cordyli of Laodicea, and was the first to elucidate their structure
and homologies; and from the standpoint of morphology his de-
scription of Dichotomia cannoides in the Proceedings of the Amer-
ican Philosophical Society of Philadelphia, 1903, may well serve
as a model for those who essay to describe medusa?.

It was in cooperation with Brooks that Conklin discovered that
in Physalia only male gonophores are found, while in another
siphonophore, Rodalia, only female gonophores occur, the infer-
ence being that in both forms the opposite sex is so different from
the one known that it may have been classed as a wholly different
genus. Another of his students, Rittenhouse, while working under
Brooks, gave an excellent account of the early stages of the devel-
opment of Turritopsis.

Facts interested him but little unless they led toward generali-
zations, and thus it is that he wrote but one purely systematic
paper upon ccelenterates, and that all of his other work was directed
toward the study of developments and homologies as indicating
what has been the path of evolution. Such a problem as the rela-
tionship between ccelenterates and bilateral animals was accord-
ingly very attractive to him, and he was disposed to lay stress
upon the (somewhat masked) bilateral symmetry discovered by
himself in Eutima and by Hamahn in other hydroids.

Brooks' views were not seldom in conflict with accepted theories,
as when in 1886 he came to the conclusion that the remote ances-

Reproduced from Appletons' Popular Science Monthly. Copyright, 1899, by D. Appletori and Company

WILLIAM KEITH BROOKS

l.SUS

A SKETCH OF HIS LIFE

tor of the hydromedusse was a solitary free-swimming hydra or
actinula with no medusa-stage but probably with the power to
multiply by budding. Finally, however, becoming more perfectly
adapted to a swimming life it was converted into a medusa with
pulsating bell, and with sense-organs. After this the larva derived
an advantage through attachment, and thus the hydroid stage
was secondarily produced, and then perpetuated through natural
selection . It may be said of this theory that while it has gained
no important following, yet nevertheless it has never been dis-
proven. It is logically sound, presents the direct development of
certain medusae from a new point of view, and the future may pos-
sibly show that it rests on a basis of truth.

Researches on the Mollusca and the Molluscoidea.16 Two of
Brooks' first papers deal with the lamellibranchs, one(1874)
with an " organ of special sense" in Yoldia, while in the other
(1875) the development of Anodonta implicata is described in out-
line, and the conclusion is reached that the larva, Glochidium, is
a specially modified stage and has no bearing on the question of
the origin of the group. In a paper " On the Affinities of the Mol-
lusca and Molluscoidea" (1876) he again approached phylogenetic
problems, and concluded that the Brachiopoda have been derived
from Vermes, Polyzoa from Brachiopoda, and the molluscan
veliger (prototype of the Mollusca) from Polyzoa. Later in his
paper on the development of Lingula (1879) he held that the Roti-
fera, Polyzoa, and Veliger were three branches which early diverged
from the vermian stem. The Brachiopoda he held to be the most
highly specialized members of the polyzoan branch, the Mollusca
the most highly specialized of the Veliger branch. For these three
branches he proposed the name Trochifera.

In his " Observations on the Early Stages in the Development
of Fresh- Water Pulmonates" (1879) he observed the rhythmical
nature of the process of cleavage, and devoted considerable atten-
tion to the origin of the germ layers, to the fate of the blasto-

15 Professor G. A. Drew, University of Maine.

THE JOURNAL OF EXPERIMENTAL ZOOLOGY, VOL. 9, NO. 1.

34 WILLIAM KEITH BROOKS

pore, and the origin of the digestive tract. The technique necessary
for the successful sectioning of such small bodies as snail eggs
had not been developed at this time. Brooks' observations were
therefore made exclusively on material studied in toto, and it is
interesting to find that this method of study led him into several
serious errors. In his paper on the " Acquisition and Loss of a
Food Yolk in Molluscan Eggs," Brooks devoted much attention
to what is now known as the yolk lobe, or polar lobe, which he
regarded as a food yolk which is disappearing in some forms, while
in others it is being acquired. In a brief paper on the " Develop-
ment of the Digestive Tract in Mollusks" he reiterates his mis-
taken view that in gasteropods and lamellibranchs the blastopore
is converted into the shell gland. Not until 1908 did he return to
the gasteropods, publishing in that year in association with Bartgis
McGlone, one of his students, a paper on the origin of the lung
in Ampullaria.

Brooks has two papers on the development of cephalopods pub-
lished in 1880. His important conclusions in these papers deal
with the homologies of the cephalopod yolk sac, siphon, and arms.
Numerous publications deal with the development and propa-
gation of the oyster. In 3 878, during the first session of the Chesa-
peake Zoological Laboratory, he attempted to find young oysters
in the gills of the female, as had been described for the European
oyster, but without success. In May, 1879, he went to Crisfield,
Maryland, the center of the oyster industry on the Chesapeake,
and settled down to study the problem of the development of
the oyster. He soon learned that artificial fertilization was possi-
ble, and that the American oyster normally discharges its eggs
and sperm into the open water, where the processes of fertiliza-
tion and development go on independently of the parents. The
results of his embryological studies on the oyster were published
in full in a report to the Maryland Fish Commission (1880).
This paper was very favorably received and was republished
in whole or in part in many American and European journals.
In recognition of the importance of this work he was awarded a
medal by the Socie'te' d'Acclimatation of Paris.

A SKETCH OF HIS LIFE 35

Economic Work on the Oyster Fisheries.1* Brooks' economic
work on the oyster began in 1882 when the Governor of Mary-
land appointed him Chairman of a Commission to examine the
oyster beds and to advise as to their protection and improvement.
While occupying this position, he was excused by the Johns Hop-
kins University from practically all duties as teacher and investi-
gator and for two years he devoted his talents and energy to the
work of the Commission. He organized and carried on an exten-
sive investigation of the actual condition of the natural oyster
beds of Maryland and studied carefully the results of the policy
then pursued by the State in its work of supervising and policing
its oyster resources. He also compiled statistics from the history
of the oyster industries of France and the North Atlantic States
in order to ascertain and to show the possibilities of oyster pro-
duction possessed by the tide waters of Maryland and the condi-
tions under which some of these possibilities may be realized.

A detailed account of these investigations was published in
January 1884 under the title " Report of the Oyster Commission
of the State of Maryland" (a quarto volume of 193 pages), and
carefully prepared plans for inaugurating a system of oyster cul-
ture under private ownership and for increasing the supply of
oysters from the public oyster grounds, were submitted to the
General Assembly for its consideration and approval.

The plans worked out by Brooks by which the oyster resources
of the Chesapeake Bay and its tributaries could be husbanded and
developed, were far in advance of public sentiment in Maryland
and were rejected. Not until 1906, twenty-two years later, did
the Legislature enact a general oyster culture law for the entire
State.

Professor Brooks' active interesc in the Maryland oyster
problem did not end when his connection with the State Commis-
sion expired. He realized from the character of the discussion and
opposition which brought about the rejection of his plan for oyster
culture, that the oyster problem in Maryland is in reality a
social and political one, and he therefore set about conducting a

16 Professor Caswell Grave, Johns Hopkins University.

36 WILLIAM KEITH BROOKS

long campaign of education. In this he had in mind to bring the
State to realize the value of the Chesapeake for oyster production,
and the worth of proper methods of supervision and cultivation.
He was available for semi-popular lectures, wrote magazine and
newspaper articles on the subject of oysters, and in 1891, published
a treatise entitled "The Oyster," a little book that had wide
influence and which was characterized by President D. C Gil-
man as ''a memoir in natural history and a chapter of political
economy," in which the life history of the oyster is described "in
terms scientific enough to be accurate, not so scientific as to be
hard of understanding."

Dr. Brooks' efforts during this period resulted in the creation
of an intelligent appreciation on the part of the general public
not only throughout Maryland but in all of the Atlantic States
as well, of the value and possibilities of the natural resources of
tidal waters for the production of oysters, and men of large influ-
ence were enlisted in the cause of oyster culture. This deep cumu-
lative influence of Professor Brooks on the public mind made him
one of the most valuable citizens Maryland has ever had. Others
carried to completion the task of crystallizing sentiment in favor of
oyster culture, and finally in 1906 the Maryland Legislature
passed a law for the protection and propagation of oysters along
substantially the line that had been advocated by Brooks. The
long campaign was thus happily terminated.

Contribution to Anthropology.1'' Brooks' paper "On the Luca-
yan Indians" embodies the results of an excursion into the field
of physical anthropology made during two visits to the Bahama
Islands in connection with his summer laboratory. Very charac-
teristically, he became interested in the history of the islands
and in the people who dwelt there when they were discovered
by Columbus. The skeletal fragments which there is reason to
believe represent remains of the aborigines are very few. The
material which Brooks had, and which was found in caves on
the islands, consisted of three well preserved skulls and some

17 Professor H. H. Donaldson, The Wistar Institute.

A SKETCH OF HIS LIFE 37

other bones and fragments of bones. From these he was able
to determine several of the more important physical characters
of that ill-fated people who have left but a single monument, the
word " hammock."

In the paper in question, which was read before the National
Academy in November, 1887, Brooks gives a series of admirable
plates artistically illustrating the skulls described, and reaches
several conclusions, which may be summarized as follows: The
bones are thick, massive and dense; the skulls are of good size.
They are highly brachycepha'ic but at the same time artifically de-
formed in a way which would increase their brachy cephalic shape.
There is no reason to think that the people were gigantic, though
they were probably of large size. From certain similarities to the
remains of the inhabitants of southern Florida, it is probable that
the Lucayans belonged to the same race.

Studies on Heredity.18 As early as 1876 in a paper entitled
"A Provisional Hypothesis of Pangenesis" Brooks began to deal
with questions of heredity and variation. His thinking in this
direction took shape and led in 1883 to the publication of a volume
under the title of "The Law of Heredity." The central point in
the theory here presented is the conviction that the reproductive
elements are, contrary to the usual opinion, not alike in function.
In support of this conclusion the author draws arguments from
the facts that hybrid offspring resulting from reciprocal crossings
are often very different; that the offspring of a male hybrid and
the female of a pure species is much more variable than the off-
spring of a female hybrid and the male of a pure species; that a
structure which is more developed or of more functional impor-
tance in the male parent than it is in the female parent is very
much more apt to vary in the offspring than a part which is more
developed or more important in the mother than it is in the father.
These and other facts convince Brooks that the ovum and sperm
cell are not only different morphologically, but that they differ
profoundly in function as well.

18 Professor H. V. Wilson, University of North Carolina.

38 WILLIAM KEITH BROOKS

In developing this idea into an explanatory theory of the way
in which hereditary transmission is accomplished, Brooks borrows
from Darwin's hypothesis of pangenesis, and assumes the exis-
tence of material particles, "gemmules," which are thrown off
from the body cells. Unlike Darwin, however, he assumes that
such particles are only thrown off at particular periods, when the
body cells are disturbed in function through some change in their
environment. The gemmules may penetrate an ovum or a bud,
but it is the male germ cell which has gradually acquired during
the evolution of the metazoa the peculiar power to gather and
store up gemmules. The ovum on the other hand has acquired
a very different nature. It contains material particles which
correspond to the hereditary characteristics of the species. Thus
in the case of a fertilized egg, as in that of a parthenogenetic egg,
the great bulk of the development is due to the properties of the
ovum itself. The gemmules brought in by the sperm cell unite
with homologous particles in the ovum and so composite particles
are produced which, as the egg segments and develops, give rise
to cells that are strictly hybrids and which therefore exhibit
variation. The ovum thus is the conservative element which
transmits the characteristics that have already been acquired.
The male cell is peculiarly that which stores up the disturbing
effects of a changing environment. It especially leads, therefore,
to variability in the offspring, to the production of individual
differences.

This ingenious hypothesis enables Brooks to explain a great
variety of inheritance phenomena and to overcome several ser-
ious objections to the unassisted selection theory. Whatever
truth there may or may not be in the special ideas of the book, it-
remains to-day a stimulating and suggestive contribution, and it
is properly looked on as one of the factors that have in recent
years focussed the attention of the biological world on the prob-
lems of heredity.

Minor papers dealing with heredity and evolution, the causes
of variation, and the determination of sex, appeared from time to
time. Sections of the "Foundations of Zoology" (1899) show, too,
that Brooks' interest in the questions discussed in the "Law of

A SKETCH OF HIS LIFE M

Heredity' ' remained active during life. Two of his last addresses
(1906-1909) deal with our concepts of heredity and variation.
In these he emphasizes the fact that the nature of an organism
is not implicit in the egg, or in the organism indeed at any time
of its life, but that it depends on a continuous reciprocal interac-
tion between the organism and its environment. Such interac-
tion leads in any particular case to a result which could not be
calculated from a knowledge, however complete, of the egg itself
since it is dependent not only on the organism but on the action
of the total environment. The outcome of such interaction is
the production of individuals which are never quite alike, although
they may resemble one another closely. The occurrence of like-
nesses, or inheritance, and the occurrence of differences, variation,
are thus not two processes but two views of the single process of
reciprocal interaction. The idea that they are distinct is an error
into which we fall through concentrating our attention at one time
on the resemblances, and again on the differences between individ-
uals. These considerations, he thinks, show the uselessness of
theories which postulate an inheritance substance and explain
individual differences as the result of various combinations of its
particles.

These addresses show that Brooks has in some measure shifted
his standpoint since the time of the "Law of Heredity." He no
longer is in a mood to employ evolution (determinant) hypotheses
to account for development. He now looks on the development
of the individual, and that of races also, as epigenetic in nature.
What will be the outcome of an individual egg depends on the
interaction between egg and environment, not on a determinate
mechanism in the egg. The pre-cambrian fauna has given rise
to the living beings of to-day. But the latter were not implicit
in the former, for with the same ancestors the course of evolution
might have been different had the sum total of environmental
influences been different.

Writings on the Principles of Science.19 Brooks dwelt often in
conversation and in minor writings, and always with an earnest

19 Professor H. V. Wilson, University of North Carolina.

40 WILLIAM KEITH BROOKS

pleasure, on the nature and intellectual value of what we can learn.
His thoughts in this field of the principles of science were event-
ually embodied in his lectures on the "Foundations of Zoology"
(1899). This remarkable book "belongs to literature, as well as
to science. It belongs to philosophy as much as to either, for it is
full of that fundamental wisdom about realities which alone is
worthy of the name of philosophy."20

Brooks was distinctly the philosophic type of naturalist. He
was fully informed, critical, and constructive in special fields,
but always aware that such fields were merely parts of a larger
whole. Thus through the bent of his mind Brooks, the keen-
sighted pioneer, and influential biologist, was also interested in
and in thorough sympathy with life and living in all aspects, past,
present and future, intellectual, emotional, and religious. Per-
haps for that reason, too, he was a great teacher and inspirer
of men.

The "Foundations" is essentially a discussion of the nature
of scientific knowledge. It is the wise talk of an experienced,
reflective naturalist of ripe years addressed primarily to younger
fellow-workers in the fields of science. The argument which makes
its way through pages and sometimes whole chapters of illustra-
tions and digressions, interesting and suggestive in themselves,
proceeds about as follows :

Our only knowledge of nature is through experience. Through
experience we learn that one sort of event follows another, and
this sequence, which we come to expect, constitutes for us the
order of nature. Nevertheless there is no reason to believe that
there is an inherent necessity in this order, for we never perceive
the presence of any intrinsic causal connection between the pre-
ceding event (cause) and the succeeding one (effect) .

When our knowledge of any part of nature has so far developed
that we know the order of events, and so can predict the later
steps in the series of occurrences, once the earlier have been noted,
we say that we understand and can mechanically explain that
particular set of phenomena. At present a gap separates vital

20 President D. S. Jordan.

A SKETCH OF HIS LIFE 41

from non-vital phenomena — to say that life is the sum of the
physical properties of protoplasm is to make a dogmatic assertion,
although to gainsay it is to make another. But with the progress
of science this gap may be bridged over at some time. Should it
be bridged over, and life in all of its aspects be found to be "pro-
toplasmic, " still we should not know why synthesis of compounds
results in an organism or why a vital action is the outcome of
protoplasmic changes. In respect to organisms and vital actions
we should still be where we are now in respect to simple gravi-
tation phenomena, for with respect to them all that we can say
is that the stone will fall (if the future be like the past), but
why it should fall we do not know.

This being the nature of our knowledge, present and future,
what should the biologist seek to discover, and what are the
problems that peculiarly concern him? Life is defined as a
continuous adjustment of internal to external relations (Spencer),
and it is pointed out that synthesized protoplasm, even were it
capable of nutrition, growth, reproduction, and contraction,
would not be a living thing if it were not also able to maintain
persistent adjustment to the shifting world around it. The
essence of the living thing and that which distinguishes it from
other forms of matter is this very adjustment. Fitness, adaptive
response, is therefore what we should seek to study in biology. The
mechanism itself is of subordinate importance. Study it as we
may, we cannot thus go far forwards, since our knowledge of nature
never includes a perception of any necessary causal connection
between events, such as would make it possible to discover vital
phenomena by reasoning deductively from protoplasmic pecu-
liarities. A corollary of practical import is that the naturalist
should endeavor to study living things in connection with their
environment.

Biology being thus defined as the study of adaptive response, the
nature and evolution of man's reason and knowledge fall within
its scope. For these are conceivably but the outcome of adaptive
responses in the beginning as simple as the geotropism of a seed-
ling's radicle. The ability, for instance, to make a distinction
between what in practical life we call a truth, a real occurrence,

42 WILLIAM KEITH BROOKS

and an error or illusion, is to be looked on as a useful response
that has been acquired through selection. Man's knowledge,
then, is of the peculiar kind that is useful to him. He may not
yet know as much as is good for him, but he at least has acquired
a store of the kind of knowledge that preserves him in the struggle
for existence.

Viewing man thus from the biological standpoint Brooks at-
tempts to deal with two human characteristics, the consciousness
that the will is free and that the individual carries a moral respon-
sibility. These, like all other vital characteristics, he thinks,
may possibly sometime be shown to be part of the order of
nature and in that sense mechanical . ' ' Rational action may some-
time prove to be reflex from beginning to end." And yet in the
face of this possibility, Brooks would still maintain that the will
is free and moral responsibility real. To some this will seem a
difficult thesis.

Underlying the scientific inquiry as to the character of our
present knowledge and of that which possibly we may acquire
about nature, is the metaphysical question, "what is nature?1'
This question Brooks does not attack in the fashion of construc-
tive technical philosophy. He makes no attempt to define reality.
His purpose in dealing with the matter is plainly the practical
one of showing us what we need not believe. He says in effect,
if then our knowledge of all nature is and will continue to be of one
sort, viz., that phenomena follow one another regularly and
(supposing the future to be like the past) in predictable fashion,
but without our ever learning why they so follow one another,
there is not now nor will there be in the future any necessity drawn
from science to believe in a fixed, necessary, determinate nature.
If in any quarter it is imagined that the progress of science neces-
sitates or may necessitate such a belief, this is a grave error: in
his own words, "The belief that the establishment of scientific-
conceptions of nature shows that after the first creative act,
the Creator has remained subject, like a human legislator, to
his own laws, is based upon utter misapprehension of science, and
upon absurd and irrational notions of natural law." In the second
place we are in nowise forced to believe by anything in science that

A SKETCH OF HIS LIFE 43

protoplasm and life are necessarily linked together: " . . .if
it be admitted that we find in nature no reason why events should
occur together except the fact that they do, is it not clear that we
can give no reason why life and protoplasm should be associated
except the fact that they are? And is it not equally clear that this
is no reason why they may not exist separately ?"

The next step in this survey and analysis of fundamental as-
pects of nature brings us to positive belief itself. As so often said,
science quite fails to find in matter and motion any intrinsic
virtue which sustains and directs the sequence of phenomena, and
is absolutely restricted to thediscovery of the mere sequence which
itself calls for (metaphysical) explanation. Hence there is nothing
in science which has any bearing on the causal origin or on the
reality of anything in nature, and we must go elsewhere for the
foundations of the belief that we may entertain in respect to such
matters. Brooks believes that "nature is intended" to be as it
is, and is a language which a rational being may read. Since the
rational being is perhaps himself a part of nature's mechanism, this
is equivalent to saying that one part of the mechanism is cognizant
of the purpose that animates the whole. This purpose is the effect
of a power, a sustaining and directing intelligence outside nature,
to which both the origin of nature and its maintenance from day
to day are due. It is not something which once for all set a deter-
minate cosmos spinning along the path of time with a full comple-
ment of " eternal iron laws." It is something which is at work
now, under every phenomenon. This is obviously Brooks' belief,
although being no propagandist he is far from enforcing it, indeed
leaves it in a measure to be inferred. What he wishes to make
plain is that science does not tell us why events happen a< we
learn they do, and so it tells us nothing of 'ultimate reality. The
question why the events we expect (from experience) should be
those that come to pass concerns not science but "the natural the-
ologian; for it is the same as the question, What is the Cause of
Nature? To this all must seek an answer for themselves; for
each has at his command all the data within the reach of any
student of science.'

44 WILLIAM KEITH BROOKS

LIST OF PROFESSOR BROOKS' WRITINGS21

1864
Do animals reason? Wilkes' Spirit of the Times.

1874
A feather. Pop. Sci. Monthly, April, 1874.

1875

On an organ of special sense in the lamellibranchiate genus Yoldia. Proc. Amer-
ican Asso. Adv. Sci., Hartford meeting, August, 1874, 3 pp. 2 cuts. Printed
Salem, 1875.

Embryology of Salpa. Proc. Boston Soc. Nat. Hist., vol. 18, November 17, 1875,
7 pp., 1 pi.; also Monthly Microscopical Journal, London, July 1, 1876.

1876

Embryology of the fresh-water mussels. Proc. American Asso. Adv. Sci., Detroit >

meeting, 1875, 3 pp. Printed Salem, June, 1876.
A remarkable life-history and its meaning. American Naturalist, November,

1876, 16 pp. 17 cuts.

On the development of Salpa. Bull. Mus. Comp. Zool., vol. 3, March, 1876.
On the affinity of the Mollusca and Molluscoida. Proc. Boston Soc. Nat. Hist., (

vol. 18, February 12, 1876, pp. 225-235.

1877

A provisional hypothesis of pangenesis. American Naturalist, March, 1877, 4 pp.
(Abstract of paper read at Buffalo meeting. American Asso. Adv. Sci., Aug-
ust, 1876).

Parthenogenesis in vertebrates and molluscs. American Naturalist, October,
1877.

Instinct and intelligence. Pop. Sci. Monthly, September, 1877.

1878

Differences between animals and plants. Pop. Sci. Monthly, November, 1878. ^
The condition of women from a zoological point of view. Pop. Sci. Monthly,
June and July, 1879.

21 Compiled by Prof. E. G. Conklin. Professor Brooks made no list of his
publications and the following list has been compiled from many sources and
may not be entirely complete.

A SKETCH OF HIS LIFE 45

1879

The scientific results of the Chesapeake Zoological Laboratory, session of 1878.

Baltimore, Murphy, 1879, 168 pp., constituting part 1, vol. 1, Studies Bio-
logical Laboratory, Johns Hopkins University, containing the three following

papers by W. K. Brooks:
Preliminary observations upon the development of the marine prosobran-

chiate gasteropods; 47 pp., 1 pi.
The development of Lingula and the systematic position of the Brachio-

poda; 70 pp., 6 pis.

The larval stages of Squilla einpusa Say; 5 pis.
The development of the digestive tract in molluscs. Proc. Boston Soc. Nat. Hist., :

vol. 19, 1879, 4pp.
Abstract of observations on the development of the American oyster. Zool.

Anzeiger, 1879.
Abstract of observations upon the artificial fertilization of oyster eggs, and on the

embryology of the American oyster. American Journ. Sci., December, 1879.

3pp.
Observations upon the early stages in the development of the freshwater pul-

monates. Studies Biol. Lab. Johns Hopkins University, vol. 1, 1879, 26 pp.,

4 pis.

1880

Observations upon the artificial fertilization of oyster eggs and on the embryology
of the American oyster. Ann. and Mag. Nat. Hist., London, 1880.

The biology of the American oyster. N. C. Med. Press, 1880.

The artificial fertilization of oyster eggs and the propagation of the American
oyster. American Journ. Sci., 1880.

The development of the American oyster. Maryland Fish Commission Report,
1880, 101 pp., 10 pis.

The development of the oyster. Studies Biol. Lab., Johns Hopkins University^
vol. 1, 1880, 115 pp., 11 pis. (Reprinted from the preceding.)

The acquisition and loss of a food yolk in molluscan eggs. Studies Biol. Lab.,
Johns Hopkins University, 1880, 7 pp., 1 pi.

Budding in free Medusae. American Naturalist, 1880.

Embryology and metamorphosis of the Sergestidse. Zool. Anzeiger, Jahrg. 3, <
1880.

Amphioxus and Lingula at the mouth of the Chesapeake Bay. American Natural-
ist, vol. 13, 1880.

The young of the crustacean Lucifer, a Nauplius. American Naturalist, Novem-
ber, 1880.

The development of the squid. Anniversary Mem. Boston Soc. Nat. Hist., 1880,
21 pp., 3 pis., 40.

The homology of the cephalopod siphon and arms. American Journ. Sci., vol.
20, October, 1880, 3 pp., 1 cut.

The rhythmical character of the process of segmentation. American Journ. Sci.,
vol. 20, 1880, p. 293.

4f) WILLIAM KEITH BROOKS

1881

Du dcveloppement de la lingula et de la position zoologique des brachiopods. Arch.

de Zool. Exp. et Generate, 1881.
Lucifer,: A study in morphology. Proc. Royal Soc., April, 1881, pp. 1-3. (Ab- y^

stract.)

1882

Origin of the eggs of Salpa. Biol. Studies, Johns Hopkins University, vol. 2,

1882, pp. 301-312, 1 pi.
Lucifer: A study in morphology. Phil. Trans. Royal Society, London, vol. 173,

1882, 80 pp., 11 pis.

Handbook of invertebrate zoology. 400 pp., 202 figs. Boston, Cassino, 1882.
Chamisso and the discovery of alternation of generations. Zool. Anzeiger, Jahrg.

5, 1882.
The metamorphosis of Alpheus. Johns Hopkins University Circulars, vol. 2,

no. 17, 1882.
On the origin of alternation of generations in Hydro-medusse. Johns Hopkins

University Circulars, vol. 2, no. 22, 1882; also Ann. and Mag. Nat. Hist.,

vol. 2, 1883.
The metamorphosis of Penseus. Johns Hopkins University Circulars, vol. 2,

1882; also Ann. and Mag. Nat. Hist., vol 2, 1883.

y. Speculative zoology. Pop. Sci. Monthly, December, 1882, and January, 1883.
On some methods of locomotion in animals. A lecture delivered to the employes

of the Baltimore and Ohio Railroad Co., Baltimore. Printed by I. Frieden-

wald for free distribution among the employe's of the Baltimore and Ohio

Railroad Co., 20 pp., 10 cuts, 1882.

1883

v( List of Medusae found at Beaufort, N. C., during summers of 1880-1881. Studies

Biol. Lab. Johns Hopkins University, vol. 2, 1883, pp. 135-146.
Report of the Chesapeake Zoological Laboratory, summer of 1882.
The law of heredity. Baltimore, Murphy, 1883, 336 pp.
Notes on the Medusae of Beaufort, N. C., II: Turritopsis nutricula (McCrady).

Studies Biol. Lab. Johns Hopkins University, vol. 2, 1883, pp. 465-475.
The first zoea of Porcellana. Studies Biol. Lab. Johns Hopkins University, vol.

2, pp. 58-62, pis. 6-7. (With E. B. Wilson.)
Alternation of periods of rest with periods of activity in the segmentation of eggs

of vertebrates. Studies Biol. Lab. Johns Hopkins University, vol. 2, 1883, pp.

117-118.
The phytogeny of the higher Crustacea. Science, vol.2, pp. 790-793;also NewZea-

land Journal of Science, vol. 2.
Reviews of work on ccelenterates in Weekly Summary of the Progress of Science.

Science, vol. 1, pp. 50, 81, 230, 287, 344, and 553; Science, vol. 2, pp. 54, 692, 773,

and 832.

A SKETCH OF HIS LIFE 47

1884

Is Salpa an example of alternation of generations? Nature, vol. 30, 1884.

On the life-history of Eutima and on radial and bilateral symmetry in hydroids.

Zool. Anzeiger, Jahrg. 7, 1884.
The development and protection of the oyster in Maryland : Report of the Oyster

Commission of the State of Maryland, Annapolis, Maryland; 183 pp., 7 maps,

13 pis., 4°-
On a new law of variation. Johns Hopkins University Circulars, vol. 4, no. 35,

December, 1884, pp. 14-15.
A new law of organic evolution. Science, vol. 4, 1884, pp. 532-534.

1885

On the artificial propagation and cultivation of the oyster in floats. Johns Hop-
kins University Circulars, 1885, vol. 5, no. 43, p. 10; Also Science, vol. 6, 1885,
pp. 437-438.

Oyster farming in North Carolina. Forest and Stream, New York, 1885.

Influences determining sex. Pop. Sci. Monthly, January, 1885, pp. 323-330.

Can man be modified by selection? Pop. Sci. Monthly, May, 1885.

Abstract of researches on embryology of Limulus polyphemus. Johns Hopkins
University Circulars, vol. 5, no. 43, October, 1885, pp. 2-5. (With Adam
Bruce.)

Notes on Stomatopoda. Johns Hopkins University Circulars, vol. 5, no. 43, Octo-
ber, 1885, pp. 10-11.

A note on inheritance. Johns Hopkins University Circulars, vol. 5, no. 43, Octo-
ber, 1885, pp. 11-12.

1886

Life on a coral island. Pop. Sci. Monthly, October, 1886; also Baltimore Sun,
August 16, 1886.

The anatomy and development of the Salpa-chain. Studies Biol. Lab. Johns
Hopkins University, vol. 3, 1886, 22 pp., 15 cuts. 2 pis.

Notes on the Stomatopoda. Ann. and Mag. Nat. Hist., vol. 17, 1886.

Report on the Stomatopoda collected by H. M. S. Challenger. Challenger Re-
ports, vol. 16, 1886, 114 pp., 16 pis., 4.°

The life-history of the Hydro-medusa : A discussion of the origin of the Medusae
and of the significance of metagenesis. Mem. Boston Soc. Nat. Hist., vol.
3, 1886, 67 pp., 8 pis.

The zoological work of the Johns Hopkins University, 1878-86. Johns Hopkins
University Circulars, vol. 6, no. 54, December, 1886, pp. 37-39.

The Stomatopoda of the Challenger expedition. Johns Hopkins University Cir-
culars, no. 49.

Development and alternation of generations of the Hydro-medusfe. Proc.
Acad. Nat. Sci., Philadelphia, March 3, 1886; also Science, vol. 7, no. 163.

48 WILLIAM KEITH BROOKS

1887

The scientific work of Adam Todd Bruce — A sketch. (In "Observations on the
embryology of the insects and arachnids," by Adam Todd Bruce.) Memorial
Volume, Johns Hopkins University Press, Baltimore, 1887.

1888

V\ The life-history of Epenthesis McCradyi, n. sp. Studies Biol. Lab. Johns Hop-
kins University, vol. 4, 1888, 15 pp., 3 pis.

The growth of jellyfishes. Pop. Sci. Monthly, September (pp. 577-588, 7 cuts),
and October, 1888.

On a new method of multiplication in hydroids. Johns Hopkins University Cir-
culars, vol. 7, no. 63, February, 1888, pp. 29-30.

Note on the ratio between men and women. Johns Hopkins University Circu-
lars, vol. 7, no. 63, February, 1888, pp. 30-31.

1889

Artificial propagation of sea fishes. Pop. Sci. Monthly, July, 1889, pp. 359,-367.
A preliminary abstract of the researches of W. K. Brooks and F. H. Herrick on the

life-history of Stenopus. Johns Hopkins University Circulars, vol. 8, 1889.
The Lucayan Indians. Pop. Sci. Monthly, November, 1889, pp. 88-98.
^ On the Lucayan Indians. Mem. National Acad. Sci., vol. 4, no. 10, 1889, 9 pp.,

12 pis.
What conditions are necessary for the establishment by selection of a deaf variety

of the human race? Report of the Royal Commission on the Blind, the Deaf,

and the Dumb, etc; London, 1889.

1890

On the relationship between Salpa and Pyrosoma. Johns Hopkins University
Circulars, vol. 9, no. 80, April, 1890, pp. 55-56.

The structure and development of the gonophores of a certain siphonophore be-
longing to the order Auronectse. Johns Hopkins University Circulars, vol.
9, no. 88, 1890. (With E.G. Conklin. )

Course of reading for graduate and special students in morphology at the Johns
Hopkins University. Johns Hopkins University Circulars, December, 1890,
p. 37.

1891

The oyster. Johns Hopkins University Press, Baltimore, 1891.
On the early stages of echinoderms. Johns Hopkins University Circulars, May,
1891, p. 101.

1892

The embryology and metamorphosis of the Macrura. Johns Hopkins University
Circulars, vol. 11, no. 97, April, 1892, pp. 65-72. (With F. H. Herrick.) (In-
troductory chapter of the following work.)

A SKETCH OF HIS LIFE 49

The embryology and metamorphosis of the Macrura. Mem. National Acad. Sci.,

vol. 5, 1892, 135 pp., 57 pis. (With F. H. Herrick.)
The English plan for the Columbus Marine Biological Station in Jamaica. Letter

to New York Daily Tribune, April 4, 1892.

1893

The origin of the organs of Salpa. Abstract of chapter 14 of the Memoir on the

genus Salpa. Johns Hopkins University Circulars, vol. 12, no. 106, January,

1893, pp. 93-97.
Aspects of nature in the West Indies. From the notebook of a naturalist. Scrib-

ner's Monthly, July, 1893.
The genus Salpa. Memoirs Biol. Lab. Johns Hopkins University, 1893, 303 pp.,

46 pis., 4.°
Salpa in its relation to the evolution of life. Studies Biol. Lab. Johns Hopkins

University, vol. 5, 1893, pp. 129-212.
Maryland, its resources, industries, and institutions.

Chap. 7: Fish and fisheries.

Chap. 8: The oyster.
The nutrition of the Salpa embryo. Johns Hopkins University Circulars, January,

1893, pp. 97-98.

1894

The origin of the oldest fossils and the discovery of the bottom of the ocean. Jour.

Geology, July and August, 1894; also Johns Hopkins University Circulars,

January, 1895.
The origin of the food of marine animals. Bull. United States Fish Commission,

1894; The World's Fisheries Congress, Chicago, 1893, 5 pp.
Address, in proceedings of the convention to consider the oyster question, held at

Richmond Chamber of Commerce, Richmond, Virginia, January 12, 1894, pp.

33-37.

1895

An old naturalist — Conrad Gesner. Pop. Sci. Monthly, May, 1895, pp. 49-59, 12
cuts.

A review of Huxley's essays. The Forum, November, 1895.

An inherent error in the views of Galton and Weismann on variation. Science,
vol. 1, February 1, 1895, pp. 121-126.

Can an organism without a mother be born from an egg? Science, vol. 1, Febru-
ary 8, 1895.

The tyranny of the monistic creed, a review. Science, new series, vol. 1, April
5, 1895, pp. 381-384.

The sensory clubs or cordyli of Laodicea. Journ. Morphology, vol. 10, 17 pp., 1 pi.

Science or poetry. Science, vol. 2, no. 40, October 4, 1895, pp. 437-440.

THE JOURNAL OF EXPERIMENTAL ZOOLOGY, VOL. 9, NO. 1.

50 WILLIAM KEITH BROOKS

1896

The study of inheritance. Pop. Sci. Monthly, February (pp. 480-492), and March
(pp. 617-626), 1896.

Woman from the standpoint of a naturalist. Forum, November, 1896.

Is there more than one kind of knowledge? Science, April 24, 1896.

The origin of the oldest fossils, and the discovery of the bottom of the ocean.
Smithsonian Report for 1894, pp. 359-376, Washington, 1896.

Logic and the retinal image. Science, March 20, 1896, pp. 443-444.

Lyell and Lamarck: A consideration for Lamarckians. Johns Hopkins Univer-
sity Circulars, vol. 15, no. 726, June, 1896, pp. 75-76. (Reprinted from Na-
tural Science, February, 1896, vol. 9.)

Lyell. Johns Hopkins University Circulars, vol. 15, no. 726, June, 1896, p. 78.

Budding in Perophora. Johns Hopkins University Circulars, January, 1896, p.
79. (With George Lefevre.) (Abstract of paper in National Acad. Sci.,
April 23, 1896.)

Note on anatomy of Yoldia. Johns Hopkins University Circulars, January,
1896, p. 85. (With Oilman Drew.)

Zoology and biology. Science, May 8, 1896, p. 708.

The retinal image once more. Science, April 24, 1896.

Lamarck and Lyell: A short way with Lamarckians. Natural Science, vol. 9.

Lyell and Lamarckism : A rejoinder. Natural Science, vol. 9.

1897

The expedition to Jamaica in the summer of 1897. Johns Hopkins University

Circulars, November, 1897.

Testimony versus evidence. Science, vol. 2, no. 49, December 6, 1897, p. 771-773. v
William Harvey as an embryologist. Johns Hopkins Hospital Bulletins, nos. 77 >

and 78, 1897, 20 pp.
Anglo-Saxon versus Graeco-Latin. Natural Science, vol. 10, no. 63, May, 1897,

p. 360.

1898

Migration. Pop. Sci. Monthly, April, 1898, pp. 784-798.

Zoology and the philosophy of evolution. Science, new series, vol. 8, no. 208,
December 23, 1898, pp. 881-893.

1899

The foundations of zoology. The Macmillan Co., New York, 339 pp.

Review of the "Wonderful Century," by A. R. Wallace. Science, April 7, 1899.

The Wonderful Century — A Review. Pop. Sci. Monthly, November, 1899, pp.

25-31.

Thoughts about universities. Pop. Sci. Monthly, July, 1899, pp. 349-355.
Mivart's Ground work of Science — A Re view. Pop. Sci. Monthly, February, 1899.

A SKETCH OF HIS LIFE 51

Scientific Laboratories: An address delivered at the dedication of the biological
laboratory. Bulletin Western Reserve University, October, 1899, 20 pp.
(Reprinted in Bull. Johns Hopkins University, vol. 10, no. 104, November,
1899.)

Truth and error. Science, vol. 9, no. 213, January 27, 1899, pp. 121-126.

1900

The lesson of the life of Huxley. Smithsonian Report, 1900, pp. 701-711.
Review of "Marriages of the Deaf in America, by E. A. Fay, Volta Bureau, Wash-
ington, D. C., 1898." Pop. Sci. Monthly, January, 1900.

1902
Is scientific naturalis-rh fatalism? Prbc. •'American Philos. Soc., vol. 41, 1902.

:

.

The intellectual conditions for the science of embryology. Science, new series,
vol. 15, nos. 377 and 378, 1902, pp. 444-454, pp. 489^92.

1903

On a new genus of hydroicl jellyfishes, Dichotoma. Proc. American Philos. Soc.,
vol. 42, 1903, 3 pp. 1 pi. (Read April 4, 1902).

1905

The oyster. A popular summary of a scientific study. (2d ed. revised.) John
Hopkins University Press, 1905, 225 pp., 16 pis.

1906

Heredity and variation, logical and biological. Proc. American Philos. Soc.,

1908, 7 pp. (Read April 20, 1906).
The affinities of the pelagic tunicates, No. 1 : On a new Pyrosoma. Mem. National

Acad. Sci., vol. 10, 1906, 5 pp. 2 pis.
Dipleurosoma a new genus of Pyrosoma. Johns Hopkins University Circulars,

1906, no. 5, pp. 98-99, 2 cuts.

Evolution. Article in the "Reference Hand book of the Medical Sciences," 1906,
pp. 733-736.

1907

Joseph Leidy. Pop. Sci. Monthly, April, 1907.

Joseph Leidy. Pioneers of American Science. American Museum of Natural

History, New York, April, 1907, pp. 23-25; also in Anatomical Record, No. 5,

January 1, 1907.
On Turritopsis nutricula (McCrady). Proc. Boston Soc. Nat. Hist., vol. 33, no. 8,

1907, pp. 429-460, pis. 30-35. (With Samuel Rittenhouse.)

52 WILLIAM KEITH BROOKS

The homologies of the muscles of the subgenus Cyclosalpa. Johns Hopkins Uni-

versity Circulars, 195, March, 1907, pp. 1-2.
The foundations of zoology (2d. ed.) Macmillan Co., New York, 1907.

1908

Biographical memoir of Alpheus Hyatt. Biographical Memoirs, National Acad.

Sci., vol. 6, 1908.
The origin of the lung of Ampullaria. Carnegie Institution of Washington, Pub.

102, 1908, .pp. 95-104, pis. 1-7. (With Bartgis McGlone.)
The pelagic Tunicata of the Gulf-Stream; Parts 2, 3, and 4: 'On Salpa floridana,

the subgenus Cyclosalpa, and on Oikopleura tortugensis, sp. nov. Carnegie

Institution of Washington, Pub. 102, pp. 73-94, pis. 1-8. (4withCarlKellner),
The province of science, -Popular Science Monthly, 1908, pp. 268r271. .
*~»vi*l b*~»V.Ui* c-'nTii*,*™ C«-V»M, *"«"**$

v

1909

Are heredity and variation facts? Address at 7th International Zoological Con-
gress, Boston, 1907; also reprinted as a memorial publication.

UNPUBLISHED WORKS

The axis of symmetry of the ovarian egg of the oyster. In press 1905, but still in

MS., not published.
An extensive work on Salpa.
Lowell Lectures.