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1) Contributions from the Botanical Laboratory, vol. 1
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PUBLICATIONS OF THE
UNIVERSITY OF PENNSYLVANIA.

i

CONTRIBUTIONS

■.i\ -

i-ti ■

FROM THE

BOTANICAL LABORATORY.-

VOL. I.

PUBLICATIONS OF THE
UNIVERSITY OF PENNSYLVANIA.

CONTRIBUTIONS

FROM I'HE

Botanical Uaboratory

VOLUME I.

WITH XXXVI. PLATES.

1892-97.

~f-^ — ■ — «--

• • < •

PHILADELPHIA:

PUBLISHED FOR THE UNIVERSITY.

1897.

iii

sr

INDEX.

I'AGE.

Aniphicarpica monoica. Life History of 270

Brunella vulgaris^ Nascent Variety of 64

Cross, L. B., on Structure and Pollination of Eiipatoritim 260

Dioftcra viuscipn/d. Contributions to History of 7

Dioniea viuscipula, Abnormal Intloresence of 45

Epigiea rcpt'ns. Observations on 5^

Eupatorium agetatoides, and E. calestinutn^ Structure and Pollination

of Flowers of 260

Greenman, J. M., on Movements of Leaves of Melilotiis alba 66

Harshberger, J. W., on Maize 75

Macfarlane, J. M., Contributions to the History of Diotuea juuscipula . . 7

Maize : A Botanical and Economic Study 75

Mangrove Tannin, Irimble on 50

Melilotus alha^ Movements of Leaves of 66

Pennington, Mary E. , on Spirogyra nitida ... 203

Rothrock, J. T., Monstrous Specimen of Rudbfckia hirta 3

Rothrock, J. T., Na^cetil -Variety of- Prunella vulgaris 64

Schively, Adeline F., on A*»phicarpiea monoica 270

Spirogyra wV/V/tfj.a Cheraico-Physiologioal study of . 203

Triml)le, H,, on Mangrove Tannin 5°

Wilson, W, P., on Epig^c r-etens .- ; 56

Wilson, W. P., on Movements of Leaves of Melilotus alba 66

a"

publications

OF THK

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Jastrow, Jr., Professor of Arabic. 60 cents.

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By William A. Lamberton, Professor of the Greek Language and
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Pak kop pill. By Stewart Culin, Secretary of the Museum of
Archaeology and Paleontology. 40 cents.

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1. Recent Archaeological Explorations in the Valley of the Dela-

ware River. By Charles C. Abbott, Sometime Curator of the
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Volume ill.

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$1.50-

2. A Primer of Mayan Hieroglyphics. By Daniel G. Brinton, Pro-

fessor of American Archix,'ology and Linguistics. ^1.20.

i(«9&?^

i !|

til

Or:

568:^9

Volume IV.

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Easton, Professor of English and Comparative Philology. 60 cents.

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English. (Just appearing.)

Volume V.

Two Plays of Miguel Sanchez (surnamed "El Divino "). By

Hugo A. Rennert, Professor of Romance Languages and Literatures*

Volume VI.

a. The Antiquity of Man in the Delaware Valley.

/'. Exploration of an Indian Ossuary on the Choptank River, Dor-
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E. D. Cope ; and an examination of traces of disease in the bones, by
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SERIES IN

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(

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M-

2.

CONTKIISIJTIONS FROM

The Botan ical Laboratory.

Volume I— No. i. 1892.

(Plates I-XIII.)

1. A Monstrous Specimen of Rudbeckia hirta.L. ByJ. T. Rothrock,

13. S., M.D.

2. Contributions to the History of Dionaea Muscipula, Ellis. By J.

M. Macfarlane, D.Sc.

3. An Abnormal Development of the Inflorescence of Dionaea.

By John W. IIarshbkrger, A.B., B.S.

4. Mangrove Tannin. By H. Trimhli:, Ph. M.

5. Observations on Epigaea repens, L. By W. P. Wilson, D.Sc.

6. A Nascent Variety of Brunella vulgaris, L. By J. T. Rothrock,

B.S., M.D.

7 Preliminary Observations on the Movements of the Leaves of
Melilotus alba, L., and other Plants. By W. P. Wilson, D.Sc ,
and J. M. Greenman.

Volume I No. 2.

(Plates XI V-X VII.)

4. Maize: A Botanical and Economic Study. By John W. Harsh-
berger, Ph.D.

CONXRIL3UTIONS KROM

The Zoological Lal^oratory.

Volume 1— No. i.

The Correlations of the Volumes and Surfaces of Organisms.

By John A. Ryder, Ph.D. (Plate I.)

The Growth of Euglena Viridis when Constrained Principally
to Two Dimensions of Space. By John A. Ryder, Ph. D.
(Plate II.)

Descriptions of Three New Polychseta from the New Jersey
Coast. By J. Percy Moore. (Plates IlI-lV.)

Volume I— No. 2.

t

On the Embryos of Bats. By Harrison Allen, M.D. (Plates
V-VIII.)

(r/

A Monstrous Specimen of Rudbeckia hirta, L.

By J. T. Rothrock, B. S., M. D.

(WITH PhAT^iS I, II, AND III,)

THROUGH the kindness of Mr. Francis Windle, of
West Chester, Pa., a most remarkable illustration of
proliferation and floral modification in the above-named
plant has come to my knowledge.

There were two specimens, taken from a field which
had been previously mowed. He removed them from the
ground and planted them in a flower-pot near his home,
where he had them under observation. We may fairly as-
sume that if the stems had not already produced flowers that
season, they were well advanced toward that period when cut
down. Hence they represented a weakened second-growth.
In the larger of these two heads there were nineteen well-
developed secondary heads and four others less well de-
veloped, but following in the course of the larger ones.

The order of development was acropetal. Hence the m.ost
of the departure from a typical condition was evinced by the
ray flowers. Of the latter there was not a single normal one
in the entire floral mass, which is represented in the unnum-
bered illustration accompanying this brief sketch.

In general, it may be stated that while the proliferation of
the tubular flowers was not so marked, still instances were
not wholly wanting.

The most striking fact was, that from the central part of
both ray and disk flowers there were found proliferated
foliar and floral axes in the position which belonged to the
styles. This is none the less singular when one remembers
that the ray of Rudbeckia is neutral. From these proliferated
masses, in many instances, there were secondary : ;^ifera-

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MUTILATED TEXT

4 Rothrock\—A Monstrous Specimen of

tions similar to the first, except that they were smaller and
less plainly differentiated.

The tendency in the stigmas of the tubular flowers to be-
come enlarged, green, and leaf-like was very positive. Fig.
6 shows this. Not only so, but in that special instance the
hairs were characteristic of the species, and an upper and a
lower leaf-like surface could readily be observed.

One tubular flower had five stamens, which though inserted
properly on the tube of the corolla, were absolutely separate
as to their anthers from each other. Its stigmas had each an
evident midrib bordered by a narrow wing of parenchyma.

In some instances, where the stigmatic enlargement was not
great, there were traces of a normal stigmatic surface. Unfort-
unately examinations of the ovaries was not always made,
but in a few, ovules, probably unfertilized, were seen. It
should also be stated that in one tubular flower, whose
stigmas were green and decidedly enlarged, a well developed
ovule was found. It could not have been fertilized.

It is, of course, hard to admit that these proliferated styles
and stigmas represent axial structures ; but it appears equally
difficult to avoid that conclusion, if one may reason from a
single, or from a few monstrous specimens.

It is not clear, however, that Linnaeus had not some
suspicion that such cases were of an axial nature, in other
than the aggregates. It is not easy to see how he could, with
his views, have excluded this Rudbeckia,^ with its enlarged,
leaf-like stigmas. There is another question raised by this
specimen, /. e., whether after all the Rudbeckia is not to be
regarded as reducing to an absurdity that form of morpho-
logical reasoning which relies so implicitly upon a few
monstrous specimens to furnish a clue to the essential nature
of normal structures. In other words, is it a safe concession
to allow that, in science, an exception proves the rule }

My attention has recently been called to some monstrous
flowers of Digitalis, which, appearing at the summit of the
stem, had resulted in the production of an unusual number
of stamens.

i See Phil. Bot. 124.

Rudbeckia hirta^ L.

5

The Digitalis and the Rudbeckia seem to support the view
that such departures from the type are most likely to occur
in individuals which are exhausted, or are from some cause in
a condition of reduced vitality. The last energy of the plant
is extended in the line of self-perpetuation. This energy,
however, is often wasted in the faulty attempt to multiply
the reproductive organs inordinately, rather than to perfect
a few. Zoologists long since recognized the fact that multi-
plication of similar parts was an indication of a lower place
in the scale of animal life.

The illustrations probably will explain the structural
peculiarities better than any lengthened descriptions, and
a brief explanation is appended.

The large unnumbered figure represents the whole head of
the Rudbeckia, enlarged two diameters.

Fig. I shows, on the left, a cluster of tubular flowers
from the side of the head. Outside this are two involucral
bracts. Between the tubular flowers and the bracts rises the
tube of a ray flower, out of which three attempted ray flowers
spring. From the uppermost one of these is an attempted
secondary head of flowers. The dotted lines on the upper
side of this secondary head indicate the point from which two
flowers arise. The upper of these flowers is tubular, and
contains two distinct stamens and a three-cleft stigma, with
two of the lobes again parted. These stigmas grow from the
top of the rudimentary ovary below. The lower flower shows
four distinct stamens and a much enlarged and thickened
style and stigmas, the latter being distinctly green and leaf-
like.

The lower dotted branch (Fig. i) shows the point of origin
of the cluster of imperfect ray and tubular flowers with
which it is connected, and the extreme terminal, enlarged
illustration shows more plainly the character of one of the
flowers of the last clump. The artist, however, has in that,
and in Fig. 5, failed to show the rudimentary ovary which
appeared in each case.

Figs. 2, 3 and 4 show front and back views of a tubular
flower and its contents.

m

m

<

6 Rothrock.—Riidbeckia hirta, L.

•

Fig. 1 is another ray flower. Beside it and to the right
one recognizes a perfect tubular flower. From this ray,
in the centre, apparently, there arise two organs which
probably represent stigmas ; one of these, however, is enlarged
above as though it were attempting to form another ray; the
other remains thread-like. To the right of these stigmas,
indicated by the dotted lines, are two smaller abortive heads.
Fig. ^ shows the remarkable foliar expansion of the stigmas
arising from a well-marked tubular flower.

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I

Contributions to the History of Dionsea flus-

cipula, Ellis.

By J. M. Macfarlane, D.Sc.

{WITH PTAT^ IV.)

IT is not surprising that a voluminous literature already
exists regarding Dioncea. The movements and digestive
action of the leaf as studied specially by Darwin and T. A.
G. Balfour, coupled with the electrical conditions shown to
exist by Burdon Sanderson have invested the plant with more
than ordinary interest. But while Darwin harmonized the re-
lation of the plant to others that catch and digest insects, the
interest excited by this seems to have prevented his reaching
some of the more striking phenomena. And similarly Burdon
Sanderson, concerned mainly with electrical action, has not
followed out many of the suggestive lines which he touched.
Six years ago, while examining some leaves during a warm
forenoon in June, I was rather surprised to notice that appar-
ently in all cases two touches of the sensitive hairs were
needed to effect closure. This observation, often verified dur-
ing subsequent years, was made in forgetfulness of Sanderson
and Page's statement : " If one of the sensitive hairs of a
leaf" . ..." is carelessly touched (i. e.^ when full open)
the leaf usually closes. If, however, a hair is touched very
cautiously, with the aid of a camel's hair pencil, it can be pre-
dicted with certainty that no visible effect will be produced,
and a similar gentle contact maybe repeated several times be-
fore the leaf begins to answer to the irritation by any move-
ment." The writer was led, however, to a closer study of
the plant, and the results now given are the outcome of that
study. »

The subject may best be discussed under the following

8

Macfarlane. — Contrtbutions to the History of

heads : (i) Leaf closure ; (2) Leaf structure; (3) Leaf secre-
tion ; (4) Leaf opening ; while the keynote to the whole is
given in the words used by Burdon Sanderson in his lecture
before the Royal Institution/ "We have to do here not merely
with contractility, but with irrito-contractility."

L Leaf Closure.

{ol) By Mechanical Stimuli.

Except for the partial limitation contained in the statement
by Sanderson and Page above given, all observers have hith-
erto asserted that the leaf closes after a single touch. Thus
Darwin says,'^ ** These filaments are remarkable from their ex-
treme sensitiveness to touch, as shown not by their own move-
ment, but by that of the lobes." Sachs' says, '' any ungentle
touch of one of these bristles effects an instantaneous closure of
the two halves of the leaf." Detmer* says, " Werden die Fila-
mente, die auf der oberseite des Dionaea blattes vorhanden
sind, beriihrt, Z. B. mit einem kleinen Holzstiickchen, so
schliesst sich das Blatt sofort." Drude, Batalin, Munk and
others, speak in similar terms. Now we can only explain this
practically universal consensus of opinion to be due to obser-
vers having supposed that after the first touch of a hair with
non-closure, they imagined that the leaf was torpid, or that
they had missed touching one, and repeated the touch with
desired result. In any case it can now be asserted that under
such conditions as the plant is normally exposed to, two touches
are needed to cause closure. It matters not whether these
are communicated to the same hair, or to distinct hairs on the
same half, or on opposite halves of the leaf.

But though no apparent movement ensues after the first
touch, if a hair on the leaf of a vigorous plant be once touched
during warm weather, by steady attention one can readily see
that a peculiar rhythmic wave motion traverses the leaf halves
in line with the length of the blade for about five seconds

1 Nature; Vol. X, 1874, P "7.

2 Insectivorous Plants, p. 287.

■5 Physiology of Plants ; Eng. ed. 1887, p. 376-
* Pflanzen physiologische Practicum ; 1888. p. 64.

ii

Dioncea Miiscipula^ Ellis.

9

after contact, so that some molecular disturbance is being
propagated through the leaf substance, though not for the
time producing very manifest results.

Experiments were conducted to ascertain the length of time
which can intervene between first and second stimulus, before
the impression of the first was lost. In other words, the mem-
ory power of the protoplasm was tested. On bright days,
with temperature at 65°-72° F. in a cool house, healthy speci-
mens were selected. A leaf hair was stimulated once but the
leaf did not close. A second stimulus after ten seconds
caused closure with a latent period of less than two seconds.
Another leaf was once stimulated, and after an interval of
thirth-five seconds a second stimulus was applied when the
leaf closed rather slowly. A rather small leaf was restimu-
lated after lapse of twenty-five seconds, and a slight degree
of closure occurred. A third shock was given after thirty
seconds which caused closure to about one-third of its extent.
A fourth shock after twenty seconds produced further closure,
and after lapse of ten seconds a fifth shock completed the
process, though the movement of the halves was slow and
deliberate, about fifteen seconds being required.

Three leaves were operated on by stimuli at intervals of
thirty seconds, and after the third stimulus no closure-effect
was observable, but when the time-interval was reduced to
twciity seconds two successive shocks closed two of the leaves,
and another shock after ten seconds closed the third. All of
the above were smaller leaves, by about half, than the average
size attained in the wild state in North Carolina, and leaves
were therefore chosen nearly like the latter. A large healthy

leaf was acted on as follows :

ist stimulus no effect.

2d

\%J ■••• .••• a.

after sixty

seconds

3d

u

(( *i

(4

44

4th

((

(( ((

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44

5 th

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" fifty

u

extremely slight closure.

6th

((

44 44

4(

decided closure to extent of
about half.

7th

((

i( (t

44

closed loosely, and moved in
doing so for eighteen sec-
onds.

• -J

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m

10

Macfarlane. — Contributions to the History of

Another leaf was acted on as follows :
ist stimulus no effect.

2d

3d

4th
5th
6th
7th
8th
9th

after sixty seconds,

<(

(I

fifty

u

((

((

((

((

scarcely perceptible closure,
little if any closure,
no effect.

(4

{(

decided though slight closure.

leaf now closed fully one-
third and movement pro-
longed through twenty-five
seconds.

leaf closed after movement
of sixteen seconds.

A rather small leaf was then acted on as follows :
ist stimulus no effect.

loth

((

it

((

((

2d

3d

4th

5th

6th

7th

8th

9th

((

u

after sixty seconds,

((

((

((

((

it

u

t(
t(

fifty

u

u

((

u

((

((

u

((

<<

extremely slight closure.
no visible closure.

loth

u

u

l(

i(

(I

((

((

very slight closure.

leaf fully half closed after
movement of fourteen sec-
onds.

leaf closed after movement
of thirteen seconds.

A medium-sized healthy leaf was acted on thus :

ist stimulus no effect.

2d " after forty-five seconds

3d " " forty

4th " « thirty-five " extremely slight closure.

5th " " thirty " about half closed.

6th " " twenty-five " entirely closed.

Another of similar size as the last behaved as follows

ist stimulus no effect.

2d " after sixty seconds, "

u

({

3d
4th
5th
6th
7 th

u

((

((

((

u

((

(k

fifty

u
(I

(i

((
(i
(i

(I

very slight closure,
leaf half closed.
" closed.

Dioncea Miiscipnla^ Ellis.

A leaf of the same size as the last two was acted on thus :
ist stimulus no effect.

II

2d

3d

4th

5th

u

t(

(t

((

after forty-five seconds

({

i(

forty

thirty

" twenty-five *'

((

scarcely visible effect,
closed to about one-third,
closed slowly, the movement

occupying about thirteen

seconds.

Experiments were made on several large strong leaves. One
was sharply stimulated by pronounced bending of a hair, and
after lapse of forty seconds was restimulated when it closed,
after a movement of about three seconds. Another was
strongly stimulated, and restimulated after forty seconds with
scarcely visible effects, but after thirty-five seconds another
stimulus caused rapid closure.

The above experiments were all conducted at the Edinburgh
Botanic Garden, in a temperature of 65° F.'to 76° P., but from
August to October of the past year the writer experimented
in Washington and Philadelphia, on a liberal supply of healthy
plants kindly placed at his disposal by Mr. Oliver, of the
Washington Botanic Garden, and on three strong green plants
similarly provided by Prof. W. P. Wilson, of the Pennsyl-
vania University, Philadelphia. Recently, also, the writer had
the pleasure of studying Dioncea in its native haunts. The
plants were experimented on mostly at a temperature of 87
F. to 92° F. in the shade. All showed a more extended mem-
ory period. Thus the following cases may be selected as typ-
ical. A leaf was stimulated without effect, but on second
stimulus, after sixty seconds, it at once closed. This was fre-
quently verified.

A leaf was acted on as follows :

ist stimulus no effect.

2d

3d

4th

5th

6th

u

u

(t

i(

(<

after eighty seconds,
seventy-five
seventy
sixty

((

((

((

t(

((

((

((

((

((

((

((

slight effect.

leaf half-closed, by move-
ment extending over four-
teen seconds.

leaf closed after movement
of eighteen seconds.

12 Macfarlane. — Contributions to the History of

Repeated experiments were made on the strong green
leaves of the plants mentioned above in the laboratory green-
house of the University of Pennsylvania during October when
the temperature varied from 75° to 83° F., and these always
closed after time intervals of sixty to sixty-five seconds, but
when extended to eighty-five seconds and then reduced, three
to six stimuli could be given, thus :

ist stimulus no effect.

2d " after eighty-five seconds "

3d •' '• eighty " slight effect.

4th " " seventy-five '* slight added effect.

5 th " " seventy " leaf closed.

From tables, like the above, we conclude that in rather
weak plants exposed to considerably lower temperatures than
the normal — /. ^., from 60° to 75^ F., as compared with 80° to
95° F., or 98"" F. in North Carolina — sharp memory of the
first stimulus is retained for from thirty to forty-five seconds ;
but that a decided fall is then experienced, and from fifty-five
to sixty seconds the effect of the first stimulus is greatly lost. A
summation of one or more stimuli, varying in number with the
time-interval between these, is then needed to effect closure.
That in strong plants the effect of the first stimulus is sharply
retained for from fifty to seventy seconds, the variation being
determined by temperature and vegetative strength of the
plants.

But the very important results obtained by Sanderson and
Page, supplemented by those now recorded, prove that exactly
similar summations of stimuli can be recorded in the contrac-
tion of Dioncea leaf as in contraction of muscular tissue, except
that the contracting protoplasmic substance in the former is
of a greatly less specialized quality, and is to a corresponding
degree more convenient and easy for demonstration. The
above observers experimented on leaves by applying stimuli
at intervals of two minutes, and the following table is given
by them to show the result of successive mechanical excita-
tion of the hairs at intervals of two minutes, continued until
the leaf closed :

Dioncea Mnscipulay Ellis.

13

Number of
excitations.

I to 7 .

8 .

9 •
10 .

II

12 .

13 •

14 .

15 •

16 .

17 •

18 .

Angular measure-
ment of effect.

Time in seconds
which elapsed
between con-
tact and the
first perceptible
approach.

o
o
O

19
20

21

22

23

24

25
26

3

2

7

8
10

00
00

GO

10.8

7-3
5.8

5-0
4-5

5-2

2.5

7.6

3.8

3-7

3-3
4.0

2.7
. 2.5
not observed.

2.2

27 at 27th excitation the leaf closed.

" From this experiment, which was repeated several times
and always gave similar results, it was learned : (i) that the
first half-dozen excitations were absolutely without mechanical
effect; (2) that the first effectual excitation was followed by
so slight a movement that if it had not been enlarged by the
lever it would have been imperceptible, and (3) that after this
each successive approach of the lobes in most cases exceeded
its predecessor."

"The time measurements, as will be seen at a glance, stand
in a remarkable relation to the mechanical effects, showing
that the delay between excitations and effects diminishes as
the extent of the effect increases, like facts having the same
meaning, viz., that in the plant, as in certain cases well known
to the animal physiologist, inadequate excitations when re-
peated exercise their influence by what has been termed
summation, i. e., that when any number of such stimulations,
say Uy b, c, dy e, etc., follow each other in succession, the effect

f

iiii

' I

14

Macfarlane. — Contributions to the History of

of each is prepared for and aided by its predecessors ; so that
although, as in the present instance, ^, b, c, d may seem to
produce no effect whatever, each of them really produces a
change in the excited structure, and each contributes when
summed with its predecessors and successors to the bringing
about of the visible effect which follows e. During the re-
mainder of the process the operation of the same law shows
itself in the gradual augmentation of the increments — the
last contraction — that by which the leaf closes, being the re-
sult of the summation of the excitation which immediately
preceded it with all the previous excitations. Our conception
of the nature of the process may be otherwise expressed by
saying, that under the influence of successive excitations the
latent excitability of the leaf gradually increases, for whereas
before it either made no response, or postponed its response
indefinitely, it now answers to the same stimulus by a visible
motion, of which the promptitude and the extent increase
together."

It is noteworthy that when a leaf closes, which has been
twice stimulated, with short time interval between the first
and second shocks, the closing movement is a rapid one,
being limited within two seconds. When a leaf closes which
has received five or six shocks with a pretty wide time in-
terval between each, the period of actual closure movement
of each leaf-half may extend over ten to fifteen seconds, while
one which has been frequently stimulated may even require
forty-five to fifty seconds to effect final closure of half the
movement limit. Now this is what we would expect from a
contractile tissue which has been fatigued by repeated stimuli ;
a reduction of the high degree of contractility which it nor-
mally possesses.

It has hitherto been supposed that only the sensitive hairs
and the triangular area between these are irritable on surface
stimulation, but both outer and inner leaf surfaces exhibit a
marked degree of sensitivity, though greatly inferior to that
of the sensitive hairs. The latter are, therefore, localized
centres of irritability, set apart in an otherwise sensitive or
irritable leaf for receiving external impressions that would be

{

DioncBa Miiscipnla, Ellis. 15

of too delicate a nature to effect movement of the halves as
they now exist. Ready proof of the general irritability of
the leaf can be adduced by scraping or tapping the outer or
inner surfaces ; but the best demonstration can be given by
use of ordinary steel forceps.

It can be predicted, that if one give a sharp but rather
gentle forceps-snip to any part of a leaf-half, there will be no
closure of the leaf; but, according apparently to the strength
of the snip, a more or less pronounced propagation along it
of the rhythmic undulations already referred to. If a second
snip be given within fifty to seventy seconds rapid closure,
within at most ten seconds after stimulus, will ensue. Every
part of each leaf-half may be tested in this manner with
uniform result.

Summation stimuli may similarly be communicated. A
medium sized leaf was chosen and operated on as follows :

First snip stimulus no effect.

Second snip stimulus after ninety seconds "

Third " " " seventy " "

Fourth " " " sixty " very slight closure.

pifth " " " " " slight closure.

Sixth " " " fifty " leaf half closed.

Seventh" " " forty five " leaf closed.

A vigorous leaf behaved thus :

First snip stimulus no effect.

Second snip stimulus, after three minutes "

Third " " "two " "

Fourth " " " " " very slight closure.

Fifth " " " ninety seconds " "

Sixth " " " " " slight closure.

Seventh " " " sixty "

Eighth " " « " « leaf now half closed.

Ninth " " " forty-five" leaf closed.

These and other experiments performed at Edinburgh show
what was more strikingly verified in Philadelphia, that the
effect of forceps-snip stimuli was more powerfully retained
than when irritation— even the most violent— was communi-
cated through the hairs. The extremely slow closing move-
ments of the leaf-halves was equally marked. Thus after the
eighth stimulus in the last experiment, the visible range of
the movement occupied twenty-one seconds.

If lilt

"If

^

i6

Macfarlane. — Contributions to the History of

One hair stimulus followed by a snip stimulus or vice versa
acts equally as if two of either had been applied.

But though at least two stimuli must be communicated
through the hairs, and at least two mild forceps stimuli
through the lamina to effect closure, a single sharp firm
stimulus usually acts similarly. The writer recently experi-
mented on twenty-three leaves attached to plants growing
wild, and eighteen of these closed after first shock when this
was strong. Several of the five that did not close, as well as
many of those that received a mild snip-stimulus, showed a
peculiarity that had repeatedly been noticed during laboratory
experiments. On application of a second stimulus, instead
of almost immediate closure, as is ordinarily the case when
the hairs are acted on, a latent period of from five to nine
seconds elapsed previous to movement, which was then as
rapid as usual. It should be noted that the leaf tissue was
not ruptured.

The mechanical action of water was referred to by Darwin
as follows : ^ " Drops of water, or a thin broken stream falling
from a height on the filaments, did not cause the blades
to close ; though these filaments were afterwards proved to
be highly sensitive ; no doubt as in the case of Drosera^
the plant is indifferent to the heaviest shower of rain."'
" Although drops of water, and of a moderately-strong solu-
tion of sugar, falling on the filaments, does not excite them,
yet the immersion of a leaf in pure water sometimes caused
the lobes to close." We have repeatedly experimented with
a thin but sharp jet of water, and the leaves have invariably
closed after a short exposure to it. But the behavior of
leaves that are suddenly immersed in water, at a temperature
like the surrounding atmosphere of from 60° to 80° F., is
more puzzling as Darwin found. Many observations that the
writer has made point to the conclusion that if a leaf attached
to a plant is suddenly immersed with the upper surface next
the water, no change ensues, but if the process is repeated the
leaf closes, /. ^., that the two mechanical stimuli, given by the

1 Insectivorous Plants, p. sQt.
*p. 292.

' ■^•..^C*;^^

MUTILATED TEXT

DioncBu Muscipulay Ellis.

17

water, act like the two shocks communicated through the
hairs or by forceps snip or otherwise. But while this is indi-
cated by twenty-one out of twenty-nine leaves, it must still
be emphasized — as showing the need for continued experi-
ment— that leaves have repeatedly closed on first immersion
in water at, or very near, the same temperature as the sur-
rounding air. Still more puzzling is the effect of gradual
immersion in water. Round Wilmington, N. C, the writer
dug up thirteen plants in succession, each surrounded by large
balls of- soil. He carefully and slowly immersed these, in
natural position, under water. Every leaf closed on all of the
plants, with the exception of two which did so when the
plants were gently removed from the water. This behavior
led him to attempt to localize the centre of movement more
exactly. Selecting twelve fine leaves, and holding them by
the petiole, he slowly immersed these from the tip of the
blade inwards, and invariably when the water reached the
level of the first pair of opposite hairs rapid closure followed
Some of these were repeatedly withdrawn when the water had
reached to about one-third of a line or more from the hairs,
but no change followed. Seventeen leaves were then gradu-
ally immersed obliquely, in a vessel, so that the water would
first touch a single hair. Every leaf closed when the water
wetted the hair. We thus learn that while rain drops or a
slight water current falling on the upper leaf surface is with-
out etfect, a sharp water impact, or immersion so as to cover
one or more hairs, starts contraction. But while water impact
is purely mechanical, a totally different explanation is needed
in the latter case. We know from Sanderson's electrical
researches that the upper leaf surface is positive to the under,
and that the hair region is specially sensitive in its electrical

condition, so that whe
might act as a con
and lower surfac
by immersing in
after repeated dippi^
for when irritated af^^
however, set up rapi<

T

leaf was immersed, the water
e electricity between the upper
Uids act differently was proved
^^n no change occurred even
, however, was still sensitive,
^ it closed. Petroleum oil,
*^- At the present stage we

«

P

Mil

I

' I

I

'S.Aki.

'.<'■'

i8

^^acfarlanc.—Cont}^bntio7lS to the History of

cannot attempt to decide what meaning attaches to these
phenomena, unless on the lines of Sanderson's work.

The above observations, coupled with Sanderson's state-
ment, that electrical disturbance is first "appreciable by
the electrometer at the external surface at about one-sixth
of a second after mechanical excitation," and that "the
excursion attains its maximum in one second, and that its
return occupies about the same time," having satisfied us tha't
propagation of a mechanical stimulus is comparatively slow, we
determined to try whether two stimuli, very rapidly applied,
with a time interval of about one-fourth second or less, might
not be propagated through the protoplasm as a practically con-
tinuous impulse. Two rapid touches, with a time interval of
about the fourth of a second, were given to a hair without
causing visible result other than rhythmic movement of the
halves. A third touch then effected closure. The above
has been frequently verified with uniform results. Similarly
two forceps-snips, applied very rapidly in succession, required
to be followed by a third to effect closure. If, however,
the interval between the first and second shocks be expanded
to a third of a second or more, the two shocks are separately
propagated, so that the leaf closes. The above phenomena
in contraction of Dioncea tissues agree with those of animal
muscle where a definite strength of stimulus must act for a
certain time. There has not been opportunity as yet for using
an apparatus which would give three, four, or more stimuli
within the fourth of a second, but there is strong probability
that such could be applied previous to another stimulus that
would cause closure. It is evident, however, that three dis-
tinct stimuli can be given previous to leaf movement. It
has not as yet been determined whether the double stimulus,
propagated as a single wave, caa-t^'r^byshorten the duration
of a summation series when wiik|l ' tafcvals are allowed to

intervene. iB^

We can now refer to what rnpy^l^Vv'' r^?
and given rise to the stat^?4il l»^is,t
to cause closure. If a h|tl 'IWM"fi:^H «^ m a rather slow and
deliberate manner, the leaJ^I^^.Vt Mi5 it must be remembered

observers hitherto,
le touch is sufficient

V

Dioncea Mnscipiila, Ellis.

19

that even in the hand of a steady operator such a stimulus
really resolves itself into a series of two or more delicate
stimuli applied in succession.

The leaf of Dioncea then is truly sensitive throughout its
halves to mechanical stimulation, but the capability of receiv-
ing sensation impulses is highly concentrated in the hairs.
These are so modified for insect catching that the first touch
fs without visible effect, but prepares the leaf by summation
action for a second stimulus which, when applied, causes
rapid closure. The advantage of this preparatory warning to
the plant is evident, for any adventitious particles blown
against it will not cause useless movement. Further, like
animal muscle, the contractile tissue of Dion(Ea must have a
mechanical stimulus of a certain strength prolonged through
a certain time to effect contraction ; while a prolonged sum-
mation series can be applied, the extent of the series varying
according to the time interval between each stimulus. These
experiments further prove that a minimal contraction stimulus
cannot be propagated through the leaf by one touch of a
hair, no matter how sharp and powerful the touch is. In
other words, the hair mechanism is such in relation to the
contractile protoplasm that the latter is not sufficiently
altered by one blow, and the minimal amount of alternation
necessary for closure can only be obtained after the second
stimulus.

We would here emphasize, however, the considerable diffi-
culty one experiences in attempting to place a piece of meat
or other solid on either half of a blade without due caution.
Very often two or more touches are unconsciously given to
the hairs.

As has been often pointed out, when a leaf is closed by
mechanical irritation of the hairs, the halves interlock
loosely at first by partial crossing of the marginal bristles,
and then slowly firm up till the bristle-bases press against
each other. But when an insect or some nitrogenous matter
is enclosed, there is a gradual tightening, and the leaf margins
recurve, owing to tense pressure of the halves, against
a glabrous non-glandular or slightly glandular area that runs

^'ll!

I

I

;:>,iiS«f«'«SK-,..>t.-<»s(flBo».»»8-

20 Macfarlane. — Contributions to the History of

round each margin about one-sixteenth of an inch from its
edge. By repeated mechanical irritation of the hairs after
the leaf had closed, the same reflexing movement of the
margins has occurred. Instructive results were got from a
lot of plants that the writer expressed from Wilmington.
Most of them during transit suffered severe concussions
though the leaf substance was not ruptured. They were at
once planted out, but seven-eighths of the leaves had shut
tightly, and their margins were reflexed, though nothing was
enclosed. A very copious viscid acid secretion, moreover,
had been poured out. Other plants, whose roots had been
washed clean of earth, had been placed loosely in a large
preserve bottle, and at least half of the leaves were in the
same condition as the last, due also to knocking against each
other in transit. The above, coupled with similar behavior
under chemical and electrical stimulus, points to the belief
that complete tightening of the leaf is of a tetanic nature,
and can only be accomplished by repeated stimulus, either of
a mechanical, chemical, electrical or other kind.

Leaves that were isolated from plants in an atmosphere at
76° F.were found to retain their normal irritability from twenty
to thirty minutes. Thereafter they became flaccid, and about
one hour after removal did not respond to stimulus. Whole
plants, from whose roots the soil had been washed away,
showed irritability in their leaves for one and a half to two
hours, according to the size, vigor, and root development of
each. Entire plants, that had been washed clean of soil and
kept in a moist, tightly-sealed glass vessel, were highly irrit-
able on removal after six days.

Leaf Closure.

(U) By Heat Stinuili.

My experiments on the effect of dry heat have not been so

extensive as to permit expression of a definite opinion, but

the action of water, at different temperatures, has been tried

on twenty-seven leaves.

Darwin says : " A leaf was cut off and suddenly plunged
perpendicularly into boiling water. I expected that the lobes

¥

!

Dioncea Muscipida^ Ellis.

21

would have closed, but instead of doing so they diverged a
little. I then took another fine leaf with the lobes standing *
at an angle of nearly 80° to each other ; and on immersing it
as before, the angle suddenly increased to 90°. A third leaf
was torpid from having recently re-expanded after having
caught a fly, so that repeated touches of the filaments caused
not the least movements ; nevertheless, when similarly
immersed, the leaves separated a little. As these leaves
were inserted perpendicularly into the boiling water, both
surfaces and the filaments must have been equally affected,
and I can understand the divergence of the lobes only by
supposing that the cells on the lower side, owing to their
state of tension, acted mechanically, and this suddenly drew
the lobes a little apart, as soon as the cells on the upper
surface were killed and lost their contractile power." One
may best lead up to the above by considering the effect of
gradually increasing temperature. All of the experiments
described below were made on leaves attached to living
plants.

Water was heated to 50° C, and a dropping pipette was
similarly heated by immersion in the water. Three drops
were then let fall on an open leaf, but produced no effect after
ten seconds. A second and then a third application was made,
when the leaf closed. Four other leaves behaved thus, but
a sixth required four applications. Naturally, the water on
exposure to the atmosphere and colder leaf surface, would
lose some of its heating power, so that we may regard the
actual temperature for stimulation as having ranged from
45° to 48° C.

Water at 58° C. produced no movement after ten seconds,
but on a second application four leaves closed after an
interval of five to eight seconds. At 65° C. six leaves were
tried, and two of these closed, one after seven seconds, the
other after nine, while the remaining four closed at once on a
second application. At 75° C. five leaves were once treated,
and all closed after intervals of six to nine seconds.

Boiling water was then used on the remaining ones, but as
regards its action on the leaf substance, it should first be

%

I

I

22

Macfarlane. — Contributions to the History of

stated that Darwin's observations, as quoted above, had been
verified by the writer some months before, and that com-
parison of the minute structure of the leaves then used with
fresh leaves, and others preserved in various media had con-
vinced him that immersion for even a second or two in boiling
water causes coagulation of the protoplasmic substance and
disorganization of the starch that is so abundantly present in
many of the leaf cells. With one exception all of the leaves
closed within from two to five seconds after application of
the first few drops. The exception proved to be an interesting
one, and such as one might scarcely have hoped to obtain.
The leaf was rather small— about half the usual size — and
like the others, began to close shortly after a few drops of
the boiling water had been let fall on the upper surface, but
just as it had half closed, movement stopped for an instant,
and it then slightly relaxed to retain the heat-stiffened position
for the remaining half hour, during which the plants were
under observation.

As regards after-effects the last mentioned leaf was the
only one which showed speedy and pronounced death changes,
for next day it had a yellow flaccid aspect and in three days
was dry, brown, and shrivelled. The other leaves of the same
series retained their green appearance for nearly a week,
except for an area about one-eighth to one-quarter inch across,
marking where the water had been first applied and which soon
became yellow and then brown. All of these leaves gradu-
ally became yellow, and in a fortnight were dead. Four
of those acted on at from 50° to 75° C re-expanded, but as the
individual leaves had not been marked in relation to water
temperature we are un^le to say which persisted.

The above facts prove that a gradual increase in tempera-
ture produces quickened stimulation up to a point where the
contractile protoplasm and food materials are so affected that
disorganization and death ensue. But the subsequent fate of
most of the leaves points to a permanent injury, either local or
general, to the living cell contents.

Six leaves were selected on two plants standing in a room
with a temperature at 72° F. Small pieces of ice were care-

\

DioncBa Muscipulay Ellis.

23

fully placed on each; one closed in sixteen seconds, another
in twenty-five, a third in forty-two, and a fourth in ninety
seconds, by which time the ice had melted. The remaining
two did not close. These results were so diverse that little
value could be attached to them, and the experiments were
made in January, when the plants were at their worst. But,
recently, in the natural haunts of the plant, the author was
able to prove that **cold " stimulus is powerful in its action.
On an afternoon, with the temperature at 79° in the shade,
small pieces of ice were placed on twenty-three leaves. Care
was taken to place these on the base or apex of the lamina,
so that even the chance of wetting the hair bases might be
avoided. Nineteen of the leaves closed sharply in from five
to eleven seconds ; three closed after sixteen to twenty-one
seconds, and one closed after thirty-six seconds, by which
time the ice had just melted. Small drops of ice-cold water
were placed in similar positions, and these acted like the ice.

Leaf Closure.

{c) By Light Stimuli.

Darwin states' that concentrated light-rays are unsatisfac-
tory in action, and the writer's experiences corroborate this
so far as investigation has gone. A large leaf had the sun's
rays concentrated on it in a glass house with a temperature
at c^"" F. After three and a quarter minutes the leaf very
slowly closed to about one-third of its extent, and about half
a minute later it suddenly closed completely. The leaf had
expanded when visited two days after, but where exposed to
the light a circular area of a yellowish color and thin in
texture showed that the leaf substance in that region had
been destroyed. Another leaf similarly treated, but with the
light more widely diffused over the lamina, closed to about
one-third of its extent, and it after expansion showed no
burning effect. Three others were tried but gave negative
results. Many careful studies must therefore be made before
definite conclusions can be reached as to light stimulus.

> p. 294-

t

i

'

24

Macfarlane. — Contributions to the History of

Leap^ Closure.

(d^ By Chemical Stimuli.

The consideration of this opens a very wide field, the
border of which only we are persuaded has as yet been
touched. All who have looked into the subject from Curtis
and Canby onwards have noted the great differences in action
which different substances possess. But Darwin and Dr.
T. A. G. Balfour are the two writers to whom we are indebted
for the most extended observations. I have repeated nearly
all of their experiments, and have added others which sug-
gested themselves as likely to extend the line of work. Gar-
diner's suggestive results have also been of use.

Balfour found that chloroform, chloride of ammonium,
carbonate, sulphate, and borate of soda, sulphate of copper,
meat, and various nitrogenous food stuffs, also pepper, all
caused closure, though he does not state the strength of any
of the solutions used. Darwin found that a moderately
strong solution of sugar, chloroform, ether and nitrogenous
compounds caused closure. Chloride of strontium and sul-
phate of iron, however, were without effect, but killed the leaf,
according to Balfour ; while according to Darwin hydro-
cyanic acid paralyzed two leaves for a time though they
continued to remain open.

Though somewhat arbitrary in method substances will be
treated of in the order that they appear to .stimulate the leaf.
At 9.30 A.M. on a clear day, three minute shreds of roasted
meat were wetted and very carefully laid on as many leaves in
the axil of a hair with the leaf surface. None of the leaves
closed. One of them was then twice touched, when it closed
and remained so for eleven days and poured out an abundant
secretion. One was slightly closed at 4 p.m., the third remained
unaffected. Next day at 9.15 the second leaf had closed, the
latter was still open. The former remained shut for eight
days, the latter never closed, and its meat particle was finally
washed off in watering. Three leaves were chosen at 9 15
on another day, and on one a minute shred of meat was
placed on the outer end. It showed no change at 5 p.m., but

DioncEa Miisciptda^ Ellis.

25

at 9.10 next day the end part of the half on which it was had
slightly inflected, exactly as in a like case recorded by Dar-
win (p. 297). Secretion had gone on in the region next to it,
and gradual digestion and absorption proceeded for five days
till scarcely a trace of it remained. On the other two leaves
pieces about three times as large were placed, and both had
closed by 4.30 p.m., and remained thus till digestion was
accomplished. But wetted pieces of meat have repeatedly
been placed on leaves, and have failed to set up closure, while
on three occasions, leaves with meat, that were artificially
closed, did not tighten up, but continued to show the digesting
meat between the bristles.

Meat, therefore, we view as a decidedly weak stimulant,
and it appears likely that were a careful and exhaustive set of
experiments conducted with shreds of varying size a toler-
ably close approximation as to the amount of a given kind of
meat necessary to effect complete closure could be arrived at.

Mainly with the object in view of trying to get an unaltered
leaf for microscopic study, one attached to a living plant was
dipped into a one per cent, solution of chromic acid and re-
mained unaltered for ten minutes. It was then carefully cut
off and dropped into the liquid, where it showed no sign of
closing for two hours ; when looked at, however, after two
and a half hours it had closed. This behavior seemed so re-
markable that the experiment was repeated on several occa-
sions, and with the same result ; but the time between
immersion and closure varied from one to two and a half
hours. But the writer had forgotten at the time the behavior
of some tadpoles which he placed a few years ago in the same
strength of solution under the supposition that they would
speedily die. On returning to the jar about one and a half
hours after he was horrified to find the animals still alive and
wriggling about. Several gentlemen have had like experi-
ences.

A leaf was immersed in a one per cent, solution of chromic
acid, and after five minutes cut off and allowed to lie in the
solution for thirty minutes. It was then passed into a twenty-
five per cent, alcohol solution and at stages lasting over an

I

'.^ I

I lili

26

Macfarlane. — Contributions to the History of

hour into increasing strengths till forty-five per cent, was
reached, when it closed.

Glycerine like strong sugar solution causes contraction
and like it in a rather slow manner. Strong solutions of
ammonium carbonate and citrate stimulate to closure in
from twenty to forty seconds, while dilute sulphuric, hydro-
chloric, and nitric acids act in from two to ten seconds.
Three minute crystals of solid chromic acid were laid on a leaf
and left for a minute without producing movement. They
were then wetted by a minute drop of water, when closure
occurred after eighteen seconds. A crystal about half as
large as a pin's head was then arranged and wetted, when the
leaf closed in five seconds. Minute fragments of potash and
soda stick were arranged and dissolved, when the leaves
operated on closed in from six to thirteen seconds.

Absolute alcohol, ether and chloroform are all rapid stimu-
lants if dropped on in the liquid state, but the vapor acts
according to its amount, contracting the leaf slowly but com-
pletely if abundant; contracting it only partially if less so,
and rendering it insensible and powerless if the quantity be
still less. But of the substances tried corrosive sublimate
and one per cent, osmic acid—notably the latter — are the
most powerful stimulants, and a leaf responds to their pres-
ence in one to three seconds.

Different substances, therefore, stimulate to very different
degrees, and even the same stimulant can cause closure in a
time ratio that is proportionate to the strength or concentra-
tion of it. These and other substances not mentioned above
are exactly comparable in their action to that on muscular
tissue. Now, in the case of many of the above agents there
was ample time for osmotic action to be set up. But while
some set up endosmotic flow, others, such as sugar solution and
glycerine, would cause vigorous exosmotic flow. Though we
may not be able as yet to explain fully their action on the con-
tractile protoplasm, it seems to us that the phenomena here
are identical with those of muscular tissue, and that we have to
deal in the leaf substance with an organized material iden-
tical in its fundamental behavior with muscle, though greatly
less sensitive in its response to stimuli.

♦

Dioncea Muscipulay Ellis,

Leaf Closure.

n

{e) By Electrical Stinuili,

After the elaborate researches of Burdon Sanderson, it is
not necessary that we should do more than refer to the fact
that closure follows application of this form of energy. We
have not yet determined accurately whether two sharp and
distinct stimuli are needed or whether one suffices, but there
are grounds for believing the former to be true. In all cases
the electrical terminals were delicately applied to the lower
external parts of the leaf halves, after moistening of the sur-
faces, and closure has invariably been effected. It seems
difficult to determine, from Sanderson's account, whether he,
in all cases, inserted the terminals into the leaf substance, or
only in connection with certain experiments, but closure by
electrical stimulus from the external surface is another proof
of the general sensitiveness of the leaf.

II. Leaf Structure.

Oudemans, T. A. G. Balfour, Darwin, De Candolle, Frau-
stadt, Kurtz, Batalin, Sanderson, Gardiner and Goebel have
all described the histology of the leaf, but it is curious, indeed,
that the descriptions and figures of the great centres of irri-
tation— the hairs — are of the most general and imperfect
nature. Oudemans, De Candolle, Fraustadt and Kurtz have
given more attention to the general histology and develop-
ment of the leaf. The two last are at variance as to the
stomata on the marginal bristles, the former stating that
they are present, the latter that they are absent. They are
present in small numbers along the lower or external faces,
but absent on the internal or upper, as might be expected.
The brown stellate hairs referred to and figured by Oudemans,
and noted by his successors, occur on the upper and lower
surfaces of the expanded petiole, but only on the under
surface of the blade. We would point out here that their
structure and development essentially agree with those of
the secreting glands on the upper leaf lamina. They fur-
ther exhibit considerable variability in the mature state, and

I

28

Macfarlane, — Contributions to the History of

many of the deviation types approach the typical glands in
showing fusion of the radiating hair processes throughout the
greater part of their length. (Plate IV, Figs. 5<^-5^.) Darwin
experimented to ascertain whether these would show aggre-
gation of the protoplasm, but got negative results. He insti-
tutes no comparison between them and the glands, however.

The glands consist, in all cases, of two elongated basal cells,
with their long axes placed parallel to the course of the veins
(Plate IV, Fig. 7), and these are surmounted by two tiers, the
lowermost of two cells, the upper of a considerable number.
The last is covered by the surface cell layer of the gland, the
appearance of which has often been figured. The illustration
of a gland given in side view on Plate IV, Fig. 6, conveys a
slight idea of the beautiful intercellular protoplasmic con-
nections that pass through the pores in the thickened trans-
verse partitions of the lower cell tiers. Similar connections
with the surface cell layer have not as yet been traced. Gar-
diner has fully described the position and relation of the
nucleus in the surface cells before and after secretion has
commenced. Considerable discussion has taken place on the
subject of cell vacuoles and their mode of origin. Gardiner
states' that '* in each cell the protoplasm closely surrounds
the cell-wall, leaving one large central vacuole filled with the
usually pink cell sap." This is true of most leaves, but it is
not difficult to find healthy expanded leaves whose surface
gland cells enclose two to five vacuoles of varying size. Such
leaves, however, are mostly of small size and of a green color,
but are irritable and secrete as usual.

The irritable Jiairs are disposed in threes on each half
of the blade, but Errera'^ has seen four or five, and a leaf
that came under the writer's observation during 1891 had
seven on one half and six on the other, and these were
arranged in an irregular manner over each half. During a
day's hunt even for Dionaa^ one often encounters leaves
with eight to thirteen hairs. Such facts give countenance
to the view that the sensitive hairs were once more numer-

' Proc. Roy. Soc, Vol. 36, p. 180.

2 Bull. Soc. Roy. de Bot de Belgique, xviii, pt. 2, p. 53.

Dioncea Mtisciptila^ Ellis.

29

' \

ous and diffuse in distribution, a condition still retained
by Droscra.

Each hair is an emergency, and consists of three well-
marked regions, the base^ the joint and the shaft} The hair
base consists externally of four to five tiers of epidermal cells
that gradually rise up from the leaf blade. Each tier is a cyl-
inder of eighteen to twenty-two cells with thickened walls that
are traversed by intercellular threads of protoplasm. These
enclose loose and slightly elongated cells with rather thick,
clear walls, the cells being continuous with those of the meso-
phyll. (Plate IV, Fig. 3.) The protoplasmic masses of all of
these are connected by threads with each other, and with
those of the mesophyll cells. The joijit or special irritable
centre is remarkable. It is a cylinder of elongated quadrangu-
lar epidermal cells, each three and a half to four times longer
than broad. These enclose a central cylinder of mesophyll
cells that are similarly elongated. In all hairs yet examined
— and this applies to twenty-three — the cuticle that is strongly
developed over the general epidermis gradually thins out over
the basal cells, and is either quite absent over the irritable
joint cells or so very delicate as to escape detection when
acted upon by cuticular tests. The middle part of each epi-
dermal joint cell is, in all hairs yet examined, creased or puck-
ered upon itself, so that it at first gives the impression of trans-
verse dividing walls on surface view. (Plate IV, Fig. i.) These
creasings have evidently misled observers into giving imper-
fect views of the hairs. The question naturally arises whether
these exist in the unstimulated hair or are due to collapse of
the cells, which, up to period of stimulation, are turgid. One
would regard the latter as the more likely view, but so far as
we have been able to bring the microscope to bear on open
leaves, the joint cells appear always to be puckered. But a
structural feature of some interest is the presence over the
free face of each cell of minute pits which seem exactly to
agree with those noticed by Gardiner' on the terminal hair

1 We are unable to agree with Goebel (Bot. Schild. ii, 1891), in dividing the hair into two
parts, since the relation, structure and behavior of the three areas that we have indicated
prove them to be distinct aUke in structure and function.

2 Proc. Roy. Soc, Vol. XXXIX, p. 229.

30

Macfarlanc. — Coiitributions to tJie History of

cells of Drosera. His account leads one to believe that he
regarded them as closed pits. Sections of Dioncea hair (Plate
IV, Fig. 2) show that the free face of each cell is slightly thick-
ened, but that where the pit occurs the internal thickening is
absent. We cannot as yet say whether each has a pore aper-
ture or is a closed membrane, but the knowledge we now have
of intercellular connections suggests the former as a likely
condition.

One naturally desires to know the use of the minute pits.
In proposing an hypothesis we would refer to some of the
views that have been advanced to account for leaf closure and
subsequent re-expansion. Darwin suggested ** that the sev-
eral layers of cells that form the lower leaf surface are always
in a state of tension, and that it is owing to this mechanical
state, aided probably by fresh fluid being attracted into the
cells that the lobes begin to separate or expand as soon as the
contraction of the upper surface diminishes." Similarly, Bata-
lin* regards both opening and closing of the leaf as due to a
migration of liquids from one zone to another, and considers
that the cells of the lower surface are always extremely tur-
gid, but that in the expanded state those of the upper surface
are even more tensely distended than the former, and he
agrees with Darwin that measurements made before and after
closure prov^e that contraction of the upper side takes place
after stimulation, and he further regards the shutting as a
result of disturbance of the tension equilibrium through irrita-
tion. In the setting up of this disturbance, he suggests two
distinct causes : either {a) an active contraction of the plas-
matic substance on the inner, and passive expansion of that
on the outer surface, /.r., molecular translocation, or (^), dis-
turbance of equilibrium in the tension of the tissues result-
ing from expression of water through the walls. He entirely
favors the latter or mechanical view, without seeming to
think that a combination of the two hypotheses might explain
matters.

Recognizing the difficulty of explaining such results as
those of Sanderson unless the action of the living protoplasm

1 " Flora,' 1877, Nos. 3-10.

DioncBa Muscipiilay Ellis.

31

be taken account of, Sachs concluded ' " that the condition
of turgescence of the cells depends upon the protoplasmic
utricle opposing the expulsion of the endosmotically absorbed
water, even under high pressure," and he follows Sanderson'
in expressing the opinion that "the extensibility of the
cellulose walls plays an important part " in effecting con-
traction after the permeable protoplasmic utricle has per-
mitted an escape of liquid. Gardiner's experiments further
prove that plasmolysis is the important factor, and the follow-
ing statement of his sums up the position : " From certain
observations on Dioncea and Mimosa the author is led to
believe that there also movement is made possible by the
establishment of sudden and different conditions of turgidity
of different cells, such differences being occasioned by the
induced porosity of the protoplasm of certain of these cells.
These phenomena occur perhaps in all cases of movement."

But the water that escapes after stimulation from the cells
on or near the upper leaf surface must be transferred to some
other region, and the amount necessary to be transferred need
not be great. The presence, in large leaves especially, of
loose intercellular passages between every pair of bundles
suggests that liquid may escape into these. But we would
suggest for further consideration the possibility that the pores
on the surface of the irritable joint cells are open, and that
through these the protoplasm can rapidly eject, or allow the
passage of, minute quantities of liquid sufficient to disturb
seriously the equilibrium ; since we have indicated that the
protoplasmic masses of the joint cells are connected by inter-
cellular threads with those of the base, as are the latter again
with the epidermal and gland cells. It might be imagined
that minute drops escaping could readily be seen, but sev-
eral difficulties stand in the way. . In attempting to bring
the objective of the microscope into focus on the joint cells
the leaf as a rule closes ; failing this the shining cell sur-
faces reflect light so much that excreted liquid would read-
ily be overlooked. The experiments performed by Darwin

1 Physiology of Plants, Etig. ed., 1887, p. 653.

2 Roy. Instit. Lectures, " Nature," Vol. XXVI, p. 356 et seq.

1*

32

Macfarlanc. — Contributions to the History of

of removing the hairs do not aid us here, as he does not
state whether the leaf that had opened after removal of all
the hairs was still irritable in the areas from which they had
been removed. But even were such true — and Balfour's ex-
periments favor it — it is still possible that minute drops of
liquid might escape through pore apertures in the cut cell
walls, just as we suppose that these escape from the pores on
the unthickened areas.

The lower part of the shaft consists of three tiers of shal-
low, closely-packed, cells, succeeded by another tier, the cells
of which are more elongated and have their upper septa
more or less obliquely placed, and wedged in with the lower
ends of the shaft cells above. The terminal shaft cells are
elongated, thick-walled, and taper into each other. Their
walls are traversed by numerous pore canals, and their cavi-
ties are filled with finely granular protoplasm, but we have not
succeeded in tracing intercellular connecting threads. The
internal or central cells of the shaft greatly resemble those of
the epidermis in size, shape, and structure.

The epidermal cells of the upper leaf surface are elongated
and nearly quadrangular in outline, and are covered by a thick
cuticle that is in proportion to the cuticle of the lower leaf
surface as 3 : 2.

Of all the plants examined by the author for intercellular
protoplasmic connections Dioncca yields the finest results in
its epidermal cells. Such are best attained, however, by a
modification of the ordinary mode of treatment that was grad-
ually arrived at during the course of the present inquiry, and
is described in the footnote.^ One readily notices then (Plate
IV, Fig. 8) along each side wall eighteen to thirty proto-
plasmic bridges which are slightly constricted on either

side of the cellulose wall, and form a central swelling at

»

^ The following has been found invariably to give better results than the methods rec-
ommended by Gardiner, Keinitz-Gerloff, and others. After iodine treatment the fresh
sections are placed in twenty-five i)er cent, sulphuric acid, and left for one to two hours.
They are then removed thrown into water, thoroughly washed in changes of it, and there-
after stained in a strong solution of watery eosin for at least an hour. Rapid washing
in water then removes the stain from the swollen walls, and brings out sharply the cell
protoplasm and threads of a rich crimson-red color. These show very clearly if mounted
in a cell with two per cent, glacial acetic solution which fixes the stain.

Dioncea Musciptilay Ellis.

33

the passage through the pore aperture (Plate IV, Figs. 9, 9^).
The transverse or oblique walls are traversed by five to
eight similar processes, so that the protoplasm of each epider-
mal cell is linked to that of neighbor cells by fifty to seventy-
five fine connecting threads, and these again collectively
are united with the cylinder of sensitive cells in the irri-
table hairs.

But the lower or internal wall surfaces of the epidermal
cells have a clear shining white aspect due to colloid modifi-
cation (Plate IV, Figs. 6, 7). After the most careful treat-
ment and study, we have failed to trace a single process
traversing these. On the other hand, where the lowest pair of
cells of each gland unites with the sub-epidermal cells their
walls are thin and traversed by protoplasmic threads. Gar-
diner has already mentioned ^ the occurrence of connecting
threads in the mesophyll, and we have succeeded in tracing
these throughout most of the cells; the threads are less
abundant, however, than are those of the epidermal cells.

Taken as a whole, then, the irritable hair cells communicate
by their epidermal and mesophyll portions with the leaf epi-
dermis and mesophyll generally, while the mesophyll cells of
the leaf substance appear to be cut off from the epidermis
by a thickened wall, but communicate directly with the gland
cells. The advantage of this is probably considerable even
from a mechanical standpoint, if we remember the amount of
tension to which the surface cells are exposed under the vary-
ing conditions of expansion and contraction of the leaf, as
well as the secretion of the digestive liquid.

Several of the observers already mentioned have described
the structure of transverse sections, but they have overlooked
points that appear to be of considerable importance. When
the leaf is open and undisturbed, the upper and under epider-
mal and subjacent three cell-layers of the mesophyll contain
chloroplasts and large starch granules. But further, little
patches of chlorophyll cells unite these with the bundles or
are irregularly disposed in patches amongst the clear meso-
phyll cells, and all of them contain starch. The bast cells of

1 Proc. Roy. See., Vol XXX VI, p. 181.
3

IN

V.

^

34

Macfarlane. — Contributions to the History of

the bundles also show granules though of smaller size. Large
quantities of starch likewise lie in the cells that make up the
triangular area between the midrib bundle and its side
branches. Very few granules occur in the cells along the
sides of the bundle, and practically none in those beneath it.
This relation the author has found to remain unchanged in
non-secreting leaves even after these have been kept in the
dark for several days. If now serial sections be made from
the base of the blade down through the narrowed process
connecting blade and winged petiole, the transition from the
former to the latter is sharply marked by absence of stored
starch around the bundle. As stated below, we believe that
the starch is largely utilized during contraction and secretion,
for then it is replaced in the bast cells by an oil and at the same
time it increases greatly in amount in the upper epidermal cells,
probably due to transference of the oil to the epidermal cells,
and temporary storing of it previous to excretion through the
glands. It is specially worthy of note that while surface
pieces of epidermis become deep blue from the amount of
starch they contain, not a trace of starch can at any time be
detected in the gland cells.

As regards the starting of the closing movement, and
the mechanism that effects closure, various views have
been advanced. It is generally conceded that after
stimulation the protoplasm becomes permeable to the out-
ward flow of water, but there are grounds for believ-
ing that it is penetrated by minute pores through which
the migrating water can escape. It is quite possible
that these might exist, though our microscopes or methods
of manipulation might be such as to fail in demonstrating
them. But in examining various leaf sections very minutely
under a one-ninth objective, several of the clear cells
were noticed in which the protoplasm showed an evident but
extremely fine striation at right angles to the leaf surface.
These cells were chiefly noticed in the third and fourth layers
beneath the upper epidermis. One must be careful not to
confound them with very similar appearances that are shown
by the cell walls when these are rather strongly illuminated,

Dioficea Miiscipula^ Ellis.

35

for then the membrane breaks up light in parallel waves that
look greatly like the condition of the protoplasm now described.
Examination of material preserved in absolute alcohol and
stained in eosin caused me to express the opinion, at the 1891
meeting of the American Association of Science, that the proto-
plasm had delicate transverse striations,and this was also stated
in abstract in the " Gardeners Chronicle^' (Oct., 1891). Finer
preparations, and the use of higher powers and additional re-
agents incline us to the opinion that it is due to rows of ex-
tremely minute globules or pores in the protoplasm. Each
globule or pore is less deeply stained by aniline dyes, such as
eosin, heliosin, and methyline blue, than is the intermediate sub-
stance ; iodine solution gives to them a pale bluish-yellow as-
pect. We are not prepared to say whether these are pores or
liquid globules, but the optical appearance they give to the cells
coupled with the movements of the leaf suggest possible cor-
relation with the structure of striped muscle. The discovery
by Haycraft that many of the optical phenomena of striped
muscle are due to puckering of its surface does not militate
against this, for the coexistence of puckerings along with pro-
toplasmic pores or globules is not improbable.

The nucleus of the epidermal and mesophyll cells of the
blade is mostly of a fusiform shape, but by the action of swell-
ing agents it at times assumes an oval or spherical outline.
Each is bounded by a clear, highly refractive nuclear mem-
brane, from which processes radiate out chiefly from the poles,
but occasionally also from the sides. Sections of leaves that
have been hardened in chromic acid and alcohol, and stained
with strong solution of eosin show these processes stained,
like the nuclear membrane, of a deep refractive pink hue.
As they run through the protoplasmic utricle they divide up
and are connected with the chloroplasts, some at least seem-
ing to terminate in these. DioncEU thus presents the same
relation of the nuclear threads with the starch centres that is
shown by Spirogyra. The nucleolus is a small spherical, highly
refractive body, lying inside each nucleus. Rarely there are
two nucleoli. Each nucleolus encloses a minute but very
sharply defined endonucleolus. Numerous minute leucoplasts

36

Macfarlana. — Contributions to the History of

exist also in the protoplasmic utricle, and their function is
probably explained by the appearance of large quantities of
starch during digestive secretion.

Gardiner's " rhabdoid " has been demonstrated with great
clearness by directly treating surface sections with strong
solution of watery eosin. They should then be fixed and
examined in acetic solution. No evidence has been obtained
to support Gardiner's statement that the rhabdoid decreases
during secretion. A series of measurements indicate that it
remains the same, or slightly increases in size, but detailed
results will be given in a future communication.

III. Leaf Secretion.

Hitherto all observers have agreed in stating that if the
leaf shuts through artificial mechanical stimulus, or owing to
the irritation of the hairs by a dry body or inorganic sub-
stance, what we may call ** non-tetanic," closure ensues; that
is in such cases the marginal bristles intercross more or less,
but do not subsequently become everted by reflexion of the
leaf margin ; and that no secretion is poured out unless nitro-
genous matter is present. As regards the first point we have
already shown that it is wholly due to non-continuance of
stimulus, and that prolonged irritation does bring about re-
flexion of the leaf margins. But as it seemed possible that
the leaf secretion might correspond in the vegetable kingdom
to what is known as the waste metabolic material of animal
muscle, numerous experiments were arranged to ascertain
whether continued or intermittent stimuli might not cause
the secretion to flow. Fresh vigorous leaves on several
plants were carefully washed with water and left to dry for a
day. Small glass beads, fragments of quartz and fragments
of pot-crock were then laid on the leaves which were made to
close. A few minutes later the hairs were repeatedly irritated
by insertion of a blunt needle, and on examination an hour
after, the leaves were found to be tightly closed. They were
then restimulated, and the action was repeated every two
hours for sixteen hours. In from eleven to fifteen hours
after closure the gland surfaces were moist, and after twenty
hours were secreting freely.

DioncBa Muscipiilaj Ellis.

37

The secretion had all the chemical, physical, and optical
characteristics of that poured out round a nitrogenous body.
It at once gave a decided acid coloration to litmus test paper,
had a thick mucous consistence, so that it could be drawn out
into threads, and when a little was placed on a slide and
treated with alcohol it coagulated and assumed under the
microscope a delicate myxoid or amoeboid granular a^e jlation.

But to prevent the possibility of tissue rupture during suc-
cessive irritations, a piece of glass dipping-rod was heated and
shaped so that a smooth spindle-like thickening was left in
the middle of two thin elongated handles. The swollen bulb
of the rod was then placed on a clean dry leaf, the handles
projecting from the lower and upper ends of the closed blade.
After successive intervals of two hours the rod was re-
peatedly raised and lowered so as to irritate the hairs. When
the leaf-halves were pulled slightly asunder after sixteen
hours, copious secretion was going on. In this as in the
above and succeeding experiments the leaves remained closed
from eleven to fifteen days, and during the greater part of
that time were bathed with the secretion. Strands of clean
cotton thread have been inserted, and pulled back and for-
ward at intervals of one to two hours for eleven to thirteen
hours. The leaves secreted freely after fourteen to twenty
hours. One that was similarly treated, but stimulated at
intervals of two to four hours, began to secrete after thirty-
one hours, but only poured forth copiously after three days.
The secretion steadily increased till at least the fifth day. It
is a mistake, therefore, to suppose that the secretion ceases
after one or even two days activity. This has been verified
by numerous other experiments.

With such results before us it seemed highly probable that
continued electrical stimulus would also excite secretion.
The terminals of a battery were slightly bent so as to accom-
modate themselves to the external, slightly convex, basal
surfaces of a leaf after being shut. On application of the
terminals to the external surface the halves closed by electrical
stimulus and gradually tightened up. Three experiments were
thus made and proved entirely successful. The secretion

I

38

Macfarlane. — Contributions to the History of

began to pour out in nine hours from one, in eleven hours
from another, and in twelve from the third. In its behavior
it resembled the ordinary fluid.

It thus appears that the secretion is entirely due to irrita-
tion of the protoplasm, and is poured out alike by mechanical,
chemical, and electrical stimulus. Now any irritant stimulus
applied to protoplasm causes rearrangement of its molecules.
In the doing of this work, decomposition of its substance
occurs with the setting free of decomposition products, either
into the cell cavity or outside the cell. The best and most
accurately investigated cases of this are derived from mus-
cular tissue among animals, where definite decomposition
products such as sarcolactic acid, carbonic acid, acid phos-
phate of potash and various nitrogenous compounds can be
detected. But in the secretion of Dioncea several products,
including some acid body, can be detected. Dr. T. A. G.
Balfour* states that Prof. Dewar determined the presence of
formic acid as well as various chlorides.

Gardiner states that after absorption various new substances
can be observed, and says, ** sections of leaves which were
placed in alcohol thirty-six hours after feeding show that the
cells contain a very large number of tufts of crystals, which
are present in the cell vacuole, and adhere to the inner sur-
face of the cell protoplasm. The tufts are formed of fine
acicular crystals, which crystallize out with great regularity
and radiate from a central point. The tufts are of a yellow-
green color. They are insoluble in alcohol, and in one per
cent, acetic acid. The formation of these crystals may be arti-
ficially produced by wetting the surface of a fresh leaf with
the fluid from a leaf which has fed for a period of from thirty-
six to forty-eight hours." Before perusing his account I had
experimented with the secretion and found the substance
described by him, but in all cases it crystallized out on addi-
tion of absolute alcohol with startling rapidity. Thus, when
a secretion-drop on a slide was placed under the microscope,
the secretion appeared glairy and indistinct as already de-
scribed, but a few drops of alcohol caused formation of the

^ Transactions of Botanical Society, Edinburgh, Vol. XII, p. 340.

Dioncea Muscipula, Ellis.

39

minute needle-like crystals almost more quickly than the eye

could trace.

On surface view each gland cell in the resting state, or just
previous to secretion, shows one large vacuole of a reddish
color, or 'more rarely several small ones, surrounded by finely
granular protoplasm. As secretion proceeds a clear refractive
viscid globule pushes or oozes out from the protoplasm into
the purple vacuole, and at times divides it up. It presses •
against the free wall face and gradually oozes out as a sur-
face excretion. The amount and continuity of the secretion
depend largely on root absorption, as it may cease soon after
first appearance if the soil becomes dry, or if reduced in
amount on this account it can be again increased by watering.

As secretion proceeds the starch grains that are abundant
during the resting stage in the mesophyll cells seem to
change into a yellowish oil that dissolves readily in ether.
This travels along the bast cells or related elements of the
phloem, and is distributed radially from these. It is possible
that this oil may be so acted on that it may become the
source of the excreted formic acid already spoken of.

IV. Leaf-opening.
We do not attempt at present to discuss the changes sub-
sequent to secretion, and the absorption of food materials that
have been digested by the secretion. In from ten to fifteen
days after closure the leaf re-expands, but remains rather
torpid for several days after doing so. A remarkable pecu-
liarity, however, has been observed in leaves that are just
opening after artificial stimulation, or after secretion and
digestion. When the leaf is a healthy one, and has not been
greatly exhausted by repeated acts of digestion, if one or two
of the marginal bristles on each leaf-half are firmly caught by
forceps, the halves very gently and steadily pulled asunder
and held in this position for one to two minutes, it will then
be found that the leaf remains in the expanded state and can
contract on stimulus. Thus, a leaf which had been mechani-
cally irritated was slowly opening fourteen hours after, and
when expanded was thrice irritated when it slowly closed.

w

40 Macfarlane. — Contributions to the History of

Numerous such experiments were repeated with similar re-
sults, and others in which a series of summation stimuli were
communicated behaved in the ordinary way. The only par-
allel case to this that we know is that termed "contraction
remainder" in animal muscle, where namely as contracted
muscle is relaxing a weight applied to it will cause rapid and
permanent expansion.

We may now attempt to unify the results already given.
We conclude that a leaf of Dioncea, previous to secretion, is
in a state of tetanic contraction. This tetanic state results
from a series of stimuli that m'ay either be partially or entirely
mechanical, thermal, luminous, chemical, or electric. Tetanic
contraction of a leaf growing in the wild state, is due to two
mechanical stimuli by an animal, which thereby cause leaf
closure, succeeded by repeated mechanical stimuli as the
captured animal struggles to escape, and continued by numer-
ous chemical stimuli as the digested excretions of the animal
act on the gland protoplasm, and through it on the general
cell protoplasm of the leaf.

Rarely it may happen in the wild state, but can readily be
demonstrated in the laboratory, that two rapidly applied
stimuli are propagated as one shock, and a third is then
needed to cause closure, the subsequent results being the
same. Any form of energy, alone or conjoined with others,
causes closure and tetanic contraction.

Secretion succeeds tetanic contraction, though it is not
dependent on it, but the amount of secretion seems largely to
depend on the amount of stimulus. Thus, in the case of
leaves that secreted feebly when small cubes of roast meat
were placed on them, the absorption by, and stimulation of,
the protoplasm in the gland cells, through presence of nitro-
genous material, might be sufficient to cause activity in the
protoplasm that would set up a limited waste excretion. In
tetanic closure the stimulus being correspondingly greater,
the tissue waste exuding as a secretion is correspondingly
greater in amount.

These phenomena are only explicable in terms of the pro-
toplasmic activity. From a careful study of like phenomena

Dioncea Muscipiila^ Ellis.

41

inDrosera, Gardiner concludes that **the protoplasmic utricle"
is traversed by pores, and that one effect of contraction ** is
an increased impenetrability of the primordial utricle and a
consequent decrease in the size of the molecular pores." Any
contraction must ultimately be referred physically to aggre-
gation of certain molecules at the expense of others, though it
does not follow in the case of Dioncea, that the main contrac-
tion changes can be traced to visible pores. But the writer
has already stated that he believes the protoplasm of certain
cells exhibits appearances which may point to such a possi-
bility, though as yet the observations are insufficient to war-
rant special importance being attached to them. It has been
shown further, that the protoplasm of most cells is continuous
with that of neighbor cells by twenty to seventy-five intercel-
lular processes. Proof has been adduced by Batalin, Burdon
Sanderson and others, that closure is due to the inner side
of each leaf, becoming less turgid than the outer, owing to
migration of liquid from the former. In a Royal Institntion
Lecture Sanderson' further states: "It has, I trust, been
made clear to you that the mechanism of plant motion is
entirely different from that of animal motion. But obv^ious
and well marked as this difference is, it is, nevertheless, not
essential, for it depends not on difference of quality between
the fundamental chemical processes of plant and animal pro-
toplasm, but merely on difference of rate or intensity. Both
in plants and animals, work springs out of chemical trans-
formation of material, but in the plant the process is relatively
so slow that it must necessarily store up energy, not in the
form of chemical compounds, capable of producing work by
their disintegration, but in the mechanical tension of their
elastic membranes. The plant cell uses its material continually
in tightening springs, which it has the power of letting off, at
any required moment, by excitation. Animal contractile pro-
toplasm, and particularly muscle does work only when
required, and in doing so uses its material directly."

Now, it may well be asked here, does the cell wall play a
specially important part, or is it not rather the case that the

» Nature, 1882, p. 486, et. seq.

• • • •

42

Macfarlane. — Contributions to the History of

living cell protoplasm is the direct and active agent ? First, it
is to be remembered, that since the cell walls are traversed
by pores, the protoplasmic threads either must act as perfect
plugs to prevent general diffusion of liquids between the
cells, or the cells that are traversed by pores must be cut off
in the unexcited state from the intercellular spaces, or from
other cells that are not provided with pores, if such exist.
Otherwise none of the walls could become tense.

It appears to be more in accordance with our present
knowledge if we regard the protoplasmic utricle as the layer
which can retain or give off its contents according to its
molecular condition, and that the cellulose membranes are
merely secondary strengthening sacs that act much like the
netting bags which surround rubber bellows. Our reasons
for this opinion are that the summation results, the effects of
different chemical agents, and of energy in varying forms
demonstrate a gradual and very exact contraction of the pro-
toplasm, with corresponding contraction of the leaf substance.

Now, if this be accepted as a working hypothesis only, the
question arises as to how the protoplasmic utricle of the cell
is affected by stimuli. It is manifest that some change in the
leaf cells follows stimulation of the hairs, and in view of their
structure and relations this seems to be largely propagated, or
distributed from the columnar cells that form the joint of the
hair. We would suggest that on first mechanical stimulation
of the leaf, or stimulation of it by chemical, thermal, or other
action that the protoplasm of each cell at once rearranges local-
ized groups of its molecules so as to form little permeable areas
for the contained sap. This may constitute the change that
succeeds first mechanical stimulus. But on second stimulus
or continued chemical or electrical stimulus, alteration and
aggregation of all the molecules causes contraction of the
utricle and squeezing out of liquids through the pores or
permeable areas, already established, a certain quantity being
speedily expelled through the pores of the hair-joint cells, if
such exist. In any case, we agree with Gardiner that the
cause of movement is to be sought for in protoplasmic
activity, which exhibits itself in permitting or prohibiting

» • • '

{

Dioncea Muscipida, Ellis.

43

the passage of the sap enclosed within the protoplasmic sac
of each cell.

Were animal histologists agreed as to the minute structure
of muscle, and the changes that occur in it during contrac-
tion, it might be possible to make some comparison of veget-
able and animal contractility. Meanwhile the facts and state-
ments to hand, suggest that contractility, alike in the veget-
able and animal kingdoms, is accompanied by migration of
liquids through the protoplasmic substance, and that this is
wholly determined by the molecular condition of the proto-
plasm irrespective of cell walls.

When one attempts to trace how such a complicated
mechanism as that of Dioncea leaf has been evolved, difficulties
appear on every side. Several definite points, however, may
be referred to. From Darwin's first statement to Lindsay's
recent one, the poverty of root development has arrested the
attention of observers. The writer dug up a lot of plants
entire, and carefully washed them. He found that the roots
never branch; that from below each cluster of fresh, or
recently decayed leaves, three to, seven, or in strong plants,
nine roots arise; that the average number is four; that the
roots are from a half inch to three inches long, according to
age ; that the average total length of functional root-system on
a plant at any one time, is eight inches, and of this only a part
is covered by functional root -hairs. It is a mistake, however,
to suppose, as some have done who never visited the locality,
that DioncBa usually grows among moist Sphagmnn. A very
few plants occasionally occur in such situations, but at least
seven-eighths grow in a loamy sand. It is a rather significant
fact, also, that they often grow in the midst of such plants as
Seymeria tenuifolia, Gerardia purpurea and G. tenuifolia, that
are equally poor as Dioncea in root development, but have
formed strong, parasitic root-suckers, by which they draw sap
from the densely interlaced roots of ericaceous shrubs, as well
as grasses and other herbs. The fly-catching capabilities of
Dioncsa, and parasitic connections of Seymeria and Gerardia
are parallel physiological advantages that one can vividly see
the benefit of for each, when studied in relation to natural
surroundings.

^4 Macfarlanc. — DioncBa Miiscipula, Ellis.

The relatively large number of leaves provided with more
than six hairs, that the writer has encountered, probably points
to a condition when the hairs were more diffuse in their
distribution; while the close similarity in development and
structure between the brown stellate leaf hairs and the
secreting glands, favors the view that the latter are merely
specialized examples of the former.

EXPLANATION OF PLATE IV.

Illirstrating Dr. J. M. Macfarlane's paper on Dioncea.

Fig. I. Surface view of lower part of sensitive hair, a, base of hair j
b, sensitive joint ; c, shaft. The cell wall surfaces of b exhibit minute pit
areas, x 400°.

Fig. 2. Longitudinal median section of lower part of sensitive hair •,.
X 400°.

Fig. 3. Transverse section of hair at junction of base and joint cells ;
X 400°.

Fig. 4. Outline surface view of secreting gland ; X 400°.

Fig. 5. a,b,c and d. Forms of hair from lower leaf surface ; X 400°.

Fig. 6. Vertical section of gland parallel to secondary leaf bundles ;
X 400°.

Fig. 7. Vertical section of gland at right angles to the last.

Fig. 8. Surface view of epidermal cells, after slight treatment with
dilute sulphuric acid and eosin staining ; X °75o.

Fig. 9. Surface view of epidermal cells after iodine and sulphuric
solution treatment and eosin staining. The continuity of the intercellular
protoplasmic threads is traceable ; X 750°.

Fig. 9a. Portion of swollen cell wall penetrated by intercellular pro-
toplasmic threads. The thickened areas in the three figures are rather
exaggerated in amount.

Vol. I V\l\W'\.

Bol. Colli. I

'^'' Msvlvajiia.

9

Madarlnnc on l)i()iia\t

|l

MUTILATED TEXT

V

An Abnormal Development of the Inflorescence

of Dionsea.

By John W. Harshberger, A.B., B.S.

{WITH PI, ATMS V AND VI.)

THE peculiarly developed plant of Dioncea now to be
described grew in the greenhouse of the Biological
Department of the University of Pennsylvania, under
ordinary conditions of heat and moisture. The specimen
flowered about the usual period, but the abnormal con-
dition displayed by it was not observed till after the flowers
had somewhat withered.

The scape rose as usual from the centre of the rosette of
leaves and produced normal flowers at the end of the branch
that is cut off and placed at the side in the illustration for
sake of clearness (Plate V, Fig. 2). This branch breaks up
into two main shoots, each of which divides into several
flower-bearing stalks.

One of the main shoots of the scape is subtended by a long
lanceolate bract, and a smaller bract is found on it a little
above the insertion of the first. In the axil of the smaller
bract two structures have developed, the lower being an elon-
gated branch, the upper evidently a metamorphosed flower,
(Plate V, Fig. i, d).

The elongated branch organ has a central rhizomatic axis
that produces leaves arranged in a flat spiral. Sections of the
leaves when magnified show normal digestive glands, and
sensitive hairs occur on the upper laminar surface. In these
and all other respects, therefore, they agree with vegetative
or foliage leaves. True roots (Plate V, Fig. i, c), developed at
the leaf bases, grow out from the rhizome. Sections of one
of these roots reveal radially arranged bundles, in which
phloem areas alternate with xylem strands.

II

46 Harshbcrger. — An Abnormal Development of

A third bract is reached in acropetal succession from the
first (Plate V, Fig. i, g^), and in its axil a growth arises that
corresponds in every particular with the branch-organ already-
described as springing from the axil of the second bract.

The axillary position of the foliar shoots and their relation
to the flower-like shoot give an insight into the morphological
relations of the teratological parts. In the abnormal floral
branch (Plate V, Fig. i, g)an indurated collection of nodular
bodies strikingly resembles ovarian tissue. Bract G, as we
have seen, subtends two branches, the lower being a veget-
ative shoot, the upper or superposed one resembling a flower.
This condition corresponds to the superposed buds met with
in such plants as Loniccra tartarica, which produces three to
six axillary buds on some shoots ; ox Juglans cinereUy in which
the primary axillary bud remains latent, and several accessory
axillary buds push out, of which one elongates into a branch.

The histology of Dioncsa has been studied and described by
Kurtz,' C. de Candolle' and Fraustadt,^ though most observers
in their desire to study the specially interesting leaf traps,
have to some extent neglected the anatomy of the underground
parts and of the scape. The vegetative shoots above referred
to seemed well suited for a study of the fibro-vascular bundle
distribution.

A cross section of the scape two inches below the point of
branching is illustrated in Plate VI, Fig. 4. The section
shows externally an epidermis made up of large polyhedral
cells. Beneath are five or six rows of loose parenchymatous
elements with intercellular spaces. The phloem or soft bast of
the bundle consists of sieve tubes and companion phloem cells,
while a few xylem cells are intimately connected with the
small phloem areas. A zone of parenchyma intervenes
between the small external bundle and the larger fibro-vas-
cular area within. The outer portion of this area is occupied
by the phloem, the inner by xylem, made up of pitted vessels,
also spiral and scalariform tracheids. The centre of the sec-
tion consists of cellular tissue.

' Archiv. fiir anat. und Physio- wissen. Medicin, 1876.
sArchiv. de Sc. Physik. et Nat. Geneva, 1875-76
3Cohn's Biologic der Pflanzen, Band II, pp. 5o-59-

)

the Inflorescence of Dioncea.

47

S

A cross section of the branch G of Plate V. made a little
above its insertion into the scape, is represented in Plate VI,
Fig. 5. Beneath a single layer of epidermal cells is a wide
zone of fundamental tissue, succeeded by a layer of scleren-
chyma interspersed by white, glistening, sclerotic cells. In-
ternal to these is a zone of phloem tissue surrounding a
central mass of xylem. The course of the bundles in the
branch can now be referred to. The leaves are arranged in a
flat alternate spiral (Plate VI, Fig. 3). Fraustadt {op. cit.)
says : " The whole rhizome is short and broad, the leaves are
placed on it with broad, flat insertions without internodes."

" The phyllotaxy I have not been able clearly to define, the
younger leaves are apparently two-ranked ; the leaves overlap
one another with their foliaceous petioles." " Frequently the
leaves show an open spiral arrangement." The abnormality
with lengthened internodes (Plate VI, Fig. 3) is especially
valuable, as it indicates more plainly the phyllotaxy and the
distribution of the fibro-vascular bundles. Fraustadt, in his
study of the underground short rhizome, encountered a mass
of fibres. A cross section of the abnormal branch, half-way
between its proximal and distal ends, appears in Plate VI, Fig.
6. Fig. 2 shows the relationship of the parts. The central
tissue of the branch shows the main bundle trace (A) two
leaves (C and E) come off from either side of this central
area, a third leaf (O) arises above.

A very thin section, at this point (A, PI. VI, Fig. 2), is
magnified and shown in Fig. 6. We have in this figure the
central tissue (A^) with the two leaves arising on each side
(C and E'), the third leaf O' arising above. A puzzling
ramification of bundles here occurs, complicated by the dif-
ferent directions of leaf growth. The bundles at the proxi-
mal end of the rhizome are represented in Fig. 5, PI. VI.
When we reach the middle section, Figure 6, a change takes
place. Before dividing into separate strands for the leaves
(C\ ES O'), a twist in the stem bundles must occur for the
three phloem areas come together at A', with the xylem circum-
ferential. The leaves in their origin run toward the growing
point a little way, then take a sharp bend, and point backward

46 Harshbcrgcr. — An Abnormal Development of

A third bract is reached in acropetal succession from the
first (Plate V, Fig. i, g^), and in its axil a growth arises that
corresponds in every particular with the branch-organ already-
described as springing from the axil of the second bract.

The axillary position of the foliar shoots and their relation
to the flower-like shoot give an insight into the morphological
relations of the teratological parts. In the abnormal floral
branch (Plate V, Fig. i, g)an indurated collection of nodular
bodies strikingly resembles ovarian tissue. Bract G, as we
have seen, subtends two branches, the lower being a veget-
ative shoot, the upper or superposed one resembling a flower.
This condition corresponds to the superposed buds met with
in such plants as Lonicera tartarieay which produces three to
six axillary buds on some shoots ; or Jnglafts einereUy in which
the primary axillary bud remains latent, and several accessory
axillary buds push out, of which one elongates into a branch.

The histology of Dion^a has been studied and described by
Kurtz,^ C. de Candolle^ and Fraustadt,^ though most observers
in their desire to study the specially interesting leaf traps,
have to some extent neglected the anatomy of the underground
parts and of the scape. The vegetative shoots above referred
to seemed well suited for a study of the fibro-vascular bundle
distribution.

A cross section of the scape two inches below the point of
branching is illustrated in Plate VI, Fig. 4. The section
shows externally an epidermis made up of large polyhedral
cells. Beneath are five or six rows of loose parenchymatous
elements with intercellular spaces. The phloem or soft bast of
the bundle consists of sieve tubes and companion phloem cells,
while a few xylem cells are intimately connected with the
small phloem areas. A zone of parenchyma intervenes
between the small external bundle and the larger fibro-vas-
cular area within. The outer portion of this area is occupied
by the phloem, the inner by xylem, made up of pitted vessels,
also spiral and scalariform tracheids. The centre of the sec-
tion consists of cellular tissue.

^ Archiv. fur anat. und Physio- wissen. Medicin, 1876.
SArchiv. de Sc Physik. et Nat. Geneva, 1875-76.
3Cohn's Biologic der Pflanzen, Band II, pp. 50-59.

the Inflorescence of DioncBa.

47

A cross section of the branch G of Plate V, made a little
above its insertion into the scape, is represented in Plate VI,
Fig. 5. Beneath a single layer of epidermal cells is a wide
zone of fundamental tissue, succeeded by a layer of scleren-
chyma interspersed by white, glistening, sclerotic cells. In-
ternal to these is a zone of phloem tissue surrounding a
central mass of xylem. The course of the bundles in the
branch can now be referred to. The leaves are arranged in a
flat alternate spiral (Plate VI, Fig. 3). Fraustadt {op. cit.)
says : " The whole rhizome is short and broad, the leaves are
placed on it with broad, flat insertions without internodes."

" The phyllotaxy I have not been able clearly to define, the
younger leaves are apparently two-ranked ; the leaves overlap
one another with their foliaceous petioles." ** Frequently the
leaves show an open spiral arrangement." The abnormality
with lengthened internodes (Plate VI, Fig. 3) is especially
valuable, as it indicates more plainly the phyllotaxy and the
distribution of the fibro-vascular bundles. Fraustadt, in his
study of the underground short rhizome, encountered a mass
of fibres. A cross section of the abnormal branch, half-way
between its proximal and distal ends, appears in Plate VI, Fig.
6. Fig. 2 shows the relationship of the parts. The central
tissue of the branch shows the main bundle trace (A) two
leaves (C and E) come oflF from either side of this central
area, a third leaf (O) arises above.

A very thin section, at this point (A, PI. VI, Fig. 2), is
magnified and shown in Fig. 6. We have in this figure the
central tissue (A^) with the two leaves arising on each side
(C' and E*), the third leaf O' arising above. A puzzling
ramification of bundles here occurs, complicated by the dif-
ferent directions of leaf growth. The bundles at the proxi-
mal end of the rhizome are represented in Fig. 5, PI. VI.
When we reach the middle section, Figure 6, a change takes
place. Before dividing into separate strands for the leaves
{C\ E', 00, a twist in the stem bundles must occur for the
three phloem areas come together at A', with the xylem circum-
ferential. The leaves in their origin run toward the growing
point a little way, then take a sharp bend, and point backward

II

48

Harshberger. — A71 Abnormal Development of

proximally. This reverses the direction of the bundles, and
brings the phloem again into its true position on the lower side
(V, PI. VI, Fig. 6) of the leaves Q\ E\ 0\ The three bundle
traces must have twisted, and divided a little back of the point
AS which thus gives the peculiar bundle anatomy at A* (PI.
VI, Fig. 6). We have in Fig. 6 a central rhizomatic tissue (A^),
with leaf tissues in the centre above and on each side. Each
wing of tissue receives a bundle trace from the main axis cylin-
der. A solution can be found apparently for the curious bundle
arrangement if it be considered that a twist in the fibre occurs.
Fraustadt found the whole mass of parenchymatous tissue in
the underground stem and leaf petioles crowded with oval
starch grains. The sections of "aerial rhizome" showed the
whole parenchyma densely packed with starch, which made
the section somewhat opaque.

The axillary structures now referred to are clearly homo-
logues of the buds or bulbils encountered in such plants as
Liliiim Bidbiferuniy atid Raniinailus Ficariay and they suggest
the possibility of vegetative or non-sexual reproduction in this
particular case. It might have been possible to have grown
new individuals from the axillary vegetative branches, but the
suggestion of this idea came after the plant had been killed
and preserved in alcohol. Authorities can only therefore be
cited which seem to sustain this belief in non-sexual propaga-
tion in DioncEa. Miss Elizabeth H. Willis records an interest-
ing case in point, that was noticed on a plant of Dioncea in her
possession. "The leaves have continued to increase by
sending out runners, which have taken root all around the
parent plant, until now the group of independent branches
(except for the creeping recumbent stem, which seems to unite
them together), numbers about twenty-five."^ A curious
record is made by Hog?:,^ "that the leaves of Dioncea^ if
placed in damp moss, will take root and produce young plants
on the margin." Nitschke^ has recorded the production of
adventitious plants on the leaves of Drosera^ a near relative

1 Botanical Gazette, Vol. X, 1885, p. 214

2 Nat. Hist, of the Vegetable Kingdom, 1858, r 84.
8 Botanische Zeitung.

|i

MUTILATED TEXT

[Vol. I, Plate v.]

Bot, Cont. Univ. Pennsylvania.

Harshberger on Dion.^a.

[Vol. 1, Plate VI.]

Bot. Cont. Univ. Pennsylvajiia.

the Inflorescence of Dioficea,

49

of the Venus Fly Trap, while additional literature on the
subject is to be found in Annales des Science Naturelle^
Science Gossip,'^ and Nature.^

From the above data we may conclude :

(i) That the production of individuals by bud propagation
on the leaves of Dioncea and Drosera occasionally occurs.

(2) That the condition described above in Dioncea is a case
of non-sexual reproduction supplanting the ordinary sexual
process.

(3) That the bud axis may be considerably elongated, and
provided with evident roots and leaves previous to separation
from the parent axis.

EXPLANATION OF PLATES V AND VI,

Illustrating J. W. Harshberger's paper on Dioncea.

Plate V.
Fig. I. Plant enlarged.

G, G^ Vegetative axillary branches.

D. Flower-like branch.

F. Leader figured at the side.
Fig. 2. Main leader or branch.

Plate VI.

Fig. I. Leaf detached showing origin of the root.
Fig. 2. Ideal representation of the outgrowth half way between distal
and proximal ends.

A Central tissue.

B Middle leaf.

C & E Lateral leaves.
Fig. 3. Outgrowth showing phyllotaxy.
Fig. 4. Cross section of the scape.

Fig. 5. Cross section, vegetative branch at proximal end.
Fig. 6. Middle section of same branch.

A' Central tissue.

Q\ E», O' Leaves.

1 Annales des Sci. Nat., XIV, 1840, p. 14. •

2 Science Gossip, 1873. P- 239; '883, pp. 44 and 91.

3 Nature, XV, p. 18.

. i

l!H

Harshbkrgrr on Dion^,a.

Mangrove Tannin.

By Prof. Henry Trimble, Ph.M.

(WITH PLATSi VII.)

[The following interesting paper by Professor Henry Trimble on Man-
grove Bark needs no word of introduction from anyone. It may, however,
not be out of place to call attention to the illustration of a mangrove thicket
near Port Morant, Jamaica, and to add that, if the extraction of tannin
from mangrove bark can be made a commercial success, the new
industry so created must react most favorably upon our hemlock forests.
For the hemlock trees, there are so many uses, that it seems unwise to
cut them down, as has often been done solely to obtain the bark. This
is especially so, when one remembers that our supply is by no means inex-
haustible, and that the rate of reproduction is extremely slow. The man-
grove (Rhizophora Mangle, L.) is a common inhabitant of tropical sea-
boards around the world, and save for the possible tannin producing ca-
pacity of its bark gives promise of no great commercial value to us.]

J. T. ROTHROCK.

\y\ ANGROVE bark from Rhizophora Mangle, L. has
/*'\ been frequently mentioned as a possibility in the
i V tannin industry. Probably the first reference to it
was made by Dr. James Howison ' in 1804, when he received a
gold medal from the Society of Arts, for his description, and a
sample of the extract made in India from 400 pounds of the
bark. The method of preparing this extract, however, was
not such as to give the largest yield or the most satisfactory
product, since the extraction was carried on without the aid
of heat, and the resulting liquor was evaporated by exposure
to the sun until quite concentrated, when it was finished by
the application of heat. There must have been considerable
deterioration of the product by fermentation. This sample
of fifty-four pounds cost eight shillings in Bengal, and the

> •• Preparation of Tan made in the East Indies from the Bark of the Mangrove Tree."
Transactions of the Society of Arts, 22, 201.

'

I

Trimble. — Mangrove Tannin.

51

author thought it could be prepared for ten shillings per hun-
dred weight.

More recently, in 1846, we find a description of mangrove
by W. Hamilton,' in which he gives an interesting account of
its manner of growth and possible uses.

It appears, however, not to have come into use, except in
the localities where it grows. The probable reasons for this
are(i) the fact that almost all parts of the world are able to
supply tanning material for home consumption, and (2) that
the mangrove produces a leather of a bad color and a spongy
texture. It is, therefore, of interest chiefly as a possibility,
for, in the event of its being needed, the color, and spongy
character which it imparts, could, no doubt, be corrected. It
has been thought that an investigation of the constituents of
the bark as well as a study of its peculiar tannin might be of
interest. F'or, if not for immediate use, it would, by giving
us a knowledge of the individual tannin, increase our infor-
mation on the whole class of tannins, of which we at present
know so little.

The material for this investigation was supplied to me by
Professor J. T. Rothrock, who collected it during his southern
scientific expedition in the winter of 1890-91.

The bark, after powdering, was found to contain 12.04 per
cent, of moisture, and 6.10 per cent, of ash. Sodium salts
predominated in the ash, chiefly as the chloride.

Only insignificant amounts of fat, wax, and compounds of
that nature were found.

Stronger ether removed 0.40 per cent, of a substance insol-
uble in water, but which caused a green color with ferric chlo-
ride. Gallic acid was proved to be absent.

Absolute alcohol extracted 20.32 per cent, of a very astrin-
gent substance with a narcotic odor. Of this an amount equal
to 13.42 per cent, of the bark was soluble in cold water, and
the remainder was almost completely soluble in hot water.
The alcoholic extract was found to consist largely of tannin,
red coloring matter, and a small quantity of glucose.

1 " On the medical and economic properties of the Rhizophora Mangle, or Mangrove

Tree."

Pharmaceutical Journal and Transactions, 6, ii.

N

52

Trimble. — Mangrove Tannin,

The other more important constituents were determined to
be, mucilage 172 per cent., ghicose 0.81 per cent., albumen-
oids 7.02 per cent., starch 4.27 per cent., and cellulose 27.49
per cent. Tannin was determined by gelatin and alum, and
found to be, in the air-dry bark, 23.92 per cent. ; this was an
average of three closely agreeing results. For future com-
parison it may be well to note that this by calculation indi-
cates 27.19 per cent, of tannin in the absolutely dry bark.

Preparation and Purification.
A quantity of the tannin was prepared by extracting the
finely powdered bark with commercial ether (specific gravity
0.750). After recovering most of the solvent, the remaining
extract was distilled to dryness under reduced pressure. The
residue was dissolved in ether of the same specific gravity,
and the solution concentrated to dryness under reduced
pressue, which had the effect to render the tannin quite por-
ous and in a condition to be dissolved by water, which was
next used. The filtered aqueous solution was carefully treated
with solution of neutral lead acetate. In this operation it is
to be noted that the first few drops of the reagent caused a
precipitate, which on stirring disappeared. A further addi-
tion of the reagent caused a precipitate of the coloring
matter, and on filtering a yellow filtrate was obtained.
This filtrate was agitated successively with three portions of
acetic ether, which removed the tannin. The precipitate was
suspended in water and decomposed with hydrogen sulphide,
the mixture filtered and the filtrate, after heating under re-
duced pressure to remove hydrogen sulphide, was treated
with lead acetate as before and the filtrate agitated with
acetic ether, which removed an additional quantity of tannin.
This acetic ether solution was mixed with that from the pre-
vious agitation and the whole distilled to dryness, the residue
dissolved in ether with some water, and again distilled to dry-
ness under reduced pressure.

Properties.
The tannin obtained by the above process was in light red-
dish-yellow porous masses, completely and readily soluble in

'

Trimble. — Mangrove Tannin.

53

water, alcohol, and commercial ether. It was partly precipi-
tated from its aqueous solution by saturation with common
salt. That not precipitated was removed by agitation with
acetic ether, and, although lighter in color, it was found to be
identical with the darker portion precipitated by the salt.

The following are the reactions of the mangrove tannin, in
one per cent, solution, with the usual reagents. There is
added for comparison the behavior of a similar solution of
gallo-tannic acid.

Reagent.
Sulphuric acid (i to 9 of

water),
Bromine water,

Ferric chloride,

and
Ammonium hydrate,

Tartar emetic,

and
Ammonium chloride,

Calcium hydrate,

Concentrated sulphuric
acid,

Lead nitrate,

Cobalt acetate,

Uranium acetate,

Potassium bichromate,

Ferric acetate,

Mangrove-tannin.

Red deposit on cooling.
Yellow ppt.

1

Dirty-green ppt.
Purple ppt.

I
j

I No ppt.
1 No ppt.

Pink ppt.

Red on surface.

Gallc-tannic Acid.

No change.
No ppt.

Blue-black ppt.
Purple ppt.

White ppt.
White ppt.

{

{

White ppt.
Turning blue.

Deep-red color. Yellow color.

No ppt. White ppt.

Faint cloudiness. Flesh-colored ppt.

Red-brown color and ppt. Crimson color.

Brown ppt. Brown ppt.

Olive-green color and ppt. Blue-black ppt.

These reactions agree closely with those given by Procter
for the tannin of Mimosa or wattle bark.

A further examination of the above mangrove-tannin failed
to reveal the presence of sugar. This was accomplished by
precipitating a solution of 0.5 gramme of the tannin with lead
oxyacetate, removing the lead from the filtrate by hydrogen
sulphide, and, after removal of the latter, testing with Feh-
ling's solution.

The Action of Heat.— By heating another portion of the
tannin in glycerin to 215°, shaking out the products of de-

^ Text-book of Tanning, p. X13.

54

Trimble- Mangrove Taufiin,

composition with ether, and, after removing the latter, dis-
solving in water, and applying the usual reagents, a catechol
tannin was indicated.

Hydrolysis.— O.S gramme, after boiling for three hours with
a two per cent, solution of hydrochloric acid (absolute HCl),
yielded a red insoluble decomposition product, and a sub-
stance capable of reducing Fehling's solution to an extent
which indicated 10.35 P^r cent, of glucose. The above red
product was readily oxidized by nitric acid yielding a red-
brown solution which rapidly faded, and which with water
became paler, precipitation taking place at the same time.
The original product was not completely soluble in cold or
hot alcohol nor in ammonium hydrate. That which did dis-
solve in alcohol was not precipitated by a large quantity of
water. These properties indicated the presence of gallic acid,
ellagic acid and phlobaphenes.

Composition.

After repeated precipitation of the tannin until it was of
a light reddish-yellow color, it was dried at 120° and sub-
mitted to elementary analysis with the following results :

(i) .2353 gramme gave .5188 CO2 and .1023 H2O.

(2) .2475 " " .5414 CO2 and .1027 H2O.

(3) -2556 " " .5577 CO2 and .1070 H2O.

indicating the following percentages :

(I)

(2)

(3)

Average.

c.

60.13

59-66

59-51

59-76

H.

4.83

4.61

4-65

4-69

0.

35-04

35-73

35-84

35-55

, Calculated for v

C25H25O11 C20H20O9 C27H24O11
59.88 59-4° 60.93

4-99 4-95 4-68

35-13 35-65 34.39

The formula C25H25O,, comes the nearest to representing
the average percentages obtained ; the formula C20H20O9 is
that given by Dragendorff for rhatania-tannin ; and the for-
mula C^eH^^Ox, is the one given by Rochleder for horsechest-

[Vol. I, Plate VII.]

Bot, Cont. Univ. Pennsylvania.

\

TRiMni.E ON Mangrove Tannin.

I!

1

Trimble— Mangrove Tannin.

55

nut-tannin, with which ^l^-^-">-^^r"' ^""J^d'cloSy
bowski, and tormentil-tannin, according to Rembold, closely

'^Tt; conclusion naturally reached by .^Ws investigation is
that we have in mangrove a tannin which is identical with
those from horsechestnut. rhatany and tormentil, and pos-
sibly also with that from mimosa or wattle bark.
Philadelphia, 1892.

I'

|f t

Wilson.— Observations on Epigcea repens, L.

57

Observations on Epigsea repens, L,

By W. p. Wilson, D.Sc.

E

( WITH PI, A T^ VIII. )

; PIG^A rcpcns or Arbutus is one of our North Amer-
ican plants, which has a very wide distribution. It
flowers early in the spring, in some places soon after
the snow has disappeared.

Owing to its early appearance, its often beautifully tinted
and sweet scented flowers, and its trailing evergreen leaves it
is a great favorite wherever it grows.

In New England it passes under the name of Mayflower.
As soon as it opens in the spring it is brought into Boston,
tied up into little bunches, with most of its attractive leaves
stripped off, and exposed for sale on the streets.

In the same way it is sold in New York, Philadelphia and
other cities. It has been nearly exterminated within fifty
miles of Boston. It still grows in great abundance in the
Pme barrens of New Jersey, but with the small army of col-
ored women actively engaged in pulling it up for the Phila-
delphia market its extermination in this locality is only a
matter of time.

Its range of growth extends from Canada to Florida
along the coast, and west to Minnesota, Michigan and Ten-
nessee. In all these localities it is much sought after by
lovers of flowers, and is picked and exposed for sale, or used
for home decorations.

In 1796 Micheaux, while journeying in the Alleghany Moun-
tains, picked up this little plant and made the following entry
in his journal: "Le2 Avril. Epig^a repens en pleine fleur
comme les jours precedents: sur plusi. individus toutes les
fleurs femelles sans rudiments d'Etamines et sur d'autres in-

dividus fleurs toutes les fleurs hermaphrodites." ' We also
find in Michaux's Flora " Flores omnes in nonnullis individ-
uis abortivi." Michaux was therefore the first to record the
different forms of this flower and to call attention to the fact,
noted many years later by Gray, that all the flowers do not

produce fruit.

In 1868 Meehan'^ recorded a number of observations ot

which the following is a brief summary :

There is much variation in the size of the corollas in dif-
ferent plants.

There are flowers without stamens, (Pistillate).

Some flowers have five-lobed stigmas that are widely diver-
gent : In others the five lobes are closed. In such the stamens
are present, (hermaphrodite.) ,

The ovaries are larger in corresponding states in the pistil-
late forms than in the staminate.

The pistillate forms shed their corollas first. The corollas
on the hermaphrodite forms dry up without dropping.(??)

Cope communicated to Meehan at this time that the corol-
las of the pistillate forms are recurved and vasiform and may
thus be distinguished from the hermaphrodite form.

Nearly ten years later the following facts were commu-
nicated to the Boston Society of Natural History ' by Gray

and Goodale.

There are four kinds of flowers :

1. Those with long styles and perfect stigmas.

2. Those with short styles and perfect stigmas.

Both of the above kinds with aborted stamens.

3. Those with long styles and imperfect stigmas.

4. Those with short styles and imperfect stigmas.

Both of the above kinds with perfect stamens.
The modified, stigmas on the one hand, and the aborted
stamens on the other are looking toward dioecism ; the differ-

^ From the Journal of Andre Michaux, written during his travels in the UnitedStates
and Canada. 1785-96. Proceedings American Philosophical Society, Vol. xxv... 1889, No.

''^^ ProcLings Philadelphia Academy Natural Sciences, Vol. xx May, 1868, p. i33-
3 Proceedings Boston Society Natural History. Vol. xxiu., 1876.
< Siliman's Journal, July, 1876.

58 Wilson. — Obsen'attofis on Epigcea repenSy L.

ence in the lengths of the styles, and the differences in the
lengths of the stamens are looking toward dimorphism.

It is not known whether the flowers with small stigmas are
ever fertile or not.

In 1891 Halsted examined sixty flowers and came to the
following conclusions : ^

That there is only a tendency towards dimorphism :

There is no difference between the pollen grains taken
from the anthers of long or of short stamens.

He finds the strong tendency to become unisexual in large
part sufficient to account for the differences in length of
styles and stigmas.

During the present year the writer has examined about
1,000 plants of Epigcea repens at the time of flowering. Most
of them were from western North Carolina. A few were
from localities in Canada, Massachusetts, New Jersey and
Pennsylvania.

These flowers readily fall into two groups :

(i) Those having perfect pistils, with or without rudiment-
ary stamens.

(2) Those having pollen-bearing stamens, with rudimentary
pistils.

Occasionally flowers are found in the first group without a
trace of a stamen. Often there are bare vestiges of filaments
at the base of the corolla. In rare cases the anthers may be
present, but without pollen.

In the second group the pistil is rudimentary through an
undeveloped stigma only. The stigma in the first mentioned
form is a five-lobed, star-shaped, terminal body, opening its
lobes out nearly at right angles to the style. In the second
form the stigma is five-lobed, the lobes being closely appressed
into a terminal, functionless, oval enlargement.

The first form is without perfect pollen-bearing stamens,
and is, therefore, pistillate. The second form, having a
functionless pistil through an imperfect stigma, but bearing
perfect stamens with pollen, is therefore, staminate. The

* Bulletin of the Torrey Botanical Club, Vol. xviii., p. 249, 1891.

Wilson. — Obsey-vations on Epigcea repens, L.

59

two forms do not occur on the same, but on different indi-
vidual plants. . u •
Epigcea repens is therefore, not becoming dioecious, but is

already so. .

If a large number of flowers be examined in any given
locality, a few will be found with long styles that carry the
fertile or infertile stigma, as the case may be, considerably
beyond the throat of the corolla (Plate VIII, Figs. 3, 6). In a
much larger number of flowers will be found an extremely
short form with stigmas only half elevated in the tubular

corolla (Figs, i, 7).

All intermediate forms may be found if one examines a
sufficient number of plants (see Plate VIII, Figs. 2, 3, 4, 8).
Two of the most common forms are Figs. 2, or 8 and 5. One
can make out a tri-morphic condition of this plant, and quite
as easily show, with a still farther examination of flowers, a
polymorphic condition. In some of the flower sections on
the Plate bare rudiments of stamens exist, as in Figs. 2 and
3 In Fig. 4 the anthers are present without pollen. Only
one plant with flowers in this stage of development was found
while examining a large mass of material from North Caro-
Una, Pennsylvania, and New Jersey.

In Fig. 5 the stamens are short ; in Fig.^ 7 they are long.
In others they occupy an intermediate position.

Figs. 5 and 7 show the predominating forms. At the
present time cross-pollination is secured with absolute cer-
taintv through dioecism.

The rudimentary stamens and pistils in the different kinds
of flower show that this has not always existed, but that the
flower was once perfect. In this early and perfect condition
of Epigcea it is evident that it developed, as many other
flowers have, these varying lengths of styles and stamens to
aid in securing cross-pollination.^

There are differences between these two kinds of flower
other than those which relate to the essential organs, the
stamens and pistils. These differences indicate that while

1 Any one not familiar with di-and tri-morphic flowers and the way in which they
aid cross-pollination, can consult Lubbock's British Wild Flowers.

6o

Wilson. — Observations on Epigcea repenSy L.

dioecism is of a later development than polymorphism, it is
still of no very recent date.

The corollas of the female forms are as a rule from one-third
to one-seventh smaller than those of the male. Figs. 9 and
10 show an average female and an average male flower drawn
in relative proportions.

The throat is somewhat more open in the female than in
the male form. In color, too, the smaller pistillate flowers
seem to have gained an advantage over the staminate form,
the latter often having little or no color, while the former
develops beautiful shades of pink or rose color. It is not
strange that the smaller and less showy female flowers should
acquire a deeper hue than the larger male flowers in order to
be equally conspicuous and attractive to the insects which
must carry the pollen from one to the other if the seeds are
to be formed.

The whole male plant, under most conditions of growth,
presents a decided appearance of vigor which does not seem
to belong to the female. In the male there is less color the
flowers often being white, the leaves are larger, the vegetating
shoots are longer and more thrifty ; while in the female the
color is brighter and deeper, the leaves are considerably
smaller and the creeping branches much shorter. The male
plant often looks thrifty in localities where the female looks
dwarfed.

In examining Epigcea in different localities this year the
writer has in several cases made a careful numerical estimate
of the ratio of the sexes, to each other. The results obtained,
although necessarily rather indefinite on account of varying
conditions that are difficult to estimate, may be of interest to
some, as touching on the question of development and per-
sistence of sex.

Quite a number of observations and investigations made on
animals seem to show that where the struggle to maintain
life is a hard one, a preponderance of males are produced ;
while on the contrary, when food is abundant and there is no
struggle, females predominate. The very few observations

Wilson. — Observations 07i Epigcea repens, L.

61

which have been made on plants seem to exemplify the same

principle.^

At Blowing Rock, in the mountains of North Carolina, in
a luxuriant forest, in every way the natural home of Epigcea,
the writer made an estimate of the relative numbers of the
two sexes. On a walk of four miles through the woods a
cluster of flowers from every plant met with was plucked,
provided the last plant picked from was not nearer then ten
feet. Conditions were as nearly natural in soil, shade and sur-
rounding vegetation as could well be imagined. In all 98
separate plants were examined ; 45 of these were females and
53 males. This was not very far from an even distribution
of the two sexes, there being only eight more male plants

than female.

On the next day a similar estimate was made on a rocky
knob, over 4,000 feet high, from which most of the trees had
been cut, the bushes burned and more or less of the soil
washed away. On the whole it was an unfavorable place for
the growth of the plant. Only 67 clusters could be found.
Of these 40 were females and 27 males. Again, on the fol-
lowing day an estimate was made for ten miles along a
mountain road, from the sides of which much of the timber
had been cut, often the bushes burned and the plants exposed
to the full rays of the sun. From the adjacent forest Epigcea
had struggled into the exposed places. The locality was not
a favorable one for the plant.

Three hundred and twelve separate plants were examined.
Of these 173 were females and 139 males. Here it will be
seen again that the females predominate. Ten days later
the last examination of this kind was made in Pennsylvania,
near West Chester. Here the plants were picked mostly on
the roadside and on the edge of a sunny, grassy field, where
the facilities for their growth were not as good as in the open
woods. One hundred and thirty-seven plants were examined.
Of these 92 were females and 45 males, an excess of 47
females over the male plants. The writer gives these notes
for what they are worth, being well aware of the fact that a

1 Dusing, Jen. Zeitsch. f . Naturw., XVII, 1883- Heyer, Ber. d. landwirthschaftl. Inst.,
Halle, V. 1884. Meehan, Proc. Acad. Nat. Sci., Phila., 1884.

62

Wilson.— Obsavations on Epigcea rcpens, L.

sufficient number of localities, with character of soil and sur-
roundings have not yet been examined to give them value.
The fact has also been noted that Epigc^a, in some localities,
seldom sets seeds, the plant propagating itself largely in.
such places by stolons. A consideration of such habits must
be taken into account in estimating the predominance of one
sex'over the other. Tabulated the results appear as follows :

•saiBW JO •

juaD jaj

•

^9

1

•saiBiu3j JO juaD J8J

•

•

■0

•

•saiBtuaj jO ssao

00

-xa ui 881? w JO Ja<l«»"NI

•sa^Bj^ JO ssaDxa

S

ui ssiBiuaj JO aaqiunj^

•S91BW JSquinN

^

lO

0

r>

Ss

•saiBiuaj jaquin>j

"*

•*

pauiuiBxa

%

»4

p>

s^UB|d JO jaqiutiM

ro

h and

ithout

Caro-

I 4^ I

C V o

Blow-

noir,

road;

Penn-
West

1

locality
woods
rush; N

locali
ob,

; No

local
k to
moun
irolin

=3"

0

J

-g^csU

able
ania,
ster.

1

.X3

-Sc^fi^

s

-5'g'C rt

0 = ^5

0 bo S 0

S-S.^

>

c:

ft

D

;:5

P

f I

[Vol. I, Plate VIII.]

Bot. Cont. Univ. Pennsylvania.

N

8

10

Wilson on Epig.ka.

Wi/son. — Observations on Epigcea repens, L.

63

Whether this higher percentage of females under severer
conditions of environment, indicates an actual production of
more female forms than males ; or whether an equal number
of both forms are not produced in the first place, the males
being reduced later by stress of surroundings, will necessitate
more careful experimentation for determination.

Observations from any one living in localities where E,
repens grows, are kindly asked for.

EXPLANATION OF PLATE VIII.
Illustrating Dr. W. P. Wilson's, paper on Epigcea repens L.
Figs. I, 2 and 3 show sections of pistillate flowers with varying length

of styles.

Figs. 2 and 3 show rudiments of stamens.

Fig. 4 shows a vertical section of a pistillate flower with complete
stamens. The anthers are, however, without pollen and shrivelled. All
four figures show the star-shaped stigma of the pistillate flowers.

Figs. 5, 6, 7 and 8 represent vertical sections of staminate flowers
with varying lengths of filaments and styles. They show the closed
character of the stigma in the staminate form in which it never opens.

Fig. 9 represents a male flower, with its large and generally less

colored corolla.

Fig. 10 represents a female flower. Figs. 9 and 10 show the relative
size of the two sexes. The female is generally about one-third smaller
than the male.

A Nascent Variety of Brunella vulgaris, L.

By J. T. RoTHROCK, B.S., M.D.

AS commonly seen in the eastern United States, Brunella
vulgaris, Z, is an erect or ascending perennial herb
with its spike or head of flowers raised from five inches
to a foot above the ground. More commonly the stem is
erect and simple ; frequently, however, one finds lateral shoots
arising low down and often creeping along the ground. Both
the erect and creeping shoots produce similar terminal spikes.
It is important, however, to bear in mind that the erect form
is with us much the more common, and that the plant is
usually found in open woods or along the road sides. Within
the past few years Brunella has invaded certain shady lawns
in the eastern part of Pennsylvania, and from the fact of its
rapid multiplication by rooting laterals is becoming a most
serious pest. The flowering shoots in these lawn specimens
have been much reduced in length, so that the flowers are
seldom raised more than two inches above the ground ; more
frequently they are on the ground. The same peculiarity has
also been noticed in the common dandelion when in similar

situations.

From a consideration of all the facts, the case of Bru-
nella vulgaris appeared to be an illustration of the prompt
action of natural selection in producing a variety of the
species which could perpetuate itself in spite of the lawn-

mower.

Within a year this special form of low-flowering and low-
seeding plant has been noticed in many places, and has occa-
sioned much surprise from the striking deviation it offers
from the type of the species.

But the most remarkable feature of the problem is the
promptness with which it seems to have appeared. Its

Rothrock.—A Nascent Variety of Brunella Vulgaris, L. 65

adaptation to the situation is most decided. The question
still may be considered an open one as to whether it has
developed as suddenly as it appears to have done.

Brunella vulgaris varies greatly in England in height, shape
of leaves, etc. It has long been known here in meadows and
open places, but nowhere, so far as my observation goes, has
this low-flowering variety been found in quantity save in
lawns which are frequently and closely "mowed off."

The late Dr. Darlington, who was a close observer, speaks
of the plant as common in meadows, but expressly declares
that it is not pernicious. The low form, however, is distinctly
pernicious on the shaded lawns of eastern Pennsylvania. A
point to be decided is whether it could have spread so rapidly
as it has done but for the disadvantage under which the
shade has placed the grass.

There is the further fact to be noticed that this variety, on
the whole, produces fewer flowers and consequently fewer
seeds than the typical form; and in like measure that its in-
crease by shoots rooting from the nodes is correspondingly
greater. The plants produced in this manner are so numerous
as to form dense mats. This is interesting, not only because
it illustrates forcibly the relation between diminished repro-
duction by seed and increased bud or shoot reproduction, but
because it in part explains the rapidity with which the variety
seems to have been developed. The chance of return to the
typical form has been reduced to a minimum by growth from
buds instead of from seed.

!l

Preliminary Observations on the flovements

of the Leaves of Melilotus alba, L.

and other Plants.

By W. p. Wilson, D.Sc,

ASSISTED BY

Jesse M. Greenman.

T

{WITH PI. ATE S IX, X, XI, XII AND XIII.)

'HE leaves of many plants may, under different condi-
tions, take three distinct positions, each one of which
may be assumed to give some advantage to the plant.
These three positions may be designated as (i) a nonna/ day-
light position ; (2) a hot S7in position ; and (3) a night position.
Nearly all the genera of LegmninoscB furnish examples which
may take any one of these positions when the surroundings
are favorable. Many other widely separated families of plants
furnish scattered illustrations of the same movements and
changes in the position of leaves. Oxalis, Pyrus {Americana
Del Sambucus, Rhododendron, Croton, Myriophyllum, and Mar-
silia give examples sufficiently separated to show that family
relationships have nothing to do with it. In some of these
the hot sun and night positions are the same. This is illus-
trated in the genera Oxalis, Myriophyllum and Marsiha.

In the sleep of plants, or night position, the leaves are inva-
riably so disposed or folded together as to lessen the leaf sur-
face displayed in the daytime. Darwin came to the conclu-
sion, supporting it with interesting experiments, that these
night positions lessened the surface exposed to radiation and
thus protected the plant from cold.

I wish to emphasize the fact that the hot sun positions,
over 200 of which I have already examined, occurring so fre-

Melilotus alba^ L. and other Plants.

67

quently at high altitudes, in strand vegetation, and in tropical
and desert areas are special adaptations to prevent a too
rapid transpiration.

In these hot sun positions the plant accomplishes the object
of lessening the leaf surface in many different ways. It may
be by folding the two halves of the leaves together as in Cro-
ton glandulosum, by rolling the leaves up as in Rhododendron
Catawbiefise, or by folding the leaflets over each other as in
Marsilia, or by elevating all the leaflets so that their apices
point directly at the sun as in Apios. In this last genus the
leaflets become parallel with the sun's rays and thus make
little or no shadow. In these different ways the leaves
escape much of the heat caused by the sun's rays, and, con-
sequently, increased transpiration which would be created
thereby.

Melilotus alba (along with many other genera of Legunii-
noscB, such as Robinia, Wistaria, Amorpha, Phaseolus, Amphi-
carpcea, Gleditschia, Cassia and others), is extremely sensitive
to its surroundings, and very readily puts itself, in accordance
with external conditions, into any of the three positions men-
tioned above. (PL XIII, Figs, i, 2 and 3.)

The day position may always be seen early in the morning.
The leaves are spread out in a plane at right angles to the
sun's rays. (PI. XIII, Fig i.)

The night or sleep position (PI. XIII, Fig. 2, and PI. IX,
Fig. i) is a very interesting one in this plant. From the
normal day position the leaflets first sink down so as to make
an angle of about forty-five degrees with the surface of the
earth below. The terminal leaflet rotates itself on its long
axis, either to the right or to the left, through an angle of
ninety degrees ; the lateral leaflets now each rotate on their
long axis until their edges are toward the zenith. They then
approach the terminal leaflet until their upper surfaces nearly
or quite touch it. In this position the terminal leaflet has
both its faces nearly covered, and the lateral leaflets present
their under surfaces only to the external air. A little over
one-third of the normal leaf surface is exposed. The torsion
takes place in the pulvini of the minor leaf petioles. Thi^

\

68 Wilson and Greenman. — Movements of the Leaves of

night position is assumed in a different manner by nearly
every genus of Legnminosce. Nor do the leaves always sleep
in the same manner in a given species. In the plant we are
examining, the very young leaves sometimes elevate them-
selves on the general petiole, instead of sinking down, and
then bring the apices of all three leaflets up together to the
zenith until they touch each other. If the young leaf hap-
pens to be at, or near the growing bud, the leaflets will often
rise up and encircle it at night. Darwin ^ has called attention
to the fact that the sleep of the young leaves of certain plants
resembles the sleep of the adult leaves in other genera. This
may indicate relationships of descent not otherwise easily
seen.

From the night position, the leaves as daylight approaches
gradually change into that of the day, in which the plane of
the leaves are generally so placed as to receive the rays of
the sun at right angles to their surfaces. Most of the leaves
are quite as low below the horizon as in their sleep positions,
but their faces are turned broadly to the light.

On a cool day, with atmosphere nearly saturated with mois-
ture, the leaves may retain this relative position to the sun's
rays (PL IX, Fig. 2). If, however, the air and soil are dry and
the sun hot, the leaves quickly take another position ; the gen-
eral petiole becomes slightly elevated and the leaflets rise up
above the plane of the horizon. By eight o'clock in the morn-
ing we may have the position indicated in PI. IX, Fig. 3. At
nine o'clock the leaves will have still further changed. The
leaflets will now be parallel with each other and also with the
sun's rays. PI. XII, Fig. i was photographed at nine o'clock
in the morning on a hot, dry day. The leaflets have all taken
a position parallel with the incident ray, and therefore cast
the least possible shadow. In order to get a profile view of
the angles made by the leaves with the general direction
of the stem the camera was set south thirty degrees west of
the plant. The leaves continue to rotate with the sun, keep-
ing themselves parallel with its rays. At twelve o'clock the
leaves will point vertically up to the zenith, as exhibited in

J Movements of PJaijts, Eng. ed., 1880, p, 345.

Melilotus alba, L. and other Plants.

69

PI. IX, Fig. 4. In this case the photograph was taken directly
from the south. On a hot, dry day all the leaves still follow
the course of the sun up to as late as six o'clock, as seen in
PI. X, Fig. 4, photographed from the north at that hour.

The preceding will show what is meant by the hot sun posi-
tion of leaves. Of the many plants which may be classed
with Melilotus in possessing this movement, not all are as
extremely sensitive, for all do not possess sensitive pulvini,
which render such movements easy. Not a few plants move
their leaves into a single hot sun position, in which they re-
main during the heat of the day. Others give their leaves a
position which will cause them to receive the least of the
sun's rays at the hottest time of the day, and then remain
rigid from this time on. (Species of Chenopodinm^ SmilaXy
Laguncnlaria and others.)

It is generally supposed by plant physiologists and those
who discuss this question, that these movements are protect-
ive in character, and that they shield the chlorophyll from too
intense illumination. A few experiments with Melilotus alba
have been instituted to test this view, the author believing
that loss of water is the cause of these motions and not the
effect of light on the chlorophyll.

Experiment No. i was conducted by taking a given lot of
potted plants, dividing them into two sets, giving both sets
precisely the same conditions of light and heaty but watering
one liberally, while the other was allowed to become quite
dry. A plant from each lot was photographed from the same
direction, and at the same time of day. PL X, Fig. i shows
the one with insufficient water, while PL X, Fig. 2 shows
the one plentifully watered. Although the day was not a hot
one, yet the plant in dry soil has its leaves very well pointed
toward the sun, while the plant in wet soil has only here and
there a leaf, some of the younger ones, turned toward the
sun. The results here obtained are given on a very large
scale in a gravel pit behind the laboratory. Hundreds of
plants are growing both on the high, dry ridges, and also in
the much moister excavations below. The plants on the
ridges are very active in taking the hot sun position, while at

I

70 Wilson and Greenman. — Movements of the Leaves of

the same time those in the moister locations below place their
leaves so as to receive the sun's rays much more nearly at
right angles with the plane of the leaves.

Experiment No. 2.— Two vigorously growing plants were
placed near together on the lawn. Over one was built a glass
case, the other was left standing in the open air. The one in
the case was soon in a nearly saturated atmosphere. The light
was the same in each case. PI. XI, Fig. i shows the one in
open air; PI XI, Fig. 2l the one surrounded by glass. The
stomata were nearly all closed in the first case, while in the
latter most of them were open.

Experiment No. 3. — Three healthy plants were placed near
each other on the lawn. The first was allowed to stand free
in the open air ; over the second was built a double glass case
with two inches space between the walls, filled with alum
water to absorb the heat rays of the sun ; over the third was
built a single glass case, which was packed with ice in the
interior to keep the temperature down. On an extremely hot
morning at nine o'clock all three plants were photographed.
The temperature was about 8° C. lower under the alum water
and in the iced box than around the plant in the open air.
The conditions of light were as nearly as possible the same
for all three plants. PI. XII, Fig. 1 shows the one in open air,
photographed, as are all the others, from the south thirty de-
grees west. The leaves are all pointing rigidly toward the
sun. They make in this way the least possible shadow, or,
in other words, expose the least possible surface to the sun
and receive less of its heat rays. In this way transpiration
becomes greatly lessened. The stomata are all closed with-
out exception. PI. XII, Fig. 2 shows the plant under alum
water. The leaves are not nearly so elevated as in the first,
many of the old ones being so disposed as to receive all the
light they can get, while only the very young ones are well
elevated and pointed toward the sun.

In the third one under glass and packed in ice (PI. XII,
Fig. 3) the leaves are somewhat more elevated than in the
last, but not to be compared with the one in open air. The
temperature was a little higher than the one under alum.

Melilotns alba, L. and other Plants.

71

This may account for some of the difference. The stomata
were well open both in the plant under alum and in the one

packed in ice.

PI. X, Fig. 3 shows a photograph of a plant taken from
the west on a clear evening at six o'clock, after a shower.
The lower leaflets have turned on edge, and others dropped
down vertically in order to receive the sun's rays at right an-
gles to their surfaces. It is to be compared with PL X.
Fig. 4, taken on an excessively dry evening at the same hour.
In the latter all the leaves are pointing directly west. The
photograph was taken from the north.

Experiment No. 4.— Four plants were placed on the lawn
and covered with red, blue, opaque or black, and white glass

respectively.

The one under blue glass seemed quite as active in the
movement of its leaves as the one under white glass. The
one under red glass lost very soon nearly all of its directive
motion from the sun, and its leaves made a slight advance
toward a sleeping position. The sleep position was assumed
nearly two hours earlier under the red glass than under the
blue or the white. It was also found that plants under red
glass put their leaves in a very different sleep position from
plants under either blue or white glass. PI. XIII, Fig. 8
shows a leaf under red glass, and Fig. 2 the normal sleep posi-
tion for comparison. In the former the leaflets are passed
down beyond the vertical, then twisted at the pulvini until
their under surfaces are all uppermost. PI. XII, Fig. 4 shows
a complete plant photographed at twelve o'clock at night from
red glass. PI. IX, Fig. i was taken from white glass at the

same time.

The plant under opaque or blackened glass lost m a short
time its power of motion as stimulated by the sun, and soon
went into a semi-sleep position and there remained with some
few undulations of motion, which, however, grew less and
less and soon disappeared.

PI. XI, Figs. JLand 4 show photographs from blue and red
glass taken on a rainy afternoon at three o'clock. The plant
from blue glass with leaves outstretched is trying to get all

)

72 Wilson and Grccninajt. — Movements of the Leaves of

the light it can, while the one from red glass is putting itself
rapidly into its sleep position. The leaves under the arrows
have already tucked themselves up in the peculiar manner
previously indicated in the sleep of this plant under red glass,
shown in PL XIII, Fig. 8, and in PI. XIJ Fig. 4.

The red glass must cut off certain rays of the spectrum
which serve to determine the ordinary sleep positions. No
difference in the night positions could be found under the
blue or white glass.

The following conclusions seem obvious from the work
done and the mass of material under observation :

(i) That there are great numbers of plants which put their
leaves in a special or hot sun position.

(2) That these Jiot sun positions have come to exist in order
to protect the plants possessing them from a too rapid trans-
piration.

(3) That these hot snn positions are not dependent on light
alone, but that the heat rays play a very important part in
determining them ; and also that the water supply of the
plant, both in the air as well as in the soil, exercises a direct
influence.

(4) That for some reason, not yet well understood, the
leaves of Melilotns alba take a different position at night
under red light from the one ordinarily assumed in the so-
called sleep of this plant.

DESCRIPTION OF PLATES.
Illustrating paper by Dr. Wilson and Mr. Greenman.

Plate IX.

Fig. I. Ordinary sleep or night position of leaves of Melilotus alba.

Fig. 2. Day position of the leaves.

Fig. 3. Leaves slightly elevated. First state of the hot sun position.

Fig. 4. Hot sun position at twelve o'clock on an extremely hot and
dry day. The leaves are pointed directly at the sun. They make the
least possible shadow in this position. The transpiration caused by both
heat and light is reduced to a minimum.

[Vol. I. Plate TX.]

Bot. Cont. Univ. Pennsylvania,

Fig. I.

Fk;. 2.

Fig. 4.

Wilson on jMkulotus.

[Vol. I. Plate X.]
Fig. I,

Bot. Cont. Univ. Pennsylvania.

Fig. 3.

Fig. 2.

il

Fig. 4.

Wilson on Mklilotus.

[Vol. I. Plate XL]

Bot. Cont, Univ. Pennsylvania.

Fig. I

Fig. 3.

Fig. 2.

Fig. 4.

Wilson on Mklilotus.

[Vol. I. Plate XII.]

FiCx. I.

Bot. Cont. Univ. Pennsylvania,

Fig. 3.

Fig. 2.

Fig. 4.

Wilson on Mklilotus.

[Vol. I, Plate XIII.]

Bot. Cont. Univ. Pennsylvania,

Is^ 8

Wilson on Melilotus.

Maize: A Botanical and Economic Study*

By John W. Harshberger, Ph.D. (Univ. of Penna.),

Instructor in Botany, University of Pennsylvania.

r

{WITH PIRATES XIV, XV, XVI AND XVII.)

CHAPTER I.

Botanical.

^OUR hundred years have passed since Columbus made
his celebrated voyages, and carried back to Europe
many strange plants and animals from the new world.
Maize seems to have been one of the plants which the Great
Navigator showed to Queen Isabella on his return to Spain.
Many prominent botanists, however, assert that maize is indi-
genous to the Asiatic Continent and the Eastern Archi-
pelago.

De Candolle, in his " Origin of Cultivated Plants," says :
"The certainty as to the origin of maize will come rather
from archaeological discoveries. If a great number of monu-
ments in all parts of America are studied, if the hieroglyph-
ical inscriptions of some of these are deciphered, and if
dates of migrations and economical events are discovered,
our hypothesis [Nicaraguan origin] will be justified, modified
or rejected." The following is a contribution to that end.

A. Gross Anatomy.

Culms several from the same fibrous root, ascending,
branched; internodes alternately furrowed. Plant five to
eight feet high.

Foliage ample ; leaves broad, long, tapering to an acumi-
nate point, horizontal, tip pendulous ; ligule short, hyaline,
ciliate.

76

Harshberger. — Maize :

Flowers monoecious, proterandrous, sometimes synacmic :
Male inflorescence, a panicle of spikelets, terminal on central
stalk, and its side branches,' branches pendulous; male
spikelets two to four to each joint, one or more short-pedi-
celled, two-flowered, flowers sessile ; glumes sub-equal, herba-
ceous, ciliate, sub-acuminate, concave, three- to five-nerved,
\yiZ2X\vi^\.^\ flowering glumes two, hyaline ; /^/^/^ two, hya-
line, concave; lodiculce two, cuneate, truncated obliquely.
Female inflorescence axillary, spicate, branched at times, with
a number of perfect ears on each branch. The spikes are
fasciated into a continuous spongy cob, so that the ripened
ear breaks readily at any point into its several joints, each
bearing two opposite pairs of kernels; ears two to six
inches long, three-quarters of an inch broad, with two,
four, eight to ten rows of kernels; spikelets many, imbricated
on a cylindrical rhachis, spikelets paired in alveoli, strongly
margined and cupulate, the margin becoming hard and
corneous; female spikelets two-flowered, with outer ones
neutral; glumes membranaceous; palets membranaceous,
concave, glabrous ; squamulce and stamens none ; ovary
slightly stalked ; grain white, hard, corneous, smooth,
ovate, pointed, constricted at the base, three- eighths of an
inch long ; style terminal, compressed, pubescent with com-
pound hairs, filiform, point bifid. The grain belongs to the
race of soft corns {Zea amylacea, Sturtevant).''

The relation of Zca to nearly allied genera, in the tribe
MaydeyE, becomes intelligible on examination of the Mexican
plant collected by Professor Duges. The plant may represent
the original wild form, or may be the reverted form of an
agricultural variety, but the latter supposition seems a highly
improbable one. The grains in the Mexican plant are placed
in alveoli, the margins of which are hard and corneous. AH
the genera in the tribe MaydEvE have the grains enclosed in a
hard, stony case {Pariana, Coix, Polytoca, Chionache, Schler-
achne, Tripsacum, Euchloena\ Tripsacum and Zea show

1 The illustrations in Plates xiv and xv show the side branch in a compacted form
♦he terminal tassel has not developed in the plant represented.

« Sturtevant, New York Agric. Exp. Stat. Rep., 1884, 124 ; 1886, 64.

A Botanical and Economic Study,

77

interesting similarities. The female spikes in maize, in all
probability, are fasciated into a continuous cob, and when
ripe the ear has a tendency to break into joints or pieces.
The spikes in Tripsacufn are axillary and terminal, separating
spontaneously at maturity into joints. The pistillate spike-
lets, two-flowered, with inner flower fertile, the outer flower
abortive, are imbedded in an oblong joint of the thickened
rhachis, occupying a boat-shaped recess, which is closed by the
cartilaginous outer glume. Zea has two spikelets, spikelets
two-flowered with inner flower fertile, the outer aborted,
placed in a cucullate depression of a fleshy cob. It seems
that the fleshy cob has been formed by the union of several
distinct spikes ; this conclusion is strengthened on compar-
ing Zea with Euchlcena and Tripsacum, for in the two latter
genera the joints are trapezoids, and easily disarticulated,
with the fruit set in a cartilaginous capsule, forming a
false fruit. A study of depauperate ears supports this
view. A bifurcation of the tip frequently occurs, when
the rhachis is prolonged into two axes. The tissues some-
times separate sufficiently to show the different spikes which
compose the fleshy cob. The arrangement of the grains
corresponds to the separate spikes of the consolidated cob.
These structural and teratological arrangements point to the
probable union of several spikes into a thick, fleshy axis, with
grains on the circumference, each paired row limited at the
side by a long, shallow furrow, a row corresponding to a single
spike of Euchlcena or Tripsacum.

The branch with alternate arrangement of ears (Fig. 9,
Plate XV), seems to be the more primitive, for cultivated
forms with one ear enormously developed have frequently
two or three ears placed in the axils of husks enclosing the
larger fertile ear.' One ear, in the cultivation of corn for
centuries, has enlarged at the expense of the others, furnish-
ing another illustration of the law of compensation in growth.'

Professor Duges found this corn called by the natives "maiz

1 Cornell Agric. Exp. Stat., Bui. 49. Dec, 1892, P- 333 See Bibliography, end
Section A.

2 St.-Hilaire called it "correlation of growth."

78

Harshberger. — Maize :

de coyote," at Moro Leon (otherwise Congregacion), about
four Mexican leagues north of Lake Cuitzco, on the boun-
dary line between the States of Guanajuato and Michoacan.^
The Duges plant, raised at the Cambridge Botanical Garden,
is probably the same as that found by Dr. Roezl, in 1869, in
the State of Guerrero, and described as a plant with ears
very small, in two rows, truly distichous, the grains small and

hard.^

Wild maize must have ready means of seed dissemination.
The cultivated forms would disappear, if man did not sow the
kernels, for the grain is too large to be carried by the winds,
and the sheathing husks prevent animals from reaching the
ripened achenes. The grain is smaller than the ordinary culti-
vated varieties, but large and wholesome enough to attract wild
beasts and birds, and in all probability this is one of the ways
in which corn was distributed. The following observation is
to the point:' "Yesterday, while at work, I saw a flock of
chickadees (Parus atricapilhis L.), one of which I saw had
something in its mouth, which upon inspection proved to be
a kernel of sweet corn. He was on an apple tree when I
first saw him, apparently trying to find a storehouse, but fail-
ing flew to a board fence, and running along found a split,
where he deposited it." Several birds have taken their names
from their liking for corn. An American blackbird, of the
family Icterid^, one of the marsh blackbirds, is fond of Indian-
corn, and devours it greedily. P. L. Sclater calls Pseudoleistes
viresccfts the South American maize bird, and Wilson desig-
nates Agalceus phceniceus the maize thief. The Indians may
have learned the use of maize from the wild animals, for
Professor Otis T. Mason refers to the fact that half-starved
Indians robbed the stores of nuts and corn hoarded by the

animals.

The original form, in the wild state, was propagated prob-
ably by lateral offshoots. All the cultivated forms produce
suckers. "The [Mexican] plants began to grow vigorously

» Watson, Proc. Amer. Acad. Arts and Sci., xxvi. 158.
2 Brewer, New York Agric. Soc, 1877-1882.
8 American Entomologist and Botanist, 11, 370.

A Botanical and Economic Study.

79

and to send off numerous offshoots from the base. These
suckers grew as rapidly as the main stem, so that the plants,
which had been placed fortunately some feet apart, had the
appearance of two hills, one of the two having nine and the
other twelve stalks ascending from a common base." ^

The production of suckers on annual plants in the north,
was probably a perennial habit in a more southern latitude,
so that in the semi-tropics the non-sexual development of
suckers was the ordinary method of propagation, the vigor
of the stock being rejuvenated by an occasional distribution
of seed by birds.

BIBLIOGRAPHY.

1. Ascherson, Ueber Euchloena Mexicana, Schrad. Verhand. des Bot.

Ver. Prov. Brand., 1875-76.

2. Ascherson, Die botanische Verwandschaft des Mais. Magyar.

Novent. Lapok. Klaus., 1877, 19- Anger's Bot.
Zeitschr., 1877, No. 2.

3. Ascherson, Bemerkungen uber astigen Maiskolben. Sitz. d. Prov.

Brand., XXI, 133.

4. Reibisch, Ueber Maiskolben mehrfach entwickelt. Isis, 1875, 29.

B. Histology.

Maize is easily grown in all its stages, and it serves, there-
fore, as a type specimen with which other monocotyledons
can 'be compared. A complete histological description is

desirable.

;e^^^._janczewski, in his classification of angiospermous
roots, places the root of Zea in his second category defined
by the general tissue differentiation: "Sharply marked
plerome cylinder and calyptrogen layer. Between the two,
at the apex of the punctiim vegetationis, is an initial group
only one layer of cells thick, which splits immediately behind
the apex into periblem and dermatogen {i.e., cortex and
epidermis)." Most monocotyledons which have been inves-
tigated agree in this.^ A longitudinal section of the root
apex shows the following arrangement of tissues. The root

» Watson, Proc. Amer. Acad. Arts and Sci., xxvi, 158.
2 De Bary, Comparative Anatomy, 10, Fig. 3.

So

Harshberger. — Maize :

cap of loose dead cells is constantly renewed from active lay-
ers within, so that the cap can be divided into two portions,
the outer dead, the inner active layers. The dermatogen,
continuous with the epidermis or piliferous layer of the root
above, is covered near the apex by the actively dividing
calyptrogen. The periblem occupies an intermediate posi-
tion between the dermatogen or proto-epidermis and the
plerome cylinder inside. The three meristematic layers, der-
matogen, periblem and plerome, take part in the construction
of the root, while the root cap is added as a fourth protective
element at the growing point. The central bundles are
formed from the plerome ; the calyptra is formed from the
calyptrogen.

A cross section of the primary root shows the tissues in a
different aspect. Two regions are defined as the central
axis cylinder and the encircling parenchymatous tissue. The
cylinder's centre is occupied by woody elements with radiat-
ing arms reaching the pericycle ; the phloem portions alter-
nating with the xylem wings. The bundle system is a radial
one. Large dotted or scalariform ducts are prominent
elements, readily identified. Spiral tracheids are found
between the large ducts and the smaller external annular
tracheids. The sieve portion of the phloem is obscure, except
when the wood elements are reduced to a minimum. The
cell walls of the pericycle show scleroid thickenings. The
endodermis is distinguished by its position on the circumfer-
ence of the axis cylinder. The remaining parenchyma is
irregular with large intercellular air cavities. The primary
root soon disappears, as in monocotyledons generally, and is
replaced functionally by the secondary roots, which differ in
the arrangement of some of the more important elements.

The layers in the secondary root are arranged as follows
(Fig. I, Plate XVI) : The large vessels of the bundle are
uniform in size ; the annular ducts and spiral tracheids have
increased until they occupy a large part of the space between
the pericycle and the wide dotted vessels ; the phloem areas
are small (Fig. i, P); the pericycle is formed of irregular
elliptical cells with long diameter anticlinal ; the endodermis

A Botanical and Economic Study,

8l

differs somewhat from that of the primary root in havmg the
rear walls of the cells crescentically thickened. The paren-
chyma is in regular rows outside, but near the epidermis it
becomes irregular and consists of loose cells.

The secondary roots arise from the stem, and in their later
origin are aerial. They appear at first as nodules, which
grow larger until the epidermis is finally ruptured by the
emergence of the rounded tip of the root. The root is posi-
tively geotropic, and grows downward into the soil. Before
entering the soil, however, gum is formed in large quantity
on the tip, and is of thick, treacle-like consistency. In the
aerial roots of palms, similar gummy matter is formed by a
breaking down and enormous swelling of the external cell
walls Cell walls which have undergone this mucilaginous
modification take, when placed in water, the consistency of
gelatin, and when warmed appear to dissolve, forming thick
mucilage. It is apparent by analogy that the secondary
roots in Zea form a mucilaginous covering by a change m
their external cell walls. The formation of these roots is

as follows :

Fig. 2, Plate XVI, shows the developing secondary root
before the point breaks through the epidermis and hypoder-
mis Three superimposed hollow cones are found immedi-
ately beneath the two outer protective layers, the outer and
middle cones being separated by a cushion of parenchyma^
The outer cone is composed of actively growing cells with
the nucleus and nucleolus plainly visible. It corresponds evi-
dentlyto the calyptrogen layer of Janczewski. The mner
cones correspond to the periblem and plerome cylinders with
the outer layer of the periblem as the dermatogen or proto-
epidermis. The cells of the plerome, destined to form the
central vascular system, are much longer than broad, the

lone: axis anticlinal. . . j^

i/m.-Comparison of an upper and a lower internode
of the stem shows the following differences :^ The upper in-
ternodes have a larger number of small peripheral bundles
than the lower, and the epidermal layers have comparatively

1 Strasburger, Botanische Prakticum, 109-

82

Harshbergcr. — Maiae :

A Botanical and Economic Study.

83

thicker cell walls. The bundles of the stem run parallel
along the internodes without intercrossing. A typical
central bundle is chosen in describing the closed collateral
bundle of maize, because at the surface the bundle elements
are reduced to a minimum, and because here a union
frequently occurs between two or three bundles. The sur-
rounding layers of the bundle consist of strongly thickened
sclerenchyma, the external and internal cells having thicker
walls than those on the lateral faces. A large intercellular
space surrounded by comparatively thin-walled cells attracts
the attention. An annular tracheid outside, and a spiral
tracheid between two large reticulated vessels are elements
forming the wood, xylem, hadrom or vascular portion of the
bundle. External to it we have the bast, phloem, leptom
or sieve portion of the bundle. The phloem portion colors
a bright violet with chlor-iodide of zinc. The first elements
of importance are sieve tubes with " companion " nourishing
cells. The proto-phloem {Cribral primanen), or actively
growing leptom, is found in this area.

The bundles at the periphery of the stem are crowded
together, so that the large intercellular space disappears ; the
phloem portion is also reduced. The bundle sheath, which
alone remains to any extent, is continuous with a layer of
thick-walled cells, called by Strasburger the hypoderm. The
hypoderm and bundle sheath function as protective and
strengthening tissues, and are elements in the mechanical
system of the plant.^ The separate cells of the hypoderm
have been called stereids, and the whole tissue taken together
has been called a stereome.

The epidermal cells in a radial longitudinal section are
longer than broad. The ordinary parenchyma cells are round,
or nearly so. The cells of the bundle sheath are long, with
contracted lumen. The intercellular passage follows the
length of the bundle without a break. The annular tracheid at
the inner border of the space is one of the most characteristic
elements of the stem. The large dotted ducts to the right
and left are shown in the section figured. Between the two

1 Pfitzer, Pringsheim's Jahrb., Bd. vni.

large vessels a spiral tracheid is occasionally found. Cells
with reticulated markings occupy the area between the larger
dotted vessels. The phloem appears as a bright area with
sieve tubes and companion cells, while the quantity of bast
increases or diminishes in different areas, but is found in
largest development in the interior of the stem.

X^rt/.—The leaf is divided into sheath and blade, which are
separated anatomically as well as morphologically from each
other by the ligule. A partial section across the base of the
sheath is shown in Fig. 3, Plate XVI. Examining the upper
inner surface first, we find the stereome immediately below
the epidermis. The stereome breaks up into discontinuous
patches near the overlapped margms of the sheath. The
outer lower surface is essentially different. The superficial
bundles are covered by an epidermis, the cells of which are
strongly thickened on the outer wall for protection against
extremes of heat or cold. Stiff, thick-walled, unicellular
hairs, associated with the long, narrow guard cells of the
stomata, no doubt serve the same purpose. The larger,
deeper-lying bundles are normal. The smaller, superficial
ones have a less number of cells, noticeably bast cells. The
smaller bundles are at places near the margin surrounded by
a circle of active parenchyma cells, which contain chlorophyll,
and called by De Bary and Sachs the starch sheath, or starch

ring.^

The upper inner midrib area of the blade at the base shows
five or six layers of sclerotic cells (stereome or water tissue)
continuous for some distance on each side of the middle Ime ;
near the margin, however, the strengthening layers are dis-
tributed in discontinuous patches with reference to large cells
between the so-called bulliform cells. Near the tip of the leaf,
the fibro-vascular bundles alone strengthen the leaf ; the stere-
ome disappears. The swelling of the bulliform cells (Fig. 4,
Plate XVI, B) by imbibition of water causes the blade to open
out in those leaves which are folded in the bud. They are
found only on the upper side of the blade on each side of the

1 De Bary, Comparative Anatomy .416; Sachs, Botan. Zeitg., 1859, i77.Taf. viii,ix ;
Pringsheim's Jahrb , iii, i94-

\

84

Harshberger.— Maize :

midrib. According to Duval Jouve,' these cells have thin
walls and watery interior, and are found on the upper surface
between the marginal nerves of the leaf. In grasses which
are devoid of such fan-shaped cells, the blade always remains
folded up, rush-like, as in Stipa, Festtica, Nardiis. In steppe
grasses, the blades roll up whenever these cells lose their
turgescence by rapid evaporation, and they open again when
the^'water has been restored. The lower side of the leaf has
especial protective arrangements against transpiration, m
strong cuticle and hypoderm.'

The lower surface of the blade shows a larger number of
small bundles than the lower surface of the sheath. Either
the fibro- vascular bundles enter the blade separately, or a
number at first unite to form a strong midrib ; later the single
bundles separate one at a time and pass toward the edge of
the leaf, giving firmness to the broad lamina. A ring of
parenchyma surrounds the small bundles (Fig. 4, Plate XVI).
The stomata are placed on surface view between two long
cells called accessory cells, which were formed when the
stomata were differentiated. The process of stoma formation
in Indian corn is essentially as follows. A vertical septum is
made across one of the long epidermal cells, the small cell
cut off being called the stoma mother cell. The two acces-
sory cells are now cut off from the mother cell on either side,
and the mother cell divides in two, forming the guard cells-
Finally, the lamellae of the wall separating the guard cells split
and the opening is formed.=^ According to von Mohl,* the
stomata on the uninjured leaf of Zea jnays widen the slit to

y|^ millimeter.

' Mention must be made of gaps in the epidermis of the leaf.
Cracks occur regularly at the apex of the leaf, from which
drops of water are expressed. The cracks arise by irregular
tearing of the original cowl-like apex of the leaf, when this
spreads out flat as it unfolds.

> Duval Jouv^, Histoire des Feuilles des Gramin^es, Ann. d. Sc. Nat., vi s^r.,tom i,

1875, 294, tab. 4.

- Hackel, True Grasses.

3 Campbell, D. H , American Naturalist, xv, 1881, 764.

< Mohl, H. von, Botan Zeitg., 1856, 697.

A Botanical and Economic Study,

85

Flower,— T\i^ style is long, reaching beyond the tip of the
enclosing husks. It is double morphologically. Two fibro-
vascular bundles, greatly reduced, run its length. The style
of Zea agrees with the styles of grasses in general. The
entire surface of the filamentous style is covered with com-
pound hairs, which catch and hold the smooth pollen grains.

The round, smooth pollen grains (Fig. 4, Plate XV) are pro
duced in great abundance, as many as 2500 being formed in a
single anther. " Each panicle of male flowers [the tassel] was
found by careful estimates to contain about 7200 stamens, so
that the number of pollen grains produced by each plant is
18,000,000. Allowing two ears of 1000 kernels to each plant,
there are still 9000 pollen grains for every ovule to be ferti-
lized."^

The epidermal layer of the grain is simple, with long
diameter anticlinal. The internal cells contain starch and
protein granules."' The starch granules closely aggregated
are polygonal, with extremely delicate circular and radial
markings. In size and general appearance, they are inter-
mediate between the starches of wheat and rice. They are
from .0002 to .0012 of an inch in diameter.'

BIBLIOGRAPHY.

1. Potonie.H., Entwickelung der Leitbundel Anastomosien in den

Laubblattern von Zea Mays. Ber. D. B. G., IV,

no.

2. Reibisch, Ueber Mais-Kolben. Sitzungsb. d. nat. Gesellsch. Isis,

1875, 29.

3. Delbruck, M., Des Gehalt des Maises an Starke, Centralblatt fur

Agric. Chemie, 1880, 155. Botanische Jahresbericht,
1879, 384.

4. Poisson, Sur la coloration des grains de Mais, Compt. Rend. Assoc,

fr. p. I'avanc. d. Sc. Paris, 1878, 688.

5. Kreusler, Beobachtungen uber das Wachsthum der Maispflanze.

Landwirthschaftliche Jahresbericht N. in Thiel.
1877, 7591787; 1879,517,617.

1 American Naturalist, XV, 1881. 1000. » ,. j uu,,«,

sSimmonds, P. L., Tropical Agriculture. 299 ; Meyer. Arthur, Archiv. der Pharm.,

231 0I2>

'» Bell, James, The Chemistry of Foods, 11, i73i London, 1891.

Mi.

86 Harshbefger. — Maize :

C. Bibliography^ Synonyms^ Names.

ZeA mats. LiNN/EUS.

1583. Dodoens, Stirpium historia, 509.

1591. Hernandez, F., Nova Plantarum Animalium et Mineralium

Mexicanorum Historia. Quarto, i65i,Rome,

1628, t. p. 242.
1623. Bauhin, Pinax theatri bot., 25 1 prumentum Indicum.
1658. " " " 490^

1694. Camerarius, Epistola, 186.
1729. MichelU Nov. pi. gen. 35; g. pi, Class XIV.

1745-
1748.

1749-
1751.

1753-
1759-

1663.

1764.

1767.

1769.

1774.

1777-
1778.

1784.

1788.

1789.

1790.

1791.

1797-
1799.

1802.

1805.

1807.
1812.
1815.
1817.
1818.

<<

((

(t

Ltnnaus, Gen., p. 279, n. 703; Hort. Cliff., 437 I Syst., 30.
Haller, Rupp Fl. Jen., 303.
Linncpus, Hort. Ups., 281.
Gleditsch, Berl. V., 123.
LinncEUs, Phil, bot., 28.

Species, II, 971.

Syst., ed. X, 1260, n. 926.

Ht. Trian. g. Gramin.

Sp. ed., II, 1378.
Adanson, Fam., II, 39.
LinnceuSy Gen. ed., VI, 480, n. 1042.
LimicEus, Syst. ed., XII., 615.
•' Hort. Kew., 406.
" par Murray Syst., 702.
Scopoli, Intr., 78 (spurior).
Reichard, Gen., 475, n. 1133.
Thiinberg, Fl. Jap., 37; g. Triand. Monogyna.
Gaertner, De Fruct. et Sem. Plant., VI, 6 t., i f. 9.
JussieUy Genera, 33.
Loureiro, Fl. Cochin., II, 550.
Necker, Elementa, III, 229; n. 1613.
Schreber, Gen., II, 621, n. 1043.
GmeliHy Syst., 148.
Persoon, Syst., 885.
Ventenat, Tableu II, p. III.
Batsch, Tabula., 156.
St.-Hilaire, Expos., I, 88.
Willdenow, Sp., IV, 200, n. 1636.
Per soon. Enchiridion, II, 533, n. 2029.
Beauvais, de, Agrost., 136, t. 24, f. 5 ; 20 t., f. 3 et 4-
Kunth, Mem. Mus., II, 75. g- Olyrear.
Sprengel, Aul., II, 183.
Nuttall, Genera, II, 203.

A Botanical and Economic Study. 87

1820. Trinius, Fund., 200.

1822. Link, Enum., II, 377.

1823. Loiseleur, Diet. V, 28, 103.
Agardh, Aphor., 153.

1825. Sprengel, Syst., I, 238, n. 242.

1827. Link, Hort., I, 253.

1828. Reichenbach, Conspectus, 55, n. 1059.

1829. Nees von Esenbeck, Agros. Bras., 311.
Link, Handb., I, 96.

Kunth, Gram., 12.

1830. Reichenbach, Fl- exc, 54.
Gaudin, Helv., VI, 2, 15.

1 83 1. Sprengel, Gen., II, 687, n. 3468.

1832. Roxburgh, Fl. Ind., 111,567-

1833. Kunth, Enum., I, 19.

Trinius, Mem., Ac. Pet., 6e, S^r. II. 78.

1836. Bonafotis, Hist. Nat. du Mais.
Endlicher, Gen. Plant., 80, n. 742.

1837. Koch, Syn., p. 769, ed. II, 889.

1838. ;^^tf^/.r, Linn. Trans., XVIII, 26.

1839. Dietrich, Syn., I, 131, 234.
1841. Reichenbach, Norn., 36, n. 1246.

1843. Meisner. Gen., 416.
Bongard, Eng., 11.

1844. Reichenbach, Fl. Sax., 21.

1846. Spach, Veg. Phan., XIII, 1172.

1847. Lindley, Vegetable Kingdom, 115.

1848. Koch, Linnaea, XXI, 376.

1849. Duchesne, Orb. Diet., VII, 595.

1854. Steudel, Glum., I, 9.

1857. Miguel, Ind< Bat., Ill, 477-

1878. Seringe, Monogr. des Cereales. 8vo, Berne.

Synonyms.

Generic: . r-vcc

1735. Thalysia, Linnsus' Syst. Nat. Fund. Bot, 244; Hort. Clitt.. 437-

1729. Mayz. Micheli, Nov. pi. gen., 36.
1763. Mais, Adanson, Fam., II, 39.
Mirbel.
Mays, Tournefort.
1808. Mays zea, De Candolle, Icones Plant. Gal. rar., 3, 98.

Specific :

Mays Americana, Baumgarten.
vulgaris, Seringe.
zea, Gaertner.

'^1

8S Harshberger, -Maize:

Zea alba, Mill.

altissima, Gmel., Hort., Carlsr.

Americana, Mill.

canina, Sereno Watson, 1890.

Caragua, Molin.

hirta, Bonafous.

minor, Gmel.

macrosperma, Klotsch., Bot. Zeit., 1851.

praecox, Persoon.

rostrata, Bonafous, Ann. de Lyon, V, 97.

The word Zea is probably derived from the Greek words
^a<o ** I live," or t^tia. (^ca), a sort of grain (spelt) applied as the
term for maize when ^ca was left unused by the creation of
the genus Tritictim. Scholars think that the Greeks first
called spelt okvpa, afterward ^cta, names which we find in
Herodotus (Hdt. 2, 36) and Homer (II. 5, 196; 8, 564).
Dioscorides (Diosc. 2, 113) distinguishes two sorts of ^cta,
which apparently answer to Triticuni spelta and T. mono-
coccurn.

Maize is an Arawak word of South American origin (see
Philology), from mahiz, marisi, mariky, etc. The name is
derived from the Haytian word mahiz, which Columbus
adopted when in Hayti.

Indian Corn.— Corn comprehends all the kinds of grain
used for the food of men or of horses ; in Scotland it is gen-
erally restricted to oats. In the United States it is applied
to maize. Those who first landed in America found a new
cereal used by the aborigines, and they naturally extended
the word corn to this grain also, specifically limiting its
use by the prefix *' Indian." In the United States, Indian
corn is called simply corn. This specific application has
been confirmed by a judicial decision in Pennsylvania, in
which it was ruled by the court that the word corn is a
sufficient description of Indian corn.

Names for Maize in Various Countries.

Abyssinia : Mashela bahry, /. e., millet from the sea.
Africa: Maheende (Central), mahindi, mas^ (Northern),
imitations of the word maize.

Belgium : Mays, Turkisch koorn.

Probably

A Botanical and Economic Study.

89

Brazil (Portuguese): Zabemo, milho de Guine, milho grande, maiz,
milho da India.

Burmah : Pyoung-boo.
Ceylon: Muwa.

China: Yii-shu-shu, jade sorgho, yuh-kau-ling.

Egypt : Doura shammy, dourah de Syrie.

Fiji : Sila-ni-papalegi.

Formosa : Fanmeh (foreign corn).

France : Le mais, bl^ de Turquie, bl^ d'Inde, bl^ de Rome (Vosges),
h\€ de Barbarie (Provence), bl^ d'Espagne (Pyrenees), h\€ de Guine.

Germany: Mais, maiz, Turkischer korn, Turkischer weizen.

Great Britain : Maize. Indian corn.

Greece: Arabosite (Arabian corn).

Holland : Tursch korn, mays.

India: Mokka, bhoot, muk-jowaree-boota.

Italy : Grano Turco, grano d' India, grano Siciliano.

Japan: Nan bamthbi (foreign corn), sjo-kuso, too-kibbi.

Malays: Jagang (indigenous).

Moldavia: Kuku-rusa.

Persia: Ghendum.

Portugal: Milho d'India, maiz, milho grande.

Russia: Tureskorichljeb.

Spain: Maiz, trigo de Turkina, zara, trigo de Indias.

Sweden : Turkish hvede, korn.

Turkey: Misrbogdag (Egyptian wheat).

United States: Maize, Indian corn, corn.

90

Ilarshberger. — Maize :

<(

<(

((

((

(<

(t

CHAPTER II.
Origin.

A. Meteorological Pi'oofs,

THE conditions most favorable to the development of the
maize plant are long summers with sunny skies, hot
days and nights, and sufficient rain to supply the
demands of the rapidly- growing crop. The following table
gives an idea of the best temperature for growth. With a
temperature of.

45°-50° the yield was 40.8 per cent, of total crop.

45°-55° " '' 75-9
45°-6o° " " 87.3

These figures refer to the year 1880.' The range of tem-
perature for the best development of the crop seems to be
between 45° F. {f C.) and 65° F. (18° C), calculated in
terms of mean annual temperature, but the largest returns
resulted with a temperature in the month of July corre-
sponding to from 75° to 80° F.; 961,123,938 bushels of the
1,754,861,535 bushels being the return in those regions having
the July temperature between 75° F. (24^ C.) and 80° F.
(26.7° C). The largest return was obtained in those localities
where the rainfall amounted to from thirty to fifty inches.

30 to 45 inches yielded 63.4 per cent, of total crop.

30 to 50 " " 86.8

The largest absolute yield corresponded to a rainfall in the
spring and summer months of twenty to twenty-five inches.
The return was 1,143,239,093 bushels of the total crop, i,754r
861,535 bushels. Frost kills the plant in all its stages, and
the' crop does not flourish where the nights are cool, no

1 Census Report U.S., Agricultural Productions, 1880.

A Botafiical and Economic Study

91

matter how favorable the other conditions.' With the varieties
grown in the United States, in ordinary culture, an elevation
of over 2000 feet shows the yield of maize to be very small
indeed.'^

Elevation and Yield.

500 to 1000 feet, 54 per cent, total yield.
500 to 1500 " 82
Above 1500 " 4.4

((

(<

((

(<

((

^^

This was not the case, however, with maize grown in
Mexico and Peru. Humboldt' mentions that 'Wast maize
fields are to be found on the plateau of Mexico at a height of
8680 feet, and in Peru, on the road between Lima and Pasco,
maize is cultivated as high as 12,000 feet (3824 m.)." Sim-
monds* states that it is raised in tropical countries at a height
of 9000 feet and more.

Professor Duges sent, in 1888, to the Cambridge Botanical
Garden, Boston, several maize plants, which he collected at
Moro Leon, otherwise Congregacion, near Uriangato, four
Mexican leagues north of Lake Cuitzco, and, therefore, near
the boundary line between the States of Guanajuato and
Michoacan. Grains from the Mexican plants sowed in 1892
in Philadelphia, in the middle of May, perished for some
reason, but one developed into a specimen about five feet high,
which showed great hardiness, the growing period extending
to the fourteenth of November, when the stalk was cut. The
plant stood well the frosty days, and a snowstorm which
came before it was cut did little harm. The exceptionally
dry autumn suited the plant well, which throve better in the

iDarwin, Variation of Animals and Plants under Domestication, 1,322.

2 The absence of large areas of tillable land in the United States at the elevation of
2000 feet explains this distribution of crops in altitude for Dr. William P. Wilson mforms
me that some of the largest and most luxuriant crops of corn he ever saw were raised on
the mountains of North Carolina at an elevation of over 4000 feet. ,^^18 exception to the
above statement enforces the view hereafter expressed as to the elevation at ^h ch Uie
wild corn grew. Broadly speaking, high vertical thermometric '■^'^^^^'^''^^^^^'^^^^
latitudes, low vertical zones to low latitudes ; the elevation (4000 feet) in North Carohna.
therefore, represents a much higher altitude in the tropics.

3 Meyen. Geography of Plants, Ray Soc, 1846, 304-
* Simmonds, Tropical Agriculture, 1877-

92

Harshberger, — Maize :

dry weather than the ordinary varieties, which were harvested
a month before. The so-called wild corn possessed greater
constitutional vigor than the ordinary cultivated forms.

The tables which follow, from the records of the United
States Signal Office in Philadelphia, may prove of value in
this connection. The first table shows the rainfall for the
month of September, 1892 :

Rainfall, September, 1892, Philadelphia. (Inches.)

Less than .01

01 to .10 .11 to .25 .26 to .50

.51 to I

Days.

II

September and October were very dry. The thermometer
stood for the thirty days of September above 59° F., a tem-
perature favorable for the plant's growth, but on only one
day did the rainfall exceed one-half inch. October showed
no days with the temperature below 32°. A comparison with
the October rainfall of the previous twelve years showed that
this was the dryest October in that whole period of years ;
0.30 inch of rain fell. The thermometer did not vary so
greatly ; the mean of the whole month was 56.4° F.

November was also dry until the fifteenth, when two
inches of rain fell. The greatest precipitation during the first
fourteen days occurred on the ninth and tenth days, when
.52 and .86 inch fell respectively. These figures show the
power that the possible wild species had of thriving well with
little rain, and are valuable as showing the conditions favor-
able to the growth of the plant in its Mexican home. Data
collected by the Mexican Observatory from numerous sta-
tions in central Mexico are compared with the Philadelphia
temperature and rainfall, as showing the localities in Mexico
meteorologically suited to the wild plant's growth.

The following table, compiled from the Boletin Mensual^

» Boletin Mensual Observatorio Meteorol6gico-Magnaico Central de Mexico, Resfi-
tnen del Ano, 1889, 37.

A Botanical and Economic Study.

93

gives the altitudes of the places, and the mean temperature
and mean rainfall for the months of April, May, June, July,
August, September and October, 1889:

Aguas Calientes
Guadalajara
Guanajuato .
Huezutla . .
Leon . . .
Mazatlan
Saltillo. . .
San Luis Potosi

Puebla

Zacatecas . .

!

Altitu

DE.

t.

Temperature.
Mean.

5590 f

2.1° C.

1

70° F.

4700

u

21.2° " i

70° "

6180

u

18.1° "

(,f -

II28

((

; 25.4^ "

78° "

5400

((

' 21.4' "

70° "

20

u

1 28° "

^t " 1

4900

((

19.9° "

68° "

5670

((

19.5° "

67° " 1

1

6520

((

1 17.6° "

64° "

1

7488

((

21.2° "

1

i 70° "

1

Rainfall,
Total.

it

K

72.1 mm.

1043
91.2

258.6 "

104.7 "

96.9 "

51.3 "

35.3 "

110.3 "

61. 1 "

2.2 m.

3-1

2.7

7-7

3-1
2.9

I

((

u

((

((

u

(i

((

u

3-3
1.8 "

The temperature of all the places is favorable to the growth
of the wild cereal, but the rainfall in Aguas Calientes, Saltillo,
San Luis Potosi and Zacatecas is so small that these places
are discarded, as being outside of the limit of the possible
original home of the wild form. Mazatlan, on the Pacific
coast, is also omitted, because it is hardly probable that corn
was a sea-side pl^nt. The other stations answer the require-
ments better. On consulting a map of Mexico, it is seen
that Guadalajara, Guanajuato, Huezutla, Leon and Puebla lie
below the twenty-second degree of north latitude, at eleva-
tions ranging from 1128 feet, that of Huezutla, to 6520 feet,
that of Puebla. From other considerations {ante), it is likely
that maize was a highland plant; Huezutla is therefore dis-
carded, as below the limits in altitude. The other places li«i
above a level of 4500 feet, and in all likelihood the original
form of corn grew at this altitude and higher. The area is
narrowed, therefore, to the region near Guadalajara, Guana.

94

Harshberger. — Maize :

juato, Leon and Puebla, and a glance at a map will show that
these localities are situated along the backbone of the conti-
nent, below the twenty-second degree of north latitude. A
curious coincidence in places must be mentioned. Leon,
5400 feet above the level of the sea, is identical with the
locality which Professor Duges explored. From the meteor-
ological facts we conclude :

(i) That maize was a highland plant ; (2) that the original
home was south of the twenty-second degree of north latitude ;
(3) that the Duges' plant in Philadelphia still preserved the
habit of withstanding dry, rainless weather, which the Mexi-
can charts show is the summer condition at Leon, its native
home.

/). Botanical Proofs.

It is a principle of geographical botany that the occurrence
of two or more species or genera of close relationship, in any
region, indicates the probable origin of those forms within
that area. This factor was of little importance to systematists
before the days of Darwin and Wallace, but the studies of
these two naturalists have served to throw light upon many
heretofore obscure problems.

There is tendency to vary, and variation proceeds along
definite lines ; in the mean time, certain forms disappear
from the evolutionary series, until it is broken into groups
representing new species and new genera of divergent char-
acter and form.

A clear relationship, in many particulars exists between
Zea, Tfipsaaim and Eiichloena ; and it is probable, therefore,
that Indian corn and these two latter plants originated from a
common remote ancestor, which grew in Mexico. Maize has
diverged most widely from the common type, in having the
separate spikes fasciated into a spongy cob.

The discovery by Professor Duges of the so-called wild form
in Mexico adds weight to the argument in favor of a Mexican
origin for the plant. The branching habit, the reproduction
of the plant by suckers, the small size of the grain, clearly
indicate a very primitive condition of the plant. With refer-

A Botanical and Economic Study.

95

ence to the offshoots, it is necessary for perennial growth of
a plant so sensitive as maize to frost, to live in a tropical
climate, where the suckers can be produced with the greatest
advantage to the plant.

Otis T. Mason believes that the original habitat of the
maize is to be found in regions where the grains remain
unharmed out of doors over the winter months ; for the grain
decays if left out in the fields in the United States exposed
to the destructive effects of the cold and ice. He belie^^es
that a semi-arid region answers the conditions most satisfac-
torily. The anatomy of the leaf points to the same conclusion,
for the lower epidermis is thickened to protect the leaf and
prevent too rapid transpiration, when the leaf is rolled up by
the loss of water in the bulliform cells.

The Maya civilization was not a growth of chance, but was
dependent for its origin on the surrounding circumstances,
propitious to such a development ; the physical and biological
features of the country influenced the settlement. The pla-
teau of Mexico has a favorable climate and naturally yields a
variety of products. On it grow the agave from the bud of
which a drink is brewed, cacao which furnishes the beverage
chocolate, the potato, the tobacco plant, the sweet sop {Anona
mnricata), papaw {Carica Papaya), nopal {Opuntia cochinelii-
fera) on which the cochineal insect feeds, guava and caoutchouc
{Hevea brasiliensis). The plants mentioned are all natives
of the warmer parts of North America, and it is likely that
maize also occurred in the same region in the wild state, and
was one of the first plants of economic importance to be cul-
tivated. The Mayas depended chiefly upon maize and honey
for their food, and there is hardly a doubt that the natives
were attracted to the warm plateau of Mexico, where the
country was so inviting and rich in natural vegetal products.
These facts point to the Mexican origin of maize.

C. ArchcBological Proofs.

The accounts are numerous as to the discovery of Indian
corn in places of undoubted pre-Columbian antiquity, but
they are general, and for the most part unsatisfactory. When

96

Harshberger. — Maize :

archaeological excavations have been made, no particular atten-
tion has been paid to the remains of maize, and the remarks
as to the discovery of the cereal in ancient mounds, barrows
and ruins have been, therefore, mostly incidental. The
attempt will be made in this section to sift carefully the evi-
dence which archaeology affords as to the cultivation of maize
in prehistoric times. For convenience, North America will
be surveyed first.

The mound-builders were described ten years ago as the
oldest inhabitants of the northern portions of the North
American continent. The discussion relative to the antiquity
of this race of men will be left to a following section ; but it
is well to notice, in passing, that they are now considered to
have lived comparatively recently.

By far the most celebrated unearthments were made near
Madisonville, Ohio. The terrain, in which the discoveries
were made, lies in the valley of the Little Miami, southeast of
Madisonville, where the space covered by the principal mound
was from four to five acres. Virchow^ says; " Jedenfalls halt
man es fiir pracolumbisch." He describes the surroundings,
that over the grave mound stood trees more than 300 years
old, particularly an oak, six feet two inches in circumference.'^
Diggings revealed 185 human skeletons, twelve pipes, three
of catlinite from Minnesota, stone rubbers, axes, lance heads,
needles of perforated teeth, two small cylinders of rolled cop-
per, two feet long, two plates of copper, and near by ash-pits
filled with ashes, and sand mixed with pipes, bones, mussels,
stone implements, the tooth of a mastodon, bones of the wild
ruminants (buffalo, elk, deer).^ At one place was found a
kind of sacrificial altar, where ashes were mixed with numerous
animal bones, deer, elk, opossum, turkey, weasel, woodchuck
and bear, evidently sacrificed to the all-powerful Manito.

1 Virchow, Zeitschrift fiir Ethnologic, 1879, 446.

* Cincinnati Daily Enquirer, 1879, April 24; Cincinnati Commercial, 1879, August 31;
Short. North Americans of Antiquity.

3 Notice the buffalo bones, as they throw light upon the antiquity of the Madisonville
mounds and surrounding structures in Ohio. Cf. Shaler, Nature and Man in America
183.

\

\

A Botanical and Economic Study.

97

Maize was found in quantities.' The account published in the
Journal of the Cincinnati Society of Natural History is worth
quoting : " On Tuesday, August 26, one of the most interest-
ing discoveries in this cemetery was made. In excavating an
ash-pit a large deposit of several bushels of carbonized maize
was found." Leaf mold, gravel and clay covered the hole to
the depth of thirty-nine inches, and contained animal remams,
implements of flint, stone and bone, and an unfinished pipe.

- 3 ft. 7 in. -

Leaf Mold

24 in.

---- 3ft.

Leaf Mold,

24 in.

Gravel and Clay.
15 in.

A.

Ashes with Animal Re-
mains, Fragments of
Pottery, Shells, etc.

-6 in.-
Clay.

Ashes, Animal Re-
mains, Shells, Sherds,
10 in.

B.

Bark Matting,
4 in.

Shelled Corn.

Ear Corn.
Boulders.

The second clearly defined stratum, in pit B, contained
deer, elk, raccoon, opossum, mink, woodchuck, beaver and
turkey bones, shells of Unio mixed with ashes, and potsherds,
to the depth of ten inches. The third layer was made up of
coarse matting, twigs, corn-stalks and bark, completely car-
bonized. The fourth contained shelled corn, probably three
or four bushels, and immediately below ear corn, completely
carbonized. The maize was turned over to Dr. Wittmack for
study. He describes'^ the ears as eight-rowed (in four double
rows).

t Verhandl. Gesellsch. Anthr. Ethn. Urg., 1881, «6; Witlmack, ^^^^^^''^^'^J:'^-
nologTe! xiMibSo, 85!; Journal Scientific Society. Madisonville. Ohxo ; Cmcmnat. Soaety

^'*« WUtmacll^erhandl. Gesellsch. Anthr. Ethn. Urg., ,88., a.6. Monatschrift f. Gar-
tcnbaues, i879.r54i«

98

Harshherger. — Maize :

The garden plots, so-called, of the mound-builders described
in earlier publications are of uncertain value.^ Locke' states
that from the Cincinnati mounds was obtained "carbonized
maize, with even the cob, leaves and stalk of the plant." In
the Museum of Archaeology, University of Pennsylvania, is
some charred corn found in a mound in Portage County, Ohio,
together with celts, gorget, arrow-points, clay pipe, copper
beads and charcoal. Mr. Stewart Culin informs me that the
specimens are authentic and undoubtedly pre-Columbian. A
quantity of Indian corn and fragments of the cob with the
grain still in place, and all very much charred, was sent to
Lucien Carr by William P. Bales and Rev. S. B. Campbell, of
Rose Hill, Va. They were found by drifting into the face of
the southern wall of the central shaft of a mound about fifteen
inches, and were on a level with the bottom of that shaft
about eleven feet from the surface.' Cyrus Thomas ' describes
the Etowah mound, on the Etowah River, a few miles south
of Cartersville, Bartow County, Ga. : " When the first white
man visited it, a stately forest covered the works, as well as
the area.*^ Although the Cherokees made use of it as a fort
against the Creeks, they always denied having any knowledge
of the race by whom the mound was constructed." •" In the
refuse layer, west and east of the three large mounds, four
feet below the present surface, were found some partially
burned corn-cobs. They were in a little heap surrounded by
charcoal. Proudfit ' states that in 1879 Mr. Stillman made an

1 Peet, S. D., American Antiquarian, vii, 15.

« Locke, Trans. Amer. Assoc. Geol., 1, 231 ; Trans. Chicago Acad. Sci., i, 242.

3 Carr, Peabody Museum, 11, 80.

* Thomas, Cyrus, Amer. Anthropol., iv, 109, 237-

5 Jones, C. C, Antiquity Southern Indians, 139; First description, Amer. Journ-

Arts and Sci. i, Ser. i, 322 (1814).

oThe studies of Dr. Seler are very significant in connection with the mounds ot
Georgia and the Ohio valley. He analyzed with care the mode of wearing the head-
dress, the clothing and weapons represented on the copper work of the Etowah mound-
builders in Georgia, and compared them with the figures on the shells from the Ohio
mounds. He concludes from his careful study that the Etowah builders and the Ohio
artists were in all probability related, and that possibly the mound-builders and copper-
working tribes were destroyed or driven to the sea-coast by the invasions of tribes [Iro-
quois and Algonquins] from the north and west, at a period not very remote from the
discovery of the continent. Seler, Globus. Bd. lxii. No 11; Science, xx, 260. See page

107.

7 American Antiquarian, ill, 278.

A Botanical and Economic Study.

99

interesting discovery. Mr. Stillman's attention was attracted
by ashes on the face of an exposure of a cut made for the
passage of the Mynster Springs ro.ad, one and a half miles
north of Council Bluffs, Mo. In company with Mr. Jacque-
min and Mr. Burke, he opened the face of the bluff, and found
what might be called a kitchen heap. The opening extended
into the hill four feet, and was five feet below the surface.
They found a fragment of an elk's antler, a shoulder-blade
fashioned into a rude agricultural implement, fragments of
bone, a pipe, a piece of deer's antler, flint scrapers, fragments
of pottery, a charred corn-cob, several large mussel shells, fish
bones, vertebral joints, and a stone paint-pot or mortar, of
rough quartzite.

At the advent of the Europeans, the Indians were found
cultivating the maize. That they practiced the first prin-
ciples of agriculture before that is shown by the remains of
agricultural implements of primitive form. When the whites
came, they gave up their clumsy instruments and adopted the
better and more lasting European tools. In a burial mound
near Illinoistown, opposite St. Louis, was discovered a stone
hoe seven and a half inches long, nearly six inches wide, and
about half an inch thick. The fastening was facilitated by
two notches. A similar but smaller hoe was found in a
garden in the city of Belleville.' McAdams' says that "the
majority of these ancient implements of husbandry were made
after definite patterns, each kind to be used for special pur-
poses, being similar to six of the deeply notched hoes." They
tilled the ground with hoes made of clam shells.^

In the southwestern United States, numerous discoveries
have been made of corn in the deserted cliff-dwellings, pueblos
and mounds. Dr. Edward Palmer* found, in the summer of
1869, in a mound in the neighborhood of St. George, Utah,
charred ears of maize, wood ashes and pieces of maize-cob
mixed together. The caiion of the Rio Mancos, in south-

1 Smithsonian Report, 1863. 379; '868. 401-
sMcAdams. Proc. Amer. Assoc. .\dv. Sci., 29, 718.

' Mass. Hist. Soc. Coll., vii, i93- Wood, New England Prospects, 106; Bureau ot
Ethnology. 11 Report. 207.

< Palmer. Monatschrift f. Gartenbaues. 1874, 163

100

Hafshberger. — Maisc :

western Colorado, contains numerous ruins of the cliff-
dwellers.^ " Above the cliff village was found a crevice
evidently used as a kitchen and storehouse, as is shown by the
beans and corn, which were found in a good state of preserva-
tion." Members of the Peabody Museum,^ in an exploration,
found a few cobs in a cliff-house on the Mancos River, which
corresponded with the "Conejos" maize in that they were
five inches long, with kernels small and flinty. Dr. Hayden,^
in exploring the ruins in southwestern Colorado, states in his
account that '* in some of the rooms [of the ruins, which he
examined] were found human bones, bones of sheep, corn-
cobs, raw hides and all colors and variety of pottery ware." *
BirdsalP describes the discovery of maize in the cliff-dwellings
of the Mesa Verde, in southwestern Colorado and northern
New Mexico. Stalks, husks, tassels, silk, cob and kernels of
corn were found. **That some of this material is as old as
the building is proved by the fact that the stalks were used in
the construction of the floors, being actually imbedded in the
adobe ; cobs being also used to chink the walls with, an
impression of the cob in the now hard adobe being found on
detaching one from its bed." The corn was a small dent.
The cob was about three inches long. Bandelier*' made some
very interesting discoveries on the side of the Arroyo de
Pecos. Human bones, walls of ancient structures, and a
grave were found on the bluff much above a layer of white
ashes, charcoal, corn-cobs and corrugated pottery, in a con-
tinuous seam 327 feet (100 m.) from north to south. "Con-
sequently the walls and graves must have been built over the
remains of a people which appears to have made indented and
corrugated pottery [and used corn, as the remains show], and
consequently the latter must be older, in time, than the
former." It does not appear that the sedentary Indians of
New Mexico made anything within recent times except

'Gannett, Henry. Pop. Sci. Month., xvi, 671.

2 Peabody Museum, n. 552, note.

« Hayden, Pop. Sci. Month., xii, 21.

* See page loi-

6 Birdsall, American Antiquarian, XI v, 135.

c Bandelier, Archeol. Inst. Amer., Rep, on Ruins of Pueblos of Pecos, 1881.

A Botanical and Economic Study.

lOI

painted pottery. Caspar Castano de la Sosa,^ when he made
his trip, in 1590, into New Mexico, mentioned that they used
painted pottery, red-figured and black. The indented ware is
almost identical with that found in the Rio Mancos and from
southwestern Utah, and it seems to be the oldest and most
primitive form. The cliff -dwellings of the Mancos are, there-
fore, comparatively very old. The cob found at Pecos, in the
ashes, or rather cut out of the bluff, is charred and small-
That these remains were not left by the later Indians is
proved by the fact that the Navajos and Apaches are terribly
superstitious concerning the cliff-dwellings, and can never be
persuaded to go near one, nor, indeed, have they ever been
known to enter one.^ The dwellings have been deserted a
long time, how long it is hard to establish, probably 500 years
or more, as the Navajo Indians relate. They are the farthest
north, and, therefore, the oldest, for as we go southward we
find indications of more and more recent habitation,' until we
reach the " Land of the Living Cliff-Dwellers," * lying be-
tween the Mexican States of Chihuahua and Sonora, in the
Sierra Madre, along the course of the Bacochic. "The timid
Tarahumari, a savage race, live mostly on the cliffs, or in
caves, and are worshippers of the sun, and while they plant a
little corn without cultivation on the steep hillsides, are not
otherwise tillers of the soil, but sustain themselves by the

chase."

The Nahuas had reached a comparatively high plane, as
compared with the more savage tribes about them. They
were a warrior race, built houses of quite skillful construc-
tion, and were agriculturists. They worshipped various
deities, Quetzalcoatl occupied the chief place, but minor gods
and goddesses were also objects of adoration. The goddess
Centeotl, corresponding with the Ceres of the Romans pre-
sided over horticulture, the fields and harvests. Her favor
was invoked both at the sowing and at the gathering of the
grain, and elaborate ceremonies were devised at these seasons

' Memoria del Descubrimiento, 238 ; Holmes, Geographical Survey, iii. 404, PL 44-
2 Monatschrift f. Gartenbaues Kgl. Preus., i^74 163
s Barber, E. A., American Naturalist, 11, 591-
'•Schwatka, Century Magazine, XLiv, 271.

102

HarsJiberger. — Maize :

of religious observance to gain her approval. She v^^as the
first woman, the ideal, the matchless' Sahagun^ pictures her
with a crown upon her head, and in the right hand a vessel,
her feet clothed wholly in red. In the ethnographical museum
at Berlin is preserved a stone image of the goddess with two
maize ears in her left hand. The Museum of Mexico ^ has a
vase ornamented with the emblems of Centeotl. It is twenty-
one inches high and nineteen inches in diameter above.

Of more questionable importance in the discussion are the
various codices of the Nahua people, in scraps and fragments,
preserved for us by the early Spanish writers, who translated
them along with many of very doubtful authenticity, but the
concurrent opinion of scholars seems to point to a pre-Co-
lumbian (if not very remote) origin of some of the better
known writings. An important record, written in Aztec
with Spanish letters, by an anonymous native author, and
copied by Ixtilcochitl, which belonged to the famous Boturini
Collection,* is the Codex Chimalpopoca, which, unfortunately,
has not been published, and references to which are to be
found only in the works of Brasseur de Bourbourg. The
quotation given by Bancroft ^ is clearly related to the Maya
narrative of the same event — the search for maize.

Until within recent years, the incriptions on the ruined
Maya edifices were a sealed mystery to philological archaeol-
ogists.*' It appears that the key at last has been found.

In 1876, H. T. Cresson visited the ficole des Beaux-Arts,
in Paris, and there saw photographs of the left-hand doorway
of Casa No. 3, Palenque. "The design and technique of this
masterpiece of the Maya scribe sculpture art is especially
fine, particularly the ikonomatic ornament, the figure of the
god Kukuitz. The head-dress of the figure represents feathers,
maize leaves, the quetzal head, and other decorations, notably
the heron (Baac-ha) in the act of pinching a fish in its pow-

» Steffen, Max, Die Landwirthschaft beiden Alt. Americanischen Kulturvolkern.

2 Sahagun, Historia General, Casas de Neuva Espana, NKxico, 1829, cap. 7.

» Brocklehurst, Mexico To-day, 195. PI- xxxv.

* Una Historia de los Reynos de Culhuacan y Mexico, etc. Boturini CatAlago 17-18.

& Bancroft, Native Races Pacific States, v., 193.

c Science, xxi, 1893, 8. Figs. 38, 39, 40. P- 9-

A Botanical and Eco7iomic Study.

103

erful bill."^ The inscription on the top of the left-hand slab
of Casa No. 3, interpreted by the Maya alphabet, reads as
follows: "the gods — earth, sky, water, maize — Kukuitz,
Kukulcan, Cauac, Muluc." '

Many interesting finds were made in South America. That
the Peruvians were agriculturists, and cultivated maize as
the staple crop, is proved by the numerous works that they
have left ; large terraces and irrigation canals testify to the
extent and development of the cultivation. The tombs,
graves, or huacas, have been relied upon generally for evi-
dence as to the Peruvian culture. These huacas nearly all
contain remains of maize, either in the ear or the grain. But
it has been shown conclusively that the graves, especially at
Ancon, have been used since the conquest, so that we must
be very judicious in the selection of our evidence and illus-
trations. The bodies of the Peruvians were buried in a
squatting position, with the thighs flexed on the pelvis, and
the legs drawn up parallel with the thighs. The face was
covered with cotton flock, sometimes llama wool ; the whole
body being rolled in a mat, and then tightly tied up. Nuts,
needles, heads of maize, and copper agricultural implements
were included in the rolling.' With the body were placed a
water-vessel, and a pot with grains of corn.* All along the
coast of Peru, for a distance of 1200 miles, are scattered here
and there thousands of ruins and huacas, while nearly every
hill and mountain have some upon them. We can say with
a reasonable amount of certainty that the Peruvians knew the
maize and used it. Darwin ^ unearthed some ears on a sea-
shore in South America, in a stratum which had evidently
been raised from nearer the sea-level, and to which he
assigns a great antiquity. Marcay " states that in the tombs
of the Aymara Indians the grain was found the color of old
mahogany, but it preserved its gloss, which he believes, from

1 Science, xx, 1892, icx).

« Science, xx, 1892, 78

8 Journ. Anthr. Inst., in, 3", 1874-

< Heath, E. R., Journ. Sciences, 3 Ser , 1879, 90

^ Darwin, Variations of Animals and Plants under Domestication, i, 320.

0 Marcay, Travels in South America, 1,69.

bMHU'.m-:?^

104

Harshberger, — Maize :

the construction of the huaca, to belong to a period long
anterior to the conquest.

Wittmack' describes the maize taken from the graves at
Ancon by Reiss and Steubel as in no way like the form
found in the mounds. But a later study led to a partial mod-
ification of his views, for Steff en '' quotes him as expressing
the opinion that the small, round, spindle-shaped grains found
in the graves were similar to those found in the mounds of
North America. The previous statement must have been
based on his third form {genabelter) dent maize. His second
form, pointed (spitz kornigeti), is found in many varieties in
Mexico, and Wittmack believes clearly points to a union
between Peru and Mexico (Steffen, 102). The nimiber of
varieties being greater in Mexico than in Peru, shoivs a greater
length of time in zvhich variation took place.

Heads of maize carved in stone are found in certain ruins.'
They are mentioned in Juan and Ulloa's work, nearly a cen-
tury and a half ago, and seem to have been better known at
that time than they are at present.* Squier gives in his book
on Peru (91) a bad representation of one of these stone maize
heads, and says that they were specially mentioned by Padre
Arriaga, in his rare book, "Extirpation of Idolatry in Peru,"
under the name Zara-tnama, and were the household gods of
the ancient inhabitants. "The Indians derived their idols
from those events which had influenced their course through
life, and which they thus commemorated."' They had their
lares, or conopa. Corn was called Zara-conopa. Stones cut
in the shape of maize ears were called Zara-7nama, and were
considered sacred, although not personified as deity.

The architectural and archaeological evidence points to
the Mayas as higher in civilization than the Peruvians, for
their sculptured figures and hieroglyphical inscriptions are
much beyond anything which the South American semi-civi-

» Wittmack, Monatschrift f. Gartenbaues, 1880, 121.

2 Steffen, Die Landwirthschaft beiden Alt. Amerikanischen Kulturvolkern, 1883.

Zeitschrift f . Anthropologie, xii, 1880.

3 Whymper, Travels Among the Greater Andes, 1892, 275. Illustration.

* Relacion hist6rica del Viaje A la America meridional, Madrid, 1748, § I047.
s Tschudi, Peruvian Antiquities, 171.

A Botanical and Economic Study.

105

lized tribes ever produced. The Mayas excelled in many
particulars: for instance, the Incas recorded their ideas by
quippas, or knotted cords, a much cruder and less satisfactory
method than the Maya glyphs. This indicates a much greater
antiquity for the civilization in Yucatan and Guatemala than
for that along the Pacific coast of the Cordilleras in South
America. We conclude that from the archaeological data it
seems very likely that maize originated north of the Isthmus
of Tehuantepec, and was carried south by barter or trade.

D. Ethnological Proofs.

The American race, as a whole, is singularly uniform in
phvsical traits, and individuals from one part of the continent
might easily be mistaken for individuals from other parts.
This uniformity is due, without a doubt, to the conformation
of the continents, which are within themselves geographical
units, the trend of the mountains and the situation of the
plains being identical in both North and South America.
The direction of the mountain chains from north to south
permitted a free and easy communication between the tribes
along the meridians of longitude.

Attempts have been made to classify the American
Indians by their ethnic traits, but the attempt so far has
proved a failure. Language affords a more satisfactory
method. We can, however, use ethnologic facts as a test of
the correctness of results obtained in other ways.

The mound-builders were a puzzle to students, and it is only
within the last few years that light has been thrown upon
their ethnic relationship. Scholars previously assigned a
great age to the works in the Mississippi Valley, but later
research shows this opinion to be erroneous. The consensus
of opinion points to the relative modernness of these tribes.
Our most distinguished ethnologists and archaeologists agree
that the mound-builders were Indians. Cyrus Thomas,
Powell, Hale, Brinton, Mooney, all state that the Indians
made mounds of greater or less size, and that after the white
settlers came the custom still prevailed.

•i

1 06

HarsJiberger. — Maize :

Brinton' says, ''A trip to the Ohio confirms the opinion
th:it the Southern Indians represent the mound-builders. It
woultl probably be hasty to point to any one of the southern
tribes as being specifically the descendants of the nation
who constructed the great works in the Scioto and Miami
Valleys." The evidence shows that the tribes of the Gulf
States and lower Mississippi constructed similar mounds, and
of even greater magnitude than the Ohio mounds. The
Choctas, the Natchez, the Cherokees, the Creeks, all built
mounds. Do Soto states that most of the Indians visited by
him built mounds. The Cherokees assert that they once
lived in the upper Ohio Valley, and that they built Grave
Creek and other mounds, and this tradition is supported by
historic data. Cyrus Thomas' asserts that the Cherokees
were the mound-builders. The agriculture of the mound-
builders was, therefore, scarcely older than that of the adjacent
southern tribes, who got maize from the west. What caused
the desertion of the mounds ? The Iroquois and Algonquins
stayed their old-time strife, and united, as tradition goes,
against a common and powerful foe. This foe was the
nation, or confederacy, of the Alligewi, the " semi-civilized
mound-builders" of the Ohio Valley. A desperate war
ensued, which lasted about 100 years, but eventually the
Alligewi fled southward, and seem to have mingled with the
more southern tribes.' Heckewelder* gives the Algonquian
tradition of the same event. The legend relates that the
Algonquins in their southeastern migration learned the culti-
vation of maize after they had reached a comparatively low
latitude, southern Indiana or Ohio."*

We have data as to the relative age of some Ohio
mounds. " In the pre-European state of the country, probably
some time after the year 1000, the American bison or buffalo
appears to have been absent from all the region east of the

» Brinton, Trans. Anthrop. Soc. Wash., in, ii6.

2 Thomas, Magaz. Amer. Hist., 1884, xi, 396-407; American Anthropologist, iv,

137-

3 Hale, Iroquois Book of Rites (Brinton's Library), 11 ; Amer. Assoc Adv. Science,

1882; American Antiquarian, 1883, Jan. and Apr.
* Heckewelder. Indian Nations.
6 The Lenape and their Legends (The Walam Olum), 187.

A Botanical and Economic Study,

107

Mississippi. It is doubtful if the creature existed for any
distance east of the Rocky Mountains."^ The Indians burnt
the prairies to make pasturage for wild herds, and the fires
communicated to the forests in the east, killed the growth
and extended the distribution of the herds to the east of the
Mississippi. Buffalo bones have been found in the mounds
at Madisonville, Ohio, and it is probable, therefore, that the
mounds at that place were raised after the year looo A.D.
The conclusion is that some of the Ohio earthworks were
erected in comparatively recent times by Indians identical in
many respects with the Indians occupying the Southern
States."^

The Huron-Iroquois ' are evidently a comparatively^old
race, and date back to a period before the arrival of the
Algonquins in the same region. Traditions seem to locate
the original home of this people in the country between
Hudson's Bay and the St. Lawrence River, a region where
maize does not grow. As they moved south, they probably
derived the use of the cereal from the mound-builders with
whom they came in contact. The Huron-Iroquois about this
time met the Algonquins, who were likewise spreading their
territory to the eastward and southward. The two nations
fought in a deadly feud, but eventually united their forces
against a common enemy, the mound-builders. History and
the traditions of both nations seem to point to this alliance
for offensive purposes.* It is probable that the Algonquins
borrowed a part of their culture at the time they came in
contact with the Iroquois and the mound-builders.

The Algonquian family occupied an area extending from
Labrador to the Rocky Mountains, and from Churchill
River, Hudson's Bay, as far south as Pamlico Sound in
North Carolina,"^ and was the most extensive stock in North
America.

The Muskogean family claims our attention next. They

' Shaler, Nature and Man in America, 183.

2 See pages 96, 97 and 98.

3 Annual Report, Bureau of Ethnology, 1885-86, 78; Brinton, American Race, 81.
^ Hale, Iroquois Book of Rites, 11.

5 Bureau of Ethnology Rep., 1885-86, 47.

8

io8

Hmshberger. — Maize :

occupied an extensive area from the Atlantic to the Missis-
sippi River, and from the Tennessee River south to the Gulf.
The Choctas were farthest to the west, along the Missis-
sippi ; the Muskokis were farther eastward. Their artistic
development was somewhat similar to that of the mound-
builders,' who have left such interesting remains in the Ohio
Valley, and there is, to say the least, a strong probability
that they are the descendants of those ancient builders
driven to the south by the irruption of the wild tribes of

the north. "^

The general trend of the Siouan migration has been west-
ward. In comparatively late prehistoric times probably most
of the Siouan tribes dwelt east of the Mississippi. The later
Siouan territory came in contact with the Comanche, a
Shoshonean family, on the west. The Mandans attained a ^
certain degree of culture, but the majority of the tribes pur-
sued the herds of buffaloes and lived on the bounties of
nature generally. The Sioux cultivated the soil ' spasmodi-
cally, for when they moved west across the Mississippi, maize
was not cultivated to the east, but was subsequently intro-
duced.

The Kioways, occupying a central position in the west,
were given to a wild hunting life,* and considered agriculture
a degradation (Whipple). They were the Arabs of the
American desert, depending on hunting and robbery for sub-
sistence.

The Caddos, or Pawnees, occupied an intermediate terri-
torial position between the tribes east of the Mississippi and
the tribes to the southwest, along the Mexican border. Their
nation extended from the Mississippi on the east to western
Texas, and from the Kioway and Siouan dominions on the
north to the Gulf of Mexico on the south. Agriculture was-
more in favor among them than generally on the plains.
Maize, pumpkins and squashes were cultivated, each family

1 See page 98.

2 Brinton, American Race, 88 ; Essays of an Americanist, 1890, 67.

3 Bureau of Ethnology Rep, 1885-86, 112; Dorsey, American Naturalist, 1886, 220 ;.
Brinton, American Race, 98; Bancroft, Native Races, i, 49i-

^ American Race, 10 1.

A Botanical and Economic Study.

109

having its own field of two or three acres. For about
four months of the year they were sedentary, dwelling in
houses built of poles and bark, covered with sods, while
the remainder of the time they wandered over the hunting
grounds. The Wichitas, Caddos and Pawnees were tribes of
this stock.

The Pacific Coast tribes were not agriculturists. Their
food consisted of fish, roots, berries and game, the spon-
taneous products of land and water.' "They reject nothing
that their teeth can chew or their stomachs are capable of
digesting, however tasteless, unclean or disgusting it may
be." ''■ We have evidence from other writers to the same
effect.*

The Pueblo tribes have long attracted attention, and they
may be divided for convenience into the following groups :
Shoshonean {Tanoan, Moqiii\ Zuiiian and Keresan.

It is probable that further research will result in proving
the radical relationship of the Tanoan people to the Shosho-
nean stock. The Keresan pueblos are probably the oldest.
" Die Keres sind nach Pike der hauptsachlichste Bestand der
civilisirten eingebornen Volker in Neu Mexico, welche die
Ueberbleibsel von so alten Stammen sind."' According to
Frank H. Gushing, the Nahua nations are a younger people
in civilization than the Keres, and as they moved south they
gradually absorbed the culture of their neighbors. The Zuni,
from their own traditions, borrowed their culture from the
Keres to the east. They occupy but a single pueblo on the
Zuiii River in New Mexico.

The Yuman stock occupied the valley of the Colorado and
the peninsula of Lower California. They were split up into
numerous tribes. Those on the peninsula were in the lowest
stage, without agriculture of any kind, and it is likely that
they represent the primitive condition of the Yumas, and
have always lived as at present. The principal tribes

• Smithsonian Report, 1887, 621.

2 Rau, Trip Across Peninsula of California, Jesuit Priest, 1773.

3 Proc. Amer. Antiq. Soc, New Ser., 11, 327.

* Buschmann.

I

no

Harshberger. — Maize :

are the Yumas proper, Maricopas and Mohaves. The
Yumas, Maricopas and Mohaves were long acquainted with
agriculture, and grew corn, using a pointed stick in planting

it.^

The Athapascan race, spreading over a vast territory in the
north and south, were a warlike people and the main agents
in destroying the civilization on the Gila and its affluents.
Intellectually they were below the average. The Apaches
were redoubted warriors, non-productive, subsisting wholly by
plunder and the chase. They lived entirely by hunting.'' The
Navajos were the most highly cultured, yet it turns out that
their culture was due to captured members of more gifted
tribes. '* Agriculture was not practiced, either in the north
or south, the only exception being the Navajos, and with them
the inspiration came from other stocks." "" The best blanket-
makers, smiths and other artisans among the Navajos were
descendants of captives from Zuiii and other pueblos.' The
Apaches and Navajos were the principal tribes.

The Shoshonean family was wide-spread and extended
from the Columbia River in the north to Nicaragua in the
south. Buschmann and Brinton consider the Nahuatl races
related to the Shoshones of the Bureau of Ethnology,' and I
am inclined to follow their lead in this matter. The tribes of
this stock present the greatest diversity of traits. The
Nahuas were highly cultured in comparison with the other
nations about them, yet they belonged essentially to the same
race as the poor root-digger Ute, with the lowest type of
skull on the continent.^ Living on the arid plains of the
interior, the Utes had been for generations half starved.
They were not agricultural; but lived on fish and wild seeds.
Indeed, the Ute branch, including the Comanches, with the

' Emorv, Rep. U. S. and Mex. Bound. Siirv., i, 112; Indian OflRce Report, Special
Com.. 1867,337; Merriwether. Ind. Off Rep., 1834. 172; American Race, 109; American
Antiquarian, viii,276.

« Delgado, Ind. Off. Rep., i860. 1864.

8 Brinton, American Race. ^\.

* Bourke, John G., Journ. Amer Folk Lore, 1890, 115; Bandelier, A. F., Indians
S. VV. United States. Boston. 1890, I75-

5 Powell. Bureau of Ethnology Rep , 1885-86.

« Virchow, Crania Ethnica Americana

A Botanical and Economic Study.

Ill

exception of the Moquis, who probably borrowed their culture
from the Keres, were averse to agriculture, but they were
more willing to accept a civilized life than the Apaches or
Kioways.

The Piman* or Sonoran branch (Brinton) of this family
comprised the Pimas, Cahitas, Coras, Tarahumaras and Tepe-
huanas, as the principal tribes. Buschmann and Gatschet
favor the opinion that the Pimas are Shoshones. The Pimas
occupied the region of the Gila Basin and head of the Bay of
Lower California. The remarkable cliff-dwellings are in this
region, and it is probable that the ancestors of the Pimas
were the builders and inhabitants of the rock shelters. There
is nothing to warrant a contrary statement, for a culture
higher than the Piman is not necessary to explain the struc-
tures. As Schwatka has pointed out,' the timid Tarahu"
maras still live in cliff-dwellings. The Pimas have a tradition
that they were driven south by the Apaches.' These people
were agricultural and irrigated their fields with canals and
ditches. The Coras of this group reached farthest south into
the State of Jalisco. These tribes were not in culture earlier
than the Keresan, for Mindeleff has explained the probable
stage of progress toward pueblo building; that the circular
house of stone and mud derivable from the circular wigwam
still used by the wandering tribes of the Ute branch, became
square by aggregation of the buildings. The people driven
to the mountain fastnesses by the warlike tribes, still built
the square form of house, but lor lack of sufficient building
sites the families built their houses one above the other on
the mountain slope (the cliff-dwelling stage), and after the
danger was over, and the people returned to the plains to
live, they built, as the result of their mountain experience,
the pueblo with compartments one above the other, and

' Powell, Bureau of Ethnology Rep , 1885-86.
2 Schwatka, Frederick, Century Magazine. 1892.
•Grossmann, F. E., Rep. Smithson. Inst., 407-10.

3 1'l

3 i'

h

ir

II

HarsJibcrgcr. — Maize :

reached by ladders/ Lieutenant Simpson ' records an obser-
vation, which seems, to support this view. He found, in close
proximity to one of the ruins, an excavation in the cliff which
had been closed with a front wall of well-laid stone and mortar,
thus associating one of the simplest cave-dwellings and a per-
fect pueblo building, a fact of no little importance.

Before leaving the territory of the present United States,
it may be well to draw a few conclusions from the facts
already presented in this section. The practice of agricul-
ture was chiefly limited to the region south of the St. Law-
rence and east of the Mississippi. In this region it was far
more general, and its results were far more important, than
is generally supposed. To the west of the Mississippi, only
comparatively small areas were occupied by agricultural
tribes, and these lay chiefly in New Mexico and Arizona, and
along the Arkansas, Platte and Missouri rivers. The rest of
the region was tenanted by non agricultural tribes. ''The
practice of agriculture, to a point where it shall prove the
main and constant supply of a people, implies a degree of

iMindeleff, Bureau of Ethnology Rep., 1882-83, 473- The Navajo hut, a bee-hived
conical structure, made of turf, etc., in Zuni is ham-pon-ne, from hawe, dried brush, sprigs
and leaves, and po-an-ne, a covering or shelter. When the term was formulated, the
Zunis were probably acquainted with this form of building. A walled enclosure, he-sho-
ta-pon-ne. from he-sho, gum, was round rather than rectangular, and was found in the
southwest lava wastes. The lava resembles asphaltum, hence ahe-sho, gum rock. The
rectangular hut was derived from the round by aggregation. The flat and terraced
roofs, in all probability, were derived from sloping mesa sites, and this overlapping is

due to the decrease in the number of available sites on the hill. The name of an upper
story, in a pueblo, is osh-ten-u thtan, from osh-ten, a shallow cave or rock shelter, and
u-thla-nai-e, placed around, embracing, inclusive.

2 Simpson, North Americans of Antiquity (Short), 292; Journal Mil. Recon., 34, 131 ;
Domenech, Deserts, i, 199, 379-81,385; I5aldwin, Anc. Am., 86, 89; Bancroft, Native
Races. 1, 652-62; liarber, American Naturalist, xi, 1877, 591.

A Botanical and Econumic Study.

113

sedentariness to which our Indians, as a rule, had not
attained, and an amount of steady labor without immediate
return which was peculiarly irksome to them. Moreover, the
imperfect methods pursued in clearing, planting and cultivat-
ing sufficiently prove that the Indians, though agriculturists,
were in the early stages of development as such, a fact also
attested by the imperfect and one-sided division of labor
between the sexes, the men, as a rule, taking but a small
share of the burdensome tasks of clearing land, planting and
harvesting. It is certain that by no tribe in the United
States was agriculture pursued to such an extent as to free
its members from the practice of the hunter's or fisher's art."
The facts collected in these few pages indicate that the
tribes to the east of the Mississippi borrowed their agricul-
ture from the west, for along the Mexican border the Indians
were more sedentary and farther advanced in the arts.
Again, the culture along the Mexican border had one or two
probable centres of distribution, the one, the Keres, the
other, the Pimas, who both derived it from farther south.

The Nahuas formed by far the most important branch of
the Shoshonean family. They occupied the territory from the
Rio Grande south to Nicaragua, and possessed, as it went in
America, a comparatively high grade of culture. They had
an elaborate government, presided over by a Montezuma ;
they were skillful as builders; their utensils were of copper
and tin; gold and silver were worked into ornaments; their
religious system was elaborate and minute, and the litera-
ture, preserved on parchment of maguey plant, in the ikono-
matic characters, was large and important; they had a calen-
dar ; they tilled the soil, as was necessary for a dense popula-
tion, raising maize, beans, pepper, gourds and other fruits.
Finally, they had organized armies. Each king was absolute
in his own country (Mexico, Tezcuco, Tlacopan) until war
broke out, when they acted jointly. The country was so
densely populated that floating gardens, or chinampas, were
constructed. All the agricultural products of the country,
particularly maize, chile and beans, were sown in abundance
on the chinampas, constructed of logs, with brush and earth

IP

114

Harshberger. — Maize :

on the top. Prescott says : " Fairy islands of flowers over-
shadowed occasionally by trees of considerable size." "That
archipelago of wandering islands."^ The Nahua nation
were probably no more cultured than their relatives the Utes
and Comanches, until they came in contact with the Kere-
sans in the north, or the Mayas and other cultured tribes,
the Totonacos, Zapotecs and Chinantecs, in the south. The
Aztecs did little more than copy the works left by their pre-
decessors.^ A general ethnic rule, applicable in many cases,
may be laid down, that less civilized tribes coming into con-
tact with more civilized nations imitate the customs of the
more successful race, and a struggle ensues, in which the
more cultured race, if a stronger one physically, has the advan-
tage, and its culture obtains the ascendency. The fact that
the Nahuas preferred the highlands of Mexico to all other
districts as a dwelling-place, is accounted for by the surround-
ings, which were attractive and wholesome. Maize grew
well, and on the plateaus, in addition to this cereal, grew the
maguey (Agave Atnericana) ; in the neighborhood lower down
the coast grew a great variety of tropical fruits, amongst
others the cacao and the vanilla.'^ They were in an already
long-settled country ; the fields were cleared, and the country
inviting, with its ancient buildings and arts. With the first
germs of civilization instilled into them, it is no wonder that
they developed into a great and powerful nation. The same
conditions determined the location of the ancient civilizations
iij the rich and fertile river valleys of the Euphrates, Nile,
Ganges and Yang-tse-Kiang.

The Mayas were the oldest in civilization of any race on
the North American continent.* Copan, Palenque, Uxmal,
were built by them, and these ancient architectural remains
were of a high order and indicate a long process of evolution
from more primitive forms. They excelled in architecture,

* Prescott, Conquest Mexico, ii, 70, 107-8; Cortes, Cartas, 79; Sarmiento, Heredia,
95, 96; Torquemada, Monarq. Ind.. 11, 483; Carli, Cartas, r, 38,39; Bancroft, Native
Races, 11, 344

-' Tylor. E. B., Anahuac, London, 1861, 188.

3 Peschel, O., Races of Man, New York, 1876, 449.

* Nadaillac, Prehistoric America, 262; Contributions N. A. Ethnology, 75; Short,
John T., North Americans of Antiquity, 203; Standard Natural History, vi, 219.

A Botanical and Economic Study,

115

for their sculpturings are bold and strong, as the facades of
the edifices, covered with curious designs, attest. Their
boats were seaworthy, and a trade was established between
Cuba and Yucatan, for Columbus was shown wax from
Yucatan, and was told about the countries toward the sunset.
Cacao beans and shells served as media of exchange. They
had an extensive literature ; they used tablets and covered the
walls of their structures with hieroglyphics. *' Their speech
forms one of the rare examples of an American language
possessing vitality enough not only to maintain its own
ground, but actually to force itself on European settlers and
supplant their native speech." Berendt states "that whole
families of pure white blood do not know Spanish, but use
Maya exclusively." The calendar attracts attention, and
appears to have been the basis for that of the Nahuas. They
were agriculturists. Maize, beans and pepper were culti-
vated, and bees were domesticated, from which both wax and
honey were collected. This nation extended over the penin-
sula of Yucatan and into Guatemala and Tabasco.

The Chibchas extended in both directions from the Isthmus
of Panama, and thus have representatives in both North and
South America. Most ofthe tribes in New Granada were of
this stock. The Chibchas proper were highly cultured.
Their home was one of the most southern of the entire
family. A number of tribes in the States of Panama and
Costa Rica were probably branches of this nation. Agricul-
ture was pursued, the produce being maize, potatoes, yucca
and cotton. Irrigation was practiced to some extent. It is
doubtful whether they had any means of writing.

The Kechuas next claim our attention. There are many
reasons for believing that the tribes comprised in this stock
appeared in South America at the extreme north of the
region they later occupied, near the great trade highway
across the Isthmus of Panama. Their seat seems to have
been near the present city of Quito. Later the Incas extended
their sway to the thirtieth degree of south latitude. The
Peruvians were highly cultured. They were governed by an
Inca, or war chief, elected by a council of the gentes. The

i fll

%:-

ii6

Harshberger. — Maize :

land was owned by the gens. Agriculture was highly devel-
oped. The soil was fertilized with guano, and extensive
systems of irrigation were used. Maize, potatoes, yucca,
peppers, tobacco and cotton were raised. Architecture
reached a^high stage. It was cyclopean, erected upon tumuli,
or pyramids. The Peruvians were greatly deficient in decora-
tive skill and sculpture. They recorded their ideas by the
quippas, or knotted string, a system far inferior to the hiero-
glyphics of the Mayas. Their religious system was elaborate
and constituted a worship of the sun. A comparison of
the two greatest civilizations on the American continents,
the Maya and the Peruvian, leads to the opinion that the
Mayas excelled in those traits which make a nation great.
Their architecture was more artistic, their literature fuller
and richer, as a comparison of Maya glyphs and Peruvian
quippas alone shows.*

The Amazon and Orinoco basins were largely occupied by
tribes of Tupi, Tapuya, Carib and Arawak stocks.

The Tupis were found along the seaboard from the mouth
of the La Plata to the Amazon, and far up the banks of the
latter. Here they are called the Guarani. " The general
culture of the Tupis was superior to the that of the Brazilian
tribes generally, but inferior to that of the Incas. They were,
to a slight extent, agricultural, raising maize, manioc and
tobacco." Some fowls, monkeys and peccaries were tamed
and used as food.^ The Tupis in Ecuador evidently profited
by their nearness to the Peruvians, for they lived in perma-
nent villages, had good roads, knew gold, silver and copper,
and cultivated large fields of cotton, maize and various food
plants. " The art forms which they produced, and the preva-
lence of sun worship, with rites similar to Peru, indicate the
source of their more advanced culture."

" The Arawak stock was the most widely disseminated of
any in South America. It began in the south with the
Guanas, on the head-waters of the River Paraguay, and with
the Baures and Moxos on the highlands of southern Bolivia,

^ See Standard Natural History, vi, 219.
'Brinton, American Race, 233.

«i

A Botanical and Economic Study.

117

and extended in continuity to the Goajiros peninsula, the most
northern land of the continent." '

The Antilles and Bahamas were peopled by its members.
They were the first, therefore, to welcome Columbus to the
Bahamas, Cuba and Hayti. The Arawak stock was above
the stage of savagery. They cultivated maize, potatoes,
manioc, yams and cotton. They made gold ornaments, idols
of rude form, and canoes constructed of hollow logs. The
Guiana Arawaks cultivated maize, but cassava afforded the
chief food. The forest tribes planted in clearings inter-
mingled with wild seedlings and sprouts from stumps. Brin-
ton places the original home of the Arawaks somewhere in
the Bolivian highlands, where, no doubt, they learned the use
of maize.

The Timucuas inhabited the State of Florida, where they
have been extinct for a century and over. Gatschet ^ and
Brinton incline to the opinion that the Timucuas were related
to the Caribs in the Bahamas and the greater Antilles.
There is no objection to holding this view, for the Timucuas
were isolated as regards the surrounding tribes, and the Caribs
extended to the Bahamas, near the coast of Florida.

This brief ethnological survey of the North and South
American tribes is useful in showing the comparative age
and cultural position of the agricultural races of Indians.
That the Mayas were the superiors of any other race on the
North or South American continent, and were the source of a
large part of the indigenous American culture is proved : (i)
by the fact that the tribes and mound-builders in the present
territory of the United States were just entering on the
agricultural state ; (2) because the Pueblos built structures
of scarcely higher order than the rock shelters of the cliff-
dwellers, who were driven by the invasion of wilder tribes
to build in the mountain fastnesses instead of living on the
plains in round or rectangular huts of stone and mud ; (3)
because the Nahuas evidently borrowed their agriculture
from the more advanced Maya tribes in the south, with whom

' Brinton, American Race, 241.

- Bureau of Ethnology Rep., 1885-86, 123.

ii8

Harshberger. — Maize :

they came in contact in their settlements and migrations ;
(4) because the Incas occupying a northern location, at first
near the [great Panama trade route, at a later date spread
their sway to the far south, carrying the germs of agriculture
and the rudiments of the arts to the barbarous tribes in the
Bolivian and Chilian highlands, and this argues for a com-
paratively recent development of their civilization and agri-
culture, for in Mexico all the tribes practiced agriculture, but
in South America, among the wild tribes, agriculture was yet
in its incipient stage.

E. Philological Proofs.
Language is important in determining the culture, migra-
tion and evolution of races of men. The study of the Aryan,
or Indo-European, tongues has thrown a flood of light upon
the early history of the European and Asiatic races. When
the American languages shall have been studied with the
same thoroughness and care, equally valuable results will be

obtained.

De Candolle has shown that the study of language aids m
determining the origin of our cultivated plants and their
subsequent distribution. It is likely, therefore, that a com-
parison of Indian names for maize will aid in determinirg
the primitive home of the cereal and its carriage through the
American continental areas.

The Algonquian family, numerous in tribes, was spread
over a vast territory in Canada and the United States. All
the tribes have a common root for maize. This points to the
knowledge of corn by the Algonquins while they still formed
one nation. It is probable that the primitive seat of the
undivided Algonquins was near the great lakes, and the dis-
persion occurred in two directions, one wing extending into
the northwest, the other to the east and northeast. The
use of maize was learned, in all probability, while the nation
occupied their;undivided home. " The root min or mun is a
generic suffix applied to all sorts of small fruits,' and when

» [3rinton, The Lenape and their Legends, 48.

A Botanical aud Economic Study.

119

maize became known to these Indians they extended the use
of the word to the cereal, and combined other terms with it
to define more positively the kind of small grain meant."
Thus the Chippeway word for corn, mandamin, the Ottawa,
mindamin, the Cree, mittamin, contain the radical min in full,
combined with an abbreviation of the word manito, divine,
meaning, of course, that the grain was divine, supernatural,
mysterious. In the Delaware, the combination is different.
The Delaware word jesquem (Campanius), chasquem (Zeis-
berger), contains the radical ask, Chippeway ashk, Delaware
aski, meaning green. The reference is to the green, moving
plant in the fields during the summer months. The Piegan
Blackfoot word drops the radical min, but retains the root
ask, thus, esko-lope. A classified list of the tribes with their
names for maize will show that all the tribes, however widely
separated, had a common root for that important cereal :

Abnaki skamen (Father Rasle) (Journ. Antiq. Soc, 11, 305).
Algonquin, mitamin (Journ. Antiq. Soc, n, 305) (La Houtan).
Chippeway, mandahmin (Keating) (Journ. Antiq. Soc.) (Long).

mittawmin (Herioti.
Cree, mittamin (Brinton).
Delaware, khasquem (Journ. Antiq. Soc , 11, 305 .

husquim (Whipple, Ewbank, Turner, 1855, 9).
jesquem (Campanius"^.

Lenape. mesittewalL corn boiled whole,
scheechgamin. coarse shelled,
schesquim, bran, husk,
schesquasquim, hulls,
winamin, corn is ripe,
vvinaminge. month of August; literally, time

of roasting ears,
achpocm, roasted corn,
simaquon, corn-stalk.
Illinois, micipi (Duponceau'.
Massachusetts eachim-meneash (R. Williams).
Menominee, waupim-meenuc (Doty) (Journ Antiq. Soc, 11, 305).
Miami, ment-sheepeh (Thornton).

Micmac S. T. Rand, Dictionary Micmac Language, 1888 .
peaskumun.

peaskumuskw, corn-cob, corn stalk,
peaskumunwees, corn-hill.

f'ti-

II

120

Harshheroer.— Maize :

Micjuac peaskumun, nac-aktook, ear of corn.

oosaboon, corn-silk ; literally, head of hair,
peaskumuna, egadakim, corn-field.
Narraghansett. ewachim-neash (R. Williams).
Ottaiva^ mindamin (Brinton).
Passamaqtwddy, piaskomin (Sturtevant).
Sac, tamin (Long's Sec. Exp. St. Peter's River, ii).
Shawnee, tame ( Johnson K

dame ijourn. Antiq. Soc, i, 287).
tarmi (Whipple, Ewbank, Turner),
tami (Jefferson) (A. S. Gatschet).
lena-wi-wi tami, Indian corn,
tami-skui, corn-cob.
nikutik-kah-knimi, corn-ear.
kulciskwa, corn-stalk,
kti-ka (Gatschet).

There is an apparent similarity between the Algonquian and
Iroquoian words for maize. This either establishes an earlier
connection between the two nations, or points to a derivation of
maize from a common source. The latter view is supported
by tradition, which relates that both stocks united their forces
against a common foe, the Alligewi, or mound-builders.
Conquest changes not only the conquered, but also the con-
querors. Insensibly, it may be, but deeply, they are affected
by the character of the absorbed races. A powerful mter-
tribal imitation must have been constantly at work, imitation
of the culture of the more highly evolved though less warlike
tribes; and this imitation and absorption prove very powerful
when conquest takes place. Duponceau^ long ago compared
the words for maize in the two languages :

Algonquin.
chas-quem.

There seems to be a probable connection between chas
(ask, aski), Algonquian, and atschia, Iroquoian, for the Seneca
drops the last portion, and calls maize onaa.

The following list gives the words for corn in Iroquois:

Cherokee, allo-selu (Journ. Antiq. Soc, n).
selutikatunung.
kungwisitung (S. A. Worcester).

' Duponceau, M^moire des Langues de 1' Am6r. du Nord, Paris, 1838.

Iroquois.
on-atschia.

A Botanical and Economic Study. I2r

Mohawk, onusti (Journ. Antiq. Soc, 11).
Nottoway, ohnehahk (Journ. Antiq. Soc.) (Wood).
Oneida, ohnloto (Journ. Antiq. Soc.) (Jefferson).
Onondaga, onatschia (Journ. Antiq. Soc.) (Zeisberger).
Sefieca, onaa (Journ. Antiq. Soc.) (Parish).

onoohquaw.
Wyandot, nayhah (Journ. Antiq. Soc) (Archseologia Americana).

The Muskogean family consisted of three principal tribes,,
the Chocta, between the Mobile and Mississippi rivers,
the Chicasa, at the head-waters of the Mobile River, and the
Maskoki, or Creek, between the Mobile and Savannah rivers.
The words in Maskoki, or Creek, are atchi (Gallatin), atshi
(Journ. Antiq. Soc), and adshi (A. S. Gatschet).

The Creeks may have been the original mound-builders, or
closely allied to them. A curious similarity appears in the
Creek word atchi, adshi, the Algonquian aski (chasquem),
and the Iroquoian onatschia, which, if so joined, philologically
enforces the theory advanced that the Algonquins and Iro-
quois obtained corn from a powerful southern tribe. The
Chocta word is tanchi (Whipple, Ewbank, Turner), tandshi
(A. S. Gatschet), tonche (A. Wright). The Chicasa word is-
tuncha (Gallatin). All these words are related to the terms
used by the Indians west of the Mississippi River.

The Caddoan family had three principal tribes, the Pawnee,
Wichita, and Caddo. The Pawnee word for corn is task (W.
E. T.), the Wichita is tash (A. S. Gatschet), the Caddo word
is kish-ee-ee (G. Gray), showing a close relation between all
the words. A. S. Gatschet thinks that the Pawnee task, the
Wichita tash, and the Chocta tanchi, are related, which would
show that the Choctas borrowed their use of maize from
the Caddos to the west. The Siouan tribes, in all likelihood,
borrowed their words from the same source, for we have
Ponca, watanzi, Omaha, wattanza (Journ. Antiq. Soc), Osage,
watanshee (Journ. Antiq. Soc), Ottoe, watooja (Say), and
this carried farther in the Winnebago wachoa (Journ. Antiq.
Soc). The first half of the word is evidently wakan, mean-
ing superior, or supernatural.V The Tutelos had Algonquian
loan-words, mandaqei, mataqe. The Catabas (in South Car-

1 Roehring, Language of the Dakotas, Washington, 1872, 14.

ii

11

II

122

Harshberger. — Maize :

olina), clearly related to the Siouan tribes, apparently derived
their word from the Caddo proper, kus, kush (A. S. Gatschet),
koos (Journ. Antiq. Soc), of the Cataba, seems to be related
to kish-ee-ee, and the tash of the Caddo stock. The Dakotas
have an entirely independent set of words, the affinity of
which is unknown. A list of Dakota words follows :'

Dakota, wamnahesa, wahinske, corn.

huwapa, wahuwapa, ears of corn.
Omaha, watanzi, corn.

wahaba, corn-ear.

The words huwapa, wahaba, may be connected with the
Chemehuevi hahwib (W. & T.), although this is merely con-
jectural.

A dance at the words for maize as used by the California

Indians will show that they learned the use of maize after the

advent of the white man into their country.'

Maize = maiz i Father Sitzar, San Antonio Mission, 1861).
Maize = maiz, Yukis, Huchnom, Porno, Gallinomero, Yokaia, Yokuts,
Mariposa, Timlinneh.

The Pueblo tribes and Yumas, which lived on the Mexican
border before the Shoshonean invasion, next claim attention.
A list of words will prove of advantage in a comparison.

Yu?na, Yuma proper, tiyatch.

terditch (Whipple).
Cocomaricopa, terditz.
Mojave, terdicha.
Moqui, ka-ah (Bourke).

karuk (Buschmann).
Zuni, melah (Eaton).

ta-a or a-ta-a, seed of seeds.^

"^"^^^ I (Buschmann),* (Eaton).^
muwe f

Tesuque, kiihn.

Isleta, i-e, corn (Gatschet).

Keres, ya-oca (Whipple).

yachi (Kiwomi Clan).*

>Dorsey, J. O., Geol. Survey Rocky Mts. Ethn., vi ; Bureau of Ethnology Rep.,
1881-2, 304; Dakota Grammar and Language, Wash., 1852, 293.
« Powell, Geol. Survey Rocky Mts , iii. 476-
3 Gushing, Indianapolis Millstone, October, 1884.
* Buschmann, Abhand. Akad. Wissensch., Berlin, 1857, 288.
5 Eaton, Schooler's Indian Tribes, iv, 1854, 416.
*i Buschmann, Abhand. Akad. Wissensch., Berlin, 1857. 3oi.

A Botanical and Economic Study.

123

Notwithstanding Gatschet's positive statement, that there
is no connection between the words of the different Pueblo
.stocks for maize, I cannot forbear comparing the Zufii ta a or
a-ta-a with the Moqui ka-ah. The statement made by Gushing,
that the Moquis have words which show that they borrowed
a portion of their culture from the Zuiiis or Keres, is sig-
nificant. The Keresan word yatchi, or yaoca, seems to be the
primitive form of the Yuma tiyatch (terdich, terditz) ; but too
much stress cannot be laid on this curious correspondence of

radicals.

The Uto-Aztecan stock of three distinct branches extended
from the Golumbia River south to Nicaragua. A list of words
for maize in the various families is given :

ShosJionean Branch.

Chemeheuvi, hahwib iW. E. T.).

Comanche, hunibist (W. E. T.\ (Buschmann, 1857, 258)

Moqui {ante).
S<i)ioran Branch.

Cahita, bachis (Buschmann, 1854).

Cora, yurit (Buschmann, 1854).

Tarahumara, schunucu, sunu (Buschmann, 1854).

Tepehuana, June (Buschmann).

Pima, ou-in, oo-on (W. E. T.}, (Buschmann, 1856, 356 L
A'ahuan Branch.'^

Maize stone,

Tortilla,

Ears,

Hulled maize,

Full maize,

Unripe maize,

PiPILS.

medat.

tax.

cinti.

tayugal.

ulut.

elot.

Aztecs.

mctlatl.

tlaxcalll.

cintlf.

tlaolli..

olotl.

elotl..

' StoU, Zur Ethno^raphie der Republic Guatemala, Zurich, 1884.

if

ll

f *

124

llarsJibergcr. — Maize

It is easy to see that the Shoshonean branch learned their
use of maize from other tribes. The Moquis were the most
advanced, but they were copyists. Buschmann says : ' " Die
Namen fiir Maiz harnewista, hahnebeteh aus welche Formen
im vergleich mit pohewista [p:isen cf., also pohewista, gold]
und dem Namen hainena-una eines Stammes harne, hahne
und haine als das eigentliche Wort fiir Maiz, bekleidet mit
einen starken Endung ; hervor springt Tarahumara schunucu,
sune, Tepeguana June, Cora yurit." In other words when they
learned of maize, the grain was golden or glistening, like the
precious metals, and they therefore aflfixed the term wista to
describe the grain, or seed, more particularly. The Sonoran
group links together the region of the Colorado River, where
the Pimas dwelt, and the State of Jalisco, in latitude twenty
decrees north, where the Coras dwelt. It is safe to suppose
that an inter-tribal communication was always kept up between
the members of this branch ; this is sufficient to account
for the distribution of maize from the south into the present
territory of the United States, and for its presence among
the Zufiis and Keres, who afterwards taught the Moquis of
later migration agricultural practice' It appears that oo-um,
oo-oon, the Pima for maize, are related to sunu through
hainena une (une, oo-oon, oo-um, ou-in). The Pimas must
be reconciled to the Uto-Aztecan stock. Buschmann ' says :
** Ich gehe daran aus dieser Wortsammlung die Resultate
zu Ziehen, welche die Pima Sprache, als ein wichtiges fiinftes
Glied des Sonorischen Sprachstammen ervveisen ; voll Sonor-
ischen Stoffes, in welchen sie die Tepeguana hiiufig sehr und
auffallend nahe steht ; denoch voll eigenthiimlicher Worter,
clurchzogen von Aztekischen Resten."

The Mayas occupied a central position in Mexico. Stoll *
gives a list of words in the Maya :

» Buschmann, Abhand. Akad. Wissench., Berlin, 1854. 396.

'-' Cf pages 123 and 147.

•■' liuschniann, Die Pima Sprache, Abhand. Akad. Wissench,, Berlin, 1856, 371.

■• Stoll, /ur Kthnographie Guatemala, 53.

A Botanical and Economic Study.

125

Huasteca, .

Tortilla.

Ear.

Grain.

Unripe.

I

bacam.

guai.

isis.

ajam.

2

Maya, .

vuaj.

nal.

ixim.

nal.

3

Chontal, . .

1 «

i(

chojno.

4

Tzental, . .

..

((

ajan.

5

Tzotzil, . .

(i

((

6

Chaneabal, ,

jal.

<(

((

7 •

Choi, . . .

. ( <

( 1

sal.

8

Quekchi, . .

vua.

«'

(i

och.

9

Pakonchi, .

vuic.

<i

raxjal.

10

Pokomam, .

( <

<(

ajm.

1 1

Cakchiquel,

vuay.

(<

iiz.

12

Quiche, . .

vua-lej.

( 1

raxjal.

13

Uspantica, .

vua-lej.

((

t (

cux.

14

Ixil, ....

< (

matzinjal.

15

Aguateca, .

vua.

ti

xeha.

16

Mame, . . .

i

• ixin.

The word aima in the Xinca language of Guatemala, also
found in Chontal, is probably derived from ixim, the universal
word for maize in the Maya. Later, we have for corn-field,
uaya'a, which is close to the Cakchiquel ouan, corn-field, or
aue.x, when the corn is young. If this is correct, it indicates
that the neighboring tribes learned the cultivation of maize
from the Mayas.*

The Costa Rican tribes, the Boruca, Terraba and Guatuso,
have words for maize,^ as follows : •

' Brinton, Proc .\mer. Philos. Soc, 1884,91.

- Archiv. fiir Anthropologie, 1886, 591; Revue d'Ethnographic, torn. 16, 1887, 128.

:■!

i .

I

126

Harshbcvircr. —Maize

BoRUCA. Terr ABA.

GUATUSO.

Maize, ^^P- ^P' 'P'

I

:Maize-field kup-kah. te.

j.^ars, I^UP- I ip-^ro^-

ain.

ani-ki-tokufa.
ai ki-kacheijo.

The Zapotec word is xupaac, which is comparable to kiip
and kupac, the Costa Rican for maize-field. The word kupac,
in Costa Rican. is equivalent to the word shipyac in the
language of the Timotes, a South American tribe, showing a
connection between the two continents. Nor does the evidence
cease here/ for the Mazatecs, in the northwestern part of the
State of^Oaxaca, were related to the Mangues and Chapanecs
near Lake Managua, in Nicaragua. The Mangues had as
neighbors, beyond the Cordilleras, in Costa Rica, a group of
related tribes, the Talamancas, Bribris and Vezertas, which
have been shown by Dr. Max Uhle, Dr. Ernst '^ and other
students to be not distantly connected with the important
Chibcha stock of New Granada. The Chapanecs were
largely infiltrated with the blood of Costa Rican tribes,
with relations in South America, enabling us to to trace
influences of a South American linguistic stock as far north
as the northern border of Oaxaca, a discovery full of signifi-
cance for the history of aboriginal culture. The Maya word
ixim agrees apparently with the Mazatec nama as shown in the
comparison of nouns.

Maize.

Maya.
ixim.

Xinca.
aima.

Mazatec.
nama.

C/iapanec.
name.

It was stated in another section, that the Peruvians origi-
nally lived in the neighborhood of Quito, where they came
in contact with the Chibchas and other tribes who carried on
a trade with the Indians across the Isthmus. The Kechua
(Peruvian) word for maize is sara, or zara, and cherchi signi-

1 Proc. Amer. Philos. Soc, 1892, 67; Zeitschrift fiir Ethnologic. 1SS5, iQi-
- Brinton, Proc. Ami^r. Philos. .Soc , 1892, 23.

A Botanical and Economic Stndy.

127

fies roasted maize.' The Peruvians were surrounded on all
sides by wild and barbarous tribes, who learned from the
Incas a little agriculture and a few arts. The Tacanas,
Maropas and Araunas, situated on the Mamore River, have
words— dije, shije, zia, which are related to words of the
Pano stock, on the Ucayali River (see any good map). The
Pano word shequi is undoubtedly the Peruvian cherchi,
roasted maize, and clearly points to the source whence the
Panos and the Tacanas got maize.' It is probable that all the
inland and forest tribes borrowed the cereal from the Peru-
vians, as did the Indians along the Ucayali, the Mamore and
the Beni rivers. The following list of words shows this :^

Pano, Ucayali River, schequi.
Culino, Juvary River, tschiiky.
Moxoruna, Tapichi River, schuky.
Canawary, Purus River, schischy.
Araicu, Jurua River, metschy.
Carysuna, Madeira River, schroki.
Bare, Negro River, macanaschy.

The Tupis, or Guaranis, extend from the mouth of the La
Plata to the Amazon, and far up the stream of the latter.
The word for maize is abati in Guarani, auaty, abaty in Oma-
gua, awati in Cocama.* This word, or the aba of the Chib-
cha, clearly appears in Guiana, and is found apparently in
P'lorida, for the Timucua words are abo, corn-plant ; abopaha,
corn-crib; aboti, stick staff;' this similarity in words, how-
ever, may prove superficial to philologists competent to speak
on the subject. That the Arawaks carried these Tupi words
north with them is evident from the fact that certain Ara-
wak tribes had Tupi loan-words. Thus the Arawak tribe
Mariayos, on the Rio Negro, had yua-naty ; the Manaos,
on the Rio Negro, auaty ; the Maranhos, on the River Jatahy,
uaty.

Columbus when he landed found maize on the West India

> The most archaic forms of Kechua are to be found in the Chincasaya (Northern).
There the word for chicha is asua, and in Kechua proper, axa. Cf. American Race, 205
2 Brinton, Proc Amer. Philos. Soc, 1892,47; KansasCity Review, April, 1883
' Martins Brasilian Sprache.

< Ruiz, Lengua Guarani, 1876; Martius Brasilian Sprache, 427.
^ Proc. Amer. Philos Soc , 1883 15.

ifl

128

llarshbcrgcr. — Maize

A Botanical and Economic Study.

129

islands, and adopted the word mahiz or mayz. The Arawak
word for maize is marisi, and various forms of this word are
found among the other tribes.

{Arawak, marisi, Guiana.
I Cauixana, mazy, Rio Jupura.
Arawak. \ ^^^y/^^^ maique. Goajiros Peninsula.
yj'asses, mary, Lower Jupura.

Puri, maky, Rio Paraiba.

Coroado, maheky, Rio Paraiba.

Carib, marichi, marisi (female use).

Cuba, Jamaica, Lucayo, maysi.

The Caribs clearly borrowed their word for maize from the
Arawaks, for the Arawak radical is used by the females of
the Carib stock, the males using an entirely different noun.
The Arawak word was used largely through northern South
America and the West Indies, and is the one which the
Spaniards adopted as the word for the new and unfamiliar
grain which they saw and described.

The linguistic evidence shows: (i) That maize was intro-
duced into the United States from two sources, from the
tribes of northern Mexico and the Caribs on the West India
islands ; (2) that the Pueblos and northern Mexican tribes
deiived maize from central Mexico; (3) that tribal connec-
tions existed between the North and South American con-
tinents, and that an interchange of products was carried on
by way of the Isthmus of Panama ; (4) that the wild tribes
living along the Andean system, in the El-Gran-Chaco
and elsewhere used Peruvian loan-words for maize ; (5) that
South American words for maize were used throughout the
Greater and Lesser Antilles and Florida, and that the Arawak
word for Indian corn, adopted by Christopher Columbus, was
used by tribes of that stock in the impenetrable and luxuriant
Brazilian forests.

F. Historical Proofs.

The historical section has been divided for convenience of
treatment into four parts, as follows : Division A, History
Proper ; Division B, Aboriginal Cultivation ; Division C,
Indian Use ; Division D, Mythology. The material has been

arranged in geographical sequence, starting with New England
and concluding with South America, and where possible the
facts have been arranged chronologically.

Division i. History Proper.— About 1002 A.D ,' Thor-
wald, brother of Lief, wintered in Vinland. The following
summer, in proceeding ''occidentale latus circumire," Thor-
wald found the sea " valde insulosum " (Mingan Islands),
and on an island far westward saw a wooden crib for corn
(kornjhalmer).' Karlsefn, in 1006, in coasting along our
northern coast, brought back to the ship a bunch of grapes
and a new-sown ear of wheat (corn). At Hop they found
self-sown fields of wheat where the ground was low, and
vines where the ground was high. ''While lying against the
peninsula of Cape Cod— Furderstrand, Nausett Beach— Thor-
fin, that he might know the quality of the neighboring land,
sent out his fleet-footed servants to run for three days over
the region and return and report what they had seen, the
ship lying at anchor during their absence. They brought
back, one a bunch of grapes, the other an ' ear of corn.' They
had two months earlier seen the ' new-sown ' (Beamish) young
corn at Hop. Ear of corn is the translation of the Icelandic
word hveiti-ax (J. Tomlinson Smith). Beamish and Rev. Dr.
Slafter ' translated the same expression ' ear of wheat.' " Was
it Indian corn.? Ax, by itself, in Icelandic, is ear of korn
(Vigfusson), hveiti is wheat. Skeat, in his Etymological
Dictionary, states that the word wheat is Teutonic, and sig-
nifies white, and hveiti, no doubt, refers to the color of the
grain or kernel of corn of whatever kind. Hviti, is white, as
applied to the White River, in Icelandic (Henderson's map),
and hviti-ax would be white ear of seed.* It is doubtful if
this seed was maize."

1 Pickering. Chronological History of Plants, 665.

-Trans. New York Agric Soc, 1878, 46

"Slafter, Prince's Society.

^ Horsford. E. A., Discovery of .America by Northmen, 1888

•' We are told in a tradition of the Tuscarora Indians, who claim they arrived on the
Virginian coast about the year 1300, that they found there a race who knew nothing of
maize and were eaters of raw Hesh The Northmen, in the ye; r 1000, found the natives
of Vinland. probably near Rhode Island, of the same race as those they were familiar with
in Labrador. Eskimo is from the Algonquin word Eskimantic-eaters of raw flesh. -The
Archaeologist, i, 13

!

fi

■m

130

Harshbe}gcr. — Maize :

The French arrived in Canada in the year 1534. Cartier
sailed up the St. Lawrence, and in 1535 reached Montreal
(^sitii) in the midst of extensive corn-fields/ He found every-
where maize,' ''mil gros comme poix, parcil a celui qui croit
au Bresil, dont ils magent au lieu de pain." In the brief
record of the second voyage mention is made of its use by
the Indians. Champlain arrived in 1603 on the St. Lawrence,
and found the Five Nations at war with the Adirondacs.
The Adirondacs were hunters, but the Five Nations made
the planting of corn their business.'* He says,* "■ that when
coasting eastward from the River Ouinibequey (Kennebec)
he saw the Indians planting their ' bleds d'Inde,' and that
in every hill they put four Brasilian beans which grow of
many colors, and as they grow they wind about the corn,
which rises to the height of five or six feet. The soil was
feebly scratched with hoes of wood or bone."'^

The Puritans landed on the bleak and inhospitable New
England coast in 1620. Captain Miles Standish, with fifteen
others, at once set out to explore the country. We are told that
they very soon found ''much plain ground," about 500 acres fit
for the plough, and some signs where the Indians had formerly
planted their corn.'"' Later, they discovered a cache. Indian
corn carried them over the long dreary winter of 1620-21, for
it is mentioned in an early narrative "that they bought
greate stores of venison and eighte hogsheads of corne and
beanes." In the spring of 1621 the Puritans began to plant
their corne, in which service Squanto (an Indian; stood them
in great stead, showing them both ye manner how to set it,
and after how to dress and tend it. Also he tould them
excepte they got fish and set within these olde grounds, it
would come to nothing, and he showed them yt in ye middle
of April)."'' "We set the last spring some twenty acres of

' Delatield, Trans. New York Agric Soc, 1850, 382.
- Trumbull, J., Torrey Botan. Bull., vi, 86.

' Transactions Canadian Institute, i, Part 1, 90; Golden, History of Five Nations,
London, 1747; Bailey, J. M., Ensilage, 77 , Slafter, Ch-'mplain, Prince's Soc. Publ.

* Champlain, Narrative (final edition), 1632.

f' Champlain's Account. American Antiquarian, vii, 18.
•' Mourt's Relations, 125-130.
" Mourt's Relations, 79.

* Mass. Hist. Soc. Coll., 4, 111,1856, 100; Goode, G. 15 , American Naturalist, xiv,473.

A Botanical and Eavioinic Study.

131

Indian corne, and sowed some six acres of barley and pease."
. " Our corne did prove well, and, God be praised,
we had a good increase of Indian corne and our barley indif-
ferent good."' Before the Indians learned from the English
the use of a more convenient instrument, they tilled their
ground with hoes made of clam shells, for which purpose
they were well adapted.' In Edward Johnson's "Wonder
Working Providence of Sion's Saviour in New England,
being a Relation of the First Planting of New England, 1628,
London, 1854," he says : " Many thousands of these [herrings]
they used to put under their corn, which they planted in hills
five feet asunder, and assuredly when the Lord created their
corn He had a special eye to supply these his people's wants
with it, for ordinarily five or six grains doth produce six
hundred." Winthrop, in 1630, obtained 100 pounds of corn
from the sea side of Cape Cod.

The Connecticut colonies, founded at a later date than the
one at Plymouth, soon came into unfriendly contact with the
Indians, and in 1637 the Pequot war broke out. In the
history of the Pequot war, it is recorded " that the Pequots
had two plantations three miles asunder, and above 200 acres
of corn, which the English destroyed."' Roger Williams
states that in the war between the Narraghansetts and the
combined forces of the Mohegans and Pequots, the latter
committed extensive robberies and destroyed twenty-three
fields of corn. The Puritans in King Philip's War, in 1675,
took "what he had worth, spoiled the rest, and also took
possession of one thousand acres of corn, which was har-
vested by the English and disposed according to their direc-
tion."* Everywhere the Puritans found maize.'*

The Dutch PLast India Company sent out Hendric Hudson
on a voyage of exploration. Hudson, when anchored off the

^ Mourts Relations, London, 1622.

- Mass. Hist. Soc. Coll., vii, 193; Holmes, Bureau of Flthnology, n Rep., 207; Wood,
New England Prospects, 106.

•• Mourt's Relations, Drake, 116; 'Jrans. Wise Academy of Sciences, vi, 87.

< Drake, Old Indian Chronicle 209

•• Morton, New England Memorials. 1S26, 68; Gooken. Mass. Hist. Coll., Chap, iii;
Bradford History Plymouth Plantation, 82, 100; Mourt's Relations, Wood, New Eng-
land Prospects; Williams, Roger, Key.

iB:

132

Hatshbergcr. — Maize

Catskills in 1609 bought cars of Indian corn, pumpkins and
tobacco.' New Amsterdam was seized by the English, but
the French still claimed the northern portions along the St.
Lawrence. Marquis de Nouville, in his celebrated expedition
against the Seneca Indians, says: **0n the fourteenth of
July, 1685, we marched to one of the larger villages of the
Senecas, where we encamped. We remained at the four
villages of the Senecas ten days. All the time we spent in
destroying the corn, which, including the old- corn that was
in cache, which we burned, was in such great abundance that
the loss was computed at 400,000 minots, or 1,200,000 bushels.
This was in Ontario County, New York.' The French army,
under Frontenac, spent three days in 1696 destroying the
corn of the Onondagas."*

The Swedish settlements in New Jersey and Pennsylvania
were obliged to buy maize of the Indians for sowing and eat-
ing, as Kalm writes.' Quaint old Gabriel Thomas, writing
about 1696, tells us that ** they live chiefly on maze or Indian
corn, roasted in the ashes, sometimes beaten, boyled with
water, called homine."' Heriot refers to the cultivation of
maize in Virginia, in 1586, called by them pagatour.

Jamestown was founded in 1607. During this year, Captain
Newport, who commanded the colony, went up the Powhatan
River to visit the chief Powhatan, who he relates had exten-
sive fields that came down to the river's edge, in which fields
Powhatan cultivated corn, beans, pumpkins, tobacco. The
English at Jamestown were sustained by the liberality of the
natives. Pocahontas in person accompanied "conductas" of
grain.*

The Indians taught the settlers the use of maize." Captain

John Smith, in his *' Indians of Virginia," says: " The greatest

labor they take is in planting their corne, for the country is

. naturally overgrown with wood. To prepare the ground, they

' Trumbull, J. E.. Torrey Botanical Bulletin, vi, 86; Wise Academy of Sciences, 30.

-Aboriginal Monuments of New York, 63-66

^ Documentary History, New York, i, 212

^ Trans. New York Agric Soc, 1850-

■' Public Ledger, Dec 27. 1892, 3.

'Jones Antiquity of Southern Indians. 296.

" Jefferson's Notes on Virgiaia ; U.S. Patent Office Report, 1854, 98.

A Botanical and Economic Study.

133

bruise the barke of the trees neare the root, then doe they
scortch the roots with fire, that they grow no more. The
next year, with a crooked piece of wood, they beat up the
weeds by the roots, and in that mould they plant their corne.
Their manner is this : They make a hole in the earth with a
sticke \cf. "John Pope, his Tour," 1792], and into it they put
foure foote one from another the graine. Their women and
children do continually keepe it with weeding, and when it is
growne middle high they hill it about like a hopyard. In
Aprill, they begin to plant, and so they continue till the midst
of June. What they plant in Aprill they reap in August ; for
May in September ; for June in October. Every stalke of
their corne commonly beareth two ears, some three, seldom
any foure, many but one, and some none. Every eare ordi-
narily hath betwixt 200 and 500 grains. The stalke being
green hath a sweet juice in it somewhat like sugar cane, which
is the cause that when they gather their corne they sucke the
stalks."

Cabega de Vaca landed in Florida in 1528, near Tampa Bay.
He found there maize, beans and pumpkins in great plenty
and abundance.' De Soto set sail for the New World in
1538. He landed in Florida in 1539. De Soto frequently
speaks of the Indian villages, that contained from 150 to 200
dwellings, guarded sometimes with tall palisades, and sur-
rounded by extensive fields of maize, pumpkins and beans.
In one instance, he narrates that his army passed through
. continuous fields of maize for two leagues. His band soon ran
short of provisions, and the Indians were robbed to furnish a
supply. At one place, they took enough corn to feed the
Spanish company for five days. He writes: "On October
18, we came to Mobile, a walled city, which we captured, and
where we rested forty days. On March 3, we departed north
with maize enough for sixty leagues." ' From Tampa Bay,
De Soto addressed a letter to the justice and board of
managers in Santiago de Cuba, informing them that Baltazzar

• Torrey Botanical Bulletin, vi, 86; Cabe?a de Vaca Relations, 1528, translation by
Buckingham Smith, New York, I071
- Wise. Academy of Sciences, vi, 87.

m.

134

Ilarshbefgcr. — Maize :

de Galligos, who with eighty lancers and one hundred foot
soldiers he had sent to reconnoitre the country, had seen
fields of maize, beans and pumpkins, with other fruits and
provisions in such quantity as would suffice to subsist a very
large army without knowing a want. De Soto's officers
found in one place 500 measures of ground maize, besides a
large quantity of grain.'

Captain Ribaut and the French Huguenots sailed up the
St. John's River, Florida, in 1562, and founded Port Royal.
Captain Ribaut' says: ''About their houses they labor and
till the ground, sowing the fields with a grain called mahis,
whereof they make them meal, and in their gardens they
plant beans, gourds, cucumber, citrons, peas and many other
fruits and roots unknown to us."' In the narrative of Rene
Laudonniere (1564), he says of his' expedition from Fort Caro-
line, at the mouth of the River May (St. John's) : '* I departed
with fifty of my best soldiers in two barks, and arrived in the
dominion of Utma. distant from the river, where we took him
prisoner. They [his tribe], therefore, brought me fish in
their little boats and their meal of maize." Le Moine, in his
"Narrative," illustrated with drawings and written in 1564,
mentions maize. "I sent a second expedition with two
shallops, having soldiers and sailors aboard, with a present to
be given in my name to the widow of a deceased chief named
Hionacara, who lived twelve miles north of us. She received
my men kindly, and loaded both of our shallops for me with
maize and nuts, and sent in addition some baskets of cassine
leaves, of which they make a drink."

In Plate XXI, Brevis Narratio/ six Indians are seen pre-
paring the ground and sowing corn. The explanatory note
reads :

" Diligenter colunt terram Indi ; earn ob causam ligones e
piscium ossibus pararc norunt viri quibus manubria lignea
aptantes, terram fodiunt satis facile, nam mollior est : ea de-

' Lawson. History of the Carolinas. 296.
2 Jones, Antiquity of Southern Indians, 299
' The Whole True Discovereye of Terra Florida, London, 1563.
^ A Brief and True Report of the new found land of Virginia (Florida is meant here).
Thomas Hariot, 1590

A Botanical and Economic Study.

135

inde probe confracta et aequata feminae fabas et milium sive.
Mayzum serunt, pr^euntibus nonnullis, quae defixo in terram
baculo foramina faciunt, in quee fabee et milii grana injicia-

antur." •

A Jesuit missionary named Marquette, with a trader named
Joliet, and five other Frenchmen, started out to reach a great
river in the far west, of which much had been heard. The
explorers reachea the Mississippi, and sailed to the mouth of
the Arkansas. Marquette says: "The first was a dish of
sagamite, that is, some Indian meal boiled in water. "^ La
Salle, another Frenchman, built a canoe on Lake Erie and
paddled through the Lakes, in 1679, as far as Green Bay. From
there, by Lake Michigan, they went to the mouth of the St
Joseph, crossed the Illinois and made their way back by Lake
Ontario. Father Hennepin and another priest, during La
Salle's absence, went down the Illinois to the Mississippi,
and up this river to the Falls of St. Anthony. La Salle and
Hennepin met on the Illinois, for Hennepin, in his narrative,
states: "We continued our course (up) this river very near
the whole of December, toward the end of which we arrived at
the village of the Illinois, about one hundred and thirty leagues
from Fort Miami. We found nobody in the village, yet we
durst not meddle with the corn they had laid under ground
for their subsistence, and to sow their land with, it being the
most sensible wrong one can do them, in their opinion, to take
some of their corn in their absence. However, our necessity
being very great, and it being impossible to continue our
voyage M. La Salle took about forty bushels of it hoping
to appease them with some presents. We embarked again
with this fresh provision, and fell down the river the first of
January. 1680. We took the elevation of the pole, which
was 33° '45'."' Hennepin evidently accompanied La Salle in
1682 to the lower Mississippi. In his narrative he says :
" I was surprised to see their Indian corn, which was left very
green, grown already to maturity, but I have learned since
that their corn is ripe sixty days after it is sown." M. Le Page

• Contributions to North American Ethnology, v, 53.
2 Hennepin's Account, Reprint Trans. Antiq. Soc, i, 73-

136

IlanJiheixer, — Maize :

Dii Pratz ' states (page 226) that maize is the natural product
of this country. "Louisiana produces several kinds of maiz,
namely, flour maiz, which is white, with a flat and shriveled
surface, and is the softest of all kinds ; homony corn, which
is round, hard and shining — of this there are four sorts, the
white, the yellow, the red and the blue ; the maiz of these
two last colors is more common in the highlands than in
lower Louisiana. For the grinding of their corn they use
large wooden mortars, formed by hollowing the trunks of
trees with fire."

General Wayne, writing in 1783 about the Miamis, states
that along the river were endless maize fields, the like of
which he never saw from Canada to Florida."^

Cabe^a de Vaca traveled westward through Texas,'' and the
Indians supplied him with prickly pears and occasionally
maize, but after crossing a great river coming from the north
(Rio Grande), he came into a country "whose inhabitants
lived on maize, beans and pumpkins." On August 4, 1528,
" incursions were made with the people and horses that were
available, and on them were brought back as many as 400
fanegas of maize" ' [3200 bushels]. ^^ Friar Marco de Nica, in
1539, traveled through northeastern Mexico, and New Mexico.
"And they presented unto me many wilde beastes, as Conies,
Quailes, Maiz, Nuttes of Pine trees and all in great abun-
dance." " Coronado (1540) immediately set to work to explore
the adjacent country near Cibolo (Pueblos). Hearing there
was a province in which there were seven towns similar to
those of Cibolo, he dispatched thither Dom Pedro de Tobar
with seventeen horsemen, three or four soldiers and Friar
Juan Padilla, a Franciscan, who had been a soldier in his
yoath, to explore it. "The rumor had spread among the
inhabitants of a very ferocious race of people who bestrode
horses that devoured men, and, as they knew nothing about
horses, this information filled them with the greatest aston-

' Du Pratz. History of Louisiana, 1758, translation. London, 1763.

-' Arcli. fiir Anthropologie, 1886, 535; Carr, Lucien, Kentucky Geol. Survey, 11.

•' Trumbull, J. E., Torrey Botanical Bulletin, vi, 86.

■• Relation Cabeva de Vaca. Smith, N. \'., 1871, 47.

'' Fanega represents eight bushels.

" Hakluyt's Early Discovery and Voyages, 1600, 441.

A Botanical and Economic Study.

137

ishment."' They, however, made some show of resistance
to the invaders, in their approach to their towns, but the
Spaniards charged upon them with vigor ; many were killed,
when the remainder fled to the houses, sued for peace, offer-
ing as an inducement, presents of cotton stuff, tanned hides,
flour, pine nuts, maize, native fowls and some turquoises."'

Ferdinand Alargon discovered and entered the Colorado
River in the year 1542. He states "that he went up the river
eighty-five leagues, when his pirogue was arrested by lofty
mountains, through which the river ran, where it was impos-
sible to draw their boats ; and as far as they went they found
the Indians cultivating maize."

Columbus,' in his letter to Ferdinand and Isabella, dated
May 30, 1498, writes, "the most remarkable inference is that
they used maize, which is a plant that bears a spine (or awn)
[silk] like an ear of wheat, some of which [wheat] I took with
me from Spain, where it grows abundantly ; this they regard
as most excellent, and set a great value upon it." Columbus
saw the bread of the plant called mahiz.' In one of his letters,
he speaks of his brother : " During a journey in the interior
he found a dense population entirely agricultural, and at one
place passed through eighteen miles of corn fields." "

When the eastern coast of Yucatan was discovered maize
was found cultivated. Cordova, in 15 17, sailed from Cuba,"
and explored the north coast of Yucatan, where "the Indians
sowed maize, and were in possession of gold.'" The Spanish
Governor of Cuba later sent out an expedition (i 518), in the
same direction under the command of Grijalva, He explored
Yucatan and the southern shore of Mexico, and verified the
belief that there was a rich empire in the interior. The
Indians presented Grijalva with the bread of maize.' Cortez
landed in Mexico, and at Cempaolla ate bread of maize. He

> Castaneda's Relations, Ternaux Compans, 59

2 Smithsonian Report. iS'jg. 316.

• Ford. Paul L., Writing of Christopher Columbus. 1892, 125.

" Bailey. J. M., Ensilage, 77

■• Aborigines of the Isthmus of Panama; Popular Science Monthly, xxii. 427-

<' Bailey, J. M , Ensilage n-

' Agricultural Production, U.S. Census. 1880. Maize.

>' Agricultural Production, Tenth Census, 1880, Maize.

h i:

it

138

HarsJibcrg er, — Maize

found Geronimo, a Spaniard, digging as a slave in the maize
fields. On the road to the City of Mexico they passed
through large fields of waving corn, where the Aztecs
ambushed themselves, and shot arrows at the cruel and hated
invaders.*

The following extracts taken from a variety of sources
will prove sufficiently the general cultivation of Zea mays.

"The Indians of Yucatan put the maize into water with
lime to steep over night. It is then given to carriers, who
make it into large balls or pellets, which they take with them
for food." '' Villagutierre^ says: ''Grandes milperias, en que se
dans dos cosechas de frutos, consecutivos al aiio y las mazorcas
y granos de maiz en extremo gruessos." Clavigero * contains
the substance of the earlier writers. He says:^ '* In the labor
of the fields the men assisted the women. It was the business
of themen to dig and hoe the ground, to sow, to keep the earth
about the plants, and to reap [contrast the Iroquois, among
whom the squaws tilled the soil] ; to the women it belonged to
strip off the leaves from the ear ; and to clean the grains ; to
weed and shell it was the employment of both."

** In the market places in Nicaraugua the women sell
slaves, maize, fish, deer, etc." " Maize and cacao are the
principal objects of barter, and form media of exchange."'*
" The men were above all things farmers, and cultivated
maize and other crops." ^ There was a mountain tribe who,
in burying a corpse, crooked the legs, and put the head on
the knees, and tied it tightly, so that it would remain in this
posture; then they digged a round hole and put the body in
it ; round the body they put food, a chocolate cup, a calabash
with atole, salvados of maize, and large maize tortillas ; the
whole was then covered with earth. The salvados were
destined for the animals that the dead ate ; the tortillas for
the dogs that were killed and eaten.

* Cortes, Cartas, 64; Torqiiemada. Monarq. Ind., 1, 515.

- Landa, § 21.

' Villagutierre, Hist. Itza y Lacandon, Lib. 8, cap. 12, Madrid, 1761.

^ Clavigero, Storia Antica de Messico, 1780.

*'• Bk VII, ch 28.

«'• Oviedo, Hk. xi.ii, ch. 3 (1535).

" Landa, § 21.

A Botanical and Economic Study.

139

"The food of the poor Indian was maize." ' '* Among the
Indians of Huancavilcas, the best-flavored maize bread is
made in all the Indies." ' " As no maize can be raised at the
elevation of Callao the people obtained their supply by the
Titimaes, who brought up loads of maize, cacao and fruits of
all kinds, besides plenty of honey" (Cieza, ch. 99). The
Incas extended the maize land over the newly conquered
provinces by the construction of irrigating canals and ter-
races.^

Garcilasso de la Vega, the last of the Incas, whose accounts
are highly overdrawn and must be received with caution, says*
that the palace gardens of the Incas were ornamented with
gold and silver figures of maize, and in one case he tells us
that there was an entire corn-field of considerable size, repre-
senting maize in its erect and natural shape.

Division 2. Aboriginal Cultivation. — The Indians of
North America followed very closely one method in the
sowing and cultivation of their maize; in the hot, arid districts
the practice necessarily differed from that pursued in more
favored localities, but broadly speaking the methods were

one.

The Navajos and Mojaves, of New Mexico, planted their
corn with a pointed stick.' The Moquis, of Arizona, in rais-
ing corn, are beset with annoyances, like drought and flood.
The soil is thin and sandy, with very little moisture at the
top, because of the evaporation induced by the heat and dry-
ness of the atmosphere. The under strata of clay and sand-
stone, however, retain the moisture a long time. The Moqui
farmer consequently buries the seed deep in the ground. He
takes his planting stick (see illustration) in his right hand,
and presses on the horizontal bar with his foot, making a

1 Pizarro, P., Relacion de la Descubrimento y Conqueste Peru, 1529, 379-

2 Cieza, Travels, 1532, ch. 114.

8 Garcilasso de la Vega, 1609, Bk. v, ch. 5.

< Garcilasso, Com. Real, Lib. 8, cap. 26; Sarmiento Relacion, MSS., cap. 24; Sturte-

vant, New York Agric Soc, 1878, 45- . , ^rr r. c. i-

^ Emory, Rep. U. S. and Mexican Bound. Surv., i, 112; Ind. Off. Rep. bpec Com.^

1867, 337; Merriwether, Ind. Off. Rep., 1854, 172
10

<

140

HarsJibcrger, — Maize :

hole in the ground from twelve to eighteen inches deep, in
which he drops the kernel of corn. As a guard against

A Botanical and Economic Study,

141

floods and winds the corn is planted in bunches. In the
Moqui fields five, six, and even ten stalks will be seen
2:rowinjr close to^xether with another cluster ten feet off,
and so on ; each cluster is almost surrounded at the foot
by small branches, wisps of hay, and little piles of mud
brought down by rain currents.*

The Mexicans, having neither plows nor oxen, used a hoe
called coatl, made of copper, fitted with a wooden handle.
The fields, in many cases, were surrounded by stones and a
hedge of aloes. The maize sower drilled the ground with
a stick and dropped the grain, covering the kernel with his
foot.'

The production of maize was manifold. Gomara"^ cites the
yield as 100 to 150 fold : " Suelo dar una hanega de maiz en
sembradura seys, diez, veynte, treynta, cinqiienta, ciento e
aun ciento e cinqiienta c mas e menos hanegas segund la
fertilidad e bondad de la tierra donde se siembra." ' Hum-
boldt says : '' In the fine plain between San Juan del Rio
near Queretaro, one bushel produces eight hundred bushels.
It can be laid down, as a general rule, that the production of
this cereal in New Spain was about 150 fold."

Division 3. Indian Use. — Dr. Franklin mentions one of
the methods that the Indians adopted in the preparation of
their maize as an article of food. A vessel of sand was heated.
The corn was then mixed with this sand and slowly heated

^ Bourke, J. G., Moquis of Arizona, 96; Loew, Oscar, Popular Science Monthly,

V, 354-

s Clavigero, Bk. vii, ch. 28.

■'Steffen, Die Landwirthschaft beiden Alt Amerikanischen Kulturvolkern, Leipzig,

1883.

< Oviedo, Hist. Gen., Lib. 7, cap. i.

until the grain burst. It was then taken out and ground to a fine
powder, which kept fresh for a number of years. Heckewelder*
called this preparation psindamocan. Captain John Smith,
in his narrative states that **they rost their corne in the eare
greene, and bruising it in a mortar of wood with a polt, lap it
in rowles in the leaves of their corne, and so boyle it for a
dainty." "They also preserve their corne late planted that
will not ripe, by roasting it in hot ashes, the heat thereof
drying it. In winter they esteem it, being boiled in leaves,
for a rare dish they call pausarowmena." Heckewelder' says:
"That the bread is of two kinds, one made of milk corn, the
other of the dry and fully ripe. The last is finely pounded
and sifted and kneaded into dough. This dough is made into
round cakes about six inches in diameter and about an inch
thick. These cakes are baked in the cleanest of wood ashes.
Frequently they mixed with the dough pieces of pumpkin,
beans, chestnuts, whortleberries and other palatable ingre-
dients. They make an excellent pottage of their corn by
boiling it with fresh or dried meat (the latter pounded), dried
pumpkins, beans or chestnuts. The pottage is sometimes
sweetened with the sugar or molasses from the maple tree.
A very good dish is made by boiling well-pounded hickory
nut kernels with the maize. The nut liquor gives a rich and
airreeable flavor to the food." LoskieP mentions that the
Indians frequently mixed smoked eels and shell fish, chopped
fine, with their corn-meal in the making of bread. Their
common food, says Captain Bernard Romans,* is Zca, or
Indian corn. "They make meal; they boil it; they parch
and then pound it, taking this pounded material on long
journeys." "They also have a way of drying and pounding
their corn before it comes to maturity ; this they call boota-
copassa, i. e., cold flour. This, in small qu?.ndties, thrown
into cold water, boils and swells up, as much as common
meal boiled over a fire. It is a hearty food, and, being sweety

1 Heckewelder, Indian Nations, 195.

-' Heckewelder, Indian Nations, Mem. Hist. Soc Penna., xii, 195.

3 Loskiel, Mission North American Indians, Lond., 1794; Abbott, C. C , Primitive

Industry, i88i, 149

^ Romans, Concise Natural History of East and West Florida, New York, i775i w.

142

HarsJiberger. — Maize :

they are fond of it." Du Pratz describes the making of what
^he calls farina froide. The corn is first parboiled in water;
the water is drained off and the grain is dried. The dried
kernels are roasted on a plate, ashes being mixed with them
to prevent burning. The grains soon take on a red color
with constant stirring, and are then removed and well rubbed.
They are then mixed with the ashes of the dried stalk of
kidney bean and a little water added, and the mixture is
thoroughly pounded into meal. The dough is thoroughly dried
in the sun, and when wanted for use is mixed with about two-
thirds water. It affords a nourishing and satisfying food.

Dr. Edward Palmer' describes the use of maize among the
Indians of the southwestern States. The Apaches cook their
maize by dropping hot stones into the vessel containing the
ears or grain. They ferment a drink from the grain which
is strong enough to affect the Indians powerfully. He
describes the uses of the cereal among the Pueblos. The
blue grains are rubbed in a stone mortar and give a meal of a
bluish-gray color. A thin dough is then kneaded with this
meal. A hot fire is kindled, over which a flat stone or iron
plate is laid. The women, with the fingers of the right hand,
together dip them into the dough and draw them out again
thickly covered, and spread the paste in a thin layer on a
hot stone or plate. The mass quickly puffs up, a sign that
one side is done, and it is then turned over, while the second
cake is placed on to bake. Finally the flap-jack is rolled
over and finished. When eaten it appears at first dry, but is
sweet and easily chewed. A second method of preparation
is to cook in lime water until the hard covering is removed.
It is then pounded to a white flour and made into bread.
The " enthiilste korn " is often cooked with pieces of meat,
and red or green pepper is added. When this is baked in
the husks it forms what is called by the Mexicans tomale,
. The maize meal, when cooked with the sugar from the mes-
quite {Prosopis juliflora D. C), constitutes the dish called
pinole. The crude meal is often made into a kind of bread,
wljich the Spaniards call tortillas.

1 Palmer, Monatschrift f. Gartenbaues, 1874, 163, cited.

A Botanical and Economic Study.

143

The Mexicans made their bread in the following way : 1 hey
put a large pot filled with water on the fire, which they
allowed to remain until the water boiled ; then they put out
the fire and poured the grain into a pot. A little hme was
added to destroy the skin of the corn, and next mornmg they
carefully cleansed the grain, ground it by stones, and, moist-
ening the meal, formed it into a paste, which they kneaded
and baked into tortillas.' Tortillas of maize, accompanied by
the inevitable f rijoles, or beans seasoned with chile or pepper
and washed down with drinks prepared from the maguey and
cacao, were the all-sustaining diet of the Nahuas.^

The Mexican drink, -chicha," is made from maize. A
quantity of grain is soaked in water, and is taken out and
sprouted. The sprouted grain is bruised and placed in a
large vessel filled with water, where it stays until it begins
to ferment. A number of old women then collect and che^^
some of the grain until they have a sufficient quantity
which is added to water and allowed to ferment. The fluid
is drawn otf after fermentation, and a strongly intoxicating
liquor is produced.

Division 4. MvTHOLOGY.-It has been frequently asserted
that the Indian had no religion excepting what has been called
- the meaningless mummery of themedicine man. /^is is
the very reverse of the truth. The Indian was essentially relig-
ious and contemplative, and it might be said that every act of
his life was regulated by his religious belief, as the following
accounts of ceremonies, myths and legends show. George
Catlin ' has described the green corn dance, as practiced by the
Minnetaree : - The green corn is considered a great luxury by
all those tribes who cultivate it. This joyful occasion is one
valued alike and conducted in a similar manner by most of
the tribes who raise corn, however remote they may be from
each other. It lasts but for a week or ten days, being
limited to the longest terms that the corn remains in this

1 Brocklehurst, Mexico of To-day, 1883, 200.

2 Bancroft, Native Races Pacific States, 347-

••' Smithsonian Report, 1885, Part 11, Catlin Indian Gallery, 314.

144

Har'shbeixc7\ — Maize :

tender and palatable state, during which time all hunting, and
all war excursions, and all other avocations, are positively
dispensed with, and all join in the most excessive indulgence
of gluttony and conviviality that can possibly be conceived.
The fields of corn are pretty generally stripped during this
excess, and the poor, improvident Indian thanks the Great
Spirit for the indulgence he has had, and is satisfied to ripen
merely the few ears that are necessary for his next year's
planting, without reproaching himself for his wanton lavish-
ness, which has laid waste his fine field and robbed him of
the golden harvest which might have gladdened his heart,
with those of his children, through the cold and dreariness of
winter."

The time of the harvest was also a season of festival and

rejoicing, when elaborate ceremonies were performed. The

festival was known among the Creeks as the Boos-ke tau

(time of maturity), or, for short, in English, the " Festival of

the Busk." Colonel Benjamin Hawkins has left us an

account of this feast -} " On the morning of the first day the

warriors clean the yard of the square and sprinkle it with

white sand. The acee, or decoction of the cassine yupan

(Jlex cassine), is made. The fire-maker kindles the fire. as

early as he can by friction. Four logs, each as long as a

man can cover by extending his two arms, are cut and

brought by the warriors and placed in the centre of the

square, end to end, thus forming a cross. The outer ends

indicate the cardinal points of the compass. In the centre

of the cross the new fire is made. These four logs are burnt

out during the first four days. The Pin-e-bun-gau (turkey

dance) is danced by the women of the turkey tribe, and

while they are dancing the possau is brewed. This is a

powerful emetic. From twelve o'clock to the middle of the

afternoon the possau is drunk. After this four men and four

women dance the Toc-co-yuh-gau (tadpole) from evening

until daylight. E-ne-hau-bun-gau (the dance of the people

second in command; is danced by the men. About ten

1 Hawkins, Sketch of Creek Country, Coll. of Georgia Hist. Soc , in, 75 ; Jones'
C, C. Antiquity of Southern Indians, 303.

A Botanical and Economic Study.

145

o'clock the second day the women dance Its-ho-bun-gau (the
gun dance), and after twelve o'clock the men go to the new
fire, take some of the ashes, rub them on the chin, neck and
body, jump head foremost into the river, and then return
again into the square. The women having prepared the new
corn for the feast, the men take some of it and rub it between
their hands, and on their faces and breasts, and then feast."

'' During the third day the men sit in the square. Early
in the morning of the fourth day the women get the new
fire, clean out the hearths, sprinkle them with sand, and
kindle their fires. The men finish burning out the first four
logs, and then, rubbing themselves with the ashes on their
chins, necks, and bodies, go into the water. This day salt is
eaten, and they dance O-bung-gau-chap-co (the long dance).
The fifth day four new logs are brought and placed in the
same position as on the first. They drink also acee, the
strong decoction of the cassine yupan. During the sixth
day they remain in the square, and on the eighth, they get two
large pots and their physic plants, and beat up with water.
The chemist (E-lic-chul-gu) blows into the decoction through
a small reed, and then the men drink it and rub it over their
joints until afternoon. They then collect old corn-cobs and
pine-cones, and placing them in a pot burn them to ashes.
Four virgins bring ashes from their houses, and having put
them intl) the pot, stir all together. The men take white
clay and mix it with water in two pans. A pan of this clay
and one of ashes are carried to the cabin of the Mico. Two
pans similarly filled are taken to the cabin of the warriors ;
with the clay and ashes they rub themselves. Two men,
appointed to that office, bring flowers of tobacco of a small
kind (itch-au-chu-le-puc-pug-gee), or, as the name imports, the
old man's tobacco, which was prepared on the first day, and
putting it in a pan in the Mico's cabin, give a little of it to
all who are present. The Mico and the counsellors then go
four times around the fire, and every time they face the east
throw some of the flowers into the fire. They then go and
stand to the west ; the same ceremony is repeated by the
warriors. A cane is stuck up in the cabin of the Mico, with

146

Harslibci'ircr. — Maize :

two white feathers in its end. A member of the fish tribe
(Thlot-lo-ul-go) takes it just as the sun goes down, and moves
off toward the river, all following him. When half way to
the river he gives the death whoop. This he repeats four
times between the square and the water's edge. Here they
all locate themselves as close together as they can stand.
The cane is stuck up at the water's edge, and they all put a
grain of old man's tobacco on their heads and in each ear.
At a given signal, four times repeated, they throw some of
this tobacco into the river, and every man upon a like signal
plunges into the stream and picks up four stones from the
bottom. With these they cross themselves four times on the
breast, each time throwing a stone into the river and giving
the death whoop. They then wash themselves, take up the
cane with the feather, return and stick it up in the square,
and visit through the town ; at night they dance the mad
dance, and this ends the ceremony."

The Zuiiians were divided into fifteen clans, organized
chiefly for social intercourse and amusement, which are of
very ancient origin. These clans corresponded in no way
with the division of the people into gentes. The corn clan
was especially sacred among the Zuiiians. The Moquis and
Pueblo Indians of San Felipe, Santa Anna, Zia, Zemez,
Cochiti and Isleta had corn clans.^ During the frequent
ceremonies carried on by this clan, the songs sung were in a
language long since lost.^

Gushing' has vividly described the dances of the Zuiii, and
the important use that corn has played in all their mystic
ceremonies. He has described the mythological creation
and origin of corn as narrated by the Zunians. " Yet, not
less precious was the gift of the seed people or Ta-a-kwe.
This was Tchu'-e-ton, or the medicine seed of corn, for from
this came the parents of flesh and beauty, the Solace of
Hunger, the Emblem of Birth, Mortal Life, Death and Immor-
tality. Born before our ancients had been other men, and

' Journ. Amer. Folk Lore, iii, 1890.

'Smithson. Miscel. Col. Philos. Soc , Wash, iv, v ; Trans Anthrop. Soc,
XXV. 88.

3 Gushing, Zuni Bread Stuffs, The Millstone, Indianapolis, Vol. ix, Jan, 1884.

A Botanical and Economic Study,

H7

these our fathers sometimes overtook and looked not peace-
fully on them. It thus happened, when our ancients came to
their fourth resting place on their eastward journey, that
they named Shi-po-lo-lon-kaia, or The Place of Misty Waters
there already dwelt a clan of people called A'-taa, or Seed
People, and the seed clan of our ancients, challenged them
to know by what right they assumed the name and attributes
of their own clan. It is well, replied the strangers, yet life
ye did not bring. Behold! .... Behold, indeed where
the plumes had been planted and the tchu'-e-ton placed, grew
seven corn-plants, their tassels waving in the wind, their stalks
ladened with ripened grain. These, said the strangers are the
severed flesh of seven maidens, our own sisters and children.
The eldest sister was yellow corn ; the second, blue ; the third,
red ; the fourth, white ; the fifth, speckled ; the sixth black ;

the seventh, sweet corn Aye, we may, replied the

strangers, and of the flesh of our maidens ye "^^Y ^^t, no
more seeking the seeds of grasses." Gushing thinks that the
strangers A'-ta-a. or Seed People, were the Keres to the east
of Zuiii, who taught the Zuni the use of corn.

Bourke^ describes the snake dance as practiced by the
Moquis: ^* The women extend their line fully, all the while
scattering corn-meal. A portion occupy the terrace directly
above the arcade, a few on ladders near the archway ; the
main body, however, stand in the space between the sacred
rock and the sacred lodge. Two or three, reinforced by a
lot of old cronies, do effective work at the eastern end of the
rectangle Nearly all carried the beautiful woven baskets,
ornamented in yellow and black with the butterfly and thun-
der-bird painted on the side. The baskets were filled with
finely ground corn-flour, which was scattered with reckless
profusion into the air and upon the reptiles as fast as thrown

down." . ... • ut u^

" This corn-meal has a sacred meaning, which might be
well to remember. Every time the corn was scattered the
lips of the squaws moved, as if in prayer. A sacred meal is
prepared of corn-meal and chalchihuitl, called cunque. All

1 Bourke, Moquis of Arizona, 163

i

148

Harshbenrer. — Maize :

the Pueblo Indians along the Rio Grande use it, and upon
rising in the morning throw a pinch of it to the east. The
Zunis and Moquis are never without it, but carry it in little
bags tied to their waists." The use of this meal resembles
the crithomancy of the Greeks, but is not identical with it.
Space will not permit any further account of the use of maize
in the ceremonial religious observances of the Pueblo Indians,
but the reader is referred for more detailed accounts to

A Journal of American Archcuology and Ethnology, Vol. ii ; (A Few
Summer Ceremonials at the Tusayan Pueblos ; Hemenway Expedition.)

The Millstone, Indianapolis, Ind., 1884— Article, Zuni Bread Stuffs;
Cushing.

The Aztecs worshipped several deities. Tlaloc, for instance,
was the god of waters, of rain, and of the fertilized earth,
Ixtlixochitl represents him in the picture of the month Etzalli
with a cane of maize in one hand and in the other an agricul-
tural implement.^ Centeotl (Ceres) was the goddess of maize,
and, consequently, from the importance of the grain, the
goddess of agriculture and of production generally. Many of
her names seem dependent on the varying aspects of maize.
The fruits of the field were consecrated to her.- The feast to
this goddess, begun April 27, was elaborate, and maize was
used in a variety of forms at v^arious stages of the perform-
ance. Bancroft has given a detailed account, and those who
wish a fuller description can find it in his colossal work, *'The
Native Races of the Pacific States."

The Mayas similarly used maize in their festivities. The
Cakchiquels, in Guatemala, were a Maya tribe. " A little
more, and they would make a god of maize," says an old
writer. All the labors of the field were conducted with
religious rites. The men, for instance, who did a large part
of the field work, refrained from approaching their wives for
some days before they planted the corn. Incense was burned
at the corners of the field to the four gods of rain and wind

' Bancroft, H. H., Native Races Pacific States, in, 325.

2j0s.de Acosta, Nat. Hist. West Ind., Lib 4, cap. 16, 236; New Vork Agric Soc,
184S, 682; Trans. Illinois .^gric Soc, 1856-7, 473; Trans. New Vork Agric Soc . 1878, 47.

A Botanical and Economic Study.

149

before weeding the plot. The first fruits were consecrated

to their deities.^

The Peruvians, during the feast Capacraqui, in the first
month Raymi, permitted no stranger to lodge in Cuzco. In
the early portion of the month Hatuncuzqui, corresponding
to our May, the Peruvians gathered their maize, and kept the
feast Aymorai. " They returned home singing from the
fields, with them a large heap of maize, which they called
Perua, wrapping it up in rich garments. They continued
these ceremonies for three nights, imploring the Perua to
preserve their harvest of maize from any danger."'

This historical review shows conclusively that maize was of
all plants the one used universally by the Indians.

G. Summary and Recapitulation.

A glance at the American continents a century before the
Columbian voyages shows the greater portion of the con-
tinental areas occupied by hunter tribes just emerging from a
wild nomadic life and entering upon a partial sedentary agri-
cultural condition. In the eastern United States, the trees
were girdled by the stone axes of the aborigines, seed was
sown between the trees, and corn planted in these forest clear-
ings and on the rich river bottoms grew luxuriantly. The
prairie was inhabited by nomadic tribes, who made buffalo
hunting their principal business, while here and there over
the broad surface of the central plain, tribes more sedentarily
inclined, as the Mound-builders of the Ohio and the Pawnees
of Louisiana, cultivated maize. A little further to the west,
in the arid tracts of the West, lived tribes in storied structures
of adobe, who raised their crops by irrigation of the simplest
description. Closely allied to these Pueblo Indians, in the
common derivation of their house styles, were the timid cliff-
dwellers, who hid themselves in the caves and pockets of the
canon sides. Far to the south, on the plains of Anahuac,

1 Ximenez, Francisco, Las Historias del Origin de los Indios, London, 1857, "91 ;
lirinton. Annals of the Cakchiquels, 13. , , .

2 Trans. New Vork Agric. Soc, 1878, 47 : Browne, D. J., Trans. New ^ ork Agnc.

Soc, 1848, rxjo.

fl

I

ISO

HarsJibcrgcr. — Maize :

dwelt a people with established armies, central government
and populous cities, with temples, palaces and market places,
the latter supplied with the fresh produce of the surrounding
country. The Nahua civilization, which reached so high a
plane, was, nevertheless, preceded by one which in many
respects excelled that of the Aztecs, and excavations clearly
attest to the vigor and numerical strength of this peaceful
agricultural race. /

The different tribes of the American race all showed pecu-
liar individual idiosyncrasies, but linguistic study shows that,
with all this diversity, American agriculture was borrowed
from a common source — the Mayas of central Mexico.

Philological comparisons show that the Indians east of the
Mississippi, the Iroquois, the Mound-builders, the Algonquins
and the Muskogees obtained maize from across the " Father
of Waters," probably from the Caddos, who in turn derived it
from the northern Mexican tribes. The Pueblos, as archae-
ology and ethnology seem to prove, are only a few centuries
removed from the wild state exhibited by the roving Apaches
and Navajos, and, therefore, as compared with that of the
Mayas, their agriculture is comparatively recent.

Philology places the Nahuas in the Shoshonean stock with
the poor root-digging Ute of the plains. Their warlike pro-
pensities and love of conquest carried them south, until they
reached the plateau of Anahuac, when, attracted by the
peaceful and promising surroundings, they laid aside their
savage life and copied the superior civilization and agriculture
of the tribes (Maya) about them.

Archeology and ethnology both place the Mayas in the
vanguard as husbandmen, and to reach this development
required considerable time ; the cultivation of Yucatan and
southern central Mexico, as the permanent seat of this race,
antedates the tilling of the soil by the Peruvians' on the one
hand, or the pueblo-builders on the other.

Hieroglyphics' on the monuments at Palenque, indicate
that maize was the chief food of the people of Yucatan ;

» The Standard Natural History, Vol. \ i, 219, corroborates this statement.
'- Science, xxi, 1893. 8.

A Botanical and Economic Study.

151

here it was first used and distributed to the surrounding
tribes, who by barter carried it to the farthest limits of the

continents. , "^

The evidence of archeology, history, ethnology and (
philology, which points to central and southern Mexico as J
the original home of maize is supported by botany and /
meteorology. All of the plants closely related to maize are
Mexican. It is an accepted evolutionary principle, that
• several species of the same genus, or genera of the same
tribe though dispersed to the most distant quarters of the
o-lobe must originally have proceeded from the same source,
as they are descended from the same progenitor. It is also
obvious, that the individuals of the same species, though now
in distant regions, must have proceeded from one spot where
their parents were first produced; for it is incredible that
individuals, identically the same, should have been produced
from parents specifically distinct. Applying these pnncipes
to maize, we reach the conclusion that maize was originally
Mexican. Monotypes and genera, which contain but a few
species have, as a rule, a very restricted area. Zca is mono-
typic, and is singularly unprovided with means of dispersal,
so that the area of its original home must have beeil espe-
cially circumscribed. The discovery of a very primitive form
in Mexico aids in determining the wild limits of the species.
Meteorology helps in fixing the area more definitely. Th^ ques-
tion naturally arises, in what part of southern central Mexico
did the Indians first find the maize plant } Its original home
must not be looked for in low-lying districts nor in forests, for
maize does not thrive in warm, damp climates, where manioc
is grown.^ The region above 4500 feet altitude and south of
twenty-two degrees north latitude, and north of the River
Coatzalcoalcos (ninety-four degrees west, seventeen degrees
north ) and the Isthmus of Tehuantepec, fulfills more nearly the
conditions which the wild form required for its development.
The evidence to the present date places the original home
of our American cereal, maize, in central Mexico.

1 Sogot, Cult, des Cereales de la Guyane, Franc. Journ. de la Soc. Centr. d'Hortic.
de France, 1872, 94-

II

h ^\

152

HarsJiberger. — Maize :

Notwithstanding the indisputable fact that maize is of
American origin, and the statements of such men as Dodoens,
Camerarius, Matthioli, Gerard, Ray, Parmentier, Discourlitz,
Bonpland, De Candolle, Humboldt, Darwin, F. Unger, von
Heer, James, Targioni-Tazzetti, Hooker, Figuer, Nuttall, Mrs.
Summerville and Flint, the contrary that the cereal is of east-
ern origin has been asserted. Bock, 1532, Ruellius, 1536,
Fuchsius, 1542, Sismondi, Michaud, Gregory, Lonicer, Reg-
nier, Viterbo, Tabernamontanus, Bonafous, St. John, De- *
Herbelat and Klippart have argued for an Asiatic origin. A
discussion of this question is interesting historically.

The ancient authors on agriculture, Theophrastus, Varro,
Columella, Pliny, Palladius, Galen and Dioscorides, do not
mention maize. The principal argument for an eastern origin
is based upon a charter of the thirteenth century, the Chart
of Incisa, according to which the Crusaders, in 1204, gave to
the town of Incisa a piece of the true cross, and a purse con-
taining a seed of a golden color. Comte de Riant' has shown
this charter to be a fictitious fabrication of a modern impostor.

Such names as Turkish wheat, Sicilian corn, Spanish corn,
Guinea corn, Roman corn, have been used in the argument
for the eastern origin of maize, but they prove no more than
the word turkey (coq dTnde) argues for the Turkish origin of
the American fowl.

The total silence of the travellers who visited Asia and
Africa before the discovery of America, also militates against
an eastern origin. Notwithstanding this, China has been
frequently called the original home of maize ; this im-
pression became current because of an illustration in an
ancient Chinese work on natural history. Dr. Bretschneider,^
an authority on Chinese cultivated plants, does not hesitate to
say that Indian corn is not indigenous to China, and Shigeno
Aneki has undertaken to prove how in Japan certain historical
episodes were "cooked" under the Tokugawa dynasty of
Shoguns. *' A little reflection will show that such manipula.
tions of history are likely to be the rule rather than the

' Riant, La Charte d'Incisa, 1877.

'-' Bretschneider, Study and Value Chinese Botan. Works, 7, 18.

A Botanical ajid Economic Study.

153

exception in Asiatic countries. The love of truth for truth's
sake is not a general human characteristic, but one of the
exceptional traits of the modern European mind, developed
slowly by many causes, chiefly by those of habits of accuracy,
which phvsical science does so much to foster. Outside
Europe and her colonies, it is easy to manipulate records,
because such manipulation shocks no one deeply, because the
people are told nothing about the matter, and because, even
if they were told, they have neither the means nor the inclina-
tion to be critical."' Siebold places the home in Japan, ^ but
Rein ' speaks unhesitatingly against a Japanese origin. Maize
is not used largely as a food in Japan, and but two varieties
are known there. The Japanese name (to-morakoshi or nau-
bau-kibi, grain of the southern barbarians, Portuguese or
Spaniard) clearly indicates a foreign origin.

All these arguments come to naught when thoroughly
sifted, and do not in the least militate against the American
origin of maize.

1 Chamberlain, Things Japanese, 164.

- Siebold, Verhandl. Batav. Genotsch, xii.

'■' Rein, Peterniann's Geogr. Mittheil, 1878, 215-17.

I

154

Harshlyerger. —Maize :

CHAPTER III.

Geographical Distribution.

\A AIZE originated in all probability in a circumscribed
/ \ locality, above 4500 feet elevation, north of the Isthmus
of Tehuantepec and south of the twenty-second degree
of north latitude, near the ancient seat of the Maya tribes.
There is hardly a doubt but that the Mayas first cultivated
maize and distributed it in every direction. The time that
this people emerged from obscure savagery is not known, but
it was not earlier than the advent of the Christian era. This
places a time limit on the cultivation of maize. From the
Mayas, the use of the cereal spread north and south. The
Nicaraguan and Isthmian tribes obtained it from tribes
farther north. The Isthmian Indians traded with the Chib-
chas, who were in close commercial intercourse with the
Peruvian State, in the region of Quito. The indomitable
Inca race enlarged its territory by conquest until its influ-
ence, dominion and agriculture extended to the farthest limits
of Chili. Comparative philology affords definite proof that the
wild tribes in the El-Gran-Chaco and on the Cordilleras learned
their use from the Incas, for the tribes along the Ucayali,
Mamore and Beni rivers have Peruvian loan-words for maize.
The Arawaks, who later peopled the West Indian Islands,
knew maize when still in their primitive home in the Bolivian
highlands, and it is probable that their knowledge of agricul-
ture was derived from their more cultured neighbors on the
Pacific coast plain. The Arawak words for maize used by
the tribes on the islands and in central South America are
identical in form in many cases, and it is surely safe to say,
therefore, that Indian corn was carried by the Caribs and
Arawaks from the South American continent by way of
Guiana and the Greater and Lesser Antilles to Florida.

A Botanical and Economic Study.

155

The Nahuas borrowed their arts, sciences and agriculture
largely from the Mayas. Agriculture, the chief occupation
of the Aztec race, spread to the Pueblo Indians on the Rio
Grande River, and from there it extended eastward to the
Mississippi. A comparative linguistic study shows that the
Chahta-Maskokis had loan-words, indicating that the cereal
came from the West across the *' P'ather of Waters." Some
of these Southern tribes are identical with the mound builders,
who were driven south by the Iroquois-Algonquian eruption.
During times of movement and conquest the impulse is power-
ful toward imitation, and inter-tribal imitation is even more
strong than that between one individual and another. It is
easy to show how, under the bracing influence of race com-
petition, the forces of change would operate to initiate new
habits and progress toward a higher state of existence. This
tribal interchange of culture happened when the Algonquins
and Iroquois moved southward, where they simultaneously
learned the germs of agriculture.

The northern extension was limited by the isothermal line
of 50° F. (13° C), or the latitude of the great lakes. Corn
can be raised in Maine with certainty a few miles south of
Umbagog. It is raised with less certainty on the lake shores,
and on the upper stretches of the Penobscot the corn crop is
precarious.

In South America, east of the Andes, the cultivation was
limited to districts comparatively free of forest, and where
the ground was sufficiently elevated for the best growth of
the plant.

The map of the western hemisphere (Plate XVII) accom-
panying this chapter displays the original home of maize and
its distribution in space and time.' It is probable that maize
reached the Rio Grande about 700 A.D., for Humboldt states
that the Aztecs learned of maize in 6(^ A.D. By the year
1000 A.D. it had reached the coast of Maine. The Incas used

• The squared areas on the map (Plate XVII) show the position of the agricultural
tribes in North and South America. It is evident that the position of the agricultural
triljes and the area of maize distribution are identical. See explanation accompanying
map. See also chapter on Ethnology for position of said tribes and the grade of their
culture.

II

I

156

Harshberger. — Maize :

it before 700 A.D. Agriculture was practiced on both con-
tinents and on the islands in the Gulf of Mexico by the year
1492, for when the Europeans arrived it was found every-
where. The Europeans carried it to Europe, from whence it

sDread.

Maize was introduced first into Spain. Gerard, in his Herbal,
states that ''these kinds of grain were first brought to Spain,
and thence into the other provinces of Europe, not (as some
suppose) out of Asia Minor, which is the Turk's dominion,
but out Of America and the islands adjoining, as out of Florida
and Virginia, or Norumbega." M. E. Discourlitz says posi-
tively that maize was brought to Europe by the Spaniards
from Peru/ and Matthioli, in 1645, affirms that Turkish wheat
is not a proper name for maize, but that it should be called
Indian wheat, because it came from the West Indies.

The names in Spain, Belgium, France, Germany, Greece,
Italy, Russia, Sicily and Sweden indicate that it was received
•from Turkey, but this confusion was due to associating the
newly-discovered islands and continents with the East Indies,
the trade with which was carried on by way of Turkey and
the eastern Mediterranean. These mistaken geographical
notions were hot rectified until 1522. when the globe was first
circumnavigated, and the lands to the west were proved to be
wholly distinct from the Asiatic continent. But the name
given to corn naturally lingered and became part of common
language, and as time passed it was impossible to correct the
mistaken impression. France appears to have derived the
plant from Spanish and West Indian sources. Grains of it
were sown in the sixteenth century in Spanish, Italian, French,
German and English gardens, and soon the plant was culti-
vated on a larger scale in the fields. Under the name kukuruz,
it was naturalized in Turkey, the Danubian countries and

Hungary. .

It came to Germany from Italy, as Turkish wheat, or walsch-
korn. " Our Germany," says Hieronymus Bock (Tragus), in
his Neu Krciitcrbuch, Strasburg, I539, "will soon be called

1 Peruvian word zara was used in Spain (see pages 89 and 126), which shows that the
word mahiz, maize, was not yet generally adopted.

A Botanical and Economic Study

157

felix Arabia, because we accustom so many foreign plants to
our soil, from day to day, among which the large walsch-korn
is not the least important." Italy probably obtained seed
from Sicily and Spain, and Sicily from Spain and the Amer-
icas. The confusion of names is great. Maize is called in
Lorraine and in the Vo.sges, Roman corn ; in Tuscany, Sicilian
corn ; in Sicily, Indian corn ; in the Pyrenees, Spanish corn ;
in Provence, Barbary or Guinea corn. The Turks call it
Egyptian corn, and the Egyptians, Syrian dhourrah. But the
widest spread name was Turkish wheat, which came from a
misconceived notion as to its origin. Rucllius uses it first in
1536^ All that can be said with certainty, however, is that
Indian corn reached northern and central pAirope from the
countries bordering on the Mediterranean. It was introduced
into Africa by the Portuguese in the sixteenth century, and
is cultivated more or less from the Middle Sea and the Libyan
Desert to the Cape of Good Hope.' It is particularly deserv-
ing of attention that the greater number of the plants culti-
vated on the Congo, and among them nearly all of the most
important species, have been introduced from other parts of
the world — maize, manioc, or cassava, and pine-apple.'^

Maize early reached India and Burmah. Baden Powell
observes in his ** Punjaub Products," that maize grows every-
where throughout the hills, and appears to flourish as well in
a temperate as in a tropical climate at 7000 feet or more. It
is the favorite crop of the people, and for six months of the
year forms their common food. It is supplanted in the val-
leys by rice, but even here there is always a little plot of
maize about the cottages of the peasant classes.

The Chinese used it early after the discovery of America,
for the Portuguese reached Java in 1496, and China in 15 16.*
Mayers' believes that it was introduced into China from
Lower Mongolia in the sixteenth century. The introduction
through Mongolia is highly improbable, but the date of a

' KuelHus. De Natura Stirpium, 428.

2 Simmonds, Tropical Agriculture. 1877, 295.

■"' Brown, Robert, Miscellaneous Botanical Works of, Kay Soc. i, 155.

^ Malte Brun, Geographie, i, 493.

•' Mayers, Journ. Bot., Seeman's, 1871, 62.

/

158

HarsJibergcr. — Maize :

treatise on Natural History, published in 1597 (Pen-tsao-
kung-mu), furnishes us abundant proof that the Chinese did
not know it before the discovery.

1557, introduced into the Cape Verde Islands.

1593, Hawkins finds it in the Canaries.

1595, maize introduced into the Marquesas Islands.

1775, Thunberg enumerates maize among the edible plants

of Japan.

The present cultivation of maize extends throughout both
American continents. In Europe, M. de Gasparin ' assigns as
the region of maize the plains which border on the Pyrenees,
the valleys which descend from the Jura, all Italy, Corinthia,
Austria and Hungary. It is cultivated in Asia Minor, India,
China, the Phillippines, the Malay Archipelago and Australia,
and furnishes in all these places a most important food for
man and beast. The surprising and rapid extension, in such
a short period of time, has depended on the merits of the
plant as an important article of food.

1 Gasparin, Lecoiiteux, " Le Mais" Paris. 1853, 260

A Botanical and Ecojioniic Study.

159

CHAPTER IV.
Chemical.

IT is important to study the chemistry of a plant for the
following reasons: (i) It is necessary to know the value
of the plant as a human food. This has been determined
by practice, but scientific analyses are essential if we desire
to compare a new food with older and more established
ones. The Germans have made painstaking analyses to dis-
cover the cheapest and most economical foods for use in the
army, and the commissary department, after much experimen-
tation, has turned its attention to maize. (2) Different
plants take from the soil a varying proportion of plant food.
It is necessary to estimate this loss in order to supply
deficiencies. (3) Products once wasted are now saved. This
is essentially a utilization of bye-products. Chemical inves-
tigation is useful in ascertaining the value of refuse.

The chemical analyses which follow have been selected
and arranged with great care from the most reliable sources.

Table I.
Relations in Weight between Different Portions of Maize in

the Green State}

Leaves

Tassel

Ear, stem

I
Upper stalk '

Middle "

I

I
Lower " ;

29.20

.66

18.01

7-56
14.86
30.01

47-87

I- 52-'3

1 00.00

100.00

1

1 ("loffart, Culture and Ensilage of Maize. 41.

i6o

Harshbcrger. — Maize :

The analyses which follow display in a summarized form
the constitution of the plant at different seasons of growth.
Boussingault' gives an analysis of seedlings of maize, after a
period of twenty days' germination. The table has been
adapted for the chapter.

As the embryo grows the reserve materials in the seed
diminish in quantity. They are conveyed to the seedling,
and are used by it to form new protoplasm and new cell walls.
" The effect of the absorption of these substances by the
embryo is that the cell-sap of the cells of its ground-tissue
become charged with them, for the absorption is much more
rapid than the consumption in the formation of proteid ; con-
sequently the seedling soon comes to contain a larger per-
centage of them than does the organ in which they are being
formed. If the seedling is growing under favorable condi-
tions these substances gradually disappear, and this is accom-
panied by an increase in the amount of proteid contained in
the seedling." The following table shows this exactly :

Table II.

Analysis of Seeds and Seedlings After Tiventy Days Ger-
mination. Absolute Weight in Grammes .

Total

Dry
Weight.

Gr.

Carhohy-

drates,

Starch,

Sugar.

Gr.

Oil

Gr.

Fihre, N. Sub- * Extrac-

Cellulose, stance., " i tives.

Gr.

Gr.

Gr.

Gr

Seeds. . . S.636 6.386 0.463 0.516 | 0.880 0.156 0.235
Seedlings . 4.529 1.730 0.150

1 .316 0.880 0.156 0.297

1 Vines' Physiology of Plants, 176.

A Botanical and Economic Study.

161

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« CO

ci "O

u

00 CO Lo CO « «

-TNO

<

ci CI CI CO -TO

LO 10

^»

0 ^

w

"V»

^

•!

0
tn

bd

5

H

Z

^

g

Oz

oi

• • • • •

^

ai.0

1

1 . .

H Z

«<

X<

1

'J

' *- 0 ■■ 0 fi On

vO CO 0 NO "^. f^

0 -J-

H 1 w n n ri

— CI CO —CI

►-" CI

^

•

/;

^1

lli

l62

HarsJibeixer. — Maize :

The foregoing table gives analyses at different periods of
growth.

This table shows the general tendency of the plant to
increase rapidly in all its parts up to August 6. After this the
stores are emptied into ear and grain, and additional increase
comes from the activity of the husks rather than from the ^
leaves. These, in so far as they are situated below the
eighth or ninth node, the usual seat of the ear, become
productively inactive and dry up, whilst those above them
yield their substance to the ear, and after drying represent a
less valuable material than the lower ones. Both become
very brittle when dry and are usually lost in the process of
harvesting.

The following table, prepared for the United States De-
partment of Agriculture, collected from the reports of forty-
nine experiment stations and many other sources, including
the publications of schools, colleges and agricultural societies
in the United States and Canada, is a compilation of all
analyses published before September i, 1890. A variety of
methods were used in the determinations by the earlier
chemists, so that a slight discrepancy exists between the
results of the older and later published analyses.

Taijle IV. (a)
Analysis of Maize Fodder of Different l^arieties^ Cut

Green.

/

^eree7itage

■

.

In Fresh

OR Al

R-

DRIED Material.

_

Water.

Ash.

Protein
(N. X 6.251.

Crude
Fibre.

N. P'rke
Extract.

Fat

'^2

<

i

Flint . . .

40

79.8

1 .1

2.0

4-3

12. 1

1
0.7

Dent . . .

53

79.0

1.2

1-7

5.6

12.0

05

Sweet . .

21

79.1

'•3

1.9

4.4

12.8

0-5

All variet.

129

79-3

1.2

1.8

S-o

12.2

0-5

' Experiment Station Record, ii, ;o2.

A Botanical and Economic Study.

Table IV. {b)

Analysis of Maize Fodder of Different Varieties, Cut

Green. Percentage.

163

Calculated to Water-free Substance.

Flint . . .
Dent . . .
vSweet . . .
All varieties

Number

of
Analyses.

Ash.

1

P

1

rot EI N.

Fibre.

' N. Free
Extract.

Fat.

40

5-2

9-7

21.3

60.6

3-2

63

5-7

^■Z

26 3

57-J

2.6

21 f

6.0

89

21 2

61.7

2 2

129

56

<S.8

1

24.1

58.9

2.6

An analysis of the roots is given. The roots, especially in
their later stages of growth, are covered by a brown and
exceedingly tough incrustation of sand and clay, which can be
removed only by much washing and rubbing, and thus retain
much earthy matter, as the analytical data show.

Table V.

Proximate Composition of Roots at the Close of the Groiving

Period.^

Crude ash ' 2 93

Ether extract 048

Crude fibre 41 94

Crude protein 6 43

Carbo hydrates 48 22

Nitrogen 103

> Schweit/er, P., Missouri .^gric Exper. Stat Bui. 9 1889 5

164

Harsh berger. — Ma ice :

Tarle VI.

Percentage Composition of the Ash of the Root of One

Plant:

A Botanical and Economic Study.

16S

Silica

Ferric oxide ....
Phosphoric-pentoxide

Lime

Magnesia

Potassa

Soda

Total . ■

Missing

June 21.

Sept. 10.

16.73

32.10

8.21

0.22

4-38

14.41

10.12

5.57

5.37

4.02

37-95

37-69

2.58

4-38

85-32

i 98-39

14.68

1.61

Table VII.
Composition of the Different Parts of the Maize J^'lant.'

«
Stalk

j

Tassel.

Ears.

Leaves.

1

_„ _ —

Entire
i Plant.

6.28

Upper
434

Mid.
3.86

Lower.

6.27

1 1.09

1

6.47

1.90

2.50

1.30

1. 00

.40

•30

1.28

, 4-70

8.30

6.50

17. CO

20.60

21.00

11.77

1
25.23

73-51

64-33

39 49

38.65

35-79

56-35

56.70

2.90

10.60

33- 10

I

33-^0

38.00

18.37

i

5.20

1.70

10.99

4-57

2.69

1.74

5-74

Nitrogen .
Fats, Sol. in
Saccharine
Starchy
Cellulose .
Mineral

* Schweitzer.

"Goffart, Composition and Ensilage of Maize.

Table VIII.
Centesimal Composition of the Ash of the Different Parts.

Entire
Plant.

71.70
3.81

'•35
4.41

8.26

Phosporic Acid .
Sulphuric "
Chlorine ....

Potash

Soda

Lime ". : 12.96

Magnesia . . . 6.60

Iron

Silex

CO and Wa.ste

Ears.

33 50

358

3-52

27. 1 1

21 36

3 46

7.04
0.51 traces.

54-75 i 0-34

Leaves. Tassels.

Stalk.

Upper Mid. Lower

0.18

0.09

3-97
3.21

1.04
123
6.78

1378

5 64

0.46

63.76

0.13

10.01 9.07 14 02 7. 17

6.13 : 5.61 8.65 381

I

2.73 i 2.15 traces. 1.35

7.88 14 61 2.41 4 41

10.37 ' 12.57 8.39 8.26

11.87 , 1029 14 31 12.96

15.03 10.52 8.73 660

o.ii 2 08 0.63 0.51

35.83 I 29.83 4i.:,7 54-75

0.03 3.27 1.49 o 18

100.00 100.00 100.00 ico.oo 100.00 100.00 100.00

i ' ' i I J

In a subsequent chapter the manufacture of maize paper
will be discussed. It is, therefore, necessary that the chemi-
cal composition of the maize husks, out of which the paper
pulp is principally made, should be ascertained. (Schweitzer.)

Table IX.
Proximate Analysis of Maize Husks. September 10.

Crude Ash . .
Ether Extract .
Crude Fibre

" Protein .
Carbohydrates
Nitrogen . . .

6

23

0.

83

1>7>

77

4

Zl

54

80

0

70

1 66 HarsJiberger. — Maize :

Table X.

Percentage Composition of Husks. September lo.

Silica

Ferric oxide ....
I'hosplioric pentoxide

Lime

Magnesia

Potassa

Soda

Total

Missing

36.22

037

2.03

5.58

6.62

37.66

2.68

91.16

8.84

The table below gives the average of 500 analyses of corn
kernel of dent, flint, sweet and pop varieties, raised in Con-
necticut, Kansas, Michigan, Missouri, Texas, Wisconsin,
Massachusets, New Hampshire and Pennsylvania.

Tai

Analysis 0^

3LE XI.

f Corn Kernel}

In Fresh or Air-
dried Material.

ri.o

1-5
10.8

2.1

69.5

5-5

Water-free
Substance.

Water

Ash

1.8

Protein x 6.25 . .
Crude fibre . . .

12.1
2 4

N. free extract .

77-4
6.2

Fat

' Experiment Station Record, n, 706.

A Botanical and Economic Study.

167

The nutrient ratio for the first half of the table is i : 7.5 ;
for the second, i : 7.5.

Chittenden and Osborn have made an extended investiga-
tion of the proteids of the kernel of maize. The variety
used was a white dent corn.' The grain contains several
distinct proteids.

(i) Three globulins separable from the kernel by extrac-
tion with 10 per cent, salt solution ; (a) myosin coagulable
in 10 per cent, salt solution at 70° C, and directly dissolvable
in water from the meal ; (J?) vitellin, non-coagulable in dilute
salt solution, insoluble in water, 10 per cent, salt solution
dissolving it after water extraction ; (c) globulin separates
from the above rcsidient only after prolonged dialysis. It is
coagulable in 10 per cent, salt solution at 62^ C.

Table XII.
Composition of Maize Globulins.

Carbon
Hydrogen
Nitrogen .
Sulphur .
Oxygen

Myosin.

V

itellin.

G

1
1

LOHULIN

52.66

517'

52.38

7.02

6.85

(>.?>Z

16.76

18.12

15-25

1.30

0.86

1.26

22.26

t

22.46

24.29

(2) Two albumins unlike in chemical composition and of
uncertain structure.

(3) The chief proteid in the maize kernel is the peculiar
body maize fibrin (zein), soluble in warm dilute alcohol, insolu-
ble in absolute alcohol or water. Zein acts characteristically,
is yellow resembling beeswax, is soft, tenacious, elastic and
heavier than water. When heated it swells, turns brown,
and leaves a bulky charcoal. It burns, but not rapidly.' The
insoluble state is formed when zein is heated with water or a
weak alkali.

1 Amer. Cheni. Journ., xiii, 453. 529 ■ XJ^, 14 ; Exp. Stat. Rec, m, 768.

2 Gorham, Jno., Philos. Magaz., i.vii, 311.

i68

HarsJibergcr. — Maize .

Table XIII.

CoDiposition of Zeiii.

Carbon .
Hydrogen
Nitrogen
Sulphur .
Oxygen .

j Soluble Z

EIN.

Insoluble Zein.

55.28

55-15

7.27

7.24

16.09

16.22

0-59

0.62

20.77

20.77

The ferments in maize are: (i) diastase; (2) invertase ;
(3) ^ glyceride enzyme ; (4) proteo-hydrolyst.'^

Taulk XIV.
Composition of Ash of Kernels.^

Carbon dioxide . .
Silicic acid ....
Sulphuric acid . .
Phosphoric acid )
Peroxide iron )
Lime ....
Magnesia . .
Potassa . . .
Soda ....
Chlorine . . .
Organic acids

^ ,„ ,„ Rhode Island

Chinese Tree. I uscarora, ii. Sweet. 15.

race.

1
trace.

trace.

1.700

0 77S

.125

1.075

1-^75

0550

49- '85

0.620
16 200
12.930

15-365
0.440

2.125

44 135

44 050

0395

0335

I2.S75

12.810

14 240

12.867

20545

22 968

0.450

0.270

3520

3025

^ .Salisbury, J- H., Journ. New York Agric. Soc, 1849.
-Green, Annals of Botany, March, 1893. 91-116.

if

A Botanical and Economic Study.

169

The cob yields a rich and abundant supply of potash. One
hundred parts of dried cobs yield after drying at 212° F. the
following :

Table XV.
Potash in Corn Cobs}

—

1
1

Ashes.

i
K CI.

K

,C03.

Silica, Iron,
Lime.

Loss.

I

1. 120

.820

.750

.140

.230

2

1.040

.805

•745

1 .180

•115

3

1.015

.840

.755

•245

.605

4

1. 115

.830

•795

.300

.020

BIBLIOGRAPHY.

Parmcntier, Le Mais, Paris, 181 2, 8vo, i vol.

^;//^wr^?^A-, Memoire sur le Mais, 1784.

Bonafous, Histoire Naturelle, Agricole et Kconomique du Mais, folio,

Paris, 1836.

Duchgsfie, E. ^., Traits du Mais,ou B16 du Turquie contenat .son His-
toire, sa Culture et ses emplois en Hconomie
domestique et en Medicine, 8vo, Paris, 1835.

Cobbett, Wm., A Treatise on Indian Corn, i2mo, London, 1828.

Neuf chateau. Supplement au Memoire de M. Parmentier, Paris, 181 7.

Sturtevant, E. Z., Indian Corn, Transactions New York Agricultural

Society, 1877-82, 37.

Brewer, H. //., Paper on Maize, Yale College.

Riant, C, La Charte du Mais, Paris, 1877.

Enfield, Edward, Treatise on Indian Corn, i2mo, New York, 1866.

Washington {D. C), Special Report on Cereal Production, 1882.

Steffen, Max, Die Landwirthschaft beiden Alt. Amerikanischen Cultur-

volkern, Leipzig, 1883.

Peckolt, Th., Monographia do Milho (Zea mays) et do Mandiocca,

sua Historia Variedades, Cultura, Uso; Rio
Janeiro, 1878.

Rein,;., Zur Cieschichte der Verbreitung des TabaVcs und Mais in Ost

Asia, Petermann's Mittheil, 1878, 215.
Eine Pflanzengeographische und Culturgeschichte Skizze,
Reis, und Mais, Jahresb. Verf. Geog., 1878, 67.

I Hazard, U., Amer. Journ. Pharni., 4 ser., 11, 152.

I/O

Hars/ibcn^cr. — Ala ize

Lecouteux, Ed.. Le Mais, Paris. 1883.

Salisbury, /. //.. History and Chemical Investigation of Maize,

Albany, 1849.
Bro7Ufu\ D.J., Memoir on Indian Corn, New York, 1856.
Browne^ P. A., Essay on Indian Corn, Philadelphia 8vo, 1837.
Helier, II. Carl, U. S. Patent Office Report, 1847.
Salisbury, J. //., Transactions New York Agricultural Society, 1848.
Flint, C. L., Transactions New York Agricultural Society, 1849.
Betnent, C. A'., Transactions New York Agricultural Society, 1853.
Brendel, Transactions Illinois Agricultural Society, 1856-7.
Klippart, Agriculture of Ohio, 1858.
Arena, Symposium on Maize as the National Flower, June. 1893.

A Botanical and Economic Study.

171

CHAPTER Vs
Agriculture — Physiological.

THE cultivation of Indian corn has been greatly improved
within recent years. New machines have been devised
for drilling the kernel into the ground, and new culti-
vators of improved pattern have been introduced. It is not
essential, in describing the maize plant, to decide such ques-
tions as the merits of deep or shallow cultivation, the value
of removing the tassels as a means of increasing the yield,
the value of root-pruning, etc. These are questions of agri-
cultural practice and extra limital.

This chapter will deal with practical physiological consid-
erations, such as the effect of fertilizers and the measures
necessary to restore soil fertility. The chemical changes
which the soil undergoes during plant growth, and the proper
rotation to be used in intensive agriculture will be discussed.
The object will be to investigate rather the principles under-
lying fundamental practices, and it is hoped that the results
will be of general and wide application.

From experiments at Rothamsted, on a large number of
cereal and non-cereal plants, Messrs. Lawes and Gilbert have
deduced many important principles. The German stations
and the experiment stations of the United States also fur-
nish material at hand for a general agricultural survey. A
few general rules are given.

It is certain that the increased growth of our staple starch-^
yielding grains is greatly dependent on a supply of nitrogen
in the soil.' The better the cereals are matured, the lower
is their percentage of nitrogen, the explanation being that
maturation means the greater formation of non-nitrogenous,

1 Gilbert, J. H., Agricultural Investigation, Rutgers College, N. J., 1S84.
12

\

172

llarsJiberg er. — Maize

substance— starch.' Tt is equally certain that the increased
production of sugar, as in sugar cane, is also greatly depend-
ent on the supply of nitrogen.- With root crops, the amount
of sugar is increased by the use of nitrogenous manures.
Nitrogen applied to leguminous crops has comparatively little
effect in increasing the product of such crops.'

Potash is essential for the formation of the chief non-
nitrogenous matters, starch and sugar.

The results with leguminous crops show that mineral ma-
nures (particularly potash) considerably increased the early
crops. Ammonia salts were of little or no benefit, and were
sometimes injurious. It may be added that the beneficial
effects of long previous applications of potash were apparent
whenever there was any growth at all. When the land is
'* clover sick," none of the ordinary manures, whether artificial
or natural, can be relied upon to secure a crop.

In experiments on the effect of various manures, applied
on fields at Rothamsted for forty years, it was found that the
plats which received ammonia salts alone, gave the smallest
yield in bushels, twenty and one-quarter of wheat, and
twenty-nine of barley. The sulphates with 600 pounds of
ammonia, gave a yield of thirty-six anckthree-quarter bushels.
It is seen that a mixture of mineral and nitrogenous manures
worked to the best advantage.

Will principles laid down for England and English cereals
hold for American cereal productions, especially maize } The
results attained in the United States are summarized as
follows:

Nitrogenous fertilizers materially increase the crop,' and
the yield was poor when nitrogen was not applied. ' Numer-
ous experiments indicate that corn thrives well and yields
fair returns when the fertilizer contains one-third or one-half
the nitrogen removed by the crop.** The crops on the plats

' Gilbert J. H., On Growth of Potatoes, Rothamsted, 20.

2 Gilbert, J II , Lecture at Rutgers College, N. J., n.

■■ Lawes. Sources of N. of Leguminous Crops, 1892, 4.

* Georgia. lUtl. 10. 1890, 20; Exp. Stat. Rec, 11, 550.

■' Massachusetts Stat.. 7th Annual Rep., 18S9, 148; Exp. Stat. Rec, ii, 579.

" (-'onnecticut. Storr's Stat., 2 Rep., 1889.

A Botanical and Economic Study.

173

receiving no nitrogenous manures were of a light green
color, and during the first half of the season the same con-
dition of the plant was noticeable on the plats receiving
ammonium sulphate.' The light green color of a crop will
indicate, generally, that the soil is deficient in available nitro-
gen. It is evident that chlorophyll formation has a close
connection with the proper assimilation of nitrogen, but that
the increased carbon assimilation does not take place unless
with this additional chlorophyll sufficient mineral matter is
present. The corn plant does not respond to heavy applica-
tions of nitrogenous manures. It is therefore easy to apply
too much."

The use of large quantities of nitrogen above the quantity
utilized by the plant is a direct waste of money.^ The use of
superphosphates alone on corn is unsatisfactory.* The use
of potash alone is unadvisable,' and of doubtful value. "The
yield of ear corn on the two unmanured plats were respectively
thirty-four and forty bushels per acre ; on the three plats re.
ceiving fertilizers containing no potash, from thirty-three to
thirty-six bushels, and on those receiving potash fertilizers,
from sixty-five to seventy-six bushels ; the largest yield was
with the combination of potash and nitrogen." " When pot-
ash was used, there was a marked increase of crop (twenty,
eight to thirty-nine bushels per acre), and the greatest in-
crease was with a manure composed of potash and nitrogen
(thirty-nine bushels). There was a profit in every instance
where potash was used, and a loss (financially) where that
element was left out." Mixed superphosphate, potash and
nitrate of soda yielded the best results. ** The yield in all
cases was larger when potash and superphosphate were ap-
plied with nitrogenous fertilizers.'"* Nitrogenous manures,

' Massachusetts Stat., 7 Rep., 1889, 148 ; Exp. Stat. Rec, 11, 579.

- South Carolina Stat., 2 Rep., 1889, 210-268 ; Exp. Stat. Rec, n, 550.

■ Connecticut, Storr's Stat., 2 Rep., 1889.

* Georgia, Stat. Bui., 10, 1890, 20.

'" South Carolina Stat., 2 Rep., 18S9, 210-268.

*^ Kentucky Stat. Ihil., 33, 1891.

" Exp. Stat Rec. 11, 725.

** Georgia Stat. I5ul., 10, 1890, 20.

^'Georgia Stat. Bui , 15, 1891: Exp. Stat. Rec, 111,604,

\l

%

174

HarsJibergcr. — Maize :

with" potash alone, or phosphoric acid alone, or all three com-
bined, increased the yield materially/

The results prove that nitrogen combined with some min-
eral salt (potash preferably) materially benefits maize, and
this essentially agrees with the physiological habit of the
other cereals, wheat, oats, barley and rye. Notwithstanding
this, William Frear and H. P. Armsby raised the question '
as to the truth of the statement that maize, as a cereal,
responded in increased starch to the increase of the nitro-
genous manures applied. '* Upon comparison of the results
ft was found that the nitrogen free extract in the grain pro-
duced upon plats receiving complete fertilizers, was the same
in percentage no matter what form of nitrogen had been
applied, and that in all cases where this ingredient was
applied there was less nitrogen free extract than was found
in the crops from the unfertilized plats." This argument is
weakened by the fact that maize is a gross feeder, and must
have its food in a shape to be readily absorbed by its deeply
penetrating roots, and it seems that some time must elapse
for the action of recently applied manures especially nitro-
genous, which become slowly available, to take place. The
Kentucky station records the fact " that the fertilizers applied
in 1888 were of benefit to the crop of 1890," which clearly
shows that it takes time for the nitrifying process to take

place.

The nitrogen in the soil is prepared for the plant by the
process of nitrification. The older theories explained this
oxydizing process as a merely passive chemical reaction,
taking place when suitable compounds were in contact. The
newer theory, however, which is gaining ground, ascribes
this process of nitrite and nitrate formation to living organ-
isms—minute bacteria. The historical development is inter-
esting. Pasteur stated long ago that probably the nitric acid
was produced in soil by a living organism similar to those
which cause fermentation and putrefaction. Schlosing and

» New York Stat., 8 Rep , 1892, 56-260.

■i Nitrogen Supply of Maize, L>oc. Promotion Agric Sci. Proc, vii, 33, Rep. Penna.

State College, 1889, Part 11, 199

A Botanical and Economic Study.

175

Miintz established by experiment the true nature of nitrifica-
tion.' Since these earlier trials additional proof has accumu-
lated, showing that nitrification in water and soils is due to
bacterial agents. Winogradsky ^ isolated by culture the
nitrifying organism. Frank, Wilfarth, Hellriegel and other
investigators have made many experiments which serve to
show that the nitrification of ammonia in the soil, and prob-
ably the nitrification of other nitrogenous matters, takes place
in two stages, and is performed by two distinct organisms.
One converts ammonia into nitrite, the other changes the
nitrite into nitrate. In the soil both organisms are present in
large numbers. The action of the two proceeds together.
The conditions favorable to their growth are alkalinity of soil,
oxygen, and a proper base with which the acid can combine-
Calcium carbonate usually plays this role in the ground. A
soil deficient in such a base is generally infertile and needs
dressings of chalk or lime.

These discoveries, combined with the equally important
one that the utilization of free nitrogen by leguminous
plants is due to bacteria which form nodules on the roots,
will eventually revolutionize modern agricultural methods.
Mr. Mason, of Eynsham, Oxfordshire, England, commenced
some experiments in 1889, with a view of applying to
practical agriculture the knowledge accumulatedconcerning
the minute organisms. -His method is to grow nitro-
gen accumulating crops for home consumption, and after-
wards nitrogen consuming crops for sale." He grows mixed
crops of LeguminoscB, liberally fertilized with basic slag and
kainit. He converts the first year's crop into silage, which
he feeds to his cattle, returning the manure to the soil, and
converts the second year's produce into hay. The land thus
produces highly nitrogenous crops without manure, and is
left in a high condition for potatoes or grain, which need
nitrogenous manuring. The fact that it is necessary, in order
to prevent "clover sickness," to grow in sucession a variety
of Legnminosce, will lead to radical and important changes in
our present system of rotation.

' Compt. Rend., Lxxxiv, 301.

•- Winogradsky. Ann. de I'lnst. Pasteur, iv (1S90-911. 213, etc.

>l

176

HarsJibiigc)'. — Maize :

A Botanical and Economic Study.

177

The question is how best to conserve the nitrogen, the
supply of which in sufficient quantity is so invaluable. The
nitrogen is lost to the soil in three principal ways : (i) By
drainage ; (2) by removal of crops ; (3) through the air. This
waste goes on in all soils, in some more than in others. Sir
John Lawes, in a paper on fertility, concludes that 3000
pounds of nitrogen have been lost in the Broadbalk wheat
field at Rothamsted in the last 250 years. It is generally
conceded that this waste can be prevented directly, to some
degree, by growing crops through the season of the greatest
production of the nitrates. Rotations properly arranged
conserve the soil nitrogen. Thus, if red clover is sown in a
growing crop of barley the land is covered immediately after
the barley is cut, and the effect is as nearly perfect as pos-
sible, the growth of the second crop following the first without
a break in the continuity. At Rothamsted it was found that
on sampling the drainage waters and calculating the amount
of nitrogen as nitrates contained therein, that the maximum
discharge takes place from October to February, and the maxi-
mum nitrification takes place during the autumnal months.

The cereals, after a wet winter, begin to grow in the spring
in a soil drained of nitrates, and the growth is over before
the greatest production of nitrates takes place. These crops
are, therefore, greatly benefited by nitrogenous manures. As
the growth of maize takes place in the late summer months,
it is more independent of nitrogenous fertilizers^ than wheat,
barley, etc.

The nitrogen, therefore, is best conserved with a crop
whose period of growth extends into the period of maximum
nitrification and maximum drainage of nitrates from the soil.
Maize is just such a crop. Cereal crops whose growing
periods are confined to the spring and early summer are very
])oor conservers of nitrogen. This accounts for the fact that
maize does not exhaust the soil nearly so rapidly as wheat,
which has its period of growth much earlier in the year.

' i^ee Pa«:e 174.

CHAPTER VI.
Utility.

T\\\\ maize plant subserves many important uses. The
study of our plants from a utilitarian point of view is
recent.

Professor Goodale, in his presidential address at Wash-
ington, in 1 891, before the American Association for the
Advancement of Science, said': ''Improvement of the good
plants we now utilize, and the discovery of new ones, must
remain the care of a large number of diligent students and
assiduous workers." ** One phase of it is being attentively
and systematically regarded in the great experiment stations ;
another phase is being studied in the laboratories of chem-
istry and pharmacy, while still another presents itself in the
museums of economic botany."

As population increases, and the centres of mart and trade
become more closely aggregated together, economic use of
our food supplies 'must be inaugurated, and foods of small
cost and great nutritive power must be substituted for those
of great cost and very little food value.

Human Food. — Does maize possess the quality of a
perfect food 1 This question can be answered only by a
reference to analytical tables, and a comparison of the
nutritive ratios of maize with other foods which enter into
our dietaries. Iwidently a proper food must minister to all
the requirements of the human frame. A good food must
supply the waste which takes place and maintain the system
in a condition of vigorous health.

' Goodale, Pop. Sci. Month., Novemlier and December, iSgr.

178

HiirsJiberger. — Maize :

A Botajiical and liconomic Study.

179

Table XVI.

Nutrient Ratios and Nutritive Values of a Seiies of Foods,

for Comparison witJi Maize.

Ratio. Value.

Ratio. Value.

Wlicat I : 6.5

Indian wheat ... : 5.2

Macaroni .... : 5.6

Oats :5.7

liarley ^ 12.25

Rye : 7

Rice : 10

Maize : 7.5

•■\Iillet 15.57

Buckwheat .... : 4.75

Peas ....... I : 2.50

Haricots 12.5

Lentils : 2.4

Peanuts : 5.2

vSoy : 2

Potatoes : 17

Turnips : 6

Carrots : 14

Parsnips : 12

Beet root : 29

Jerusalem artichokes : 8

Onions : 3.5

Sweet potatoes . . 113

82.0
84.6
89.6
102
8s

^S

84

87

84.7
86

79
So

86
'51

lOI

4

7-5
16

12

16

Yams * 7-1

Cabbages ... 14

Vegetable marrow : 5

Tomatoes ... : 5

Iceland moss . . : 8

Iri.sh moss ... : 5.5

Celery 1:4-5

I

Apples : 27

Gooseberries . . : 20

Grapes ... : 20

Figs : [o

Bananas .... 14

Carol) beans . . : 8.5

Walnuts . . . . , :6.q

!

Coconuts . . . . : 16

Milk 1:4

Cheese : 2.4

liggs : 1.9

Calves' liver : .3

Beef :.8

\^eal :.i

Pork : .7

Mutton i : I.I

7-5
3-5
8.5

79
64

5

9
16

68

24

68

94
90

99

40

The nutrient ratio is a comparison of the albuminoids, or
flesh-formers, and the carbo-hydrates, or fat-formers. It is
obtained by adding the quantity of starch, sugar, and the
starch equivalent of fat together, and comparing the sum with
the protein substances present. F'or a man at moderate work
this ratio should stand as i : 5. The foregoing table gives the
nutrient ratio and nutritive value of a large series of foods for
comparison with maize.

The sum of the albuminoids, starch, dextrin and sugar, and
the starch equivalent of fats, is called the nutritive 'value ;
this value is that of 100 parts (grains, ounces or pounds). It
is seen from this table that the various food materials depart
more or less from the standard ratio that we have adopted,
I : 5, as most nearly expressing the proportion which the nitro-
genous should bear to the starchy food. Maize departs con-
siderably from the standard, and is poor in protein substances
at the best. This fact rather unfits it for a standard article
of food, unless combined with some other products which are
richer in albuminoids. Thus peas, with the nutrient ratio of
I :2.5, can be combined with maize, and the deficiencies of
both be equalized.

Animal Food.— The food requirements of an animal are very
similar to those of man. A proper proportion of flesh producers
must be combined with fat producers, in order to nourish the
animal satisfactorily. The digestive systems of the different
d(3mcsticated animals, however, differ from that of man, so
the material fed must be of a different kind ; for although
bread principles can be deduced from experimental work on
man, these principles must be modified and corrected by
experiments carried on with farm-yard animals. Again, the
kind of feeding must be conditioned by the purposes for
which the animal is grown. Mutton for table use must be
lean, and the shepherd must have a knowledge of the principles
of nutrition, in order to prevent fat accumulation. On the
other hand, if the sheep are to be grown for tallow, then
fattening foods must be used. When these principles become
more thoroughly recognized and observed, it will be possible

%

i8o

HarsJiboxcr.-

Maif^c :

to obtain more condensed protein matter for human food
without the necessary loss in fat.

MilcJi Coivs. — To determine the value of maize for all-round
feeding reference must be made to a few experiments. The fol-
lowing is significant :' "The nutritive value of our dry corn
stover and of a good corn ensilage, taking into consideration
pound for pound of the dry vegetable matter they contain, has
proved in our case fully equal, if not superior, to that of the
average I^nglish hay." "The total cost of the feed for the
prodirction of milk is lowest whenever corn fodder or corn
ensilage have replaced in the whole or in part English hay."
The Wisconsin station made some experiments on the yield
of butter and milk by animals fed on fodder corn. For a
period of forty-two days, 1227 pounds of corn supplemented
with 672 pounds of bran and corn meal produced 1487.72
pounds of milk and 58.68 pounds of butter.

A discussion of ensiled corn vivsus fodder corn has been
engaged in. If it can be shown that the nutritive value of the
ensiled material is higher than the field-cured, then the loss
which occurs in the silo is immaterial.

Table XVII.
Results of Feeding Ensiled and Fodder Coni.'^

Food

Co

NSLME

■

C

Bran.

Corn
Meal.

Knsilaj^e . .

4960

504

168

Fodder corn

1227

504

168

Proolcts.

Watkr.

MlI.K.

BUTTMK.

2376 1688.14 62.3

5235 1487.12 58.11

This table indicates that the largest product both of milk
and butter occurred when the ensiled material was used. The
Maine station (Rep. 1889) compares the value of corn silage
and hay : " In these experiments the addition of ensiled corn

' Massachusetts Agr. Exp IStit. Rep., 1888.

* Wisconsin Agric Stat. Rep., iSSS ; Exp. Stat. Bui., U. i?. D. A.. 11, 193.

A Botanieal and Economic Study.

181

resulted in an increased production of milk solids over that
produced by hay." Again, with steers, the results showed
that a pound of digestible matter from the corn silage pro-
duced somewhat more growth (flesh) than a pound of timothy
hay. In comparing ensiled maize with hay, we find that the
gain in live weight is the greatest when the ensiled material
was used, as the following table shows : '

Table XVIII.
Slunving Gain per Ani^nal {Steers).

First three weeks
Second three weeks
Third three weeks .

With

SiLA(ii-:

66.3 pounds

57-7

a

38.0

u

With Hay.

43.3 pounds
46.3 "

2 1.0

u

Pigs.— It has been largely the practice in the Western States
to fatten pigs with Indian corn. The farmers have selected that
breed of pigs which will take the greatest live weight (chiefly
fat) to market. Experiments have been tried with the idea of
determining the best and quickest way of fattening hogs for
sale. The conclusions reached at the Kentucky station - were
that fat is more rapidly produced by feeding shelled corn, and
that corn affords the cheapest material for this purpose. The
breeders in the great corn belt have considered it profitable to
feed their corn instead of shipping the corn itself. Experiments
at the Illinois station indicate "that during the first five or
eight weeks of each trial, when one lot received only the half
feed of corn with pasturage, the gain per bushel of corn was
best in the case of the pigs on the full ration of corn, either
with or without pasturage."' A mixed diet is not only
cheaper, but more beneficial. At the Wisconsin station,

1 Exp. Stat. Rec, iii, 180; Virginia Stat. Bui , 10, June, 1891.

2 Kentucky Stat. Bui., 19, May, 1889.

•' niinois Stat. Bui., 16, May, 1891 ; Exp. Stat. Rec. in. 149-
* Wisconsin Stat., 7 Kep , 1890; Exp. Stat. Rec. n, 438

I82

Ha rs/i bcrger. — Ma ize

experiments were made with hogs feeding for fat and for lean.
The experiments showed the following points in favor of the
hogs that had been fed on corn, shorts and bran, over those
getting corn alone : (i) a more rapid growth ; (2) a much more
economical gain for food consumed ; (3) much more blood in
the body ; (4) larger livers ; (5) stronger bones in proportion to
the weight of the body. Notwithstanding, a large part of the
pork produced in the United States is grown on corn, and in
consequence is excessively fat. With nitrogenous food swine
have better developed organs and leaner flesh. Lean pork is
more valuable as a human food, and commands better prices.

Sheep. — Facts indicate that corn as an exclusive grain
ration does not give the best results when fed to growing
or fattening sheep. The production of wool is very greatly
dependent upon the nitrogen in the ration.

CJiickcns. — A consideration of the feed of chickens, which
consists largely of maize, a highly carbonaceous food, shows that
this diet works to the detriment of the fowls. ''The fowls
having the more nitrogenous food were always in better
health, and their plumage, except during a short moulting
period, was always full and glossy, while those having the
more carbonaceous ration were oftener sick, and their plumage
was always ragged and dull."' The gain in weight was the
largest for the nitrogen-fed chickens.'^

Live Wekjht.

Gain.

July 26. Nov 27. Pounds. Per Cent.

I. Xitrogenou.s food
II. Carbonaceous food

S.94
9.06

17.89
12.63

8.95
3-57

100. II
3940

"But the eggs laid (E. S. R., 11, 506) by nitrogenous-fed
hens were of small size, having a disagreeable flavor and
smell, watery albumen, an especially small, dark-colored yolk.

1 Exp. Stat. Rec, iii, 37.
- Exp. Stat. Rec, 11, 506.

A Botanical and Economic Study.

183

with a tender, vitelline membrane, which turned black after
being kept several weeks, while the eggs of the carbonaceous-
fed hens were large, of fine flavor, natural smell, large, normal
albumen, an especially large, rich yellow yolk, with strong
vitelline membrane, which was perfectly preserved after being
kept for weeks in the same brine with the other eggs." The
following table is interesting in this connection. In the case
both of the larger and smaller breeds the number and weight
of the eggs were larger with the corn-meal ration than with
the more^iitrogenous mixture, the difference being greater
with the smaller fowls. The fowls having the corn-meal
ration continued to lay for a longer period.

Table XIX.

Average Number of Eggs and Weight ivith Nitrogenous and

Carbonaceous Foods .^

Smaller Fowls, ^ ^^^^^ nitrogenous
Larger " )
Smaller Fowls, )
Larger "

Food Per
No. E(-GS. Wehjht. Day.

S

Corn- meal ration

43-7
48.9

68.7

50.1

91.48
108.24
136.29
112. 16

2.43
3-30
2-57
3-27

For chickens, feed nitrogenous materials; for eggs, carbonaceous.

Maize, with a nutrient ratio of 1:7 'S> does not fill entirely
the requirements of a well-rounded dietary. Unless com-
bined with other more highly-charged nitrogenous foods, it
works detrimentally. American agricultural production is
one-sided. We are a generation of fat and sugar-eaters.
Corn, our great staple, is poor in protein, at the best, and
is poorly adopted for a staple food, unless combined with
other materials. American dietaries range from i :6to i :8,
while the European standard is i :4.5 to i :55- The high con-
sumption of carbo-hydrate vegetal food is largely augmented

t New York Stat. Bui., 29 N. S., April, 1S91 ; Exp. Stat. Rec, m, 36.

184

HarsJibero^cr. — Maize :

by the use of fat meats. The ranchmen and swine-growers
convert a large part of the product of the soil into the fat of
beef and pork. The European farmer cannot afford to prac-
tice these wasteful methods, as the soil soon would become
exhausted.

" Not only must Americans develop American resources,
but Americans must adjust themselves to American condi-
tions. American people in the end must live upon those
articles of food for which American soil is most productive,
and must cease to consume in large quantities, as they do,
those articles for which our soil is poorly adapted. New
articles of diet will find their way into use, and habits and
customs will develop which will make the American of the
future a man utilizing all the resources of our country."'

Principal Maize Products.

I^

Whiskey.

10

2

Beer.

1 1

3-

Medicine.

12

4

(ilucose.

'3

5

Alcohol. ■ 1

M

6

Oil.

'5

7

Soap.

16

8

Paper.

»7

9

F^abrics.

18

{a) Fibres.

'9

{b) Yarn.

20

(c) Cloth.

1

Fuel.

Fire Lighters.

Packing Material.

Mattresses.

Pipes.

Baskets.

Thatch.

Corks.

I^otash.

Substitute for Coffee.

Green Manures.

Medicinal. — The stigmas of maize (corn silk) are used in
medicine as maydis stigmata. They are diuretic and lithon-
triptic. The corn smut of fungal origin is used as an em-
menagogue and parturient. An <:xcessive use of ergot is

' Patten, S. N., Economic Basis Protection, 103.

A Botanical and Economic Study.

•8s

attended by a shedding of the hair, or even of the teeth of
man. Mules fed on it lose their hoofs, and fowls lay eggs
without shells. Its action is as powerful, and even more so,
than the ergot of rye. An infusion of maize leaves has been
used as an anti-febrile, but its action is unreliable.'

Sugar. — Experiments were conducted in 1879 to determine
the yield of sugar from the maize plant. The stems contain,
if taken at the proper time, a great quantity of saccharine
juice. The sugar is crystallizable cane-sugar, and is not the
worthless corn-sugar expressed from the plant. Professor
Collier, in a special report, states that the sugar obtained was
in a satisfactory condition in every respect, as shown by its
high polarization, 90° {cf. sorghum 94°).

Table XX.
Comparison of Maize and Sorghum Sugar.

Stalk.

Corn Butt Ends .
Corn Tops . . .
Sorghum Butts .
Sorglnim Tops .

PkrCknt. Juice. Sp. Gr. Juice. ^^'V^oTuick^ '*^''

29.04
19.94
44.49
38.62

'053

1050

1060.5
1058

14.62
13.4G
16.44
14.48

Paper. — The moment must come sooner or later, when it
will be absolutely impossible for the paper-makers to keep pace
with the paper consumption, if they should not discover a sat-
isfactory substitute for rags. Experiments have been made
with the various vegetal fibres as a substitute for rags. Only
plants produced in large quantities can satisfy the demand.
Of the plants tried, maize seems to be the best adapted to the
l)urpose. In the last century two maize-straw paper manu-
factories were in existence in Italy, according to Dr. Johann
Christ Schaeffer's *' Sammtliche Papierversuche " (Regens-

' Anier. Journ. Pharm., 3 ser. v, 315.

1 86

liars Jib crircr. — Maize :

burg, 1772). Moritz Diamant, a Jewish writing master, in
1856, called the attention of the Bohemian Minister of
Finance, Baron Bruck, to the value of maize fibre in the
making of paper-pulp. The imperial paper factory at Schlo-
gelmiihle, near Glognitz, was authorized under Diamant's
direction to make a certain amount of maize paper. The
paper produced was not of a satisfactory quality, the cost
was too great, and the manufacture forthwith stopped. Moritz
then tried to interest private parties in his enterprise, but
without success. In 1859 he again applied to the govern-
ment, and Baron Bruck, at the advice of judicious men, again
permitted Diamant to try his hand. It was found that the
chief expense lay in the transportation of the crude material
to the seat of operations. A half-stuff factory was erected in
i860, at Roman-SztMihaly, near Temesvar, where the culti-
vation of Indian corn was extensive. The half-stuff was so
poor that operations were again suspended. The first period
in the manufacture of maize paper closed. These failures led
to important results, for it was found upon further experi-
mentation that the husks yielded a fibre which could be spun
and woven. All the fibre and gluten wastes can be used in
the manufacture of paper. Dr. Alois Ritter von Welsbach,
Director of the Royal State Printing Establishment, discovered
the process. The catalogues of the Austrian exhibition at
London (1862), in German, French and English, consists of
such paper. The manufacture of paper in Vienna from maize
is, at the present time, in extensive operation. The "Allge-
meine Zeitung," a scientific paper of importance, is made of
maize paper. The yellowish tint is restful to the eyes.' The
advantages of using Indian corn in the manufacture of paper
are many: (i) Very little sizing is required; (2) it bleaches
well; (3) it has greater strength than rag paper; (4) no ma-
chinery is necessary for tearing up the leaves.' In the
manufacture of paper, the leaves are digested in hot water
for two days. They are then separable into three parts : (i)
The large veins and ribs, which serve for coarse gunny

1 Intellectual Observer, iii, 4<'vS.

- Amer. Journ. Pharm.. 3 ser., ix, 232.

A Botanical a fid Econojnic Study.

187

bags, cordage and the finer fibres for cloth ; (2) the material
between the ribs is made into bread of an agreeable taste ;
(3) the coarse paste, which finally separates, is used in the
making of paper. The perfection of the process is largely
due to the efforts of Pfob, Jung, Marsanich and others, men
who deserve the highest commendation for their industrious
perseverance.

Oir.. — The oil is not obtained by direct expression, but the
grain is malted, and the germ is separated by careful crush-
ing and winnowing. The germs are then submitted to hy-
draulic pressure, and yield 15 per cent, oil, and a press cake
rich in albumen, containing 4 to 5 per cent. oil. Maize oil
is of a pale golden-yellow color, and has a peculiarly agree-
able taste and odor. It is a thick liquid, and has a specific
gravity of 0.9215 at 59^ F. In consists of olein, stearin, pal-
mitin, and contains some volatile oil. It solidifies to a quite
solid mass at 10° C. (14° P\)'

Maize oil is well adapted for illuminating purposes, giving
a bright white flame, and in burning it develops a high degree
of heat. It is used advantageously in the dressing of wool,
as a lubricant for machines. The yield is sixteen pounds to
every 100 bushels of grain.

Fuel. — In the Western States, where the supply of fuel is
precarious, the whole ear of corn has been used as fuel, but
preferably the cob deprived of its kernels. Three tons of
corn-cobs equal one ton of hard coal as fuel.' In France, the
cobs, saturated with resinous matters (sixty parts melted
resin, forty parts tar) are used as fire lighters, and are bought
at prices ranging from twelve to twenty francs ($2.40 to
$4.00) per thousand, according to the size used.^

The husks are used in packing oranges and cigars, in the
stuffing of pillows, mattresses and lounges. The cob is used
as a stopper for bottles. The toasted meal is substituted for
coffee.

263.

^ Amer. Journ. Pharnu, 4 ser., XV, 403; Bronnt, Animal and Vegetal Fats and Oils,

* Journ. Soc. Arts, xx[, 235 ; Council Bluffs Nonpareil.
■* Journ. Soc. Arts, xxiii. 887.

I

;i

i88

HarsJibergcr, — Maize :

The cobs yield an abundant supply of potash. Large mills
can furnish a big product of corn-cob potash. A mill shell-
ing 500 bushels an hour turns out 7000 pounds of cobs an
hour, or equal to 70,000 pounds per working day.^ The cobs
are used for fuel in the mills, and the refuse ashes are col-
lected for the extraction of the potash, 1000 pounds of cobs
yielding 7.62 pounds of potassium carbonate, or in a factory
of the above capacity 535 pounds per day.

It seems strange that during the century which has elapsed
since our birth as a nation no adequate conception has been
reached as to the true value of the commonest product of our
soil — our native Indian corn.

1 Hazard, Amer. Journ. Pharm., 4 ser., 11, 152.

A Botanical and Economic Study.

189

CHAPTER \TI.
Economic Considerations.

THE importance of agriculture in laying the groundwork
of a true national prosperity is recognized by all. A
historical retrospect proves that with the decay of
agriculture in the States of Greece and Rome and the growth
of cities, the political habits of the people underwent a
decided change for the worse. Many writers have pointed,
therefore, with justifiable alarm to the last decennial census,
which shows that one fourth of the entire population of the
United States dwells in large towns and cities, and in many
places the most progressive part of the rural classes have
moved to the large industrial centres. Many forces conspire
to produce this change. A few of them are the diminishing
returns of agriculture, exorbitant transportation rates, natural
monopolies, unsatisfactory financial conditions and state
interference by partisan legislation.

The law of diminishing returns is that the product propor-
tionally decreases with increased application of labor ; that a
time will be reached when, with intensive cultivation, the
land will be finally and irremediably impoverished. Some
decades ago very bad agricultural practice prevailed in the
United States; one crop was grown exclusively on the same
piece of land. The cereals, wheat, rye and barley, which were
largely raised, sown successively for years in the same field,
rapidly exhausted the soil of the most valuable ingredients.
It is characteristic of starch and sugar-forming plants to take
large quantities of nitrogen from the soil. The plants are so
constituted physiologically as to need large supplies of nitro-
gen, potash and phosphorus for the proper storage of the
carbo-hydrates. These three essential elements are the most
difficult to restore to the land in sufficient quantities when

190

HarsJibergcr. — Maize

once the soil is exhausted of them. There has been an
over-production of the cereals in America, with the natural
deteriorating effects upon the soil. The enormous exporta-
tion of 55,131,948 bushels of wheat in 1891 illustrates vaguely
the annual drain upon our soils. The American system of
agriculture is imperfect, slovenly and wasteful in the ex-
treme. A change for the better has been manifested of late

years.

Modern experiments prove that the production of the
nitrogen-consuming plants (cereals, root-crops and fruits)
should be alternated with the production of the nitrogen-
storing plants, such as clover, beans, peas, vetches and
lupins, which accumulate atmospheric nitrogen by the agency
of the nodules on the roots, inasmuch as the soil is best con-
served with a properly adjusted rotation. Mr. Mason's trials
at Eynsham, Oxfordshire, are referred to a second time, as
illustrating this important principle. His idea is to grow
mixed crops of leguminous plants, liberally manured with
basic slag and kainit, and to convert the produce of the first
year into silage and of the second into hay. The land is thus
occupied for two years and the assumption is that in this way
highly nitrogenous crops will be obtained with mineral, but
without any nitrogenous manure, and that the land will be
left, so far as nitrogen is concerned, in a high condition for
the growth of saleable crops, such as potatoes or grain, which
need nitrogenous manuring. In other words, the plan, as he
puts it, is "to grow nitrogen-accumulating crops for home
consumption and afterwards nitrogen-consuming crops for sale
or export to foreign countries."

This system should be extended so as to comprehend the
whole country in a complete and perfect system of rotation.
The adaptability of particular districts of the United States
for certain kinds of agriculture, should be observed in com-
bination with the most scientific succession of crops, one
crop following another in such a manner as to yield the
largest returns, and at the same time maintain the fertihtyof
the soil. The plant best suited to any rural section, that is, the
one from which best financial returns are expected, should be

A Botmiical ami Econoviic Study.

191

grown as the major crop, and the other plants subordinated as
minor crops. In the growth of maize as a major crop, for
instance, in any locality, the minor crops should be associated
with each other and with maize, so as to permit of the largest
production of maize, and, at the same time, prevent rapid soil
exhaustion. The major crop, again, in the New England
States, for example, should be associated with the major crop
in the Central States, and that with the major crop of the
Southern States, to constitute a national system of rotation.
The national rotation will permit the best use of the whole
country, and will make it an agricultural unit.

A partial division of labor in agriculture will be feasible,
for the farmer becomes a specialist in the growth of that
plant for which his region is especially adapted. He will
study, more scientifically, the physiological requirements of
his major crop, and be able better to solve the numerous
questions of practice which constantly arise. It will be
necessary, however, for the central government to study the
especial adaptability of the physiographical sections of the
United States before an agricultural subdivision of the country
can be scientifically made. These regions, at the present
time, can be designated only roughly.

Thus the physical conditions of Florida make it plain that
this peninsula is to develop its life on the lines of agriculture
and marine industries. The agriculture is destined to be of a
peculiar sort, gardening and fruit-growing rather than ordinary
field tillage. Such tropical and sub-tropical fruits as the orange,
the lemon, the mango, the sapodilla and tender vegetables,
are easily raised, and assure the agricultural position of the
district. The low-lying portion of the Gulf States, an old sea
bottom, which has been elevated recently above the ocean,
contains soil of only moderate fertility, but well suited to
the great staple, cotton ; rice can be grown to advantage
along the river bottoms. The Blue Grass region of Ken-
tucky and Tennessee lies in the range of the Silurian lime-
stones, and the soil possesses great fertility. This region,
which grows very nutritive grasses, will be given up to
stock raising. The northern Central States, Ohio, Indiana,

192

HarsJibcrger. — Maize :

Illinois, Missouri, Iowa, Kansas and Nebraska have soils and
climates especially adapted for the growth of maize. They
form collectively the great corn belt, and are also known as
the surplus States. Regions protected from late frosts will
be eminently suited for most large and small fruits. Such
places are situated in western New York, Vermont, Pennsyl-
vania, peninsular Michigan, Delaware, peninsular Maryland
and California, including the irrigable lands. The irrigable
lands have very great fertility, and are singularly enduring to
tillage. Their vast extent permits a great diversity in the
product of the watered fields. Thus in Montana and Idaho
the natural products, grasses, are grown, while in New
Mexico and Arizona the finer fruits may be advantageously
cultivated. Future research will reveal the particular regions
where definite crops, such as cotton in the South, will be
raised exclusively by reason of the climatic, geological and
meteorological conditions.

The national system of rotation will cause the supply re-
quired for use in any section to be drawn from the region
where the crop, as a major one, is raised most advantage-
ously. The production of maize illustrates this. The seven
States of Ohio, Indiana, Illinois, Iowa, Missouri, Kansas and
Nebraska are the "corn surplus States," practically furnish-
insr all that enters commercial channels. Outside of these
seven States the yield is practically of only local interest.
The crop is consumed where grown, and it exerts an influ-
ence on commercial corn owXy as it supplies Jioinc requirements
or 7nakes necessary a demand on tJie surplus States. The
great bulk of the corn crop is used at home, in fact, is con-
sumed upon the farms where grown, and but a very small
proportion is ever shipped abroad.

A Botanical and Ecomnnic Study

193

Distribution and Consmnption of Maize y 1891.

Section.

Retained for County
Consumption.

Bushels.

Percent.

Distribution beyond
County Lines.

Bushels. Per Cent.

Eastern ' 8,336,000

Middle 64,331,190

Southern 352,019.990

Western ' 859,505,700

Pacific 3,992,330

99.4
91-3

94-3
84.4

87.4

53,120

6,125,810

21,155,010

159,211,300

576,670

.6

8.7

5-7
15.6

12.6

Suppose the supply of maize required for consumption in
New England to fall below the quantity raised, which may
happen, since in New England maize is a minor crop. The
deficiency is met by drawing upon the regions where there is
a surplus of maize, which will be where it is grown most ad-
vantageously as a major crop. The price is regulated by the
cost at the locality where it is raised at the greatest disad-
vantage. The farmer in District A disposes of his major
crop at the same rate which prevails in District D, where the
product is grown as a minor crop at a greater relative disad-
vantage. Likewise, the farmer B derives from his major
crop what the farmer C obtains for it. An equalizing and
comi)ensatory action, therefore, is established in agricultural
production.

Crops.

Districts

a + 2b + 3C + 4d -f 5e = '5x.

5a + b -h 2c 4- 3d + 4e = I5X-

3a + 5b + 4C 4- 2d -f e = 15X.

2a 4- 3b + c -h 5d + 46 = 15X.

4a -f 2b 4- 5c + 3d 4- e = 15X.

= b = c = d = e = x; x = unit of usefulness (udlity).
Figures and letters in heavy-faced type indicate major crops.

The above diagram represents the national system of agri-
culture. The algebraic letters indicate the separate crops

A.
B.
\ C.
D.
E.

Legend. — a

194

Ha rsh bcrger. — Ma ize :

grown ; the coefficients indicate the relative importance of
the crops a, b, c in terms of the utility x. It is seen that the
sum of the utilities in each region, A, B, C, D and K,
is the same. The lands of the United States by this system
become of equal usefulness, for the relative disadvantage of
District A, for instance, in one crop is equalized by its rela-
tive great advantage with respect to some other crop. This
equalization of usefulness will have a material influence on
rent. If the productiveness of the various agricultural re-
gions of the United States can be equalized, rent will be
equalized. If the Ricardian theory, that rent depends on
the difference in the productiveness of different soils, is true,
then this equalization will result in the lowering or total
abolition of rent.

Districts and farms, besides varying in fertility, vary also
with respect to their nearness or remoteness from centres of
population. Even if we succeeded by a proper national rota-
tion in making all agricultural lands of equal productivity, yet
rent would arise owing to the differences in the distance from
market. A farm in proximity to the market would be better
situated and would command a higher rent. A national pro-
tective system, which builds up local centres of trade and
industry, materially diminishes market distance and corres-
pondingly lowers rent. The products of the soil are consumed
in the vicinity of the farms, and the farmer has at hand the
means of making such a return to the soil as will maintain and
even increase its fertility.

What shall be the motive force that shall inaugurate a
wiser system of agriculture and distribution t Increase of
intelligence must be the main factor in accomplishing a pro-
gressive and substantial advance. Man rises superior to his
environment. Progress is possible because the intellectual
superiority of man enables him to modify infinitely the forces
of nature. Progress should begin subjectively in the men
themselves. They should become familiar with the idea of
self-government, of moral restraint, of the beautiful and of
the good. This change will first become apparent in the con-
sumption of individuals. Old articles, which before satisfied

A Botanical and Economic Study

195

the cravings of their appetites, will be replaced by new
articles, which are better adapted to intellectual and enlight-
ened men. Nowhere will this change be more apparent than
in the consumption of food.

Instead of consuming such large quantities of food (par-
ticularly of fats and starches), a less quantity better adapted
to the needs of the human system will be substituted. Mus-
cular labor requires large supplies of proteids ; intellectual
labor, light food with condeflsed nutriment. Americans
should observe these physiological laws, for, as a rule, their
diet is poorly adapted to the labor they perform. The large
consumption of carbo-hydrate vegetal food is largely augmented

by the use of fat meats. It is evident from principles already
advanced, that the excessive production of fats (in animals),
starches and sugars impoverishes the soil. The American
people must live upon those articles of food of which the
American soil is most productive, and must cease to consume
in large quantities those articles for which the soil is poorly
suited. It is possible with this change to get relatively the
same nourishment at a fraction of the previous cost, because
food is produced cheaper. A more extended dietary will follow
such an alteration in the tastes of individuals, for there are
many articles which are nourishing and cheap which are easily
substituted for the dearer forms of food.

A high standard of life means a variety in consumption.
The first condition of such a standard is the reduction of
primitive appetites and passions. "So long as a few primi-
tive wants are intense the standard will be limited to the few
articles which gratify them." ' The relative urgency of the
other wants can only increase when the primitive wants sink
to a level with the new desires of civilized life. These changes
in the consumption of men will work eventually enormous
changes in production.

Agricultural production will be greatly improved. "The
best use of all our land will come when this change in the
habits of the American people is [accomplished]. There are
lar^re tracts of land which cannot be utilized because the

1 Patten S. N., Dynamic Economics, 129.

196

HarsJiberger. — Maize :

American people do not want the plants which can be grown
in these areas. So long as the home market does not demand
any other articles of food than those staple ones to which
our ancestors in Europe were adjusted, there can be but little
use made of districts of our country for which [certain plants]
are most fitted. A much greater improvement in the condi-
tion of the American people could be made by adjusting our
consumption to American conditions than by all the machines
that it is possible to devise. '^ These changes in the con-
sumption of food make it possible to construct a national
system of agriculture.

The national rotation makes it possible to have many arti-
cles upon a plane as regards price and nutritive value. Meat,
beans, peas, cheese, for instance, will be placed essentially on
the same level, as regards marginal utility or value. When a
scarcity of any one of these articles occurs, another can be
substituted immediately without effecting any hardship.

In the standard of life old articles will be replaced by new
ones which have the same ratio of cost to utility as the old
ones. This renders it easy to construct a scale of total con-
sumption. The total consumption of an individual consists
of groups composed of isolated articles, which stand higher
or lower in the scale according to their marginal utilities.
Articles of the same marginal utility (value) and same nutri-
tive ratio can replace old articles of the same value in the
group without in the least affecting the general usefulness of
the group. The association of the articles together in a food
group can be represented by a curve. The highest point of
the curve represents the position of the articles which give
the most satisfaction and pleasure, and the lowest point
represents the position of those articles which give the least
pleasure :

' Patten, S. N., Economic Basis of Protection, 121 I am indebted for many of the
views here presented, concerning a dynamic society, to the lectures of Professor Patten,
deliverfd at the University during 1892-93, and to his books and numerous of his articles
on the subject, but the responsibility for their connection in this chapter rests with me.

A Botanical and Economic Study.

197

(a) Bananas; {!)) Oranges; (r) Apples; {d) Peaches; {e) Pears; {f )
Plums; [g) Figs.

(a') Beef; (^') Mutton; (<:') Cheese; {d) Pork; (^M Eggs.

4i\

• {a"-) Wheat; (A') Maize; (^-) Oats: {d') Rice.

3:

{a^) Potatoes; {b") Tomatoes; [c') Beets.
^1 (a») Water; {b') Milk; {c') Beer.

• I:

The articles on the numbered horizontal lines in the diagram
separately indicate the complimentary goods which can be
substituted the one for the other, because article a=^b = c =
rt^ = ,r marginal utility. The articles on line 5, which give,
suppositionally, the most pleasure, have the same marginal
utility, and have been produced in the national system of
agriculture at relatively the same cost. The articles a\ b\ c
etc., on line 4, also have the same marginal utility, as also
those on the lines i, 2 and 3, respectively. Articles can be
equal in value only when the same degree of utility is im-
puted to them. When every individual shall recognize, for
instance, that cheese in definite proportions has the same
nutritive value as meat, i. c, supplies the same bodily wants,
when meat no longer satisfies the taste, cheese, beans, eggs
or lentils will be substituted for meat, and the usefulness of
the group remain the same.

Agricultural production directly responds to this change
in consumption, and the national .system is possible. The
general rotation is feasible, because the demand is for a large
variety of goods arranged in scientific proportions. The
farmer is directly benefited, because he derives a larger
surplus, which can be devoted to social and agricultural
improvement.

198

HarsJiberger. — Maize :

Cheaper food is also possible to the industrial population
with the better agricultural system, for a division of labor
into particular specialties ot agricultural production is pos-
sible. This means a larger surplus for the laboring classes
and increase of efficiency, because the bodily system and
mind are improved by the use of better food. With increased
efficiency the hours of labor will be shortened, and, as a con-
comitant of the greater efficiency of the laborer, the cost of
manufactured goods will be lowered. The farmer is benefited
as well as the city man by this reduction in costs.

The larger surplus applied to education increases the
intelligence of the community.' The increase of intelligence
checks the rapid multiplication of the population. *' Only
when the increase of mankind shall be under the deliberate
guidance of judicious foresight can the conquest made from
the powers of nature by the energy of scientific discoveries
become the common property of the [race]." With the im-
proved industrial conditions and decrease of selfishness, by
philanthropic sympathy, a larger return goes to the laborer
as wages. Higher wages mean more comfort, and with
cheaper food the laboring man will make enormous strides
toward a high standard of life.

The full development of the agriculture of a country fol-
lows a strong protective policy, for by it new markets are
created and fostered. New centres of industry mean a short
haul for the farmer. The soil is not exhausted as rapidly of
its ingredients, because the waste of large centers of popula-
tion is returned to the fields in the immediate vicinity of the
market.

The stability of the social fabric, tJic agricultural prosperity
and the future industrial achievements of the nation, de-
pends upon a strong national spirit, which renders a society
dynamic.

> The very suggestive report of Dr. Harris, United States Commissioner of Education,
should be read in connection with the above. It shows clearly what a diversification of
industry and agriculture means for the social and moral education of the masses. Kep.
Com. Educ, 1889-90. Vol. i, pages 22-26.

A Botanical and Ecojwviic Study,

199

CHAPTER VHI.

Future.

^r^HE use of maize promises to increase in the near future.
I The plant subserves so many important purposes that
it cannot fail to occupy a prominent position in the
future agricultural production of our country.

Not only will Indian corn be raised for home consumption,
but also for foreign export. The people of Europe must
be taught to enjoy maize as a food. President Harrison's
administration, in the person of Secretary Rusk, made a wise
movement when it appointed a special agent to look after
corn interests in Europe. His efforts have met thus far with
a due measure of success, but Colonel William J. Murphy
finds it difficult to overcome the prejudices and conservatism
of the peasants who need most sorely a cheaper food. Ex-
cept an insignificant amount, exported corn is used chiefly in
Europe as a food for animals, as a grain for distillery pur-
poses and starch manufacture. - The only form of corn at
all known abroad is corn starch, which is sold principally in
the British Isles under the name of corn flour." ** A better
knowledge of maize as a human food, in addition to bringing
into use its other forms, will increase the demand for all its
products, which will call forth a supply that will cause the
price in Europe to fall from its present high artificial point.
- There are multitudes of half-starved toilers in Europe who
would welcome the golden grain if only they were taught its

merits."

To supply the wants of these people would be to render a
philanthropic service worthy of our best endeavors, and the
increased export of our cereal maize would be a sure, practical
and speedy benefit to the farming interests of our country.
Great crops and a demand for the products of the soil

200

Ilarslibergcr. — Mairjc

strengthen credit, expand the volume of manufacture, speed
the wheels and fill the sails of commerce, and make a nation
prosperous.

Maize, the greatest arable crop which we grow, occupying
the largest portion of our cultivated area, has never been
known to fail. It is destined to occupy, in America, the
place that rice fills in India, China and Japan, that cassava
fills in South America, and that sago occupies in Borneo,
Java and the Indian Archipelago— the staple food of man.

Maize lends itself thoroughly to use in architecture and
mural decoration ; for industrial designing it has unrivaled
preeminence. ''Let the rose, queen of flowers, bloom for
England ; let Ireland honor the shamrock ; Scotland her
thistle bold ; let the lily unfold her pure white petals for the
joy of France; but let the shield of our great republic bear
the stalk of bounteous golden corn."

The author is indebted to the following gentlemen, who
materially aided him: Major J. W. Powell, A. S. Gatschet, F.
Webb Hodge and Frank H. Gushing, of the United States
Bureau of Ethnology, for suggestions as to the North Amer-
ican Indians ; Professor Otis T. Mason, National Museum ; Mr.
Stewart Gulin ; Professors Simon N. Patten, whose economic
works have been drawn upon, J. T. Rothrock, W. P. Wilson,
John A. Ryder and J. M. Macfarlane, of the University of
Pennsylvania; Dr. William Garruthers, British Museum, Dr.
J. H. Gilbert, Rothamsted, England, Mr. John Redfield and
others.

PHn.ADELi'Hi.A, June, 1893.

A Fyotanical and Economic Study. 201

EXPLANATION OF PLATES.
Illustrating Dr. John W. Harshheiujek's paper on Maize.

PLATE XIV.

Wild Mexican Plant g;rovvn in Philadelphia. Shows terminal pendu-
lous male panicle and contracted lateral branch, with ears set alternately
upon it. In the axils of other leaves, as shown in the plate, ears are
developed. Size of full-grown plant, five feet.

See Cornell Agricultural Experiment Station Bulletin (Bui. 49, Dec,
1892, p. 332) for photograph of a plant with fully developed lateral
branches with ears set upon them, each branch terminated by a small
male tassel.

PLATE XV

F
F
F
F
F
F
F
F
F
F

g-
g-
g-
g-
g-
g-
g-

3-
4-

5-
6.

1-

Details of Gross Anatomy.

Branch of male panicle showing spikelets.
Paired spikelets removed.
Male spikelet dissected showing two flowers.
Cross plan of male spikelet with pollen below.
Dissection of pointed grain of Mexican corn.
Sprouted grain.

Lateral branch with husked ears. Front view,
ig. 8. Lateral branch with husked ears. Back view,
ig. 9 Branch with alternate arrangement of ears. Husks removed.
. ig. 10. Ideal longitudinal section of a portion of an ear with female
spikelets in a hardened depression of cob with ovary, glumes and palets.
Fig. II. Cross plan of female spikelet, showing empty flower.
Fig. 12. Cross plan of lateral branch with four ears. Husks, or leaves,
lettered in order from base of branch to top.

PLATE XVI.
Histology.

Fig. F. Cross section of secondary root.

P. Phloem patches.
Fig. 2. Longitudinal section of emerging secondary root {t. e. aerial).

C. Calyptrogen layer.
Fig. 3. Cross section of leaf

S. Stereome.
Fig. 4. Cross section of leaf-blade near margin.

B. Pulliform cells.

Vi :j

:

202

Harshbergcr. — Maize :

[Vol. I, Plate XIV.]

Jn)t. Cont. riiiv. Pcfutsylvania,

PLATE XVII.

Map showing the original home of maize with its geographical dis-
tribution in space and time. The heavy contour lines represent elevation ;
one set shows an elevation of looo feet, the other 4500 feet and over, as
indicated by the letters. The arrows show the direction of maize distri-
bution. Note that corn entered the United States from Mexico and the
West Indies.

The squared area on the map shows the position of the agricultural
tribes in North and South America. It is evident that the position of the
agricultural tribes and the area of maize distribution are identical. See
for discussion the Chapter on Ethnology.

Legend in squares at the side :

1. Original home of maize.

2. Limits of primitive cultivation by the Mayas and allied tribes.

3. Limits of distribution in North and South America prior to the
year 700 A.D.

4. Limits reached in North and South America by the year 1000 A.D.

Harshberger on Maize.

[Vol. I, Plate XV.]

Bot. Cant. Univ. Pennsylvania.

H.\ksin5KK(ii.K ON Maize.

[Vol. I, Vlitc \VI ]

/uft. Cant. Unw. Punisylvatiia.

\

1

;

I I.\k>iii'.i:K(.i.R ox M.\i/i-;

p/ol. I, Plate XVII.]

Bot. Cont. Univ. Pennsylvania.

4 l_

I

.1

IlAKSHIiEKCKK ON MaIZE.

A Chemico= Physiological Study of Spirogyra

nitida.

By Mary Engle PenninCxTon, Ph.D.

Research Fellow in Botany.

AS the physiologist investigates more and more mi-
nntely the problems of plant life, he is confronted
on every hand by chemical qnestions, to many of
which he can still give no answer. The pharmacist has
introduced us to a great variety of substances obtained
from vegetable sources, but he has told us nothing of
their origin or use in the plant economy.

Together the chemists and physiologists, working in
this field, are little by little surmounting the difficulties
which beset the path leading to our more perfect under-
standing of the complex results as we find them in plant
life. It was with the hope of adding a few additional ob-
servations to our knowledge of chemico physiology that
the following investigation was undertaken.

Spirogyra nitida was selected for this work because of its
quick response to stimuli and because of its simple struc-
ture. A considerable quantity of easily accessible and un-
usually pure material which appeared in a pond in the
Botanic Garden of the University of Pennsylvania made
the selection possible.

By far the greater share of the knowledge which we pos-
sess of the laws of growth has been derived from a study
of the higher plants, where we have a division of labor.
In this simple alga each cell functions, so far as we
have observed, as an independent organism, building up
the necessary compounds for its maintenance and giving
off its waste products. The study of such simple forms
14 '^^

•I

I!

!
4

i

204 Pennington— A Cheniico-Physiological

should be rich in suggestions for the explanation of the
life-history of higher types.

The results of the investigation here offered are, in many
respects, incomplete. But it is believed that the work so
begun may suggest further study along the lines which are

here touched upon.

The research was carried out under the direction of Prof.
John M. Macfarlane, of this university, and it gives me
pleasure to express my appreciation of his never-failing
kindness throughout the entire progress of the work. I
am also indebted to Prof. Edgar F. Smith, who has placed
at my disposal all the facilities of the John Harrison Lab-
oratory of Chemistry for the prosecution of the purely
chemical part of the research.

Analysis of Spirogyka Nitida.
For the analyses made during this investigation Spiro-
gyra nitida, in pure culture, was used. The material was
obtained from a pond in the Botanic Garden of the Uni-
versity which was singularly free from other species of the
crenus. Some FJodea and a large water lily were the only
plants growing in the pond. The depth of water m the
deepest part was about four feet. The material was always
collected during the morning of clear days and was pre-
pared for use immediately. It was washed in running
water until any mud which might have adhered to the fila-
ments was removed. Distilled water served to free it from
all inorganic salts which the river water held m solution.
The clean threads were then laid on glass plates which
were inclined at a sharp angle, so that all water clinging
to the threads might run off. The threads were accepted
as dry when they assumed a light green color and their
individualitv could be distinguished.

In order to determine the water, dry substances, and ash
of Spirogvra, about sixty-eight grams of pure material were
dried at ^io° C until constant weight was obtained. It
was then found to have lost 89.94 per cent, of water. The

Study of Spirogyra nitida, 205

dry substance remaining was carefully burnt, when another
loss in weight occurred corresponding to 7.54 per cent. The
ash, after this treatment, was 2.52 per cent, of the original
material. Spirogyra 7iitida then contains : —

Water 89.94%

Dry matter 7-54%

Ash 2.52;^

Total 100.00

G. Mann,* gives for a mixture of S. nitida and S, jugalts

Water 96.8 %

Dry matter 2.72%

Ash 48%

Total 100.00

which differs widely from the results obtained in this labo-
ratory with pure Spirogyra nitida. Several determinations
were made with material gathered on different days, but
the results were practically in accord.

Combustions.

While Mann, in the paper cited, gives the water, dry
matter and ash, he makes no mention of an analysis for
carbon, hydrogen, nitrogen and oxygen. In fact such an
analysis has not been found in the literature on algae.
Therefore, in order to determine the primary composition,
a series of combustions was made under the following
conditions : —

The plant was removed from the pond on a clear, cool
day in early October. It was prepared for use as previously
described, then dried at 110° C. to constant weight. The
same material was used for the entire series of combustions,
and after each portion had been removed from the weigh-
ing bottle the remainder was again heated to 110° C. This
precaution is necessary in order to secure uniform samples,
since the finely divided material is exceedingly hygroscopic.

*Proc. Bot. Soc, Edinburgh, Vol. 18, (1890).

II

!i

ii

2o6 Pennington — A Chemico-Physiological

The weighing bottle, for additional protection, was kept in
a sulphuric acid desiccator.

The combustions were made in an open tube in a cur-
rent, first of air, then of oxygen, because the substance
proved exceedingly hard to burn. A period of eight hours
was required and the highest temperature available in the
combustion furnace. Very hard glass tubing was used.
This difficulty in burning is noteworthy, since it is rather
opposed to preconceived ideas.

A white sublimate was invariably found in the front part
of the tube at the close of the operation. This proved to
be volatilized alkaline chlorides. The ash remaining in
the boat was light gray, but could not be weighed because
the loss of alkali introduced an error.

The heating should be very gradual at first, so as to drive
over the water slowly but steadily. When this has been
collected in the calcium chloride tube the heat is increased
and maintained until all the carbon has been burnt to oxide.

The oxygen in the potash bulb and calcium chloride
tube was displaced by a carefully purified air current and
the weighings made in this gas. Seven analyses of the
material yielded the following results :

Maximum Minimum Mean

Substance taken . . . 0.2398 0.0931 0.1810

Carbon found . . . .46.66% 46.15% 46-35%

Hydrogen found . . 5-69% 5-005% 543%

Accepting for nitrogen 2.61 per cent, as was found by
a Kjeldahl determination, and for the percentage of ash in
the dry substance 15.43, the oxygen by difference is shown
to be 30.21 per cent. The primary composition of Spiro-
gyra nitida may then be formulated as

Carbon 4^-35

Hydrogen 5-43

Nitrogen ^-^^

Oxygen 30-2I

Ash • ^540

100.00

Study of Spirogyra nitida.

207

Analysis of the Ash.

During the preparation of considerable quantities of ash,
the apparatus best suited to obtain a perfect combustion
was found to be a broad platinum dish, over which was
suspended a wide glass tube a little longer than an ordinary
lamp-chimney, (Schultze method.) The air current so pro-
duced causes an even and rapid oxidation of the carbon,
with the expenditure of a minimum amount of heat. The
ash so burnt was of a pale gray color and perfectly homo-
geneous. It was preserved for analysis in closely stoppered

weighing bottles.

The method of analysis was essentially that of Bunsen
as described in his ^'Aschen Analyse.^' According to
this analyst the weighed ash should be brought into a
cylinder with water, and the latter saturated with carbon
dioxide until the liquid becomes colorless. The solution
should then be transferred to a platinum dish and evapo-
rated to dryness on the water bath, heating finally at 160° C.
By this means all the lime should exist as neutral carbon-
ate, and a water extraction should give only the salts of the

alkalies.

The aqueous extract was made up to a known volume
and divided into five portions. In the first the chlorine
was determined, in the second the sulphuric acid, m the
third the alkalies, in the fourth phosphoric acid, and m the
fifth carbon dioxide. Tlie insoluble residue was analyzed
for silicic acid, phosphoric aqid, calcium, magnesium, iron
and aluminium, sulphuric acid and carbon dioxide.

An analysis of Spirogyra ash, conducted as above out-
lined, gave the following percentage composition :

I. II. "I-

sio, 5.29 5.38 5.30

S03 9.13 9.II

Ql ^ ^ ^ 24.24 24.08 24.51

P,o/ \'' °-898 0-9I

CaO 9.01 9.20

MgO 2-2« 2-'^

208

Pennington — A Chemico-Physiological

AljOa
NajO

}

2.06 2.CX)

32.15 32.00

3.78 3.93

CO2 ii-io i°-90

Total 99.86 99.69

A glance at the recorded analyses of plant ash shows im-
mediately the great variation in their quantitative, though
not in their qualitative, composition. Not only do plants
of different genera vary largely, but those of different
species are quite as changeable.

Fucus
vesiculosus.

Fticus
nodosus.

Fucus
serratus.

La mill aria
digitata.

TT 0

15.23

10.07

1

4.51

22.40

NajO

24.54

26.59

31.37

24.09

PflO

9.78

12.80

16.36

11.86

MgO

7.16

10.93

11.66

7-44

FcjOa

0.33

0.29

0.34

0.62

P2O5

1.36

1.55

4.40

2.56

Of)

28.16

26.69

J 21.06

13.26

Ov/j ••••.••••

SiO^

1.35

1.20

0.43

1.56

Cl

•
15.24

12.24

11.39

17.23

T

0.31

0.46

1.13

308

Total ash

Total

13.89

14.51

1389

18.64

103.46

103.32

102.65

104.10

But few analyses of the ash of aquatic plants are to be
found in chemico-botanical literature, and these consist

Stndy of Spirogyra nitida.

209

largely of salt water forms. Fucoid plants were analyzed
by GiJdechens,* and his results are given in the precedhig

tablet

A comparison of the ash of these aquatic plants, Fucus

and Spirogyra, one a salt-water form, the other growing in
fresh water, is in several respects, interesting. The sodium
content of the ash is in both cases higher than the potas-
sium, and this is even more exaggerated in Spirogyra than
in sea forms. Chlorine attains a percentage of 17.23 in
Laminaria, but in Spirogyra we have 24.24 per cent. It is
exceedingly difhcult to account for the unusually large
quantity of silica, considering the soft and almost gelatin-
ous consistence of the plant. The amount is even greater
than we find in many hard, wiry land plants. Sulphuric
acid is considerably less in Spirogyra than in Fucus, and
phosphoric acid is in very small quantity. The magnesia
here falls below the magnesia content of Fucus, likewise
the lime ; but, on the other hand, iron and alumina are

much higher. . 1 , • •

We have in Spirogyra a plant which, though inhabitnig
fresh water that contains essentially the same constituents
as the soil through which it flows, yet differs essentially in
its ash composition from the land plants, and approaches
closely to the sea plants in its high sodium and chlorine
content, and its small amount of potash.

This great similarity to the salt water types led to a care-
ful search for bromine and iodine, but these elements could
not be detected.

* Von Gorup.Besanez has analyzed Trapa naians and finds for it an ash
content as given in this table. (Chem. Pharm. Centralblatt, 1861.)

Total ash KoO

13.69 6,06

NaoO

2.71

CaO

17.65

MgO

5.15

Fe,03

23.40

SOa

SiOa

2.53

Cl

27.34

0.46

Both iron and silica, in this analysis, are extraordinarily high,
t Ann. d. Chem. und Pharm., (1854) Vol. 54. p. 351-

i^ ■!

til

2IO

Pennington — A Chcniico-PJiysioIogical

Study of Spirogyra nitida.

211

Analysis of the Dry Material.

The material used in this analysis was prepared as
already described in this paper, and was dried at iio° C.
It was then ground to a fine powder.

O. IvOew. and Th. Bokorny,* state that an analysis of
the dry substance of Spirogyra (species not mentioned)
yielded them 6 per cent, to 9 per cent, of fat, 28 per cent, to
32 per cent, albumenoids, and 60 per cent, to 66 per cent,
cellulose and starch. These authors also consider lecithin
to be one of the plant's constituents, as well as cholesterine
and succinic acid.

Because of the large chlorophyll content of the plant it
was deemed advisable to extract the fat with petroleum
ether in such quantity that for each gram of substance
there was 10 cc. of solvent. It was allowed to stand, with
frequent shaking, for eight days. The supernatant liquid
was then decanted into a weighed flask, the residue well
washed with petroleum ether, and both extract and wash-
ings were evaporated to dryness in a vacuum desiccator.
The substance remaining was semi-solid and deep brown ;
it had a resinous odor. In quantity it amounted to 3.45
per cent, of the substance taken. Saponification yielded a
body resembling a resin. It was reddish brown, soluble in
potassium hydrate, from which it was reprecipitated by
hydrochloric acid. It was also soluble in sulphuric acid,
forming with this agent a deep red solution. Water pre-
cipitated the substance unchanged. Its melting point lay
between 85° and 90° C.

Probably a fatty acid was mixed with this resinous sub-
stance, since the filtrate, after extracting with alcohol and
ether, gave 0.43 per cent, of glycerol. The resinous body
itself amounted to 3.02 per cent.

IMicro-chemical tests failed to show the presence of a
resin in the cells. Fats, likewise, were not detected in the
vegetating cells. It is therefore likely that these sub-

'^Journ. f. Prakt. Chem., Vol. 36, p. 273.

Stances were incorporated with the chlorophyll. Indeed,
the presence of a resinous substance in a plant of this
character is hardly to be expected. Yet if we consider the
close relationship between the tannins and the resins some
licrht is thrown upon the matter. The tannin content of
the cell is comparatively great^S.;! per cent -and accord-
ing to Bastian * and others, resin is, in some plants at least,
produced from tannin.

The residue from the petroleum ether extraction was
treated with ethyl ether until everything soluble in this
menstruum had been removed. The extract was evapor-
ated to dryness in a tared flask and weighed. The dry
material so prepared was treated with cold water, but
nothing dissolved. It was then taken up with absolute
alcohol and proved to be totally soluble in this solvent.
The solution was clear, dark green, and seemed to consist
of very pure chlorophyll. The quantity amounted to 2.62

^"^Arextraction of the dry residue with absolute alcohol
gave a tannin content of 8.71 per cent. The method used
tas that of Lowenthal, as modified by Procter,! ni which
the tannin solution to be determined is titrated with a per-
manganate solution of known strength, after the addition
of an indigo carmine solution.

The aqueous extraction yielded 1470 per cent, of a gum
soluble in water, but insoluble in dilute alcohol. The addi-
tion of strong alcohol to the filtrate produced a precipitate
of dextrine-like carbohydrate, which amounted to 7-24
per cent This high content of mucilaginous substance is
perfectly accounted for by the appearance of the plant and
bv its soft mucilaginous character. ^

The acid content was found by precipitating a portion of
the aqueous extract with lead acetate, weighing the precip-
itate on a tared filter, then igniting. The loss in weight
represented the organic acids and amounted to 3.01 per cent.

* Am. Jour. Pharm., Vol. 68. p. 137.

t Journ. Soc. Chem. Imlustry, Vol. 3, p. 82.

2 1 2 Pennington — A Chcmico-Physiological

An effort to determine the individnal acids of the mixture
failed because of the small quantity of material available,
but succinic acid could not be found, and tartaric acid
yielded its characteristic reactious.

The sugars were determined by the reduction of Fehl-
ing's solution and the electrolytic estimation of the copper.
Starch was hydrolyzed and determined in like manner.
The sugar yielded 4.8 per cent, the starch 10.7 per cent, of
the material taken.

The substance insoluble in hydrochloric acid was, when
dry, a perfectly white, houiogeneous powder. It was
accepted as cellulose and the insoluble portions of the plant
ash, and upon weighing was found to equal 9.76 per cent.

A Kjeldahl nitrogen determination, couducted according
to Gunning's modification of the method, gave a nitrogen
content of 2.61 per cent. Calculating the albumenoids on
this basis we have 16.31 per cent, of such compounds.

The ash content of the water-free plant is 15.43 P^^ c^^^-
If this be included in the analysis of the dry material we
have :

Resin 3-02

Glycerol o.43

Coloring matter 2,62

Tannin 8.71

Mucilage i4-7o

Gums 7-24

Acids 3.o»

Starch • lO-?

Sugar 4.8

Cellulose . . 976

Albumenoids 16.31

Ash 15.43

Total 96.73

It is greatly to be regretted that the material could not
be obtained in pure culture in sufficient quantity to permit
of an investigation into the exact nature of the individual
substances of which it was composed.

Study of Spirogyra nitida. 2 1 3

Micro-Chemical Investigation.

Fresh, healthy material was examined under the micro-
scope for the substances usually present in plant cells.

As was expected, starch was present in abundance, and
was easily seen even without the aid of iodine solution.

Vannin was shown by ferric chloride solution, which
gave a color varying from deep green to black, according
to the quantity of the substance in the cell. Copper acetate,
in a saturated solution of which the threads were allowed
to remain for several days, gave a brown precipitate.

If a vegetating cell be tested for a reducing sugar by
Fehling\s''solution there will be seen in the interior of the
cell one, or more generally two, areas over which the copper
oxide is deposited. These areas have the appearance of
contracted pellicles carrying the cuprous oxide in their
walls. They are situated inside the chlorophyll bands, and
if the test is properly carried out, the position of the bands
in the cell is not materially altered.

The appearance and position of these structures indicate
that one of the innermost protoplasmic layers bounding
the vacuole of the cell carries in its substance the glucose.
By the prompt reaction of the Fehling's solution and the
glucose, the cuprous oxide was deposited where ongmally

the glucose existed.

The most characteristic form for the pellicle to assume
is that of a flattened cone having the base directed toward
the end of the cell. One such cone is placed on either side
of the nucleus. In no other part of the cell can any
deposit of cuprous oxide be detected, though careless hand-
ling may cause these pellicles to break into a number of
pieces when these will be found scattered over the cell.
Since 'only a small quantity of glucose is present m the
vegetating cell the pellicle bearing the copper deposit was
colored a light red-brown when viewed by transmitted

' The microscope revealed the presence of two varieties of

214

Pennington — A CJiemico-PJiysiological

crystal, one form having, normally, four pointed arms of
equal length radiating from a centre. Sometimes these
arms were broken off or splintered, or the crystals other-
wise maltreated, so that fragments could be seen lying
about. The size of these crystals was very variable.
They proved to be calcium oxalate, and as such were first
described by Mann.* He gives seventeen as the number
found in a single cell. In the material studied here more
than twenty were sometimes counted.

Another form had four short arms, two or three times as
broad as the oxalate crystals. The arms were smooth and
did not show any tendency to splinter. Though they
varied considerably, some attained a much larger size than
the oxalate crystals ever reached. Occasionally one was
found having a diameter more than half that of the cell.
They wxre dissolved by a lo per cent, solution of caustic
potash; likewise by hydrochloric and dilute sulphuric acid.
Sulphuric acid in alcohol gave a precipitate of calcium
sulphate. A 2 per cent, solution of acetic acid took these
crystals into solution, while a concentrated solution did
not affect them. This behavior corresponds to calcium
tartrate, and furnished an evidence of the presence of tar-
taric acid, which was later confirmed in the gross analysis.

I have not been able to find any mention in the literature
on this subject of the presence of calcium tartrate crystals
in Spirogyra. While not nearly so numerous as the oxalate
crystals, their presence was so constant that they may be
safely taken as a normal constituent of the plant

Trimethylamine Content of Spirogyra.

While preparing Spirogyra for use, that is, freeing it
from foreign matter, washing, drying, etc., it was noticed
that the hands of the operator acquired a fishy odor, as did
also the vessels in which the air-dried substance was pre-
served, and that, while drying, the same odor could be
distinctly detected. Drying at ioo° C. to constant weight,

*Proc. Bot. Soc. of Edinburgh, Vol. i8, (1890).

Study of Spirogyra nitida.

215

caused a total disappearance of the trimethylamine, which
proved to be the source of the odor. It did not reappear
even on boiling with potassium hydrate.

Fresh material boiled with caustic alkalies yields trnne-
thylamine in considerable quantity. Air-dried Spirogyra
also gives it, but in small amount, showing that loss has

occurred by drying. , .1 1 .

Loew in the paper before cited, asserts that the plant
contains lecithin because of the trimethylamine evolution
when acted upon by caustic alkalies, and states, as a further
proof, that he obtained phosphorus from the ether-alcohol
extract. He does not refer to the evolution of trimethyl-
amine at the ordinary temperatures, and indeed this begins
almost immediately upon removing the plant from water.

While we have considerable evidence to show that
chlorophyll is either a substitution product of, or closely
allied to, a lecithin-like body, we believe that the com-
pound so formed possesses greater stability than is indicated
by such a ready evolution of trimethylamine as we find m
spirogyra. Such behavior is more generally attributed to

a proteid. ^ . . , . ,

We have, among plants, a number of instances in which
this amine is given off. notably in the Stinking Goosefoot
iChenobodium Vulvaria), where it is produced by the
leaves and in the Hawthorn {Cralcrgus Oxyacantha) m
which' the small white flowers are the active parts. In
these cases the amine is regarded in the light of an alka-
loidal waste product, which, being gaseous at ordinary
temperatures, is not stored up in the plant tissues.

Trimethylamine is one of the simplest mtrogenous com-
pounds with which we are acquainted. Hence its presence
in plant tissues is exceedingly interesting, and the question
arises, Is it always present as a katabolic product, or is it,
L some cases if not all, produced as one of the primary
products in the synthesis of nitrogenous bodies?

Borodin * has found from his investigations with grow-

*Bot. Zeit., (1878).

1

2l6

Pennington — A CJicmico- Physiological

ing shoots of certain plants that etiolation tends to pro-
duce a large amount of asparagine, and accounts for the
accumulation of the amide by the carbohydrate having
been used up, while nitrogen assimilation can proceed
independently of light. From such experiments we may
suppose that either the proteid decomposes in yielding
the amide, or that it is produced synthetically and not
further used because of a lack of suitable combining
material. As there can be little doubt but that amides
are, under normal conditions, synthetically formed as inter-
mediate products toward the building up of proteid, the
latter supposition is more likely to be the correct one.

In order to determine if possible the role of trimethyl-
amine in Spirogyra, the plant was de-starched and kept in
darkness for about thirty-six hours. A portion was then
placed in a large test tube containing sufficient water, and
the tube tightly corked. After standing in the dark for
about half an hour, the cork was removed and the odor of
trimethylamine was plainly detected. Heating with potass-
ium hydrate gave a larger amount than was obtained
from starch-containing cells, showing that the amine had
accumulated in the cells which were kept in the dark.

While the less complex nitrogenous substances can be
found in the leaves and stems of healthy plants, neither
sugar,. starch nor amide is to be detected in the actively
growing tip, the energy here being so great that there is
immediately a union of carbohydrate and nitrogen com-
pound, the amount of each constituent being present in
exactly the correct proportion to form proteid. In other
parts of the plant, where the activity is not so great, the
separate constituents can be detected in the cell. If tri-
methylamine is a synthetic product, does it ever, when
growth conditions are favorable, accunuilate in the cell, or
is it given off in the water by the plant?

Bokorny* has found that Spirogyra remained healthy in
a 0.05 per cent, solution of trimethylamine neutralized with

* Chemiker Zlg., 1894, No. 2.

\

Study of Spirogyra nitida.

217

sulphuric acid, but that a deposition of starch did not occur
until the eighth day. It is not, then, injurious when in
small quantity.

Since the amine is readily absorbed by mineral acids,
yielding with them stable salts, we have a ready means by
which to detect and estimate it, and to answer some of
these questions for Spirogyra.

Newly formed threads of Spirogyra were placed in a
flask of distilled water which had been rendered ammonia-
free. A culture mixture, containing the nitrogen as nitrate,
provided the necessary inorganic food. As the plant grew
the gases were pulled out of the flask by means of an air
current and caught in sulphuric acid, from which the
trimethylamine was isolated. The apparatus in which this
work was carried out was arranged in the following

manner :

Air was admitted to the flask by means of a tube extend-
ing almost to the bottom of the liquid, and having a short
arm attached to a U tube containing sulphuric acid. This
served to free the entering air from ammonia and organic par-
ticles. Another tube, having a short limb, carried off" the
air which by this contrivance was compelled to travel
through the entire depth of liquid. This tube was attached
to a Liebig potash bulb containing sulphuric acid, which
effectually prevented any amine from passing through and
so occasioning loss. Between the absorption bulb and the
drop-aspirator, by which a steady air-current was mani-
tained, was fixed another U tube holding sulphuric acid,
to prevent any gases from decomposing organic matter
(which the river water might contain) pushing back into
the bulbs. The air current was regulated by means of
ground-glass stop-cocks.

As soon as the threads showed any tendency to collect
and sink together in the bottom of the flask, the long
masses which waved above were carefully lifted out and
transferred to a new culture solution, so that all decompos-
ing cells should be eliminated.

2l8

Pen nington — A Chemico-PJiysiological

t;

The air current was maintained steadily for three weeks.
At the expiration of this time the absorption bulb was
removed, the acid transferred to a round-bottomed flask,
rendered alkaline with potassium hydrate, and distilled.
The volatilized substances were collected in a large quan-
tity of distilled water which was afterward examined for
trimethylamine, and also for ammonia, but none could be
found. This experiment was repeated three times, and in
no case did the amine result. From one experiment the
culture solution was poured off through a filter, then dis-
tilled, in order to determine whether the amine remained
dissolved in the water in spite of the air current. The
result was the same — no amine was present.

Having proved that trimethylamine is not given off from
the cells when they are exposed to light, the next step was
to determine whether it is given off if the plant be kept in

darkness.

Accordingly the culture flask was covered by an opaque
screen and the gas collected as before. At the expiration
of twenty-four hours the plant was de~starched. This cul-
ture, starch free, remained in good condition, so far as the
morphology of the cells indicated, for seven days. At the
end of this time the absorption bulb was removed, its con-
tents made alkaline and distilled as before. The method
used for the separation and estimation of this amine was
that described by Fleck,* which is based upon the solubility
of trimethylamine sulphate in cold absolute alcohol, ammo-
nium sulphate being under such conditions perfectly insolu-
ble. Tlie amine is weighed in the form of its sulphate
(CH3)3N. H^SO,.

From the culture solution which had been maintained in
darkness, very appreciable quantities of the amine were
obtained, and the merest trace of ammonia.

It was mentioned above that trimethylamine began to
come off almost immediately upon removing the plant
from water, and that it was completely expelled by drying

* Jouru. Am. Chem. Soc, Vol. i8, p. 670.

Study of Spirogyra nitida.

219

at 100° C. After drying the plant, even alkalies caused no
further evolution of the gas. This fact was indicated by
the low percentage of nitrogen as determined by Kjeldahls
made with dry material, and others made with fresh
material. To obtain the entire nitrogen content it was
necessary to work with fresh threads which had been sim-
ply drained. The loss of nitrogen was not very great,
hence if it was due to proteid decomposition there was,
relatively, only a small amount of the substance which
yielded trimethylamine.

To determine whether the loss of nitrogen by drying
was due to its exit as trimethylamine only, the fresh plant
was drained, weighed, and introduced into a large flask
containing potassium hydrate. The substance was dis-
tilled until the gases were no longer absorbed, then the
amine was estimated as before. In this case, however,
considerable ammonia was produced.

From this analysis the plant was found 'to contain 0.45
per cent. (CHs^iN, which is equivalent to o.i per cent,
nitrogen. The nitrogen found in the fresh plant amounted
to 2.61 per cent. ; that in the dried material, 2.52 per cent.
Apparently, then, we have the loss of nitrogen perfectly
accounted for by the trimethylamine content.

If, as Loew believes, it is the lecithin only which pro-
duces trimethylamine, we should find some agreement
between the quantity of phosphorus organically combined,
and the quantity of trimethylamine evolved, since they
exist in lecithin in the proportion of i : i.

The phosphorus was determined by extracting the
material, dried at 30° C and powdered very fine, with
absolute ether until it failed to yield any green coloring mat-
ter to the solvent. Absolute alcohol was then allowed to
act until all the green substance had been removed. The
united extractions were evaporated in a platinum dish,
then ignited with calcium carbonate and ammonium chlor-
ide, the last traces of carbon being finally oxidized by the
aid of ammonium nitrate. The white residue consisted of
15

220

Penning ton — A Chcmico-PJiysiological

pliospliorus, as calcium phosphate, and alkalies, in the
form of sulphates, carbonates, and chlorides. It was taken
up with water and thoroughly boiled out, the phosphorus
being estimated in the insoluble portion by separating with
ammonium molybdate, and weighing as magnesium pyro-
phosphate.

Loew * states that mono-potassium phosphate is soluble
in absolute ether to the extent of 3 mg. in 100 cc. This
solubility introduces an error into the estimation of
lecithin as determined by the phosphorus content of the
ether-alcohol extract, provided this phosphate occurs in
the plant. Another, and more general opportunity for
error lies in the f^ict that in the estimation of lecithin
the material is preferably dried at very low temperatures.
Much water is in this way retained in the tissues,
and so carries out the soluble salts during the extrac-
tion. Both sodium and potassium were found in the
extract in quantities far beyond those required to combine
with the phosphoric acid. A trace of magnesium was also
detected. This was probably from magnesium sulphate,
which is somewhat soluble in absolute alcohol. Sulphuric
acid was present, perhaps arising from the oxidation^ of
organic sulphur, perhaps in combination with the alkali as

sulphate.

Lecithin contains 4 per cent, of phosphorus and 7.56 per
cent, of trimethylamine. Calculating the lecithin from
the phosphorus found, we have 0.19 per cent, contained in
the plant. If we make the calculation upon the basis of
trimethylamine, we have 5.97 per cent, or a quantity over
thirty times as great as that indicated by the phosphorus

content.

Many attempts were made to isolate crystals of lecithin

from the plant, but all were fruitless.

Though these experiments do not absolutely prove that
lecithin is entirely absent from the plant, they do indi-
cate that the evolution of trimethylamine is from another

* rfliiger's Arch., Vol. 79-

Study of Spirogyra nitida.

221

source, and that it probably plays an important role in the
synthesis of the plant proteid.

Chemical Changes in the Conjugating Cells.

Having obtained some insight into the chemical charac-
ter of the vegetating cell, it was considered advisable to
investigate the changes in composition, if such changes
take place, in the conjugating cell. Loew, in the paper
before cited, states that during conjugation a decrease in
the starch content takes place, with a corresponding rise in
the sugar content. This observation was confirmed during
the present investigation — not only does the glucose and
starch vary considerably, but the entire cell seems to be
fundamentally altered.

It was found impracticable to conduct the analysis ot
this material as was done in the case of the vegetating,
since the conjugating threads were inextricably mixed
with those not conjugating, and even in the individual
threads, conjugating cells were more frequently separated
by several vegetating cells. The work, therefore, was
accomplished by the aid of micro-chemical reactions, and
comparisons were made with the normal material. Obser-
vations were started when the tubes had just begun to push
out, and were continued until the entire act had been com-
pleted. The conjugating material was first noticed in the
pond in the latter part of April, 1896, after a week of very
warm weather had stimulated vegetation.

In the very early stages it was found that the chlorophyll
bands, alike in their matrix and ground substance, had
very materially altered. If the cells be treated with carbon
disulphide or chloroform, a disintegration of the band
results, because of the solvent action of the reagent upon
the green substance. The reagent first destroys the char-
acteristic contour and arrangement of the pyrenoid centre
and starch grains, and collects the green chlorophyll in
large drops at the edges of the bands. Ultimately, if the
action of the solvent be continued, the green solution is

0-7 -7

Pcfiuhi<rton — A Chemico-PJiysiological

diffused over the whole cell. Before this stage is reached,
the droplets at the edges of the bands of conjugating
material show more of a yellow tint than the corresponding
droplets in the vegetating bands. This difference in color
is plainly marked and indicates that we have here, in all
probability, a componnd which leans toward etiolin rather
than toward noruial chlorophyll. So marked is the differ-
ence, that the masses of conjugating threads have a
distinctly paler color than the others.

Not only is the green snbstance in an unstable condi-
tion ; the framework of the band seems to have nnder-
gone a change also. This framework in the conjugating
cells is entirely effaced by continued treatment with
carbon disulphide or chloroform, while in the vegeta-
ting cells the outline of the band can be plainly seen, even
after the chlorophyll has been completely dissolved out.
Just what the fundamental difference is has not been
ascertained. It is probably closely correlated with another
observation which was made at this time. A microscopic
examination of those cells which had distinctly developed
tubes, showed clear refractive bodies lying on, or closely
against, the chlorophyll bands. The diameter of snch a
droplet was sometimes as great as that of the band,
decreasing until it was represented by only a refractive
point. The substance was insoluble in cold and hot water,
but .soluble in the usual solvents for fats. Osmic acid
colored it deep brown or black. This test must of
course be made after removing the tannin. It was accom-
plished by immersing the threads for a moment in boiling
water, then exposing them for a short time to the vapor of

osmic acid.

Because of the resin found in the vegetating material,
and which was supposed to be contained in the chloro-
phyllaceous substance, these droplets were tested for such
compounds, following the idea that owing to the high state
of activity of the cell, and the marked chauges which had
taken place, it was possible that the resin had been excreted

Study of Spirogyra nitida.

223

froui the band. Such tests, however, yielded only negative
results. The substance corresponds in every respect with
a true fat. It persists in the cell through the primary
stages, and even after the contents have balled off, dark
spots are seen on treating with osmic acid. These show a
tendency to collect in the interior of the mass.

The relation of the tannin to the conjugating cell is very
complex. The quantity of tannin in the normal healthy
cells is extremely variable, and the different amounts ob-
served in the conjugatiug cells were so wide-spread aud so
unlooked for, that at first it seemed impossible to formulate
them. Close study of the plant at all stages indicates the
following history of the taunin content for the conjugating

cell.

It has been noticed that rapid division always precedes
conjugation. After this has occurred, just previous to con-
jugation, the quantity of tannin is very great. Iron salts
produce an inky-black color, while copper acetate gives
large masses of its characteristic brown precipitate within
the cell wall. This large amount of tannin does not per-
ceptibly alter during the growth of the first half of the
connecting tube. At about this stage it begins to diminish,
and by the time the tubes are in close proximity, the tannin
has almost entirely disappeared. Usually, until the tubes
actually meet, a slight reaction can be obtained /;/ the tube
itselj. Only the merest traces are to be found in cells
giving or receiving the contents of an opposite cell.
Neither was it possible to demonstrate that one cell of the
pair invariably contained more tannin than its associate,
though frequently this seemed to be the case.

In the two uniting threads it always happens that some
cells are unable to accomplish the act of conjugation,
though stored with the required food, and showing Uie
same preparatory peculiarities that the others show. This
may be due to a divergence in the direction of the two
threads, causing the tubes to be too short, or it may be due
to the fact that one thread is frequently composed of a large

224

Pennington — A Cheniico- Physiological

number of short, almost square cells, while the uniting
thread is formed of cells of the normal length. These
short cells may alternate with long ones in the same thread,
where they generally occur in chains of three or four. As
we find Spirogyra to be strictly monoecious there are, of
course, a number of cells which remain unfertilized.

These stimulated cells, which are unable to conjugate,
accumulate the large amount of tannin and retain it.
Indeed, proportionately to their size, the amount of tannin
stored by them is much greater than in the larger cells.
Frequently these cells form rudimentary or even fully de-
veloped tubes, which, however, never meet a corresponding
tube.

We may ask, does not this tannin behavior approximate
to that of malic acid in the fern? The tannin, though a
constant constituent of the plant, is formed in much larger
quantity immediately preceding conjugation. During the
first period of attraction between the cells, it maintains
this quantity, but as the cells are drawn closer and closer
together we find it disappearing and apparently by way of
the tube, since it is there that we detect its last traces.
The evidence indicates that it diff'uses through the cellulose
wall of the tube, and this may account for the fact that we
have so marked an alteration in the wall of the cell at this
period. (The modification of the cell wall will be again
mentioned.) It seems scarcely likely that the tannin
formed should act as a food supply.

The presence of a resin so closely united with the
chlorophyll would indicate a like origin for this substance.
Though the plant synthesis of both classes of compounds
is as yet unexplained, the indications are that the tannins
are intermediate between tlie oils and resins. For the
pines, at least, it has been established that the young cells
are filled with protopla.sm, which is partially replaced later
by tannin. Tlien oleo-resins appear, and increase at the
expense of the tannin and the protoplasm. According to
this view we would expect the resin content of the cells

Study of Spirogyra nitida.

225

jki.

to diminish, since the tannin formed seems to be diffused
into the water. Unfortunately this could not be deter-
mined ])y micro-chemical tests, since the resin is so closely
bound with the chlorophyll. But the oil drops exuded by
the bands probably represent the material which, when the
cells are not subjected to such a stimulus as conjugation
affords, yield with the tannin the resinous body.

The carbohydrates of the cell show decided modifications.
As has been already mentioned the glucose increases, ap-
parently at the expense of the starch. There is always,
however, a considerable amount of starch in the cell along
with the glucose.

The behavior of a vegetating cell with FehlingVs solution
has already been discussed, the cell in this case having
only a small amount of glucose. In the conjugating cells,
even before the tubes have reached any great size, the
quantity of glucose has .so increased that the cuprous oxide
deposit appears quite black under the micro.scope. When
the bands of the cell begin to change their normal position
the great mass of glucose is seen to be directly in front of
the tube, and to be pushing into it. This mass of glucose
persists in the cell, and when the giving cell transfers its
contents, the glucose in large quantity can still be detected.

No trace of cane sugar could be found in these cells.

Every one who has handled fresh Spirogyra is familiar
with the smoothness and apparent slimine.ss of the mass of
threads. This is readily explained by the large amount of
mucilage united with the cellulose. In the conjugating
material this sliminess has largely disappeared. The
threads are far more adherent than at other times, and show
a strong tendency to collect and hold the small particles of
disintegrated organic matter with which they come in con-
tact. There is here a modification of the cell wall, causing
an increase of the gum, hence rendering the cell more
adhesive.

The proteid of normal Spirogyra gives off very readily
trimethylamine. The conjugating material did not yield

'f

226

Pcmiington — A Chejuico-Physiological

Study of Spirogyra fiitiila.

227

the characteristic fishy odor to the hands of the operator,
nor to the vessels containing the air-dried substance, in any-
thing like tlie quantity usually observed. Some of the
threads, as free as possible from vegetating cells, were accord-
ingly heated with potassium hydrate. The odor of trinie-
thylaniine was plainly detected, but comparison with a like
quantity of vegetating threads, under the same conditions,
showed that the amine was relatively in very small amount,
and in all probability this was due entirely to the admix-
ture of vegetating cells, from which it was impossible to
free the material.

Morphological study has demonstrated that the cellulose
wall is intact over the ends of the connecting tubes until
after they have met. Watching closely the behavior of the
giving cell, it was noticed that after the mass of material
had begun to enter the tube a clear, refractive, colorless
layer of protoplasm was sent in advance. It was granular
in composition and very motile. The dividing wall of the
closely pressed together tubes having been reached, the
substance remained flattened against it for about one-half
minute. The wall then suddenly dissolved, and the trans-
fer of material from one cell to the other began.

This clear particle of protoplasmic material behaves like
a cellulose ferment, and doubtless contains such an enzyme.
Whatever be its nature it certainly is the active agent in
dissolving the septum between the cells. It can be easily
seen that a simple rupture does not take place, the passage
being clearly cut with sharp boundary lines, and of the
same size as the protoplasmic mass which pressed aj^ainst
the cell wall. The dissolved area may be in the centre of
the plate or at one side. It is, however, fully formed by
the substance above described, and does not increase in size
during the passage of the remainder of the cell contents.

Starch passes over as such. Treatment with iodine shows
blue masses in the act of transference. Many of these are
too large for single grains. It would seem, therefore, that
the pyrenoid, with its surrounding starch granules, is only

dislodged from its position in the chlorophyll band, and
not disorganized.

Chemical ChanCxES in the Celus of Spirogyra Due
TO THE Action of Monochromatic Light.

Since 1864, when Sachs performed his now classic ex-
periments on the relation of blue and yellow light to plant
assimilation, many investigators have not only repeated,
but also widely extended the work done by him.

To obtain the desired conditions, Sachs used double-
walled bell jars containing a saturated solution of potas-
sium bichromate for the yellow rays, and a solution of
ammoniacal copper sulphate for the blue. Others, work-
\\v^ in this field, have used frames of wood, provided with
glass top and sides, the glasses being of the color desired.
Frequently light-tight, blackened boxes, having one side
closed by a flat bottle filled with the colored solution, are
employed. Engelmann obtained monochronuitic light by
means of the micro-spectroscope, examining an algoid
thread in the various portions of the spectrum, and noting
the oxygen evolved. But by this method only small quan-
tities of material can be used, and the conditions, other
than light, are abnormal.

Preparatory to the work done in this laboratory, a num-
ber of colored glasses have been tested spectroscopically,
but not one has been found to be monochromatic, even in
the broadest sense of the term. Most of the aniline dyes
and various inorganic salts, yield impure spectra. A satu-
rated solution of potassium bichromate, which has been
accepted as affording yellow light, gives in addition all the
red and orange and part of the green. It will be shown
later that these individual colors have specific actions on
plant growth, hence the light obtained, after passing through
bichromate, gives to the plant the combined results of the
red, orange, yellow and part of the green. Ammoniacal
copper sulphate, if the solution be too dilute, lets through

1*

m

228

Pen nington — A CJicmico- Physiological

Study of Spirogyra 7iitida,

229

red and green, as well as bine, while if too concentrated
nearly all the light is cut out.

For accurate work, then, such conditions will not suffice.
Hence it became necessary to devise a means by which such
defects as are above mentioned should be remedied. After
many trials the method about to be outlined was adopted
as most satisfactory.

Description of Color Screens.

White earthenware bowls, having a diameter of 9^^
inches at the top, ^Y^ inches at the bottom, and 4>^ inches
in hei^rht, served to contain the material to be investi-
gated. Into these bowls were fitted dishes of white glass,
two inches in height, and having straight sides. Into
these the desired color solution was poured. Tin foil was
fitted over the edge of the glass dish and made to extend
some distance below the top of the bowl, all white light
being in this way excluded. The dish was covered by a
glass plate to prevent evaporation and the entrance of dust.
It is advisable to mark the original level of the liquid by a
diamond scratch, and see that this level is maintained by
adding the proper solvent from time to time. If two
layers of the colored liquids were necessary, a second glass
dish was placed on top of the first, and the joint made tight
as before.

By such a contrivance, all the light reaching the interior
of the bowl must pass through the entire depth of colored
medium, since the side light is excluded by tin foil. The
light intensity is increased by the reflection from the white
walls of the bowl.

The colored solutions were tested by a No. 7 Kriiss
spectroscope, and the limits of the colored band determined
as closely as possible. Monochromatic light is, in the strict
sense of the term, the light obtained from a single wave
length. But it is quite obvious that such light is not prac-
ticable for physiological experiments. Hence the term
"monochromatic light" has been adopted for a compara-

tively narrow band isolated from all the other visible rays
of the spectrum, and in which the human eye, aided by
the spectroscope, can distinguish only one color.

Such a band was isolated from each fundamental color of
the spectrum, namely, violet, blue, green, yellow, orange
and red. To produce these bands the following solutions
and mixtures were found to be the most satisfactory.

Yiolct.— Dissolve 128 grams of copper suli)hate in 1000
cc. of water. Add to this 0.08 grams of Hoffman's violet
(blueish) in 40 cc. strong alcohol. A layer one inch in
thickness gives a band extending from 449 ///^. to 417 /v^.

Blue.— Dissolve 35.5 grams of copper sulphate in 1000
cc. of water. 100 cc. of strong ammonia water renders
this a clear, deep blue. A layer of one inch gives from

476 /i/i. to 435 iLfi.

(;,.^.^.„._ Victoria green (3(C^H^NX1). 2ZnCl2 + H,0) was
taken as the basis for this color. In pure solution, how-
ever, it transmitted some of the blue and some red. Ani-
line yellow removed the former, and copper sulphate the
latter. The quantities used were: 0.32 gram Victoria
green in 1000 cc. of water, 0.0454 grams aniline yellow in
22.7 cc. alcohol, 16 grams of copper sulphate in 100 cc. of
water. The mixture is nsed in a layer of one inch, yield-
ing a band extending froui 535 ita. to 5 10 ////.

Yellow.— This band, being so §hort, and also the
brightest color of the spectrum, proved exceedingly diffi-
cult to obtain. Yellow dyes and .solutions were very imper-
fect, yielding red, orange and green invariably, and fre-
quently some blue.

It has been seen that copper sulphate cuts out the red
end of the spectrum, either wholly or in part, according to
its concentration. An aniline dye, "mandarin," having
the formnla C.eHuNASNa and showing a deep orange
when in aqueous solution, removes the more refrangible
rays. If, then, a ray of white light be allowed to pass
through both these liquids we have reuiaining only the
yellow rays, provided the solutions be of the proper con-

ii

230

Pennington —A Chcmico- Physiological

Study of Spirogyra niticia.

231

centration. This is obtained by dissolving 0.888 grains of
mandarin in 1000 cc of water. A layer of \^/k inches is
necessary. The copper snlphate is a solntion satnrated at
the ordinary temperatnres, and is nsed in a i5^ inch layer.
For this screen, therefore, we reqnire two glass dishes,
one above the other. The wave lengths obtained are from

603 Hfi. to 579 N'-

Orauf^e.— This screen was made mnch like the preced-
ing one. For the orange, however, a stronger solntion of
the mandarin orange is necessary, and a weaker copper
snlphate solntion, since the portion of the spectrnm desired
now inchides rays which are less refrangible than the yel-
low. These solutions are :

1. 3.333 grams mandarin in looo cc. of water. The

layer should be iYq inch.

2. 160 grams copper sulphate in 1000 cc. of water. The
layer should be one inch.

The light passing through these two solutions is entirely
absorbed with the exception of a narrow band which
extends from 634 fin. to 599 ftfJ..

Ijea.— The band in this case maybe obtained by the use
of an aqueous solution of a scarlet aniline dye. This dye
consists of a mixture of equal parts of (i) the soda salt
of xylidin-azo-B-naphthol-sulphonic acid, and (2) sodium
mono - sulphonate of amido - azo - benzol - azo - B - naph thol.
Technically, it is known as "Scarlet, [(SRRB) (BASF)],'>
and serves as a wool dye. Of this 12 grams should
be dissolved in 1000 cc. of water, and the glass dish
filled to the depth of one inch. The red band extends
from 7 1 8 fill, to 643 /i//.

It will be .seen that, with the exception of the yellow and
orange screens, a single layer suffices for the attainment of
the desired wave lengths. The yellow and orange, occupy-
ing so short a portion of the visible spectrum, have been
found to be unsatisfactory unless produced as above de-
scribed. The wave lengths as given are for the orange
from 634 /i/i. to 599 /V^-, and for the yellow from 603 fin. to

579 f'i^- There is here an overlapping of 4 f^W., but this is
too small a band to make any appreciable difference in the
physiological results. In the blue and violet, likewi.se, the
bands overlap somewhat. The blue extends from 476 ^/^.
to 435 ^^.^ while the violet is from 449 i^i"- ^0 4^7 i^^i ^
lap of 14 iiiJ-.

After the foregoing work had been completed it was
found that Landolt * had constructed what he terms a
*' color filter," for which he makes use of inorganic
solutions almost exclusively, and uses not le.ss than two
lavers for each colored band. For the yellow which
extends from 614 f^qi, to 574 |W|L^. he finds three layers are
necessary, viz.: i. Nickel sulphate; 2. Potassium bichro-
mate; 3. Potassium permanganate. Landolt does not give
an orange band distinct from red and yellow. He fills
tightly stoppered, flat, glass bottles with these solutions,
standing them closely against one another in metal frames.
While this form of apparatus might be constructed on .such
a scale that it would be available for biological work, it is
believed that the simpler scheme above given is moie read-
ily adapted to the conditions required for such investiga-
tions. The glass dishes and earthenware bowls may be
obtained in so many sizes, that large or small objects can be
accommodated with equal facility.

Experimental Study of Spirogyra nitida.

Conditions having been obtained, under which the plant
could be grown for a long period in light compo.sed of but
a small number of wave-lengths, the effect of these narrow
bands upon the growth and composition of the individual
cells remained to be determined.

To this end Spirogyra nitida was placed in the culture
bowls and its development watched. The most favorable
conditions for the growth of the plant were found to pre-
vail when a layer of clean, washed .sand covered the bottom
of the bowl, having planted in it about six actively grow-

* Sitzungsber. d. Kgl. Akad. d. Wissenscliafteiizu Berlin, (1894) Vol. 38.

»'

ll>

J

Poniiiigton — A Che ink o- Physiological

Study of Spirogyra nitida.

233

ing shoots of Elodca, This plant grows easily and rapidly,
and fnrnishes, therefore, a good snpply of oxygen. Whether
it be this fact or some deeper reason there can be no doubt
that Spirogyra grows more rapidly and makes healthier
threads when growing with an actively vegetating plant
than when growing alone.

The bowls so stocked were allowed to stand two days in
white light before being covered with colored screens in order
that possible injurious results from moving the Spirogyra
mio-ht be outo-rown. The bowls were then covered as de-
scribed in the previous section of this paper, and placed in
windows facing the south and in close proximity to eastern
windows, so that they secured the maximum amount of
direct sunliiiht. A crreenhouse was found to be too warm
to produce the best results. A temperature averaging about
22"^ C. gives the most desirable growth. Occasionally the
water was siphoned off and a fresh supply introduced in
the same manner.

When the experiments were begun the cells were in good
condition and well supplied with starch. While they con-
tinued a record of the temperature and amount of sunlight
was kept. The work was repeated many times, the condi-
tions, as far as possible, being strictly similar. They ex-
tended over an entire winter. The results as given here
set forth more especially the chemical, but also, to some
extent, the morphological changes which the cells undergo
when certain rays only are allowed to act.

The special changes induced by each colored band will
be given separately, the order being that of the colors in
the spectrum, beginning wnth the most refrangible rays.

Violet.— If the plant be exposed under the violet screen
to bright and continuous sunlight, the cells rapidly become
abnormal. The first noticeable change occurs in the chlor-
ophyll bands. These, at the end of twenty-four hours, are
seen to have a vacuolated appearance, and are of a pale green
color. At this time sugar, starch and tannin are still pres-
ent in the usual quantity. The protoplasmic pellicle ap-
pears normal.

By the close of the second day, provided the sunlight
continues, the chlorophyll bands are balled together more
or less tightly in the centre of the cell. This balling causes
the protoplasmic pellicle to recede from the cell wall, so
that it can be distinctly seen around the entire cell. When
such a cell is treated with a solution of iodine in potassium
iodide this pellicle takes on a violet-blue color. The fluid
in the cell-vacuole assumes the same tint, and the starch
grains, which are in considerable quantity, instead of show-
ing a pure deep blue, incline toward a violet. A test made
with Fehling's solution shows the presence of very little
sugar. The color which these cells yield with iodine, is
intermediate between the true starch blue and the violet of
erythrodextrine.

These peculiar phenomena indicate that we are dealing
with a cell in which elaboration and metabolism are .so re-
duced, at least in respect to .starch transformation, that only
a very small proportion becomes soluble, and that little is
far from a true sugar. Judging from the coloration pro-
duced by iodine it is just beyond the amylodextrine stage.
This product does not seem to be utilized by the cell, nor,
so far as could be determined, is it carried to the sugar con-
dition.

Ivudwig Klein * has proved that the conidial stalks of
Botryiis cinerca do not grow during the day because of the
presence of blue-violet rays. A paper by Ward f on the
reduced growth oi Bacillus anthracis in blue light, hints that
the reason may lie in some deep-seated chemical changes,
and the work of Macfarlane J on the sensitive i)lants, would
indicate that the chemical equilibrium of the cells is, for
the time being, much disturbed.

The absorption spectrum of chlorophyll shows a dark
band in the violet. Hence, according to the theory of
Timiriazefif and his school, work is done by these rays.
Engelmann, too, reports an evolution of oxygen in the

*Bot. Zeitung, 1885. \Vxoz. Roy. Soc, Vol. 53-

X Botanisches Centralblatt, Vol. 61.

I

234

Pennino;ton — A Chcmico- Physiological

Study of Spirogyra 7iitida.

235

W\

blue violet. It may be then, that all the available energy
furnished by the violet rays is only sufficient to produce
this imperfect hydrolysis. Or, judging from the experi-
mental work cited, the violet rays are so detrimental to the
cell that its normal activity is reduced almost to zero.
Under the usual conditions, that is the less refrangible rays
being present also, their activity may overcome the injuri-
ous effects of the violet rays. Botrytis cinerea, as shown
by Klein, is too sensitive to these rays to have their action
overcome by the presence of the red end of the spectrum,
and doubtless other plants would show the same phenom-
enon.

The third day generally finds the cells attacked by bac-
teria. The very rapid increase of these organisms causes
complete disintegration of the cell contents, with the simul-
taneous production of an inky-black compound which fills
the cell cavity. By the sixth day the cells are quite empty
save for the black substance.

The bacterial organisms never failed to make their
appearance under the violet screen, though the other cul-
tures were quite free from similar growths. They formed
an iridescent pellicle over the surface of the water, and
caused a foul odor, suggesting butyric acid.

Through the kindness of Dr. Alexander Abbott, Director
of the D^^epartment of Hygiene, this pellicle was investi-
gated and found to contain four spirilla which did not
liquify gelatine, and whose characters do not agree with
those of\ny known form. Unlike other organisms these
flourish under violet light Three of them are chromo-
genic, producing a pale yellowish-green color.

Having proven that the cell under violet light could not
transfornt its starch into sugar, the question arose. Can the
de-starched, but otherwise healthy, cell produce starch
when exposed to violet light only?

To answer this query material was rendered free from
starch and placed under the colored screen. At the end of
twenty-four hours the cells were still quite free from starch,

but apparently otherwise unchanged. Later the cell-con-
tents balled together as before, but now treatment with
iodine sohition did not cause any blue- violet color. The
organisms above mentioned promptly attacked the cell,
which, weakened from starvation, succumbed to their
ravages in about four days. During this time no starch
formation could be detected.

Blue. — The phenomena observed, when the plant was
grown under blue light, were exceedingly interesting and
suggestive, when compared with the results obtained with
bands from other parts of the spectrum.

For five days after submitting Spirogyra to the action of
blue rays the cells were, morphologically, in good con-
dition. The starch grains were slightly diminished in size.
Chemical examination showed that sugar was in nmch
smaller quantity than that normally present.

At this time the protoplasm was seen to take on a gran-
ular appearance, which grew con.stantly more marked.
By the seventh day the chlorophyll bands had lost their
characteristic disposition in the cell and had become some-
what balled, though not to such an extent as those under
violet light. The pyrenoid centres, with their surrounding
starch granules, were scattered promiscuously over the cell,
while here and there the dense granular protoplasm could
be seen between the green masses. Treatment with iodine
caused the starch granules to become deep bine, while
the dense protoplasm assumed a rich red-violet color.

When Fehling's solution is used we find the areas
corresponding to the red-stained protoplasm precipitating
cuprous oxide.

Exposure to blue light pushes metabolism a step ahead
of that obtained under violet light. There is sufficient
energy here to ])roduce a reducing carbohydrate, and one
also which can be assimilated by the cell. The blue rays,
like the violet, cannot produce starch. But starch hydro-
lysis is possible, though the process is so slow that the
granules originally present become from day to day almost
16

236

Pen nington — A Chemico-PJiysiological

imperceptibly smaller, until at the end of the fifth week
they have entirely disappeared from many of the cells, and
in the others are very few, and so small that strong mag-
nification is necessary in order to distinguish them. After
the starch is completely used up the cell contents rapidly
disappear, leaving finally only the empty cellulose walls.

Green.— Under the green screens the growth of the
threads and the general appearance of the cells is excellent.
So far as the carbohydrate of the plant is concerned we
have a condition closely approximating that of the normal.
The starch granules are somewhat larger than those formed
under white light and are very plentiful. Sugar can always
be detected and is in the usual quantity. Tannin, likewise,
can be observed in the cells, but never in large amount.

Though these green rays have so slight an action upon
the carbohydrates of the cell, the protoplasm is strongly
modified by their action. The quantity very materially
increases, so much indeed that the protoplasmic pellicle is
doubled in size. This pellicle is a denser mass than is
usually seen and is filled with small, dark granules.

The most striking phenomenon is the exaggerated
motility of this protoplasmic layer, soon after placing the
plant under the colored screen. In about three or four
days after the commencement of the experiment this
activity is noticed, and it prevails for five or six days.
While actively motile the protoplasm shows a strong ten-
dency to collect at the ends of the cell, either pressing
closely against the cell wall, or lying some distance back
from it. Frequently protoplasmic currents are seen flowing
across the cell. These may run quite straight, or more or
less diagonally, or they may take a zig-zag and tortuous
path. The large granules are carried along rapidly in this
moving mass, while some granules can at times be followed
entirely around and then across the cell.

Green light induces the formation of crystals of calcium
tartrate, and is unfavorable to the production of calcium
oxalate crystals. A very noticeable increase in the number

Study of Spirogyra nitida.

237

and size of the tartrate crystals takes place by the fourth
day, and after this time a rapid decrease in tlie number of
oxalate crystals is observed. Many break into small pieces.
The diameter of the tartrate crystals is occasionally quite
as great as that of the cell. This peculiar variation pre-
vailed vSo long as the culture was maintained.

No breaking down of the chlorophyll bands occurs, and
the healthy green color remains unaltered when the plant
is exposed to green light. But a flattening of the band,
with a corresponding increase in width occurs, and the
irregularity of the outline is so marked that the term
'* amoeboid" may well be applied to it. In some cases the
bands run together so completely that it is difficult to dis-
tinguish any boundary line between the individuals. In
these bands the pyrenoids are remarkably distinct. They
are very refractive and larger than usual.

Yellow. — A comparatively short exposure to yellow light,
that is two or three days, causes the green substance to become
much paler in color, and the bands show a slight tendency
to ball together. Under these conditions the starch rapidly
disappears, until by the close of the fifth day it is entirely
gone. The sugar, meanwhile, has increased far beyond the
normal amount. The cells, when treated with Fehling's
solution, become quite covered by the cuprous oxide result-
ing from the reaction. Although about five days are
required to free completely all the cells from starch, many
of them are emptied by the second day.

While this conversion of starch into sugar is taking
place, there is a sinuiltaneous and very marked growth in
the length of the cell. So rapidly does this elongation
take place that the chlorophyll bands are pulled out of their
spirals, and made to lie quite straight in the cell, which is
ultimately three times as long as when grown under white
light, though its diameter is not increased.

These cells were tested repeatedly for tannin. In the
great majority of them none was found, though occasionally
a trace could be seen. •

238

Pennington — A Chcniico- Physiological

Another marked and snrprising consequence of the
growth nnder the yellow screen, was the almost total elimi-
nation of the crystals normally present in sncli large nnm-
bers. These disintegrate, oxalate and tartrate alike, and
finally disappear entirely from the cell. The small frag-
ments into which the crystals break are at first angnlar,
then they become ronnded off and gradually smaller until
they are quite dissolved. What part these substances play
in the nutrition of the cell could not be determined, but
there seems every likelihood that a solution does take place.

The protoplasmic pellicle surrounding the cell wall be-
comes much reduced in size, and is quite clear and refrac-
tive. Few granules are to be seen in it. The nucleus,
too, with its suspending threads, is very clear. This pro-
nounced refractiveness is most striking, and gives to the cell
a starved appearance. Even the chlorophyll bands partake
of it, becoming narrow and of a clear pale green tint.

Treating the long sugar-filled cells with a very dihite
solution of iodine in potassium iodide caused a sudden rup-
ture of the cell, and the consequent scattering of the cell
contents. This rupture was liable to happen to any part
of the wall, the cell-plate being quite as frequently pushed
out — so breaking up the threads into short lengths, or even
single cells — as was the true wall torn. The force was
sufficient to break the protoplasmic pellicle, and even to
push the chlorophyll bands and the nucleus through the
aperture, throwing out the latter, and carrying it for some
distance.

We have here a large cell in which the sugar content,
and consequently the specific gravity of the cell sap, is
much increased. The osmotic pressure of a sugar solution
is comparatively small, while the pressure of the inorganic
salt, potassium iodide, is almost twice as great. The sud-
denly-increased endosmosis being greater than the proto-
plasmic utricle can withstand, it is violently ruptured,
tearing also the cellulose wall.

The culture could never be maintained for more than

Study of Spirogyra nitida.

239

seven days because of the attack of a fungus, which pene-
trated and completely demolished the cell contents. Soon
after the formation of a large quantity of sugar, and the
lengthening of the cell, the presence of this fungoid growth
could be detected. The conditions in the cell were favor-
able for its rapid growth, namely, a large quantity of soluble
nutritive substance, with a lowered vitality, which per-
mitted the inroads of organisms.

Some investigators, notably Pfeffer, consider the yellow
rays most active in causing assimilation. Judging from
the experiments just cited, the energy expended in the cell
is, indeed, great, but the ultimate growth is very abnormal,
there being a tendency toward an extensive production of
carbohydrate in a soluble condition. Is this energy such
that sugar can be produced in the cell without the appear-
ance, so far as micro-chemical tests can show, of starch ?
If so, starch-free cells placed for a time under yellow light
should have the sugar content increased. After this stage
their behaviour should correspond with that described in
the previous experiment.

This was found by experiment to be perfectly true. De-
starched threads placed under the yellow screen were exam-
ined after 24 hours, but no starch had been produced, and
the quantity of sugar was not excessive. The cells were
of the usual length. Forty-eight hours showed the cells
much lengthened, and the sugar content was also greater.
On the fourth day the cells had attained to their full length,
they contained much sugar, and many of them had been
attacked by the fungoid growth. In control experiments,
destarched material when placed in white light, showed
always a prompt starch formation, and made a good growth.
Oraiiffe.— Though following so closely upon one another
as do the orange and yellow rays, indeed slightly over-
lapping in the experiments herein described, the results of
their separate action are widely different. The narrowness
of these bands makes their dissimilar actions all the moYe
striking.

240 Pennington — A Chcmico-Physiological

Two or three days under the yellow screen sufficed, as
we have seen, to free the cells from starch, fill them with
sugar, and in many ways alter what we would consider to
be distinctive characters. On the contrary, five days
under the orange made no appreciable differences. The
plant's behavior w^as normal. By the eighth day the
number of crystals had greatly diminished. The oxalate
crystals suffered more than did the tartrate, the former
almost entirely disappearing from the cells, while the
latter, though few in number, were in fairly good condition.
Au examination for sugar showed this substance to be in
larger quantity than in the control threads grown under
white light. However, it was still far below the quantity
of sugar contaiued in the culture made in yellow light.

The protoplasmic pellicle was modified in that it had
lost its granular structure, and was clear and refractive,
though not so much so as in the case of the yellow. The
nucleus and its threads partook of this change also. The
nucleus stood out clearly in the cell, while the nncleolus
was very large and prominent. No change in shape or in
the relative positions of these bodies could be observed.

Tannin was always present, and in fairly large amount.
Its quantity varied without recognizable cause, just as it
does when the plant is exposed to white light.

Occasionally cells were seen which were longer than the
normal. These were not very frequent, and were not more
than twice the length of the other cells.

The cultures in orange light were sometimes kept under
observation for five weeks. At the end of this time they
were still healthy, showed very few crystals, contamed
starch and sugar and tannin, and had made a fairly good

growth.

Kert.— Under the red screen the plant approached still
closer to the normal. Assimilation went on rapidly, so
rapidly in fact, that the cells became gorged with starch.
At the end of five or six days so many granules, and such
large ones, surrounded the pyrenoids that the bands were

Study of Spirogyra nitida.

241

crowded closely together, almost or quite hiding the
nucleus from view. This piling up of starch was not at
the expense of sugar, the presence of which was always
demonstrable, and in quantities agreeing with the normal.

The tannin formation was excessive. Copper acetate
threw down a heavy precipitate in the cells, and ferric
chloride colored them inky black.

The crystals, on the other hand, were not so many nor
so large as when grown in white light. Eight days caused
a breaking down of oxalate, while by the tenth day very
few of these remained. Calcium tartrate crystals were not
many, but here and there one did survive. These, when
solution took place, had the arms dissolved first, leaving
behind a square plate which was at first mistaken for
another variety of crystal. In time the.se likewi.se dis-
appeared.

The protoplasm seemed to be normal in every way. The
nucleus, when it could be di.stingni.shed in the dense mass
of starch-laden chlorophyll bands, was very refractive, but
contained dark particles as usual. The nucleolus was also
very refractive.

These cultures, like the orange, can be kept for an in-
definite period in good condition. The growth is even more
rapid than in zvhite light.

Growth Under Colored Glass.
Having carefully cultivated and watched the develop-
ment of the plant under pure light, some experiments car-
ried on under colored glasses may well be compared with

them.

The apparatus was the same as that used by Macfarlane
in his ^' color screen^' experiments on the sensitive plants,
and consisted of a .square wooden frame, into the sides and
top of which were fitted glass plates of the desired color.
The healthy Spirogyra was grown in white glass jars hav-
ing a capacity of about one litre. These were placed

* Bot. Central. VoL 61.

242

Pennington — A Cheuiico-Physiological

inside the box, and the whole apparatns exposed to bright
light, either in a green-house, or, as these experiments
were made in the early autumn, in a sheltered place in the
Botanic Garden.

Three screens were used, viz.: blue, green and. yellow-
orange. These glasses, tested by the spectroscope, were
found to allow the following rays to pass:

/. Blue glass :

Red, from 738 /i//. to 703 aft.

Yellow-green, 566 jlji. to 552 fffi.

Green blue, blue and violet, 517 ////. to 408 nfx.

2. Green glass :

Orange, yellow, green and a little blue, from 629 ///i.
to 458 itti.

J . Yellozv-ora nge glass :

Red, orange, yellow, green, aud green-blue, from
687 \L\Ji. to 464 \x[i.

When grown under such conditions, it is obvious that
the plant is acted upon by a considerable part of the spec-
trum, since these glasses divide it roughly into three por-
tions with much overlapping in the less refrangible rays.
Hence the changes observed in the plant are due to a com-
bination of forces, these forces being represented by the
rays which pass.

Blue Glass. — The blue light, having a small band of red,
some yellow-green, and a little blue-green, in addition to all
of the blue and the visible violet rays, acts upon the plant in
a much modified form, the red and green tending to over-
come the evil effects produced by the blue and violet. As
a result of this the plant made a nnich slower growth than
under normal conditions, but a growth far beyond that made
under pure blue.

Starch and sugar diminished in quantity, and frequently
the starch disappeared entirely, only to reappear in a short
time. The tannin fell considerably below the normal
amount, but was never completely lost. The chlorophyll

Stndy of Spirogyra nitida.

243

maintained its deep green color, and was perfectly healthy.
Morphologically, it was modified in that it tended to form
dense aggregations over the nucleus, giving the appearance
of a protective covering. In this mass it was difficult to
distinguish the individual bands. The cells were much
shortened — being not more than half the usual length, and
the spiral bands were wrapped very closely around them.

The threads grew straight upwards, the folding upon
itself which Spirogyra is so apt to show, and which Mann*
ascribes to heliotropic action, being entirely absent. Since
the blue-violet rays are those which, more than any others,
induce heliotropic curvature, this fact is noteworthy.
Further mention will be made of it when the growth under
the yellow-orange is discussed.

Greeii j-iiiss. — Under green glass the results were more
abnormal. Orange, yellow, green and a little blue cannot,
apparently, compensate for the absence of the strong red
rays. In twenty-four hours from the time of the begin-
ning of the experiment, the chlorophyll bands had begun
to break down, while at the end of forty -eight hours many
cells had the pyrenoids connected by only a fine green
filament. The spiral position of the bands was quite lost,
and the remains lay hap-hazard in the cell. The condition
of the cells composing a single thread varied nuich. Some
were quite free from starch, others had unusually large
starch grains around clear large pyrenoids. In those cells
which were free from starch iodine frequently, though not
invariably, produced a violet coloration, closely approach-
ing the color obtained in like manner when the plant was
grown under violet light.

Another phenomenon, which was observed only a few
times, was the presence of oil droplets, on or close to the
chlorophyll. These were blackened by osmic acid, were
soluble in the usual solvents for fats, and behaved in every
respect like the droplets observed in the plant when pre-

* Proc. Bot. Soc. Edin. Vol. 18.

244

Pen nington — A Chemico- Physiological

paring for conjugation. Actual conjugation was not, how-
ever, attempted.

The quantity of sugar and tannin is very small, the
latter constituent occasionally entirely disappearing from
some of the cells. Crystals of both kinds could be detected.

Yollow-orauj^e ji^lass. — This glass, while it permitted
the passage of all the orange and part of the red, gave
results which were strikingly like those obtained under
pure yellow, though the time required to produce them
was nnich lengthened.

The starch disappeared about the ninth day, and the cells
were then filled with sugar and were very long. The
tannin, unlike the pure yellow culture was in large quantity.
No crystals could be seen in the long cells, and the chloro-
phyll was in rather a bleached condition. The bands here
lie straight in the cells just as do those in yellow light.
Treatment with iodine in potassium iodide causes the same
rupture of the cell wall.

It was stated that under the blue glass no bending of the
filaments took place, even though these rays are so posi-
tively heliotropic in their action. Under the yellow-orange
screen the threads were bent sharply upon themselves, as
many as '^\^ distinct folds being assumed. Since the rays
which this glass transmits are supposed to be almost with-
out heliotropic action it may well be asked, Are not these
folds due simply to the weight of the long weak threads?
The plant under the blue screen was made up of short,
strong cells, and the filaments were short, as compared with
the long filaments under the yellow. This may account
for the fact that no bending occurred under the blue.

Behavior of the Cells in Darkness.

Some endeavors to de-starch Spirogyra nitida by simply
placing it in the dark produced rather peculiar results,
which led to a more careful study of the behavior of the
plant when all light was excluded. It was found in this
preliminary work, that the threads did not de-starch readily.

Study of Spirogyra niti<Ia.

245

even when, so far as could be determined, the plant grew
in perfect darkness. A quantity of fresh threads placed in
a beaker glass, and covered with tap water, the whole
being maintained in darkness, showed in the cells very
appreciable amounts of starch even after two weeks had
elapsed. The condition of the cells at the expiration
of this time was not very good. Here and there the bands
had lost their normal positions, and in .some a distinct
breaking down of the cell could be seen. By the time the
starch had been converted into .soluble products the cell
was so altered in other respects that it was quite useless
for physiological experimentation.

In tiiese darkened threads a solution of the cry.^tals was
very apparent. They broke down into small fragments, and
were finally dis.solved as under the action of yellow light.
This solution became more rapid as the amount of starch
grew le.ss, but in no case were the crystals wholly used up.

Since starch-free material was much needed for experi-
mental work it became necessary to find a method by which
this end could be gained, and still have comparatively
healthy cells. It was thought that running water might
perhaps prevent decomposition and enable the cells to
survive until all the starch had been metabolized. Accord-
ingly a small stream from a laboratory tap was led into a
jar containing the plant, and the whole covered with a tight
box. At the end of twenty-four hours scarcely a trace of
starch was found in the cells which were otherwise perfectly
normal. Thirty hours will sufftce to remove all the starch
from spirogyra nitida, while the more slender species

react even sooner.

To determine whether this result was due to the plenti-
ful supply of inorganic salts furnished by the running
water, a quantity of the plant was placed in distilled water
to which a culture mixture was added. The .starch was
removed very slowly, and the cells behaved as they did in
a limited .supply of tap water. The inorganic salts, then,
are not responsible for this behavior.

246

Pefinin^^toti — A Chcniico-Physiological

Fresh Spirojryra was next placed in a flask fitted with a
doubly perforated cork carrying glass tubes arranged as in
an ordinary wash bottle. The short tube was connected
with a suction pump, and by means of the long tube a cur-
rent of air was drawn through the flask. The air must not be
drawn through with such force that it churns the Spirogyra
in its passage. If the flask is kept quite dark, the plant
under such treatment behaves precisely as it does when
running water is supplied to it ; the starch is rapidly meta-
bolized, and the cells soon become starch free.

P'roni the above facts we are led to believe that in this
starch transformation the gases of the air play an import-
ant part. Since all experiments have shown that carbon
dioxide is acted upon by chlorophyll only when light is
present, we must look to the oxygen as the active agent.
This element seemed, by its presence, to so stimulate cer-
tain functions of the cell that it was enabled to convert
starch into soluble products.

An explanation of this peculiar behavior lies in the fact
that the simpler nitrogenous substances, such as amides,
are built up in the dark, and that these substances there
unite with the carbohydrate present, to form the more com-
plex compounds or proteids.

While the absorption of free oxygen by the plant is
generally followed by a breaking down of the organic sub-
stances, just as in animal respiration, it is probable that
being deprived of the nascent oxygen, which results from
the decomposition of carbon dioxide in sunlight, and also
from the free oxygen of the air or water, the plant is un-
able to combine the first-formed nitrogenous compounds
with the carbohydrate already present. Or, since we know
that in time the starch is dissolved, this synthesis takes
place so slowly that the equilibrium of the cell is destroyed
before all the carbohydrate is used by the nitrogen com-
pound.

The nature of the hydrolytic ferment in Spirogyra is as
yet unknown. But from the study of this class of ferments

Study of Spirogyra nitida.

247

we may readily suppose that free oxygen, in greater quan-
tity than is supplied by still water, is necessary in order
that it may produce hydrolyzed products from starch.

The Action of Light on Diastase.

A consideration of the foregoing results led to an inves-
tigation of the behavior of unorganized hydrolytic fer-
ments in rays of varying refrangibility. The information
which we possess upon this point is very limited, and its
relation to plant metabolism is quite untouched.

St. Victor and Corvisart * state that sugar is formed
from starch more rapidly in light than in darkness.
Green t exposed solutions of diastase to the action of
electric light and of sunlight. The diastase so treated
was then allowed to act upon starch paste, when he found
that light exercised a destructive influence upon the fer-
ment, this influence being more marked at the violet end
of the spectrum. But the light with which he experi-
mented was polychromatic, and if diastase is affected by
light rays, it is probable that the portions of the spectrum
affecting plant growth will also influence the rapidity of

diastatic action. J

The relation of light rays to the ferment is, as deter-
mined by the previously described color screens, very
marked, and in perfect accord with the changes observed in
Spirogyra when grown under corresponding conditions.
An account of the experiments whereby this conclusion
has been reached, will make clear the above statement.

Potato starch was purified by washing with very dilute
potassium hydrate, then with i per cent, hydrochloric

* Coinptes Rend., Vol. 49- t Annal.s of Botany, Vol. 8.

X After this investigation had been passed to press another article by
J. R. Green appeared in the Proceedings of the Royal Society, Vol. 61,
p. 25. Many of his results agree with those above given, but some are
strikingly different. In the above investigation narrower, and therefore
purer colored bands were used than those from which Green drew his
conclusions. It is noticeable that he fails to give the action of the yellow
rays as distinct from the orange.

I

248 Pemiington — A CJicmico-Fhysiological

acid, and finally with distilled water nntil the washings
did not react with silver nitrate. It was then dried in the
air, and was preserved for nse in tightly-stoppered glass

bottles.

A preparation of diastase was obtained from Messrs.
Bullock & Crenshaw, Philadelphia. This was found to be
only partially soluble in water, yielding when filtered a
clear liquid of a pale straw yellow.

The starch paste, made by pouring starch, mixed with
cold water, into boiling water, contained i gram in 100 cc.
It was freshly prepared each day. The strength of the
diastatic solution soon changed, hence it was necessary

to prepare this daily.

The tests were made by placing the beaker containing
the starch and diastase in a white bowl, and covering
with a colored screen. These bowls, when a temper-
ature of 45° to 50° C. was desired, were partially filled
with heated water. The progress of the reaction was
noted by removing from time to time a drop of the liquid,
and applying the iodine test. The control experiments
under white light were conducted in like manner, except
that the glass screens contained only distilled water. The
work was done in direct sunlight. As it was found that
the light intensity caused a very material variation in the
time required for this hydration, the experiments were
made only on very bright days, and then only in the mid-
dle of the day when the sun's rays were most direct.
Diastase solutions of varying strength were used, though
if too dilute the end reaction was hard to determine.

The experiments were carried on from the middle of
February to the end of April.

For the six screeus the following ratios were found, the
time in white light being taken as unity :

White .
Darkness
Violet .
Blue . .

1. 00
1.28

2.CK>
1. 21

Green 'H

Yellow 0-76

Orange 0-^5

Red

Study of Spirogyra nitida.

249

A few examples, taken at random from the series of ex-
periments, indicate the time in minutes. Each experiment
conducted in colored light was checked by a similar test in
white light, the time required in both cases being here
given :

Darknkss.

Exp. I.
White, 10 minutes.
Darkness, 13

< <

ViOIvET.

Exp. 1.
White, 9 minutes.
Violet, 18

Exp. 2.
White, 10 minutes.
Darkness, 12.75 minutes.

Exp. 2.

((

White, 22 minutes.
Violet, 45

i(

B1.UE.

Exp. r.
White, II minutes.
Blue, 15 '•

EXF. 1.

White, 14 minutes.
Green, 15

Exp. I.
White, 12 minutes.
Yellow, 9

Exp. I.
White, 15 minutes.
Orange, 13 **

Exp. I.
White, 9 minutes.
Red, 9

Exp. 2.
White, 9 minutes.
Blue, 12 '*

Green.

Exp. 2.

White, 10 minutes.
Green, 12

<(

YELI.OW.

Exp. 2.
White, 13 minutes.
Yellow, 9.5

( i

Orange.

Red.

(<

Exp. 2.
White, 15 minutes.
Orange, 13 "

Exp 2.
White, II minutes.
Red, 1 1

<(

1. 00

The light rays, from the above data, attain their maxi-
nnim for diastase in the yellow, and in darkness the action
is slower than in light. Under orange the change from
starch into sugar is slightly slower than with yellow, while
red light requires the same length of time as white. Pro-
gressing toward the more refrangible end of the spectrum
we find the green band a little slower than white light, and

250

Pennington — A Chcmico -Physiological

Study of Spirogyra nitida.

251

the blue still more unfavorable to sugar production, while
the violet rays are only one-half as rapid as the red.

In diffuse daylioht the action of these rays is much less
marked, the ratio between yellow and white bein<^ as i : 0.92.
As diffuse yellow light exercises less influences for good
than does bright yellow light, so is the destructiveness of
the blue and violet lessened by diffuse light.

In the experiments made by Green, the enzyme was ex-
posed first to the influence of light, then mixed with starch
paste and its activity noted. As the conditions in these
experiments were slightly different, starch and diastase
being mixed first, then exposed to light, it was deemed ad-
visable to expose diastase in solution to light, as did Green^
then test its activity under the colored screens. Such tests
showed that the solution became rapidly weaker in light,
but in colored light maintained its relative strength.

This behavior indicates a protective influence of starch
on diastase when exposed to light. It is even more than
that, since in the one case light accelerates, and in the
other retards its hydrolytic activity.

Interpretation.

The relation of light to plant growth being one of the
most interesting and wide-spread of all the many questions
with which the plant physiologist has to deal, it has nat-
urally received much attention. And while each portion
of the spectrum has been considered by the various workers,
the greatest amount of interest centres in the red-orange
and yellow, because of the theories which have been built
upon the study of the behavior of these rays in regard to
plant assimilation. It is the portion of the spectrum con-
taining the less refrangible rays which, from Sach's time
onward, has been recognized as best promoting plant growth.
According to Lommel, Miiller and Timiriazeff this is be-
cause the rays, absorbed by chlorophyll, have their energy
converted into some other form of energy, which is then
capable of bringing about the decomposition of carbon

dioxide and water and causing their subsequent combina-
tion to form organic products. These writers argue, that
only the rays which are absorbed are capable of causing
chemical changes in the plant, and therefore the absorption
bands of the chlorophyll spectrum correspond to the regions
in which growth takes place.

Pringsheim, taking exactly the opposite view, considers
that the ra}s absorbed are those whicli would be prejudicial
to the growth of the plant, and that they are in this way
removed. Pfeffer believes that the yellow rays — and there
is no absorption of these — are those most active in assimi-
lation. But, the conditions under which he worked being
inexact, too much reliance cannot be placed upon his de-
ductions.

Engelmann's experiments tend to confirm the work of
the school of Timiriazeff, namely, that the region of most
active assimilation is situated at the junction of the red and
orange, which region is also that of the densest absorption
band in the chlorophyll spectrum.

The theories regarding the action of light have been
based upon the rate of decomposition of carbon dioxide
and upon the corresponding evolution of oxygen as deter-
mined for the different parts of the spectrum. The results
obtained with Spirogyra show the general condition of the
cells, and the changes in quantity and in kind of the sub-
stances produced, by the growth of the plant.

The spectrum of living chlorophyll as determined by
Mann* gives four absorption bands. The first two, which
extend from 678 /i/^ to 662 nn, and from 654 /i// to 638 /i/i, are
both included in the band furnished by the red screen, which
lies between 718 ///^ and 629 /i/i. What are connnonly
known as the third and fourth bands of Kraus, Mann does
not find in Spirogyra, but the fifth he places between 531
fui and 459 liii. This band is partly covered by the blue
screen (from 476 /i/i to 435 /i/i.) and a very small portion is
covered by the green (535 ///^ to 510 /i/i.) The sixth band
*Proc. Bot. Soc. of Edinburgh, Vol. 18, (1890.)
17

^

252 Pennington^A Chcmico-lhysiological

he finds to extend from 459 N^ to 445 N'-^ with the centre
at 450 iifi. The violet screen, from 449 W to 4^7 /V^^ i^^"
chides part of this band, which Mann says is not a well-
defined one, and also inclndes the absorption band which
we invariably find at the extreme end of the spectrnm.

Under the orange screen we have no absorption band ;
the yellow, likewise, is not absorbed, and nnder the green
screen a few, only, of the rays coincide with those which
are absorbed. Bnt plants grown nnder these screens show
striking variations from one another and from the normal,
and the orange, where, according to the absorption theory,
growth shonld not occur, gave, next to red, a growth more
nearly approximating the normal than did any other.

Such facts cannot be reconciled with either of the theories
above given, since it is apparently not so much the absorp-
tion of the rays, as the amount of energy which they pos-
sess which influences assimilation for either good or evil.

According to Langley the greatest amount of energy, as
measured by the bolometer,, is furnished by the orange rays.
Following this statement, growth should here be at its
maximum, but the foregoing experiments with Spirogyra
gave a more normal growth under red, in which the energy

is somewhat lower.

The very peculiar behavior under the yellow screen I am
inclined to attribute to the action of these rays on the
hydrolysis of starch. This is converted into soluble carbo-
hydrate so rapidly that the nitrogenous products, which
normally are in sufficient quantity to combine with it to
form higher compounds, are in this case unable to do so.
The sugar, therefore, accumulates in the cell, upsetting the
perfect balance of power which should prevail. The in-
roads of the fungus cannot then be withstood, and the

plant dies. ^-

If we accept the red, as the region of greatest growth, or
the junction of red and orange, we must conclude that the
maximum hydrolytic activity is not conducive to the wel-
fare of the plant ; and we must also, from the experiments

Study of Spirogyra nitida.

253

upon the action of diastase on starch paste in the different
light rays, place the two centres of activity, the assimilative
and the hydrolytic, some distance apart. The activity of
the latter decreases, from the yellow toward both the more
and the less refrangible end of the spectrum, and it is to be
noted, that where the diastatic activity approximates that in
white light the plant makes a good growth, if not a per-
fectly normal one. This condition is seen under the red
screen, which in its action on diastase is exactly equivalent
to white light.

The green, standing to white light, as measured by
diastatic activity, in the relation 1:1.14 has its starch
hydrated rather more slowly than in white light, yet the
decrease is not sufficient to stop nor to materially injure
the growth of the plant.

The strange appearance and unusual activity of the pro-
toplasm produced under this screen suggests that the cell
is in a condition exactly the reverse of that induced by
yellow light. There, soluble carbohydrate predominated
over nitrogenous compounds ; here, the speed of production
of soluble carbohydrate is slightly lessened. The nitrogen
assimilation is then, probably, in excess, and it may be
that the proteid formed contains a larger proportion of
nitrogen than that formed under the usual conditions.

Mendeleeff, in his '' Principles of Chemistry," has called
attention to the fact that nitrogen content and protoplasmic
activity seem to go hand in hand through all forms of life.
Animals, whose tissues are so largely nitrogenous, possess
this activity in the highest degree. The higher plants,
with their great quantity of stable carbohydrate, have it
only feebly. But, on the other hand, in lower plant forms,
as for example in the zoospores, where the rapidity of
movement is quite comparable with that of animal life,
the nitrogen content is high, and remains so until the
active period is passed. When the spore fixes down it
develops a cellulose wall, loses its high percentage of nitro-
gen, and becomes in all respects a true plant.

254 Pennington — A Chemico- Physiological

If protoplasmic activity depends upon tlie multiplication
of unstable nitrogen atoms the increased movement of the
protoplasmic pellicle may be due to the synthesis of such a
product under these abnormal conditions.

By the time we reach the blue rays the diastatic activity
has greatly diminished. Yet enough energy still remains
to convert the starch grains, little by little, into soluble
products, though these seem to differ chemically from those
compounds generally obtained by diastatic action. The
cell energy, too, has fallen so low that carbon dioxide and
water are no longer decomposed to yield starch as the first
visible product. When the supply with which the cell
was furnished is used up, its resources are at an end and
death ensues. Under blue light we may say that meta-
bolism is at a minimum, while carbon dioxide assimilation,
so far as we can determine, has ceased. The density of the
protoplasm shows that a very pronounced alteration has
taken place in its composition, and this is shown likewise
by the color which iodine imparts to it. This color would
indicate that even though the starch is slowly hydrated the
energy is not sufficient to carry it to a true sugar, and can-
not link together nitrogenous and carbohydrate substances
into the proteid found in the plant under white light.

The violet screen, which furnishes the more refrangible
rays, does not seem capable of changing the starch which
is already in the cell, into any product which can be used
in plant metabolism, though it is indeed somewhat altered,
since we find a blue color diffused through the cell after
treatment with iodine. Probably both this fact and that of
the greatly reduced cell energy combine to produce a con-
dition in which proteid cannot be formed in Spirogyra.
This same energy, on the other hand, is not only sufficient
for, but favorable to, the growth of certain bacterial organ-
isms. The deleterious effects of this screen are seen also
in the action of diastase on starch where the time required
for hydration is twice as long as the time required when
white light acts.

Study of Spirogyra nitida.

255

Not only do we find the fundamental constituents of the
cell undergoing modifications, but other products, such as
the cell crystals, are much altered by different light rays.
While there is, in all probability, a direct connection be-
tween the formation or disappearance of these bodies and
the general activity of the cell, I have not been able to for-
mulate it from the above experiments.

In yellow light the obliteration of the crystals is almost
complete, while it is less marked in orange and red. When
the plant is kept in darkness without a sufficient oxygen
supply the .same breaking down occurs. As etiolation
implies also starvation, it might be supposed that the solu-
tion of the crystals provided nutritive material. Yet in
orange and red light there is a diminution in the number
of cry.stals, though the cells are well nourished.

Green light tends to promote the production of tartrate
crystals, increasing them both in number and size, though
here also oxalate crystals were used up. Weber* finds the
greatest absorption of ash constituents to take place when
the plant is in white light; it is somewhat Icssni yellow,
then follow red, blue and violet, decreasing in activity
as the refrangibility of the rays, but green gives the
lowest ash content. These results were based upon the
action of the light which passed through colored glasses,
none of which were monochromatic. One would infer
from the large crystals of the green culture that an exten-
sive absorption of inorganic constituents was here taking
place, while results from the yellow, showed either very
few or no crystals whatever, leading to the opposite view.

A number of observations have been made which show a
similarity between yellow light and darkness. Sachs and
Kraus have found that etiolation tends to produce long in-
ternodes ; and rays of low refrangibility do the same. Cor-
responding to the internodes we have the individual cells
of the Spirogyra thread, and, as we have seen, these under
yellow light attain to three times their normal length. It

^Landw. Versuchstat., Vol. 18 (1875).

256

Pennincrton — A Chemico-Physiological

is interesting to note in connection with tlie statements
made by these observers that, in the experiments made on
Spiroi^yra, it was the yellow only which cansed a marked
lengthening when the red, orange and yellow rays were
separated, thongh nnder the orange-yellow glass the action
of the yellow rays was not snfficiently retarded by the
presence of the others to prevent the cells elongating.

Ranwenhoff * states that less tannin is found in etiolated
leaves and plants than in green ones. Here, too, the
behavior of the yellow culture accords ; tannin is entirely

absent.

Looking at the question of the action of light on plant
growth, from the chemical changes noticed in Spirogyra,
we see that assimilation, though in a more or less modified
form, can occur when none of the light rays are absorbed
by the green coloring matter. Also, that such absorption
may or may not be accompanied by assimilation.

The blue and violet rays seem to act as checks to the
great activity of the yellow, and perhaps to the orange,
preventing the too rapid transformation of starch into sugar,
upon which the rapid growth in length apparently depends.

The Action of Palladious Chloride on the

Living Cell.

While studying Spirogyra micro-chemically palladious
chloride was used as a reagent. The action of this sub-
stance upon the living cell demonstrated so clearly certain
morphological features, that a more careful examination
of its behavior was made. It was found that a solution
containing 0.1478 grams palladious chloride in 100 cc. dis-
tilled water fixed the cell instantly, and with the minimum
amount of distortion. The protoplasm was slightly browned,
making the nucleus and its threads the more easily seen.
If the solution be diluted until it contains only 0.001478
grams palladious chloride in 100 cc.,the morphological

* Ann. d. Sci. Nat., Ser. 6, t. v., (1877).

Study of Spirogyra nitida.

257

changes are very gradual and show plainly the structure of
the nucleus and its contents.

This weak solution of palladious chloride was run grad-
ually under the cover glass, its action on the cell being
carefully watched. The first visible result was a sharp
rounding off of the nucleus from the cell, though the nuclear
threads were not broken. By this rounding oft the nuclear
membrane became plainly visible, and appeared to be a
clear, homogeneous, doubly refractive bounding layer.
Even by aid of an oil-immersion lens it was impossible to
detect any structure in it. The nuclear substance mean-
while assumed a brownish appearance, which was probably
due to a deposit of metallic palladium, and its granular
structure became very marked. In many nuclei there
appeared to be a mass of tangled threads in the meshes of
which lay the more fluid substance. The threads suspend-
ing the nucleus could be distinctly traced through the
nuclear membrane and into the granular mass of the

nucleus.

The nucleolus showed a dark bounding layer of double
contour. Its substance remained homogeneous, but became
nnich more refractive than is usual. The dark layer is
undoubtedly a true membrane dividing the nucleolus from
the nucleus, as stated first by Macfarlane,* and confirmed by
subsequent investigators. It was distinctly visible and in-
variably present.

These changes took place in from 8 to 10 minutes. A
continuation of the action of the very dilute palladious
chloride caused the suspending nuclear threads to rupture,
and in so doing generally displaced the nucleus. Where
this happened the nucleus was observed to be incased in
a protoplasmic pellicle which was di.stended until it was
nuich larger than the body which it enclosed. The sus-
pending threads were seen to penetrate this pellicle.

A further treatment of the cell with this reagent caused, at
the expiration of half an hour, a complete balling together

* Trans. Bot. vSoc Edin. Vol. 14.

258

Pennington — A Chemico- Physiological

of the chlorophyll bands, the inner layer of protoplasm
and the nucleus. The outer protoplasmic layer, however,
was unaffected and still adhered closely to the wall of the
cell. The balled mass was generally pulled to one end or
close to the sides of the cell. The many crystals were in
this way left quite free in the unoccupied space of the cell,
and could be easily studied. None were seen in the balled
material, hence it was inferred that these bodies lie between
the two protoplasmic pellicles.

Summary.

1. An analysis of Spirogyra nitida shows that the chlo-
rine and sodium content of the ash is comparable with the
chlorine and sodium content of the salt-water alg.-E. The
dry matter yields all the usual organic plant constituents,
tannin being in specially large quantity.

Micro- chemical analysis reveals crystals of calcium tar-
trate, as well as those of calcium oxalate, side by side in

the cell.

2. The trimethylamine, which is readily evolved from
5. nitida^ is apparently closely connected with, and aids in
the formation of a proteid body. It is not evolved when
the plant is exposed to light, but is detected in darkened
material. This amine accumulates in the de-starched cell,
but does not seem to result from the decomposition of

lecithin.

3. Conjugating cells show a chemical composition which,
in almost every essential differs widely from that of the
vegetating cell. The behavior of the tannin, and its
marked increase in quantity is now striking ; as is also the
presence of oil droplets, which are found free in the cells.

4. Under colored screens, furnishing light of definite
quality, the chemical composition of S. nitida differed ac-
cording to the light rays which it received.

[a) Violet rays prevented almost wholly the hydration of
starch, and soon killed the plant.

{6) Blue rays gave an imperfect hydration, but sufficient

Study of Spirogyra nitida.

259

to preserve life for some time. Starch was not formed in

the cells.

{c) Green rays caused an active assimilation with a con-
tinuous growth and unusual protoplasmic motility. The
protoplasm was also in increased quantity. Green light
favored the production of crystals, particularly of calcium

tartrate.

(d) Yellow rays caused elongation of the cells, which
contained abnormal quantities of soluble carbohydrate, but
no starch, no tannin, and no crystals. The cells were
short-lived.

[e) Orange rays caused a good growth closely approxi-
mating the normal. Crystals were not plentiful, however,
and the sugar was in rather large quantity.

(/) Red rays caused a growth which was even more
rapid than that made in white light. Tannin was formed
in larger quantities than under the normal conditions.

5. The action of monochromatic light, from the various
portions of the spectrum, upon solutions of diastase mixed
with starch paste, shows that yellow light causes the most
rapid hydration, while violet requires the longest period in
which to accomplish this result. The red rays required the
same length of time as white light. Wherever a marked
difference can be traced between the action of diastase in
rays of a certain refrangibility and in white light, there
too, the chemical composition and growth of Spirogyra is
abnormal; when, however, colored light and white light
coincide in their effect on diastase, the plant grows also.

6. Spirogyra cells de-starch rapidly, and without decom-
position, if, while preserved in darkness, a stream of fresh
water or a current of air be supplied to them. Cells kept
in darkness, without an oxygen supply, show a solution of
the crystal content, and a general breaking down before
the starch has all been converted into sugar.

7. Palladious chloride, in extremely dilute solution, serves
well to demonstrate the morphology of the nucleus and its
contents.

On the Structure and Pollination of the

Flowers of Eupatorium ageratoides

and Eupatorium coelestinum.

By Laura B. Cross, Ph. D.

(With rlate XVIII.)

IN the contributions that are being made to plant polli-
nation, the relative frequency of self and cross polli-
nation has received a considerable share of attention.
The fact that various flowers, once regarded as being
probably cross-fertilized have, during recent years, been
found to be systematically self-pollinated ; and also the
increasing number of plants that have been shown to bear
cleistogamic flowers, both point to the necessity for accu-
rate statistics, not only for the orders, but for the species
composing them. No order, probably, has received more
attention than the Compositrc ; but much remains to be
done in the extended study of the different genera and
species. The following is a contribution toward the study
oi Etipatoriiim suggested by Dr. Roth rock in 1891.

H. Miiller* has given some details, structural and physi-
ological, of his observations on Eupatorium cannahinum ;
and though the species I have examined differ in specific
details, the fundamental points established by him have
been confirmed by my observations.

Twelve plants of Eupatorium ageratoides and twelve of
Eupatorium coelestinum were selected and subjected to the
same treatment throughout. Four plants of each were pro-
tected by thin cotton cloth, and four by coarse Swiss
muslin, at a time when the flower parts were immature

*" Fertilization of Flowers," Eng. Ed. 1883, pp. 318-320; 363-364.
260

Cross — Eupatorium ageratoides and coelestinum. 261

and completely covered in by the involucre. The remain-
in^*- four plants were marked with bits of muslin and left
exposed. Six days later, twelve additional plants from each
species were selected. Two of these were covered by thin
cotton cloth, two with Swiss nmslin, two had one-half of
each inflorescence protected by thin cotton cloth, and the
other half exposed, two had one-half of each inflorescence
protected by Swiss muslin, two had single branches of
the panicle protected, and the remaining plants were left

exposed.

In each lot of plants selected, those remaining uncovered
so developed their flowers that the style arms divaricated
about three days earlier than those covered with muslin. The
nuislin-covere'd plants matured their flowers two days earlier
than did those covered with cotton cloth, this difference
being probably due to the greater amount of light passing
through the open meshes of the Swiss muslin, than through
the more closely woven cotton cloth. The difference in
the time of maturing was also very noticeable in the plants
which had a part of the inflorescence protected and a part
exposed, both having similar conditions, except in the
amount of light. Uncovered parts were faded, when covered
parts were just maturing. Another interesting diff'erence
between protected and exposed flowers, whether on the
same or separate plants, was that in every instance the
protected florets had their style arms developed in a very
contorted manner. Those of one floret often touched
a neighboring style or dipped into the adjacent corolla.
Hildebrand states that the style arms of chicory roll up
like a feather, the pollen being thus brought in contact
with the stigma. This he considers to occur most fre-
quently in the absence of insects. It was difficult to col-
lect a small amount of pollen from exposed plants, but in
protected ones the pollen was so abundant that it covered
the inflorescence and the inside of the muslin cover.
Indeed, it is difficult to form an adequate conception of the
amount of pollen produced unless the inflorescence be pro-

262 Cross — Eupatorium ageratoidcs and coekstiuum.

tected. There was no marked dilTerence in the time of
maturing of the fruits, whether borne on the most vigorous
early plants, or on those selected at later periods, the same
number of days bringing about, as nearly as possible, the
same results in every instance.

Iiiflorosceiioe of Eupatorium aj^oratoides and Eupato-
rhiiii cM»elestiiuiiii : Each capitulum is surrounded, and, in
early stages of development, completely covered in by the
involucre. E. ageratoidcs produces rather open panicles
of capitula with 12 to 17, commonly 15 white florets of deli-
cate texture on a flat disk-like receptacle. E. coclcstiniiin
bears compactly-clustered panicles, and for the massing of
a greater number of florets the receptacle is prolonged
upward into a sharp-pointed cone, each of which bears
from 46 to 60 florets of a bluish purple color. In both
species the pappose rudiment of the calyx (Fig. 8 m) con-
sists of a series of long multicellular hairs, the bases of
w*hich are surrounded by a very delicate tissue (Fig. 8 n)
which adheres to the torus when the pappose hair is
removed from the margin. The corolla in Eupatorium
agcratoides consists of a lower tubular portion, expanding
into an upper cup-like form and terminating in five
corolla lobes. These lobes bear on the outside, and near
their bases, multicellular hairs (Fig i //) longer than the
lobes themselves. These hairs occupy an upright position
until the corolla is expanded, when, by the recurving of the
petals, they are thrown back, become flaccid and wither
with the corolla. In structure tlie corolla exhibits not
only the long external hairs just referred to, but the inner—
or when expanded, the upper— surfaces of the petals are
covered with large, rounded, closely-set, tubular papillae,
(Fig. I p) which resemble the sweeping hairs on the
style, but differ from these in being shorter and in dimin-
ishing to slight swellings at the base of the corolla lobes,
where they finally disappear. These updirected outgrowths
may be of use to insects in affording them a firmer footing
when visiting the florets for pollen and nectar.

Cross — Eupatorium ageratoidcs ami coelestimim. 263

(b) The Corolla in Eupatorium coelestimim (Fig. 2) is
smaller and less spreading ; the lobes are shorter, more
rounded and less reflexed than in E. ageratoidcs. On the
outside, and near the base of the lobes, are found rounded
sessile hairs (Fig. 2 //) which in dried specimens are filled
with reddish purple pigment. The inner surface of each
lobe bears papillae only around its margin, and not uni-
formly over its surface, as in E.. ageratoidcs. The lower
part of the corolla in each species is composed of rather
long, straight, thin-walled cells, which extend upward as
far as the attachment of the filaments. Above this point
the cell walls become slightly waved toward the top of

the corolla.

[a) The Stamens of E. ageratoidcs are borne on the
corolla tube alternate with its lobes. Each filament in cell
structure consists of two parts, the lower being composed
of long, thin-walled cells extending through two-thirds of
its entire length (Fig. 4 a). The other part, next to the
anther (Fig. 4<^) is composed of oval or quadrangular cells
bounded by a thickened, beaded, strengthening wall, the
sauie in structure as the wall of the anther. The anther
consists of two lobes united by a connective which extends
upward between the lobes as an irregular, four-sided struc-
ture, broad on its external, and narrow on its internal face.
Each connective is expanded above into a transparent
deltoid process (Fig. 4 c) which unites with its neighbors
to form a pyramidal roof over the apex of the style, thus
protecting the more irritable parts from external influences
and the nectar from rain. The anthers are united by their
contiguous margins, and dehisce by introrse longitudinal

slits, (Fig. 5).

[b) The Stamens of E. coelestimim (Fig. 5) tliff'er from
E. ageratoidcs (Fig. 4) only in being shorter and less

firmly united.

[a) The Pistil of E. agcratoides is deeply cleft at the top
into two long style arms which are covered from their tips
downward over two-thirds their length with multicellular

264 Cross — E2(patoruiin agcratoidcs and coclcstimim.

hairs, that sweep the pollen from the anther cylinder,
hence called sweeping hairs (Fig. 6 e). Interspersed among
the sweeping hairs are occasional goblet-shaped nuilticelhi-
lar hairs. Immediately below the sweeping hairs are the
stigmatic papillae (Fig. 6 d and d') arranged laterally in
groups of three or four (^0 covering almost the entire
length of the lower third of the style arms. In this species
there is a glabrous area at the base of the style arms. The
style is slightly swollen at its base, and surrounded by an
annular nectar gland g,

(b) The Pistil of E. coelestinum (Fig. 6) is longer than
in agcratoidcs and is frequently twisted. The stigmatic
papilLx extend entirely to the bases of the erect style arms
which have enlarged club-shaped tips— Fig. 6 e. Hilde-
brand * states that in E. cannabinum the stigmatic surfaces
remain closely appressed, and that the stigmatic papilla,-
are not fully developed until after the pollen is matured,
shed, and carried away by insects, so that cross pollination
is insured. In the dried specimens of E. coclcstimim
examined, the style arms were slightly separated before
they protruded beyond the top of the anther cylinder, but
the stigma was not fully developed. In the matured flowers
the style stands more erect than in E. agcratoidcs. In
flowers not fully expanded, and with the anther cylinder
closed at the top, the style showed a distinct, loop-like cur-
vature just above the receptacle (Fig. 6 /). In fully ex-
panded flowers the style is quite straight.

The five anthers cohere to form a hollow cylinder which
is filled with pollen when the corolla begius to expand. In
the first stage with its two style arms closely applied to one
another, the pistil extends to the base of the anther cylinder
and gradually elongates, pushing the pollen before it out
of the cylinder by means of the outwardly directed sweep-
ing hairs (Fig. 6 e) which cover the upper part of the
style arms. When the stigmatic portion, which is smaller

*" Ueber die Geschlechts-Verhiiltnisse bei den Compositen." Verhand
der Leo. Carol. Acad. Dresden. Vol. 35 (1869).

(Jyoss — Fjipatoriujn agcratoidcs and coclcstimim. 265

in diameter than the area bearing the sweeping hairs enters
the otherwise empty cylinder, the pressure is reduced
and a probable transfer of liquid diminishes the turges-
cence of the filaments which bend themselves at point
a Fig. 4, and draw down the cylinder from over the
stigmatic area. In E, coclcstimim (Fig. 6) the stigmatic
branches are further carried up by the unfolding of the
loop-like curvature in the style. The pollen thus carried
out of the anther cylinder falls upon the corolla, and is at
once carried away by insects, or by the wind ; this closes
the first or staminate stage. The stigmatic surfaces readily
develop their papilhii and secrete stigmatic fluid. When
by frequent observations the pistillate stage was determined
in the first lot of plants selected, two of the first four plants
were carefully cro^s-pollinated by hand, and two were close
pollinated in the same manner. The second four plants
were left undisturbed until put in press, as were also those
exposed. In the second lot selected, two of the first four
plants were cross-pollinated, and two were close pollinated.
The second four, in which half the inflorescence was pro-
tected, and half exposed, were left undisturbed until put in
press, when ninety-five per cent, of the fruits grown on
exposed parts were well developed, while sixty-two per
cent, of fruits grown on protected parts were apparently
good— showing an increase of over fifty per cent, in favor
of exposed plants. The plants with separate branches pro-
tected were also cross and close pollinated ; while the last
two of the second choice were left exposed, to compare with
the last four of the first choice. Ninety-four per cent, of
the fruits were matured in each case.

The immature condition of the stigma at the time the
pollen is swept out of the anther cylinder clearly indicates
cross-pollination, whatever the agency may be. The pollen
is carried from the corolla by nectar or pollen-feeding
insects, or by the wind. If removed by insects, as seems
most probable, they carry it from flowers in the staminate
stage, to those in the pistillate as they pass over the capitu-

266 Cross — Eupatorium ageratoidcs and coclestifium.

lum, or from one plant to another. The results throughout
show that fertilization under these conditions did occur.
If the pollen be carried by the wind, it will as in the case of
insect agency, be deposited upon the fully matured pistils
— but the results recorded in the following tables do not
show that fertilization by means of wind did occur ; for
though the pollen was abundantly distributed over the
entire inflorescence and the inside of the covering of the
protected heads, the seeds could not be made to germinate
under the same conditions as those grown upon unprotected
plants. Moreover, the colors of the flowers and the abund-
ance of nectar secreted, would indicate entomophilous
fertilization, wliile the anthers are shorter than the corolla,
imited in a tube and dehisce introrsely — conditions unfavor-
able to wind pollination. From the time of pollination to
the time when the fruits are ripe, a period of eight to ten
days elapses. Previous to pollination the ovarian wall is
clear, pale, and delicate in appearance. In surface view it
shows very long cells, each containing a row of transparent,
circular areas at irregular intervals — Fig. 9. Cross-section
— Fig. 10 — shows five layers, each of which is composed of
cells of uniform size, except the innermost — Fig. 10 / —
which shows smaller cells. Soon after pollination the
fourth layer from the outer surface has pigment deposited
in it, which gives a most delicate ecru tint to the wall. As
the fruit matures the pigment is gradually deposited in the
fourth layer between the circular areas, whose perimeters
are much thickened, forming chimney-like openings (Fig.
10 k). The wall passes through every shade of brown,
until at maturity it is very dark brown. During the period
from pollination to maturity, no change occurs in the
circular areas. H. M. Ward * suggested in a recent
publication that the pigmenting of the maturing fruit
is a provision intended to act as a color screen against
the blue violet rays, *'as these rays would otherwise

* " Further Experiments ou the Action of Light on Bacillus Anthracis^''
by H. Marshall Ward.

Cross — Eupatoriian ageratoidcs and coclestinum, 267

destroy the reserve substance by promoting its rapid oxi-
dation.'^

E. AGERATOIDES.

Table of Statistics showing
comparative results of cross,

•<
*■*

a.
<

a. «

0 « :

UMBER OF

ERS DEVELOPED.

E NUMBER OF

ERS IN CaPITULUM.

OF ACHENES
RED.

OF ACHBNES
MATURED.

E NUMBER OF
RED ACHENES.

MBER OF DAYS UNDER
OBSERVATION.

MBER OF DAYS BLAPSED
BBT. POLLINATION AND
MATIRING OF ACHENES.

RCENTAGE OF GOOD
FRUITS.

close and natural pollination.

MBER
BXAM

TAL N
FLOW

HRAGl
FLOW

MBER
MATU

MBER
NOT I

ERAGI
MATU

'-> 10

> »

» >

" 1 >'

u

A ! H 1

1

< 2

i

Z j2

Cn

Plants matured under \

natural conditions, \ 8 1 117

14.5

I 10

7 137

23

__ \

94

Aug. 29 to Sept. 20. j '

Protected vSept. 3, cross

j)ollinated Sept. 19, \ 8

108

13-5

66

42 8.2

23

9

61. 1

gathered Sept. 27. ) i

Protected Sept. 3, close \ 1

pollinated Sept. 19, \ 8

106

13.2

43

63 525

23

9

39.6

gathered Sept. 27. J

I

245

15.3

10

235

.62

23

■

I

4.1

Protected Sept. 3, 1
gathered Sept. 26. /

16

•

Q

•

e \'t<y. .

<

H

6.

^g

2

12

t w i 2 , < z I 0

E. COELESTINUM. : h

■<

Table of Statistics showing ^ a

a. ui
comparative results of cros.s, ] 0 z

lUMBBR OF
ERS DEVELO

B NUMBER 0
ERS IN CAPI

OF ACHBN
RED.

OF AcHEN
MATURED.

E NUMBER C
RED ACHEN

. OF DAYS L
*VATION.

. OF DAYS El
POLLINATIO
'RING OF Ac

TAGE OF GO

rs.

close and natural pollination.

MBER
EXAM

FAL >
FLOW

ERAGI
FLOW

MBER
MATU

MBER

NOT ]

£ M '2 5 < Zx

3 : 0

> t> 1 --

>

3 ^

, r

A H

< 2 \^

<

z

<t!r,

!^ i

Plants matured under \

j

1

1 i

natural conditions, >- ' 8

448

56

352 96

44

23

7».5

Aug. 29 to Sept. 20. J
Protected Sept. 3, cross )

1 !

1

66.9

1

pollinated Sept. 19, [' 8 381; 481255

126

32 23

9

gathered Sept. 27. J

1

1

Protected Sept. 3, close |

1

pollinated Sept. 19, \ 8

424

53

102

322

13 23

9

24

1

gathered Sept. 27. J

1
1

1

Protected Sept. 3, )

R

4S6

«;7

1 "io

AO6

6.2 23

10.9

gathered Sept. 26. j

1

'

Referring to the preceding tables, it will be seen that
eight capitula grown under natural conditions produced 117

18

268 Cross — Eupatorium ageratoides and coelestimim,

fruits in twenty-three days, no of which were black and
swollen and contained well-developed embryos. Eight cap-
itula, selected from a portion of a panicle protected Sep-
tember 3d, cross-pollinated by hand September 19th, put in
press September 27th, produced 108 fruits. Of these forty-
two were white, shrivelled and imperfect, and sixty-six
were black and promised to be fertile. Plants protected
September 3d, close pollinated by hand September 19th,
the product gathered September 27th, produced 106 fruits
from eight capitnla, forty-three of which were black and
swollen, and when dissected showed well-developed em-
bryos, while sixty-three were pale and shrivelled, with im-
mature embryos. Plants protected September 3d, left
undisturbed till gathered on September 21st, produced 245
fruits from sixteen capitnla, only ten of which appeared
well-developed and fully matured. In order to test the fer-
tility of the embryos, I selected seeds showing the best de-
velopment, from plants grown under the four conditions
mentioned in the tables, and planted them January 7th, in
a large germinating dish between pieces of white felted
flannel. On March 31st radicles were seen protruding
from fruits grown under these conditions. This continued
until the middle of May, when at least seventy-five per
cent, had germinated. The radicle in germinating pro-
truded from the side of the fruit— Fig. 11— just above the
point of attachment to the stalk i. Twenty of the seed-
lings were potted, and made vigorous growth. None of the
seeds from plants cross or close pollinated by hand, or seeds
from plants protected and left undisturbed, germinated ;
although subjected to the same conditions as those grown
naturaUv. Reviewing the above observations, it seems
clear that self-pollination in the species of Eupatorium is
very rare indeed ; and that even when it does occur, the re-
sulting fruits are of weak germinating capacity. When
close pollination by hand is effected, a slight increase in
the production of good fruits is obtained ; but when cov-
ered flowers are cross-pollinated the increase is very strik-

Cross — Eupatorium ageratoides and coclestinum. 269

ing, and decidedly points to the conclusion that the arti-
ficial method adopted is an imperfect imitation of the action
of insects, whose visits assure the large percentage of good
fruits indicated in the table of statistics.

The foregoing brief observations point to important lines
of inquiry, which the writer may in the future be able to
follow out more fully,

DESCRIPTION OF FIGURES IN PLATE XVIII.

Illustrating L. B. Cross' paper on Eupatorium.

Fig. I. Longitudinal section of the corolla of E. ageratoides, show-
ing the external surface of two lobes of the corolla bearing papilht, /,
on their inner margins, and multicellular hairs, //, on their outer surface.

X65.

Fig. 2. Longitudinal section of the corolla of E. eoeleslinum show-
ing papillui, p, over the margins of the lobes and oval sessile hairs, //, on
the external surface. X 65.

Fig. 3. Stamen of E. ageratoides] a, junction of long, thin-walled
cells with b oval, quadrangular, beaded cell wall ; r, process of expanding

connective.

Fig. 4. Cross section of stamen of E. ageratoides. X 65.

Fig. 5. Four of the stamens of E. eoeleslinum, showing connection

and dehiscence, X ^5-

Fig. 6. Pistil of E. eoeleslinum ; g, nectary ; /, loop in the style ; d,
stigmatic surface; d\ high power view of stigmatic papillie ; c, sweeping

hairs. X ^S*

Fig. 7. g, nectary enlarged. X *50-

Fig 8. Longitudinal section of fruit, showing, /, receptacular stalk ;
ov, ovule ; ;/, transparent tissue surrounding ba.se of the pappus ; o, base
of corolla ; g, nectary in section ; r, base of style. X 65.

Fig. 9. Surface view of seed coat showing brown, elongated cells,

with pitted areas. X 350-

Fig. 10. Cross section through the seed coat showing, i, innermost
layer of small, thin-walled cells; fi, pigmented layer with pitted mark-
ings. X 350-

Fig. II. Germinating seedling with fruit wall attached ; g, shrivelled
nectary ; r, remnant of style. X 35'

Contributions to the Life History of Amphi-

carpsea Monoica.
{witu plates xix-xxxvi.)

By Adeline F. Schively, Ph.D.,

Assistant Teacher iu the Department of Biology, Philadelphia Normal School.

THE Genus Amphkarpcca was established by Elliott'
in 1 817. An account of his observations may be
found in the Journal of the Academy of Natural
Sciences of Philadelphia. He mentions in "The Botany
of the South "" two species— .4. monoica and A. sarmentosa.
In the '^Genera Plantarum" are enumerated seven spe-
cies inhabiting North America, Japan and the Himalayas.
The Genus is stated to contain fifteen species 111 " Die
naturlichen Pflanzenfamilien." These are distributed as
indicated above, except that two additional are mentioned
as occurring in Tropical America. These are A, pnlchella
and A. angustifoUa-hoWx found in Mexico, the latter also

as far south as Peru.

Both publications give A. Edgcivorthiixn the Himalayas,
also A, monoica in North America. Neither records A.
sarmentosa and A. Pitclicri, which are mentioned in some
of the leading American Manuals of Botany.

The investigation of A, monoica was undertaken at the
sucrgestion of Professor Macfarlane, who studied the sensi-
tiv'e movements of the plant, and also the production of
cotyledonary buds. My work has progressed under his

supervision. . ,,. . :n

The scientists who have worked upon Amphicarpcza will

now be mentioned in chronological order.

Darwin'^ experimented upon the movements of the leaves
and also upon the shoots producing subterranean legumes.
270

ArnphicarpcBa monoica.

271

A very short account appears in " The Power of Move-
ment in Plants."

In 1878, Farlow^' reported investigations upon Synchit-
riiim fiilgens—VTsx. decipiens—w\\\Q\\ badly infests the
plant in some localities.

During 1886 and 1887, Meehan^' studied the flowers and
fruit. The account of his investigation is found in the
'' Proceedings of the Academy of Natural Sciences (Phila-
delphia, 1887).'' The results of his observations are sum-
marized at the close of his paper. The statement given
below is not an exact quotation; the arrangement of
facts, however, is that used by him. His conclusions are

as follows :

1. Climbing as well as trailing stems bear apetalous

flowers.

2. These produce a third form of legume.

3. Petaliferous flowers under suitable conditions of nu-
trition produce legumes as freely as leguminous plants gen-

4. Petaliferous flowers are adapted to close fertilization ;
the apetalous are fertilized from these.

In 1890, HutV^ published a paper, entitled, *' Uber geo-
karpe, amphicarpe, und heterokarpe Pflanzen." He classes
AmphicarpcBa under the second heading indicated in the

above.

Macfarlane" reported to tlie Botanical Section of the
Academy of Natural Sciences (Philadelphia), his observa-
tions upon the production of cotyledonary buds. About
the same time (1893), in a lecture delivered at the Woods
Holl Marine Biological Laboratory,"" the sensitivity of
Amphicarpaa was compared with that of Mimosa, Cassia,

Oxalis^ etc.

This list comprises the known contributions to Botanical
Literature concerning AmphicarpcBa up to the present time.

2/2

Schively— Contributions to the Life History of

Ainphicarpcea nionoica.

273

General Conditions of Life.
AmphicarpcEa monoica is a plant quite widely distributed
tlirougliout the eastern United States. It is, indeed, one
of our most common weeds. Its favorite habitat is the
woods, where its fresh, green foliage presents an attractive
appearance during the spring and summer. It seems to
grow more luxuriantly, if it obtains a considerable amount
of sunlight, and if the roots receive abundant moisture.
Continued exposure to the sun's rays has an injurious
effect upon the delicate leaves. It thrives best in a rich
loam, but vigorous plants grow in clay or gravel, provided
the water supply is fairly plentiful.

The roots are fibrous and copiously branched. Upon
these occur singly or in clusters, the tubercles that are char-
acteristic of the Leguminosce. They are nearly spherical,
but slightly flattened, varying in diameter from one to five
millimeters. They are found when the plant is but a few
inches high. The number varies according to the char-
acter of soil, and whether the plant is grown in a pot or in
the open. (Plate XIX.) The comparative production of the
tubercles in different soils has not been studied experi-
mentally ; but they seem to be more numerous, as well as
larger, if the plant is grown in a pot.

Wood, Gray, and other botanists refer to A, monoica as
" perennial.' ' In this latitude there is no evidence of this
being typically the case. The plants appear in the spring
as the result of the germination of the seed. A very few
instances which indicate variation from this rule have been
found. While gathering underoround seeds in the autumn
of '95 I collected a plant possessing long thickened roots,
which could be described as tuberous. Unfortunately the
fact was not noticed, until too late to ascertain whether the
plant would really have perennated or not. In the spring
of '96, several specimens were obtained, in which, from
thickened root and lower stem region, arose the new plant ;
at one side persisted the dried remains of last year's stem.
(Plate XXXII., Fig. i.)

The stem is herbaceous, but often becomes very woody
toward the end of the season's growth. It is usually green,
though often it may be quite purple. In habit, the plant
is a twiner, but in the manifestation of twining, great differ-
ences are observed. A vigorous plant will ascend rapidly
to the top of any weed, whose firm stem offers a sufficient
support. If no such sturdy neighbors are near, several
plants may twine around each other, forming a thick coil,
which for a long time may assume a procumbent position.
So delicate is Amphicarpcea, and the stem so weak, that
the protection of other plants seems a necessity. In at-
tempting to grow specimens out of doors, either in the
ground or in pots, it was noticed that they suffered mark-
edly from even slight winds. Though provided with a
stick, the twinings were uncoiled ; finally shoots and even
leaves were injured.

Legumes and Seeds.

Legumes are produced both above and below ground.
A description of these and of the seeds is essential to assist
in explaining the accounts of germination.

Terrestrial Legumes vary much in size, shape and thick-
ness. They are ellipsoidal, much flattened laterally, and
might best be described as irregularly pyriform. Mature
specimens are on the average three-quarters of an inch in
length, and about three-eighths of an inch in thickness.
They vary greatly, measuring, however, from a half inch to
occasionally an inch in length. (Plate XIX.)

There is considerable diversity in color. Some are pale,
others are rich pink-purple, and still others are dark purple
or reddish brown. It is suspected that the character of the
soil exerts an influence in this respect. Those found in a
very moist locality are usually darker than those found in a

dry situation.

The entire surface of the legume is very hairy. For a
long time the calyx persists (Plate XX., Fig. i), but in the
larger and older legumes, it is scarcely discernible. It is

m

274 Schively — Contributions to the Life History of

probable, therefore, that It gradually decays. Usually
there is a single seed, filling the entire cavity ; but a
two-seeded condition is not uncommon (Plate XX., Fig 2).
The walls of the legume are at first comparatively thick,
but the tension consequent upon the growth of the seed
causes them to become quite thin. Histological peculiar-
ities will be described later in this paper.

The seed is smooth, of a mottled pink purple color, and
varies in size and shape ; the statements made concerning
the legume, practically applying here. The coat consists
of the following layers : a row of strongly indurated cells,
prismatic in shape, with long diameter lying at right angles
with the outer surface of the coat (Plate XX., Fig. 3, a\
These cells merge externally into a homogeneous layer of
similar tissue— the cuticle. Purple coloring is diffused
throughout many of the cells of the prismatic layer. Imme-
diately underneath these, are several rows of parenchymatous
cells, many of which contain chloroplasts (Fig. 3, b). In
the neighborhood of the hilum the layers are increased to
form a chalaza, though there is no change in character, nor
are new types of element introduced. In that portion of the
coat, directly bordering upon the cotyledons, a row of flat-
tened elongated cells is seen, forming the inner epidermis.

(Fig. 3)

The seed is exalbuminous, and contanis a large propor-
tion of starch ; the granules measure 6 /i, their prevailing
form being ellipsoidal. The cotyledonary cells contain
numerous refractive protein granules, the nature of which
has not yet been determined.

The aerial legumes for the present will be referred to as
{a\ {b) and {d) ; {a) resulting from purple flowers ; {b) from
green inconspicuous ones, produced in late summer; and
{d) from somewhat similar green, inconspicuous ones pro-
duced in winter only.

[a) Legumes of this type are about one and a half inches
long, stipitate, and decidedly tapered toward the apex,
where the remains of the style persist. The shape might

Amphicarpcea monoiea.

275

be described as falcate; the surface is smooth, excepting
the dorsal and ventral sutures which are hairy. The calyx
persists and remains quite conspicuous until dehiscence.
(Plate XX., Figs. 4, 5.) The seeds vary in number from two
to four ; three, however, are generally found. The walls
of the legume are firm, resisting and quite thick.

[b) The legumes of this type are about an inch in length,
and are not s'tipitate, or but very slightly so. They are not
tapered, their shape being nearly oblong; the style extends
from the apex of each as in [a\ and its calyx is smaller than
that in (a\ but like it persists. The surface is slightly
hairy; the sutures resemble in condition those of [a\
Usually there are two seeds, but the number varies from
one to three. (Plate XX., Fig. 6.)

{d) The legumes of this type are still smaller than those
of (/;), and are sessile. They are oblong, the style remaining
as a hook which often lies close to the suture. The sur-
face as well as sutures are noticeably hairy. The seeds num-
ber one or two, the latter being most common. (Plate XX.,

Figs. 7, 8.) ^ .

Histological features will receive attention toward the

close of the paper.

In the aerial types (^, b, d) the seeds are reniform, varying
from one-quarter to three-eighths of an inch in length.
When ripe the color is grayish green, flecked with dark
purple. Upon exposure to the air they become dark
brown, with purple black spots ; quite often the coloring is
a uniform purple brown or black. Type [d) bears the
largest seeds; {a) and {b) bear seeds of about the same size.
In the general aspect of these but little variation is seen.
Those of {a) seem darker, but it is doubtful if this distinc-
tion wall hold.

The structure of the coats of the aerial seeds will be de-
scribed according to the sequence adopted for those of
the terrestrial. No histological differences in the three
types of aerial seed have been observed. The outer indu-
rated laver of cells resembles in shape, arrangement and

276 Sclav dy— Coitributions to the Life History of

general coloration, that seen in the terrestrial, bnt the
depth of the layer is nearly twice as great as that in the ter-
restrial (Plate XX, Fig. 9, a). Beneath is a row of cnrionsly
shaped cells, that are strongly thickened and possess flanges
abutting upon each other. Six or eight rows of cells he
below tiiis, showing a gradual or sudden change from indu-
rated to parenchymatous type, according to the region of
the seed from which the section is taken. In the region of
the hilum all of the above layers increase remarkably in
size and thickness. Forming the chalaza are numerous
pitted and reticulate tracheids, of the character usually
found toward the termination of vascular areas. In this
part of the seed, Iving between the tegmen and the cotyle-
dons, is a narrow tapering band of sclerenchyma, limited
in extent. Bordering directly upon the cotyledons is the
inner epidermis of the coat ; this consists of narrow elongated
cells. Immediately above these epidermal cells are two,
occasionally three, layers of strongly indurated elements

(Fig- 9 ^^)- r 1

The seeds of the three aerial types, like those of the ter-

restrial, are exalbuminons. In the aerial seeds the prevail-
ing shape of the starch granules is spherical ; they are
smaller than those of the terrestrial. The proportion of
proteid seems greater in the former than in the latter.

Weight.

Equal numbers of terrestrial and of aerial seeds were
weicrhed, and then dried in the oven for from two to four
hours at a temperature of about 80° C The resnlts have
been arranged so that comparisons of the weights of a
single seed of each class may be made.

Small terrestrial seeds weighed about -345 grammes;
after drying, .181 grammes. Medium sized seeds v^^\^\\^^
1. 143 grammes ; after drying, .605 grammes. In a certain
locality near Burmont, several exceedingly large specimens
were found. The weight of one of these was 2.601 grammes ;
after drying, 1.1711 grammes. Thus it will be seen how

Amphicarpcca monoica.

277

great is the proportion of water present in this class of seed ;
certainly from forty to fifty per cent, of the entire weight

is lost in drying.

The aerial seeds do not contain so much water as the
preceding. Seeds of type {a) weighed before drying, .0291
grammes; after, .0274 grammes. Specimens of type [b)
weighed .0611 grammes; after drying, .0564 grammes, while
those of type [d) weighed .0741 grammes, and afterwards

.0726 grammes.

In the '^Fertilization of Flowers^' (Miiller, Eng. edit,
page 214), a reference is made to some of Darwin's results.
These were published in a certain edition of "Different
Forms of Flowers in Plants of the Same Species." As a copy
of this could not be found, no more exact information can be
given. He found the weight of the aerial seeds to be 7V, that
of the terrestrial form. There is no statement as to the size
of the terrestrial ones— whether small or medium. The
figures above will give a much larger fraction.

Germination.

Such remarkable differences in size, weight, and struc-
ture of seed-coats prepare us for more or less striking pecu-
liarities attendant upon germination. When the investiga-
tion of Amphicarpcca was begun in the autumn of '94, a
quantity of both aerial and terrestrial seeds was collected,
and placed in boxes to be used as occasion required. The
terrestrial ones became exceedingly hard, almost horny in
texture. Both varieties were soaked for twenty-four hours
previous to planting. Very few aerial seeds germinated ;
and only a few terrestrial ones, which had been placed m
pots of earth almost immediately after gathering. Upon
investigation it was found that those which had been dried
and later soaked, had decayed. The same result was ob-
tained each time the experiment was repeated. No germi-
nation records were possible during the succeeding winter.
In the following autumn, all terrestrial seeds were put
in pots of earth, slightly moistened and kept in a cool

278 Schively — Contributions to the Life History of

place. As quite satisfactory results were obtained in gerini-
iiating these, this method of preserving the seeds is recom-
mended. All experiments described, were conducted in the
greenhouses of the Botanic Garden of the University of
Pennsylvania. The temperature ranged from about 20° C.
to 36° C, probably averaging 27° C.

Upon October 14, 1895, twenty terrestrial seeds and the
same number of aerial were planted. Several aerial ap-
peared in the course of three weeks, but no terrestrial. All
of the latter germinated, however, during January, 1896.

Upon December 16, 1895, thirty-two seeds of each kind
were planted. An aerial seed first appeared within ten
days ; about December 30, several terrestrial seeds showed
themselves above ground. By January 18, 1896, twenty-
six terrestrial and four aerial seeds had developed. After
this, at irregular intervals, the remainder of the terrestrial
germinated. Result, 100 per cent, terrestrial; I2>4 per

cent, aerial.

On January 4, 1896, twenty terrestrial seeds were planted
as follows: Ten, one inch below the surface; ten, two
inches below the surface. On the same date, twenty-five
aerial seeds were planted, ten, one inch below the surface ;
ten, two inches below the surface, and five, one-half inch
below the surface. On January i8th, one aerial (i inch)
had appeared above ground ; investigation showed that one
terrestrial (i inch) had germinated, though no shoot was
visible above ground.

After this results were as follows :

Plants above f>: round.
Jan. 2ist.

I Aerial (i inch).

1 Terrestrial (i inch).

Plants above ground.
Jan. 29th.

1 Aerial (i inch).

4 Terrestrial (i inch).

2 Terrestrial (2 inches).

Plants above ground.
Feb. 1st.
2 Aerial (i inch).

5 Terrestrial (i inch).

6 Terrestrial (2 inches).

Plants above ground.
Feb. 5th.
2 Aerial (i inch).
6 Terrestrial (i inch).
10 Terrestrial (2 inches).

Amphicarpcea monoica.

279

Plants above ground.

Feb. 8th.

3 Aerial (i inch).
6 Terrestrial (i inch).
10 Terrestrial (2 inches).

Plants above ground.

Feb. nth.

3 Aerial (i inch).
10 Terrestrial (i inch).
10 Terrestrial (2 inches).

Results, 100 per cent, terrestrial, 12 per cent, aerial.

Keeping aerial seeds in water at a temperature of 35"" to
40° C. did not materially increase the rate. Considering
the results obtained from filing Cajina and other seeds with
hard coverings, I concluded to try the method here. The
percentage then increased to 50, sometimes 70.

About February i8th, seedlings from terrestrial seeds be-
gan to appear above ground ten to twelve days after
planting, and this time-interval between the planting and
germination decreased as spring advanced.

In the autumn of ^95, a pot containing one hundred seeds
was ])uried out-of-doors a foot below the surface, where it
remained during the winter. April ist the weather became
quite warm. April loth the pot was dug up ; every seed had
germinated, and within three or four days all plants came
above ground. From the middle to the end of April is the
time for the appearance of AmphicarpcEa seedlings in their

native haunts.

It would seem, however, that terrestrial seeds require a
certain period of rest even if the requisite heat and moist-
ure are provided. For instance, seeds taken upon April
19th, from plants grown in the greenhouse during the win-
ter, were planted immediately, but failed to germinate until
early in June. While collecting seeds, on October 13,
from plants grown in the greenhouse during the summer, it
was impossible to gather all ; many remained among the
cinders. Notwithstanding the favorable conditions to
which these seeds were exposed, no indications of plants
were seen until the latter part of December.

Similar results to those obtained in the autumn of '95
were got in '96. Again, after the latter part of December,
germination proceeded rapidly.

28o Schively — Contributions to the Life History of

Having learned by the autumn of '96 that there was more
than one variety of aerial seed, efforts were made to ascer-
tain their respective powers of germination. In the case
of type [a] the percentage ranged from naught to twenty.
With seeds of type [b) the same percentage variation was
obtained. But filing in each case raised the percentage
to seventy. Seed of type {d), (found only during the
winter) all germinated within six weeks, if planted imme-
diately after being removed from the pots. If allowed to
dry, results were no better than those previously mentioned,
but with these also, filing raised the percentage.

Very satisfactory results have been obtained by germinat-
ing between layers of moist flannel, kept near steam pipes
(temperature of course quite high). Within four or five
days many had germinated ; at the end of two weeks there
was a well-formed stem and root. All terrestrial seeds re-
sponded quickly to this treatment, whether legumes or seeds
only were used. Unfortunately this method was not tried
until after January i, 1897, so that I cannot say whether it
would be possible to shorten the supposed resting period or
not, had the experiment been made in the early autumn.

With all seeds a temperature of 28° to 30° C. is essential
for rapid results.

It would seem from the above data, that it is rare indeed
for terrestial seeds to fail to reproduce the species.

Aerial seeds were likewise placed under the above con-
ditions; germination was nnich slower than for terrestrial
ones. Type [a) gave 2 per cent. ; but with 90 seeds of type
(b\ not one germinated.

Some aerial seeds gathered in '94, gave results but little
different from those of '95, when subjected to similar con-
ditions at the same time.

Doubtless the peculiar structure of the seed-coats of the
aerial ones is an obstacle to their more general germination.
In nature, few, if any of these are likely to produce plants
in the succeeding spring.

The seeds of type (/;), while in general appearance and

Amphicarpcea nionoica.

281

histological structure not distinguishable from type (^), do
not scan to possess the same capacity for responding to
conditions suitable for germination if both sets are planted
without filing. It may be that (b) requires a longer resting

period than [a\

An interesting feature has been noticed in the behavior
of the cotyledons, of both terrestrial and aerial seeds, but
particularly the former. Normally these, as has been stated,
remain below the surface; if however, the seeds happen to
be in such position' that the cotyledons are wholly or partly
exposed to the light, chlorophyll is developed with astonish-
ing rapidity. Nor does it seem that much light is required ;
seeds which were germinated between layers of flannel, in
a deep box placed under the shelves near the floor of the
greenhouse showed this phenomenon well within three
weeks. Aerial and terrestrial seeds behaved similarly in

this respect.

During the winter a plant raised from a terrestrial seed,
whose cotyledons remained half above the soil, attained
the height of two and a half feet, and produced numerous
legumes. The size of the cotyledons was but little reduced;
they were firm and not at all shriveled in appearance. The
exposed portion of the cotyledons was an intense dark green,
really deeper than that seen in the leaves ; the subterranean
half was quite white ; a sharp demarcation showed plainly
how the seed had been placed.

Sections through the very green portion showed chloro-
phyll granules, closely crowded around the cell walls;
in the deeper tissue of each cotyledon, these granules were
seen, sparingly scattered. Still deeper, no granules were
found. The iodine test revealed the presence of the large
starch granules characteristic of the terrestrial seed. These
were most numerous in the colorless portion, but were not
by any means absent in other parts. In addition, minute
starch centres were detected in the chlorophyll granules

themselves.

Seeds planted in normal position soon develop a small

282 Schivcly— Contributions to the Life History of

shoot in the axil of each cotyledon. The main stem of
the plant at this time is eight or ten inches high. If, how-
ever, the cotyledons are placed, so that they are exposed to
the light, longer shoots, fonr in nnmber— two for each axil
—are developed by the time the plant reaches the height
mentioned above. Cutting off one of the cotyledonary
shoots caused the production of two more buds of unequal
strength. This operation has been repeated three times in
succe'ssion, upon the same plant, with a like result. I am
not prepared, however, to state how frequently there will^

be similar response.

It is true, that the cotyledonary shoots developed in nor-
mal position give somewhat similar results, when experi-
mented upon in this manner, but the response to the
operation of cutting is not so rapid.

In the capability to develop such a quantity of chloro-
phyll, Awphicarpcm seems unique. The cotyledons of
Vicia Faha^ normally hyopgean, when exposed to the light,
do behave similarly; but the depth of color is not so pro-
nounced.

Eihvardsia chilensis is the only seed mentioned by Ivub-
bock,'Miaving subterranean cotyledons, which are greenish.
Nothing more explicit is stated; one cannot tell whether
that is "the natural color, or whether it is produced by
accidental exposure to light.

At present no other member of the Leguminosae is known
to exhibit such a marked tendency to form chlorophyll in
the cotyledons, unless those organs naturally arise above
the surface of the ground.

Young Plants.

As has been stated, the cotyledons of both terrestrial and
aerial seeds remain underground. The radicle has grown
quite long, and the petioles of the cotyledons have elon-
gated considerably, before the bent epicotyl issues from their
protection. For some time the epicotyledonary region
remains in this position. From the moment of germina-

Amphicarp(2a mofioica.

283

tion, the contrast between the seedlings of aerial and
terrestrial seeds is evident, and becomes more pronounced as
growth increases (Plate XXL) The stem from the terrestrial
seed, as it appears above ground, is of a deep purple hue; it
is strong, vigorous and grows rapidly, soon carrying the first
pair of opposite green leaves to a height of four or six inches.
After this, compound leaves are produced ; the internode
between the simple leaves and the first compound leaf is
short. When the stem is about ten inches in height,
it exhibits an inclination to twine ; if a support is within
reach, the plant soon becomes a vigorous twiner and
the internodes increase in length. The stem of the plant
from the aerial seed is slender, feeble and is usually
green. It grows during the first ten days almost to its full
height, which is much below that attained by the preceding
plant. The simple and compound leaves are about half
the size of those of the first described. Twining does not
take place ; even tying to a stick failed to induce twining
movements. During the winter variations from such types
were seen. A terrestrial was found which was low growing,
although it still twined. (Plate XXXV., Fig. i.) Occa-
sionally an aerial grew taller, or more strictly perhaps,
longer ; but reclined upon the ground. This striking dimor-
phism continued throngh the winter. Plants produced from
• filed seeds behaved as the others. In the late spring a good
many of the aerial seedlings twined; their general appear-
ance was feebler than that of the terrestrial form. Reference
to these facts will again be made toward the close of this
paper.

In the descriptions which follow, plants produced from
terrestrial seeds are intended, for they furnished specimens
apparently most normal in behavior.

Below the first pair of green leaves the .stem is nearly
smooth, above these, the stem exhibits all degrees of hairi-
ness. The hairs are always retrorse, and vary from a rather
scanty growth to a thick brown felt. The vigor of the plant
19

284 Schivdy— Contributions to the Life History of

and the place of its growth seem to exert an influence upon
this characteristic.

COTYLEDONARY BUDS.

The germination of seeds of xhnphiearpcca by Macfar-
lane'^ had been interfered with by mice; and the same
difficulty confronted the writer. Even the raising of plants
out of doors was not without serious draw-backs. More
than one plant several inches high was visited by rabbits
or by sparrows, which found the fresh green herbage agree-
able. The main shoot was often removed close to the
ground. Upon examining the plants a few days later, in
nearly every specimen several cotyledonary buds had
developed and were manifesting apogeotropic tendencies.
The number varied from two to five; they were thick and
white. It is not usual for more than two to appear above
ground, but occasionally three have been observed. It was
found that plants from three to six inches in height that
were thus injured sent up within ten days two shoots arising
in the manner described.

These occurrences suggested experiments in which the
plant was deliberately injured by cutting off" the main stem
close to the ground. Many of these experiments were
performed with results exactly as given above. A plant
fifteen or more inches in height, so treated, either never
developed cotyledonary shoots, or else after many weeks
one appeared above ground— a tiny feeble thing.

If four or five cotyledonary buds have developed, and a
vigorous shoot resulting from one is then cut off, the
immediate result seems to be the formation of two more.
For example, one so treated had finally ten buds.

These experiments were not a success when plants pro-
duced from aerial seeds were used. Sufficient vitality
seemed to be lacking.

It is worthy of notice for comparison with statements
soon to follow, that these cotyledonary shoots never bear
simple leaves, but compound only. They soon begin to

AniphicarpcBa monoica.

285

twine, however, and the growth of the cotyledonary shoots
in all respects is about equal.

Axillary Shoots.

Let us now turn attention to the cotyledons of a plant
which has developed without accident. Examining the
conditions below the soil, it is found that from two to five
runners arise in the axils of the cotyledons. These usually
remain underground, and are therefore white, bearing
stipules, but no leaves. For a time their growth is com-
paratively slow. Plants in the woods, developing from
seeds in the spring, showed about July ist, cotyledonary
axillary shoots from three to six inches long. Those raised
in pots were not so long, but were more branched. As the
summer advances, these runners elongate rapidly, and
eventually may become twenty to fifty or even sixty inches

in length.

In the axils of the green leaves both simple and com-
pound, arise branches, which may be referred to as axillary
shoots (Plate XXII.). In the case of the simple leaves, two
to four, occasionally six, of these are found. Possibly here
the term "runner" maybe more appropriate. The com-
pound leaves show one, often two axillary shoots. All of
the axils of leaves upon these original branches possess
similar possibilities. One may easily imagine therefore the
mass of vegetation resulting from a healthy plant, from
which axillary shoots three or four feet in length are

actively growing.

This development in the axils does not begin until the
plant reaches the height of ten to fourteen inches. Some-
times their increase is so rapid, that growth of the main
stem is evidently retarded.

Apogeotropic and geotropic tendencies.

Plants resulting from terrestrial seeds twine more or less
vigorously— and the main stem is decidedly apogeotropic.
The axillary ''runners" 2.1^ geotropic ; as soon as the bud

286 Schivcly— Contributions to the Life History of

develops, this tendency becomes manifest. When the
gronnd is reached the tip does not seem to penetrate the
soil bnt growth takes place and the shoot may extend along
the'snrface for some distance. Many of these rnnners are
quite leafy ; others are devoid of those organs, and show
stipules only. From the axils of the stipules grow numer-
ous branches of varying length which exhibit geotropism
in beautifully curved tips.

The extremity of the runner is often turned upward m
the most curious manner. In order to ascertain if these
runners possessed any apoi^reotropic tendencies, many of
them were given an opportunity to twine. A support was
placed for them, they were even assisted by being artificially
twined around the stick and then tied to it. In all cases
they persistently refused to twine, gradually turned so that
the tip inclined downward, and as growth contmued they
finally reached the soil and behaved as previously described.
Experiments, made upon the runners when about four
inches long, consisted in carefully cutting a sixteenth of an
inch, and also an eighth from each apex. The shoots were
then supported and inclined upward. In the course of two
days, they had grown an inch and showed a decided geo-

tropic curve. .

It has been noticed that these runners from the simple
leaves appear to be negatively heliotropic. No special
experiments have been made which positively prove that
this characteristic is stronger than the geotropic one. These
axillary runners and their branches continue their growth,
finding now a crevice, now a space under a flower-pot—
both of which are suitable places for the maturing of ter-
restrial fruits. In the green-house, the plants were placed
upon a shelf about a yard in width, raised several feet from
the floor. Many runners from the plants finally extended
across the shelf either directly or diagonally, and continued
their growth in the dark space below, many of them

maturing fruit. ,. r u

In the woods some of the best developed and most Iruit-

Amphicarpcea monoiea.

287

ful specimens of axillary runners have been those growing
among the dead leaves, which excluded nearly all the light.
AiraxilUiry shoots from the axils of compound leaves
exhibited for a time geotropism. Plants which were
inverted gave proof of this. Care was taken that plants
selected for this experiment should have short axillary
shoots (not more than an inch and a half). Within twenty-
four hours, the portions of the specimen had adjusted
themselves to their altered relations; the shoots were

geotropic.

Ordinarily this condition does not last long, for as growth
increases, a great curve is noticed near the tip, sometimes
extending for several inches. All that is needed is the
placing of a support near the apex. In a few hours twin-
ing begins, and consequently apogeotropic tendencies are
manifested. If no support is given, the shoots will twine
around any neighboring support, or indeed around some
adjacent individual of the same species.

We may now recapitulate briefly. Axillary runners from
simple leaves are geotropic and negatively heliotropic. In the
young condition, the axillary runners from compound leaves
2iX^ geotropic; this characteristic is usually lost quite soon,
for they become apogeotropic and twine readily. The last
mentioned shoots usually appear to resemble the main stem
in their habits of growth.

Occasionally exceptions to this last statement are seen in
certain plants that, differing in no apparent particular from
others in their early life, exhibited later the peculiarities
which will be described below. The axillary shoots from
compound leaves, as well as the branches from these, were
purplish, rather than green, and leaves were very few;
often entirely absent. Such shoots seemed always geotro-
pic. I have now in mind, one plant which grew in the
greenhouse during the summer; it became fully five feet
high ; the main stem twined vigorously. The long purple
shoot's were numerous, but they did not exhibit the slightest
inclination to twine, even around neighboring shoots ; they

288 Schwely — Contributions to the Life History of

grew directly downward. Yet just beside this specimen
were plants with many leafy branches which twined upon
any available support.

Truncation Experiments.

Having observed these phenomena, a series of what may
be termed " truncation experiments" were made. These
consisted in cutting off the main stem at different levels,
and noting the result upon the future development of the
axillary shoots. Nearly all of these were performed in the
green-house during June and July, a few, however, during
the winter months. The results of some of these are stated
below.

I.

Cutting off Main Shoot above Simple Leaves.

On June 6, 1 896, the main shoot was cut off above the
simple leaves; in the axil of each was a branch one inch

long.

On June 16, both branches had lengthened considerably ;
they formed almost a right angle, and were growing up-
ward.

On June 29, the axillary runners were about eight inches
long, and although growing upward, had not yet twined.
Sticks were placed for them.

On July 6, the runners were over a foot long, and both
were twining. (Plate XXIII.)

On June 19, the main stem of a plant was cut above the
simple leaves ; also all buds from the axils.

On June 29, additional accessory buds had developed,

and the shoots were one inch, and one and one-half inches

respectively. Another bud was appearing in the next axil.

On July 7, the branches had turned upward, and were
three and six inches respectively. The bud remained in
about the same condition.

On July 15, the branches were ten and fourteen inches.

Amphicarp(Ea monoica. 289

As no support had been given to them, they twined around
each other.

On Jul v I, the main stem of a plant was cut off above
the simple leaves. One of the axillary branches was taken
away ; the other about ten inches long was left.

On July 15, it was seen that the runner had increased to
fifteen inches, and in the axil of each small leaf upon it
had appeared a little branch. The main runner did not
twine nor exhibit any inclination in that respect. An ac-
cessory bud was seen in the axil of the simple leaf.

In some cases the shoot declined for some distance out
of the pot, then twined feebly around some support that
was near. It did not take the one provided in its own pot.
The results obtained in this series bear a direct relation to
the age of the shoot at the time when operations are begun.
One may readily see, then, that if the true stem is de-
stroyed, its place is taken by the axillary shoot, and that
which in its normal condition possessed pronounced gcotropic
characteristics, noiv exhibits apogeotropic ones. Yet, if the
injury takes place after the shoot is tolerably long, it is
impossible to change its strongly-impressed habits; its
inherited tendency remains.

Cutting off Main Stem above First Compound

Leaf.

On June 6, the main stem of quite a tall vigorous plant
was cut off above the first compound leaf. Axillary
branches were removed from the simple leaves. A single
bud remained in the axil of the compound leaf.

On June 29, the branch which had developed in the axil
of the compound leaf was vigorously twining. The plant
was about two feet high. Axillary runners from simple
leaves were ten inches in length. • , • 1

On July 8, the plant was two and a half feet in height ;
the axillary runners, eighteen inches.

290 Schivcly — Contributions to the Life History of

On June 12, the main stem of a plant was cut oflf above
the first compound leaf, and all the axillary buds from the
simple leaves were removed.

On June 29, the plant was one foot high, and the newly-
developed shoot in the axil of the compound leaf was

twining.

On July 8, the plant was one and a half feet high, and
twining well. Axillary runners had developed.

On July 13, the new twining shoot was cut off above its
first compound leaf. A bud remained in its axil. All
other axillary buds and branches were removed.

On July 15, the bud had developed into a branch an inch

long.

Unfortunately the last date has been omitted from my
notes, but in a short time this new branch twined and con-
tinued the upward growth of the plant.

Cutting off Main Stem above Second Compound

Leaf.

On June 6, the main stem of a vigorous plant was cut off
above the second compound leaf. One bud remained in
the axil of this ; all others were removed.

On June 29, the plant was two feet high. The new
shoot was twining ; the internodes were rather longer than
those on main stem. One axillary shoot ten inches long
arose from the axil of the first compound leaf. Along the
twining shoot a bud was seen in the axil of every leaf.

The plant continued to increase in height, and the axil-
lary shoots developed rapidly.

Experiments with Cotyledonary Shoots.

Roots of young plants were carefully washed, then re-
placed in pots, so as to bring the cotyledonary shoots above
ground. Occasionally they were found already appearing
above ground. Usually they were white and finely pubes-
cent ; then they soon acquired a green color. For some days
they grew slowly ; they were supported by small bits of

Amphicarpiea monoica.

291

wood, and inclined upward, in order to prevent a continua-
tion of their tendency to return to their natural abode. A
record of a few of these is now given.

On June 27, a plant was arranged so that the cotyledo-
nary shoot was above ground. The shoot was short and

white. . 1

On July 8, the cotyledonary shoot was quite green, ana

was about six inches long. It had branched, and was

growing out in a horizontal direction. It was turned up

and tied to a stick.

On July 15, the shoot was about twelve inches long, pos-
sessed branches and leaves. It was longer than the original
stem, but no twining had taken place.

On June 27, a plant was found having the cotyledonary
shoots above ground. There were two about four inches

long.

On July 6, the shoots had increased in length. One was
tied to a stick (the other was accidentally broken). The
main stem of the plant was twining.

On July 15, the plant was eighteen inches high, the
shoot about fifteen inches, but did not twine.

On July 22, no twining had occurred.

On June 27, two small cotyledonary shoots, already
green, were supported as described above.

On July 8, the shoots were four inches in length, and
had made a decided right angle, for they were now grow-

ing upward. rr tm i »

On July 15, the main plant stem was cut off. Tlie shoots

five and seven inches respectively bore tiny green leaves.

They were tied to a stick.

On July 22, no twining had occurred.

About this time, a plant having four cotyledonary shoots
above ground was noticed. It was observed again early in
Sentember. Extensive growth had taken place, but no

292 Srhhclv — Contributions to the Life History of

twining, althongh the shoots varied in length from twelve
to eighteen inches. The main stem had grown but little.

When plants much older than these (about three feet
high) were similarly treated, no results were obtained.
The shoots remained white and did not seem to increase in
length, and finally died.

Another experiment was as follows: A vigorous plant
was truncated just above the simple leaves; every axillary
bud was removed, and this process was repeated, no bud
being allowed to develop. The result was that in the
course of about ten days two cotyledonary shoots had arisen
from the soil. Later on twining took place.

Referring for a few moments to the paragraphs upon the
development of cotyledonary buds, (p. 284) it will be found
that when the main stem is destroyed, and the developing
shoots take its place, twining becomes their habit. Yet if
the stem persists, and they are brought above ground, their
growth is more or less horizontal.

Thus the cotyledonary shoots, and the axillary runners,
which under normal natural conditions are geotropic, may
acquire apogeotropic tendencies, if necessity places certain
responsibilities upon them.

Illustrations need not be multiplied; enough has been
said to indicate the deep resources of the plant. In the
'* struggle for existence " many, if not all, of the conditions
artificially produced, no doubt do occur. Very beautifully
does Amphicarpcea show its ability to cope with adverse cir-
cumstances, and by various growth-compensations to pro-
long its existence, and even to continue its accustomed
habits of twining.

ClRCUMNUTATION EXPERIMENTS.

In his writings, Darwin includes under the term circum-
nutation, the movements of heliotropic organs, those of geo-
tropic organs, as well as those of the twining stem. Under
the last heading, belong the principal experiments to be
described in this section.

Amphicarpcra monoica.

293

It is not the intention to discuss here in detail, the various
theories which have been advanced concerning the behav-
ior of the twining stem, the phenomenon of its circunmu-
tation, and their probable relations.

Von Mohl, Palm, Darwin, Dutrochet, De Vnes, Pfeffer
Sachs Kohl, Noll, and other authorities have contributed
\o the literature dealing with these problems. To a cer-
tain extent most of them believe that the peculiar manifes-
t'Uions of the movement are due to variations of turges-
cence in the growing cells. Numerous hypotheses have
been advanced to explain the movement.

Dutrochet suggested that twining stems possessed pro-
perties peculiar to themselves, but his efforts to theorize
concerning their activity are not particularly lucid. Von
Mohl, and later. Kohl, asserted their belief that twin-
inc. plants are endowed with irritability. More recent
auUiors concur in this opinion. The stimulus afforded by
contact with the support is of vast importance in causing
the continuance of the normal twining activity, and the
health and vigor of the plant is doubtless dependent upon
the successful manifestation of its habit of growth. Ex-
ternal conditions too have a decided influence upon the
behavior. The force of this statement will be evident as
the results of experiments are presented.

Plants of the species under investigation twined readily
around a rough or smooth stick, varying from one-quarter
to three-quarters of an inch in diameter; also around glass
rods, string, or wire, and if nothing else was available,
they utilized each other. Amphicarpcm twines in dextrose
fashion— against the sun.

DarwiuVs experiments recorded in - Climbing Plants,
though numerous and embracing many plants and repre-
sentatives of many orders, seem to the writer to ack cer-
tain essential requisites of information. Much of the move-
ment exhibited is unquestionably due to the annate char-
acteristics of the species. This again is modified by the
height and vigor of the individual under observation.

294 Schively — Contributions to the Life History of

These facts to a certain extent are considered by Dar-
win.'- He distinctly refers to the increased amplitude of
the circuninutation, noticeable by comparing the young
plant with the older stage, although the height of the
specimen is not always recorded in his tabulated results.

May not the varying periods of movement be due to the
environment? Is it not due to certain external conditions
that the manifestations of activity are now quickened, now
retarded ?

Various writers, among whom is Dr. Macfarlane,-"' have
shown in reference to experiments made upon Oxalis,
Cassia^ Mimosa^ etc., how important is the temperature
alone. Quick response to stimuli cannot be obtained, unless
care is taken to see that the thermometer registers from
75° to 105° F., according to the species. While this kind
of irritability is not here considered, we may rightly extend
the truth taught in this special case, to all phenomena
which may be classed under the manifestations of irrito-
contractility. To none are the above words more appro-
priate than the species now under consideration.

If we watch the tip of the long stem passing through
space, as if reaching for a support, we may actually see it
move. Hour after hour it proceeds steadily onward like
*' the hand of a clock." If we are fortunate enough to
see it touch a stick, we notice in a few hours a change
in the manifestation of its activity. It continues its course
now in a "beaten path," as it were, the influence of the
action still felt below, and being seen in the tightening
coils which clasp the stick.

If the support be removed, the upper part at least of the
plant, slowly straightens and repeats the same searching
movements. My reader may say, '' This is inherent in the
plant." True ; these statements have only been repeated
in order to make very plain the exceedingly active proto-
plasmic conditions that exist in a climbing or twining plant.
Then, how much more easily does the physiological
equilibrium change with the variations in environment.

Aniphicarpcra monoica.

295

Judging only from what has been learned of one plant, I
conclude that in addition to height of specimen, information
concerning temperature, conditions of moisture, exact time
of day at which the experiment was made, and also light
intensity should be stated.

A few items of detail may not be out of place here.
Amphicarpcea positively refused to circumnutate if exposed
to a temperature of 15° C. for any length of time. If re-
tained at such a temperature for a day or two, the effects of
the injurious environment remained for some hours after
the plant was placed under more favorable conditions.

On the other hand, a temperature of 38° C. or over seemed
to have a similar paralyzing effect. If supported, the plant
remained almost stationary, and often drooped. If free, the
movements were irregular and spasmodic, partly due to the
plant's adjusting its equilibrium, which is more or less dis-
turbed by the para-heliotropic position of the leaves. Upon
one occasion, a plant behaved as above indicated, tempera-
ture 40° C. ; it was carried into a cooler greenhouse, the
temperature there being 24° C. Apparently undisturbed
by being moved, in a few moments the plant recovered
from the heat, and proceeded in its usual circular path.

Many of the diagrams given by various authors to illus-
trate circumnutation, indicate that an erratic or sometimes
a sinuous course, has been taken by the plant. It is pos-
sible that these results are due to a temperature which is
too high for the species. The exceedingly long time
required by certain plants may partly be caused by an in-
juriously low temperature.

The most favorable temperature for AmphicarpcBa is
from 26° to 32° C. ; good results are obtained with the
thermometer above and below these figures, but the most
rapid circumnutation usually occurs within these limits.

If a plant is taken from the moist greenhouse and placed
in a dry room, marked retarding of movement occurs,
though the temperature is about the same.

Placing a lighted lamp or candle during night, before a

296 Schively—CoHtribntions to the Life History of

plant which has been circnmnutating well, though perhaps
slowly, will cause it to become stationary and remain so
for a time. The last two statements have not been put to
the test of many experiments.

Some of this work was done during the winter in the
greenhouse. Difficulties were encountered in the varying
temperature and moisture of the greenhouse, also the
supply of fresh air ; but on the whole, results were very
satisfactory. Experiments were continued during May,
June and July, when the greenhouse was not artificially
heated. Out of doors, even a slight wind was a disturbing
factor. The greenhouse being well ventilated, the tem-
perature varied little from that outside. As the glass above
was very lightly white-washed, the shade produced a con-
dition similar to that in which the plants normally grow.

Specimens under observation were placed as close as
possible (without touching) to the glass upon which records
were made. No special effort was made to obtain a con-
tinuous path traversed by the plant, though all records are
fairlv accurate. The time in which the revolution might
be completed was the information desired. Few observa-
tions were made as to the number of internodes circnmnu-
tating at one time. Two or three seem to be in activity at
the same time; no difference in the rates could be distin-
guished; the movement seemed uniform.

Plants under eight inches exhibited slow, irregular
movements, rarely forming a circle. When nine to ten
inches long the curving of the tip became more evident,
and from this time on, as the twining inclination increased,
the path described became circular. The rate of elongation
is rapid — from one to two inches a day.

There is a certain marked periodicity. Beginning with
the early hours of morning there is a gradual acceleration
until II or 11.30 a.m. The greatest rapidity occurs from
this time until 2 or even 3 p.m. After that time there is
a gradual decrease in the rate, until several hours after
midnight. The maximum period may be much extended,

Amphicarpoea monoica.

297

beginning earlier and continuing until 4 or perhaps 430
P.M., if the day is very hot.

The quickest rate obtained was 51 minutes; the longest
3 hours 36 minutes. In the former case, the specimen was
unsupported and was 14 iuches high. It was observed
upon February 8, the temperature being 26>^° C; it com-
pleted a circle 7% inches in diameter between 11.42 a.m.
and 12.33 P.M. In the latter case the specimen was 9
inches high. The observation was made upon June 30
between 1.32 a.m., and 5.08 a.m. The temperature was
i6°-i4° C, and the circle but 2>^ inches in diameter.
Both days were bright and clear.

It is true that the smaller plant had not yet exhibited a
twining inclination. But from the two extremes may be
gained some idea of the varying conditions under which
these experiments with Awphicarpcea have been performed.
From what I have observed, I am confident that so long a
time would not have been required by the second plant,
had the records been made under conditions similar to those
of the first. It may be best, therefore, to state the longest
time noted for an unsupported plant whose height was
twelve or fourteen inches.

A plant, 14 inches high on February 14, completed a
circle from 6 p.m. to 8.15 p.m., temperature 21° C. The
time was 2 hours 15 minutes. On July 2, another specimen
12 inches high, completed a circle in 2 hours and 10
minutes, the temperature being 23^ C. The time of day was

6 35 P.M. to 8.45 p. M.

The series which follows is selected as showing average
rates of circumnutation and also the periodicity to which al-
lusion has been made. The specimen was 22 inches high, and
supported, 9 inches being free; the observations were made
during February. Beginning at 9.30 a.m., a circle was
completed in i hour and 20 minutes; another immediately
followed and was completed at 12.01 P. m., in i hour and
10 minutes. An interval of a few moments unfortunately
occurred ; at 12.30 p.m., it was placed in position again, and

298 Schively — Contributions to the Life History of

each circle was completed at the expiration of i hour and
10 minutes. The next period began at 1.45 p.m., and was
completed in i hour and 20 minutes. The date was
February 5 ; weather was clear and sunny. The tempera-
ture was 26° during the greater part of the time; it began
to decrease during the last one recorded. The last circle
required 2 hours and i minute, and during this time water-
ing of greenhouse took place; the temperature having
fallen to 21° C. Observations ceased at 5.21 p.m.

A plant was observed June 20; the height was 16 inches
and it was not supported. From 6.21 a.m. to 8. 10 a.m.
a circle was completed in i hour and 49 minutes; tempera-
ture 23° C. At 9.13 A.M. observations began again; in i
hour and 27 minutes another circle was made, then fol-
lowed another in i hour and 7 minutes. The slight
increase of time required for the circle made from 11.49
A.M. to 12.59 P-^^-> ^s probably due to a fall of 11° in the
hygrometric conditions ; since at this part of the day, a
more rapid movement should have resulted. The tempera-
ture averaged 29° C. The circle was 12 inches in diame-
ter ; observations could not be continued, as the plant after
this, became caught in the frame. The main stem of this
plant had been truncated, and the actively circumnutating
shoot was one which had developed in the axil of a com-
pound leaf.

One plant on a hot July day, interested me greatly. It
was about 14 inches high; before daylight it required 2
hours and 15 minutes for a revolution, the temperature
being 21° C At 12.47 p.m., I observed it once more;
from that hour until 4-49 P-M., four large circles were
made; the first in 55 minutes, the remainder in 60 minutes
each. The temperature was very regular, 31° to 32° C,
but it fell a little, just before the close of the last period.

In some cases, I have been able to study one plant
through several successive days,. at least for a portion of
the time. Thus the varying phases of individuality,
growth, time of day and temperature, are more emphati-

Ai}iphicarpcea monoica.

299

cally shown. Watering the greenhouse always retarded
movement. In the middle of the day, I feel sure great
heat, and want of fresh air affected the rapidity, even when
moisture was sufficient.

The above may serve to indicate general conditions. For
further information, the reader is referred to the tables on
succeeding pages, where have been recorded the results of
some of the experiments with necessary details.

On July 22, a clear, sunny day, four plants were care-
fully observed in the greenhouse for twenty-four hours.

Plant I was .supported; it was 15 inches high, 8 inches
being free. At the close of the twenty-four hours the height
was itYi inches.

Plant 2 was unsupported. It was I2>^ inches at the be-
ginning and 15 inches high at the close of the series.

Plant 3 was supported. It was i8>< inches high on July
22, and 2\yz inches on July 23. Nine inches were free.
Plant 4 was unsupported. It measured xoYi inches at
the beginning, and I2>^ inches at the clo.se of the series.

The necessary handling for measuring, arranging, etc.,
probably affected the results of the first circumnutation.
The time records for the night hours show a gradual in-
crease until about 2.04 A.M.; after that a .steady decrease.
The temperature, it will be noticed, varied very little for
nearly twelve hours, and yet the length of time required
was longer at 3 a.m. than it was at 7 p.m. The free portions
of these plants frequently curved, so that the time of move-
ment was remarkably increased at some times, perhaps di-
minished at others. This is the only explanation of tho.se
results which tend to depart from the beautifully graded
series. The size of circle increased rapidly, with elonga-
tion of the stem, but as the preceding table gives an idea
of the varying sizes, together with the time required for
their completion, I have omitted those details here.

During the hours between 9.30 a.m. and 12, it will be
noticed that several results of less than an hour have been
obtained.

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302 Schively—Contrilmtwns to the Life History of

The results are given in separate series for each plant, so
that comparisons of time periods may readily be made.
Tlie reader will appreciate, too, the favorable character of
the temperature of the day and night, and the slight varia-
bility in the moisture of the air.

TABLE II.— PLANT I. !

Thermometer. ! Hygrometer. Time begun.

28— 29

29—31—29
29— 3(J

30—31
31 -28

28—26

25

25

24—23

22

22

22

22*

22

23
23-24

25—27

28 — 29

29
30

28—29

29—31-29

29—30

30

30
28 27
26—25

25—24
23—22

22

22

22

23—24

25
25—26

26—28

28—29

30

72—68
68—70

70

68-67

65—70

70—76

76—78—72

72

70

70

70

70

70

70

70
70

68
66- 6u

60—55
55

72-68
68—70
70—68
68—66
66

72-75
78—72
72—70

70

70

70

70

70

70

70
70—65
65-55

55

10.46 A.M.
12. 1 P.M.
1 P.M.
2.4 P.M.
3.7 P.M.

4.14 P.M.
5 22 P.M.
6.35 P.M.
7.55 P.M.

9.15 P.M.
10.50 P.M.
12.20 A.M.
2.3 A.M.
3.37 A.M.
5.20 A.M.
6.35 A.M.
7.52 A.M.
8.54 A M.
9.50 A.M.

10.45 A.M.

PLANT IL

10.47 A.M.
11.55 A.M.
1.2 P.M.
2.9 P.M.
3.20 P.M.
4.35 P.M.

5-55 PM.

7.32 P.M.
I 9.10 P.M.
' 10.55 P.M.

12.37 P.M.

2.25 P.M.

4.16 P.M.

6 P.M.

7.20 P.M.

8.37 P.M.

9.45 P.M.

10.40 P.M.

Time
completed.

12. 1
I
2.4

3.7

4.14

5.22

6.35

7-55

9-15
10.50

12.20

2.3

2.37

5.20

6.35
7-52
8.54
9-50
10.45
11.40

11.55
1.2

2.9
3.20

4-35

5-55
7-32
9.10

IU.55

12.37
2.25

4.16

6

7.20

8.37

9-45
10.40

11.40

Time
occupied.

hours, mill.
15
59
3

3

7
8

13

20

20

25
30
43
34
43
15
17
2

56

55
55

8

7

7
II

15
20

22

38
45
42
48

51
44
20

17
8

55
59

* Peculiar curving of free portion.

Amphicarpica monoica.

303

TABLE II.— PLANT IIL

Thermometer. Hygrometer.

28 —29

29—31—29
29—30

30

30
28—27
26-25
25—24
24—22

22

22

22

23—24
25—26
25—26
26-28
28—29—30

72—68
68—70

70— 68

68 66

66

72-75
78-72
72—70

70

70

70

70

70

70

70
70—65
65—55

Time begun.

Time
completed

1

1
10.48 A.M. j

12.15

12.15 P.M.

1.20

1.20 P.M.

2.37

2.37 P.M.

3.44

3.44 P.M.

4-58

4.58 P.M.

6.20

6.20 P.M.

7-50

7.50 P.M.

9.20

9.20 P.M.

10.55

10.55 P.M.

12.37

12.37 A M.

2.25

2.25 A.M.

4.10

4. 10 A.M.

5-55

5-55 A.M.

7.20

7.20 A.M.

8-55

8.55 A.M.

10

10 A.M.

11.6

Time
occupied.

hours, niin.
27-

55
17
17
14
32
30
30
35
4-*
48

45

45

25

35

5

6

Thermometer. Hygrometer.

28 — 29*

33— 37t

37t

34

33
29 — 27
27—25

24—23
23 — 22

22

22

22
23-24
25-28
28-31

33—34

72—68

45
45—50

50—55

55
55—72
72—70

70

70

70

70

70

70
68-65
50—55

55

PLANT IV.

'"\

Time

Time

Time begun.

1

completed.

occupied,
hours, tiiin

10.49 A.M.

II. 51

I 12

11.51 A.M.

1-3

I 12

1.3 P.M.

2.30

I 27

2.30 P.M.
3.41 P.M.
t^.I P.M.

3-41

5.'
6.24

I II
I 23
I 23

6.24 P.M.

8.5

I 43

8.5 P.M.

9-50

I 45

9.50 P.M.

11-45

1 I 55

11.45 P.M.
1.50 A.M.

1.50
3- 50

2 5

3.50 A.M.

5-35

« 45

' 5-35 A.M.

7-9

I 32

7.9 A.M.

8.34

I 25

8.U A.M.

9.41

I 7

9.41 A.M.

10.38

57

* Changed to another greenhouse.

t Irregular movement.

304 Schively— Contributions to the Life History of

For the species, the average time occupied is about i
hour aud 20 miuutes. Darwin'- records as his most rapid
circumnutatoi among climbing plants— ^rr//?^?////?//^— which
performed a revolution in i hour and 17 minutes. Amphi-
carpiva nwnoica must be awarded the palm ; as repeatedly
the results of i hour, and quite frequently 55 to 51 minutes
have been obtained.

Among tendrils, however, there are more rapid rates
reported by Darwin'l However, he does not note the
length of the circumnutating i)ortion. With Passiflora
-gracilis, circles are made in as short a period as 57 and 58
minutes ; others require longer time ; the average is a little
over an hour. A footnote on page 1 54 in "Climbing Plants"
calls attention to Passiflora accrifolia, whose tendrils ac-
cording to Gray complete a circle in 46 and even 38 >^

minutes.

Plants of Amphicarptm often exhibited a tendency to
curve the apex downward, forming almost a complete circle
with the part of the stem below. Observations were fre-
quently stopped by this occurrence. After remaining sta-
tionary, sometimes for several hours, the apex would
straighten out and continue its revolutions. No cause could
be assigned for this behavior. At times, the circumnutating
tip would describe a path, in which the dot recording the
place last occupied, would be internal to the continuation
of the circle, instead of in line with it.

The axillary shoots from simple and compound leaves
have both been studied. When examined in a normal un-
supported condition, they swayed to and fro in wide irregu-
lar ellipses, which rarely were complete in form. It was
not difficult to discover that circumnutation was very ac-
tive, particularly with shoots from the compound leaves.
As soon as they were six or eight inches in length, their
inclination to twine was manifest, and the apex visibly
moved through space. When gently supported, the results
were fairly rapid, but the circle was small. The shoots from
the simple leaves were not expected to show so much move-

Amphicarpica mouoica.

305

nient • but really there was little difference in the rapidity.
All fo'rmed irregular small circles in an hour or an hour
aud a quarter. I do not feel thoroughly satisfied with the
methods adopted in observing the axillary shoots, and I
trust in future studies to remedy these defects.

HlSTOLOGV OF THK STEM.

In creneral the stem exhibits the structure of a typical Dico-
tyledon. Plate XXIV, Fig. 1 , shows the transverse section of
a vervvoungstemof a plantabout six inches high; the epi-
dermal^ cells have the outer walls slightly cuticularized.
Numerous rows of cells constitute the cortex Next may
be distinguished one row of flattened, slightly thickened
cells which form the bundle sheath. About ten vascular
bundles surround the central fundamental tissue ; the pro-
toxvlem is well marked. Among the protoxylem cells
and spiral tracheae are found tannin canals. In the phloem
numerous sieve cells with comparatively large plates are
seen ; here, too, among the companion cells appear tannin
canals ; usually one in the phloem of each bundle. Small
areas of hard bast are seen forming in the region bordering
directly upon the bundle sheath. , t 1

lu sections of an older stem (Fig. 2), we find that the hard
bast has developed considerably, forming an almost complete
ring. The xylem appears to push down mto the phloem,
forcing it aside into two nearly distinct patches. Tannm
canals are now more nnmerous than in the young phloem
three or more being present in each area. From s udy of
longitudinal sections the component elements of the sec-
ondary xylem are found to be wood cells, pUted ve.ssel.s and
tracheids with reticulated walls. In the cells of the bun-
dle-sheath, just outside of the hard bast are found ?"«-
n,atic crvstals of similar appearance to those met w,th in
the leaf and other portions of the plant. Their appear-
ance will be described more minutely in later pages. At
present their position and development in the plant de-
inand attention.

3o6 Schwely — Contributions to the Life History of

If a transverse section is taken from the stem of a plant
abont fifteen inches high, there will be seen a single row of
cells in the inner cortex, in which may be distinguished
small crystals. Just how soon after germination this forma-
tion begins has not been definitely ascertained, but evi-
dently it occurs quite early in the plant's career. The in-
crease in size and number of crystals in these cells is rapid.
Finally a distinct crystal sheath (Plate XXIV., Fig. 4,) sur-
rounds the vascular area. It is considered that this crystal
sheath is equivalent to the original bundle sheath; the
position occupied by the former seems to correspond ex-
actly to that of the latter. These crystals seem to be
surrounded by a protoplasmic film of considerable thick-
ness, as this area stains very deeply. This condition is
plainly shown in a section taken from a plant stem three
months old (Fig. 2). In Fig. 3, is seen a transverse
section of a stem, a little older than the preceding,
but which, having grown during the summer, was much
more vigorous. There is a marked increase in the
xylem and also in the phloem. The hard bast appears to
have undergone some transformation, very few of the cells
are visible and these are in scattered groups. The crystal
cells referred to above have greatly increased in number,
and are abundantly present in the outer expanded ends of
the medullary rays. As yet no reagent that will dissolve
the crystals has been found ; but it is possible that treat-
ment by several chemicals in succession will accomplish it.
A transverse section of stem of a plant raised from an aerial
seed shows, when three months old, a condition similar to
that seen in Plate XXIV., Fig. 2 ; but the diameter of the
section is about \i of that represented in Fig. 2.

Yet one more feature of interest is found in the section
Fig. 3, when studied with high power. In the xylem and
to some extent in the phloem, are noticed sack-like bodies
whose contents are refractive, and gelatinous in appearance
— some protoplasmic substance possibly : Chlor-iodide of
zinc colors these a deep purple-red : iodine test gives no

Ainphicarpcea monoica.

307

reaction. No starch is found in the section, nor have tannin
canals been located.

During the life of this plant some marked physiological
changes seem to occur, the character of which is not yet
understood. Nor can it yet be stated whether there is any
connection between fruit production and the progress of the
changes or not. Further investigation may elucidate the
meaning of what has been observed.

General Appearance oe Leaves.

As has already been stated, the first green leaves of
AmphicarpcBa are simple and opposite. They are netted-
Veined, broadly ovate, with rounded apex, hairy upon both
surfaces, and also upon the margin, which is entire ; the
stipules are interpetiolar. At the base of the petiole, and
also at the base of the blade is a pulvinus. The petiole is
about one and a half times the length of the blade, and
both it and the pulvini are retrorsely hairy. The remain-
ing leaves are compound, provided with stipules, and are
arranged alternately. They are pinnately trifoliate, petio-
late and stipellate. The primary pulvinus is usually very
well developed, being at least four times the size of those
belonging to the leaflets. The terminal leaflet possesses a
petiole of its own— a continuation of the main one, also a
pulvinus and two stipels. The petiole of this leaflet makes,
with that of the main leaf, an obtuse angle ; thus its blade
lies in the same plane as the remaining leaflets. Each
lateral leaflet is attached by a pulvinus only, and is pro-
vided with a single stipel. All leaflets are ovate, with an
acute apex, the blade being longer than those of the simple
leaves ; but the terminal is larger than its fellows. The
lateral leaflets are rarely symmetrical, the outer side being
nuich wider. In other characteristics the compound resem-
ble the simple leaves. All leaves are remarkably thin, and
of delicate texture. The petiole forms with the stem an

3o8 Schivcly — Coniributions to the Life History of

angle of 4S° to 6o^ The pulvini of the leaflets stand up
at an angle of 30° with the petiole, so that the surface of
the blades lie about in the same plane.

Histology of the Leaf.

The epidermal cells are irregular, stomata occur upon
both surfaces of the leaf, but are much more numerous
upon the lower than upou the upper. Large firm unicel-
lular hairs are placed at regular intervals upon both sur-
faces and are exceedingly numerous upon the margins, par-
ticularly of voung leaves. These hairs are thick-walled
and bear tubercle-like markings. On the midvein, on
souie of the larger veins, and on the margin are found small
bladder-like hairs.

The vascular bundles branch extensively; but frequeutly
there is no evidence of anastomosing. This is often seen
with the more delicate branches, which suddenly terminate
in the mesophyll. Embedded in the larger vascular areas
(probably in all, however) have been seen crystals similar
to those occurring in the clearly defined crystal sheath,
previously described in the stem. Possibly, however,
because they are not so massed up on one another, their
distinct twin-like structure is much more easily distin-
guished, the partition, whether apparent or real, beiug
easily located. (Plate XXIV., Fig. 5.)

The transverse section shows thin upper and lower
epidermal layers, while the mesophyll consists of irregular,
but tolerably uniform cells, so that a differentiated palisade

tissue is absent.

Simple and compound leaves correspond in the above

details.

Leaf Positions.

In common with many other members of the Leguminosae,
Amphicarpcca is exceedingly susceptible to changes in tem-
perature, moisture and light. Because of the delicate foliage,
many difficulties arise, and constant watchfulness is neces-
sary to explain the condition of each plant.

AnipJiicarp(ea monoica.

309

a. Normal Position.— \r\ diffuse daylight, in early morn-
ing, or when shaded from the sun's rays, both simple and
compound leaves assume a position which will here be
alluded to as ''normal." The dorsiventral relation is
shown ; leaflets present a flat position, though occasionally
the terminal is slightly elevated. The angle of the petiole
with the main stem varies from 45° to 60°.

/;. Paraheliotropic /^^.s7//V?//.— If exposed to direct action from
the sun's rays or to a strong reflection, the lateral leaflets
turn the outer edges gradually upward, the upper surface
being inclined toward the terminal leaflet. As the intensity
of heat and light increases, the pulvini are stimulated more
and more, the angle assumed by the margin is 90° with a line
drawn through the surfaces in their normal position. The
terminal pulvinus rises about 45°, thus elevating the leaflet.
The lateral pulvini rotate the leaflets at this stage; but
there is little change in the angle they make with the main

petiole.

A further increase of the sunVs action causes the lateral
leaflets to incline more toward the terminal, which rises usu-
ally 90"" ; frequently the upper surfaces of the leaflets are in
direct contact, the under surfaces being outward. The
lateral pulvini have risen slightly ; but their angles are
difficult to estimate, as these blades have been so much
rotated. (Plate XXV, Fig. i.)

Often the terminal does not assume a perpendicular
position, but the apex points either right or left. It is
noticeable too, that the terminal leaflets on opposite sides
of the plant do not incline in the same direction, even if
the sun's rays are vertical.

A series of questions which must yet remain unanswered
are suggested. Are these movements of advantage to the
plant ? Are the pulvini protected ? Is too rapid transpira-
tion retarded ? Are the movements due to heat, or to cer-
tain rays of light?

For the present, a partial answer at least may be given
regarding the last. Plants were studied in the greenhouse

3IO Schively — Contributions to the Life History of

where the temperature often was 90° F. While there was
no sunlight, para-heliotropic positions were not assumed.
Certain it is that sunlight has a direct influence ; it has
been found that leaflets toward the east, for instance, may
be visibly affected by the rays falling upon that side of the
plant, while those toward the west still remain normal.

A plant whose leaves have taken the para-heliotropic
position, if shaded, or lifted to a shady spot, will show the
ordinary position in from three to seven minutes.

The simple leaves rise slowly ; in a very young plant
they rise 90° and bend over toward the tender shoot. (Plate
XXV, Fig. 2.) In older plants they rarely rise more

than 75°.

In all of the above movements, the angle of the petiole
seems unchanged ; leaflets and their pulvini alone being
concerned.

A very hot sun causes the under surface to turn com-
pletely uppermost, the leaflets appear limp and droop. In
some species of plants, where this condition is seen, the
blades are firm at least when they are handled ; but in
AmpJiicarpc2a^ I think the expression used, best describes
their peculiar appearance.

This movement is not apparent in all the leaves, usually
only the three or four nearest the upper part of the plant.
This behavior was noticeable upon hot days in the green-
house as well as out of doors. Macfiirlane'"^' first drew
attention to the above condition, as occurring in numerous
other plants, some of which are not at all sensitive accord-
ing to the ordinary application of this word. After the
intensity of illumination subsides, the leaflets gradually
assume the para-heliotropic position, or may even return to
the normal without the above transition.

c. Nyctitropic position. In preparing to assume the nyc-
titropic position, the terminal leaflet falls slowly, the lateral
do likewise, gradually turning their upper surfaces toward
the terminal leaflet at an angle of 45°. The complete de-
flexing may not take place until an hour after preparations

Amphicarpo'a monoica.

311

have evidently begun. Sudden changes in environment may
accelerate movements, or indeed may cause plants to take the
sleep position even in the day-time. Carrying a plant from
the moist atmosphere of the green-house into the dry air of a
heated room having about the same temperature, produced
a shock, which resulted in the assuming of the nyctitropic
position. Often complete recovery did not take place until
the plant was returned to congenial, moist surroundings, or
some time thereafter.

Exactly what may be designated as a complete nycti-
tropic position is still perhaps a matter of doubt. About
sunset the following condition may be seen : The leaflets
have fallen 90°, the terminal one points perpendicularly
downward or inclines in the direction of the petiole, the
surfaces of the laterals approach ; often the apices touch
each other.

Simple leaves also deflex 90°. When the plant is quite
young, after thus deflexing, the blade turns and lies with its
under surface below the petiole, nearly parallel with it, and
almost in contact with it. Thus, at this time these leaves
pass through 180°. (Plate XXV, Fig. 3.) This remark-
able movement disappears in plants having four or five
compound leaves, and frequently before this number is
reached.

Within two hours after all leaflets have assumed the posi-
tion described above, the terminal one rises slightly, the pul-
vinus forming an acute angle with the petiole. The blade
stands out, as shifting toward the right or left has taken
place. Gradually, though constantly, the leaflet turns its
upper surface, the pulvinus changes its angle, the leaflet
inclines again toward the perpendicular until about 1 1 p.m.;
as a result of this continuous movement, the upper sur-
face of the terminal leaflet lies in the same plane as the
surface of a lateral leaflet, and faces outward (that is to-
ward the observer or directly away from him— according
to his position in relation to the plant). (Plate XXVI.)
The petiolar angle most commonly assumed is nearly

312 Schivcly— Contributions to the Life History of

90° ; obtuse and acute angles have also been noted. Out
of seventy-five leaves upon eight plants, the petioles of
eleven showed obtuse angles, six, acute, and the remainder
were right angles. Upon one plant two different angles may
be observed ; usually, however, uniformity exists. Certain
leaves showing each of the angles were marked and
watched upon several successive nights; each night the
customary noted angle was regularly assumed. There
seems to be no reason for this difference; but in the numer-
ous sets of plants studied, it was always seen. The right
angle, then, appears to be the prevailing one. As the
normal leaf position showed an angle varying from 45° to
60°, there would here be a f^ill of 45° to 30°.

Darwin'' states that the petiole may fall through about 57°.
From the facts given, he nmst have studied one which as-
sumed an obtuse angle. No others are mentioned by him.
Those which did assume this angle seemed to the writer
to fall much more than the number of degrees estimated
by Darwin; certainly 75° is nearer the correct amount.

In the passage above referred to, the circumnutation of
the leaf was traced by Darwin". He is not himself, satis-
fied with the method employed, but his object was, he
states, ''to ascertain if the leaf moved after it had gone to
sleep.'* A diagram of his results is given, but they do not
give an exact idea of what may be observed during the night
hours. First, the apparatus used would seem to me to
have a deflexing effect upon the entire leaf. Secondly, it
records movements of the terminal leaflet only, and even
the results for it are questionable. Thirdly, his observa-
tions ceased at 10.50 P.M. What is to be seen after that
hour will be described presently. However, it may be
indicated that so great and varied is the activity of leaflets,
that no idea is given by the tracing nor by the description
in Darwin's work.

The movements until n p.m. have already been de-
scribed. From that hour till daylight, leaflets are con-
stantly in motion. Observations were made every half-

Aniphicarpcea monoica.

313

hour. Those recorded below were made during the last
week of June and early in July. I cannot speak exactly
for night movements earlier in the year, but the behavior
at that season until 10 p.m. would indicate during the re-
maining hours decided similarity to that described for the
summer months. Between 11.30 p.m. and 2 a.m. the
terminal leaflet re-assumed gradually a nearly perpendicular
position, turning again, so that the margin instead of the
surface was more in the plane of the surface of the lat-
erals, but it still stood out from the remaining leaflets, the
apex inclining away from the petiole. The laterals had
rotated upon the pulvini, so that their upper surfaces in-
clined somewhat inward.

Between 2 and 3 a.m. the terminal rose slightly, and the
laterals shifted so that their upper surfaces faced in the
normal direction. All leaflets were deflexed about 45'', but
the spreading apart of leaflets was marked.

From 3 A.M. the general attitude of a plant might be de-
scribed as that of expectation. About 4 A.M. the petioles
had arisen, and the day position was rapidly being as-
sumed. By 4.30 some few leaflets had completely ex-
panded; many at 4.45 a.m.; nearly all at 5 a.m. These
records are of green-house plants, temperature 2o°-25° C. ;
the hygrometer, Ss^-QO"^. During this month the sun rose

at 4.35 a.m.

Those out of doors varied greatly— clouds, winds, low
temperature retarded the assuming of the day position, al-
though during the night the leaflets showed similar
activity to that just described (unless it was very windy).
On a moderately warm morning, some were found in day
position at 5 A.M.; on a cool morning there was a delay
until 6 A.M., and even later.

In this connection it may not be out of place to allude to
the Mimosas which were in the green-house. While Am-
phicarpcea was in agitation half the night, the leaflets of
Mimosa were quietly folded. When the time arrived for
the day position, the leaflets expanded in a very short time,

314 Schivcl) — Contributions to the Life History of

no indications of movement being visible before 4.45.
Mimosa assumed day position about a half hour later than
A mphicarpcea.

Observations were made to ascertain what relation the
time of assuming nyctitropic position bore to the hour of
sunset. Until the early part of May, records were made
from plants in the green-house only ; after that time com-
parisons could be made with those growing in the Botanic
Garden. During the time the green-house was artificially
heated, the temperature frequently fell late in the after-
noon, and, as the final watering for the day occurred about
that time, the hygrometer record was high.

Previous to April no data of value were obtained. Dur-
ing that month, upon clear, warm afternoons, the nycti-
tropic position was assumed about nine minutes before
sunset (6.24-6.54 P.M.).

During May, ten to twelve minutes before sunset (6.55-
7.21 P.M.) was the general time. May, 1896, was a most
unfavorable month, as there were but few clear, warm days.
After that— in June and July— plants in the green-house
were often completely nyctitropic, fifteen or twenty min-
utes before those in the garden, which quite often, although
showing indications of nyctitropic position for some min-
utes, did not assume it completely until sunset hour or a
minute or two before (7.30 p.m.).

These differences may be attributed to the following cir-
cumstances :— the humidity of the green-house, and close-
ness of the air, and comparatively, the more rapid approach
of darkness, caused by .shading of the greenhouse by certain
adjoining walls. In June the position of the sun was such
that these conditions were quite noticeable. Those out of
doors had the advantage of all the light at that time as

well as fresh air.

Dark, rainy days caused rapid assuming of the nycti-
tropic position from thirty to fifty minutes previous to

sunset.

On such days, the plants out of doors gave no exact

Amphicarpcea monoica.

315

results as to time of taking nyctitropic position, for usually
the leaves remained more or less deflexed during the entire
day. Even on a clear day, a brisk wind or a fall in tem-
perature would induce the nyctitropic position at an earlier

hour.

Electrical disturbances of the atmosphere seem to have an
effect upon AmpJiicarpcea. On one hot July day a heavy
thunder-storm occurred about 5 p.m. ; the clouds were
unusually heavy. Within an hour, the sky had cleared,
and the sun was shining brightly. Yet the plants did not
recover their normal position, but retained the nyctitropic,
which had been assumed on the approach of the storm.
This statement applies equally well to those plants which
had not suffered from the force of the rain. The sunset
hour was 7.30.

So many conflicting conditions make it difficult to state
a general rule. As far as I am able to gather from these ob-
servations, there seems to be but a slight difference between
the time of taking nyctitropic position and sunset hour in
the summer months. On the other hand in the winter
season, the leaflets close from fifteen minutes to a half hour
before sunset. The rapid approach of darkness in the
latter case; the long twilight in the preceding certainly
form important factors.

Plants in an open space, upon a clear warm day, are late
in taking the nyctitropic position — just about the .sunset
time. But this statement can not be correct for the indi-
viduals growing in the woods. From the few studied in
their native haunts, the period in August (sunset about
7,) is from fifteen to twenty minutes earlier than this hour.

The above facts lead to the conclusion that there is not
a definite time-relation between the time of nyctitropic
position and the exact time at which the sun sets.

PuLviNus Structure

The pulvini are much more pubescent than the other
portions of the petiole. Both the long, unicellular hair and
21

3 16 Schively—Contributiojis to the Life History of

the bladder form are abundant; the former is tuberculate

in character.

In a transverse section of the piilvinns the flattened por-
tion indicates the dorsal region. The epidermis is strongly
cuticularized ; in the many celled cortex there are few
chlorophyll granules. Numerous crystals of the character
previously mentioned as occurring in the stem are found ;
the number is variable, but nearly all exhibit the twm
arrangement more plainly than is usually seen in other
parts of the tissue. In the secondary and tertiary pulvmi
the crystals are comparatively more numerous.

In the fundamental tissue, lying next the central mass,
the cells are similar, and possess somewhat thicker walls.

The cells of the bundle-sheath resemble in appearance
those in the centre of the pulvinus, and show clear collen-
chymatous walls. Chlor-iodide of Zinc produces a violet
coloration. Next to the sheath lies the phloem ; then the
xylem which occupies a relatively large area. Chlorophyll
is very abundant just within the bundle sheath.

All the usual reagents reveal the presence of tannin cells

in the xylem. .

Continuity of protoplasm is best demonstrated in the
phloem region and the central mass of collenchyma.

Clear refractive globules have been observed in the
cortex ; these do not consist of oil, nor of tannin, for treat-
ment with special reagents to demonstrate the presence of
either substance, yields no satisfactory results. Their com-
position has not yet been ascertained. They are not always
seen, and from a few observations which have been made,
I think their presence has some connection with the posi-
tion of the leaf at the time when the study is made ; but
whether they are most abundant during the normal, the
para-heliotropic or the nyctitropic position can not yet be

definitely stated. ,

These globules which appear to be absent in the secon-
dary and tertiary pulvini, may be distinguished m the
primary.

Amphicarpi^a monoica^

3^7

Flowers of Amphicarp^a.

Before stating observations as to the flowers and fruit of
Amphicarpcea^ it may be well to give a brief outline of the
facts to be found in the principal Botanical Manuals.

lu the '' Botany of the vSoutli," Elliott^ describes pale pur-
ple flowers found in racemes. These are couiplete, but he
says they are ** generally sterile." Indeed when, in his
report to the Academy of Natural Sciences, he describes
the shape and appearance of the legume, he quotes Walter
as his authority, stating that he himself had never seen one.

Peduncles from the root bear flowers without petals*
Near the surface of the earth racemes are produced, the
flowers of which are furnished with a calyx and rudiments
of a style. "The fruit here" he says, "is a one-seeded
ovate pod."

The preceding account refers to A. monoica. In addi-
tion he mentions A. sarmentosa. It bears filiform racemes
which are three-flowered and apetalous. The calyx alone is
described. The fruit is an oblong pod.

Darlington ^ agrees with the above statements in regard
to complete flowers, and also as to fruit. On the radical
peduncles he finds apetalous flowers, which " are often
merely pistillate." Peduncles arise from the base of the
stem ; and a solitary legume develops at the extremity.

In the " Flora of North America "* (Torrey and Gray) the
descriptions are much more detailed and indicate more
careful observation of the floral parts. The purple flowers
are described, the shape of legume also, though no reference
is made as to the fertility of the flower.

The imperfect flowers are located as the preceding
authors have doue. It is stated, however, "Stamens want-
ing or often five or ten, shorter than the ovary ; three or
four with perfect anthers, the others rudimentary— the
filaments are distinct. The ovary is nearly sessile, tipped
with a short recurved style. The legume is obovate, hairy,
one-seeded, usually maturing below the surface of the
ground."

3 18 Schively— Contributions to the Life History of

The accounts given by Wood, Chapman, and Gray^=^ are
similar to those given above; but none are so minutely
descriptive as that of Torrey and Gray.

In a communication to the Academy of Natural Sciences
by Meehan'^ quite a different statement is made. The
purple flowers, the imperfect flowers above alluded to, and
the fruit in each case are described. A legume differing in
shape from that usually described was noticed by him and
figured. The following is quoted from the paper: ''In
what may be termed the more vigorous racemes, the two
lowermost flowers, either have but a small vexilhnn pro-
jecting beyond the calyx, or none. The next half dozen
flowers are perfect in every respect and fall without perfect-
ing a legume. The apetalous flowers can scarcely be classed
as ' cleistogamous,' for there is no pollen. A few unde-
veloped stamens are found here and there. In absence of
positive demonstration, I should regard these as pistillate
flowers receiving pollen from the petaliferous one."

Summing up the previous statements, it is found that
there is a general agreement regarding the structure of the
purple flowers, and their position on the plant. Their
fertility, however, is questioned.

There is no unanimity as to the structure of the imperfect
flowers. No doubt seems to exist as to their presence upon
the plant, but the location is not definitely stated.

Excluding the observations of Meehan, and for the
present, Elliott's statement regarding A. sarmentosa, there
have been given but two kinds of flowers, and the same

number of pods.

AnipJiicarpica monoica, however, possesses no less than
four distinct varieties of flower, and as many legumes. These
may be enumerated as a, b, c and d. (a) Evident aerial
flowers of purple color may fairly be distinguished as
the normal type, (b) Aerial cleistogamous flowers, possess-
ing no corolla or a very rudimentary one, are intermediate
between the purple aerial and the subterranean, {c) The
subterranean is much reduced in structure, and may be

Amphicarpcea monoiea.

3^9

regarded as derived from the normal through {b). {d) This
form is produced during the winter months, and is a com-
pound of {b) and {c)\ it might be called a transition type.

Purple Aerial Flowers.

The lavender purple, often almost white flowers are pro-
duced in pendulous simple, occasionally compound, racemes.
These are borne in the axils of cauliue leaves (from about
half the plant's height upward), and also upon the upper axil-
lary shoots. Buds appear during the last week of July and
bloom from August lo into September. These statements
are made from observations in the neighborhood of Phila-
delphia. At Woods HoU, Massachusetts, blooming occurred
from five to eight days later. The number in a raceme
varies from ten to twenty-four, occasionally more. There
are frequently two buds, certainly one, in the axil of a
broadly ovate, partly clasping, striate, pubescent bract.
(Plate XXVII., Fig. i, 2).

Careful study has led to the conclusion that there are orig-
inally two buds in each axil. Very young as well as more
advanced racemes showed this condition. As elongation
takes place, some of the buds fall off; frequently when most
of the flowers are in bloom, there is but one in many of the
axils. No histological investigation has yet been completed
regarding this. It may be proper to state here, that if this
supposition is correct, there may be some foundation for the
statement given in Torrey and Gray that these bracts are
formed by a " union of a pair." However, it is elsewhere
stated by these authors in regard to the imperfect flowers that
the bracts are distinct. No bracts are found with these latter
flowers; the stipules certainly have been mistaken for them.
Therefore the previous statement would not be of value.

Some observations in reference to the number of buds
occurring in the axil of each bract of the raceme have been
recorded as follows : —

320 ScJiively — Contributions to the Life History of

a

a

o

I

3
4
5

6

7
8

9

lO

II
\i

13
M
15
i6

17

i8

19

20
21
22

2a

24

35
26

29

30

31
3a

33

34

35
36

%

39
40

^

'I
E2

flj !/5

W

{:

8
II
10

7

13

2

4
11
12

II

7

15
20

20

II

II

«7
14
14

13

16

f 22

22
20
10

17

14
16

II
15

be

C o

S >. X

8
10

8

6
10
10
10
10
12
10

6

14

18
20
10
10
16
12

14
10

16

18

12

22

20

8

16

12

14

8

12

7

0

6

2

15

10

13

10

18

14

«3

10

17

16

II

10

10

8

10

8

13

10

9

8

u

S !/3 A

o
I

2
I

3
2

4
I
o
I
I
I
2
o
I
I
I

2

O

3
o

4
o

o

o

2
I
2
2

3

7
4
5
3
4

3

I
I

2
2

3

Remarks*

Young.

Young.
j Little older.

Much older — still unopened.

Lower three had opened.
{Compound raceme, lower
S flowers had opened.

Young,
i Young.

Nearly all open.
i Young.
I Young.
! Young.
I Young.
! Young.
j Young.
\ Young.
I Young.
' Young.

Lower open.

Lower.

y Compound raceme.

I Young.
I Young.
' Young.

Young.
j Young.

Young.

Young.

)^ A branch and a bud in the axil
I j of I bract.

Young.

I I Compound raceme.

i Young.

Young.

j j Lower bract has in axil 2 buds
I 1 and a branch bearing 2 buds.

Young.
I Young.

Young.

Young.

Branch and bud in i bract axil.

Youne.

Amphicarpcva monoica.

321

More observations are needed on this point. At present
it cannot be stated in which portion of the raceme, rednc-
tion is most likely to take place, as the blooming season ap-
proaches.

The seasonal conditions and the general environment
exert a marked influence upon the number of purple
flowers produced. A hot, dry season is very unfavorable.
The summer of '95 showed but few racemes; in '96 the
plants were covered with them. In the shaded portion of
the woods purple flowers are rarely produced; on the other
hand, too much sun does not seem favorable. Amphicarpcea^
grown in the Botanic Garden in two places where there was
constant sunlight, showed but a very few flowers, although
the plants were provided with a fair supply of water.

The calyx is greenish-white, if the corolla is pale, other-
wise purplish- white, flecked with deep purple; in both cases
it is pubescent. It is four-parted, the teeth acuminate and
dissimilar. Thus far nothing has been seen to indicate the
presence of a fifth vascular bundle. It is possible in
younger buds, some evidence may yet be found. In the bud
the calyx parts are valvate, the tips being spirally twisted.
The calyx is about half the length of the corolla, it is
swollen at the base in the posterior region, and, as the
flower-parts expand, the appearance is that of inflation. It
persists at the base of the legume.

The corolla is pale purple, but may vary to pure white.
The aestivation is that of a typical member of the Papilio-

nacere.

The anterior margins of the carina cohere but for a short
distance. The upper portion of each is shaped like a
rounded triangle, and narrows abruptly into a long slender
claw. In the inferior region of the triangle occurs a marked
depression. The ahe are rather oblong in the upper por-
tion, otherwise they resemble the carinal members in
general outline. In each of these is seen an invagination
pointing posteriorly; the floral parts are so arranged that
the pouches thus formed upon carina and alae inter-digitate,

II

322 Schivcly — Contributions to the Life History of

and thus these petals are held firmly together as described
by Miiller for other forms. On the superior margin of
the broad portion of each wing occurs a sack-like process
or short spur whose opening is upon the exterior. The
curious auricled appearance of the petal is due to the
presence of this contrivance.

In the bud the vcxilliim encloses the remaining petals
almost completely, but, as the flower expands, it is pushed
posteriorly. This petal is obovate, tapering gradually ; there
is no true claw. It is marked transversely about half way
down, by an irregular, imperfectly semi-circular band of
deep purple hue, which may possibly be interpreted as a
"path-finder.'' The most deeply pointed portion of the
calyx is opposite the anterior petals (carina); the remaining
three are around the standard.

The stamens are ten in number, five long, five short ;
they are diadelphous and the insertion is perigynous. The
staminal tube is united for three-fourths of its length ; the
superior stamen is free. The anthers are two-celled, small,
almost spherical, versatile and introrse ; they are of a
deep orange color, and w^ell filled with pollen. In young
flower-buds, the anthers are found to be well-developed,
but the filaments are short ; for some time the style and
stigma extend for a distance above the stamens.

Plate XXVII, Fig. 3, is a transverse section through a
bud, showing the anther lobes and a portion of the connec-
tives of the ^v^ shorter stamens, also filaments of the five

longer.

In a transverse section of an anther belonging to a
purple flower, the following condition is seen : the epider-
mis (exothecium of authors) is a single row of suiall deli-
cate cells ; the hypodermal layer (endothecium of authors)
consists of nuich thickened, colunmar cells.

In developing anthers the pollen is normal. Some, per-
haps all, of the anthers dehisce when the bud is about one-
half inch long, and showing no indications of unfolding.
At that time the pollen is pale yellow, powdery, and under

Aniphicarpcea monoica.

323

the microscope appears as a shriveled husk. Upon the
addition of water it becomes slightly granular, and swells
immediately. Differentiation into exospore and endospore
is noticeable now, and a small quantity of oil is seen. The
size of the pollen grain will be discussed later on by study
of a comparative series.

The monocarpellary //^///arises from a slender gynophore.
The ovary is free, superior, unilocular, and contains from two
to four ovules. The style in a mature state is exceedingly
loug, the stigma small, capitate, and quite hairy. Micro-
scopic examination reveals the presence of beautiful tufts
of hairs surrounding the base of a rounded surface, smooth
and having a rather sharp contour; this is evidently the
stigiuatic area. The hairs project outwards and upwards.
The margins of the ovary are clothed with long unicellular
hairs, also small bladder hairs. The latter are numerous
upon the style. Stomata are present upon both style and

ovary.

Extending around the carpel and passing up both dorsal
and ventral sutures onward into the style are strongly
marked bundles. Their behavior with reagents, when the
carpel is entire or sectioned, leads to the conclusion that
tannin canals are located here.

Histological study has also demonstrated the nature of
"the sheath at the base of ovary " mentioned by various
authors. It is without doubt a nectary. A transverse sec-
tion of this is seen in Plate XXVIIL, Fig. i, to possess ten
vascular bundles. The upper portion, not figured here,
possesses numerous pits or cells of glandular appearance,
which are particularly noticeable around the free margin
of the structure.

In a young bud, the style is short, hooked, and the
stigma though capitate, shows an undeveloped condition of
the hairs, which are as yet closely appressed. For some
time after, while the style is elongating, the bent appear-
ance is retained, and thus the stigma inclines downward.
Finally the style straightens and also the stigma, the brush

324 Schivcly — Contributions to the Life History of

of hairs having gradually assumed the characteristic
appearance. The early condition of style and stigma is
seen in Plate XXXL, Fig. 8, and the later one in Fig. 9.
These drawings are made to the same scale.

Fertilization.

It will be seen from the previous description of the struc-
ture of Amphicarpcca that the purple flowers are well pro-
vided with those devices which are associated with insect
fertilization. It is probable that this is not the method
here, though there are strong grounds for believing that
the plant may be a descendant of types that were insect-
pollinated. If insects visit these flowers, the writer has not
seen them, nor has any indication of such visitors been
discovered.

As has already been stated, in the tiny bud, not more
than one-quarter of an inch in length, the style has already
grown some distance beyond the stamens, and curves the
stigma toward them. After this both stamens and pistil
elongate and finally are about the same length, the stigma
protruding a short distance beyond the anthers. Upon
opening a bud about a half inch long, but that is still
closed, it will be found that the anthers have dehisced, and
usually some pollen is found upon the stigma.

It is quite certain that the majority of the purple flowers
do not produce fruit; a number, however, are productive.
Perhaps the stigma is not in condition when the pollen is
mature. Should this be the case, the question naturally
arises, how is fertilization accomplished in any case? F'or
the present it can only be stated that it is possible that the
ten stamens mav reach maturitv at different times. It is
also suspected that there may be some explanation yet to
be discovered in the long-continued curving of the style
towards the stamens.

* Amphicarpma monoica.

325

Aerial Clkistogamous Flowers.

m

Not until the former blooms are fairly well developed is
this variety seen. They appear in the axils of cauline
leaves, lower down than the purple, and also upon numer-
ous axillary shoots. Frequently they are solitary ; sometimes
they are found in a short, closely clustered few-flowered
rudimentary raceme. Occasionally one or two flowers
develop at the base of the purple raceme. Again they
are found upon a long shoot resembling that upon which
the subterranean pods develop — that is, we see a slender
axillary branch upon which at intervals, occur solitary
legumes, often to the number of three or four. This
special shoot frequently branches and occasionally appears
to terminate in the production of two pods growing from
opposite sides of the apex. The last described structures
will be discussed in connection with the underground type

of legume.

When once recognized, the two varieties of aerial flower
need never be confused, either in flower or fruit.

The calyx is four-parted, as in the preceding, but is
smaller and the teeth are not so pointed nor so long. It is
pubescent, but the color is always greenish white.

Even the tiny bud, an eighth of an inch in length, pos-
sesses a flattened appearance quite different from that of the
purple one.

Upon dissection a comparatively large ovary is seen,
bearing recurved style, who.se stigma is not usually capi-
tate. This ovary is nearly sessile, and is somewhat hairy,
particularly upon both anterior and posterior margins.

Occasionally a rudimentary vexillum is present, but
practically the flower is apetalous.

The stamens are typically ten, and show a small filament-
tube, or they may be distinct. They are quite small and all
transitions of perfection in the anthers exist. Plate XXIX,
Fig. I, shows the pistil surrounded by the anthers, vary-
ing in the degree of perfection indicated. In Fig. 2, a

326 ScJiively — Contributions to the Life History of

well-developed anther, found in this kind of flower, is
much magnified.

The rapidity with which these flowers pass into fruit is
quite startling. One must examine a very small bud to
ascertain that fertilization takes place early, that the anthers
do not dehisce, but send forth numerous pollen tubes to-
ward the stigma. The anthers then shrivel, the ovary
presses out of the calyx and within a few days a good sized
legume has developed.

The study of these microscopic flowers has been rewarded
by the discovery of a beautiful series of transitions in the
shape, size and structure of the style and stigma. This is
probably a suitable place in which to allude to them,
although it may be anticipating slightly. The green
aerial flowers without doubt are reduced purple forms. The
styles and stigmas, shown in the nine drawings (Plates XXX,
XXXI) are enlarged in the same proportion. In each form
represented here, pollen tubes were seen passing toward
the stigma, if not already in contact with it, and there is
no doubt that the series figured in these Plates represents
a set of mature organs.

Fig. 1 (Plate XXX) belongs to a terrestrial flower, and
is yet to be described, but is here mentioned to show a very
rudimentary condition. Fig. 2 is occasionally found in the
green aerial, but belongs especially to the underground form.
In Figs. 3-7 inclusive, green aerial styles only are represent-
ed. The curving and extension downward is marked, but in
Figs. 3, 4, 5, there is no indication of a capitate stigma.
Along the inner surface, just beyond the rounded apex of
some of these, are seen, by careful focusing, thickened areas
where the cells appear different from those of neighboring
parts. Here is probably the stigmatic surface. Although
there is a great difference in the aspect of Figs. 6 and 7,
we have present the brush of hairs, still however, appressed.
Figs. 8 and 9 have previously been mentioned ; they be-
long to the purple flower and complete the series drawn.
The immature specimen in Fig. 8 bears a striking rcsem-

Amphicarpcea vionoiea.

327

blance to the mature one in Fig. 7. In Fig. 9 the style
has the characteristic erect position and the perfect capitate

stigma.

Plate XXVIII, Fig. 2, is a transverse section through the
ovary of the green aerial flower. The macrosporangium,
with the enclosed macrospore, may be readily studied.
Other sections which were examined showed variation in
height and perfection of the stamens.

These flowers have been seen upon plants bearing purple
blooms, but appear about a month later. Some of the.se
were in the green-house, others out of doors, alike in the
Botanic Garden and in the country. As late as the first
week in October, the aerial green flowers weYe found in bud
condition. There is no doubt they would have matured
fruit, had the weather continued favorable.

Plants grown in poor soil, or where there was too nuich
sun, or in too much shade, never produced any but cleisto-
gamous aerial flowers.

Meehan^''' calls attention to the two kinds of aerial
flowers ; but he does not describe their location upon the
plant, nor their structure correctly. He regards the apeta-
lous flowers as pistillate, ''receiving pollen from petalife-
rous blossoms." He denies the presence of pollen in these
imperfect flowers, and therefore does not wish to call them
cleistogamic. He does not note the time when this
latter type appears.

Subterranean Flowers.

Some weeks before AmphicarpcEa blossoms above ground,
the cotyledonary axillary runners have already formed
buds, it has been already stated that the runners from
the axils of simple leaves are geotropic, and the secondary
branches upon the.se also exhibit the same physiological
characteristic. While the cotyledonary shoots mentioned
are colorless, those from the simple leaves frequently are
deeper purple-green than the remainder of the plant. As
they lie upon the ground, often the main branch thick-

I

328 Schivcly—Contributions to the Life History of

ens for a distance of an inch or two behind the tip, some-
times becoming etiolated ; tips of the secondary branches
were often beautifnlly cnrved downward.

Before discussing the peculiarities of growth manifested
by these runners, it may be well to describe the flower
borne by them. The writer does not consider that there is
a raceme-like arrangement of the inflorescence, although
there seems to be ; the plan of inflorescence is somewhat
puzzling. Below each flower is a pair of stipules, never a
bract. It is easier to explain the arrangement by referring
first to one of the runners which we find arising from the
axil of the simple leaves. Now, this shoot grows to a
great length, ptoducing leaves with stipules at their bases,
frequently each leaf is reduced to a pair of stipules.
Each new axil possesses the possibilities of one or more
buds, each of which may be similar to the parent. In this
part of the plant the reduction of the leaves becomes the
rule, and in the axil of the stipules appears a small branch
terminating in a flower-bud. Here may arise two— even
three— branches (rarely at the same time), each bearing
flower-buds. The main shoot may increase in length in-
definitely, and likewise branches may develop from the
apical buds hidden between the stipules at the base of the
flower-bearing branch. It may be said that the flowers are
solitary ; and when occasionally two are found terminating
a shoot, it may be inferred that in the axils of that special
pair of stipules, two flower-bearing branches arose at the
same time. This condition of affairs is more easily studied
when the fruit begins to form. It often happens in the
subterranean flowers that the buds are very close together.
This is because great reduction in length of the shoot has
taken place; but the stipules may usually be found even
then, and are always readily seen upon the longer cotyledo-

nary branches.

The calyx is four-parted, but the teeth are more rounded
than in eidier of those described. As yet no evidence of
an original five-parted calyx has been seen. In this case

AnipJiicarpcca monoica.

329

very young flowers have been studied, for it is only when
in that condition that the facts which follow may be dis-
covered. These flowers are apetalous.

The stamens are distinct, the number varying from six
to two. From two to four bear fertile anthers, all others
are rudimentary. The filament is short, the anther flat-
tened, and fairly well-filled with pale yellow pollen. Often
a stamen is seen, bearing but one productive anther lobe.
In Plate XXIX., Fig. 4) is represented a series, show-
ing the variations in the character of the stamens. The
first is but a filament, in the second the microsporangia
are clearly defined, but no microspores are developed. In
the third, only one anther-lobe is perfect, while in the
fourth, the normal condition is seen.

Transverse sections made through the same flower-bud
at difl'erent levels are very instructive. That seen in Plate
XXVIII., Fig. 3, is taken from a tip which had not yet suc-
ceeded in concealing itself in a dark place. The ten fila-
ments are seen in section, but the ovary has not been
reached in cutting.

Examination of other sections show that the number of
filaments becomes gradually less. No one of the other
sections belonging to this series showed more than three
perfect anthers.

There is one carpel; the ovary is nearly oblong and ses-
sile, possessing long hairs upon the sutures; it is also
finely pubescent, though this condition is much better seen
after it emerges from the calyx. It is usually one-, but
often two-ovuled. The style is short, curved, but extends
outward and very slightly downward ; the stigma is not
capitate.

The relation of pistil and stamens is indicated in Plate

XXIX., F'ig. 3.

When fertilization takes place the flowers are about a
millimeter in length, or even less. Considerable care is
required to obtain just the right stage in which the pollen
tubes may be observed passing from the fertile stamens

\

W

330 Schivcly — Contributions to the Life History of

which are located just below the style. Sometimes a stamen,
with a well-formed anther is found upon the other side of
the ovary away from the style. It is probable that this does
not take part in the work of fertilization.

Subterranean fruits may result from the flowers originally
formed under ground, and from others, which result from
flowers produced by shoots above ground, and eventually
covered by soil, or succeed in reaching a dark place where
thev mature fruit.

Darwin '' (page 503), refers to the actual penetration of the
flower bearing apex into the soil. Repeated observations
have been made, but the writer has not been able to satisfy
herself that this occurs unaided. In the green-house, a thick
layer of loose soil was placed over the cinders covering the
shelf, but the shoots simply spread over the surface, some-
times for a distance of several feet. If a small hole were
made a short distance from the tip of a shoot, it would soon
turn down into it.

Often a favorable spot for concealment would be under a
flower pot. In their efforts to seek darkness, some runners
extended over the sides of the shelf, upon which they were
placed in the greenhouse, and continued to grow to the
floor, a distance of at least four feet. They matured fruit
here, some hiding among cinders ; upon others soil was
placed. If the tips do not find a place suited to their liking,
they either dry up or produce a tiny flat green legume.

It seems probable, that in nature, earthworms assist in
burying the flowers; there are, too, crevices, stones, or
leaves which afford the desired protection ; the beating of the
rain, perhaps, may cause the tip to become covered. Having
once secured a proper place, growth in length will continue,
for some time, even in the soil, if that is loose in character.
Indications of the presence of subterranean flowers, pro-
duced upon cotyledonary shoots, may be seen on a plant
six or eight weeks after its germination. Those upon the
over-ground branches are much later in appearing. The pos-
sibilities of production upon the former seem unlimited. As

Ainphicarpita nionoica.

331

lonf>- as the plant exists, development continues ; for branch-
in«- and re-branching do not cease. As late as October, all
stao-es may be found from the tiny bud to the large legume.
In Auo-iist, a number of shoots were taken from plants
growing in a moist spot, with plenty of dead leaves lying
upon the ground. Most of these were cotyledonary shoots,
but some may have been runners from the simple leaves,
which long ago had disappeared. The number of buds and
small legumes then present were as follows : thirteen, six-
teen, twenty-two, forty, fifteen, thirty, twenty-seven, twenty-
one, twelve, seventeen, respectively. An astonishing series,
truly ; it is probable, too, that most would have matured fruit
Observations were made to ascertain approximately the
length of time necessary for the maturing of these legumes.

On February 15, an axillary runner from the first pair
of leaves was buried. It was i8>^ inches long, and was
buried for about half its length. It bore one compound leaf,
a small runner in the axil of this leaf, and a runner nearer
the plant, in the axil of stipules. Each of these had two
tiny buds. Beyond these the main runner divided into
two.

On February 22, the ruUner was 19 inches— there was

no apparent change in buds.

On February 29, the runner was 19^^ inches long and
the changes in pods were as follows :

Pod 3 was ^ inch long, and ^ inch in circumference,
and was becoming rounded.

Pod 4 was Yj^ inch long, and was flat.

On March 3, pod 3 was ]/z inch long, and quite rounded.
Pod 4 was ^ inch long — also becoming rounded. The
others had been injured in an unexplained manner.

On March 14, pod 3 was ^ inch long; pod 4, ^ inch;
both quite full and round.

On March 7, an axillary runner 30 inches long, from the
first pair of leaves, was buried for half its length. It bore
22

ff

332 Sc /lively — Contributions to the Life History of

four compound leaves, and in the axil of each a small branch
having minute buds. Beyond the last leaf was a shoot
9 inches long, bearing three buds.

On March 14, the shoot was 32 inches long, and there
was a very slight increase in size of buds.

On March 21, the length of shoot was 3234 inches.
From the axils of the second and third leaves there were two
axillary buds each. That from the fourth leaf had two tiny
flat green legumes y^ inch long. Upon the terminal portion,
pod I was % inch long, 2, -{^ inch, 3, j/s inch long ; all
were still rather flat.

These and similar notes seem to indicate that the matur-
ing of the terrestrial legume is comparatively rapid. In
the second, certainly the third week, after fertilization, they
assume good proportions. After this, the increase is very
steady, and growth would be likely to continue during the
life of the plant, though the seed is probably mature early.

Then came the question : Will shoots from other leaves
than this first green pair, produce these subterranean
legumes?

On July 23, shoots from simple leaves, also from first,
second, third, and fourth compound leaves were buried in
separate pots. All were long, bore leaves, but- as yet
showed no indication of flower buds.

Circumstances prevented examination until September
3. Finally all were removed September 25. Every one of
the shoots had branched extensively both inside and out-
side of the pots. The tips which were thin and green
when placed in position had thickened, formed, even roots
had developed. Legumes of all sizes were here ; well-
developed ones one-half to three-quarters of an inch long
were taken from each pot, in number from two to six;
while smaller ones were numerous. In regard to produc-
tiveness of axillary shoots, those from simple leaves gave
the greatest yield ; those from compound leaves differed
little in fruitfulness ; not one failed to develop some.

Ainphicarp(Ea monoica.
Structure of Winter Flowers,

333

During the winter of 1895-96 plants raised from ter-
restrial seeds produced minute green flowers which re-
sembled the subterranean flowers previously described.
They are indeed almost counterparts of those found in
sunnner upon the tips and upon the branches of axillary
runners, and which do not become covered with soil.
They differ in the resulting legume, as well as in their
physiological behavior. While the summer ones, fail-
ing to obtain a suitable spot for development, either
produced a tiny, flat, one-seeded legume, or else dried up, the
winter-type seemed unaffected by its surroundings and
usually matured a good-sized two-seeded legume. On the
other hand, this winter-form might be regarded as a degen-
erate type of the green aerial flower of late summer, as it
is nuich reduced in several particulars.

The calyx is about the same size as that found in the
terrestrial flower, is more pointed in its lobes, but is gen-
erally much larger than that of the green aerial. The
corolla is entirely absent. Ten stamens are generally pres-
ent; they are distinct. Four of these are unusually fertile,
the remainder being mere outgrowths from the receptacle.
The pistil has a short curved style, the stigma is not capi-
tate; the ovary is sessile and typically two-ovuled. The
stamens in number suggest resemblance to the green
aerial, while the appearance of style and stigma causes
our thoughts to turn to the terrestrial, for no such varia-
tions of style and stigma as previously described for the
green flowers of summer are seen here.

Plants raised from aerial seeds during winter rarely pro-
duced any fruit above ground ; if they did, it was a tiny,
flat, one-seeded legume, of the character alluded to in a
previous paragraph. An additional reason this, for consid-
ering the winter type of flower to be most closely related to
the terrestrial form.

Within four weeks after germination, flowers were evi-

334 ScJiivdy — Contributions to the IJfe History of

dent ; legumes were ripe at the end of two months. Not
the slightest indication of purple flowers was seen upon
any plants during the winter and spring.

Summary of Views on Floral Variation.

To recapitulate, Amphicarpcea presents the case of a plant
bearing at least three ^ probably yO//r distinct types of flower ;
and these not occasionally, nor spasmodically, but simply
following regular laws of development.

In summer, purple flowers may be expected to appear upon
the upper main stem, and the upper axillary branches. A
certain intensity of the sun's rays seems necessary, for
these are not found upon plants growing entirely in the
shade. Neither do they develop extensively upon plants
constantly exposed to the sun. Here the water supply may
be insufficient ; plants grown in such a situation are not apt
to reach a fair height.

But later the same plant, having, we will suppose, as is
often the case, produced an abundance of purple flowers,
now proceeds for a month or more to bear aboveground a
cleistogamous form.

Frequently, too, plants which have not developed a single
colored flower, may, late in the season, produce this last

type.

Similar cases of this method of development are not un-
known. Among the Violacece it is quite common, but all
of the cleistogamic forms are produced close to the ground,
hiding among the leaves, and only occasionally upon spe-
cial short branches.

Speciilaria perfoliata bears inconspicuous flowers late in
the spring ; in June the bright purple ones are found.

Henslow'^ mentions the finding of flowers of Linaria
and Potcntilla in the autumn, in which he discovered pol-
len tubes passing to the stigma. These specimens, though
imperfect, however, still possessed color.

Amphicarpcea certainly bears its cleistogamic flowers in
great abundance, some of them in a very conspicuous posi-

Amphicari>cca inonoica.

335

tion upon the plant. The aerial cleistogamic form occurs
persistently and for a long period.

As for the nnderground flowers, from the middle of June
until October, unceasingly Amphicarpcea is adding to the
number. No average environment seems to prevent their
production : but there is marked difference in the size and
number of legumes produced.

Among the Oxalidaceai the special underground shoots
do not develop until the summer months are far advanced.

EpipJicgiis Virginiana shows a similar behavior to Amphi-
carpcea. During the summer of 1896 Dr. Macfarlane ob-
served at Woods Hole Polygala polygama, which grows
there abundantly. Early in the summer he found this
plant to produce subterranean flowers, many of which ripen
fruit before the aerial flowers have opened. After the
purple flowers have almost ceased blooming, colorless
flowers appear above ground, upon elongated shoots which
develop below those bearing purple flowers. A study of
these is now being made by Mr. C. H. Shaw.

Is the behavior of Amphicarpcea in winter not due to
absence of intensity in the sun's rays? High temperature
and abundant light and moisture were provided, still plants
in the green-house from January to end of April showed
no purple blossoms. Yet others placed in the same part of
the green-house in May were covered with these flowers

in August.

Linuc-eus relates a similar experience with plants taken
to the Gardens of Upsal '•' ; they produced inconspicuous
flowers only. Gray,' too, reports similar experiences with
certain plants in the Cambridge Garden. Insufficient tem-
perature might be adduced as a cause for this non-produc-
tion of colored blooms; but from my experience with
Amphicarpcea the light intensity is without doubt an im-
portant factor.

1

I

33^ Schively — Contributions to the Life History of

Microspores and Macrosik>res.

Little has been said concerning the pollen; it seemed
best to compare all flowers in this particular at one time.
When young the anthers of the purple fl'omers are quite
well filled. The appearance of pollen after the anthers
have dehisced has already been described. Dr. Macfarlane in-
forms me that it resembles the inefficacious pollen of certain
hybrids. It measures when dry, 13 fi, upon addition of
water it measures 16-18 /i.

The anthers of the gi-ecti aerial flower do not contain so
much pollen as those of the purple flower ; it resembles in
appearance the pollen of the latter wlien water has been
added. It is pale yellow, granular, and measures i J /i.

The pollen of the terrestrial flozoers is less abundant
than that of the last; but the appearance is similar; it
measures 8-9 /i. Two nuclei are frequently seen in this
pollen grain. (Plate XXIX., Fig. 5.)

I can not state so certainly the comparative size of the
macrospores, as I do not feel sure that those measured were
absolutely at the same stage — namely, ready for fertiliza-
tion. From specimens which have been examined, the
macrospore in the purple floiver measures 5 /i ; that of the
green aerial 6 /i, and that of the terrestrial 4 /i.

Before turning attention to the legumes, it may not be
out of place to refer to the comparative size of the pistils
of the three types of flowers, as shown in Plate XXXII.,
Figs. 2, 3 and 4.

Histology of Legumes.

The macroscopic appearances of the legumes have already
been described, and reference to illustrations will convince
the reader that it is not easy to mistake the products of the
different flowers. It is possibly not scientifically correct to
apply the term ^'legume*' to the fruit of the terrestrial
flower, as there is no dehiscence ; but considering it as a
modification of the others, the expression may, perhaps, be

AinpJiicarpaia monoica.

337

allowed to stand. For the present, laying aside the winter
type, Amphicarpcea fuonoica presents in nature, three dis-
tinct kinds of legume. A striking series of transitions is
readily made from the purple to the terrestrial, including
those tiny flat forms which are not successful in reaching a
suitable maturing place. If again we place the winter-type
in our list, the series is unusually complete.

With the purple, and green aerial, the fruit is green, turn-
ing brown when ripe, the style persisting and forming a
characteristic feature in each, as does also the region just
toward the apex. In the winter type, the same remarks as
to color change will apply; usually the style disappears; if
it remains, it either stands out at right angles to the legume,
or it is appressed to the ventral suture.

The terrestrial form is white, but is soon tinged with
pink purple. When ripe, the style has disappeared, the
sutures are often scarcely discernible, the color is purple of
varying vividness and density.

Appropriating a word used— possibly introduced— by
Huth,'^ we may properly class Amphicarpcea monoica among
hitevocajpic plants. In his paper, however, he places it
among ampJiicarpic forms.

The histological structure of purple, green aerial and
winter-type legumes is similar. Numerous stomata alike
in size and shape are found; these are surrounded by several
subsidiary cells. The epidermal cells are irregularly isodi-
ametric. ^ (Plate XXXIV., Fig. i.) Just beneath the epi-
dermis, lie one or sometimes more layers of sclerenchymatous
fibres, closely arranged, tapering so that they fit together to
form a mechanical device for producing the inflation of the
pod. In transverse section, in addition to what has been
mentioned, we find cells resembling mesophyll and con-
taining chlorophyll. The inner epidermis is thin, and has
no stomata. The glistening, somewhat opaque appearance
of the interior of the legume is probably due to air spaces
under the epidermis. The mesophyll tissue in the winter
type appears of looser texture than the others.

338 Schivcly — Contributions to the Life History of

When young, all are somewhat pubescent, but the purple,
and green aerial when mature, bear hairs upon the sutures
only; those elsewhere have disappeared. The winter- type
possesses these epidermal structures through life and also
when ripe. Both long unicellular hairs and the bladder
forms, previously described, occur here.

The terrestrial form presents several peculiar charac-
teristics. The purple coloring is due as elsewhere in the
plant, to a liquid either in the epidermal cells or just below
them. Stomata are quite as numerous comparatively, as
upon the surface of the overground legumes, but they are
here placed upon the apex of a small papilla raised con-
spicuously above the surrounding epidermal cells, which
resemble in size and shape those of the other legumes
(Plate XXXIV., Fig. 4). Certainly three kinds of hair
occur ; one of these has a base suggestive of a glandular
function. The others are of the same character as those
seen upon the aerial legumes, but are much more numer-
ous. A fourth kind may be the shorter unicellular seen
in the drawing. (Figs. 2, 3)

But instead of these walls being firm and unresisting,
they are thin and delicate. No sclerenchymatous fibres
exist, but there is present the parenchyma corresponding
to mesophyll in texture, whose cells also contain chloro-
phyll. The seeds occupy all the space within the legume,
and the result of the tension is seen by tiie final thin cover-
ing, as increase in size of the seed takes place.

Remembering the great contrast in legumes produced
upon the lower axillary runners, due apparently entirely
to the special environmental conditions under which they
were allowed to mature fruit, an experiment was undertaken.
Some of the winter aerial legumes (not located upon the
lower runners) were buried, after they had become quite long,
but were still flat, the seeds being small and green. At
intervals during the course of a month they were exam-
ined ; at the end of that time a remarkable transformation
had taken place— the counterpart of a terrestrial presented

Aniphicarpcca monoica.

339

itself. The whole structure had swollen enormously ; its
thickness was four times that of its former state ; its color
was now pink purple. Instead of firm walls, their appear-
ance was now thin and tightly stretched ; even the color of
the seed-coats had changed. Various accidents destroyed
some of these fruits before the observations were completed.

Experiments are now being carried on to ascertain more
regarding this remarkable physiological change, and also
what histological changes, if any, result. Winter forms in
varying stages of development, and from all portions of the
plant, have been buried in specially prepared receptacles,
containing soil and sphagnum, and these have been sus-
pended from the roof of the green-house.

It is hoped, too, that some information may be obtained
as to the possible function of the long hairs with glandular
base, whose presence was mentioned above.

As soon as possible a similar series of experiments will
be tried with legumes of purple, and green aerial flowers.

{a) Legumes from Aerial Purple Flowers.

In order to ascertain the amount of fruit resulting from
purple flowers, racemes were tagged while yet in the
bud; others were gathered and examined after legumes
had developed, or were mature. Several localities were
then studied, and some of the records are given below. As
there is at least one flower in the axil of every bract,
results are fairly accurate. It is not possible to state the
definite position upon the raceme, where fruit may occur.
This as well as the number is exceedingly variable.

1

340 Schivcly — Contributions to the Life History of

Aniphicarpcea nionoica.

341

Observations upon Plants Growing near Burmont

AND LANSDOWNE.

Observations upon Plants Growing along Wissa-
HicKON near Chestnut Hill.

1

4-1

1

owers

a

Number of fl
iu raceme.

5c
v4

Remarks.

Number of fl
in raceme.

Legumes pre

Remarks.

1
II

I

24

4

12

I

18

4

12

2

44

8

Compound raceme.

18

4

15

0

22

3

Compound raceme.

18

0

12

I

14

0

8

I

II

I

»7

I

6

I

20

8

Compound raceme.

13

2

25

I

14

2

9

I

6

3

9

2

9

8

18

I

18

3

II

^

J

30

4

Compound raceme.

10

3

13

5

10

4

24

4

17

3

18

4

II

I

10

2

16

3

20

4

Compound raceme.

17

4

Compound raceme.

21

6

12

i 3

20

5

60

j I

Compound raceme.

28

5

Compound raceme.

12

3

^

13

3

14

5

Compound raceme.

13

5

17

3

19

3

22

10

II

2

16

4

10

0

18

9

27

0

II

8

13

0

20

7

««

;

l-i

u

>

:'

0

«c .

•

v-li

;v

0 c

i,.
it.

24

22

9

23
24

II

12

5

7
10

II

5
17

a
(/)

a.

IU

IU

•4

Remarks.

O
O

o
o
o

9
8

2

5

9
6

3
II

Compound raceme.

t

4.!

a

« .

4;

a

fc J{

I

^n

a

§8

&

^•^

1;

5z;

H^

10

6

10

2

7

5

12

5

6

4

7

3

20

6

7

5

13

6

20

7

6

5

14

3

Remarks.

Many other racemes noticed in the latter locality yielded
no legnnies.

These statistics are qnite snflficient to prove that Amphi-
carpcea nionoica produces legumes from purple flowers in as
large proportion as the Leguminosa,- generally, though not
at all equal to some well-known genera. As is the case
with all plants, however, some years yield better returns
than others. The legumes are neither few nor difficult to
find. Certainly the statements found in all the Manuals
concerning the usually sterile condition of the flowers, are
erroneous.

(b) Legumes from Other Flowers.

Rarely do the green aerial (b) fail to produce fruit, these
exceptions being due to the appearance of the flowers so
late in the season, that the temperature does not permit
their maturing, ic) For the terrestrial flowers the results
are practically as given above.

!i

342 ScJdvdy — Contributions to the Life History of

Comparative Productiveness of Flowers.
The following records show the comparative productive-
ness of all these flowers, as obtained from plants cultivated
in the green- house in pots during summer. They were
twenty in number and placed some distance apart. Any
intertwining shoots were carefully separated; but the results
are not absolutely accurate in the case of the number of
terrestrial legumes for each plant, as many which were fair-
sized, though still flat, were dried by the heat in the green-
house in the latter part of September. Many of the runners
had no means of hiding the flower-bearing portion, except
among the cinders, and it was impossible to avoid breaking
these from their attachment, when pots were lifted. The
numbers, however, will give an idea of the possibilities of
productiveness in the case of single plants; this could not
be ascertained from growth in the woods.

The number of flowers producing terrestrial legumes is
not given, nearly always there were numerous tiny ones
which were yet undeveloped ; mature fruits only were con-
sidered.

Plants were placed in position late in May and early in
June ; examination of results was made in the last days of
September. Many specimens reached a height of five feet,
and developed axillary shoots abundantly, some of the latter
being six and eight feet in length.

Plant I. was raised from a terrestrial seed produced upon
a small (non-twining) plant in late winter.

It bore 66 purple flowers resulting in 3 legumes.

14 green aerial flowers resulting in 14 legumes.
7 -I- terrestrial flowers resulting in 7 legumes—

2 of them large.

Plant II. was raised from a terrestrial seed produced upon
a small (non-twining) plant late in the winter.
It bore 66 purple flowers resulting in 3 legumes.

14 green aerial flowers resulting in 14 legumes.
6 -I- terrestrial flowers resulting in 6 legumes—

3 of them large.

Amphicarpcea monoica.

343

Plant III. was similar to I and II. It bore

30 purple flowers resulting in o legumes.
13 green aerial flowers resulting in 13 legumes.
3 -f terrestrial flowers resulting in 3 legumes —
2 of them large.
Plant IV. was similar to III. It bore

38 purple flowers resulting in o legumes.
12 green aerial resulting in 12 legumes.
3 i terrestrial flowers resulting in 3 legumes.
Plant V. was produced from seed of a winter type legume.
It was rather feeble, and bore

o purple flowers resulting in o legumes.
8 green aerial flowers resulting in 8 legumes.
6 -f- terrestrial flowers resulting in 6 medium
sized legumes.

Plant VI. was similar to V., and bore

o.purple flowers resulting in o legumes.
16 green aerial flowers resulting in 16 legumes.

5 + terrestrial flowers resulting in 5 legumes ; i
of them large.

Plant VII. was similar to V. and VI., but was stronger, it

bore

6 purple flowers resulting in i legume.

21 green aerial flowers resulting in 21 legumes.
10 + terrestrial flowers resulting in 10 legumes
(small and medium).
Plant VIII. was produced from terrestrial seed upon a
plant raised from terrestrial seed. It bore

73 purple flowers resulting in o legumes.
8 green aerial resulting in 8 legumes.

7 4- terrestrial resulting in 7 legumes, small and

medium.

Plant IX. was similar to VII. It bore

4 purple flowers resulting in i legume.
24 green aerial flowers resulting in 24 legumes.
6 -V terrestrial flowers resulting in 6 legumes.

4f|
^1

344 Schivcly — Contributions to the Life History of

Plant X. was similar to IX. It bore

(Number lost) purple flowers resulting in i legume.
26 green aerial flowers resulting in 26 legumes.
18 -f terrestrial flowers resulting in 18 legumes, 5
of tliem large.

Plant XI. was produced from an aerial seed gathered in
'95, whether it was from purple or green aerial flower was
not known. It bore

43 purple flowers resulting in 2 legumes.
30 green aerial flowers resulting in 30 legumes.
14 + terrestrial resulting in 14 legumes, 3 of
them large.

Plant XII. was similar to XI. It bore

1 1 purple flowers resulting in i legume.

25 green flowers resulting in 25 legumes.

1 1 -F terrestrial flowers resulting in 1 1 legumes.

Plant XIII. was similar to XI. It bore

13 purple flowers resulting in i legume.

1 3 green aerial flowers resulting in 1 3 legumes.

13 -\- terrestrial flowers resulting in 13 legumes.

Plant XIV. was similar to XI. It bore

17 purple flowers resulting in i legume.
23 green aerial flowers resulting in 23 legumes.
13 -| terrestrial flowers resulting in 13 legumes.

Plant XV. was produced from a terrestrial seed raised
upon plant from growth of terrestrial seed. It bore
64 purple flowers resulting in o legumes.
37 green aerial flowers resulting in 37 legumes.
13 -f- terrestrial flowers resulting in 13 legumes
(small).

Plant XVI. was the same as XV. It bore

165 purple flowers resulting in o legumes.
29 green aerial flowers resulting in 29 legumes.
7 + terrestrial flowers resulting in 7 legumes.

Amphicarpcea monoica.

345

Plant XVII. was the satne as XV. It bore

90 purple flowers resulting in o legumes.
37 green aerial flowers resulting in 37 legumes.
18 + terrestrial flowers resulting in 18 legumes.

Plant XVIII. was the same as XV. It bore

o purple flowers resulting in o legumes.
31 green aerial flowers resulting in 31 legumes.
28 + terrestrial flowers resulting in 28 legumes.

Plant XIX. was the same as XV. It bore

232 purple flowers resulting in o legumes.
36 green aerial flowers resulting in 36 legumes.
12 + terrestrial flowers resulting in 12 legumes.

Plant XX. was the same as XV. It bore

40 purple flowers resulting in i legume.

21 green aerial flowers resulting in 21 legumes.

30 -f terrestrial flowers resulting in 30 legumes.

The underground legumes were then collected from the
pots and stage cinders in their immediate vicinity. As it was
almost impossible to avoid breaking some of their attach-
ments, the space was carefully examined and yielded ninety
legumes of varying size. Thus from these twenty plants
the grand total of terrestrial legumes was three hundred

and twenty.

In a plot of ground where plants were separated as much
as possible, there was a large yield of terrestrial legumes,
but comparatively few aerial, none of which resulted from
purple flowers. Although the soil was quite rich, it was
constantly exposed to sun, and moisture was not abundant.
All of the plants were low, wound somewhat around each
other, but rather trailed on the ground. This may have
been partly due to the winds which blew quite usually over
the special part of the Garden where these grew.

Three very vigorous plants deserve notice, and were so
arranged that each could be lifted out with the mass of soil
still clinging around the roots. Careful examination

346 ScJiivclf— Contributions to the Life History of

revealed the remarkable results of twenty-nine, thirty-five
and fifty legumes respectively. These were developed fiiirly
close to the original cotyledonary region. Axillary shoots
could not well produce fruit, as the soil was rather clayey
on the surface, and no leaves lay upon it.

As the legumes were being collected from a certain
plant, the curious branched appearance of the cotyledonary
axillary runners attracted attention. Investigation proved
the presence of no less than two hundred and fifty-one
hypogean flowers and legumes in varying stages of devel-
opment. (Plate XXXVI, Fig. i).

An interesting incident which occurred while raising
Amphicarpica in a small city yard, is not without value
here. In the autumn the space four feet by two which had
been used for this purpose was examined carefully, and
a number of terrestrial legumes gathered. So diligently
was search made, that the writer felt sure that all were
removed. Imagine then the surprise, when, in the following
spring, one hundred and fifteen plants appeared. The ex-
planaUon was that the bed in which they were grown was
beside a board fence underneath which the runners had
matured the fruits. An asphalt pavement in the next
yard prevented their growth in length. Several of the
plants were examined as to place of origin, and showed a
long white, rather crooked stem which had struggled
upward. It was not easy to find the seed, only its evident
location could be determined.

One certainly doubts the possibility of such a species
being exterminated.

Attention should be called, however, to one more set of
statistics. On April 18, plants which had been growing
since the beginning of February in particularly rich wood
soil, which had been kept very moist, were uprooted, and
yielded as recorded below. The only aerial fruit was the
winter type previously referred to.

Amphicarpcea monoica.

IA7

fi7 a
I 3 u

aerial.
Plant i. -( , underground.

2 aerial.
Plant 3' i 3 terrestrial.

20 aerial.
Plant 5- i 2 terrestrial.

Plant. 2.

Plant 4.

Plant 6.

Plant 7.

Plant 9.

'21 aerial.
5 terrestrial, 2 of these
two-seeded.

20 aerial.
8 terrestrial , 2 of these
two-seeded.

Plant 8.

Plant 10.

Plant ii.

Plant 13.

7 aerial.

6 terrestrial, 2 of these
two-seeded.

10 aerial.

1 1 terrestrial, 4 of these

two-seeded.

13 aerial.

I perfectly formed ter-
restrial.

Indications of 24 others
Yi inch long.

20 aerial, mainly one-
.seeded.
I terrestrial.

5 aerial.

9 terrestrial.

10 aerial.

5 below, I of these
two-seeded.

1

Plant 12.

Plant 14-

10 aerial.

6 terrestrial, i of these
two-seeded.

10 aerial.
7 terrestrial , 2 of these
two-seeded.

7 aerial.

6 terrestrial , 2 of these
two-seeded.

Total of terrestrial legumes (perfectly developed) 64, of
which 16 were two-seeded. In Plate XXXV, Fig. 3> a view
of the cotyledonary region of Plant 8 in this series may be

seen.

Having now presented some notion of the relative fer-
tility of each kind of flower, several conclusions may be

stated.

I. Plants oi Amphicarpcea growing under suitable condi-

23

348

Schively — Coiitrilmtions to the Life History of

tions of sunshine and shade, and having abundant moisture,
will produce all three varieties of flower. Tall, vigorous
plants, with such environment, seem to excel in quantity
of purple flowers.

2. Plants growing in almost perpetual shade, or those
exposed to constant sunshine, even if well supplied with
moisture, rarely produce purple flowers, and but few green
aerial ones. The subterranean fruits are quite abundant ;
but the quality and quantity depend upon the character of
the soil. Loose, rich, forest soil, with many decaying
leaves, gives excellent results, for the terrestrial fruits may
then develop from overground as well as underground
runners.

Along the banks of the Wissahickon, not far from Chest-
nut Hill, lies a certain strip of land about a quarter of a
mile in extent. It is an open space, not shaded by trees,
and is a perfectly luxuriant mass of vegetation, abounding
in tall weeds of various kinds, also a few shrubs. The
stream is narrow, and the high banks upon the opposite
side give a due amount of shade in the afternoon. The
soil is loose, very wet and sandy. Plants of AiNphicarpmi
monoica growing here are most vigorous specimens, rising
to the height of six and eight feet, and are densely covered
with ferruginous hairs. The best supply of purple flowers
was found here, and the racemes were often compound.
Strange to say the underground legumes were compara-
tively few, and nio.st of them small.

Does, then, the number of purple flowers affect the pro-
duction of underground cleistogamic ones ? Or is it due
to the character of the soil ? It has also been noticed that
tho.se plants giving abundant results in the way of subter-
ranean fruits, possessed more tubercles upon the roots.
The.se were quite insignificant, both as to quantity and size
upon the Wissahickon plants. Whether this is a mere coin-
cidence, or whether there is a real physiological connection,
can not now be definitely decided.

One need, however, pass but a short distance up the

Amphicarpica mofwica.

349

rocky hillside, covered with a dense growth of trees, to
find in certain localities, plants of A^nphicarpiua in abun-
dance, twining around each other and trailing over the .soil,
or occasionally rising higher. Only glimpses of sunlight
through the thick foliage of the trees ever reach the.se
plants. No purple flowers are borne here; sometimes a
few green aerial ones ; but the number of terrestrial flowers
must be truly .striking. If the.se localities are visited in
spring the young plants form a close bed of green ; later a
dense tangled mass of vegetation results.

CHARACrER OF PLANTS RESULTING FROM SPECIAL

Seeds.

As previously stated there are four types of seed— (a)
from purple flowers, {b) from green aerial, [c) terrestrial,

[d) winter type.

Any of the above seeds will give ri.se to a plant capable
of producing terrestrial seeds.

The terrestrial seed will, in summer, give rise to a plant
which may bear a, b, c. This is equally true, if the terres-
trial seed has been derived from a plant produced from a
purple, a green aerial, or a winter type seed.

What possibilities lie in plants raised from the other
seeds can not yet be confidently stated as a .sufficient number
of experiments has not been performed. It is likely that
the seed of the winter type produces all three kinds of

flowers.

The sharp dimorphism referred to in the early portion
of this paper, seems to disappear as the sunlight increases
in intensity, for then all transitions from the tall vigorous
twiner to low-growing feeble specimens exist.

In the localities where purple flowers fruit abundantly,
germinating seeds of this type have not been found. Hun-
dreds of seedlings have been uprooted, and terrestrial seeds
only have been seen. If none of the preceding germinate,
there would seem to be a tremendous waste of energy in the
plant, for in some cases the amount of seed produced is

3 so Schively — Contrilmtions to the Life History of

not by any means to be despised. From study of the
structure, and the experiments tried in the greenhouse, I
think it is improbable that the germination of these aerial
seeds occurs in their native haunts. Should they germinate
the feeble specimens resulting, would soon be crowded out
of existence, not only by their more vigorous relatives, but
also by the surrounding vegetation.

Comparison of Amphicarp.ka Flowers with those

OF OTHER Plants.

An examination of the Iveguminosre shows that there are
other members of the Order which share with Amphi-
carpcea the peculiarity of producing subterranean fruit. In
Huth's paper '^, quite a number are enumerated ; but with
many of those mentioned, beyond the simple statement
that the special plant produces two or more types of fruit,
little definite information concerning the life history of the
individual has been recorded. Among the most familiar,
are Trifoliuni suhtcrrancuui^ Vicia anipJiicarpa^ and Arachis
hyp ogee a,

Trifoliiim siibterraucum produces practically but one kind
of flower which may mature fruit above ground,^ if the
head is prevented from forcing its way under the surface.
Usually after the flowering period is over, the peduncle
bends, gradually lengthening, until the earth is reached.'*
Under the soil, the fruit ripens. According to Warming,^
the inflorescence commonly contains but four or five normal
flowers; the remaining ones he terms *' metamorphosed."
The calyx of this type is peculiarly developed, the function
evidently being to assist in penetrating the soil. All other
floral portions are absent in this type of flower. What
structural differences, if any, exist between the aerial and
subterranean legumes, I have not been able to ascertain ;
the subject probably awaits investigation. It is stated, how-
ever, by Belli ^* that the seeds contained in the former,
germinate with difficulty unless the integument is broken.

Vicia ajHpliicarpa bears two kinds of flower, each of which

AmpJiicarpcea monoica.

351

produces a distinct type of fruit. The aerial flower has a
papilionaceous corolla, the subterranean is apetalous.'^
The aerial flower never matures a legume similar to that
resulting from the subterranean flower; but there exists no
record of experimental evidence regarding the behavior of
either flower, if compelled to develop in a manner differing
from that which is considered normal.

Arachis hypogcEa has recently been studied by Mrs.
Pettit.'^' Arachis bears practically but one kind of flower,
which is usually found above ground, though occasionally
specimens have been seen upon the subterranean portion of
the steuL The flowers occur singly in the axils of the
leaves, and are sessile. After fertilization, the floral parts
fall, the gynophore lengthens and exhibits a decided geo-
tropic tendency. Growth continues until the ovary is
carried underground. This process is aided by the de-
velopment of a brown hardened tip upon the ovary. If
from any cause the ovary is prevented from entering the
soil, no fruit results. This statement is verified by experi-
ments conducted by M. Correa de Mello.-^' No differences in
character have been observed in the fruit resulting from
the subterranean flower. Mrs. Pettit observed the devel-
opment of a zone of hairs upon the gynophore after its
entrance into the soil; these hairs also developed where
the gynophores were placed in moist chambers. After a
series of experiments, she decided that their probable func-
tion is to assist in absorbing nourishment for the maturing

fruit.

In marked contrast to all of these plants, stands Amphi-
carpcua with its three distinct types of flower and fruit
occurring in a season. We have here a series of great
value, as study of the structural pecularities shows a
gradual transition from the aerial to the subterranean form.
It is worthy of attention, too, that the aerial apetalous
flower in the upper part of the plant may be forced to de-
velop a fruit and seed of the subterranean type, provided
the flower or a young legume is placed under the proper

ff "

li

M

352 Schively — Contributions to the Life History of

conditions. Possibly the above statement may apply even
in the case of the pnrple flower ; and experiments are now
in progress to test the matter.

The abnndance of axillary shoots, both primary and
secondary, presents a featnre qnite different from the plants
jnst discussed, as does also the habit of growth. The
cotyledonary, axillary shoots normally produce many flowers
and fruits. The runners above ground may do so likewise,
and if a darkened moist place is reached, the ovate one-
seeded legume results ; otherwise a flat green legume
resembling a diminutive aerial type is produced. These
rnnners are not provided with any device to assist them in
forcing their way into the soil. The runners from the
simple leaves are not strongly geotropic ; perhaps after a
certain period of growth, they are best described as nega-
tively heliotropic, yet the secondary branches frequently
exhibit beautiful geotropic curves. The axillary runners
from compound leaves are negatively geotropic, yet these
may be so treated that subterranean flowers and fruits are
borne. The cotyledonary axillary runners are feebly geo-
tropic, extending from the cotyledons at an angle of 75 to

90 degrees.

In Arachis and Trifoliuni the geotropic characteristics of
those portions bearing the flowers which are capable of ma-
turing underground, are quite pronounced.

The purple flowers of Ainphicarpcca evidently require
for their development, as has already been stated, consider-
able light intensity, and possibly also demand the vigorous
growth of the upper portion of the plant. When the spe-
cial period for the production of these is past, the aerial
apetalous form appears. The seeds in these legumes are
numerous, yet they possess a very hard covering, and are
difficnlt of germination. Filing these produced snch a
marked difference in percentage records of germination
that the success obtained by similar treatment of Trifolium
seeds matured above ground is recalled.

The subterranean fruit of Amphicarpcea is larger, and

Amphicarpcca monoica.

353

possesses a seed whose coat is delicate in structure, as is
also that of the legume. No difficulty whatever is experi-
enced in germinating these, whether the entire legume is
used or the seed only. It is not yet known what function
may be exercised by the hairs of gland-like base pecnliar
to this type of legume. These, as is surmised for the epi-
dermal structures of Arachis may assist in the accumula-
tion of nourishment.

Thus these seeds, though resulting from the most reduced
type of flower, are those upon which the reproduction of
the species may be said to depend. As it has been proved ex-
perimentally that these flowers do not give rise to this vari-
ety of seed, if they remain unexposed to dark surroundings
or above ground, it seems almost useless to propose pro-
tection from destruction by animals as one reason for the
subterranean seed development. It is true animals do
seek them, but there would not exist these delicious
morsels, if the seeds matured under other influences.

Darkness and moisture seem to exert some powerful in-
fluence upon these seeds, not only structurally, but also in
the nourishment stored up, and which possibly contains
certain substances valuable for the more successful growth
of the future seedling. Whether or not materials are ab-
sorbed from the surrounding soil, cannot now be answered.
Comparative chemical analyses of the aerial and subterra-
nean seeds will perhaps afford an explanation.

The phenomenon of subterranean seed-production is one
that has never been satisfactorily explained. It is gener-
ally conceded that there must be some signal advantage to
the species. Amphicarp(Ea nionoica is a plant whose flowers
at present illustrate transitional reduction. One is in-
clined to consider that the statement might be also made
for Trifolium subterraneum and Vicia amphicarpa. Was
Arachis hypogieat.\^^\\\ the same condition? Amphicarpcra
in some instances produces only the subterranean type of
legume. We have seen that the conversion of an aerial
to a terrestrial legume may be accomplished experimentally

354 Sc /lively — Contributions to the Life History of

within a short time — indeed, within the life of the indi-
vidual. Doubtless, we may picture the frequent occur-
rence of just the proper combination of circumstances ne-
cessary to cause a similar transformation of the aerial
legume. Consequent upon its production followed the
more successful germination of the seed, and certain struc-
tural changes were initiated. The characteristics once ac-
quired, were slowly, but persistently transmitted, until the
habit of the plant as regards subterranean seed-production
is now a permanent one.

It is puzzling to the biologist why in many instances
Amphicarp(Ea still yields quantities of aerial seed from
which, it appears, the species is so little benefited. Varia-
tion due to environmental conditions is in Amphicarpcea
beautifully illustrated, and in the history of few plants is
it possible to obtain a series equally valuable for demonstra-
tion of a transitional condition; for the presence of corolla,
the number, size and degrees of perfection found in the
stamens, the ovary, style and stigma — all afford material
which is deserving of careful comparative investigation.

In conclusion, the facility exhibited by certain of the
axillary runners, in acquiring new habits of growth, which
are continued through life, deserves emphasis. It has
already been stated that in case of injury to the main stem,
the cotyledonary axillary shoots and also those from the
axils of the simple leaves — themselves non-twiners — will
assume the duties of the main shoot. While the latter remains
uninjured, or should injury occur after the runners have be-
come five or six inches long (perhaps shorter) no amount of
artificial assistance and training is sufficient to induce the
twining habit. Since the experiments recorded in the early
part of this paper were performed, axillary shoots from the
simple leaves have been fastened so that they were made to
grow upward. Many of these have now reached the height of
five feet, and are nearly on a level with the main stem, yet
their free ends do not twine. Their actions indicate nega-
tively heliotropic and feebly geotropic tendencies. The

Amphicarpcea moiioica.

355

question may well be asked, from the evolutionary stand-
point, Would these peculiarities be modified or ultimately
disappear if continued cultivation under these artificial
conditions were persistently carried out?

With the readiness to change the habits transmitted by
inheritance to the various plant parts and to acquire new
ones from the pressure of external conditions, with great
possibilities in subterranean seed production, and the un-
failing germination of these seeds, it is not strange that a
luxuriant growth of Amphicarpcea greets the visitor to the
woods.

A Review ok the Species of Amphicarp.^a.

Recalling the dimorphism in plants resulting from the
germination of the terrestrial and aerial seeds in the green-
house, the variation in size, strength, and habit of the indi-
viduals growing in the open, and also the variation in the
flower production, a consideration of the probable number of
species or varieties with which we are dealing, is worthy of
our attention. A brief discussion of some of the causes
operating to produce such results will now be undertaken.

One reason why purple flowers do not appear upon plants
growing in shaded localities may be due to the non-develop-
ment of the upper stem-region and its axillary shoots where
these flowers are normally borne. Repeated injuries to the
growing main stem, and even to its successors may be an
explanation. Insects, birds and small rodents are fond of
the foliage, and the struggles for existence in a wooded spot
have doubtless something to do with the low-growing con-
dition described.

Those found upon the outskirts of the woods are taller
possibly because receiving more sunlight, but there may be
fewer enemies of the classes mentioned.

Amphicarpcea is rarely found as an isolated specimen ; it
seems incapable of developing except among other plants.
If no suitable support be available, several adjacent indi-
viduals will form a coil by the union of their stems.

35^ Schively — Contributions to the Life History of

These statements were proved by attempting to grow out
of doors, for more carefnl observation, single plants in pots
or in soil, so arranged that they were separated by quite a
distance. Even a slight breeze interfered with the attempts
of the stem to continue its normal habit of growth ;
stronger winds uncoiled it for some distance, frequently
snapping it in two, and so injuring the plant that it lay
upon the ground.

The number of types of flower produced, depends largely
upon the character of the natural conditions to which the
plants may be subjected.

As it is possible to raise from seed such different speci-
mens of AuipJiicarpcea^ the question arises, How many dis-
tinct species of the genus should be recognized for the
Eastern United States?

The Manuals mention A, inonoica^ sarmentosa and Pitcheri.
The first two certainly seem to be but varieties, resulting
from environmental conditions. This appears to the writer
to be proved by the study of plants in the greenhouse,
Botanic Garden, and several natural localities.

Twining plants of ^. monoica from Wissahickon differ in
no apparent respect from herbarium specimens of A.
Pitcheri ; for large, healthy leaves, a dense growth of fer-
ruginous hairs, and heavily-flowered racemes described for
the latter are likewise borne upon the individuals of the
former found in the above locality. Gray states that it
is not known whether A. Pitcheri produces underground
legumes or not. I have also received from F. Reppert, of
Muscatine, Iowa, plants of these species. While sufiicient
examination of them has not yet been made, the state-
ment by Gray, that has just been referred to, may be said
to be incorrect, for the cotyledonary shoots have been
seen in fresh material, and also underground pods in the
herbarium specimens. Nothing definite can be yet reported
concerning aerial flowers.

I consider, from observations that have been made, that
A. Pitcheri is an extremely vigorous A. monoica ; while A.
sarmentosa represents the poorly nourished type.

Atnpliiearpeca monoica.

357

A Ed<^e7vorthii is a Himalayan species, differing little in
appearance from A. monoica, Concernino tliose found m
Mexico, nothing explicit could be learned.

Summary.

1. A. monoica is strictly annual under ordinary condi-
tions.

2. There are two types of seed, aerial and subterranean.
The aerial may be divided into [a) those resulting from
purple flowers, {b) and [d) from cleistogamic ones.

3. Four distinct types of legume are developed {a) aerial,
green lanceolate acuminate pods, containing three to four
seeds; [b) aerial green oblong acuminate pods, containing
two to three seeds ; {d) aerial green, oblong pods, contain-
ing two seeds ; {c) subterranean purplish one-seeded ovate

pods.

4. In weight the subterranean pods exceed the aerial m

the ratio of 40 to i.

5. During winter a sharp dimorphism is noticeable m
the plants resulting from germination of subterranean
and aerial seeds. The form produced from the latter is a
low-growing non- twining plant, that from the former is a
tall vigorous twiner. During the summer, a dimorphism
still exists, but not in so marked a degree. The aerial
seed-plant is usually feebler in appearance, although it

twines.

6. The normally hypogean cotyledons of all seeds, if
placed so that they are exposed to light, develop chlo-
rophyll.

7. Injury to the main .stem below the simple leaves when
a plant is young, causes cotyledonary shoots to develop.
These normally negatively heliotropic and non- twining
shoots become apogeotropic and twining.

8. Cotyledonary axillary runners if allowed to develop
naturally, and then brought above ground, do not twine.
If the main .stem is destroyed after these shoots have
reached a length of several inches, twining does not take
place.

a

I \

358 ScJiivcly — Contyibutions to the Life History of

g. Axillary runners from the simple leaves (/. ^., first pair
of green leaves) in their natural condition are geotropic and
refuse to twine. When, however, the main stem is de-
stroyed, these runners become apogeotropic and twine well.

10. Axillary shoots from compound leaves soon exhibit
apogeotropic tendencies, and twine. If the main stem of
the plant is destroyed, axillary shoots from any compound
leaf will readily continue the twining habit of growth.

11. Circumnutation experiments show for Ainphicarpcea
the most rapid rate of movement known for a twining
plant. A complete revolution may be made in 5 1 minutes.
The optimnm temperatnre is from 26° C. to 32° C. ; the
behavior is greatly modified by environmental conditions.

12. In early development numerous crystals appear in
the cells forming the inner row of the cortex of the stem.
These ultimately constitute a distinct crystal sheath.
The crystals are somewhat prismatic in form, possessing
an apparent partition across the middle. These twin
structures occur also round the vascular areas of the leaves,
and also in the cortex of the pulvini.

13. All green leaves are capable of assuming normal,
paraheliotropic and nyctitropic positions. These positions
are absolutely dissimilar.

14. When young, the simple leaves assume peculiar
paraheliotropic and nyctitropic positions.

15. All leaflets manifest great activity during the night
hours.

16. Amphicarpita produces four types of flower: (a)
purple aerial, {b) green aerial, (<r) subterranean, (d) winter
aerial. A comparison of these leads to the conclusion that the
purple aerial is the original type, and that by a series of
transitions through {b) and (d) the most reduced type [c] is
reached. These variations are doubtless due to the environ
mental conditions under which they are produced.

17. Purple flowers are not produced during the winter,
even though the greenhouse is kept at a high temperature.
It is believed that the non-production of these flowers and

Amphicarpcea monoica.

359

also the sharp dimorphism previously noted as existing in
the winter between plants developing from aerial and sub-
terranean seeds, is due to a want of sufficient light intensity.

1 8. The purple aerial flowers (a) are produced on the upper
part of the plant from the latter part of July on to Septem-
ber. The green aerial [b) do not appear until the latter
part of August, and they may be found in all stages — flower
to fruit — as late as October. They are not so definitely
located upon the plant as the purple flowers are. The pro-
duction of the subterranean type begins in June and con-
tinues throughout the entire season.

19. The purple flowers {a) possess 10 diadelphous stamens,
the filaments of the 9 being united almost the entire length ;
their anthers are similar and perfect. The green aerial (b)
flowers have 10 stamens, distinct or with the filaments
united into a short tube; these anthers exhibit varying
degrees of imperfection. The winter flowers {d) have 10
stamens, which are distinct ; four of these usually bear
perfect anthers, the remaining stamens being quite rudi-
mentary. The terrestrial flowers [c] have indications of 10
stamens (many of them exceedingly rudimentary), those
which develop anthers vary from 6 to 4, and usually but
2 possess perfect anthers.

20. The thickened hypodermal layer (endothecium) pre-
sent in the anthers of («), is absent in anthers of the
remaining types.

21. The microspores and macrospores of the three types
of flower exhibit a gradual reduction in size.

22. A very complete series illustrating transitional reduc-
tion in styles and stigma may readily be obtained from a
study of types <7, />>, c.

23. Flowers of type {a) are probably close fertilized,
although there are present several devices associated with
insect fertilization. The remainder are cleistogamic flowers;
the pollen not being discharged from the anthers, but send-
ing forth tubes toward the stigma.

24. Flowers of type {a) do quite frequently produce

360 Schively — Contributions to the Life History of

legumes. Those of the remaining types rarely fail to pro-
duce fruit.

25. Legumes of types a, b, d, possess firm resisting walls,
being provided with sclerenchymatous fibres. These fibres
are absent in {c). But two types of hair are found upon
legumes of types (a, b, d) ; they are more numerous upon
(d) than upon the others. Three distinct types of hair are
seen upon the surface of legumes of type {c\ The stomata
of the aerial and subterranean legumes also differ.

26. Aerial cleistogamic flowers or young legumes may
be converted into the subterranean form of legume (c),
if buried in the soil.

27. A plant raised from terrestrial seed is capable of
bearing in summer, flowers of all three types— ^?, b, c.

28. The low-growing forms of Amphiearpcea found in the
woods, which may be feeble twiners or non-twiners ; and
the taller, more vigorous specimens found upon the outskirts
of the woods, are believed to be varieties due to environ-
ment and not distinct species as some have supposed.

Bibliography.

Eli.ioTT.1— '' Journal of Academy of Natural Sciences of Philadelphia,"
Vol. I, p. 372.

Ei^iyioTT."^— " Botany of the South."

Dari^ington.»— " Florula Cestrica."

ToRREY AND Grav.^— " Flora of North America."

GRAy.*— " Botany of Northern United States."

Von Mohl.«— " Bot. Zeitung," 1857, p. 730.

.T_«'Bot. Zeitung," 1863, pp. 309-328.

KUHN.8- «« Bot. Zeitung," 1867, p. 65.

Gray.'— " American Journal of Science/' Second Series, Vol. 39, p. 'oS-

Darwin.>«— *• Forms of Flowers on Plants of the Same Species."

.n *< Power of Movement in Plants."

.i2_" Movements and Habits of Climbing Plants."

HENSI.OW.''— "Nature," October 19, 1876, p. 543.

.!♦__" Gardener's Chronicle," 1877, p. 271.

FARI.OW.**— ** BuUetin of Bussey Institute," 1878, pp. 224-252.

Meehan.'«-" Proceedings of Academy of Natural Sciences," Philadel-
phia, 1887, pp. 323-333- „

HUTH E.'^— " Ueber Geokarpe, Amphicarpe, und Heterocarpe Pflanzen.
Abhandlung. Ver. Naturwiss. VIII, 1890, pp. 89-121.
Frankfort-on-Oder.

Aniphicarpica jnonoica.

361

Lubbock." — "A Contribution to Our Knowledge of Seedlings."
Mackari.ane.'*— "Proceedings of the Botanical vSection. Academy of

Natural Sciences," Philadelphia, 1894.
. ,^0 - " Lectures at Marine Biological Laboratory at Woods

Holl," 1893.
— 'i\ — «« 'file Sensitive Movements of Some Flowering Plants

Under Colored Screens." Bot. Centralblatt, 61, 1895.

Warming."— " Bot. Centralblatt," 14, 1881.

Bei.M, S."— " Sui rapporti sistematico biologici del Trifolium subter-
raneum cogli affini del gruppo Calycomorphum." Malphigia
VI. 1892.

M. CoRREA DE MEiyU).'^*— " Notes on Brazilian Plants from Neighbor-
hood of Campines." Jour. Linn. Soc. Bot. XI.

Mrs. a. S. YKtrit.'^^—'' Arachis hypogcra:' Memoirs of the Torrey
Botanical Club, Vol. 4i No. 4-

EXPLANATION OF PLATES.

PLATE XIX.

Roots of three plants, showing legumes and tubercles ; from photo-
graph.

PLATE XX.

Fig. I . Terrestrial legume ; one-seeded type, natural size.

Fig. 2. Terrestrial legume ; two-seeded type, natural size.

Fig. 3. Transverse section through seed-coat of terrestrial seed.

Fig. 4. Legume of purple flower ; immature condition ; slightly reduced.

Fig. 5. Legume of purple flower ; mature condition ; slightly reduced.

Fig. 6. Legume of green aerial flower ; slightly reduced.

Figs. 7 and 8. Legumes of winter type of aerial flower; slightly

reduced.

Fig. 9. Transverse section through seed-coat of seed of aerial type.

PLATE XXI.
Plants of the same age; one raised from terrestrial seed, the other
from aerial seed ; from photograph.

PLATE XXII.

Lower portion of a plant, showing axillary shoots, which were raised
from their normal position, and supported by means of thread; from
photograph.

PLATE XXIII.

Fig. I. Plant a few days after truncation had taken place.
Fig. 2. Plant in twining condition ; from photographs.

362 Schivcly — Contributions to the Life History of

PLATE XXIV.

Fig. I. Transverse section of very young stem.

Fig. 2. Transverse section of older stem.

Fig. 3. Transverse section of still older stem.

Fig. 4. Transverse section of stem about the age of that shown in
Fig. 2 — much magnified.

Fig. 5. Portion of leaf, showing vascular bundle, containing crystals.
All figures from photo-micrographs.

PLATE XXV.

Fig. I. A fair-sized plant showing leaves in paraheliotropic position.
Fig. 2. Young plant showing leaves in paraheliotropic position.
Fig. 3. Young plant showing leaves in nyctitropic position : from

photographs.

^ PLATE XXVI.

Plant showing position of leaves at u p.m ; from photograph.

PLATE XXVII.

Fig. I. Racemes of purple flowers, showing manner of growth.
Fig. 2. Compound raceme of purple flowers.

Fig. 3. Transverse section of a purple flower-bud. Figs, i and 2 from
photographs; Fig. 3 from photo -micrograph.

PLATE XXVIII.

Fig. I. Transverse section of purple flower-bud showing nectary.
Fig. 2. Transverse section of ovary of green aerial flower.
Figs. 3 and 4. Transverse section ; longitudinal section of terrestrial
flower-bud ; from photo-micrographs.

PLATE XXIX.

Fig. I. Stamens and pistil from green aerial flower. X 35°-
Fig. 2. An enlarged anther from green aerial flower. X 75°-
Fig. 3. Stamens and pistil from terrestrial flower. X IS° •
Fig. 4. Stages of staminal reduction from terrestrial flower ; i consists
of filament only, 2 and 3 possess imperfect anthers, 4 is perfect. X 75 •
Fig. 5. Microspores from terrestrial flower. X 35o°-

PLATES XXX AND XXXI.

Series of styles and stigmas showing transitional reduction, taken
from three types of flower. Drawn to same scale.
Figs. I and 2. From terrestrial flower.
Pigs. 3-7. From green aerial flower.
Fig. 8. Immature form, purple flower.
Fig. 9. Mature form, purple flower; drawings from photo-micrographs.

A mpJiicarpica monoica.

363

PLATE XXXII.

Fig. I. Thickened roots of plant; slightly reduced.

Fig. 2. Pistil of purple flower. X i^°.

Fig. 3. Pistil of green aerial flower. X 18°.

Fig. 4. Pistil of terrestrial flower. X 18°.

PLATE XXXIII.

Axillary shoot .showing arrangement of the winter legumes ; from
photograph.

PLATE XXXIV.

Fig. I. Stoma from aerial legume. X 35^°-

Fig. 2. Epidermal hairs from terrestrial legume. X 250°.

Fig. 3. Hair with glandular base from terrestrial legume. X 250°.

Fig. 4. Stoma from terrestrial legume. X 35^**-

PLATE XXXV.

Fig. I. Low-growing plants raised from terrestrial seeds ; from photo-
graph.

Fig. 2. Plant raised from aerial seed, bearing terrestrial seeds; from
photograph.

Fig. 3. Cotyledonary region of plant, showing the arrangement of
terrestrial legumes; from photograph.

PLATE XXXVI.

Fig. I. Cotyledonary region of plant showing all stages of flowers
and legume formation ; from photograph.

Fig. 2 Cotyledonary region of plant showing cotyledons spread open
and axillary shoots arising; from photograph.

Vol. I. I'lftte XVIII.

Bof. Contrib. Univ. Penn.

Cross on Eupatorium.

Vol. I. Plate XIX

Hot. Coiifrlh. Univ. Penn.

SCHIVELY ON AmPHICARP/KA.

Vol. I. Plate XX.

Bot. Contrib. Univ. Perm.

SCHIVELY ON AmPHICARP^EA.

Vol. I. Plate XXI.

Bof. (onfrih. (Jniv. Petni.

Schivp:ly on Amphicarp^ka

1p

I

Vol. I. Plate XXII.

Bot. Contrib. Univ. Penn.

SCHIVELY ON AmPHICARP.^.A.

Vol. I. Plate XXIII.

liitf. Coiitrlh. Univ. Pom.

SCHIVEI^Y ON AmPHICARP^:A.

Vol. I. Plate XXIV.

Bot. Contrib. Univ. Penn,

i-ig. a.

Fig 5.

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M^.^k

SCHIVELY ON AMPHlCARPyEA.

Vol. I. Plate XXV.

Bot. Conti'ib. Univ. Penii.

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SCHIVELY ON AmPHICARP^A,

Vol. I. Plate XXVI.

lU)t. Contr'ib. Univ. Penu.

SCHIVKivY ON AMPHICARPi^iA.

Vol. I. Plate XXVII.

Hot. Confrih. C^niv. Venn.

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Fig. 3

vSCHTVKLY ON AmPHICARP/KA.

Vnl. I. Plate XXVII

I inf. C'ottfi'ih. (

Hir. Icini.

vSciIIVKLV OX AmpiiicakimvA,

INTENTIONAL SECOND EXPOSURE

Vol. I. Plate XXVIIt.

Bot. Contrib. Univ. J*enn.

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SCHIVKLY ON AmPHICARP^:A.

Vol. I. riutexxviii.

Hot. (\nifrih. I nil'. I

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SCHIVKLY ON AmPHICAKP.ua,

INTENTIONAL SECOND EXPOSURE

Vol. I. riate XXIX.

Boi. Contrih. Univ. Penn.

Fig. 4

fig. 2

SCHIVEI^Y, ON AmPHICARP^A.

Vol. I. Plate XXX

Bot. Contrib. Univ. Penn.

SCHIVELY ON AmPHICARP/KA.

Vol. I. Plate XXXI.

Bot. Contrih. Univ. Penn.

SCHIVELY ON AMPHICARPi^A,

Vol. I. Plate XXXII.

J>ot. Contr'ih. Univ. Penu.

Schivp:ly on Amphicarp^,a.

Vol. I. IMate XXXIII.

But. (onfrih. (hiii\ Penn.

SCHIVKLY ON AmPHICARP.KA.

Vol. I. Plate XXXIV.

But. Contrih. Univ. Penu,

SCHIVELY ON AmPHICARP.*:A.

Vol. r. Plate XXXV.

Bot. Contrlb. Univ. Penn.

SCHIVKLY ON AMPHlCARPi^iA.

Vol. 1. IMute XXXVI.

Bot. Contrih. f^niv. Penn.

SCHIVELY ON AmPHICARP/KA.