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BOTANICAL GAZETTE

THE

BOTANICAL GAZETTE

EDITORS:

JOHN MERLE COULTER anp CHARLES REID BARNES,

VOLUME XXXI1X

JANUARY —JUNE, 1905

WITH TEN PLATES, TWO MAPS AND SIXTY-FOUR FIGURES

CHICAGO, ILLINOIS
PUBLISHED BY THE UNIVERSITY OF CHICAGO

1905

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TABLE OF CONTENTS.

The effect of the presence of insoluble substances on ete
the toxic action of poisons - Rodney H. True and C. S. Oglevee I
The relation of soils to natural vegetation in Ros-
common and Crawford counties, Michigan.
Contributions from the Hull Botanical Labor-

atory. LXVI (with map) - - B. E. Livingston 22
New and noteworthy western plants. II - - A.D. E. Elmer 42
The morphology of Monascus pur pureus - - E.W. Olive 56
The theory of respiration - - - C. R. Barnes 81

The forests of the Flathead Valley, Montana. Con-

tributions from the Hull Botanical Laborat

LXVII (with map and twenty-three aa a N. Whitford 99,194,276
spi in the Gramineae. VIII. Munroa squarrosa

Nutt.) Torr. saat twelve figures) - . Theo. Holm 123

Soke of generations in animals from a botanical

standpoint. Contributions from the Hull

Botanical sabaeieny: LXVIII “= two

figures) . C. J. Chamberlain 137
Gametophytes and embryo of Torreya ait. Con-

tributions from the Hull Botanical Laboratory. °

LXIX (with plates A, I, 17, TI) - J. M.CoulterandW.J.G. Land 161
The principles of phytogeographic nomenclature - Pehr Olsson-Seffer 179
Contributions from the Cryptogamic Laboratory of

Harvard University. LX. A new American

species of Wynnea (with plates IV and V) - Roland Thaxter 241
On the development of Hamamelis virginiana. Con-

tributions from the Botanical Laboratory of

Johns Hopkins ik tite No. 3 iis si

VI and VII) - - > D.N. Shoemaker 248
Sexual reproduction in the rusts (with _— VIR}. + A. H. Christman 267
On proteolytic enzymes. I - Arthur L. Dean 321

Development of sporangium in Botrychium. Contri-
butions from the Hull Botanical nee:
LXXI (with plate IX) - - Ira D. Cardiff 340

vi CONTENTS

Physiological properties of bog water. Contributions
from the Hull Botanical ere LXXII
(with three figures) . -

An apparatus for observing the eae se stream
(with two figures) - -

Leguminous rusts from Mexico - -

On the water-conducting systems of some desert
plants (with ten figures) - -

The effects of toxic agents upon the action of bromelin.
Contributions from the Hull Botanical Labor-
atory. LXIII - - - - -

BRIEFER ARTICLES—

Fertilization in the Saprolegniales - =- ~~ -

The sexual organs and ili generation
of the Rhodophyceae -

New precision appliances for use in plant phys
ology. II (with four figures) -

Terminology of the ita structures in the
Uredinales -

Notes on the physiology of pesdaai

Contributions from the Hull Botanical Lab-

oratory. LXX (with three figures) - -
Fertilization in the Saprolegniales - -
Polyembryony in Sphagnum (with three fig-
ures) - - < . “ . =

Adenoderris, a valid genus of ferns (with two

The early history of angiosperms = - Sa a,

CurRENT LITERATURE - -
For titles see index oak. ae name wat
Reviews. Papers noticed in ‘Notes for
Students” are indexed under author’s name

and subjects

News : = ‘ a *

DATES OF PUBLICATION.

[VOLUME XXXIx

B. E. Livingston

Otto V. Darbishire
J.C. Arthur

W. A. Cannon

J.S. Caldwell

B. M. Davis”

B. M. Davis
W. F. Ganong
J.C. Arthur

B. E. Livingston
A. H. Trow

Hi. L. Lyon

W. R. Maxon -

Ethel Sargant

PAGE

% 153, 223, 301, 370, 424

- 79, 160, 240, 319, 384, 436

No. 1, January 20; No. 2, February 17; No. 3, March 20; No. 4, April 18;

No. 5, May 20; No. 6, June 22.

i
b

Bases oe piss eg Gi

VOLUME XXXIX] CONTENTS vil

es St em Se

ERRATA.
2, line 9g, before John Milliken University read Lincoln College of.
20, line 2 from below, same 2s above.
149, line 5, for red read blue.
150, line 11 from below, the * Should be omitted.
229, line 11, for ovule read ovules.
366, in first footnote, for Teut read Tent.
368, last line, for pinnatifid read bipinnatifid.

383, line 20, for of read to.

rec: No. 1

“ Rena Sas

Che Botanical Ga3ette

A Monthly Journal Embracing all Departments of Botanical Science

Edited by Joun M. Covtrer and Cuartrs R. BARNES, with the assistance of other members of the
botanical staff of the University of Chicago.

Vol. XXXIX, No. 1} Issued January 20, 1905
CONTENTS
F OE. EFFECT OF THE PRESENCE OF INSOLUBLE ene ON THE TOXIC
ACTION OF POISONS. Rodney H. True and C. S. O ~ I

THE RELATION OF SOILS TO NATURAL gcc IN ROSCOMMON AND
CRAWFORD COUNTIES, MICHIGAN. ConTRIBUTIONS FROM THE HULL BOTANT-

caL LaBoratory. LXVI (wirn Map). Burton Edward Livingston - 22

NEW AND NOTEWORTHY WESTERN PLANTS. II. A.D.E. Elmer - - ~ - 42

THE MORPHOLOGY OF MONASCUS PURPUREUS. Edgar W. Olive - - ~ ~ 56
BRIEFER ARTICLES.

FERTILIZATION IN THE SAPROLEGNIALES. B. M. Davis - 61

RGANS - RATION OF THE REGHOFRYCEAE. B. M. Dene 64

CURRENT LITERATURE.
; BOOK REVIEWS §

* a is a pa ee i “e a 67
THE LIFE HISTORY OF PINUS
MINOR NOTICES ad - z rs ~ a 4 a = - 68
: NOTES FOR STUDENTS . _ i - - = _ © - 68
eR ee

£ Aten 24. 1 LA k Ps Pe | oe tere he

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VOLUME XXXIx NUMBER 1

bOLANICAT 9e W711i

JANUARY, 1905

THE EFFECT OF THE PRESENCE OF INSOLUBLE SUB-
STANCES ON THE TOXIC ACTION OF POISONS.

RODNEY H. TRUE and C. S. OGLEVER.

Ir has been repeatedly observed in connection with the poisonous
action of dissolved substances that the harmful effects are much
more marked when the roots of the plants under test are immersed
in the solutions than when these are grown in sand cultures which are
watered with like or even more concentrated solutions (1). Upon
this point the practice advocated by NAGELI (2) for removing traces
of metals from distilled water may be cited. When to distilled water,
rendered harmful for algae by the presence of traces of copper, crushed
graphite, shredded filter paper, paraffin shavings, and other insoluble
substances are added, the copper is removed from the water by some
process dependent to a degree upon the amount of surface offered
by the insoluble substance.

In connection with experiments carried out in 1902 by TRUE
and Grrs (3) at the Plant Physiological Laboratory at Woods Hole,
Mass., on the modification of the action of poisons by the presence
of other compounds, some preliminary experiments were made
bearing on the effect exerted by the presence of insoluble materials
on the behavior of seedlings of Lupinus albus. A concentration of
copper sulfate was experimentally determined in which growth for
twenty-four hours was possible, though at a somewhat diminished

~rate. When to duplicate cultures insoluble substances were added,
in nearly all cases a marked increase in the growth rate was noted.

t After this paper was prepared, the line of argument and the evidence were sub-
mitted to Dr. Lyman J. Briccs, physicist of the Bureau of Soils, U. S. Department of
Agriculture. He has most generously given us the benefit of his special advantages.

I

P BOTANICAL GAZETTE [JANUARY

The same volume of solution was used in each case and the same
amount. of copper was present in each culture. In some cases a
copper-containing solution, in itself harmful, gave a marked stimula-
tion when the insoluble material was added. In other cases the
increase in the growth rate was less marked or, in two cases, absent.
The general trend of the evidence, however, was clearly in favor of
NAGELI’s conclusion that insoluble bodies in some way remove from
action limited quantities of the copper in the solutions.
During the summer session of 1903, C. S. OGLEVEE, of the John
Milliken University of Lincoln, Illinois, while working in the Plant
Physiological Laboratory at Woods Hole, repeated the experiments
above described and extended their scope. He used the radicles
of seedlings of Lupinus albus as test objects.
The insoluble substances used represented several types of sub
stances, having little in common beyond their insolubility. In mak-
ing a choice of these the suggestions of NAGELI were again acted
upon. In experimenting with these different insoluble or difficultly
soluble materials we attempted to use such quantities of each sub-
stance as would offer, as nearly as possible, like surfaces to
solutions. It is improbable, however, that even approximate accu-
racy was reached on account of the different sizes and shapes of t
particles. Clean sea sand of rather coarse texture was used | :
frequently. It was selected from the cleanest parts of the b e
near Woods Hole, boiled for about an hour in dilute hydra h
acid, then thoroughly washed in fresh water and finally in we

——> eS

~ 1905] LRUE & OGLEVEE—TOXIC ACTION OF POTSONS 2

ment with distilled water. It was noted that gas bubbles were repeat-
edly found over the surface of the submerged particles of coal and
that after being removed by thorough stirring these would again
form. This evolution of gas continued during the entire time in
which the coal was under observation. The nature of the gas was
not determined.

Finely divided pure paraffin of high melting point was also made
use of. Investigations made on the effect of paraffin on the electrical
conductivity of pure water have shown that paraffin does not notice-
ably increase the conductivity and seems to be practically insoluble
in water.”

Owing to the lightness of the paraffin, the finely divided particles
floated on the surface of the solutions, but owing to the small spaces
between the pieces, the solution rose by capillary action among them
and they became, in effect, submerged in the solutions.

At the suggestion of Dr. HENRY KRAEMER, of the Philadelphia
College of Pharmacy, unruptured potato starch grains were tried.
A stock was freshly prepared from potatoes by crushing and washing
out the starch grains, care being taken to avoid other structures from
the tuber. The starch thus obtained was dazzlingly white and dis-
tilled water in which a portion of the material was allowed to stand
for some time gave no starch reaction when treated with iodine.
By rubbing up a little of the same starch with a little distilled water
a strong starch reaction appeared on adding iodine solution. Careful
washing removed all traces of other substances.

The toxic substances receiving most attention in this investiga-
tion were those which have been shown to possess great poisonous
activity. Copper sulfate, silver nitrate, mercuric chlorid, hydro-
chloric acid, sodium hydroxid, thymol, and resorcinol may be
especially mentioned. The chemicals in the case of the heavy
metals were Merck’s guaranteed reagent grade. The remaining
compounds were recognized makes of chemically pure’ quality.

_ The solutions were made up with such care as the apparatus of the

2 We are informed that investigations by F. K. CamMERon and Lyman J. Briccs
of the Bureau of Soils, U. S. Department of Agriculture, show that high grade paraffin
of high melting point, of the sort here used, has but a minimal effect on the electrical
conductivity of distilled water in which it is placed.

4 BOTANICAL GAZETTE [JANUARY

laboratory permitted, and we believe that the concentrations were
usually accurate.
The method of operation consisted essentially in suspending
- the seedlings on glass rods in the solutions concerned for twenty-
four hours each in each experiment, in such a manner as to immerse
the radicles. In nearly all cases the plants were under observation
at least forty-eight hours. Never less than four seedlings were
used in any experiment and frequently more were employed. The
average is taken as the result. The seedlings were placed with the
roots in the solutions, care being taken to use plants in like stages of
development. Growth was measured from an India ink line placed
15 ™™ from the tips of the roots at the beginning of the experiment.
During the exposure to the action of the solutions the culture dishes
stood in diffused light. The influence of variable factors, such as
_ temperature, was kept track of by check cultures in distilled water.
The water was distilled in an apparatus lined with block tin and
was not, therefore, of the highest purity. It was consistently used, how-
ever, throughout the season and forms a constant factor in all experi-
ments. Beakers of 300°° capacity and of similar shape were used.
In studying the effect of the presence of insoluble substances on _
the poisonous activity of the compounds: mentioned, a quantity of
the sand, filter paper, or paraffin was generally used equal to about
one-third of the volume of the solution of the poisonous agent.
Thus, in the beakers used for these cultures about two-thirds of the
depth of the solution was to be seen above the level of the sediment
in all but the floating paraffin, in which case the order was inverted.
After marking the roots and introducing them into the solutions
under study, the plants were set away for twenty-four hours, at the _
end of which time they were again measured and the amount of ;
growth noted. Usually, to avoid further complications arising in
the case of prolonged experiments, the results due to exposure of

eed Se ow

cn eae hours were sought.
Th it ee ee a | els ee

: r : gece
matel the maximum +

1905] TRUE & OGLEVEE—TOXIC ACTION OF POISONS 5

The first group of toxic agents tested were exclusively electrolytes,
which at the very advanced degree of dilution here seen may be
regarded as acting chiefly as ions. The possible presence of a resid-
uum of molecules, especially in the case of HgCl,, is of no particular
significance for the present discussion. Concentrations are always
cited in terms of parts of a gram-molecule of the substance in question
per liter of solution.

Mercuric chlorid.

Preliminary experiments were made to determine the toxic actiy-
ity of the solution in the absence of insoluble substances. A concen-
tration was then selected at which growth, although greatly depressed;
is still possible before death results, and parallel cultures were then
made in these concentrations both with and without the insoluble
substance. The following table summarizes the results, growth made
being indicated in millimeters:

TABLE I.
MERCURIC CHLORID.

I gram-molecule Growth in 24 hrs. | Growth in 7o hrs.
Mckee, PCS tao a eretresat ina ee Gaze 'n Se nace. GS es 2.0mm 6.5mm
OC eeOe CAN ao Boia sis. Hela lean a wasted oa a gd 6.0 21.0
oO, Rec eee ater nace sya ereree are due, mbeely ie non orice 10.0 28.5
2G,000-F Crushed glaSS 24 shaw. care Sie wteeeiore toed oe 16.9 40.0
Checkin waters: 24) newea cet cela ike. e acre 12.0 16.0

It is clear that the presence of both sand and crushed glass here
modifies very markedly the toxic influence of the mercuric chlorid.
This amelioration is evident not only during the first twenty-four
hours of the experiment, during which the roots were exposed to the
corrosive sublimate solution, but also during the following period
of nearly two days, during which the roots were placed in tap water.
The course of the check experiment shows that the presence of the
insoluble bodies exerts a clearly stimulating effect on the roots. The
same may be seen in a less marked degree in the more dilute solution
lacking the insoluble substance.

Silver nitrate.

The above methods were applied to silver nitrate in a large number

of dilutions, with somewhat less marked results.

6 BOTANICAL GAZETTE

TABLE II.

SILVER NITRATE.

2
|
;
i

[JANUARY

— es

Growth in 24 hrs.

Growth in 48 hrs.

ooo lit

,000 lit SRNR cel ri gs 2 oc eon deat Wind nono
h

250,000 HTLETS.. 2 2. ee ee ee eee ee eee ene

250,000 liters

Water, distilled

Water +sand.

Oe @ 216s eae ee es ae 6 6 ee ee ©

web) we Oe ee 0 ee Ree 6 8 eK

S50. ct WE. Sen ee deen hd ati cea eee
250,000 liters +sand...........-. 6220s eee eee es

Sa ae ak pecan eo Ot oe) Bw) cy wr ee oe Oe

ee ee ee a

AUGUST 14

ae

000000

9.0

16.0
(weak seedlings.)
32.0

19.0

A study of the numerical data shows that in four cases out of ten,
in which the active solutions with and without the solids are com-
pared, no increased growth rate appears in the magnet of the —

ilic euler exci vate were with thatcof the on mike ie .
_ lacking solids. In one case out of ten there is a retardation seen to
follow the introduction of the solid. If we examine the six cases in -
which an acceleration followed the introduction of the solid, the sum of ©
the: nieces in asian of the cultures containing the solids is 32".

_ Thus, seen tk

te growth ceratoaneten presence
“of the solids is very marked. Since no ex]

Se various solids used, we cannot
li of

1905] TRUE & OGLEVEE—TOXIC ACTION OF POISONS 7

solids present may be responsible for the differences noted. In
some degree, also, slight differences in handling during the experi-
ment and also individual differences in the roots produce slight,
uncontrolled effects.

Copper sulfate.

A greater number of experiments were made with copper sulfate
than with any other substance. This has an added interest because
of NAGELI’s (2, p. 23) experiments with Spirogyra in extremely
dilute solution. NAcELr found that distilled water prepared by means
of a copper still may contain enough metal to produce fatal results
with Spirogyra. The remedy proposed as a laboratory expedient
for making water thus prepared fit for use was to put finely divided
insoluble bodies into the water. The results obtained with copper
sulfate follow. Although there is considerable repetition and some
experiments are not completely carried out, it seems desirable to
present all the evidence, showing as it does the range of variation
in the results:

TABLE II.
COPPER SULFATE,
I gram-molecule Growth in 24 hrs. Growth in 7o hrs.
Jury 8
ESO TET i ee wick eet « viny bo on nti ae 2.9mm Dead

aie gril et iid euler Be Rleaay oes SOAS chapel pt ate 2.0 on
aA cs NOs ks ches ered See Waee mera EN, e
#552 TORS + SAMs ies Sen kes Shes eh eee Fs 4.0 *
Thtetitied! ‘writers. cas nd oe esa em eee eeese ee 12.0 17.9mm

JULY 10
50,000 li Ee cd areude ek GREE ME eran ie eee 1.0 Dead
50,000 ters sand 9 0s & de wae dren ened 14.0 28.omm
PENS OEE, os cna ec ened sass Ween o ese ce eee 72. 28.0
Sa ert sane be Soca ak eee sa Ae 16.0 32.0

JuLy 13 Growth in 48 hrs
35) Ee Pe org sae ek carts tional owne 98 Res I.0 Dead
AS Me MeL S 4 SANG, C5. yds ae Stee sence we eee 7.0 I2.9mm
RIO tenis sl PISS ak face eles 4s ae ERAS Os 8 2:0 Dead
AED CON TE 1- SAL Groh cit Gia, eee ont aia ems 5.0 g.omm
MN I I 5 12 Awan ace sb ease eee 2.0 Dead
50,000 Boies lis Mle ates ar oldie eccre oalle we eee rp ee, 17.9mm
DMI URN a ei lV aswiieie s Kirk ga ew nwa ow 10.0 13.0

istilled tics Se PMMA ig re Att Shona aK a, sive e 8.0 15.0

[JANUARY

35,000 liters + garden soil od hic ok Shik ‘ores
Distilled

8 BOTANICAL GAZETTE
TABLE IlI]—Continued
1 gram-molecule Growth in 24 hrs. Growth in 7o hrs.
JuLty 16
35:00 Re ae Sinead Mem has are eet rea 4.0 Dead
35,00 PARE Ne LP Onc att. feat cls a ee Io. I7.omm
35) 000 cdrom ADO WA io. 6 eiaw Sys Ce an axe 18.0 32.0
WME ore alors iotala oe a mdf cpa S's ada soarer abe renee 12.0 15.0
Distilled INI, SES pd eink cy on nh om de . 9.0 16.0
Distilled water-}iier Papers. in os. vee nwewcaes 12.0 21.0
Distilled water +glass wool..............+-. Ase 16.0 29.0
JuLy 20
WE SEES TREY ORIN no oi ese AE oo ae Me I2.0
35,00 liters +glass wool.............- Petes 5.0
Distilled ‘waters. soca s ce Palape ig aces 14.0
illed SSG a BT seis Safe Sed 12.0
JuLy 24 _ Growth in 72 hrs.
ph ES de aie oka at oc ioe tan (3.0 : :
zeroes Titers paraiso... os. ew ee wa sien meee 8.0
PAE WHER oes. Sarctek's oto iern es She pepe, a ¥2.0
JULY 29
eo... Sey ae ae eee ea ees iden et of £6
35,000 litess-¢-glass....< ....2. 2222+. se apres lar eau 4.0
Juty 30
Ces 8: ee ee ee ee oe Ssapeanies wists I.0
a500e Titers | paraffin... 2.5. - - By Siete 3-0
JULY 31 -
be en Geraclietsd Cee IeAkS I.0 ad
; 16.0"

1905] TRUE & OGLEVEE—TOXIC ACTION OF POISONS 9

repeatedly observed before (3, 4). When, however, to a solution
containing nearly or quite twice this amount of the copper salt, finely
divided insoluble bodies, like quartz sand, crushed glass, and filter
paper are added, the toxic action is so far diminished as to permit
not only an increased growth before death takes place but even vigor-
ous survival. In case the amount of copper is diminished, an advantage
is regularly found in favor of the cultures containing the solid body.

In the case of the culture containing the garden earth, the effect
of a common type of soil containing considerable soluble matter was
desired. Of course chemical reactions have entered here, but at
this great dilution the copper probably still existed in the ion condi-
tion. Here, too, the introduced soluble material may have had its
own direct action on the organisms.

The action of paraffin is marked in every case but in a somewhat
less degree than in the other cases noted. The paraffin offered some
slight difficulty of method which may account somewhat for the fact
observed. Although hard paraffin was used, it was found difficult
to shave or chop it up into particles of the desired degree of minute-
ness. Thus less surface was exposed to the solution than in the
other substances used. Its lightness, too, tended to buoy it up upon
the surface of the solution, and prevent a thorough mingling. This
was remedied in part by the tendency of the solution to rise by capil-
lary action among the floating particles. It was also found practicable
to hold the paraffin under, to a considerable degree, by means of
pressure on the surface of the floating mass.

The effect of solids on growth in the distilled water was tried in
control experiments, the general result seeming to indicate that the
plants are not favored by this presence. Redistilling from glass,
discarding the first half of the distillate and leaving a third of the
original volume in the retort, seems to yield a product having no
advantage over the water but once distilled.

In all of the toxic substances thus far considered, we have had to
do exclusively with electrolytes, therefore, at the high dilutions con-
cerned, with ions. Experiments were made with a. few non-electro-
lytes in order*to ascertain whether the ionization of the molecule
may be a necessity for the observed decrease in the poisonous action
of the presence of insoluble substances.

Ke) BOTANICAL GAZETTE [JANUARY

Non-electrolytes.

Those tested in this connection were phenol, resorcinol, and thymol.
The investigations by BADER (5) on the conductivity of phenyllic
compounds showed that in the substances here concerned no ioniza-
tion is to be found. We have, therefore, here to do with entire mole-
cules.

TABLE IV.
PHENOL.
I gram-molecule Growth in 24 hrs.
AUGUST 5
ONS TUNG bien 6h oh ceria wine wide es Vere 5.0mm
BS ae Cae) Ceeee eens 6.5
aoe fiters 4 samo. ooo te ae 6.0
Bustilied Water... 0c. 66s eeu 41.5
AUGUST 7
Reet ees ce, ee et sy Ber I.omm
Too nai BEAL a eee ee 2.0
CEU EERE Nace ene u sens ew sleraiy RO 2.0
150 liters isa Riga ota a hea eres ac
Distilled water..... mere ilakalt! Sn aecane 10.0

An inspection of the results shows no apparent amelioration result-
ing from the presence of the solid. It should be noted, however,

in passing, that compared with the electrolytes used, phenol is a
weakly toxic substance, and the plants are apt to make a good growth

in solutions containing a comparatively great number of molecules (Q).

TABLE V.
RESORCINOL.

1905] TRUE & OGLEVEE—TOXIC ACTION OF POISONS II

In this substance, in concentrations great enough to interfere
greatly with the growth rate, the presence of the solid seems to bring
about a slightly increased rate of growth, this effect ceasing as the
dilution is increased. Here we meet again a compound which is
tolerated in considerable concentration by the plant (9).

TABLE VI.
THYMOL.
I gram-molecule Growth in 24 hrs. | Growth in 48 hrs.
JULy 30
2600 nai Dace Pee ae PR Ad ly cei Bele she o.omm o.omm dead
BEOG HUCTH a ete e ea eae i ee Ore Sears 4.0 g-omm; 2 dead
2800 itaseade cass be ee ercteie & bani hgh cme amare oars 14.0 28.omm
JULY 31
te Dieta ge SEP O11 8 ule O erode tem rate orate eacnatee ac 2 dead
ae for nd Siig. Sung Oh axe arp @reeete: pene a eens 350 3 dead
AUGUST 5
IS TRE oc toa) divs OAS Sas Boece ca ie epee 2.0
micion WRG rS Sa arc b5 ea we oa ne cawecet elo, Wes mages I.0
ERO! URNA ora carla wes date eat ee trees 4.0
liters BI as. plas ce ev ennatele ee aleetlacts oF 5.0
Distilled, watets aur ss wren cigsinealg sina eesti L5
AUGUST 7
AGO TRUS Ie om orsiianla casero tssve molest sawn e te 52 Be oiF all killed
20 BTA BANC hid, Peel kon oe ws ea ole ss 3 2.0 all killed
PBOO- ME CE Sec ee oes Genie rate a cn te) ties Wein etecpiecele; Oks 6:5 all killed
2800 lite 1 a Barone de lel an ae 7.0 3 living
Distitled: watétgos. 2 ss ee Pa eee 10.0

The results with thymol, while not homogeneous, indicate that
in a concentration of one mol. weight in 2600 liters of solution no
steady advantage accompanies the presence of the solid. When
the dilution is somewhat increased, however, such an advantage
appears. The time at our disposal was too brief to carry out the
study of this compound or others as far as seems desirable. Never-
theless, the preponderance of evidence seems to indicate in thymol,
and less clearly in resorcinol, a favoring effect wis sane! oa the
presence of the solid (Q).

DISCUSSION OF RESULTS.
The general situation developed by the results detailed above
may with profit be summarized in order to furnish a clear view of
the facts which are to underlie our further consideration.

x3 BOTANICAL GAZETTE [JANUARY

It appears in general that the presence of a considerable body
of certain insoluble substances in solutions of strongly toxic com-
pounds, both organic and inorganic in their nature, be they electro-
lytes or not, tends to decrease the toxic activity of the solutions in
question.

On the whole this ameliorating action is more clearly marked
in case the poisonous solutions concerned are dilute solutions of
strong poisons than when relatively concentrated solutions of weaker
poisons are concerned.

In general, filter paper and potato starch grains exert a more
marked modifying action than dense sand, glass, and paraffin.

A number of questions immediately arise concerning the relation
in which the active ions or molecules stand to this phenomenon.
It seems probable that no electrical phenomena are here necessarily
concerned. In the case of electrolytes such a situation might be
possible, but in thymol and resorcinol we have to do with electrically
neutral bodies. _

A more satisfactory explanation seems to lie in another quarter.
Chemists have long known that gases form a denser layer over the
surfaces of the walls of containing vessels than in the free space

away from the walls. Certain solids have the capacity of condensing —

on their surfaces larger volumes of gases than others, seeming to —
have specific properties in this direction. For example, platinum —

in a finely divided form condenses many times its own volume of
oxygen. A similar phenomenon has long been known in a practical
way in the case of solutions. Many years ago it was noted that bone
charcoal has the property of taking certain coloring bodies out of syrups,
thereby clarifying the latter products. The investigation of this

and similar phenomena has shown that molecules of many dissolved
substances are attracted to the surface of charcoal, quartz, and filter

paper, when these bodies are placed in these solutions, and in general

the greater the state of subdivision to which these insoluble bodies

are reduced, the more clearly may this condensation of molecules

from the solution be seen. This process by which a layer of greater i
molecular density is formed on the surfaces of solids immersed in __

solutions has been denominated by DuBots-REYMoND es (6
Not all solids have been found to be Tg ete “t

1905] TRUE & OGLEVEE—TOXIC ACTION OF POISONS 13

bodies and all molecules do not seem to lend themselves with equal
readiness to the adsorbing influence of solids (6).

Applying this doctrine of adsorption to the results observed in
our experiments, we find most points noted capable of explanation.
The earlier experiments of NAGELI (2) with Spirogyra in distilled
water obtained from copper containers and in other solutions con-
taining extremely minute quantities’of heavy metals seem capable
of satisfactory explanation. He observed that distilled water con-
taining minute traces of metal could be rendered harmless by placing
a quantity of some finely divided insoluble substance in the water.
He used paraffin, graphite, filter paper, glass, and other substances,
finding that thev interfered with the toxic action of the metals present.
Doubtless the quantity of harmful ions or molecules adsorbed was
so great as to reduce the number of ions or molecules in the free
body of the solution below the harmful concentration.

In the case of our lupines this same sort of action is also probably
at the bottom of the ameliorating influence seen to attend the mere
presence of a finely divided solid in solutions or strongly toxic agents.
By drawing a portion of the ions or molecules of the poisonous sub-
stance from the free body of the solution and by retaining these
molecules or ions in a molecularly denser laver over their surfaces,
the solids have removed from the free solutions enough of the toxic
substance to decrease its poisonous properties. Since in the case
of all of the insoluble substances used, paraffin excepted, the specific
gravity is greater than that of water, the solids sink to the bottom of
the culture vessel. As adsorption takes place, a somewhat gradual
process (6), molecules and ions are renewed from the solution above,
leaving it actually less concentrated than would be the case were the
solutions not provided with the solid. As a result, when the roots
of the test plant are introduced into the solution, it is found to have
a decreased poisonous action.

In the case of certain of the solids used in these experiments, it
seems not at all impossible that a small degree of solution may have
taken place. In experiments with crushed Bohemian glass this
was doubtless the case, since it has been repeatedly demonstrated
that such glass undergoes a slight solution even in.cold water. Unfor-
tunately we were not able to make conductivity determinations to

check up this factor.

14 BOTANICAL GAZETTE [JANUARY

In the case of the starch grains, the potato tubers were crushed
in a mortar, care being taken not to grind the material and crush
the grains. The starch that settled out on washing was repeatedly
washed with distilled water. Distilled water in which the starch
grains were allowed to stand gave no trace of the starch reaction
with iodine, a reaction which appears in case any considerable num-
ber of grains are crushed, exposing the more soluble interior layers.
Some hesitation, however, may be felt as to the insolubility of the
grains.

The question suggests itself as to whether all insoluble solids

exert a like adsorptive action under these conditions (6). No attempt —

was here made to test this question, since it was not practicable
under the conditions of experiment given to work with the accuracy
necessary in such a study. Furthermore, since adsorption is depend-
ent to a considerable degree on the amount of surface offered by
the solid, some way of taking account of this factor would seem
necessary. In the case of sand this could be done, but to carry over
these values to filter paper and starch grains, consisting of masses
of organic substance supposed by some (7) to have a submicro-
scopically porous structure, seemed to be impracticable. The lack,
therefore, of any practicable unit, whether of ,weight or surface, by

which to compare the solid substances used, made it best to content ©

ourselves with a rough qualitative method.
In general, the most marked results seemed to follow when filter

paper or starch grains were used. Whether this is due to the porous —

structure and consequently immensely enlarged surface, cannot here
be discussed.
Another point of interest in this connection concerns the nature

of the contact between the solution and the solid body. In all cases
except that of paraffin the solution is capable of wetting the surface _

of the solid, thus bringing the molecules of solute and solvent into
closest relation with the adsorbing surface. In the case of paraffin

the situation is different. The solution is not capable of wetting -
the paraffin even when in as close contact as can be obtained. As@
result, a surface of quite another sort is formed, more nearly com- =
parable in some ways to that bounding the surface of the solution e

where it comes in contact with the air. me

Mot tb oes ca ete al

|
|

iis oa

1905] TRUE & OGLEVEE—TOXIC ACTION OF POISONS 15

between the solution and the paraffin, the condition of the solution
more nearly resembles the surface film. In this case this film surrounds
the particles of submerged paraffin, and any properties found to be
possessed by the surface of contact with the paraffin should be in a
manner similar to those of surface films. However this may be,
the surface of contact with the paraffin seems to be such as to bring
about a condensation of molecules with a consequent removal of a
_ portion of the poisonous substance from the free solution and a decrease
of the poisonous properties of the solution. This happens, also,
in the face of the fact that the paraffin floats and must condense
the molecules in the upper part of the culture vessel. It might seem,
therefore, that the surface portions of the solution should be markedly
more toxic than the lower free portion. If this be so, the ions or
molecules seem to be held so firmly by the adsorptive force that they
are still in effect removed from the solution so far as the roots are
concerned.

AMOUNTS OF TOXIC SUBSTANCE REMOVED FROM THE FREE SOLUTION

BY ADSORPTION.

The results seen in the tables above given seem to show that
solid bodies are capable of adsorbing molecules or ions of toxic
compounds, holding them so closely that in effect they are removed
from the solution. We see many cases in which the growth rate is
so far accelerated by the presence of the adsorbing solid as to lead
one to infer that a very considerable portion of the toxic agent may
be adsorbed. In cases it is observed that reduction has so far taken
place as to practically relieve the solution of its toxic properties.
In some instances the solution, decidedly toxic without the solid,
becomes capable of supporting a more than normal growth in the
presence of the solid. Here, an abnormal acceleration replaces a
marked retardation, judged from the growth rate made in the dis-
tilled water check culture. Such instances are seen in mercuric
chlorid, os

20,000
solid amounting to an average of 2™™. When sand is added, the
acceleration is so great as to bring the average growth rate in twenty-
four hours to 16™™, 4™™ more than in the control culture. In the
case of silver nitrate a number of instances of this sort occur. In

. A retardation is seen in the culture lacking the

16 BOTANICAL GAZETTE [JANUARY

case the solution without the solid permits a growth rate above that
seen in the control, this increase is sometimes still further augmented
by the presence of sand. The results with copper sulfate show a
number of instances illustrating these different phases of action. —

It is a well established fact of general physiology that many
compounds which at certain concentrations act as harmful agents and
depress activity become in more dilute solutions accelerators of
activity. If dilution be carried on there arrives a point at which all
effect ceases, and only the effect of the solvent appears. This has
been pointed out by Couprn (8) for certain salts. We may represent

ey ace ae
7 =
es Be
Check growth rate eee —— ee e
bse ec ioe aa ip ys. Si AGM le sam ce at a > —_—
2 oa '
= = I |
é ae
z * " i
e wee i! 1
oh TEES ae [1 \
fatal depressant neutral accelerant neutral
Dilution =
Fic. tr.

the different physiological phases resulting from the progressive
dilution of a fatal concentration by a series of terms something like

the following. Almost instant death (a) is followed by a lengthening
period of survival (6) with depressed activity when compared with —

the control in water. This depressant phase is followed by a short
phase in which the growth rate approximates the control (c). This
passes over into the more extended acceleration phase (¢d) marked

age

pee ete ens :

by activity beyond that seen in the control. This acceleration rises 2
to a maximum and with further dilution falls off, until the solution

becomes practically distilled water in its action on the organism

(e). This may be represented diagrammatically as above (jig- 1); *
growth for the twenty-four hours in this case being indicated on the
_ axis of ordinates, progressive dilution (that is, the degree of dilution) _

on the axis of abscissas progressing to the right, the growth of controls .

in distilled water being regarded as unaffected by other extern
conditions, and the roots used assumed to be i in the same phase

1905] TRUE & OGLEVEE—TOXIC ACTION OF POISONS 17

_ development. While it may not be practicable in all cases to work
out all features of this generalized scheme with the very toxic salts
tested, the most striking points hold.

It is now not difficult to understand a number of phenomena seen
in results presented above. By preliminary tests not here detailed
a concentration of each substance was found which practically sup-
pressed growth and gave a physiological starting point from .which
to follow out the action of the poison along the line of progressive
dilution and also in the presence of solids. In most cases, the solu-
tions of the toxic compounds lie in the degree of concentration marked
by a depressant phase of physiological action. As molecules or ions
are removed by adsorption, the effect is virtually one of dilution, and
the depressant action becomes less marked; or when adsorption
removes a sufficient portion of molecules or ions to bring the solution
into the accelerant phase, instead of the depressant, an acceleration
above the control rate appears. The degree of acceleration depends
on whether adsorption withdraws such a proportion of the molecules
or ions as will leave the solution in its maximum accelerant phase or
above or below this. It should also be borne in mind that at least
theoretically, after the depressant phase has been passed, two quite
differing concentrations lying on either side of the maximum may
give the same physiological result. Experiments with solutions further
diluted will enable one to decide on which side of the maximum the
result may lie. Theoretically, a careful study of a long series of
slightly differing concentrations should enable us to ascertain rela-
tively how much of a given poison is removed by a given quantity
of an adsorbing agent. This can hardly be attempted with justice
from the results at hand, since they were obtained under conditions
rendering the necessary accuracy of experimentation impossible.
Nevertheless, I believe some rough idea may be obtained from the
work presented. It will suffice to point out a few instances, since
it is manifestly impossible to point out the application of this scheme
to all of the results obtained.

In the case of the mercuric chlorid (p. 5) —— is manifestly

low in the depressant phase of its action, the introduction of the sand
carrying it a manifest step upward, though leaving the situation

1§ BOTANICAL GAZETTE [JANUARY

still in the depressant phase. A solution =

Soo Permits a growth
rate of ro™™, still less than the control vente a Pte of ro%™". The
depressant phase of dilution is still represented, but is nevertheless
but little below the neutral point.

The addition of an adsorbing body therefore should carry it
into the accelerant phase. This is the result seen, and the growth
of 16™™ compared with the check seems to represent a rate probably
near the maximum. By making a series of experiments under condi-
tions like those met in this experiment, in which a graded series of
solutions of mercuric chlorid alone is used, we should be in possession
of a calibrated standard by.means of which we could estimate more
or less accurately the amount of the toxic molecules adsorbed. Similar
methods of interpretation applied to the results with silver nitrate indi-
cate with varying degrees of clearness a similar situation.

The results obtained with copper sulfate illustrate many of the
different phases of action. In some cases the presence of the solid
served not only to clearly decrease the harmful action of the salt,
but even to make survival possible in concentrations otherwise
promptly fatal.

In this connection a series of experiments was made to ascertain

approximately in how far the quantity of solid substance present —

might influence the action of a toxic concentration of a salt. For
this purpose a series of six cultures was prepared with like quantities

of copper sulfate. To five of them prepared sand was added in

different quantities. The growth rates seen in Table III, August
ro (p. 7), have been plotted in the following diagram (fig. 2)-
The growth rate in millimeters is indicated as before on the axis
of ordinates. The progressive addition of solids is shown on the
axis of abscissas. Although, unfortunately, no control culture was
made in this case, we may fairly accept 12™™ as average growth
in distilled water on the date concerned. This value is plotted as
the control result. The copper sulfate solution furnishing a basis
for this experiment is concentrated enough to kill the roots eas
usually permitting little growth. The addition of 40%™ of

adsorbs a sufficient amount of copper sulfate to permit half the

peas |

amount of growth expected of the control culture, and in addition

to make sgnsarege the survival of the test objects. Nevertheless, 2

atior oe

1905] TRUE & OGLEVEE—TOXIC ACTION OF POISONS 1g

depressant phase of concentration. The doubling of the amount
of sand brings the growth rate nearly to the normal rate of the control,
a slight depressant action still being seen. The roots, however,
survive in good condition, so far as their appearance indicates. The
increase of the sand to 120%™ results in so advanced a degree of
adsorption that the growth raie is thrown clearly into the acceleration
stage, perhaps near the maximum. Further additions still further
reduce the molecules in the free solution and show a stage of dilution

Check growth rate

Growth rate: mm. per 24 hrs.

m/35000 + m//35000 m/35000 =m/35000 + M/35000 m/ 35000
CuSO, CuSO, CuSO, CuSO, CuSO, CuSO,
- - - ~ +
408™ sand 80g™ sand 1208™ sand 1608™sand 2008™ sand

FIG, 2.

corresponding to the descending stimulus phase. The addition of
200%™ of sand shows this reduction of the copper effect at its limit.
The free solution is, physiologically, distilled water, so far as the
duration of the experiment allows one to judge. Here then the
progressive addition of quantities of a solid produces results paralleling
the progressive dilution of the toxic medium, the underlying cause of
these results being the gradual removal of molecules or ions from the
free solution by the insoluble body present.

In view of the evidence here presented and discussed, it is likely
that the process recognized as adsorption plays an important réle in
many problems of plant and probably of animal physiology. Experi-
ments by the water culture method cannot be used as a safe basis for
argument concerning soil conditions. The so-called absorptive

20 BOTANICAL GAZETTE [JANUARY

properties of the soil toward chemical compounds which form a
demonstration experiment in the usual elementary course in plant
physiology is in considerable part a demonstration of adsorption.
Indeed, the whole relation of the plant to its soil environment is
very largely influenced by this factor.

It is also probable that in the internal processes of the plant and
of the animal, similar forces are in operation, influencing, together
with diffusion and osmosis, the movements and distribution of mole-
cules and ions within the organism. For example, it would appear
not at all improbable that in cells containing starch grains or other
undissolved substances, the distribution of the various molecules and
ions of the cell sap is to no insignificant degree influenced by adsorp-
tion, and the transfer of molecules or ions from place to place may be
modified by this tendency of the walls, starch grains, and other porous
but insoluble substances to attach to themselves condensed layers of
molecules or ions. Indeed, it would be easy to see in a speculative
way many directions in which the processes of life may be affected
by the form of activity demonstrated in the above experiments to be
in operation under certain common conditions.

In nature the roots themselves and other absorbing organs fein :
surfaces for adsorption and the layer of solution about them is molec-
ularly more dense than the free solution. This applies in a much
greater degree to minute, submerged forms of life, as bacteria.
Absorption into the cell takes place, therefore, in general from a _
denser layer at the surface of the organism by the continued operation
of the energy of ad ion. This surface layer is continually renewed,
resulting in an ada absorption.

Another relation of importance is seen in the size of the organism.
The more minute, the greater would be the efficiency of adsorptive _ :
activity. This may aid in accounting for the extreme efficiency of the
‘minuter forms of life in operating on their medium. 4

Many suggestions occur as to divers other ways in which adsorp a

tion may play a part 1 the lives of living th oe
but to give them value the accumulation of more evidence is necessary a3
Further studies in this direction are in progress. a.

U. &. Dewan or AGRICULTURE, Washington, D. = and et Mu a
Universiry, —— Tl.

ee Se

905] TRUE & OGLEVEE—TOXIC ACTION OF POISONS 21

LITERATURE CITED.

- Kanna, M., Studien iiber die Reizwirkung einiger Metallsalze auf das Wachs-
thum hdherer Pflanzen. Jour. Coll. Sci. Imp. Univ. Tokyd 19:1-37.

1904.

. NAGEL, C. von, Ueber oligodynamische nena in lebenden Zellen.
Denkschr. Schweiz. Naturforsch. Gesell. 33:1. 1893.

. True, R. H., and Gres, W. J., On the phsaklbncel action of some of the
heavy metals in mixed solutions. Bull. Torr. Bot. Club 30: 390-402.

1903.

i. KAHLENBERG, L. and TRUE, R. H., On the toxic action of dissolved salts and
their electrolytic dissociation. Bot. GAz. 22:96. 1896

Baper, R., Zeitschr. Physik. Chemie. 6: 289.

. OSTWALD, W., Lehrb. d. allgem. Chemie 1:1080. 1891; a brief general
account of adsorption up to 1890. General discussions of the subject
are contained in the following articles about to appear: Briccs, Lyman J.
(a) On the adsorption of water vapor and of certain salts in aqueous
solution by quartz (to appear in Am. Jour. Physical Chemistry); (0)
Adsorption and its relation to soil investigations (as a Bulletin of U. S.
Department of Agriculture).

. Meyer, ArtHur, Untersuchungen iiber die Starkekérmer. 1895.

. Couprn, H., Sur la toxicité de sodium et de l’eau de mer a l’égard des végétaux.

Rev. Gén. Bot. 10:177. 1898.

. True, R. H. and HunKxet, C. G., The poisonous effect exerted on living

plants by phenols. Bot. poms a 48-51. 1898.

KU
.

ay

THE RELATION OF SOILS TO NATURAL VEGETATION
IN ROSCOMMON AND CRAWFORD COUNTIES,
MICHIGAN.*

CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY.

LXVI.

BURTON EDWARD LIVINGSTON,
(WITH MAP)

In a study? of the distribution of vegetation in Kent county,
Michigan, made by the author several years ago, the tentative con-
clusion was reached that the controlling factor in the distribution of
the plant societies on the upland is the amount of water in the soil,
and that this in turn is determined by two conditions, the nearness
of the underground water level to the surface and the water-retaining
or capillary power of the surface layers. This last factor is dependent
largely upon the size of the soil particles. It was thought well to
continue this subject farther north, using the same methods. Accord-
ingly in the summer of 1902 the work here reported was accomplished.

It covers nearly all of Roscommon county and the southern half of

Crawford county.
TOPOGRAPHY AND SOILS. }

The region consists of a series of ridges and depressions. The
former are sometimes several miles wide but more often narrow;
they are always comparatively low, seldom rising more than 50 to
| oat above the level of Higgins Lake, which lies in the middle of the

t Published by permission of the U. S. Bureau of Forestry and of the Michigan ae
Board of Geological Survey. A more detailed account of the investigations here
t,

conditions in this region has been published in the Report of the Michigan Forestry

Commission, 1902.
2 LIvINGsTON, B. E., The distribution of the plant societies of Kent ——
Mich. : 81 oe |

CEA] 314),
san
?

1) ONS

%
3
jx

HF ig

||
RS-
r|
i} i

5

i

| -Kivomer

Swamp and channel margin... Bash ut hore

LIVINGSTON (ON SoILs AND VEGETATION,

Moraine margin ......
Soil margin

ii

White pine .

\\ | : eee gta, Bes
11] | | [ = oe 4 ut, * cee

zt
j
i |

VV REE Re ee ee |

duoamMvuo

FI t
KINAOS NO “O2s0N
b = ee = "
ieee ‘ania ‘i
pa ae o— pee > a = SYiee 2 AR Ye

1995] LIVINGSTON—RELATION OF SOILS TO VEGETATION 23

area. ‘These ridges are moraines and between them are lower and
more level stretches consisting for the most part of plains which slope
downward from the ridge margin to the nearest stream. The sur-
face soil of these plains is almost pure sand in many instances, while
in others it contains enough more finely divided material to be termed
loamy sand. Gravel deposits are very rare and it is seldom that one
finds in the plains even scattering pebbles. The moraines are more
heterogeneous in composition, containing clay, loam, and _ gravel
as well as sand. An accurate description of these features is to be
found in the geological report referred to, and will be here omitted.
(See map.)

The soils are nearly all sandy, the only exception to this state-
ment being a few low clay areas and certain clayey portions of the
larger moraines. The surface soil of most of the ridges is gravelly
and loamy sand, the predominating sand containing a sufficient
admixture of finer particles to produce a marked difference in physical
properties from that of the true sand plains, while they also contain
pebbles and sometimes scattered bowlders. The slopes downward
from these ridges are of sand, either pure or loamy, seldom containing
many pebbles of any considerable size. The true sand plains,
which lie at the bottom of some of the valleys, contain little or no
loamy material and no pebbles. Their surface is a fine grayish-white
sand, which drifts readily by the. wind when loosened. Obviously
difference in degree of water-washing determines these different soil
characters.

3 Sandy soils are composed of coarse particles and contain much
| silica, loamy soils are of finer particles and contain considerable
quantities of alumina, while clay soils are of still more finely divided
materials and contain a much larger percentage of alumina. Since
all of this material was transported from the north by the glacial
| ice, and since it must have been quite thoroughly mixed by this
| agency, it is reasonable to suppose that, had it not been washed by
water during and after its deposit, it would be at least fairly uniform
in its mineral constituents. The washing process sorted the soils
according to size of particles, but also according to their chemical
nature. This is partly due to the fact that alumina breaks down
into fine particles more readily than does silica. It is also due to the

24 BOTANICAL GAZETTE [JANUARY

fact that in well-washed soils even the less soluble constituents are
apt to be actually dissolved and washed out to a greater or less degree.
Thus, phosphates and sulfates are less abundant in well-washed
soils than in those less thoroughly washed.

In this glaciated region, fine soils, such as clay, were either devaada
under the ice of the glacial epoch, and hence not well washed, or else
they were deposited from deep and very slowly flowing water. The
former variety usually contains many coarser particles, as loam, sand,
and pebbles. In the case of loamy soils a good part of the fine material
has been washed out, but a considerable amount remains with the
sand, so as to give it its loamy character; this is especially true of the
ridges. Sandy soils are still more thoroughly washed; the gravel
was left farther up stream, on the sides of the ridges usually, while
the clay was held in suspension to be deposited at a lower level, where
the velocity decreased.

On account of the difference in size of particles, which results
from water washing, there naturally follows a corresponding differ-

ence in the size of the interstitial spaces of the soil; the finer the

component particles, the smaller must be the spaces between them.

And because of this there comes to be a corresponding difference in” —
water-holding power and water-lifting power. The surface tension _
of water films is greater and hence more effective over small surfaces _
than over large ones, and the film surfaces are greater in coarse than -
in fine soil. Thus, the smaller the particles of any soil the more water —
it can hold and the higher it can lift this liquid from a lower level. ©
WaRMING; quotes WOLLNY as having shown that quartz sand con- |
sisting of grains over 1 to 2™™ in diameter can hold only one-tenth —
as much water as that with grains o.o1 to 0.07™™ in diameter. |
SCHIMPER‘ states that loose sand has a water capacity of 15.7 per |
cent. of its volume, while clay exhibits this property to the extent of |

49.9 per cent.

The nature of the soil particles themselves often plays an important 4
aia in ae the water-retaining and water-lifting power. :
der dkologischen Pilanzengrographic, iibersetzt von a

-, Lehrbu
Dr. as Roctinech. Bearbeitet von P. Graebner.

w —e
a > a

en een)

tat agi

1905] LIVINGSTON—RELATION OF SOILS TO VEGETATION 25

Especially is this true in the case of humus, which is composed of
organic débris, decayed plant parts, and to some extent of animal
offal. Pure humus has a great power to hold and lift water. This
is partly because of its very fine particles, but is also to be traced in
part to the actual penetration (by imbibition) of the liquid into the
intermolecular spaces of the organic substance itself. Thus, by
admixture of humus to a coarse (and therefore porous and permeable)
~ soil, the water capacity of such a soil is increased. The filtering
' power or permeability to water of a soil increases of course with
| _ decrease in its capillary power. Also its permeability to air increases
» in the same way.
4 A general exposition of this question of size of particles, water-
retaining power, etc., is to be found in either of the works just cited.
A much better treatment, however, has appeared in the publications
of Briccs5 and WuitNneEy.° The reader is referred especially to the
writings of the former author.

Chemical analyses of a number of Michigan soils have been made
and published by Kepzie.7_ Table I, showing chemical constituents
_ and water capacity, is compiled from his pages, the samples described
| being all from the portion of the state in which the present studies
_ were made. The first column gives the chemicals found in the soil,

in per cent. of dry weight, excepting in the case of water capacity,
which is presumably given in per cent. of total volume—although
this is not stated in the original papers. The upper tier gives the
_kind of soil, the location, and the nature of the forest cover.

:

5 Briccs, L. J., The mechanics of soil moisture. U.S. Dept. Agric., Div. Soils,
Bull. ro. 1897.

——., Investigations on the physical properties of soils. U. S. Dept. Agric.,
Field operations Div. Soils, 1g00-1901. pp. 418-421.

6WuitneEY, M., The Division of Soils. Yearbook U.S. Dept. Agric. 1897. pp.
120-125.
, Soil moisture. U.S. Dept. Agric., Div. Soils, Bull. 9. 1897.
see WHITNEY, M., and CAMERON, F. e chemistry of the soil as related
to crop-production. U.S.’Dept. Agric. Bull, 22. ak

7 Kepzir, R. C., The jack pine plains. Mich. Agric. Exp. Sta., Bull. 37. Also,
27th Ann. be Secy. State Bd. Agric. 1888. pp. 207-210.

of Michigan, Bull. 99. Also, 52d Ann. Rept. Secy. State Bd.

Agric. ae PP: 403-415.

26 BOTANICAL GAZETTE [JANUARY
‘TABEE &,
SHOWING RESULTS OF ANALYSES OF SOILS OF NORTHERN MICHIGAN.
Average, 6
a Tittaba- | P " s :
= Location Munsee a Gaylord —— Lake co. Fe
I Valley ; aa and Oscoda
e
z NATURE OF SOIL Clay Loam Loam Loam Sand Sand
2 ay (
@ | WaTeR capacity | 51.40 45-40 39.60 39.10 35-30 33-00
z >
“ | Forest tyre | Hardwood | White pine | Hardwood | Hardwood | Jack pine | Jack pine
Siea Ses 67.20 75-54 QI.52 69. 39 92.48 Q4.22
% | Alumina..... 6.31 10.62 2.93 8.35 oe | Ricwes
Se ee ee 7.91 3.80 0.90 5.80 EeSo 1.88 K
= OOP ae 1.64 0.94 0.40 rvES O.35 Oo. 37
eS "33 0.48 0.13 0.98 0.30 0.06
Sec taie J 1.85 1.96 o.61 1.95 O73 0.85
oan et ae see ¥.25 0.28 Eas 0.32 0.27
g PESC),.....'55 ©. 30 0.26 0.10 0.25 0.06 0.01
5 H;PO,....--- °.49 0.44 0.14 0.28 0.14 0.08
c
materials 7-48 2.97 2.20 4-73 1.22 2.16

more marked when the humus lies as a distinct layer on ee
than when it is distributed through the underlying soil. The humus
layer acts like a sponge filled with water, and allows the water to oad ee

It will be noticed in the table that sandy soils usually exhibit a
marked scarcity of soluble salts. This fact is perhaps to be explained.
by the “leaching” action of the percolating waters, as well as by the _
thorough washing to which these soils were subjected at the time of
their deposition. The water of precipitation percolates rapidly through ©
these porous soils and may often wash the soluble salts down vege
the level of the ground water, this process being termed “leaching.”

‘In humus-covered soils, it is probably not to the point to determine J - :
humus content and water capacity after the humus has been mixed |
with the loamy layers; the effect of the organic substance is very mu

depth of 20 to 30°™; nothing is said regar
reports from which these data are derived.
—— Ir aor the water capaci

of several ail samples ok

1905] LIVINGSTON- RELATION OF SOILS TO VEGETATION 27

lected by the author in Roscommon and Crawford counties. The
determinations were made in this laboratory. By “surface” is meant
the first 5 to 6°™, by “subsoil” the next 15 to 20°™.

TABLE II.
SHOWING THE WATER CAPACITY OF ROSCOMMON AND CRAWFORD SOILS.
WATER CAPACITY VOLUME
PER CENT.
— Township Soil type See Forest type
Subsoil Surface
I 21.4 Clay loam Plain Hardwood 43-5 74.1 humus
2 22.4 Clay loam Ridge Norway 45.9 40.0
3 25.4 Plain Jack 37.0
4 22.4 Loamy sand Plain Norway 43.8 oe
5 22.4 1 P) Hardwood 56.9 Mainly humus

It is noticeable that the subsoil of sample 1 has the same water
capacity as that of sample 4, but the surface humus of the former
brings its water-retaining power up to a point far above that of the
latter. The surface of sample 4 was apparently like the subsoil,
containing very little humus.

| TYPES OF VEGETATION.
_ The vegetation of the region may be subdivided into several types
or plant societies. These grade more or less into one another, but
there are a few places where an observer would be puzzled to deter-
mine any particular type. There are to be distinguished four types
on the upland and three on the lowland. Practically all of the area
under discussion has been lumbered. A virgin pine forest is almost
entirely unknown now, though some of the finest pines of the state
were cut from here. The hardwood areas have been left almost
untouched, excepting for the removal of the white pine originally
scattered through them in many places. The hardwood is now
tee rapidly removed, however, and it will not be long before there
be none left. In the lowlands, the merchantable arbor vitae
has very largely been removed, as has been also much of the spruce
and even considerable quantities of the tamarack. In the present
: wee there will be presented first a characterization of the
original vegetational cover, as well as this can be determined at the
¥ present time; then will follow a description of the present conditions.

aA 3

28 BOTANICAL GAZETTE - [JANUARY

1. Upland types.

A. Harpwoop TyPe. There is very little hardwood in the region
studied, but it is quite typical of all northern Michigan. Areas so
covered have not been so thoroughly lumbered as those covered
with pine forests. The original form of this type comprised the follow-
ing characteristic trees:$ Acer saccharum, Fagus americana, Tsuga
canadensis, Ulmus americana, U. racemosa, U. julva, Abies balsamea,
Betula lutea, some Picea canadensis and P. mariana, often scattered
Pinus Strobus of enormous size, together with such low forms as
Rubus strigosus,. Mitchella repens, Lycopodium clavatum, Taxus
minor, Amelanchier canadensis, Lappula virginiana, Hedeoma
pulegioides, Solidago caesia, etc. Acer, Fagus, and Tsuga.make
up three-fourths of the forest, now one and new another of the three
being dominant.

Lumbering has affected this type very little, excepting by removing
the white pine, and in some cases the hemlock. Hardwood lumbering
is now going on in the areas covered by this type; in these operations
everything is being removed which is merchantable. Fires have not
injured this form of forest to any extent, and the original humus
usually remains.

B. Wuite PINE tyPE. This is typical pinery, often containing
little besides white pine. Usually, however, there is an intermixture
of Norway pine (P. resinosa), and often of hardwoods. The type —
is quite sharply distinguished from the preceding, but not nearly so _
well marked off from the following type, into which it grades in many
places. As has been stated, there is at present hardly any of this
type in the region under discussion. In lumbering all the pine was
_ removed, and the subsequent fires have killed the young growth of
this tree as well as the scattering hardwoods. Over vast stretches
originally covered with white pine there are now no trees at all.
They are regions of dwarfed Quercus alba, Q. rubra, Acer rubrum, —
and a number of shrubs. The oaks and maples are rarely more eG
than twice as high as a man, are burned down every few years, and
exist here only because of the fact that they sprout from the roots, —
which are not always killed by the fire. ee

8 ‘The semnenclatere is that of Brrrron and Brown’s Illustrated Flora.

1905] LIVINGSTON—RELATION OF SOILS TO VEGETATION 29

thus possess enormous roots which are partially dead or dying,
gnarled, contorted, and deformed by frequent burning. For an
interesting description of how maples, oaks, etc., are able to attain
great age in this manner, and still not be over a few feet in height,
the reader is referred to BEAL’s® paper on this subject, which is
accompanied by convincing illustrations.

Among the lower forms occurring here may be mentioned the follow-
ing: Rhus hirta, Monarda fjistulosa, Pteris aquilina, Gaylussacia resi-
nosa, Vaccinium pennsylvanicum, V.canadense, Comptonia peregrina,
Solidago hispida, Hamamelis virginiana, etc. The ground between
the blackened stumps is now thoroughly covered by densely growing
sweet fern, huckleberry, and blueberry, the growth of the sweet fern
being so luxuriant that the numerous prostrate logs are often entirely
hidden from sight.

C. NoRWAY PINE TyPE. At the time of lumbering, this type
consisted mainly of the species for which it is named, but usually
contained scattering white pine and more numerous though often
dwarfed red and white oaks and red maples. The present’ aspect
of this type is much the same as that of the preceding. The two
oaks, red maple, and seedling Norway are the characteristic trees
now. Seedling Norways are more numerous than in the preceding
type, perhaps because of the greater number of seed trees here, as
well as the somewhat greater ability of this species to withstand fire
than that possessed by the white pine. The low plants are much the
same as in the last. Solidago caesia of that type is replaced here by
S. juncea; and Lacinaria scariosa is common here, as it was not in
the other group.

D. JACK PINETYPE. This is the most open of the series and occurs
on the most sterile sands of the area. The only trees are Pinus
divaricata, Quercus coccinea, Prunus virginiana, and seedlings of
Populus tremuloides and P. grandidentata. All but the pine and
oak are hardly more than shrubs. Besides the trees there occur on
the jack pine areas the following low plants: Pieris aquilina, Solidago
nemoralis, the three blueberries above mentioned (but rarely the
huckleberry), Arctostaphylos Uva-Ursi, Comptonia peregrina, Prunus

9 BEAL, W. J., Observations on the succession of forests in northern Michigan.
27th Ann. Rept., Bd. of Agric. Mich. 1888. pp. 74-78.

30 BOTANICAL GAZETTE [JANUARY

pennsylvanica, P. pumila, Andropogon scoparius, A. jurcatus, Dan-
thonia spicata, Lacinaria cylindracea, Salix humilis, Cladonia rangi-
jerina, etc. This style comprises the worst part of what is called the
“plains.”

2. Lowland types.

_ For the most part the swamps which were originally wooded have
not been denuded of forest. Where they contained white pine,
that was taken out, leaving the other trees, which protected the
undergrowth and soon produced a dense, almost jungle-like formation. —_
Within the past few years the merchantable arbor vitae and tamarack
have been removed from these swamps, but almost always enough
small trees are left to produce shade. Also, the swamps have not
been subjected to burning nearly so often as the uplands, and are
generally in much more nearly. their original condition than are the
latter. The three types may be described as follows:

E. OPEN MEADOW TyPE. This is partly open hay meadow
(largely of ‘“blue-joint,” Calamagrostis canadensis), partly ot bul-
rush and cat-tail marsh, and partly of sphagnum bog. It grades
into the other two types. :

F. TAMARACK-ARBORVITAE swamp. This is the most typical
swamp of the region. It contains Larix laricina, Thuja occidentalis,
Picea canadensis, P. mariana, Abies balsamea, which form dense _
and often impassable thickets. In some localities Larix occupies
almost all the ground to the exclusion of other trees, and in other
places the same is true of Thuja. But there is not nearly so much ~
tendency here for these two trees to form separate and distinct
types as is found cae south.'® There the tamarack seems t¢ —
occupy ee portions of t p lands which are most poorly drained, 4
the arbor vitae esinae in incailities where drainage is more thorough,
yet still not complete enough for the river swamp vegetation. Here — |
the question of drainage does not appear to play so important a part. ‘

G. Mrxep swamp. This formation is found near swamp Mar
gins, especially where the underlying clay is near the surface. Thus |

it often occurs along lines where the hardwood forest reaches dowr

be Si bie tte ees ee
= : eS

1905] LIVINGSTON—RELATION OF SOILS TO VEGETATION 31

toward a swamp. It may be looked upon as intermediate between
the tamarack-arborvitae type and that of the hardwood. There is
always a great mingling of species here. Among the trees are;
Larix laricina, Thuja occidentalis, Picea mariana, P. excelsa, Abies
balsamea, Betula papyrijera, B. lutea, Fraxinus americana, Tsuga
canadensis, Sorbus americana, Acer saccharum, Prunus serotina,
Pinus strobus, Amelanchier canadensis, etc., together with such low
forms as Rubus strigosus, R. villosus, Pteris aquilina, Lycopodium
clavatum, Taxus minor, Alnus, and Ilex verticillatus. The relative
proportions of the different trees vary from one locality to another,
so that nothing definite can be stated in this regard.

DISTRIBUTION OF FOREST TYPES.

The actual distribution of the types is shown by green lines on
the accompanying map. The upland types are denoted by letters,
each area bearing a letter to denote the type which it represents.
Thus, H denotes hardwood; W, white pine; N, Norway pine; J,
‘« +k pine; and these letters stand for types A, B, C, and D, respectively.
Of the lowland formations the open meadow is represented by the
‘conventional sign for marsh where it exists over broad areas.

The main facts of distribution are presented in the following
paragraphs. The upland and the lowland types will be considered
separately.

7 U plands.

The hardwood type occurs in this region always on soils which
~ contain considerable amounts of clay. Such soils are always covered
to a depth of several inches with leaf mold or humus, and in this layer
the seedlings of hardwood and hemlock grow and thrive.

_ The white pine type occurs in the clayey Murray Hills, on the most
clayey parts of Norway Hill, and on the great southeastern moraine,
in T. 21 N., R. 2 W. These soils are often as clayey as those of
-many of the hardwood areas, but are higher and therefore better
drained. It also occupies most of the gravelly ridge in T. 25 N.,
R.2W. Very often the swamp margins are occupied by this type also,
especially where the slopes are not abrupt, a condition which gives
_ humus a chance to collect in the sand.

32 BOTANICAL GAZETTE [JANUAR

The Norway pine type occupies gravelly ridges and loamy saudi
plains. The soil here is somewhat lighter than in the locations hei! ,
by the last-named type, but it is generally too poor for profita’!
agriculture. As will be seen by a glance at the map, most of + 1°
uplands here studied were originally covered by this type

The jack pine type occupies only the most thoroughly washed
the sand plains. The localities held by this open formation lie »
the valley of the Au Sable. The parts lying about the headwaters
of the Muskegon have abundant plains of loamy sand, but these
support the type of Norway pine. This fact has no connection w''”
the rivers themselves, however, for farther down the Muskegon «'-
to be found typical jack pine barrens. The soil of the jack pine ty)
is almost worthless for agriculture; it is light and dry, and where t»«
surface is broken it is apt to be wind-blown, and often forms sm
traveling dunes.

Lowlands.

The distribution of the lowland type was not worked out wi)
accuracy. Great difficulty was experienced in studying such areas, 7:
the swamps are often almost utterly impassable. The greatest are“:
of open marsh encountered are marked on the map, as alreac)
described. There are doubtless many areas of like nature whic’ —
were not seen at all, but these cannot be of very great extent. in
all the swamps the ground is covered with a layer of humus, i: :
of the nature of peat, and there seems to be no difference in ti is”
substance between the sand and the clay areas. Neither is there any |
apparent difference in the swamp enaiee whether it is upon sa
or clay. a

RELATION BETWEEN DISTRIBUTION OF FOREST TYPES AND THAT 75 |
| SOIL TYPES. ee
The upland types seem to be very closely dependent upon "|
nearness of the underground water level to the surface and upon "8° >
nature of the soil. The former factor determines at once whet)"
the vegetation shall be classified as upland or lowland. The ©
- tinction between these two classes is more evident on the surfs
than it is after closer study; it is difficult to state just how far ©
water level may recede from the surface and still support a lowl

1905] LIVINGSTON—RELATION OF SOILS TO \ °C’ ATION 33

type of the forest. Very few determinatior: \..~ ‘cen made in this
regard. Mayr" states for northern Wise. ns.) \i.t where the water
level is less than 2.5°™ below the soil suriacc, ‘\« vegetation is of the
swamp form; while if it is lower than est to. § “ the soil bears white
pine or some other upland type. This » no sandy soil. Warm-

ING’? has determined the depth of water eee in various soils in
Denmark. He finds Juncus and Carex forms holding the ground
until the water level is about 22.5°™ below the surface; with water
at a depth of 30 to 37.5°™, grasses grow well, forming what we should
term a moist meadow. With the water from 45 to 60°™ below the
surface, all grains grow well; this seems to represent our fertile
uplands. With the water still lower the soils becomes poor for
grains. Data from natural vegetation, so far as I know, have not
been gathered.

The more water there is in a soil, the less is the access of air to
the roots of plants growing therein. This is because air diffuses
much more slowly when in aqueous solution than when in the form
of a gas. Gas diffusion is checked by the filling of the interstices of
the soil with water, and hence most of the oxygen which reaches
roots in wet soil must do so by diffusing as a solute in the water.
Some ordinary plants cannot grow without rather free access of oxygen
to their roots, and so it follows that a soil saturated with water is
very poorly adapted to their growth.ts This is probably the main
reason why saturated soils are usually occupied by a vegetation
differing entirely in aspect from that found on soils which are drier;
thus we have swamp or lowland types of vegetation contrasted
with upland types. Swamp plants are able to live with a scanty
supply of air to their roots; but since upland plants cannot, it is
possible to have too much water in the soil for the well-being of the
latter. Thus areas with much water are occupied by typical swamp
plants, often perhaps because they alone are able to live in ap
situation.

™ Mayr, H., Die Waldungen von Nordamerika. Miinchen. 1890.

12 WARMIN' , Excursionen til Skagen i Juli 1896. Bot. Tid. 21: 59-112.
figs. 4. 1897. : am A to G. H. Jensen for an abstract of the article. *

13 WoLLNY, E., U. S. Dept. Agric. Exp. Sta. Record 4:528-543, 627-641. 1893.

34 BOTANICAL GAZETTE [anvar:

1. Factors of distribution in the uplands. s F

ORIGINAL DISTRIBUTION. Throughout the uplands, excepting

in the narrow swamp borders and in the low clay plains about Hough-
ton Lake, the underground water level is far from the surface, its ‘
depth varying from 3 to 25™ or even more.
Wherever the upland group occurs near the water level, it takes the
form of one of two types, either the hardwood (on the low clay) o1
the white pine (on the lowsand andloam). Farther above permanent
water the former of these types occurs only in one locality, in the |
northwestern part of the area, and there upon loam. Where the |
water level is not near the surface, the white pine type occurs only +
on clay and loam. The Norway type is found throughout the aree :
on loamy, or loamy and gravelly sand, and the jack pine type appears
exclusively on sand which is hardly at all loamy and thoroughly
washed. The distribution of the upland types just described may be |

ET ee eae

tabulated as follows: :
POSITION OF UNDERGROUND WATER LEVEL. 7:
Soil Near surface Deep a
EE ick ea At TE ge eee One ee eee eee :
Sandy loam......... |e ae ce ea
aay Te Lc es. Ry ees on ee B (A)*4 3
oe a ee: ro a a Oe ere

ee
i as

In the above table the different types are denoted by the letters
oe used in their description. It will be noticed that from clay
to sand, with water level deeply seated, we have a series passing fror
the white pine type to that of the jack, through the Norway. 4
single exception to this is the hardwood area on clay loam in T. 25 N-
R. 4. W, to be spoken of in a later paragraph. But with the wate”
Jevel near the surface, the series runs from the hardwood to the Nor
way type, the jack pine not occurring at all. This observation seem:
to agree with those made by Mayr’ in northern Wisconsin. H«
states that sand ridges rising out of the swamps usually bear whit:
pine on the slope, then Norway, and lastly the jack on the mos
elevated parts. The same author points out that white pine wil
grow on poor sand if the water is pias near the surface. The sam —
* seems to be true here also.
4 The hardwood in T. 25 N., Raw,
ts Mayr, HL, loc. cit. p. 207.

mn
a

) 1905] LIVINGSTON—RELATION OF SOILS TO VEGETATION 35

L. To explain the distribution of the different types, either of two
hypotheses may be resorted to. As has already been mentioned,
bes the finer the particles of the soil, the greater its power to lift and hold
Nie _ water above the underground level. It is well known, too, that some
3 soils contain more of*certain salts than do others. Thus, the reason
for the observed distribution on the uplands may be sought for
either in the water-retaining power of the soil or in its chemical
constituents.1° That the depth of the water table itself sometimes

_ plays an important part in determining plant distribution is shown
by the above table. Along a swamp margin the increased amount
of water may influence the plant growth directly, but how much of
the observed influence is to be considered as indirect is an open
question. The presence of water alters a number of other soil

factors.
First, it checks free access of air. Thus, if jack pine roots need
- more air than do those of Norway, this might explain why the former
_ fails along the swamp borders and the latter takes its place.
Secondly, with increase in water content there follows a more
equal distribution of the dissolved salts, for these can diffuse only
through continuous water films, and the greater the cross-section of
\ the latter, the more rapidly will diffusion take place. As a corollary
‘to this statement, it follows that ‘“‘leaching,” the washing down of
| soluble salts out of the upper into the lower strata, cannot occur to
/ any great extent in a soil which is constantly filled with water. More-
‘ over the upward diffusion of salts during dry times would probably
F more than counteract the downward washing during heavy rains.
_. The upper layers of a wet soil are apt to have more soluble salts
- \ after they have lain for a time than when first placed. This is of
i course on account of the evaporation at the surface, which increases
s [ the concentration of the soil solutions in the upper layers. Of course

16 For A papers on this subject se
THURMANN, J., Essai de ean appliquée 4 la chaine du Jura. Berne.

849.
NAGELI, C., Sitzungsber. Akad. Wiss. Miinchen. 1865.
UNGER, Ueber den Einfluss des Bodens auf die Verteilung der Gewaichse. Wien.
1836.
\ more recent paper dealing with this penser of soil physics and soil chemistry
,as influencing vegetation is Cow1Es, H. C., influence of underlying rocks on the
,character of the vegetation. Bull. Amer. nr of Geog. 2:1-26. el
i

4

this indirect effect cannot be exhibited unless there is a sufficient
amount of salts in the deeper lying soil. But in a glaciated region
such as this, there can be little doubt as to the presence of these rela-
tively near the surface.

Thirdly, the checking of air access, coincidént with the filling of
the pores of the soil with water, must check the process of oxidation
and thus accelerate the formation of humus. This seems to be a
very important factor in determining vegetation.

Fourthly, the growth of micro-organisms in the soil, bacteria,
etc., takes place much more rapidly in a moist than in a dry soil.
However, excess of water is deleterious to the growth of many of these
organisms, so that a soil may be too wet for them; but flooding is not
so fatal in sand as in finer soils.*?__ It is well known that soil bacteria
and mycorhizal forms are very important in increasing the amount
of nitrates in the soil, and thus it appears that a moist soil, even a wet
soil, if it be sandy, will gain nitrates much more rapidly than a dry one.

The comparative rapidity of temperature changes in moist and
dry soils may have considerable importance in regard to plant growth.
A dry soil is subject to much more sudden and rapid changes of tem-
perature than is a moist one when the soils are exactly alike, as I
have shown by actual tests.

The points of difference between a moist and a wet soil may be
tabulated as follows:

;

36 BOTANICAL GAZETTE [JANUARY "

b.

Re nae
ee Re ae %

Condition of soil Dry | _* Moist Wet
Water for rocts. Too little Enough for most More than needed for
am : plants most plants
Soluble salts originally near sur- Sometimes .
face. f leached farther down Still near surface | Still near surface
- Soluble salts originally in lower é —— 2S - - ae co oe
layers Lo. cen jeac farther down artly in upper layers Partly upper layers :
Ems. content... 5024 2S None — hoch Gs i ei *
Pepa io Puenben uae stwe = Some ——
Miao eee ae ae
‘eperaare change. 22. : er i “Les raid Slow

Whether these points are Paar by nearness of the under-

ground water level or by the capillary power of the soil, there appears _

| to be no doubt that the amount of water in the layers near the sur
- practically determines the nature of the vegetation in this

sabi ee ae

:) - Besides the general discussion of this matter to be found in
¥ Gams, E. Action de Veau du sol sur Ia végtation. Rev. Gén. et a

1905) “LIVINGSTON—RELATION OF SOILS TO VEGETATION ay

; - WARMING and SCHIMPER (loc. cit.), the reader may refer to GArn™® and
e HEpGcock."?

Of the upland series, the hardwood type of vegetation seems to
need the most water, the most soluble content, and the most humus.
Probably this is the reason why this type occupies the moister soils
of the upland, no matter whether these are moist through nearness
to the underground water table or through the lifting power of the
| 2 soil itself. The types of white, Norway, and jack pines seem to
& require less water in the order of their arrangement. Probably
the Norway and jack pines require more air in the soil than either
the hardwood or the white pine. The typical tree forms of both
these last-named types occur in the mixed swamp quite commonly,

| but I have yet to see either Norway or jack pine in soil which is wet
4 the greater part of the year. Throughout the region it seems that

each type occupies soils which correspond in water content to its
ze needs. It must be remembered here that a sandy soil near the water

level may contain much more water than a loamy one where the
water is farther from the surface. This idea offers a possible explana-
E tion for the occurrence of hardwood on low loam in T.25 N., R. 4 W.
+ Addition of surface humus has also, perhaps, raised the water holding
a ' power of the soil to the neighborhood of that manifested by clay
a ‘itself. The subsoil is such here that the white or Norway type might
f be expected.
PRESENT DISTRIBUTION. The statement so frequently met with
that white pine will not come up after it has once been cut off and
_ the ground burned over seems to strike wide of the truth in this
region. I have visited almost every square mile of the uplands and
am thoroughly convinced that scattering seedlings of white pine
are now evident on practically all areas originally covered by that
species, which have not been recently subjected to the action of
fires. Seedlings of the Norway, however, are now more numerous on
these areas than are those of the white pine itself. They are plentiful
throughout the region on light soils excepting the very lightest.

é

8 Gain, E., Recherches sur le réle physiologique de ’eau dans la végétation,
Ann. Sci. Nat. Bot. VII. 20:65-215. 1894.

2 EDGCocK, C. G., Botanical Survey of Nebraska: Studies in the vegetation of
. the a Hi. Linasethe: 1902

ager.

38 BOTANICAL GAZETTE [JANUARY

Fires prevent the young growth of the white pine and also prevent
humus formation. Thus, as long as the latter are allowed to continue,
the water capacity is not apt to rise and the growth of nitrifying —
bacteria is not apt to increase; but the presence of a few white pine
seedlings is evidence that the species can grow if protected. Indeed
the best young stands of any kind that I have seen are of this tree, f
and they promise exceedingly well for reforestation.

As has been said, the Norway pine is coming in quite freely in
the areas originally covered by this species and the white pine. The
degeneration goes no further, however; I have almost never seen even |
individual jack pines appearing in any of these areas. Indeed, |
there is some evidence that the Norway pine is gradually advancing ;
its seedlings into the areas held by the jack pine.

The hardwood forest reappears quite rapidly when cut. This ©
is doubtless in part due to the fact that this material does not burn ~
so readily nor so violently as do the pines. The scattered white ,
pines which formerly characterized some of these forests in the eyes |
of the lumberman are not returning. They are perhaps only a relic
of a past generation of forest.2° Hemlock is reproducing well and
will return with the beech and maple if through lack of humus the
soil does not become too dry for the seedlings. The sugar maple is
the best for reclaiming cut-over lands. Its seedlings stand close
together and do not seem to suffer from one another’s shade. :

The work of SHERRARD?! in this region resulted in a map and |
statistical study of the tree growth of township 25 N., R. 4 W, as
well as a general discussion of the forestry conditions of the reserve-
The township thoroughly studied by this author originally contained
practically no white pine, but the other types are well represented. ;
SHERRARD’S “oak flat” and “oak ridge” are all original Norway
pine land. Forstatistics of saeisite etc., the reader is referred to his |
paper. | |
2) caeSep the ‘eutline’s Kank county paises, loc: 08; ako Wanrvoxn, BN. TH
genetic ees ee forests of southern Michigan. Bor. Gaz. 31:289-325-

2 2* SHERRARD, iT es nmnian eared sus Report Mich. Foresty :

} 31905] LIVINGSTON—RELATION OF SOILS TO VEGETATION 39
BS 2. Factors of distribution in the lowlands.

y

degree the conditions of soil moisture. In this case it seems better,
however, to arrange the types in the reverse order and to present
+ them as following conditions of drainage. As has been stated, all

three types are composed of forms which can withstand a great deal
of moisture. No positive evidence can be given as to whether or
| ‘not there is any difference in water content between the soils of the
open meadow and those of the tamarack-arborvitae forest. Noth-
ing has been made out regarding the conditions which decide in
favor of one or the other of these; but the mixed type is always found
on the better drained portions, where there are hummocks raised out
of the saturated soil, and where the general level is a few inches
: higher. Often this better drainage seems to come about merely by
accumulation of vegetable débris, a fact which suggests that perhaps
in time the conifer swamp might give way to the mixed, and at last
possibly to the hardwood upland type.

Attention has already been called to the fact that this series of
types have not been seriously altered by the hand of man. The
large white pines have been taken from the mixed swamps, as have
also many of the most valuable tamaracks and arbor vitae, but the
forest conditions have not generally been destroyed.

| The three types of lowland vegetation seem also to follow in some

RELATION OF THE VEGETATION TO THAT OF KENT COUNTY.

The predominance of the pines in the region under discussion is
an expression of the fact that the flora here is a typical northern one.
Only one pine (the white) is found in Kent county, and there it
grows in poorer soil than it holds here. The presence of hickory
_ and the better growth of the black, red, and white oaks in the more
southern area is an indication of a more southern flora. The factor
which keeps the jack pines out of the more southern county may
be a climatic one; this species occurs with the white pine on the
dunes at the extreme southern end of Lake Michigan.

The hardwood forests of the two regions are very nearly the same
in character. In the northern part of Kent county the hemlock
begins to be an important tree in this group, as it is farther north.

¥
Ie

40 BOTANICAL GAZETTE [JANUARY |

A study of the transition zone between these two areas will be i

necessary before the working out of the exact relation of the various
societies can be attempted.

RELATION OF THE VEGETATION TO THAT FARTHER NORTH.

The student of plant distribution who has formed any definite —
theories as to the relation of plant societies to topography and soils —
from the southern part of the state and from Indiana and Mlinois
will be much surprised if not actually shocked by a visit to the region |
of the straits of Mackinaw. On the island of Mackinac, for example, |.
occur most of the tree forms found in Kent county, but growing :

apparently without relation to the soil condition. Norway and ]{;

white pines, beech, sugar maple, red maple, tamarack, arbor vitae, |
balsam fir, basswood, etc., will be found growing side by side on the |/
well-drained uplands in clay or sand or loam, or even partially bare }
rock. As to the reasons for this nothing can be said before more of

the area has been studied in detail.

CONCLUSION.
It appears from these investigations that the main factor in deter-
mining the distribution of the forests on the uplands of this region
is that of the size of soil particles, the sorting of which dates back
- almost entirely to the close of the last glacial epoch. The size of
particles determines the amount of air and moisture in the soil,
and these in turn determine the amount of humus formation and
the growth of nitrifying organisms, and perhaps also to a certain
extent the amount of soluble salts in the surface layers.

3 ae rake
se ee

A factor of less importance, because applicable only over small j-
areas, is the nearness of the underground water level to the surface. )

This affects the uplands only along swamp borders.

In a broad way, physiography may be said to determine the
vegetational distribution here.77 The physiographic features are
largely ones of glacial topography or are traceable directly to these.
logical factors, in one way or another, have of course dete — ¢

aay ae 5 PaaS ae ly Har f this idea see CowLeEs, H. C., The

oS ecology of — and vicinity. Gaz. po hig Ans, The ast

mc.

1905] LIVINGSTON—RELATION OF SOILS TO VEGETATION 4I

the distribution of surface soils and the distance below the surface
of the ground water level.

It is probable that many dry soils may at length become moist
enough to support one of the more moisture loving types of vegeta-
tion, simply by increase of humus content, which must go on slowly
at first but more rapidly as the amount increases. This is merely
an application of one of the general principles of forest succession
pointed out by WHITFORD (/oc. cit.).

\ The lowlands are covered with a vegetation complex of species
iuch that they can bear excess of water and paucity of oxygen in
ae soil. From the open meadow and coniferous swamp we pass,
tvith better and better drainage, through the mixed swamp to the
flardwood or the white pine of the uplands.
} It appears that the natural reforestation of the pine areas, with
‘Norway pine, and to some extent at least with white pine, will take
place if the fires can be suppressed.

THE UNIVERSITY OF CHICAGO.

Note.— Since the preparation of this paper, a similar conclusion has been
ached in regard to sterility in agricultural soils by WHITNEY and CAMERON,
ull. 22, Bureau of Soils, U.S. Dept. Agric. 1903. Also, it has been shown by
‘actual field test that coarseness of soil particles alone can produce sterility in spite
lof a plentiful supply of water. See LrvincsTon and JENSEN, An experiment on
jthe relation of soil physics to plant growth. Bor. Gaz. 38: 67-71. 1904.

¢

NEW AND NOTEWORTHY WESTERN PLANTS. Il.

&

A. D. E. ELMER.

In the spring and summer of 1902, while engaged in a collecting,

trip in western Santa Barbara, eastern Ventura, and portion of

northeastern Los Angeles counties of California, a number of note- }

worthy plants were collected as well as new species discovered. Or
the train from Stanford University to Santa Barbara, where my
headquarters were made, I was impressed with the surroundings of
Surf as having peculiar physical characters. Having no time to
examine the vegetation then, I observed nevertheless a palmlike
bush with large golden yellow flowers which I did not notice farther
south. Curious to know this plant, I made a side trip to Surf in
May, and found its local sand-dune flora the most interesting of all
the vegetation in the counties visited. Surf is near the coast and has
a stretch of sand dunes a few miles in length and a half mile in width.
The breakers are active most of the year, and the winds at certain
seasons are and strong; at such times the sands are shifting
almost SEE The temperature is always moderately cool.
This place is one of the few points along our coast possessing more
striking peculiarities than the ordinary sea beaches. It is a place
where the oceanic influences are predominant. One can imagine
similar conditions on the Santa Barbara Islands off the mainland
of southern California. It is my conviction that if a critical study
were made, comparing the island flora with that of the mainland

such as we have at Surf, a striking similarity would be discovered.

_ Of the following species, all are described as new excepting the first

four, and the first seven were collected in the vicinity of Surf. Some |
of these seven have already been reported from the islands, and the

others may be assumed to belong to the island flora.

= the snd hills at hac and it was collected again in the Mohave’

\ CALIFORNICA Hary.—This peculiar crucifer I collected .

1905] ELMER—NEW WESTERN PLANTS _.. 43

most northern limit of its distribution along the coast, and it may be
expected on the Santa Barbara Islands.

MESEMBRIANTHEMUM CRYSTALLINUM Linn.—This_ beautiful
ice plant seems to love halophytic conditions along drifting sand
beaches continually swept over by cool winds from the ocean, though
it has also been reported from the Mohave desert by Mrs. BRANDE-
GEE, according to PARIsH. Along this coast its range extends from
Surf, Santa Barbara county, California, southward through Lower
California, and on the islands off the coast. It is an old world species,
indigenous to the coast of Africa.

LEPTOSYNE GIGANTEA Kellg.—This is the plant observed from
the moving train as a palmlike bush with large golden yellow
flowers. Aside from the original publication, the Botany of Cali-
jornia reports it from Guadalupe Island and San Miguel of the
Santa Barbara Islands. The Synoptical Flora gives it from the
mountains near Santa Barbara and San Miguel, and the islands off
the coast. This citation is rather obscure, and it is to be doubted
whether this plant was discovered on the mainland as early as that
date. In Bull. Calif. Acad. 4:402. 1887, GREENE writes that he
found it frequent on cliffs toward the sea on the north side of Santa
Cruz Isand, and reports it flowering as early as February and March.
Miss AticE Eastwoop reported it for the first time from the main-
land, and in Zoe 4:286. 1893-4, she writes that JARED found it
growing abundantly near the old wharf at Point Sal, Santa Barbara
county.

CEANOTHUS ImpPRESSUS Trel.—About twenty-four years ago
JARED collected southwest of Guadalupe toward Point Sal, Santa
Barbara county, California, a Ceanothus of which TRELEASE wrote
in Proc. Calif. Acad. Il. 1:112. 1888:

C. impressus,n.sp. Villous with short spreading hairs: leaves broadly ellipti-
cal to nearly orbicular, 6-8™™ long, loosely villous, especially on the veins below,
the upper surface deeply furrowed over the midrib and several pairs of lateral
nerves, the slightly glandular margin very revolute, appearing as if crenate:
peduncles about 1ro™™ long, scaly toward the base; inflorescence subglobose,
compact: fruit not seen.—Santa Barbara county, Cal.

About fifteen years later Mrs. IpA BLocHMAN collected it again
near the same place. While botanizing a few miles north of Surf
station I found a Ceanothus (distributed under number 3870) which

aS ae — aia. barely cleft

44 BOTANICAL GAZETTE [JANUARY

seems to answer well to the above diagnosis. Its range appears to
be restricted to this coast, and it should be looked for on some of the
northern Santa Barbara Islands. In Proc. Calif. Acad. II. 4:202.
~ 1894-5, Mrs. KATHERINE BRANDEGEE writes that in her opinion it
appears to be nothing but a stocky southern form of C.dentatus T. &
G. growing on unsheltered sandhills. In the same paragraph she
states that C. dentatus in its typical form appears to be confined to
the vicinity of Monterey. Still there are quite remarkable differences
between the two forms or species, and to one observing them in the
field they would not for one moment be taken for the same. C. den-
tatus at Monterey has not the dense prostrate habit. The leaves of
C. impressus are broadly elliptical to nearly orbicular, while in C.
dentatus they are oblong-cuneiform; furthermore, the branches of
C. impressus are rigid and spinescent, while in C. dentatus they are
flexible and not spinescent—a character also contrasting C. ihyrsi-
florus Esch. and C. sorediatus H. & A.
Malacothrix succulenta, n. sp.—Succulent maritime perennial
herb, with long fleshy creeping and numerously branched stems
forming dense mats: stem above the sand branching near the base

or at least below the middle, 1-2°™ high, rarely erect but usually reclin- -

ing, when young floccose especially on the concealed portion, soon
becoming glabrous: lower leaves persistent after they become dry,
linear-oblanceolate, sessile when mature, glabrous on both sides,
5°™ long, 8™™ across the widest part, obtusely rounded, rarely
undulate or dentate along the upper margin: scapes gradually
thickened at the apex, usually branching from a little below the
middle, its bracts minute, weak and sub-flexuose: heads solitary,
turbinate, 2°™ long, about that in width at the top: involucral bracts
chiefly in one series, 1o™™ long, thin, glabrous, linear, loosely imbri-
_ cated, subtended by numerous smaller triangular bracts of the scape:
flowers many, all with lemon-colored ligulate corolla: receptacle
naked, flat: achene smooth, 1.5™™ long, with semi-truncate or short
obtuse ends, bearing a soft silky white pappus whose bristles are
finely scabrous and equally 6™™ in length: corolla 7™™ long, gradu-
ally tapering into the 4-toothed ligulate expansion, much exceeding
the pistil, its tube 3™™ long and glabrous: stamens exserted; anthers
linear, united, 3™™ long; reaps ae — distinct, et

i ae ipsa kee e:
Pe ee ee

1905] ELMER—NEW WESTERN PLANTS 45

Dunes and terraces about Surf, Santa Barbara county, California, May 1902.
Type specimen, no. 3639, in Herb. Stanford University.

It is now just sixty-six years since NUTTALL first discovered on an island in San
Diego Bay a Malacothrix which Torrey and Gray described (Fl. N. Am. 2: 486,
1838-40) as M. incana. Fifty years later GREENE rediscovered it growing
abundantly on San Miguel, the smallest and remotest of the Santa Barbara
Islands; he also found it on the western extremity of Santa Cruz Island, a dis-
tance of 200 miles northwestward from its type locality.. My plant from Surf is
similar, but in my opinion is specifically distinct, though very closely related to
M. incana.

Carduus maritima, n. sp.—Succulent maritime biennial or peren-
nial, from strong thick tap roots: stems 1™ long, much branched
from the base and up to the middle, giving the plant a rounded
bushy appearance; branches covered with long white appressed
hairs and terminating in 1-3 heads: leaves thick, felty, densely and
thickly lanate pubescent, 1-2°™ long, the blade proper ovate in out-
line, the lower one-third abruptly narrowed, its base semi-auriculate,
the entire margin irregularly and dentately lobed and beset with
straight needle-pointed spines which are more numerous toward the
base than on the coarse lobes of the blades: heads large, dull white,
5°™ long, nearly that in width, loosely corymbose: bracts many,
imbricate, nearly equal in length; the exterior ones similar to the
leaves in lobation, spinescence, and pubescence; the interior ones
linear-lanceolate, sharply acuminate, the lower and inner surface
wholly glabrous, finely spinescent on the margins above the middle:
the dense bristly hairs of the receptacle not hispid, dark brown,
12™™ Jong: achene 6™™ long, flattened, narrowly obovoid or cuneate;
brown and shining, usually somewhat curved: pappus bristles inter-
laced with fine secondary ones except at the apex, 2°™ long, dull
white, attached as a whole to the apical rim of the achene: corolla
tubular, 2.5°™ long, segments 4™™ long, the tube funnelform or
slightly inflated immediately beneath the lobes: anthers united and
pubescent, er long: stigma barely notched.

Surf, Santa Barbara county, California, May 1902. Type specimen, no.
3631, in oa Soaked University.

The habitat of this species is strictly ee and most probably it can be found
on the islands and possibly in other localities along the coast. I found only a few
plants on the most exposed dunes, on which they formed low spreading bushes
not very dissimilar to the Russian thistle. Its low succulent halophytic habit,
remarkable pubescence and peculiar lobation and spinescence of its leaves readily
distinguish it from all other species of this —

46 BOTANICAL GAZETTE [JANUARY

Monardella crispa, n. sp.—A low strongly aromatic shrub, 3°™
high, the lower woody branches numerous, giving it a small dense
bush-like appearance; younger branches lanate pubescent, older ones
with thin coarsely and irregularly checked bark: leaves sparsely
lanate pubescent, somewhat succulent, fascicled at regular intervals,
more strongly punctiform on the dorsal surface, the larger ones
2-3°™ long and 1°™ wide, undulate or coarsely crisped along the
margins below the rounded apex, oblong or narrowly oblanceolate
with a long attenuate base; upper pair of leaves ovate to oblong,
without the slenderly narrowed base: heads solitary upon the stems
or branches or in a series of two or even three, sub-globose, 2-4°™
across, upon densely woolly scapes 2-5°™ long: bracts 1c™™ long,
apex acute or rounded, thin, purplish, the 3-7 nerves conspicu-
ously ascending, the outer surface scft pubescent, almost equaling
the calyces, the exterior ones elliptic to broadly oblanceolate, the
interior ones narrower and a trifle shorter: flowers 18™™ long, upon
pedicels 1™™ long: calyx tubular, 6™™ long, woolly pubescent, 14-
nerved, the 5 acute teeth 1™™ long: corolla 13™™ long, light pink,
obscurely bilabiate; upper lip broad, deeply emarginate; lower one
divided into 3 ligulate segments; exterior corolla tube 4™™ long,
exceeding the calyx, sparsely set with fine short recurved hairs:
stamens 4, spreading and much exserted: style included, bearing a
short-lobed stigma.

Surf, Santa Barbara county, California, May 1902. Type specimen, no.
3965, in Herb. Stanford Universi rsity.

This mint is a maritime species, and 3h Sent ee eines ee eee
beaches. Without much doubt it may also the Santa Barbara
Islands where the sea influence is strong.

Monardella robusta, n. sp.—Faintly aromatic, tall, woody, 1-2™

high, prostrate or supported by other undergrowth: stem almost

scape-like below, with a few branches toward the top, covered with a

; thinly checked but not shredded bark; younger branches dark brown,
- sparsely pubescent, usually terminated by a solitary head though

often bearing smaller axillary ones beneath them: leaves opposite,
the larger ones 5.5°™ long including the 5™™ long petiole, 15™™

_ wide, lanceolate, margins involute, coriaceous, nerves radiating

from the midrib and ascending, prominent beneath, sunken above,

entire, or eet oor’ vom ane or only sparsely

1905] ELMER—NEW WESTERN PLANTS AT

hirsute on dorsal surface, thickly white lanate pubescent beneath,
sub-scarious; axillary leaves shorter and narrower, fascicled, without
a distinct petiole: heads 2°™ or more across, less than that in height,
solidly flowered, nearly sessile or pedunculate; the lower bracts
shorter and broader, 8™™ long, broadly elliptic, acute or obtuse,
ascendingly nerved mainly from the broad base, soft pubescent,
especially upon outer surface: flowers short pedicellate: calyx tubular,
8™™ Jong, 14-nerved, terminated by 5 acute teeth 1™™ long: corolla
at least 2°™ long, its tube exceeding the calyx teeth by 4™™, scarcely
bilabiate, lower segments 6™™ long, exterior of exserted tubular part
set with fine retrorse hairs: stamens 4, exserted: style slender, mostly
included, 2-notched.

Rattle Snake Canyon, a few miles back of Santa Barbara towards the Santa
Ynez Mountains, Santa Barbara county, California, August 1902; rare, and only
found in dense thickets of underbrush. Type specimen, no. 3728, in Herb.
Stanford University.

Encelia actoni, n. sp.—Shrub 1™ high, moderately branched,
forming rather unique individual, bushes; wood soft; bark thin,
smooth, grayish white, becoming finely checked and ultimately
shredded: leaves not crowded, nor are the secondary ones fascicled,
3°™ long, 2°™ wide, ovate, acute, chiefly entire, 3-nerved at base,
alternate, without stipules, cinereous pubescent, upon petioles 1°™
long: heads 3°™ in diameter, solitary upon long cinereously pubescent
scapes barely bracteate. below the middle: involucral bracts nearly
of the same size, 7-10™™ long, 3-5™™ wide, imbricate in two or
three series, hispidly pubescent and sub-glandular on the exterior,
glabrous on the interior: ray flowers pubescent, tube bearing a pistil
and gradually expanding into a truncate ligule 8™™ long, not numer-
ous, with a sterile achene 4™™ long; ligule proper finely pubescent
- on the outer surface, light yellow, many-nerved; disk flowers perfect
and fertile, subtended by bracts; these 1o™™ long, strongly enfolding
the achene, with a prominent midnerve besides the 1o-14 lateral
nerves terminating in an acute glandular pubescent apex whose
edges are fimbriate: corolla tubular, equaling the achene, lower one-
fourth constricted, terminating in rather acutely thickened recurved
apical segments: stamens included; filaments 1"™ long; anthers 5™™
long, introrse, with triangular apex, and minutely sagittate base:
style 2-cleft, stigmatic only upon the upper outer side of the lobes;

48 BOTANICAL GAZETTE [JANUARY

achene 5™™ long, much compressed, narrowly obovoid or cuneate,
only the edges densely ciliate especially toward the apex.

Acton, Los Angeles ios California, June 1902. Type specimen, no.
3724, in Herb. Stanford Univers’

This interior species differs Fit the seacoast E. californica Nutt., and can
at once be distinguished in the field by its conspicuous light grayish bark, its
cinereous pubescence, its fewer and larger heads, and its individual habit.

Chrysopsis californica, n. sp.—Perennial, 4-6¢™ high, from
woody base: stem long silky pubescent, fastigiately branching, termi-
nating in a subcorymbose inflorescence: leaves more or less fascicled,
sessile, often clasping, the average length 2°™ and width 5™™,
though often much shorter or longer, granular surface covered with
dense white silky pubescence, edges becoming involute: heads 2°™
broad, upon short leafy ascending peduncles: involucral bracts
many, villous pubescent on exterior surface, linear or narrowly
lanceolate, regularly imbricated in several series: receptacle naked:
ray flowers: pappus 2-seriate; the upper of reddish-brown scabrous
bristles 6™™ long, the lower lighter colored; minutely paleaceous,
only 0.5™™ long: corolla 2°™ long, lower half tubular, upper expanded
into a semi-obovate or oblanceolate ligule which is closely and minutely
3-toothed at apex; opposite this primary ligule is usually present a
secondary one which is shorter, more linear and inserted upon its
throat: disk fowers: pappus bristles in one series, 2™™ long: corolla
glabrous and tubular, with rich yellow acute segments, 1.5™™ long:
filaments short, separate, inserted upon a short thick base: style
exserted and deeply cleft, the branches triangular acuminate and
stigmatic only on exterior surface; achenes apparently all fertile,
4™™ Jong, short pubescent, slightly compressed or 4-costate.

a Sandy soil —e beach, Gaviota, Santa Barbara county, California, May

— Type s, en, no. ne Eee S ge University.
e a The piace ke is rare, and w was collected at same station where kee

ee Deasanaen Ndi: n. sp.—Annual or at “most biennial: stem
Le brown, generally =, chiefly branched toward the top, hispid
wee : : ascending, leafy,the lower ones shortest,
“occasionally hedring subglomerate claites. below the terminal one:

leaves ovate, lanceolate or linear-lanceolate, short hispid pubescent

erence more ee z-5™™ long, the smaller

1905] ELMER—NEW WESTERN PLANTS 4Q

ones entire, the larger ones remotely dentate: heads turbinate, 8"™
long, upon leafy bracteate peduncles: 5 involucral bracts 6™™ long,
uniserial, narrowly oblanceolate, the apex sparsely hispid pubescent,
acuminate: ray flowers: ligulate corollas rarely more than three to
each head, the basal tubular part 1™™ long, ovate, papillose; the
expanded wing broadly obovate, 5™™ long, nearly that in width, yellow,
the apex bearing a narrow middle tooth: achenes without pappus, 2™™
long, obovate, brown or possibly black when mature, puberulent,
triquetrous, tightly enclosed by the involucral bracts; style slenderly
2-cleft, stigmatic on the edges for the entire length: disk flowers: the
5 disk flowers separated from the ray series by 5 united bracts: achenes
short and sterile, bearing a paleaceous pappus 1.5™™ long: corolla
tubular, 4™™ long, bearing 5 yellowish teeth: filaments half as long
as anthers, separately inserted upon the corolla; anthers 2™™ long,
with constricted triangular apices: style cleft, the lobes pe
ciliate.

Quite plentiful in dry half sandy soil, Santa Hdshass, Santa Barbara
county, California, August 1902. oa specimen, no. 3839, in Herb. Stanford
University.

Machaeranthera pinosa, n. sp:—Densely tufted perennial, 1.5 ¢™
high: underground stems numerously branched, those above ground
rather rigid, brown, with a short cinereous pubescence, usually divari-
cately 3-branched from a little above the middle, basal portion
marked with ring-like scars caused by the early falling of the radical
leaves: basal leaves dry and brittle, those above the middle of the
stem 3-5°™ long, linear-oblanceolate, densely cinereous pubescent on
both sides, usually conduplicate above the middle, sessile though the
lower half is narrow and petiole-like, minutely dentate, the teeth
terminating in short and sharp-pointed hyaline spinules; upper

leaves bract-like, broadest below the middle: heads turbinate, 1°™

high, solitary or upon bracteate brown glandular pubescent peduncles
4°™ long: involucral bracts imbricate, very unequal, mostly linear,
acutely pointed, the exposed surface glandular, the inner surface
straw-colored or purplish-blue: corolla of ray flowers 22™™ long,
tube minutely pubescent, 12™™ long, gradually expanding into the
rich purple-blue broadly spatulate minutely 3-toothed ligule: pappus

-in one series, 12™™ long, scabrous, reddish-brown, very unequal:

corolla of disk flowers 0.75™™ long: stamens with anthers 3™™ long

Mo. Boat. Garden
1905

5° BOTANICAL GAZETTE [JANUARY

and short filaments: style deeply cleft, the branches straight and with
the entire dorsal surface stigmatic; immature achene 2™™ long, obo-
void, finely pubescent.

Rather rare on the summit of Mt. Pinos, Ventura county, California, July
1g02. Type specimen, no. 4145, in Herb. Stanford University.

This beautiful deep blue or purple-flowered Machaeranthera, like many other
purely alpine — has the habit of apparently forcing its growth along crevices
of rocky summits

Chi yndthanieie corymbosus, n. sp.—Shrubby, o.5-1™ high,
numerously branched, forming well rounded bushes; wood dry, soft,
yellowish, with thin shredded bark : leaves on second year old branches
of varying lengths, much shorter than on younger twigs, usually fas-
cicled, soft pubescent on both sides, dry and becoming recurved;
those on the upper branches mostly of the same length, 1o—15°™
long, ascending, not fascicled, narrowly linear or oblanceolate, with
a short sparse pubescence, the surface more or less glandular, obscurely
punctiform: heads glomerated in rounded corymbs, 2—3°™ across, at
ends of branches: peduncles 6™™ long or longer, they as well as the
subtending bracts sparsely lanate pubescent, usually monocephalous:
involucral bracts persistent, broad and mainly in two imbricate
series; lower ones triangular, short and pubescent; upper ones 5™™
long, nearly glabrous and almost shining, broadly linear-obtuse, with
narrow hyaline margins: heads about 7™™ high, cylindric or sub-
turbinate, 1o-flowered: ray flowers few, 8™™ long including the
ciliate pubescent achene 2™™ long: pappus uniserial, 3™™ long,
finely scabrous, whitish and copious: tubular part of the corolla
shorter than the expanded portion, pubescent; ligule broadly elliptic,
yellow, thin, apex much folding over, bluntly toothed: style cleft:
corolla of fertile disk flowers tubular, 5™™ long, 5-segmented: fila-
ments 3™™ long; anthers 2™™ long, linear, with triangularly con-
stricted apices.

Near Lancaster, Los —_ ety, California, June r902. Type specimen,

part ofthe day this deser-loving plant =

1905] ELMER—NEW WESTERN PLANTS 51

branching above the middle and bearing few-flowered cymes: leaves
clustered at the base, longer ones 6°™ long, covered with a silky-
velvety pubescence, usually with a petiole 2°™ long; stipules 15™™
long, adnate and conduplicate, the interior surface quite smooth,
terminating in a laciniate pubescent segment 5™™ long; those of the
smaller cauline leaves broader, larger lobed, and nearly clasping the
stem; leaflets of the lower leaves 8-12 pairs, densely silky pubescent on
both sides, 5™™ long, fully that in width, sessile, merely 3—5-pointed,
the lower one or two pairs somewhat distant, those above the middle
more or less crowded and diminishing in size toward the apex: inflor-
escence loosely cymose: flowers short or longer pedunculate, sub-
tended by leaf-like bracts: hypanthium cup-shaped, 5™™ broad
and 5™™ high including the teeth nearly 2™™ long; calyx bracts in
two series of 5 segments each, the outer shorter and narrower, the
inner 3™™ long and acute: stamens 10; filaments acuminate, nearly
glabrous, 2”™ long and slightly unequal in length, petaloid in color,
flattened especially toward the base; those opposite the petals much
shorter than the others which are exceeded by the longer calyx seg-
ments; anthers 1™™ long, introrse and basifixed: petals alternating
with the outer series of stamens, cuneate or oblong to obovate, 3-4™™
long, 2™™ wide, barely exceeding the longer of the calyx segments;
between the base of filaments and receptacle proper a narrow dark-
brown smooth circular band: receptacle ciliate, ovoid to conical:
pistils smooth, numerous, sessile; style 3™™ long, erect, straight,
constricted or articulated to the ovary; achenes 1™™ long, brown,
semi-recurved.

Dry gravelly flat near Griffin’s postoffice, Ventura county, California, July
1902. Type specimen, no. 3971, in Herb. Stanford University. Named in honor
of P. A. RYDBERG.

Castilleia gleasoni, n. sp.—Perennial, 2-4°™ high, from a hollow
thick dwarfed woody base: stems mainly from the base, branched at
the middle, obscurely ridged or striate, clothed with a soft pilose
pubescence which conceals the more or less dry glandular surface;
branches usually clustered from the distal ends of the abruptly
- terminated stems, striate, similarly pubescent and glandular, bearing
erect or sub-flexuose spicate racemes 1°™ long: leaves many but not
crowded, all cauline and equally distributed; larger ones 4.5°™ long,
12°™ wide, 3-5-nerved, the pubescence on both sides concealing the

52 BOTANICAL GAZETTE [JANUARY

numerous fine sessile glands, the upper one-third divided into 3 lobes
with the middle one usually broader and longer; smaller leaves linear
to lanceolate and entire; axillary leaves few, short and inconspicuous,
not fascicled: average bract 2°™ long, strongly 3-nerved, divided to
below the middle into 3 strap-like purple-red entire segments; these
are obtuse or rounded, glandular and pubescent especially on the
outer surface above the middle: calyx tubular, 15™™ long, 4-nerved,
glandular pubescent, the bilabiate part 5™™ long, each bearing acute
teeth 2™™ long; galea 2°™ long, slightly curved, slit half way down,
apex narrowly truncate or faintly 3-toothed, exterior apical section
glandular, middle part pubescent, glabrous at base; lower lip repre-
sented merely by 3 broad and short lobe vestiges: stamens 4, the two
inner ones slightly shorter; filaments separate, thread-like, inserted
upon the corolla tube beneath the throat; anthers approximate,
included by the galea, the two cells unequal in length, united above
the middle, distinct below: style slender, gradually enlarging toward
the apex, exceeding the galea by 2™™, bearing a 2-lobed flattened
stigma: capsule ascending,.15™™ long, ovoid to lanceoloid, slightly
dorsiventrally compressed, acute or acuminate.

Cliffs and in rock crevices on the summit of Mt. Gleason, near Acton, Los -

Angeles county, California, June 1902. Type specimen, no. 3659, in Herb.
Stanford University.

This “paint-brush” or “‘red-hot-poker,” as I have heard it called, is neither
the early flowering C. foliolosa H. & A. of the seacoast nor.the glandular southern
C. Martini Abrams.

Eriogonum baratum, n. sp.—Annual, 3~5°™ high, erect, mostly
single: stems glaucescent, purptish at base, straight and erect, one-
ae the height of the plant, terete, of the same thickness throughout;
‘anches 1-5 at a whorl, smooth and also glaucescent, ascending,

mately widely branching, subtended by 3 acuminate smooth

oe _ coriaceous bracts 1-2™™ long: leaves all radical, longer ones

bs long, includjng the woolly petiole 2°™ long which is perceptibly

. ee expanded at base; blade proper Se long, ovate to broadly elliptic

1905] ELMER—NEW WESTERN PLANTS 53

from one-fourth to one-third the entire length of the involucre: pistil-
late flowers little protruding, the stalks apparently articulated, 2.5™™
long, commonly 10, becoming deflexed, intermixed with ciliate

bristles 2™™ long: corolla 2™™ long, white or pinkish, petals 6, in

two whorls; outer broad elliptic or more or less rectangular in shape,
with bases clawed; inner shorter, narrower and obovate: staminate
flowers fewer, sessile or sub-sessile, intermixed or more or less ih the
central region, also subtended by ciliate bristles, 1™™ long, petals
smaller though similar in shape and equal in number: the 10 stamens
included, upon slender filaments, bearing small yellow anthers: ovary
smooth, elongated, 2™™ long, surrounded by numerous sterile fila-
ments; styles 3, spreading, terminated by small capitate stigmas;
achene sub-terete and spindle-shaped, purplish, smooth.

These hills contain borax mines, and since this plant is strictly confined to
this region it would seem that the salt is a necessary constituent for it.

On very hot barren exposed round-topped hills between Griffin’s postoffice
and Mt. Pinos, Ventura county, California, July 1902. Type specimen, no. 3593,
in Herb. Stanford University.

Lupinus glareosus, n. sp.—An almost acaulescent annual, 3-sam
high, somewhat succulent though readily curing: stem very short,
rather thick and hollow, terminated by a spicate inflorescence;
branches with a soft short brownish pubescence, chiefly from the
stem, the outer ones curved upward from the middle: leaves clustered
at the point of branching, erect or ascending, upon finely pubescent
petioles 1-5°™ long, with broadened bases subtended by laciniate
adnate stipules; leaflets 10, sessile, only sparsely pubescent on both
sides, soft and fleshy, obovate, on an average 15™™ long and 5™™
wide, the older ones with a very peculiar dead scarious upper margin:
spikes cylindrical, 3°™ in diameter, 10-15°™ long, erect and exceed-
ing the leaves, upon pedunculate scapes: flowers in whorls of 5, upon
short pubescent pedicels, subtended by acuminate similarly pubescent
bracts which slightly exceed it: calyx cup-shaped at base, bilabiate,
pubescent; lower lip rather broad, 7™™ long, obtuse, 3-nerved;
upper one thinner, 3™™ long, notched; banner broadly elliptic, 12™™
long, finely nerved and entirely glabrous; lateral wings equaling the
length of the banner, nearly 5™™ in width throughout, basal margins
ciliate and abruptly terminating into a short narrow claw; keel at °
least 12™™ long, falcate, with a constricted ring-like portion near

54 BOTANICAL GAZETTE [JANUARY

the base and with the margin of the aperture densely ciliate: stamens
of two kinds; the 5 longer ones bearing small anthers, the shorter
ones with linear anthers: style falcate, the stigmatic portion barely
exceeding the keel; ovary covered with long rubiginous hairs; pod
equally tapering to both ends, 1°™ long, 2-seeded, silky pubescent,
terminated by a persistent much recurved style; seeds subsessile,
smooth, compressed.

Griffin’s postoffice, Ventura county, California, July, 1902. Type specimen,
no. 3588, in Herb. Stanford University.

It is wholly confined to dry gravelly soil along water courses, hence its name.

Astragalus gaviotus, n. sp.—An erect densely tufted perennial
3-57™ high: stems many from the woody base, branching above the
middle, cinereous pubescent: lower leaves falling early, becoming
recurved, 10°™ long, subtended by distinct acute triangular stipules
5™™ in length; leaflets extending nearly to the base, 7-15 remote and
scattered pairs, 15-25™™ long, 5™™ wide, the rachis usually termi-
nated by a solitary leaflet; lower ones strigulose and short-petioled,
the larger and older ones broadly obtuse or truncate, linear to nar-
rowly elliptic: racemose inflorescence 2-3°™ long, terminating the
rigid striate and strigulose peduncles 1ro‘™ long, barely exceeding
the leaves: pedicels 5™™ long, ascending, strigose, subtended by
shorter long acuminate strigulose bracts: calyx inflated, cup-shaped,
3™™ long, apex apiculate but often more or less toothed, barely
pubescent with a mixture of black and whitish hairs: corolla, stamens,
and pistils not seen: pod persistent, compressed, spreading or becom-
ing pendent upon pubescent usually recurved pedicels 8™™ long,
dull or yellowish-white, 6-10™™ wide, 20-28™™ long, gradually

a : and acuminately tapering to both ends though more often widest

_ above the middle, apex terminated by a short blunt cusp, the sides
et cranes but finely reticulated and under a lens only sparsely hairy,
straighter along the dorsal edge ; seeds commonly 6, saan! stipitate,
on ca smooth, brown, semi-reniform, strongly reticula
ne Sencty aoil ‘not distant from the beach at Gaviota, Santa Barbara county,
California, May 1902. Type specimen, no. 3759, in Herb. Stanford University.
a — A. jilipes Torr, which Sener a: ae

sus, var—A —— tufted
pecies, with man ; from

a Oe a Oe a oo 5 aa

1905] ELMER—NEW WESTERN PLANTS 55

a perennial root; branches as well as leaves subtended by short, con-
crete, and scarious bracts or stipules: leaves ascending, usually 4°™
long, the lower one-fourth petiolate; leaflets glabrous or sparsely
strigulose, 6-10™ long, 7-9 pairs though the lower ones are much
scattering, the terminal either in pairs or solitary by abortion, sub-
sessile, mostly conduplicate in the dried specimens, linear to lanceolate
to oblanceolate: racemes terminal, 2°" long, upon peduncles 4°™
long: calyx subinflated, 4™™ long, terminated by 5 short but nar-
rowly sharp pointed teeth, short-pediceled, pubescence of black and
whitish hairs: pod 3°™ long, obovoid inflated, narrowed toward the
base but not stipitate, reflexed, glabrous, reticulations prominent,
beautifully mottled with a purple or reddish-violet, falling rather early.

Summit of Mt. Pinos, Ventura county, California, July 1902. Type speci-
men, no. 4005, in Herb. Stanford University.

This variety is chiefly distinguished from the typical form by its scarious
stipules, more linear leaflets, and in not being stringulose.

HERBARIUM OF STANFORD UNIVERSITY,

September 1903.

i ee cus. Ann. Botany 1'7: 167-236. pls. 12-13. 1003.

THE MORPHOLOGY OF MONASCUS PURPUREUS.*
EDGAR W. OLIVE.

BARKER’S account of the morphology and development of Monas-
cus in the early part of 1903," as will be remembered, was shortly
afterward questioned in a contradictory paper by IKENO.? IKENO
gave, in fact, a diametrically opposite interpretation of certain of the
more important phenomena connected with the spore formation in
this fungus, and even went so far as to assert positively that BARKER’S
**Samsu” fungus did not belong to the genus Monascus. If BARKER
is right, then Monascus shows close resemblances to other Ascomy-
cetes; if, on the other hand, IkENo’s views are correct, the fungus
is an aberrant form.

The main points of difference between these opposed views may~—~

be concisely stated as follows. BARKER maintains that sexual repro-
duction results in the formation of ascogenous hyphae. The ascogo-
nium consists, according to him, of an elongated penultimate cell;
while the original terminal cell, which is pushed over to one side by
the growth of the enlarging ascogonium, constitutes an antheridial
branch. Fusion takes place between the two, the elongated tip of
the female cell functioning in this act as a kind of trichogyne. After
fusion, this tip portion is cut off by a septum and its protoplasmic
contents, as well as that of the antheridial branch, finally become
disorganized and eventually disappear. It is the cut off penultimate
cell-of the ascogonial branch, called by BARKER the “‘central cell,”
_ which still further develops and produces the ascogenous hyphae.
: This central cell becomes enormously swollen and is invested at
_ maturity almost completely by a mass of closely clasping sterile

ics: 3 atria its enlargement and development it affords protec-

| tion and nutriment to the growing ascogenous hyphae. The latter

| This work was done under a grant from the ——— Institution.

t BA RKER. aay B. T. P., od gt One eee dq f th ascocarp in Monas-

2 ee, Ss; Ueber die Sporenbi und systematische Ene von Monascus
gs perpaens Wen ie Deutsch. Bot. Gesell 21:259-269. pl. 13. 1903.

[JANUARY

Pes

1905] OLIVE—MORPHOLOGY OF MONASCUS 57

arise, according to the evidence obtained by this author, as an out-
growth of the swelling central cell, which grows over the surface of
the latter, closely applied to it, until it finally pushes in, or invaginates,
the enlarged central cell. In this invaginated cavity, or little ‘‘nest,”
thus produced, the ascogenous hyphae are formed, whether as bud-
dings of the end of the in-pushing hypha, or simply as segments of
it, the author does not make clear. At any rate, growth continues
until the closely entwined hyphae occupy almost the whole of the
interior of the enveloping perithecium. ‘‘Asci are eventually pro-
duced from the ascogenous hyphae, and in each of them eight asco-
spores are produced. When the spores are ripe, the asci and asco-
genous branches degenerate, the surrounding central cell having by
this time lost its contents, remaining as a brown cuticularized enclosing
wall. The spores are thus liberated into the cavity enclosed by this
wall, and the ripe perithecium appears to be nothing more than a
brown cuticularized sporangium-like structure.” (L. ¢., p. 204.)
IKENO apparently agrees with BARKER in regard to sexual repro-
duction in Monascus, although his evidence appears to be drawn
from dubious sections, judging from his one drawing of the process.
IKENO’S main point of dispute with BARKER lies, however, in the
method of spore-formation. He asserts that there are no ascogenous
hyphae in the Monascus which he studied, but that, on the other
hand, so-called “‘spore mother-cells” arise by free cell formation.
The cytoplasm of the ascogonium becomes, according to him, aggre-
gated about certain nuclei, resulting in the formation of uninucleate
“‘Cytoplasmaballen,” or ‘‘spore mother-cells.” According to BAR-
KER’s account, these ‘“‘spore mother-cells” are segments of a hypha,
which become rounded up into asci. IkENo further notes that the
““Cytoplasmaballen” become increased in number by fission, and
that they may contain, at the culmination of nuclear division, a greater
number of nuclei than is necessary for spore-formation, and he says
that the extra ones degenerate. Usually six or eight spores are
finally produced, which are, in his opinion, formed by free cells
formation, leaving a small amount of epiplasm between and surround-
ing the spores, thus corresponding to conditions observed in the
ascus. An agreement is noted in the final phenomena connected
| with the development of the fungus, in that in the mature spore fruit

58 BOTANICAL GAZETTE [JANUARY

the walls of the “‘spore mother-cells’” become disorganized, thus
setting free the spores within the perithecial wall. IkENo concludes
with the assertion that WENT’s classification of Monascus among
the Hemiasci is strictly correct, and that BARKER’s fungus is not a
member of the genus Monascus, but that it is a typical Ascomycete.

Several important differences will be at once noted in these two
accounts, the crucial difference being the one which concerns the
origin of certain structures in the swollen ascogonial cell, or “‘central
cell,” as BARKER terms it. IKENO conceives of these structures,

e ‘“Cytoplasmaballen,” as arising by free cell formation, simply
by an aggregation of cytoplasm around certain nuclei; other nuclei
in the ascogonium taking no part whatever in the process. BARKER,
on the other hand, thinks that these deeply staining bodies are seg-
ments of one or more hyphae which have grown in from the outside
of the central cell, and that the ultimate origin of these ascogenous
hyphae is as outgrowths of the central cell itself, which finally turn
and grow again into the cell from which they have sprung. Both
assert that these bodies bear within them later the spores, and in
other respects the two accounts in the main agree.

DANGEARD proposes now to assist us out of our difficulties by
making BARKER’s Monascus a new species. KuyPER,? the latest
writer on the subject, accepts this suggestion, and in a study of both
M. purpureus Went and M. Barkeri Dang., finds confirmation in
the main of IKENO’s conclusions, although he calls IkKENo’s ‘‘spore
mother-cells” true asci. He adds, however, the additional observa-
tion that a nuclear fusion precedes the development of the young asci.
Monascus Barkeri differs, according to his views, from M. pur pureus
in an important respect. In M. Barkeri he has the nuclei fusing ‘in
pairs before the formation of the asci, and in M. purpureus, after the
asci are formed. In M. Barkeri, his observation agrees, therefore, _|
with BARKER’s assumption that nuclear fusion in pairs occurs in the
enlarging ascogonium. Kuvyper regards this fusion as analogous
to the fusion in the young ascus of M. purpureus, but such a fusion
as he describes in the latter should be referred rather to the later —

3 Kuyper, H. P., De peritheciumontwikkeling van Monascus purpureus Went en
Monascus Barkeri Dang. Kon. Akad. van Wetenschappen, Amsterdam 13:46-294-
T pl. 1904.

x

1905] OLIVE—MORPHOLOGY OF MONASCUS 59

fusion which is known to occur in the young asci, whereas his fusions
in M. Barkeri belong to the early sexual fusions, as are seen in Pyro-
nema and other forms.

As BARKER and others have pointed out, the development of Mon-
ascus from the germination of the ascospores or the conidia up to the
formation of the fructifications may be readily followed out in hanging
drop cultures. Observations of living cultures of Monascus in hang-
ing drops show that a fusion appears to take place between the basal
cell of the antheridial branch and the tip portion of the ascogonial
branch, which BARKER likens to a trichogyne. It appears likely
that in some cases, at least, this fusion takes place after the cutting
off of the tip of the ascogonium, although BARKER asserts that the
act must take place before the wall is thrown across. The swelling
up of the central cell and later an appearance which suggests the
pushing in or imvagination of the swollen central cell by some body
which enters generally near the side on which the stalk is seen;
further, the enlargement of this invaginated part until the central
cell seems to be entirely displaced by it; and, finally, the appearance
of vacuole-like bodies within the invaginated cavity and the final
formation of spores, which adhere for a time in little groups, all may
be readily traced in hanging crop cultures.

When one attempts to explain these appearances, however, by
an examination of sections, one meets with many difficulties, as is
evidenced by the remarkable differences obtained by the investigators
above mentioned. I have seen in but few instances, in fact, a hypha-
like outgrowth from the central cell, and, perhaps somewhat less
rarely, the actual pushing in of a hypha-like bocy into the sice of
the central cell, thus forming the “‘nest,” or invaginated cavity, in
which the ascogenous hyphae cevelop. Another theory than that
held by BARKER has suggested itself to the writer, which is borne
out by many observations. Instead of the ascogenous hypha arising
from the central cell, it may have its origin in the “‘trichogyne-like”’
end of the ascogonial branch. The following observations point
to this conclusion. First, the rarity of protuberances from the
central cell which are long enough to furnish absolute conviction
that the outgrowth is a hyphal branch. They may as well be, as
BARKER himself intimates, a bulging out of the swelling central

60 BOTANICAL GAZETTE [TANUARY

cell at a point where the investment formed by the covering hyphae
is incomplete. Secondly, the fact that, in very thick sections, one
sometimes finds that the invaginated cavity of the central cell is
connected with the outside by a comparatively large, broad opening,
where we might naturally expect a small, narrow one, since the
ascogenous hyphae are relatively small. The large opening suggests
. the pushing in of a larger body, possibly the fertilized ‘trichogyne”’
cell above mentioned. Lastly, the frequency of occurrence in early
stages of a comparatively large, deeply staining cell, lying to one
side of the swollen central cell and not yet pressed into it, indicates
that this deeply staining body is not a hypha with the origin BARKER
- attributes to it, but is instead the cell above mentioned. Should
this prove true, then the fertilized ascogonial cell is in reality the end
cell, while the enormously swollen penultimate cell performs a sort
of “nurse-cell” function, ultimately becoming entirely displaced,
its contents digested and absorbed, by the ascogenous hyphae develop-
ing within it.

That the structures growing within the cavity in the central cell
are segments of ascogenous hyphae, I have no doubt; therein I agree
perfectly with BARKER. For one may see in sections of almost
every immature fructification young segments which are elongated
and twisted and hypha-like, and which sometimes show evidences
of dividing by fission, as is pointed out by IKENOo. Later, these
segments become rounded off and vacuoles appear in them; still
further development results in the formation of their eight spores.

The Monascus species which I have used was first sent from Java
some years ago by D. G. FarrcHILp to ERwin F. SmitH and was
regarded by the latter as M. purpureus Went. Professor HARPER,
who in turn received some material from Dr. SmirH, has turned the
fungus over to me for examination. So far as I am able to judge
from gross measurements, and from a careful comparison of figures,
this form agrees with BARKER’s description of his Monascus, and as
well with the accounts of WENT, UyEpDA, and others, so that my
opinion is that IkENo and Barker worked with similar forms. I
find therefore in the main a confirmation of the conclusions of BARKER, — ;
as well as ory rei gauges of the oe of IKENO:

MAbpIson,

BRIEPFER ARTICLES:

FERTILIZATION IN THE SAPROLEGNIALES.

TRow! has reaffirmed his conviction that a sexual act is present in
the water molds. His paper gives in detail the results of an investigation
of Achlya DeBaryana and A. polyandra in comparison with my own study
of oogenesis in Saprolegnia.2 Much of his paper is devoted to a defense
of his earlier work and to cain criticisms of the skepticism of myself
and others.

A priori there is of course no reason why the Saprolegniales should not
show examples of sexuality; the discussion has only been on the evidence
that Trow presented in his paper of 1899.3 This evidence consisted in

_| the varying number of nuclei present in the oosphere and oospore, expressed

by Trow in the ratio 1:2:1, and certain figures of male gamete nuclei
within the egg which did not seem to me clear enough to be convincing
and appeared capablé&of other interpretations. I found that binucleate
eggs, young and old, were “‘quite common” in an apandrous form of
Saprolegnia mixta, and that by the methods of oogenesis bi- and trinucleate
eggs might be expected in any of the Saprolegniales. Indeed all investi-
gators of this group, HUMPHREY, Hartoc, and Trow himself, have known
and described binucleate eggs. TRow in his last paper (p. 547) shite

“in

| to the binucleate eggs of my form as of ‘occasional occurrence” an

all probability of a pathological character.’ I stated expressly that they
were “quite common ” and I cannot see the slightest reason for suspecting
them to be pathological. The cultures appeared perfectly healthy and the
phenomenon was not infrequent as TRow implies. For these reasons I
held that the number of nuclei in the ratio of 1:2:1, ordinarily accepted
as evidence of fertilization when found successively in young and older
stages of eggs and oospores, was insufficient for the establishment of sexu-
ality in the Saprolegniales. To my mind proof is only furnished by clear
and unmistakable instances of fusion between eggs and antheridial fila-
ments and with thé male nuclei actually en route to the female at the point

t Trow, A. H., On fertilization in the Saprolegnieae. Ann. Botany 18:541-560.
pls. 34-36. 1904.

2 Davis, B. M., Bor. Cuan: 35: aye 1903.

3 Trow, A. H., Ann. Botany 13:131. 1899.
1905] ‘ 61

62 BOTANICAL GAZETTE [JANUARY

of fusion and within the egg. Few botanists would have been satisfied
I think with Trow’s evidence on the latter point in his paper of 1899. I
regarded his views as “‘unproven and improbable” in the face of the mass
of observations upon which most botanists have generally agreed that the
Saprolegniales were apogamous. I still consider myself quite justified
in taking this attitude, which I tried to present with due consideration to
Trow as possibly being mistaken in his interpretation of some very difficult
conditions. He seems to have taken my expressions as controversial,
which I tried hard to guard against, and has certainly answered them in
that spirit. One result at least has been an account of fertilization in
Achlya DeBaryana which in detail of description and clearness of figures
is quite a different sort of contribution from that of 1899.

Achlya DeBaryana furnished Trow the most complete series of stages
of fertilization. In his-jfigs. 23-26 the antheridial filaments are united
with the eggs by broad strands of protoplasm, and nuclei lie in some cases
almost exactly at the points of fusion, either as though they had just entered
the egg or were about to do so (fig. 23). The figures (especially 24
and 25) even show that peculiar structure of the protoplasm which indicates
a flow from the antheridial filaments into the eggs, conditions which have
been illustrated many times in accounts of fertilization among the Perono-
sporales. These statements give us what we have a right to expect of any
account of fertilization, and especially when the technical conditions are
so complicated as in the Saprolegniales. They seem to me fundamental
to the present problem, and the establishment of the ratio of nuclei in
oosphere and oospore as 1:2:1 merely a corollary to the main proposition.
If Trow will exhibit his preparation showing these protoplasmic fusions
of antheridial filaments with eggs at scientific meetings in Great Britain
or elsewhere he is not likely to have any difficulty in establishing his con-
tention.

account of a second mitosis in the cogonium and the origin of the aster-
like structures which I discovered in Saprolegnia and considered coeno-
centra. During the second mitosis, according to TRow, two centrosomes —
with radiations appear at the poles of the spindle in anaphase, structures
which were not present in the first mitosis. Some of these increase rapidly —
__ in size and become the centers of the egg origins, each one being accom-
_ panied by the nucleus that lay beside it, after anaphase of the second mitosis.
Relatively few of the nuclei in the oogonium pass through this secon

_ mitosis, the remainder degenerating, and Trow believes that some of the

” —— nuclei of this division with their asters also break down. The

Of greater interest to me than the evidence of fertilization is TROW’S_

1905] BRIEFER ARTICLES 63

surviving asters with their accompanying nuclei become the centers around
which the cleavage of the protoplasm proceeds to form the eggs. TROW
calls the egg-asters ‘‘ovicentra.”” He believes that the number of chromo-
somes shown in the first mitosis is reduced by half to form four in the
second, so that the latter is a reduction division in oogenesis.

TRow’s account of the development of his ovicentra is fundamentally
different from mine. I regarded them as simply representing dynamic
centers of the protoplasm without any relation to mitotic phenomena,
centrosomes, or asters, and as strictly comparable to the coenocentra of
the Peronosporales. ‘Trow considers them centrosomes in an aster (astro-
spheres) appearing at a final reduction division (second mitosis) in the
- oogonium. The situation has become very much complicated. If ovi-
a { centra are the same as coenocentra, why have not the latter been found

. associated with mitotic figures in the Peronosporales, where the structures
-) are large and have been much studied? Is it possible that we have a new
q *\ structure in the Saprolegniales? Tam not willing to give up my view of
oogenesis, although I can at once see the necessity of further investigation
in both the Saprolegniales and Peronosporales. I do not think that
Trow has presented enough evidence or treated the subject sufficiently
with reference to the Peronosporales to be convincing. His theory is
based, so I understand, chiefly upon the sections from one oogonium.

. There are some interesting questions suggested by the group of figures

, numbered 18, which one hesitates to ask without seeing TRow’s prepara- i
i; tions. But why are not the asters present at metaphase, the time when
4. such structures are generally most conspicuous? And again, why are the
asters sometimes so far away from the poles at anaphase (jig. 188)? Is it
_ possible that the ‘‘anaphase” is simply a resting nucleus drawn out towards

a coenocentrum and attracted by it, as is the habit in the Peronosporales ?

These questions are not put in a captious spirit, but by one who is deeply
interested in the problem and knows the difficulty of the investigations.
i! Trow has advanced a most interesting hypothesis, which he must expect
| to be keenly scrutinized.

‘“The sperm nucleus soon after its entry into the oosphere acquires a
distinct centrosome and astrosphere.” This seems to the writer the most
remarkable announcement in TRow’s paper. Neither the origin nor the
fate of this structure was followed. It looks exactly like an ovicentrum.
The male and female nuclei and their accompanying astrospheres remain
quite apart in the egg until the astrospheres disappear. In the meantime
the oospore gradually fills with reserve material and the wall thickens.
While these processes are under way the gamete nuclei fuse. The structure

64 BOTANICAL GAZETTE [JANUARY

a ae

of the egg when it contains two astrospheres with the gamete nuclei is of —
great interest, and this phase of the subject is worthy of extended study.
I have seen and figured eggs in Saprolegnia with two coenocentra, each
accompanied by a nucleus, when it was evident that the two were sister
structures included within the same egg origin.

It is evident that very much more work must be done both in the Sap-
rolegniales and Peronosporales before some of the points suggested by
Trow’s paper will be established. An ovicentrum arising from an aster
associated with a reduction mitosis is a very different sort of conception
from our present idea of the coenocentrum in the Peronosporales. The
same sort of structure is also reported to appear suddenly beside the sperm
nucleus after its entrance into the oogonium. It seems hardly conceivable
that similar structures could behave in such different ways or could hold
relationships to the coenocentra of the Peronosporales. Of course my
own view of the essential agreement of the Saprolegniales and Perono-
sporales in processes of oogenesis around coenocentra must await the
results of future investigations.—B. M. Davis, The University of Chicago.

THE SEXUAL ORGANS AND SPOROPHYTE GENERATION OF
THE RHODOPHYCEAE.

OLTMANNS’s theory that the structure developed from the fertilized
carpogonium in the Rhodophyceae represents a sporophyte generation has

received substantial confirmation in recent cell studies of WOLFE? on ©
Nemalion. This investigation also makes an important contribution to
our knowledge of the sexual organs of the red algae, establishing some *
complications of structure that are likely to prove very general in the 7
group, and quite changes our conception of their morphology. Their ©
possible relations to the sexual organs of the lichens, Laboulbeniaceae, and —
even to the Uredineae is full of interest.

carpogonium during the early development of the tiehonyee, panes
to the bottom of the carpogonium. Although no mitotic figure was obser
it seems clear that the nuclei of the trichogyne and egg are sisters, follow:
ing a division in the terminal cell of the procarp.
These observations on Nemalion confirm my account of the trichog}
* WoLrE, J. J., Cytological studies on Nemalion. Ann. Botany 18:607-
_ pls. 40-41. 1904. ;

\ ‘

}
905] BRIEFER ARTICLES 65

f.

of Batrachospermum,? which I described as a nucleated cell with a chro-
matophore, and whose long life was accounted for by these complexities
of structure. Later investigators, SCHMIDLE3 and OsTERHOUT,‘ failed to
find the trichogyne nucleus, but I am confident that my account of this
structure is substantially correct. It is probable that nucleated trichogynes
will be discovered in other red algae, and indeed they are likely to be
quite general in the group, for the great length of these structures is much
more readily understood as nucleated appendages than as cytoplasmic
extensions from the small carpogonia. We shall have to recast our con-

ception of the female sexual organ of the red algae. Instead of being a

4}. uninucleate cell (oocyst) with a filamentous cytoplasmic receptive out-
s growth, we shall probably have at least a two-nucleate structure with the
trichogyne obviously very much like a second cell in the organ.

4 These complications, puzzling at first thought, may clear up some very
\ difficult problems of morphology. I refer to the multicellular trichogynes
tie of the lichens and Laboulbeniaceae. They will not seem to be so far
-» 8 removed from the female organs of the red algae if trichogyne nuclei prove
) ” to be general, and we may come to regard them as the highest expressions

of this growth tendency on the part of a primitive type of unicellular organ.

BLACKMAN’SS recent investigations in the Uredinales indicate that female
i sexual organs are present in some forms (e. g., Phragmidium), and the
a presence of a sterile cell above the fertile is wonderfully suggestive of a
2 - trichogyne. All the female organs with multicellular trichogynes form a
-\ group quite by themselves, and apparently without relation to other mul-
ticellular sexual organs (gametangia). Since the cells are probably con-
nected with one another by broad strands of cytoplasm, the physiological
conditions may be very close to those of multinucleate sexual organs
(coenogametes).

The sperm (spermatium) of Nemalion is formed singly in the spermat-
ocyst and leaves this structure as a uninucleate naked or thin-wal
mass of protoplasm. But before fertilization the original nucleus of the
sperm divides with a mitotic figure, so that two male nuclei are formed
'- and both enter the trichogyne. SCHMIDLE reported binucleate sperms
, in Batrachospermum, and some of my own figures show the same conditions,
* which, however, I explained as phenomena of fragmentation. It is evi-
dent that the sperm discharged from the spermatocyst (antheridium)

or onstage sdeetes at
PO am, li ae ~~ udhorine. -
i, Oe Nw a “

\)

j 2 Davis, B. M., Ann. Botany 10:49. 1896.
\ 3 SCHMIDLE, Bot. Zeit. 5'7:125. 1890.

! 4 OsTERHOUT, Flora 87:109. 1900.
| 5 BLACKMAN, Ann. Botany 18: 323. 1904.
'

’ en bs oe
oo aad thlly crepe patel
le lad ia

66 BOTANICAL GAZETTE [JANUARY

involves all of the protoplasm of this mother-cell, and the mitosis is a relic
of times when more than one sperm was formed. The present binucleate
sperm is a simple type of multinucleate gamete (coenogamete).

Both male nuclei are discharged into the trichogyne of Nemalion, which
at the time of fertilization contains no organized nucleus. One male
nucleus passes into the carpogonium, in the upper portion of which it meets
the female nucleus and here fusion takes place. Several sperms may
unite with and discharge their contents into the trichogyne, but supernu-
merary male nuclei soon break down. The fertilization of Batracho-
spermum undoubtedly takes place in essentially the same manner as
described by ScHMIDLE and OsTERHOUT, but both of these observers
failed to understand the complications of the trichogyne. On the other
hand, I was misled by these complications and failed to discover the essen-
tial act of fertilization as a nuclear fusion in the carpogonium.

The development of the cluster of fertile filaments (gonimoblasts)
follows the older accounts, but WOLFE contributes a new point of impor-
tance in showing that the carpospores are developed successively at the

ends of the filaments. When one spore has been discharged the cell

behind grows into the old cavity and develops there a new carpospore.
This method of spore formation by successive proliferations is well known

in certain groups of fungi, but among the algae has so far only been described

for the Rhodophyceae.

The cytological evidence that the cystocarp is a.sporophyte generation
rests with a count of the chromosomes in mitotic figures at different peri
of the life history. The number is about eight for the gametophyte, as
shown in vegetative mitoses and during spermatogenesis. Mitotic figures
were easily found at certain stages in the development of the fertile fila-
_ ments (sporophytic), and these always presented an approximate double
number of chromosomes, about sixteen. The numerical reduction of the

;

chromosomes seems to take place just previous to spore-formation, for —

nuclear figures in the terminal cell of the older fertile filaments had quite —

_ the same appearance as those of the gametophyte and showed eight chro- —

Mmosomes. The manner of the reduction was not determined.
WOLFE gives some interesting details of cell and nuclear structure.
The chromatophore of Nemalion has the form of a hollow ellipsoid, the

center being a vacuole and not a pyrenoid. The chromatin of the resting —

nucleus is present in the form of a globular body (nucleolus). During —
prophase the chromatin passes along fibrillae to the nuclear wall. The |
spindle is intranuclear and centrosomes are present at its poles during ;

metaphase.—B. M. Davis, The University of Chicago.

CURRENT LITERAEUSE.
BOOK REVIEWS.
The life history of Pinus.

Dr. FERGuson’s detailed account of this subject, profusely illustrated with
excellent figures, will be useful as a reference work.* Much of the text and many
of the figures have been taken from her previous papers, but the investigations
have been pushed farther in every direction, so that the repetition adds to the
value of the present account. The work deals primarily with Pinus Strobus,
but conclusions are in most cases supported by observations upon several other
species.

winter, but the mother-cell stage is not reached until the following April; in P.
- Strobus the archesporium does not appear until May. Probably there is a quali-
tative reduction of chromatin during the second mitosis in the pollen mother-cell.
The air sacs arise by the separation of the exine from the intine at two definite
points. A partial wall, lying within the intine at the prothallial end of the spore,
is an interesting feature not hitherto described. The body cell (“‘generative
cell’) is not surrounded by a definite wall, and when its nucleus divides the two
sperm. nuclei lie free in a common mass of cytoplasm, never organizing distinct
sperm cells. The two sperm nuclei are unequal in size and the larger one is
always in advance.

The endosperm contains about two thousand free nuclei before walls begin
to be formed. The archegonia appear about two weeks before fertilization. The
independence of the male and female chromatin during fertilization, described
by Miss Fercuson and previous investigators, is here worked out in great detail
No cell walls are laid down at the base of the oosphere until the eight-nucleate
stage of the proembryo has been reached. The divisions which result in the
formation of four tiers of cells in the proembryo are described as taking place in
the rl nuclei, which lie in the cytoplasm of the main body of the egg.

$s FERGusON’s conclusions are based upon an adequate field study and
upon ae examination of an immense number of preparations, so that the danger

eee ee ee dies a

t FERGUSON, MARGARET C., Contributions to the life history of Pinus, with
special reference to sporogenesis, the lpr eee of the gametophytes, and fertili-
zation. Proc. Wash. Acad. Sci. 6:1-202. pls. I-24. 1904.

1905] fi

68 BOTANICAL GAZETTE [JANUARY

of oo an occasional condition as a typical one has been reduced to a
minim’

It is = doe to Miss FERGusON to announce that the manuscript left her hands
Dec. 29, 1902, after which only a few minor corrections in the latter part o the
work were possible.—C. J. CHAMBERLAIN.

MINOR NOTICES.

THE FIRST PART of the second supplement (1896-1900) of the Kew Index,
containing the letters A-L, is ready for distribution. It is printed uniformly
in all respects with the Index Kewensis itself. The advantage of having such a
record of the years 1896-1900 cannot be overstated, and the tremendous amount
of labor it involves on the part of Sir W. T. Tatsetton-DveEr and his colleagues
deserves recognition from botanists. The American Branch of the Clarendon
Press is at 91-93 Fifth Avenue, New York City; and the price of the part is 13s.—
LMC.

Koorpers and VALETON? have published another fascicle of additions to
the anh a flora of Java. The five families included are Fagaceae
(Castan p., Quercus 25 spp.), Lauraceae (71 spp., 10 of which are new), |
Masi taciee (4 spp.), Coniferales (Podocarpus 5 spp., one new), and Casuari-
naceae (2 spp.).—J. M. C.

AOTES FOR STUDENTS.

INSECTS AND FLOWER COLORS.—Before considering some very recent papers

on this sg ik brief reference may be made to the work of PLaTEav, who more
y other investigator, has raised objections to the familiar ‘‘flag’’ or “‘sig-

nal” eae as held by SPRENGEL, MUELLER, and Darwin. Among the more
striking of PLATEAU’s observations, 3 we may note that he observed insect visits |
in normal number to heads of Dahlia, which were variously hidden from vision —

indifferently flowers of Salvia and Zinnia, similar in species but differing in
sae that Pelargonium, however showy the flowers, is but little visited by insects
y, but that visits are readily induced in great numbers, if honey is placed
2 the flowers; that visits to Dahlia flowers cease if the nectar glands are removed,
but begin again if honey is added; that the removal of the outer radiant flowers
of Hydrangea has no effect on insect visits; that even such intelligent insects
as bees commit many errors, visiting buds, wilted flowers, fruits, etc. In general

2KoorpeErs, S. H., and VaLETon, oh Boomsoorten op Java. Bijdrage no. 10. —
Mededeel. ” sLands Plant. no. 68. pp. vi+287. Batavia. 1904. !
3See Bull. Roy. Acad. Belg., 895; 1896, 1897; Mém. Soc. Zool. France, 1898, —
1899, 1900; Ann. Soc. Entomol. Belg. 1901, 1902; Bull. Roy. Acad. Belg.-1902-
Reviews of these articles in Biol. Cent. 16:417; 1'7:599; 18:469; 19:349; 20:4993
21:650; 23:224; 23:311.

1905] CURRENT LITERATURE 69

it appears that PLATEAU’s views have become less positive; he held at first that
color and form have no influence in mapa insects to flowers. At present,
while all color preference is denied, and while odor and the presence ti a
are believed to be the most potent attractive agents, he seems not to deny altogether
the possibility of color attraction. PLATEAU’s work has been accepted by but
few students of flower ecology, and has been violently attacked especially by
Krenitz-GEeRLorr.4 P£REz,5 while not calling PLATEAU’S results in question,
+ thinks that his conclusions are too one-sided. He holds that odor attracts at a
} distance, but that color serves to give directional precision close at hand. Forr®
} thinks that PLareau and other authors have given insufficient attention to the
memory sense of insects.
The immediate occasion for this review is the work of ANDREAE? and Griray.®
_ ANDREAE begins his admirable treatise with the rather teleological statement
‘ that it is inconceivable that there should be so many showy flowers in the world,
~ kunless they are of some value to the plant, and that we cannot conceive of any
-}-use other than the attraction of insects! As the result of a great many ingenious
g experiments, he divides insects into two genera! groups: those with weak visual
sense, short flight, and high odor sense; and those with opposite characters.

in particular, includes the Apidae and the higher Diptera. Thus, while many
insects are attracted chiefly by odor, the bees and the higher flies are attracted
chiefly by color. Bees tried to get inside of inverted glass vessels with flowers
inside, while they paid little attention to open vessels with honey or perfume.
A very gaudy artificial poppy had more visits than natural and less showy flowers,
although it was observed that the insects soon learned to visit non-showy flowers
containing honey and to avoid showy flowers that had no honey. Since the
perception of odor is highly modified by direction of wind, moisture, and the
| presence of other odors, ANDREAE thinks that color is a much more reliable and
_ f effective attractive agent than odor.

The memory sense, touched upon by Foret and ANDREAE, has been the
special! thesis for GILTAy’s work, and his work seems to harmonize the discord-
ant results of PLATEAU and ANDREAE. A new line of experimentation is sug-
gested by his work on Papaver Rhoeas, a species whose flowers are self-sterile;
from one group the petals were removed and the flowers left free for insect visita-
tion, another group was left in the normal condition and free for insect visitation,

hie

.

EE necator cag plea She Ri

4See Bot. Zeit. 54:123; 55:84; 55:108; 56:138; Biol. Cent. 18:417; 23:557.
5 See Act. Linn. Soc. Bordeaux, 1896; Mém. Soc. Sci. Phys. Nat. Bordeaux, 1903.
6 See Zool. Jahresb. t1gor.

7 ANDREAE, E., Inwiefern werden Insekten durch Farbe und Duft der Blumen
“@ angezogen? Beihefte Bot. Cent. 15:427-470. 1903.

a 8 Gittay, E.: Ueber die Bedeutung der Krone bei be Bliiten und iiber el
z ee ecto Sereno der Insekten. I. Jahrb. Wiss. Bot. 40: 368-402.
~ 1904.

70 _ BOTANICAL GAZETTE [JANUARY

while a third group had the corollas removed and was hand-pollinated. Instead
of noting insect visits, the seeds from an equai number of flowers of the three
groups were weighed. The average for each fruit was 117™ in normal insect-
visited flowers, 115™S in the hand-pollinated flowers deprived of corollas, and
but 50™8 in the flowers deprived of corollas, which were left free for insect visita-
tion. In other words a poppy with showy flowers produces more than twice as
much seed as a poppy without showy flowers, when left free to insect visits.
Thus GiLTay was at first led to doubt PLATEAv’s conclusions. However, fur- —
ther experiments produced a modification of his views, since it was noted that
insects, and particularly bees, pay less and less attention each day to showy
flower groups where no nectar is present, while they pay more and more attention
each day to inconspicuous flower groups, where nectar is present. It would
seem, then, that PLaTrau failed to see the earlier visits of his insects, or at least (
failed to discriminate in his results between the earlier and later visits, while f-
ANDREAE, in general, seems to have concluded his experiments too soon.—H. C. §
CowLEs.

Scumipt ° defines the mangrove association as that made up of tree-like ever-

mn plants which live near the strand of sea and estuary where the ground i
constantly or at regular intervals flooded with salt or brackish water. Owing
to its peculiar development and mode of life this is the most characteristic and
well defined association known. The English “tidal forest” or the Holland
“‘vlordborsch”’ is suggested as preferable to mangrove association, the former
expressing exactly what is characteristic of the formation. The Barringtonia
association (Schimper) borders closely upon the mangrove. As a rule, however,
they are quite distinct, the Barringtonia being found on a higher level where the
tidal water does not re

Light plays an important part in determining the position of the leaves and
branches? By placing ten leaves horizontal and ten others upright in the normal |}
light of the mangrove forest it was found that disorganization of the chlorophyll
took place in one-half to two minutes in the former, while after thirty minutes
only a partial disorganization occurred in the latter. By assuming the vertical
position the buds and leaves expose the least surface to the direct rays of the
sun. ‘The relative intensity of the light according to the exposure of photographic
paper is for the open sea one-fourth to one-half second, in the mangrove two
seconds, and in the inland jungle ninety seconds. Hence it is seen that the light
. very —— in ne mene swlpeeeit In the inland jungle the undergrowth :
t light, but so as to get the most, 7. e.,

ina Saateoneal position. :
The major part of the article is devoted to taxonomic descriptions, es {
being laid upon the arrangement and character of the buds, branches, and leaves.

* SCHMIDT, Jous., Bidrag til Kundskab om Skuddene hos den gamle Verdens —
(Contribution to the knowledge of the shoots of mangroves in the

old oaks Rot. Tidsskrift 26: 1-113. Bs. 46. 1904.

fe

1905] CURRENT LITERATURE 71

No general uniformity exists in regard to the branching, although two general

may be separated: (1) the cymose, (2) the racemose. The leaves show
considerable similarity in outline. Two type groups are made, the narrow leaved
and the broad leaved type. The former type occurs only on species whose leaf-
bearing branches are short and upright, while the broad leaves occur only on
those whose branches are long

In regard to protection of aids there is a considerable uniformity. Rhizo-
phora, Bruguiera, Ceriops, Kandelia, and Scyphiphora are protected by means
of axillary leaves which surround the developing parts; Aegiceras and Xylocarpus
by means of true bud scales; Lumnitzera by red fleshy bud scales; Avicennia
and Sonneratia by means of depressions in the leaf stalks. In comparison with
northern trees, therefore, it will be seen that the majority of mangroves have
buds similar to summer buds (Platanus), while Aegiceras and Xylocarpus cor-
respond in this respect to the winter buds of northern latitudes.

The author criticises A ’s work on the anatomy of mangrove leaves,?°
in that the latter author has not studied the plants in nature and his theory that
salt-secreting hydathodes are general among mangroves is not proven in nature.

at salt-secreting glands do occur is definitely proven, but only in the one
genus Aegiceras.—G. H. JENSEN.

THE FINAL paper of BLracKMANn’s studies on the fertilization, alternation of
generations,-and general cytology of the Uredineae has appeared,"! presenting the
subject more fully than the preliminary account which was reviewed in our
April number.'?, We shall note at this time simply some additional details in
this very clear and interesting paper.

The structure of the nuclei and processes of mitosis were studied, but it is
evident that these subjects are very difficult. The chromatin remains for the
most part in masses, so that chromosomes cannot be counted; but of course while
there is but one mass throughout the gametophyte history from the teleutospore
to the aecidium, there are two in the paired nuclei of the sporophyte. The
nucleolus, always present, is extruded with each mitosis and formed anew in the
daughter nuclei. A rudimentary spindle can sometimes be observed, with
bodies at the poles which BLACKMAN calls centrosomes and which he believes
arise by division, but his evidence seems insufficient on the latter point. The
“conjugate” method of division of the paired nuclei only differs from that of
single nuclei in the fact that two divide simultaneously side by side. The daughter
nuclei remain entirely apart, so that the continuity of the chromatic material is not
broken until the fusion in the teleutospore.

LACKMAN points out the exact agreement of certain conditions in the Basidio-
mycetes with those in the Uredinales. Thus the mycelium preceding the basidium

10 Bibliotheca Botanica, heft 56.
11 BLACKMAN, V. H., On the fertilization, alternation of generations, and general
cytology of the Uredineae. Ann. Botany 18: 323-373. pls. 21-24. 1904
12 Bot. GAz. 37:320. 1904.

-

re BOTANICAL GAZETTE [JANUARY

contaihs paired nuclei, which by fusion form a single nucleus in the mature basid-
ium; the basidiospores are of course uninucleate, as is the mycelium that arises
directly from them. Somewhere in the history of the mycelium the single nuclei
must become paired. This will correspond exactly to the period of the aecidium
in the life history of a rust, and must represent the time when an ancestral gamet-
ophyte developed sexual organs, now suppressed, and passed over to a sporo-
phyte generation. The paired nuclei probably arise, as in the rust, by the union
of neighboring cells in the mycelium, a process of apogamy, at the period when
the sexuality is due.

It is important to note that the simpler types of life histories in the Uredinales
where an aecidium is omitted must be regarded as derived from the ew and opsis
forms by the entire suppression of cells which represent female sexual organs,
and such life histories are in essential agreement with those of the ae
which stand at the extreme in the process of reduction —B. M. Davi

Jutes LAuRENT’S has given a detailed account of his experiments on the car-
bon nutrition of green plants. While the old humus theory was long ago aban-
doned, largely by reason of the work of Lresic, the tendency in recent years has
been to hold to the utilization of organic foods to some extent, at least, by normal
autotrophic plants. Boum, E. Laurent, and Acton have conducted rather
conclusive experiments along this line, and a moderate amount of heterotrophic
nutrition on the part of autotrophic plants is accepted by PFEFFER and Jost.
The most conclusive experiments upon which this view may be based are those
here considered, whose publication in full has been long awaited, since a
preliminary notice was published in the Comptes Rendus for 1898. The chief
reason for reposing confidence in Laurent’s work is because he has worked
with aseptic media, according to the most approved modern technique. Experi-
ments with maize roots in contact with glucose solutions, in addition to DETMER’S
solution, resulted in an increase of dry weight in the dark, whereas control cul-
tures in DeTMER’s solution only failed to show such increase in weight. Maz&é
has secured a like result with the vetch. Cultures in an atmosphere deprived
of CO,, but exposed to light showed an increase of weight due to an intake o
glucose by the roots. Various plant species, previously deprived of starch, whose
roots were placed in 1 to 5 per cent. glucose solutions, the plants being placed under
a bell-jar, over a potash solution and exposed to sunlight, showed the formation
of papas from glucose that had been taken in by the roots. Similar results were

tained in darkness, perhaps because less gl absorbed. Autophytes
are found to differ from ‘eaptaphytes not only quantitatively, as appears from the
above, but also more fundamentally. Careful experimentation failed to disclose
any power on the part of the roots of green plants to digest starch, a power pos-
sessed by holosaprophytes. A chapter is devoted to the consideration of the

; io i eta mies J.: Recherches sur la nutrition carbonée des plantes vertes & l’aide

188-203; 231-242 1904.

de matitres organiques. Rev. Gén. Bot. 16:14-48; 66-80; 96-128; 155-1665 2

nC eee

1905] CURRENT LITERATURE 73

influence of organic substances on growth and form. In general, stem growth

is checked in concentrated solutions. While this is doubtless partly due to the
increased osmotic pressure, it is evident that chemical factors are also active,
: since glucose and glycerin solutions of equal concentration give different results.
4 A final chapter deals with the application of his results to agriculture —HENRY
4 C. Cowles.

KrrKwoop'’ has published the results of an extended investigation of Cucur-
bitaceae, chiefly relating to the development of the ovary and the embryo sac
} structures. The genera included are Fevillea, Melothria, Apodanthera, Bryon-
+ opsis, Trichosanthes, Momordica, Luffa, Cucumis, Lagenaria, Benincasa,
1 Citrullus, Cucurbita, Sicyos, Micrampelis, Coccinia, and Cyclanthera, a remark-
} able list for one investigator in these days to present in a single paper; but the
’ 4 work has been under way since 1898. ‘The mass of details will always be consulted _
’ for comparison with similar details in other families, and it is important to have so
_ compact and clear a record. Certain general results, however, should be referred
‘to even in a brief review. As is well known, the position of the family has always

a ‘been a problem because of its contradictory characters, but it is generally placed
', now among the higher Sympetalae. It is interesting to note that Kirkwood has
‘| discovered that the sixteen genera studied unanimously contradict certain charac-
F) ters of Sympetalae that have been thought to be fundamental. The ovule of this
if great group is remarkably constant in its single and very prominent integument,
4 its very much reduced nucellus, and its elimination of tapetal tissue (the hypo-
i dermal archesporial cell passing over directly into the mother-cell). In all the
_ # Cucurbitaceae studied, however, Kirkwood finds the ovules with two integuments,
| a well-developed nucellus, and tapetal tissue by no means eliminated, but often
4 very extensive. It seems to the reviewer that just as the single character of poly-
4 g Petaly arbitrarily applied, has detained the Umbelliferae among the Archi-
+chla

Be

% study of the peculiar pollen tubes reported for the family, or of fertilization and the

7) development of the embryo. e endosperm is noteworthy for its extensive
growth and nutritive activity, to which the author has given somewhat detailed
attention. ‘The general conclusions from the development of the ovary, especially
the growth of the variable “placenta” and the origin of the ovules, are used in an
; interesting way to relate the genera in phylogenetic series.—J. M. C.

f

he

FittINGc’s's preliminary report promises a paper which will be of special
» interest because he has attempted a solution of several of the most fundamental
problems of geotropism. The behavior of orthotropic organs upon a klinostat
14 KIRKWOOD, JOSEPH EDWARD, The comparative embryology of the Cucurbi-
| taceae. Bull. N. Y. Bot. Garden. 3: 313-402. pis. 58-O9. 1904.
: 15 FITTING, H., Geotropische Untersuchungen. (Vor. Mit.) Ber. Deutsch. Bot
- Gesells. 22: 361-370. 1904.

74 BOTANICAL GAZETTE [JANUARY

so arranged that the rotating plant describes a cone whose axis is inclined to the
horizontal leads him to announce the following statements. The optimum
position for geo-perception is horizontal and equal responses are induced by
equal deviations above and below the horizontal. (In a recent paper NEw-
cCoMBE”® finds that roots and stems do not receive equal stimuli at equal angles
above and below the horizontal.) The minimum perception time is extremely
brief, less than two-thirds of a second. A proportionality between reaction time
and perception time does not exist. Orthotropic organs can be geotropically
stimulated even though the rotation be very rapid and uniform. There is a
threshold-difference not only for intensity of stimulation but also for duration of
stimulation, and for position during stimulation (deviation from horizontal).
Intermittent is much less effective than continuous stimulation in the induction
of geotropic curvatures. The response is little affected by the duration and num- |
ber of individual stimulations. The presentation time is never smaller than for
continuous stimulation, and is practically the same as for the latter when the
ratio of stimulus duration to rest duration is as small as 1:5. The duration of
single stimuli is without influence upon reaction time so long as the ratio of rest —
duration to stimulus duration does not exceed 5:1. Nothing was observed which ©
would support HABERLANDT’S statolith i on H. Ponp

SPINDLE FORMATION in the pollen mother-cells of Cassia tomentosa L. has
been studied by Hus,'? who finds that the cytoplasm of the young pollen mother-
cell consists of a network of fibers more or less radially arranged, on and between
which large and small granules are found. The meshes adjacent to the nuclear
wall become smaller and elongated parallel to that wall. A granular zone appears
around the nucleus; and at the same time there appears in the cytoplasm deeply
staining rough fibers frequently arranged in conical groups. A felt-like zone,
partially or sometimes completely surrounding the nucleus, next appears, and the
deeply staining rough fibers of the cytoplasm, now united into cones, establish —
connection with the fibers of the felt-like zone. The linin threads become parallel
with the other fibers and also parallel with the axis of the larger or ascendant cone.
As soon as the rough tents of the nies become RTH ~ nucleolar —_
breaks down, the linin and th ]
into bundles. A multipolar spindle % is formed, two cones of which, opposite to
each other, are more prominent and gradually absorb the smaller cones. The

narily met with in dividing mother-cells of pollen, spore, and embryo-sac, and the
multipolar diarchal spindle Anlage described for vegetative cells.’—W. J. G
LAND.
16 NEWCOMBE, F. €., Limitations of the klinostat as an instrument for scientific
research nce 20:376-379. E904.
‘7 Hus, Henrr T. A., Spindle formation in the pollen mother-cells of Cassia —
tomentosa L. Proc. Calif. Acad. Sci. III. 2: 329-354. pls. 30-32. 1904.

1905] CURRENT LITERATURE 75

Lanc'S has described in detail the single prothallus that he refers provision-
ally to Psilotum; otherwise the gametophyte of Psilotaceae is entirely unknown.
During his expedition to the Malayan peninsula, LANG found a number of plants
of Psilotum growing on the stems of tree ferns, with rhizomes embedded among
the adventitious roots covering the stems. In close association with one of these
plants the single prothallus referred to was found. A preliminary account of its
external form was published in rgor. It is thick cylindrical, about 6™™ long
and 4.5™™ in diameter at the broad upper end. It shows the usual differentiation
of the subterranean gametophytes of Lycopodium into vegetative region below
and sexual region above. The latter appears as a thick overhanging margin in
which numerous antheridia occur. The sexual region does not contain chloro-
phyll, as in the Lycopodium cernuum type, the gametophyte evidently being
entirely saprophytic. The vegetative region exhibits considerable aetucs
differentiation, an endophytic fungus occupying a peripheral zone, and the central
mass of tissue being entirely free from it. Some very interesting details in refer-
ence to the fungus are given, especially the occurrence of some remarkable multi-
nucleate “vesicles” borne upon it. If this gametophyte is that of Psilotum, the
genus is closely related to the homosporous Lycopodiales; if it is not, it is certainly
the gametophyte of some Lycopodium.—J. M. C.

IN A RECENT number of Science MORGAN’? attempts an analysis of the phe-
nomena of organic “‘polarity.”” He opposes the “formative stuff” hypothesis
of Sacus and others, or rather that feature of it which calls for the migration
of specific formative material, and arrays considerable evidence against it.
MoreGan’s conception is that while the material which gives rise to new parts is
totipotent, it is not homogeneous, so that it is different at different levels of the
body, this difference being graded from one end of the organism to the other.
This accounts for the gradation in the facility with which new parts are formed
from one end to the other. Morgan also recognizes an organizing principle,
which he calls the property of “‘formative organization,” ‘“‘which acts on the new
and old parts as a whole and determines the relative arrangement and propor-
tions of the new organs.”

In regard to “‘polarity”’ in plants Morgan maintains that these same principles

old. Assuming that there is a gradation from the distal to the proximal end of
say, a mullen stem, in the size and vigor of the buds, the greater ability to produce
shoots at the distal end is due merely to the fact that these buds are more vigor-
ous and mature. The oe is made that the opposite may be true of the
roots.—W. B. McCati

18 LANG, WILLIAM H., On a prothallus provisionally referred to Psilotum. Ann.
Botany 18:571-577. pl. 37. 1904.

10 Morcan, T. H., An analysis of the phenomena of organic “‘polarity.””. Science
20:712-748. 1904.

76 BOTANICAL GAZETTE [JANUARY

IN A COMMUNICATION to the Royal Society December 1, Scorr?° described a

w type of strobilus belonging to the Sphenophyllales. The close affinity to

Sa Meca taal is shown by the anatomy of the axis, which has the solid triarch
wood characteristic of that genus, and by the fact that the whorled sporophylls are

are tale, forming a system of protective bracts, while the ventral lobes alone
bear the sporangia; in the new cone, dorsal and ventral lobes are alike fertile, and
no sterile bracts are differentiated. Each lobe of the sporophyll divided palmately
into several segments, the sporangiophores, each of which consisted of a slender
pedicel, terminating in a large peltate lamina, on which two pendulous sporangia
were borne. The wall of the sporangium has a rather complex structure, the most
interesting feature in which is the well-defined small-celled stomium, marking the
line of longitudinal dehiscence. The spores, so far as observed, are all of one
kind; they are ellipsoidal in form, with longitudinal crests or ridges; their dimen-
sions are 90-96 in length by 65-710 in width.—J. M. C.

THE comMMON Ithyphallus im pudicus, generally considered to be a saprophyte
only, has been found to be the cause of a destructive root rot of the vine in Hun-
gary. According to the account given by IstvANFF1** the subterranean part of
the stem is entwined by a network of the characteristic cord-like strands of myce-
lium of this fungus. From these, branches are sent into the interior of the stem.
Small roots are totally destroyed by strands which penetrate them lengthwise,
destroying all the tissues and leaving only the thin decaying cortex. In the
older roots the cortex and phloem are totally destroyed, leaving only a mass of
débris. The wood cylinder is last attacked, but this also is finally destroyed,
leaving only scattered remnants of the vessels. Destruction of the immature
fruit-bodies and watering the sai roots with fungicides is recommended as
a means of prevention.

The same paper also contains an account of a mite, Coepophagus cchrnepes,
which lives within and destroys the tissues of the roots of the vine. Not infre-
quently vines are attacked by both pests at the same time —H. HAsSELBRING.

SVEDELIUS”? has published an account of Enalus acoroides, one of the marine
Hydrocharitaceae growing on the coast of Ceylon; the subtitle being “A contri-
bution to the ecology of the hydrophilous plants.” The pollen grains are very
large (about 170 in diameter) and heavier than sea water, owing to the presence

20 Scott, D. H., On the structure and affinities of fossil plants from the Paleozoic
rocks.—V. On a new type of sphenophyllaceous cone (Sphenophyllum jertile) from
the Lower Coal-measures. » oo December 1, 1904.

2 IstvANFFI, Gy DE, Deux nouveaux ravageurs de la vigne en Hongrie (I’Ithy-
phallus impudicus et le Site O echinopus). Ann. Inst. Cent. Ampélologique
Roy. can 3: — pls. I~}. figs. T§. 1904.

VEDELIvS, N., On the life history of Enalus acoreides. Ann. Bot. Gard.
beckons mesg pl. 24. 1904.

1905] CURRENT LITERATURE "7

of numerous starch grains. The wing-like leaves of the ovulate spathe aid in
___ keeping the flower in a horizontal position, and the petals are specially adapted to
catch the floating staminate flowers. The exposure of the ovulate flowers and
the loosening of the staminate flowers occur only during low water. During
- high water the ovulate flowers are vertical; and the heavy pollen grains can scarcely _
| , avoid sinking down upon the erected stigmas. The pollination of Enalus, there-
_. fore, is both at the surface of the water and beneath it. The development of the
§ embryo is of the ordinary Alisma-type, with a very large basal suspensor cell.
At maturity the testa is an easily loosened cap around the embryo, the latter
escaping entire and developing immediately.—J. M. C

= Locx?s has been investigating the rate of growth of the giant bamboo (Den-
| _ drocalamus giganteus) in Ceylon. The stems show a gradually increasing rate
of growth up to 5™; then a nearly constant rate up to 15™; then a slowly dimin-

“4

_ ishing rate to the final height of 30" or more. The average daily rate at a height
+ of r™is ro°™ per day; at 5—15™, 30°"; and at 20-25", 15°". The greatest growth

as

- recorded in 24 hours was 46°™. The daily rate of growth is much increased by
_ rainfall, the effect being greater the greater the height of the stem. The greatest
change observed in rate of growth in two successive days was 50 per cent. of the
@ average rate. Growth is almost always more rapid at night; the average hourly
growth at heights of 3-12™ between 7:00 A.M. and 5:00 P.M. being 6™™, and
Ve A between 5:00 P.M. and 7:00 A.M. 16™™; the maximum rate being reached soon
i at after dark. The curve of sess by day follows very closely that of the percent-
4 a age moisture of the air.—J. M. C
| i THE SECOND section of Davis’s?4 Studies on the plant cell is entitled, “‘Activi-
4 ties of the plant cell.’”’ The activities described are (1) vegetative activities, and
_ (2) cell division. Cell division is treated under the subheads (a) the events of
- nuclear division and (b) the segmentation of the protoplasm. The account of

discussed in detail and considerable attention is paid to the centrosome problem.
3 The view is expressed that all divisions of chromosomes are longitudinal. The
-_ evidence in favor of regarding the chromosome as a permanent organ of the
a cell is regarded as not yet sufficient. In dealing with the segmentation of the
Bi, protoplasm, the author gives a valuable summary of recent work in this field.
—CuHartes J. CHAMBERLAIN.

_. Burns?s concludes that in Proserpinaca palustris L. the water-environment
R . is not the cause of the division of the leaf. He considers that the plant has two

k, R. H., On the growth of giant bamboos, with special reference to the
ard

4.
24 Davis, B. M., Studies on the plant cell. Amer. Naturalist 38:431-469. jigs.
- 1904.
25 Burns, G. P., a ea ie in Proserpinaca palustris L. Ann. Botany 18:
579-605. pl. 38.

~

78 BOTANICAL GAZETTE [JANUARY

types of leaves, an adult and a juvenile type, the latter being the divided form.
Under good vegetative conditions the former is produced, while under poor vege-
tative conditions the juvenile form is developed. A reversion from the adult
to the juvenile form follows a change from a favorable to an unfavorable vegeta-
tive condition—W. B. McCatium.

Lioyp** calls attention to the parallelism between the results of his studies of
the pollen tubes in the Rubiaceae and those of Lonco in the Cucurbitaceae. —
Each of these investigators has found that the pollen tubes vary greatly in behavior,
even in closely related genera or species. They seem to have come independently
to the same conclusion; that is, that endotropism and ectotropism are not distinct
from one another; both regard the behavior of the pollen tube as a physiological
character.—R. B. WyLIr

sain

Wricut?’ has published an extended account of the gross morphology, anat-
omy, and taxonomy of Diospyros as represented in Ceylon. The f
such interest among dicotyledons that so extensive a study of living eae
is welcome. In the taxonomic portion of i paper twenty species are recog- —
nized and described in great detail—J. M

SrupDEnTs of plant-breeding will be interested in the results obtained by Locx?®
in conducting a somewhat extensive series of experiments in Ceylon. In his first j
paper the details of the experiments indicate in the main a confirmation of MEN-
DEL’s law.—J. M. C

76 Lioyp, F. E., The pollen tubein the Cucurbitaceae and in the Rubiaceae.
Torreya 4:86-91. 1904.

27 WRIGHT, HERBERT, The genus Diospyros in Ceylon: its morphology, anatomy,
and taxonomy. Annals Bot. Gard. Peradeniya 2:1-106, 133-210. pls. I-20. 1904.

28 Lock, R« H., Studies in plant-breeding in the tropics. I. Ann. Bot. Gard.
Peradeniya 2: 299-356. 1904.

[1 NEWS.

Proressor H. MarsHaty Warp has been elected president of the Cambridge
' Philosophical Society.
lat Dr. J. F, GARBER (University of Chicago) has been appointed instructor in
. f botany in the Yeatman High School of St. Louis.
' j Mr. LutHER BurBANK, so well known for his experiments in plant-breeding,
_» has been appointed a special lecturer at Stanford University.
Be ¥ Dr. J. C. ArTHur has been appointed by the Botanical Society of America
as its pen to the International Botanical Congress at Vienna.
- Marre C. Stopes (Munich) has been appointed assistant lecturer and
t Rie ae in botany in the University of Manchester, England.
im /ELLESLEY COLLEGE has received $7200 from the ROBERT CHARLES BILLINGS
f font the income of which is to be applied to the needs of the department of
1 botan
bf ae cas R. Maxon, of the U. S. National Herbarium, left Washington
ir: " December 12 for Guatemala, to be gone until May. Primarily he is to be engaged
{ ~ in cotton investigation.
a PROFEssor N. Ono, of the University of Tokyo, has been visiting the botani-
bal laboratories of the United States on his way to the University of Leipzig for
research in plant physiology.
ae _ C. R. Barnes, H. C. Cowtes, and C. L. SHEAR were Pee delegates
4 rom the American Association for the Advancement of Science to the Inter-
“ hational Botanical Congress at Vienna.
gi, Mr. Francis Darwin who, upon his removal to London some months ago,
- i retired from his position as Reader in Botany in Cambridge University, has
| been succeeded by Mr. F. F. BLackman.
dp 4 Dr. R. S. Woopwarp, dean of the faculty of pure science in Columbia Uni-
' |, versity, was elected president of the Carnegie Institution at the meeting of the
trustees held at Washington, December 13.
un ; THE Roya Socrety of London has awarded the Darwin medal of 1904 to
_ Mr. William Bateson, “for his contributions to the theory of organic evolution
; 4 by his researches on variation and heredity.
a A CONTROLLING INTEREST in The Plant World has been purchased by Pro-
aft fessor F. E. Luoyn, of the Teachers College, Columbia University, the journal
a, ‘ passing under his editorial management at the beginning of the new year.
— :

80 BOTANICAL GAZETTE [anvary |

Mr. R. P. Grecory, demonstrator of botany at Cambridge University, has
been awarded the ey armen mir of the Renee for 1904, his thesis being |
‘*The reduction division in pl redity.”

physiology of here
THE Botanical Society of America Pee as Officers for the ensuing year:
president, R. A. Harper; vice-president, E. A. Burt; secretary, D. T. Mac- —
DovcaL; treasurer, ARTHUR HoLuick; councillors, L. M. UNpERWoop, WM. |
TRELEASE. ;
Dr. Burton E. Livincston will sever his connection with the University
of Chicago at the close of the winter quarter, having been appointed to the staff
of the Bureau of Soils of the U. S. Department of Agriculture. He will take up }
his new duties at Washington on April r.
ProFEssOR HENRY S. GRAVES, director of the Yale School of Forestry, has
been commissioned by the Bureau of Forestry to do “inspection work” in the
Philippines. During the winter his courses will be conducted by Professor B. E.
FERNOW, formerly director of the Cornell School of Forestry.
BEGINNING with this year and volume 18, the Beihejte zum Botanisch
Centralblait which has attained an enviable reputation for the prompt publica-
tion of original papers of a high grade, will be issued under the same edito
Drs. Untworm and Kont, by a new publisher, GEorGE Tureme, of Leipzig.
Hereafter the journal will be published in two sections, the first to cover anatomy,
histology, morphology, and physiology; the second being restricted to taxonomy,
phytogeography, and like topics. The price of each section is to be M 16.
ALBERT GaAupry, the eminent paleontologist, has headed a subscription f
the erection of a monument to the memory of BERNARD RENAULT, whose untime
death this fall brought to a close those remarkable researches which have
greatly enriched the science of paleobotany during the past thirty-six years. It
roposed to erect this monument at Autun, France, RENAULT’s native place and
the scene of most of his labors. Subscriptions may be sent to M. BERTE R,
secrétaire de la Société d’ histoire naturelle, 2 rue de |’ Arbaléte, Autun, France.
THE THREE botanical societies, Botanical Society of America, Society for”
Plant Morphology and Physiology, and American Mycological Society, through’
committees of conference, have agreed upon certain general principles, upon |
the basis of which they will fuse into one national society under the name, The |

There are to be two classes of membership, members and associates, the dis- |
tinction being based upon published work. The fees are to be $5 per year- |
Grants for research are to be made from the income. Meetings are to be annual,

or temporary sections in iles of committees. A joint committee has been
formed to prepare a constitution for the united societies, which shall embod
the principles = to, and to complete the reorganization, which is
expected to promote research and good fellowship among American botanists.

_ Of Ail Se

BUFFALO
LITHIA WATER

lo Remedy of Ordinary Merit Could Ever
Have Received Indorsations from
Men Like These.

Samuel O. t. Potter, A. M., M. D., M.R.C.P., London,
Professor ~ the Principles and P: ractice of Medicine and Clinical
Medicine in the yoliees Sait Pi cians and he. cons, San francisco.

o. H 5 H rote d q
: Bright s Disease Dr. Wm. H. Dru nd, = Yr i ossor Medical Jurisprudence,

ee s cea Miakwee t
Cyr Edso ie M., M. D.. calth Comer New
York "City and GH President Board o pe. Pharmacy, New York
Sera City, Examining Physician Corporation Comncd, ae
John V. shoemaker, M.D., LL. D., Professor Materia
Pre oe Medica and Ther ledico- -Chirurgical College, Philadelphia.
§ y Dr. Geo o Ben. Joti nston, Aichmor sii se Ex-President
Southern Surgical —_ Gynecological Association, Ex-President
Medical Soc of Va., and Swe of Smacoines and Abdominal
Surgery, ‘Wedicat College of Va

Dr. A. Gabriel Pouchet, Professor of -heseoh satin and
Stone in the Blad- Materia Medica of the Faculty of Medicine, Pari
3 Renal Calculi, Dr. J. T. LeBlanchard, Prof. Montreal Cink: il” SN., VT.
a Jas. M. Crook, A. M., M. D., Professor Clinical Medicine
. and Clinical Diagnosis, New York Post Graduate Medical School.
Inflammation Louis C. Horn, M. D., Ph. D., o— Diseases of Chil-
of the dren and Dermatology, Baltimore U; niversit
a d Dr. J. Allison Hodges, President eiak Poel ssor Nervous and
a der Mental Diseases, University College of Medicine, Rickonond: V. ‘
Dr. Robert Bartholow, M L. D., Professor Materi
1 G ut Medica and General Therapeutics, ore Medical Colege, Phila.
n Go 5 Dr. I. N. Love, Vew York City, Former Professor Diseases of
a ‘eyes Children, College iy ” Physic cians and. ‘Sarecoks, and in Marion Sims
: College o Medicine, St. Louis.

2 Hunter McGuire, M. D., LL D., £x-President America
Urie | Acid Medical Association, Late President and sith er Clinical Sea
c liti gery, University College of Wikicioe. pide nd, Va

on ns w York, Pedesioe of Surgery,

r. Alexander B. Mott, of
Bellevue Hospital Medical College, ected: Bellevue Hospital.

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| The Botanical Gazette
. wu Montbly Journal Embracing all Departments of Botanical Science

d by JoHN M. COULTER and CHARLES R. BARNES, with the ar of other members of the
botanical staff of the University of Chic

XXXIX, No. 2 Issued February 17, 1905

CONTENTS

b THEORY OF RESPIRATION. Charles R. Barnes - - - - - - . 81

| FORESTS OF THE FLATHEAD VALLEY, MONTANA. CoNTRIBUTIONS FROM THE
HULL BOTANICAL pms ee II (wir MAP AND TWENTY-THREE FIGURE 5}.

Harry N. Whitjord 99
pres IN THE GRAMINEAE. ee TADMRGK sip onan cicaians ) Le saasos insets TWELVE
FIGURES). Theo. Holm 123

E ERN, ATION OF GENERATIONS IN ANIMALS FROM A BOTANICAL STAND-

POINT. ContTRIBUTIONS FROM THE HULL er ANICAL LABORATORY. LXVIII sata

TWO FIGURES). Charles J. Chamberlain - i37
a. a ARTICLES.

EW PRECISION-APPLIANCES FOR USE IN Roca ota ng ae Il ret FOUR R FIGURES).
- Ganong

145
RENT LITERA TURE.
BOOK REVIEWS - -- - - - . - - - Eggs
4 THE TEACHING OF BIOLOGY. A MONOGRAPH ON TRANSPIRATION.
NOTES FOR STUDENTS ae - = = - ~ id - - 155
é - « . e S 4 * - “ - a + ¥66

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Copyright, r904, by the University of Chicago.

5

VOLUME XXXIX NUMBER 2

BOTANICAL, GAZETTE
FEBRUARY, 1905

THE THEORY OF RESPIRATION.°
CHARLES R. BARNES.

I Ask you to consider with me a topic which is of fundamental
interest to physiologists, whether they concern themselves primarily
with animals or with plants. I take it the basal identity of the living
matter in all organisms and of its metabolism needs neither demon-
stration nor emphasis at my hands. Nor do I need to lay stress
upon the importance of respiration as one of these metabolic phe-
nomena, since it has been recognized from the earliest period as
indispensable to life. The phlogiston theory of the composition of
the atmosphere had scarcely disappeared below the scientific horizon,
before the fact was discovered that there occur, in animals and in
plants alike, an intake of oxygen and an output of carbon dioxid
which are intimately related to their existence. This became obvi-
ous to man, of course, in his own experience, a very superficial study
of the composition of the air inspired and expired from the lungs
showing that it had lost oxygen and gained CO,. This much of
Tespiration was early recognized to occur also with the larger ani-
mals, and a few years later like observations were made upon plants
by Prrestiey, and more accurately by LAVoIsIER and INGENHOUssS.
Even this knowledge of respiration was not possible before PRrest-
LEY’S discovery of oxygen in 1774, and the very remarkable revo-
lution in chemistry that followed in the closing years of the eigh-
teenth century. Yet this disappearance of oxygen and formation
of carbon dioxid are only the external indication of respiration, as
has on aie recognized.

of the retiring president before the Botanical Society of America, Phila-
detphia Seite: December 28, 1904. Published also in Science.
81

82 BOTANICAL GAZETTE [FEBRUARY

RESPIRATION IN ANIMALS.

Upon undertaking special consideration of this topic, I found it
needful to examine the recent literature of respiration in animals,
the aspect of the general subject with which I felt myself least
familiar. I found, to my very great surprise, that animal physi-
ologists have concerned themselves very little with the essential
problems of respiration. They seem to have been diverted to the
study of the mechanism of gas movements in the higher animals.
The lungs, with their intricate structure of lobes, lobules, atria, and
air “cells;”” the box in which the lungs are located, with its com-
plex muscular mechanism, and the very complicated mechanism of
innervation for the voluntary and involuntary movements which it
executes; the blood, and the physico-chemical relation of the gases
that enter and leave it in the lungs, of those that come into it from
the tissues, and of those it gives up to the tissues—these are the
topics that one finds exploited at length when he turns to the text-
books. I diligently examined the most modern and most thorough
text-books on animal physiology; such books as Foster’s Physi-
ology; STEWART’S Manual oj Physiology; the American Text-book
of Physiology; and SCHAEFER’s Text-book of Physiology; but in
them I found no treatment whatever, indeed no mention whatever, ©
of the real problems of respiration, that is, of what is happening in
the tissues—the processes of which these external phenomena are
the sign. Yet this much-studied respiratory mechanism, which is ~
so striking in the higher animals, is entirely wanting in the lower —
animals and in plants.

Not finding even a clue to the literature in the text-books, it was

_ only after much search that I was able to discover that anything at
all had been done; and it is so little that it is almost a negligible
quantity. There is an obvious reason for this, beside mere interest

_in the more striking phenomena. I am intending, however, neither
arraignment nor excuse, but a bare statement of what were to me

RESPIRATION IN PLANTS.

aoe ‘The knowledge of respiration in plants began about the same
- time—the os of the ear century—and advanced d rapidly

ra lea hie a

1905] BARNES—THE THEORY OF RESPIRATION 83

on account of the notable revolution in chemistry which took place
about this time. INGENHOUss, the Dutch naturalist, really ascer-
tained and published in 1779 the chief external facts of respiration;
at least he was able,to state them essentially as they were known
for twenty-five years “alter his time. In 1804, DESAUSSURE showed
that growth is dependent on respiration; that respiration is more
active in growing parts than elsewhere; that it is the cause of the
loss of weight to which plants are constantly subject; and later, that
the heat set free in flowers is related to the absorption of oxygen.
Not until 1833 was respiration treated comprehensively, when
DUTROCHET expounded the subject, comparing the respiration of
animal and plant, and showing it to be fundamentally alike in both.

Now at this point there began two remarkable misconceptions.
One was the confusion that arose between respiration and the manu-
facture of carbohydrates, which DuTRocHET called “diurnal respi-
ration.” Of that I shall not speak, save to say that the great weight
of Liesic’s authority made this error persist for half a century.

RESPIRATION AND COMBUSTION.

The other misconception was engendered by the comparison of
respiration to combustion. It had been observed by LAVOISIER
that the heat of the animal body was dependent upon respiration;
the heat of the plant body was shown by DESaussvReE to be related
to a disappearance of oxygen; combustion consumes oxygen and
produces heat; therefore, respiration is a sort of combustion. So
the argument ran.

It is quite impossible to overestimate the influence that this con-
ception has had on the study of respiration. The mischief it has
wrought depends chiefly, perhaps wholly, upon a misconception of
the actual mechanism of combustion, a process that has ever been
the béte noire of chemistry, as the history of the “phlogiston” theory
well shows. To our changed conceptions of combustion I shall
return later.

The idea of combustion, however, which dominated the argument
I have cited, was that oxygen combined with carbon to form CO,
and with hydrogen to form H,O. It was most natural, therefore,
to conceive that the food taken up by the organism stood to it in

84 BOTANICAL GAZETTE [FEBRUARY

the same relation as does fuel to the engine, and that what happens
is an actual oxidation of the food immediately and directly; in fact,
a process precisely parallel to the burning of the same food outside
the body.

One evident outcome of that idea is the current classification of
foods into plastic and dynamogenous—those which are useful in
building up the body, and those that are useful in producing heat
within the body; into “‘fattening foods” and “heat-producing”
foods. You are doubtless familiar with these phrases.

But if foods are “burned” in the body, it must be important to
know how much oxygen enters it, and how much carbon dioxid
and water leave it, so as to discern the ratio which exists between
them. Plainly a basis for this must be a comparison of the differ-
ences between the combustion of foods outside the body and their
“combustion” within the body. Yet, strangely, this has not been
made until recently. Without giving the full tables, let me show
the results arrived at by two observers, regarding two of the most
common plant foods, glucose and tartaric acid. These observers
assume, you will notice, that the processes are comparable. The

‘ co
results are stated as ratios of

oO, y
By RESPIRATION
By Com-
Foop BUSTIO’
Diakonow Purjewicz
Glucose - - - - - - - aoe 93
100 100 10°
Tartaricadd - - - - 160 = 162
100 100 100

_ Dtakonow’s whole series shows that in combustion the carbon
dioxid was always less than in respiration; Puryewicz found (with
the exception of tartaric acid, and even there the difference between
his results and Drakonow’s is in the same direction) that it was ©
always gi greater, his results being absolutely different in significance
oo. frome DraKonow’ s. And this is a good type of the results to be
ae — in Uecescreans the literature! I am not now concerned in

mining which set of results is correct, inasmuch as I believe

1905] BARNES—THE THEORY OF RESPIRATION 85

both are valueless, since on the assumption upon which they are
based neither can be interpreted.

RESPIRATORY RATIO.

Long before this sort of comparison was made, however, a
voluminous literature arose which was concerned only with the
ratio between the carbon dioxid given off and the oxygen consumed,
, and how this ratio was influenced by temperature, by light, by this
. q kind of food or that, by mere hunger, or by starvation. This ratio,
, the so-called respiratory ratio or respiratory quotient, the plant
physiologists really inherited from the animal physiologists, by whom
_ __ it was devised with reference to the gaseous exchange that occurs

in the lungs. This respiratory ratio has proved a veritable will-
o’-the-wisp, leading investigators into a bog where their labors and
____ their thinking were alike futile. For as a sign of what is going on
within, the respiratory quotient is absolutely valueless, however
interesting the facts in themselves may be. I could cite an indefi-
nite number of investigations to indicate this. I select a few cases.

As long ago as 1885, RUBNER showed! that the respiratory ratio
varied in resting muscles at different temperatures.

At 8.4° - - . . - “oe = 3.28
2.2 - - - - - 1.01
33-8 = a = = - 1.18
38.8 - - - - - 0.91

Von Frey and GruBeEr? showed that in a dog’s muscle, with
artificial circulation, contractions are accompanied by an increase
in the carbon dioxid added to the blood, but they found this increase
variable (46-10 per cent.) and less than the corresponding absorption
oj oxygen, so that the respiratory ratio became lowered during con-
traction. Tuissor3 showed that the production of carbon dioxid in

* Versuche iiber den Einfluss der Temperatur aster die fap cara des ruhendes
Muskels. DuBois-Reym. Archiv fiir Physiol. 1885:3

2 Versuche iiber den Stoffwechsel des Muskels. beer En Archiv fiir Physiol.
1885: 533-562.

3 Recherches sur i respiration musculaire. Arch. de Phys. Nori: et Path. V.
6:838-844. Also, Variation des échanges gazeux d’un muscle extrait du corps. Op.
et ser. cit. 7:641-653. 1895.

86 BOTANICAL GAZETTE [FEBRUARY

excised muscles was increased if the muscles were killed by heat or
were fatigued by prolonged stimulation. The output of carbon
dioxid in such cases was not related to the rate of absorption of oxygen.
Six years ago FLETCHER,* using Blackman’s apparatus, the most
intricate and accurate apparatus yet devised for following gaseous
exchanges, showed that the evolution of carbon dioxid from excised
frog’s muscles is independent of the amount of oxygen taken up during
the period. He distinguished in the production of carbon dioxid,
first, a short period (about six hours), which he thinks dependent
upon the presence of oxygen; and second, a long-continued evo-
lution of carbon dioxid ‘due to chemical processes occurring spon-
taneously within the muscle, in which complex molecules are replaced
by simpler ones, with the conspicuous results of the appearance of
[sarcolactic] acid and of free carbon dioxid.” He adds: “Under
suitable conditions the occurrence of active contractions in an excised
muscle is not accompanied by an increase in the rate at which carbon
dioxid is yielded by the muscle,” though oxygen is abundantly sup-
plied then by the blood. He does find, however, an increased for-
mation of other decomposition products.

CHAUVEAU and KAUFMANN, as long ago as 1887, found that the
output of carbon. dioxid from the levator muscle of a horse’s upper
lip was greater during activity than during rest, and contained more
oxygen than that absorbed in same time.5

A great number of researches of the same tenor can be found
in botanical literature. A single example must suffice. In an
elaborate paper PuryEwIcz shows® that the variations in the carbon
dioxid produced and the oxygen absorbed during a given period
under various conditions are not parallel, the amount of carbon
dioxid ranging within far wider limits than the oxygen. Thus, the —
carbon dioxid varied from —14 to 120 per cent. of the average; the
oxygen varied from o to 48 per cent. of the average. PURJEWICZ,
indeed, expresses his conviction that the respiratory ratio has no value
as indicating the actual course of respiration, and would separate :

4 Survival respiration of muscle. Jour. Physiol. 23:10-99. 1898.
5 Le coefficient de l’activité nutritive et respiratoire des muscles. Compt. Rend.
Acad. Sci. France 104:1126-1132. 1887.

oe _ © Physiol. Unters. iber Pflanzenatmung. Jahrb. Wiss. Bot. 35:573-610. 1990-

1905] BARNES—THE THEORY OF RESPIRATION 87

the taking up of oxygen and the production of carbon dioxid as
two processes, only indirectly related.

It.is clear that such results as have been cited became difficult
tito reconcile with the idea that respiration is combustion, and so an
attempt was made to evade the force of the facts, while maintaining
the comparison, by introducing a qualifying term and speaking of
respiration as ‘‘ physiological combustion.” This modification, how-
ever, blinks the difficulty; it does not remove it.

Before passing from this part of my subject I may mention another
false conception, which is more or less directly dependent on the
notion that respiration is combustion. One often finds respiration
described as a gaseous exchange—the taking up of oxygen and
giving off of carbon dioxid—a trade between the atmosphere and
the body. Clearly this is another case of transferring the superficial
interpretation of our own physiological processes to other organ-
isms. The exchange that takes place between the tissues and the
blood, between the blood and air in the lungs, gives the foundation,
and the unessential phenomena of respiration become substituted
for the essential. It would be quite as correct to describe photo-
synthesis as “an exchange of gases,” for carbon dioxid is taken up
and oxygen is eliminated. Yet no one ever thinks so superficially
of this process.

Joi lay

bia

are ey

ANAEROBIC RESPIRATION.

For three quarters of the last century it was supposed that the
evolution of carbon dioxid could only occur when free oxygen was
available. But in the early seventies PrLicER discovered what
Seemed a peculiar form of respiration. He found that a frog put
into a vacuum continued to give off carbon dioxid; and presently
the same phenomenon was observed by PFEFFER and others in
plants. So firmly had the conception of combustion fastened itself
upon physiologists, that when this anaerobic respiration came to be
explained, it was supposed that certain molecules of organic matter
within the cell gave up their oxygen to others, that they might thus
be burned in the body-furnace to yield energy. Hence arose the
term “intramolecular respiration.”

_ The study of anaerobic respiration, misleading as this early
\nterpretation of it was, has thrown in late years a very great light

88 BOTANICAL GAZETTE [FEBRUARY

upon normal or aerobic respiration. Here is a process which results —
in the evolution of energy, and gives rise to one important end-—
product of aerobic respiration, viz., carbon dioxid; yet it early became —
evident that it could not be counted a process of combustion, at least —
in any sense in which combustion was then understood. Plainly, —
the changes that were going on within the organism, which enabled —
it to give off carbon dioxid when no free oxygen was to be had, could —
only be a rearrangement of atomic groups within the molecule and :
the formation of products which were simpler than those from which :
they arose.
FERMENTATION.
The process of fermentation, first thoroughly explored by PASTEUR, —

of HANSEN and many others, are evidently related to those of respi-
ration by the nature of the end products and the conditions under —
which the processes occur. Indeed, when one compares the end —

to be identical in all respects. Other sorts of fermentation like- ©
wise yield many substances that are found originating in the metab-
olism of the higher plants.

We have, then, three modes of energy-release, which are evi-
dently closely related, if not identical; aerobic respiration, anaerobic
respiration, and fermentation. Their relations, so far as was known
in 1898, were stated by Prerrer in his Pflanzenphysiologie and —
need not be reviewed.

THE COURSE OF RESPIRATION.
In translating that work (p. 519) Ewarr wrote: “The actu
course ine — — the protoplast is quite obscure.” PFEFFER

Perhaps you will say, to foresee rather than to see—but hypothesis
must outrun demonstration. The advances to which we are ind

ae for — Lapsicas! are in three pee first, aa chemistry of

eT
STS ine

1905] BARNES—THE THEORY OF RESPIRATION 89

teids; second, the course of combustion, especially at low tempera-
tures; third, the nature of anaerobic respiration, and its relation
to aerobic respiration. Let me speak of these in order.

CHEMISTRY OF THE PROTEIDS.

A knowledge of the proteids, complex as they are, could be
obtained only by a study of their decomposition products. Now
there is a very remarkable uniformity in these decomposition prod-
ucts. No matter what the organism from which they are derived,
no matter how simple they are or how complex, when broken up by
the process of digestion, or by boiling with acids, they yield invari-
ably a series of products which have become in the last few years
much better known. These are amino- or amido-acids; such sub-
stances as leucin, tyrosin, arginin, glutamin, glycocoll, etc. Materials
of this kind are invariably present, and certain ones are so invariably
present that they can be used as the basis of distinctive tests for the
occurrence of digestion or similar decompositions of proteids. This
gave a clue to the nature of proteids which was followed by several
observers, notably by Kosset, in the study of what are believed to
be the very simplest proteids, because of the fewness and uniformity
of the fractions into which they break up. These are the prota-
mines. It has become clear from the study of these simple proteids
that they are made up in somewhat the same way as the polysac-
charides among the carbohydrates, that is by linking together a
series of the amido-acids. This is possible because the amido-
acids have a peculiar construction. They are, so to speak, different
on different sides. On one side is-an acid group and on the other a
basic group; and so the amido-acids can hang together in chains,
or even be condensed or polymerized to make a simple proteid.
Among the amido-acids, as in the carbohydrates, there are certain
atomic groups, like CH,, CH,, CHOH, CH,OH, COOH, etc.,
which recur again and again, and in such groups the possibility of
replacing a hydrogen atom or a hydroxyl radicle by some other
atomic group is very great.

Note, for instance, the comparatively simple acetic acid, CH,’
COOH. If we replace one of the three H atoms by the amido group,
NH,, we have at once an amido-acid, glycocoll, CH 2(NH,)° COOH,

go BOTANICAL GAZETTE [FEBRUARY -

which is one of the sorts of material out of which proteids can be
made. Out of an alcohol or out of a sugar we get just the groups
CHOH, CH,OH, etc., from which these amido-acids may be con-
structed when nitrogenous substances are present to supply the
amido group NH,. Thus the mode of construction of the proteids
has been found to show a likeness to that of the complex carbohy-
drates, and it has long been known that the carbon groups were very
much alike in both. It further appears that when the proteids are
digested by any organism they break down into these fragments, of
one sort and another, the amido-acids, the amides, etc., which may
be put together again in new form to constitute the peculiar pro-
teids of that particular organism. We may thus get one proteid
out of any other by the breaking up of the complex molecule and the
rearrangement of its constituent fragments. This fragmentation is
readily accomplished by the proteolytic enzymes, which probably
act on these bodies as the diastases do on carbohydrates.
OXIDATION.

The second important line of progress has been in the study of
the oxidation of carbon compounds at low temperatures. For our
purpose the important facts, which have only recently been developed,
are that the oxygen of the air does not combine directly with carbon
or with carbon monoxid to form CO,, or with hydrogen to form
H,0, as has heretofore been sup

As long ago as 1893 Drxon’s eeucket on explosive gases
showed that molecular oxygen was by far the most effective of the
atmospheric gases in retarding combustion. This surprising result
could not be interpreted then, and only in the light of TRAUBE’S
2 theory and the studies of BonE and others® on the oxidation of gases

90:25. 1904.

oxygen. Chea, awe
_ Chem. Soc. a 1903.

1905] BARNES—THE THEORY OF RESPIRATION gi

not undergo direct oxidation but hydroxylation, i. e., its hydrogen
atoms are successively replaced by hydroxyl radicles, with conse-
quent splitting into various intermediate products, such as carbon
monoxid and hydrogen peroxid, carbonic acid and water being the
end products. ARMSTRONG says:

There is little reason to suppose that changes take place at high temperatures
in rapid combustions in ways very different from those in which they occur at
lower temperatures... . . The effective operation is not the mere blow due to
impact or the vibration caused by this in the molecule, but the conjunction of
compatible molecules and the consequent formation of composite systems within
which change can occur. In so far as temperature influences the formation of
compatible systems, either as regards their character or the rate at which they
arise, temperature has an influence, but probably not otherwise.

I ask you to notice, then, that the process of combustion is now
being interpreted in the light of changes like those which have long
been known in organisms under the name of hydrolysis, and are
the characteristic mode of action of enzymes. Thus, when starch is
acted upon by diastase it is probably by repeated reactions between
water, dissociated into hydrogen and hydroxyl groups, and oxygen—
in other words, by continued hydroxylation—that it becomes ready
to fall apart into a series of dextrines and finally into maltose. Dias-
tase in some way facilitates this dissociation. Maltase takes up the
task, and maltose, further hydroxylized, cleaves into two molecules
of glucose. Then zymase may lend its aid and hydrolyze the glu-
cose molecule into lactic acid, breaking the latter still further into
carbon dioxid and alcohol.

The mechanism of the digestion of starch is not known in detail,
though the various intermediate products have been fairly well
studied. The usual assumption made is merely that water com-
bines with the starch under the action of diastase. I have carried
the theory a little further into detail, as seems warranted by the
studies of combustion. It is worthy of note also that the late steps
in the process, the hydrolysis of glucose by zymase, have been desig-
nated by the term ‘‘fermentation.’”? The combustion of starch has
likewise not been examined, but as the end products are identical
with those of digestion, it is not at all improbable that the inter-
mediate steps are the same, though they succeed one another too
fast to be followed by means at present available.

I need hardly remind you that our present ideas of the dynamics

92 BOTANICAL GAZETTE [FEBRUARY

of chemical reactions forbid us to believe that such dissociation does
not go on slightly at low temperatures, even when unaided. But it
is so slow as to be ordinarily beyond our measurement. The enzymes
seem to be mere accelerators of the several processes, perhaps pre-
paring “compatible systems,” as high temperature may do in com-
bustion; perhaps entering into union with the substance they a
on and forming compounds which are dissociable at ordinary tem-
peratures in appreciable amounts.

The clue to an understanding of respiration has been found,

ologists at least will do well to drop “combustion” altogether from —

their vocabulary, as neither the past conception of it nor its probab
use in the future conduces to clearness of thought.

NATURE OF ANAEROBIC RESPIRATION.

The third line of advance has been in a study of the relations of —

fermentation and anaerobic respiration. The first step was that

long-sought discovery by BUCHNER, that the process of fermentation

certain hexose sugars into carbon dioxid and alcohol. But a further
step in advance has lately been taken. It appears from the work of

BUCHNER an NHEIMER’® that the alcoholic fermentation is not
direct, but that it occurs always in indirect fashion, as shown below:
0 aes
| | |
CHOH sa a CHOH H CH,OH
|
CHOH = nee E CH, CH;
CHOH OH OH COOH OH co,
i
CHOH CHOn CHOH H CH,OH
| |
CH,OH H CH, CH, CH,
glucose water hypothetical 2 _— nor aay carbon dioxid
: substance ethyl alcohol

10 Die eieatoche Vorginge bei alcoholischen ST Ber. Deutsch.
ere 1

sua cs sala

1905] BARNES—THE THEORY OF RESPIRATION 93

STEPANEK has reached the same conclusion,'' and Maz&"? has found
acetic acid as an intermediate product in alcoholic fermentation by
a different yeast. The interest of the discovery that inactive ethy-
lidene lactic acid is the intermediate substance in this process of
fermentation lies in the fact that one of the two acids of which that
is composed namely, d-ethylidene lactic acid, or sarcolactic acid,
is formed as a product of respiration when proteids break down in
the working, fatigued, or dying muscle. FLETCHER observed this
as a more prominent product of contracting muscles than carbon
dioxid itself. Thus a regular product of fermentation is also formed
in the ordinary course of respiration.

The analogy between anaerobic respiration and fermentation had
been suggested early—even by PAstEUR—and has thus been growing
closer with each added bit of knowledge But the precise way in
which the destruction of the living substance went on in anaerobic
respiration was still unknown. Fermentation had been shown to
be due to an enzyme. Was anaerobic respiration also due to an
enzyme ?

Of course, enzymes are known to be present in a great many of
the parts of plants, and the oxidizing enzymes seemed to be the sort
to be sought. But none seemed to answer the conditions. At last,
however, the object appears to have been attained. STOKLASA, in
a series of papers published in various journals,*’ but all dealing with
the same general problem, declares he has found in various tissues
of animals, and in considerable number of plants, an enzyme analo-

™ Ueber die aerobe und anaerobe Atmung der Eier. Centralbl. Physiol. 18:
188-205. 1904. :

72 Utilization du carbone ternaire. Ann. Inst. Pasteur 18:277-303. 1904.

13 rere Identitat anaerob. Atmung u. Garung. O6esterr. Chem. Zeit.
1903. (Not se

STOKLASA, eek and VirEK: Der anaer. Stoffwechsel der héh. Pfl. und seine
Bezeihung z. alcoh. Garung. Beitr. z. Chem. Physiol. u. Path. 3:4 c

ASA and Cern¢: Isolierun| ng des die anaer. Atmung der Zelle der héh.
org. Pfl. und Tiere bewirk. Enzymes. Ber. Deutsch. Chem. Gesells. 36: 622-634
1903.

——: Ueber die anaer. Atm. der Tierorgane u. iiber die Isolierung eines garungs-
erregenden Enzymes aus dem Tie erorganismus. Zentralbl. Phy siol. 16:652-658.
1903.

Semen! Ueber die Atmungsenzyme. Ber. Deutsch. Bot. Gesells, 22: 358-361.
4.

94 BOTANICAL GAZETTE [FEBRUARY

gous to BUCHNER’s zymase, and like it glycolytic. This enzyme he
reports in leaves and roots of beet, tubers of potato, seeds, seedlings,
and young plants of pea, seedlings of barley, and entire plants of
Paris quadrijolia. Confirmatory results have (naturally enough)
been obtained by several students or assistants who have evidently
been engaged upon portions of the problem under the guidance of
STOKLASA. It is only fair to say that Mazé has strongly criticised —
Stoxiasa’s methods from the bacteriological side, and declares
himself unable to secure like fermentation under aseptic conditions ;"*
though Sroxiasa claims to have guarded carefully against infection
and to have rejected contaminated cultures. Independently, Mazé
has found what he calls zymase, in connection with pea seedlings,
Aspergillus and Eurotiopsis. He declares it “‘an enzyme normal to
all plants, arising like all other enzymes during vegetative (aerobic)
life.’ In the higher plants, however, and in most fungi it “is oxi
dized with the greatest ease, so that one never finds more than a
trace of it.”
Mazé and STox.asa interpret their results somewhat differently,
Mazé holding the process of fermentation to be a nutritive one,"
sugar being only assimilable when fermented and the nascent alcohol
thus made available, while Sroxiasa believes fermentation to be
merely anaerobic respiration and essentially a process for the imme-
diate release of energy.
Confirmation comes also from another source, for GopLEWSKI,"* _
working with lupines, finds similar products, and concludes that th
“anaerobic respiration is identical with alcoholic fermentation, of
at least in essence dependent on it.”
Moreover, sgenekion nist = Maxtmow"® wets found

Aspergillus an Tes i
7 = yme and is responsible

*4 Various papers in Annales Inst. Pasteur 18: 1903.
_ 4s IWANOWSKY in 1894 propounded the theory that alcoholic fermentation 15 4
pathological case in the nutrition of yeast, called forth by the abnormal com mposition.
of the nutritive medium.
os as — Ze Kennt. der — Evnases Bull. Acad. Sci. Craco’
(1904: 115-158. also his earlier paper with Porzentusz. Bull. cit. kot bs
37 Ueber Pha der Scien Ber. Deutsch. Bot. Gesells.
| 207-215. 1904.
ca ar: gi itber die auiteek Ber. Deutsch. Bot. Gesells. 22: 225-235-

1905] BARNES—THE THEORY OF RESPIRATION 95

| for the formation of CO, whether in aerobic or anaerobic respiration.
Thus several independent observers are testifying to the rather
widespread occurrence of an enzyme which brings about a disruption
’ of plant substance, under most varied external conditions, whether
; the plant be fed on one food or another,'® this dissociation resulting
| in the formation of carbon dioxid and of various other products.

THE MECHANISM OF RESPIRATION.

Let us now focus the light coming from the chemistry of proteids,
the mechanism of combustion, and the physiology of respiration, to
form a picture of what goes on in the body.

First: We should conceive of the respiratory dissociation as taking
place in the living material of the body and not in a food still unassimi-
lated. Experiments with a wide range of foods have shown that
they affect the intake of oxygen and the output of carbon dioxid in
the most diverse ways, whence it has been assumed that the respira-
tory ratio varies because of the way in which the given food is oxidized.
I do not say that it is not possible for the protoplasm to decompose a
sugar directly or to oxidize a fat. But it must be remembered that
in no case has it been experimentally proved that the food is directly
attacked, and that all the facts can be explained on the other assump-
tion, and some of them very much better than on the theory of direct
oxidation. Moreover, the lability of proteids which have been raised
to the life-level is their most striking characteristic as contrasted
with their ordinary stability.

In such labile material the second step is easily conceivable.
There occurs a shifting of the atomic groups within the molecule,
perhaps as a result of the last step in their anabolism—the addition
of hydroxyl groups from the water everywhere present. Dissocia-
tion follows necessarily; very slow perhaps at ordinary temperatures
and with a scanty supply of water, yet sufficient evidently for the
maintenance of life. Such conditions may very well be those obtain-
ing in resting organs, spores, and seeds. But normally this cleavage
May go on at a measurable rate, without anything more than the

*9 See a paper is capes which has just come to hand (Ueber die normale
und die anaerobe Atm g bei Abwesenheit von Zucker. Jahrb. Wiss. Bot. 40:563-
592. 1904), showing Tas erroneousness of DIAKONOW’s idea that anaerobic respira-
tion is only possible when sugar is supplied.

96 BOTANICAL GAZETTE [FEBRUARY

inevitable dissociation when hydroxylation has progressed to a certain
point. It seems, however, that there is generally—perhaps always—
a hastening of this process, and that the highly unstable protoplasm
is dissociated so rapidly that it liberates not only the energy imme-
diately utilized in growth, movement, etc., but also an excess suffi-
cient to be easily measured by so coarse an instrument as the ther-
mometer. Catalytic agents like the enzymes are certainly (I think
I may be permitted so strong an assertion) the usual accelerators.
And it is highly probable that an enzyme identical with zymase oF
at least analogous to it, is an active though secondary agent in this
acceleration. It may very well be also that those changes outside
the protoplast (whether without the organism or not) that are called
stimuli accelerate still further the katabolism, even to an explosive |
speed in some cases.

This primary dissociation may plainly be independent of free
oxygen, though it is hardly conceivable that there will not be some
oxygen present unless the plant has grown under most unusual con-
ditions, which one can scarcely realize experimentally. The products
of this decomposition are not sufficiently known, nor is their precis€
character important for our discussion. Among them are certainly
the more complex amido-acids, carbon dioxid, and alcohol.

Third: Up to this point the respiratory processes are quite alike

whether the plants grow in the air or apart from it. If sufficient
oxygen be not present, the disruptive processes may reach an equilib-
rium, just as an electrolyte practically ceases to pass a current of
electricity unless a depolarizer be present. So in the hydroxylation
of proteids, there is needed some substance to disturb constantly,
in one direction or another, the equilibrium that tends to be reached.
‘The common agent in this is oxygen. Of course, oxygen can ha

_be the ne ‘depolarizer that can promote further action. Thus,

ts characteristic “procs are carte

1905] BARNES—THE THEORY OF RESPIRATION 97

dioxid and alcohol. Indeed, lactic seems an equally characteristic
though transient product. The fact that hydrogen has also been
often recognized among them supports the interpretation of the
function of oxygen just suggested, and accords thoroughly with the
theory of hydroxylation. In that process hydrogen atoms from the
dissociation of water would be left free in case there was insufficient
oxygen to form H,O,,.

Fourth: But if the organism can get an adequate supply of oxy-
gen, the katabolism continues, some of the most complex previous
products breaking up by hydroxylation and thermal cleavage.
Among the fragments are undoubtedly some that lose in part those
very groups in which sugars, alcohols, fatty acids, etc., are peculiarly
rich. These are rebuilt at the expense of such foods, which there-
fore disappear as a result of respiration. That ethyl alcohol does
not persist when oxygen is present may mean either that it is decom-
posed, or that in its nascent state it is assimilated in the rebuilding
of proteids, for we have seen how easily acetic acid, one of its oxida-
tion products, can be converted into an amido-acid, glycocoll, and
be thus in direct line for reconstructive metabolism.

This, in its fundamental features, is the theory I have presented
in lectures to advanced classes since 1898, though always as more or
less a speculation. For various details I am indebted to the recent
literature, already cited. Because it is capable of explaining the
observed facts, which are now sufficiently numerous to demand a
coherent explanation, I conceive it to be entitled to the dignity of a
theory. Time forbids the discussion of details, and many points
have been considered that cannot be here presented.

This theory maintains the direct relation of aerobic and anaerobic
respiration, whose genetic connection was long since advocated by
PFEFFER. Anaerobic respiration is the primary process in all organ-
isms. Whether aerobic respiration occurs or not depends upon the
availability of oxygen. The relation of fermentation to the process
is not wholly clear; for although fermentation gives rise to the same
products as anaerobic respiration, this may depend in part upon
respiratory decompositions, such as have been described, and in part
upon digestion, which, as IwANowsKY and Mazé think, render the
alcohol from sugars available for assimilation. I am inclined to

98 BOTANICAL GAZETTE [FEBRUARY

Be
believe that in fermentation we deal with an exaggerated anaerobic
respiration, the active ferments being plants in which zymase is
produced in such amounts that it can attack sugars outside the organ-
ism, and thus secure sufficient energy with a minimum destruction
of the protoplasm.

ENERGESIS.

Finally, I may suggest that for didactic purposes it is desirable —
to have a word other than “respiration” to designate the disruptive —
processes by which energy is released, leaving “respiration” to-
designate the more superficial phenomena of aeration with whi
plant physiologists are but little concerned. Perhaps the word
“respiration” is already too firmly imbedded in literature to be so_
limited. It will at least do no harm to propose that the terms
“aerobic and anaerobic energesis” be considered, to which “fer-
mentative energesis” may be added if necessary. |

Tae UNIverstrv oF CHICAGO.

ee en ee

THE FORESTS OF THE FLATHEAD VALLEY, MONTANA.*
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY.
»XVIT.

HARRY N. WHITFORD.
(WITH MAP AND TWENTY-THREE FIGURES)
INTRODUCTION.

THE factors concerned in the grouping of plants are classified by
SCHIMPER? as climatic and edaphic. When the climate of any region
is such that a particular form of plant is favored, in most instances
it gives character to the vegetation. For example, in the eastern
United States the climate favors the tree form and gives rise to a
“forest formation.” If the grass form predominates, however, a
“prairie formation” is the result. Again, if the climate be such that
the cactus form gives the tone to the landscape, the “desert formation”
is developed. .Two such formations are shown in fig. 1, which is a
view of a portion of the Flathead valley looking west from a high
mountain. In the distance on the west side of the valley is a prairie
formation; on the east side is a forest formation.

However, if one stands on a mountain top and looks down into
these formations, he will observe that isolated areas in the forest do
not contain trees, but may have a prairie, a swamp, a clearing, or a
heath. The prairie formation may contain forests along streams or
on protected hillsides. Also, in the forest formation there may be
areas occupied almost exclusively by one, two, or more species of
trees; while at a little distance there may be another area with entirely
different trees. In other words, the composition of the forest
changes from place to place. To distinguish these local groups
from the general climatic grouping, SCHIMPER has called them
“edaphic formations.’ By other authors they have been called
“plant societies,” ‘plant associations,’ or merely “plant forma-

This paper is based on work as a collaborator in the U. S. Bureau of Forestry,
to which acknowledgment is here made for permission to publish.
CHIMPER, A. F. W., Plant geography upon a physiological basis (translated
by FISHER) 159-161. 1903.
1905] 99

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1905] WHITFORD-—FORESTS OF FLATHEAD VALLEY 1OL

tions.” Jig. z will also illustrate these smaller groups. In the
foreground may be seen a pond which contains its characteristic
plants; to the right of this a meadow; and between the pond and
Flathead Lake a local prairie. These are in the forest formation.
While the soil factors determine primarily the variety of the plant
landscape, other factors must be taken into consideration. In this
paper, however, the convenient term “‘edaphic formation”’ will be
used.

CowLes* was the first to show clearly that plant societies are
not static, but dynamic. In brief, his contention is that the plant
society changes with the physiographic changes that are constantly
going on. In a former papers I have attempted to correlate the
various plant societies found in northern Michigan. In that study
it is shown that the region lies in a deciduous forest formation, and
that while the coniferous forest societies are present in the more
xerophytic conditions, there is a tendency for them to be replaced
by the climax society, the beech-maple combination. The present
study, made in a region with a somewhat different climate, was
undertaken to determine whether the various plant societies could
be correlated in the same way. For this purpose a general survey
was made of the Flathead valley, and a portion of it was selected to
study in closer detail. No attempt was made to study the con-
ditions in the higher altitudes where a different climate prevails.

PHYSIOGRAPHY.°
There exists a close relation between the development of the
physiography of a region and the life history of its forest formation;
ence the necessity of describing the topographic features of the
Flathead valley. It is a well-defined physiographic unit, situated
in the northwestern part of Montana, about long. 114° 30’ W. and
lat. 47° 30’-49° N. (fig. 2). The altitude of the valley is approxi-
For a discussion of the subject, see Cowes, H. C., The physiographic ecology
of Chicago and vicinity. Bor. GAZETTE 31:74-76. rgot.
4 Loc. cit. pp. 75-108, 145-182.
5 WHITFORD, H. N., The ie development of the forests of northern Michigan.
Bor. Gazette 31: 289-325..

6 See ELrop, M. J., The ty siography of the Flathead Lake region. University
of Montana Bull. 17: 197-20

Io2 BOTANICAL GAZETTE [FEBRUARY

Fic. 2. ae a of northwestern portion of Montana, showing the eocit)
of Pateee valley.—After Etrop.

“The main singe system of the valley om of t .

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1905]

WHITFORD—FORESTS OF FLATHEAD VALLEY

Seale of miles

s2--+- TRAILS

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map.—After ELR

compare with fig. 1 and

104 BOTANICAL GAZETTE [FEBRUARY

Lake; its principal inlet, Flathead River; and its outlet, the Pend —
d’Oreille River. Flathead River is formed by the confluence of L
three branches known as the North Fork, South Fork, and Middle
Fork, all of which rise in the Rocky Mountains. These flow for
their entire course in valleys in the mountains and unite to form the :
main river near Columbia Falls. The Stillwater and Whitefish
Rivers are the principal branches that lie in the Flathead valley.
For about the lower half of its course through the valley, the Flat-
head River is a broad, sluggish, navigable stream that is constantly
depositing sediment. Especially is this the case at its mouth, where
a delta about 3*™ in length has been formed (fig. 4).

Flathead Lake (jig. 5) is some 40*™ from north to south and
varies in width from 10 to 20*™, its altitude being 890™. It is a
remnant of a former lake of much greater extent, which probably
covered a large portion of the valley at its head. An old outlet
near Dayton (jig. 3), some 120™ above the present level of the lake,
and terraces at approximately the same height, indicate the former
distribution of the waters of the lake. The Pend d’Oreille River
(jigs. 5, 6), the present outlet, has cut its way through a huge moraine
at the foot of the lake. This moraine no doubt acted as the dam
that backed up the water over the low valley lying to the north.
The river rapidly cut its way through this moraine until its present
condition was developed. This erosive process is going on much
more slowly today because the river channel has reached bed rock.

The Mission Range lies to the east of the lake and the Mission
valley. This is a distinct range separated from the other mountains
on the east by the Swan River valley. It has an altitude of approxi-
mately 2750™ at its southern end. From this altitude the mountains
become gradually lower until near the north end of the lake, where |
they merge imperceptibly into the valley. The Swan River valley |
opens into the Flathead valley where these mountains end (map and :
jig. 1). It is in reality only an arm of the latter, and during the time —
of the greatest extension of Flathead Lake the water probably backed —
up into this valley and formed an embayment. As_ the Pend 3
d’Oreille River cut its way through the moraine at the foot of Flat- —
head Lake, the level of the water in this embayment was lowered. |
But some time before the present condition was reached, a moraine

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WHITFORD—FORESTS OF FLATHEAD VALLEY

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1905] WHIT FORD—FORESTS OF FLATHEAD VALLEY 109

was probably thrown across the mouth of the Swan valley, and
thus its connection with the main lake was cut off. A gap in the
Mission Range offered a favorable place for the water thus cut off
to make its escape. This outlet, of course, marked the beginning
of that portion of the Big Fork River known as the “rapids” (map).
Just as the present condition of the drainage system of Flathead
valley is only a stage in the history of the destruction of the former
enlarged Flathead Lake, so the existing drainage system of the Swan
valley is only a stage in the history of the destruction of the former
more extended Swan Lake. This system consists of Swan Lake;
its inlet, Swan River; and its outlet, Big Fork River. Swan Lake
is a body of water some twelve miles long and, except at its upper
end, very narrow. It passes almost imperceptibly into the Big Fork
River. This is a meandering stream which with its branches drains
the valley and the mountains lying to the east and west of it. With
the exception of small rapids here and there, and that part of it known
as the “rapids,” the river is a rather sluggish stream. The valley
(jig. 8) is approximately 945™ above the level of the sea. The Swan
Range of the Kootenai Mountains rises abruptly from the east side
of the valley to an altitude of from 1800 to 2200", and the Mission
Mountains border the valley on the west.

Such, in brief, are the main features of the physiography of the
Flathead valley and its arm, the Swan valley. More detailed
peculiarities will be noted in connection with the discussion of the
edaphic formations; for, as will be shown later, there is an intimate
relation existing between the destruction of the two lakes and the
development of the plant formations that are found in the valleys.

GEOLOGY.

Flathead valley is not, as one would suppose, an erosion valley,
but according to Witu1s’ it is due to a fault resulting in the down-
throw of the region of the valley and an uplift of the region of the
present Rocky Mountains. The northern Rocky Mountains con-
sist of limestones, quartzites, and siliceous argillites about 2 750™
thick. During pre-Cambrian times the whole was under water.
At i ts sata: of the Cambrian age an uplift brought it above

» Bartrey, Structure of the front range, northern Bock? areata:
Mota isice N. S. 15:86-87. 1902.

[FEBRUARY

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BOTANICAL GAz

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1905] WHITFPORD—FORESTS OF FLATHEAD VALLEY 1 iB

water, and during all the Palaeozoic and part of the Mesozoic the
pre-Cambrian rocks were subjected to subaerial erosion. A move-
ment at the close of the Jurassic again brought the sea over it. Then
Cretaceous sediments of considerable thickness were deposited upon
pre-Cambrian limestone and quartzite. In post-Cretaceous times
a fold was overturned to the northeast. Some time in the Tertiary,
probably during the Miocene, the faulting mentioned above occurred.
The rocks tilt as a rule to the southeast, the northwest fault face of
the rocks being very steep.
SUMMARY.

1. The Flathead valley is due to a fault, and is technically known
as a rift valley.

2. The greater part of it was formerly occupied by a lake.

3- The present drainage system of the valley consists of the Flat-
head Lake, the Flathead River and its branches, and the Pend
d’Oreille River.

4. The Swan River valley, an arm of the Flathead valley, has
for its drainage lines the Swan Lake, the Swan River, and the Big
Fork River with its branches.

5. These drainage systems are remnants of the former more
extensive lake that occupied the valleys.

I. CimMate oF FLATHEAD VALLEY IN RELATION TO CLIMATIC
FORMATIONS.

It has already been shown that the climate determines the general
plant formation of a region. It is important, then, that the elements
of the climate be analyzed thoroughly, and it should be pointed out
at the same time in just what way these elements affect the distri-
bution of plants. In order to obtain a better understanding of the
true relation of the forests of this valley, its climate will be com-
pared with that of the northern peninsula of Michigan, where
both deciduous and coniferous forests are found; and with that of
the Puget Sound region, where the coniferous type of forest reaches
its highest development.

There are meteorological reports from two stations in the Flat-
head valley. These reports, though meager in some particulars,
will give a fair idea of the climatic conditions found in a prairie

1r2 BOTANICAL GAZETTE [FEBRUARY

formation and in a forest formation. The data collected from
Kalispell, Montana, cover a period of from four to five years. This
station is located in the prairie region of the valley. From a station
at Columbia Falls partial meteorological data were obtained that
will give an idea of the climate of the forested portion of the valley.
Marquette, Michigan, was chosen as the station to represent the
climatic conditions of northern Michigan; and Seattle, Washington,
will stand fairly well for the kind of climate found in the Puget Sound
region.®
TEMPERATURE.

The following table contains the latitude, longitude, altitude of
the stations, the number of years data have been collected, and the
mean annual and monthly temperatures (degrees Centigrade) for
the four stations.

TABLE TI.

Stations Lat. Long. | Alt. |Years| Jan. | Feb. |Mar. |April | May | June] July | Aug. | Sept. Oct. | Nov.| Dec. HEE
Kalispell......... boty? Pa 904™} 5 |—4.0/—4.5| 1.5] 6.0] 10.5] 14.0] 16.5] 16.0| 11.0] 7.0) 0©-5/—?-° ba
Columbia Falls. . . ego 046 | 5 |—s.o|—4.5| 0.01 6.0] 11.5] 14.0] 18.0] 14.0] 10.0, 6.0] 0.0|\-43 #
Marquette....... { See Ww. 224 | 29 |—9.0/—8.5|—5.0| 3.0] 9-0] 15.0] 18.0] 17.5] 13-5] 7-0}—9-5}75°° "i
Seattle...........| eit = oem 1.5 | 12 5.5] 5.5] 7-0] 10.0] 12.5] 5.5] 17.5| 17-5] 15.0] 10-5] 7-0) %° "

The significance of these figures will be clearer when it is known
just how the temperature influences the various physiological pro-
cesses. An examination of Table I and fig. 9 will show that the
mean monthly temperatures during the growing season is rather
low as compared with regions farther south; also, during the months
of May, June, July, August, September, the means for the four
stations are not far. apart; so that any difference in the type of
vegetation in the four stations cannot be accounted for by differences
in temperature during the so-called growing season. If the temper

8 The data given in the tables below were obtained from the following ee

in charge of the stations at their respective cities: Kalispell, Mont., H. B. Dic ; Mar-
quette, Mich., H. R. Parricx; Seattle, Wash., G. N. Satispury. The cael for
{in

of Montana for 1900 and 1901. No sunshine records are kept at Marquette, so those
from Escanaba, Mich., are taken to represent northern Michigan in that particular.

1905] WHITFORD—FORESTS OF FLATHEAD VALLEY Niaee
ature has anything to do with the difference in the character of the
vegetation, it must be the temperature of the non-growing rather
than of the growing season.

It is too often assumed that during the non-growing season physio-
logical processes of plants are inactive.? Of course, this is true of
growth; indeed, growth is checked and ceases rather early in the
season; but photosynthesis is known to occur during the winter
Experiments by MryAkE'° show that this process is active
at temperatures in the neighborheod of o° C. Among the evergreen
leaves experimented upon were species of pine and spruce. One
of MIyAKe’s conclusions is that starch is formed in winter, though

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), Kalispell (

Fic at seeps temperature wn Seattle ( ),
eaten Falls ( +), and Marquette ( —-====— ; the figures are temperature
Centigrade; winter r temperature for nai mans high, for Marquette low,

for Kalispell and Columbia Falls intermediate; summer temperature for all stations
nearly the sam

in small amounts, and its translocation occurs in the same season.
His work was conducted at Tokyo, Japan, where the mean tem-
perature of the three winter months is as follows: December, 5.1° C.;
January, 2.7°C.; and February, 3.5° C. If trees can do photosyn-

9 Meastan « ays that ‘‘in computing the sum of the positive or effective tempera-
t a minimum of 6° C. has been assumed as marking the inception of the period
of physiological activity in plants and of reproductive activity in Sa Miaheem,
‘od ife zones and crop zones of the United States. U. S. Dept. Agric., Div.
Biol. foes » Bull. ro: 4, note 2. 1808.

Migs » K., On the starch of evergreen leaves and its relation to photosyn-
thesis during a aiaaee: Bort. GAZETTE 33: 321-340. 1902.

II4 BOTANICAL GAZETTE [FEBRUARY

thetic work in the climate of Tokyo during the winter months, it is
possible that more food is manufactured by the trees in the Puget
Sound region, where the temperature is even higher than at Tokyo.
If this be the case for the winter months, it will be even more so for
the early spring and late fall months, when the mean temperatures
at Seattle are as follows: March, 7.22°C.; April, 10° C.; October,
10.5° C.; and November, 7° C.

The evergreen trees, however, in a climate like that at Marquette
would not be able to do so much work during these months, for the
temperature is very much lower. In other words, in a climate with
warm winters like that at Seattle, the evergreen trees can work more
or less during the winter, early spring, and late fall months; while
in a climate like that at Marquette, where the mean temperature is
considerably below the freezing point during the winter months,
this work would be very much checked if not stopped altogether.
As one would expect, from this standpoint, the conifers would be
more successful in the Puget Sound region than in the northern penin-
sula of Michigan. The conclusion that is reached from the forego-
ing is that, other things being equal, an equable distribution of heat
is favorable to conifers. A reference to the temperature conditions
in the Flathead valley will show that the climate is more equable
than that at Marquette, but not so equable as that at Seattle. The
coniferous forests are better developed here than at Marquette, but
are not nearly so luxuriant as at Seattle. From the standpoint of
carbohydrate manufacture, the deciduous trees are little or not at
all affected by the temperature conditions of the non-growing season,
for their leaves are absent, and consequently photosynthetic work
is very much reduced.

A comparison of the temperature conditions at Kalispell and
Columbia Falls shows a little difference, the latter being slightly
colder. While this difference may affect herbaceous vegetation, it
would likely have little or no effect on the forests; so the fact that
there is no forest at Kalispell, while there is one near Columbia Falls
will have to be explained on other than temperature grounds.

Transpiration, which also takes place during the non-growing
season, will be discussed in another connection.

as ae aha

CE ee EC fae ae Te EO ee ee ee ee

a
it
7
3
q
:

1905] WHITFORD—FORESTS OF FLATHEAD VALLEY ces

RAINFALL.

The greatest danger to trees is an excessive loss of water, and
therefore the rainfall of a region is very important. It will be pointed
out later how the conservation of this water supply influences the
character of the forest. Other things being equal, the greater the
amount of rain that reaches the earth, the more luxuriant the vege-
tation. The mean monthly and annual rainfall for Kalispell, Colum-
bia Falls, Seattle, and Marquette are as follows, the amounts being
given in millimeters: |

TABLE II.

Stations Jan. | Feb. | Mar.|April | May | June | July | Aug.| Sept.) Oct. | Nov.} Dec, |Ann’l
Malispell.... oi.c ceo 24 25 161 ax | 66} a7 | 33 28} 50 18} 49] 35] 415
Columbia Falls...... 48 38 301 a3) 68 72 a7 | Sr] 68) ged Sa} 30 | Sas
Marquette. ei cs. 4 52 45 47 5x 76 89 77 73 99 | 83] 69] 62 | 823
ORs oe Bos orth tir 94 79 | 82 86 | 40 17 14 | 45 76 | 143 | 158 | 945

The difference in the amount of rain on the west and east sides of
Flathead valley no doubt explains the difference in the types of vege-
tation in these two situations. It has already been shown that there
is only a slight difference in the amount of heat at the two situations.
It is very probable that the other elements of climate—light and
velocity of wind—are about the same for the two places, although
there are no data to prove this. It must be remembered that the
towns are only thirteen miles apart, but the difference in rainfall is
128™™, It is believed that this difference is sufficient to make a
forest vegetation at Columbia Falls and only a prairie vegetation at
Kalispell. Kalispell is well out in the valley, while Columbia Falls
is at the base of the mountains. The rain-bearing winds from the
west sweep across the high mountains west of the valley, where they
lose a considerable portion of their moisture, and then descend into
the valley where the higher temperature they encounter enables
them to hold more moisture. This is liberated, however, when the
winds bank up against the cool mountains east of the valley. Thus
the west side of the valley probably has a rainfall of 4oo™™ or less
and is not able to support tree growth except along streams; while
the east side has a rainfall of from 400 to 543™™, a sufficient quantity

116 BOTANICAL GAZETTE [FEBRUARY

to enable forests to exist. It is very probable that farther to the east
in the mountains there is a still greater quantity of rain. At least
the character of the vegetation would suggest that this is the case,
for here the forest growth approaches in luxuriance that of the Puget
Sound region where the rainfall is much greater.

A comparison of the rainfall in the valley with that of the Puget
Sound region and of the northern peninsula of Michigan will lead
to some interesting conclusions (Table IJ and fig. 10). In this com-
parison the data at Columbia Falls will be used rather than those at
Kalispell, for reasons that are at once apparent. The character of

100
= iaaeeEn
a A si aS Eee
50 Lede =
Seen 2 A ft =
sues gensecce7 caus s
iat TT | T
Oo (HEB Bi ETAL ia name!
(eae Bs | iB iy |
PELE TT tT EE i i
(Bee oD \GRREE Cirr TT CEECELET
(atens jaune Lees
ia EMRE (Zee CEE Bers EP gh
Ap

My Je RU Ags O Nw «ja F Mr

IG. Renn: sates rainfall for Seattle ( —), Kalispell (———), Colum-
bia Falls (——-—— -), and Marquette (-—---—--— ); the figures are millimeters of
rainfall; summer rainfall highest at Marquette and exceedingly low at Seattle; in
winter rainfall the reverse is true.

hi

ia) i og
rk Ach ial
mie

mit

ii a
‘ani
Att
Bo

SET

imei

the distribution of the rainfall is of considerable importance. Thus
at Marquette and Columbia Falls the rainfall is more or less evenly
distributed thoughout the year, with the five warmest months, May,
June, July, August, and September, having about half of the mois-
ture. Thus these five months at Columbia Falls show a fall of 2747”
out of a total of 543™™; and at Marquette 414™™ out of 823”
On the other hand, Seattle with its total of 945™™ has only 202™™
during these months. It is a well-known fact that the broad-leaved
deciduous trees evaporate more moisture during the summer months
than do the narrow needle-like leaves of the conifers. It is very
possible that the 202™™ at Seattle and even the 274™™ in the Flat
head valley are not sufficient to maintain the broad-leaved deciduous
trees in these climates. In any event, they are absent in the two

1905] WHITFORD—FORESTS OF FLATHEAD VALLEY 117

regions, except along water courses, and are present in the Marquette
region where they are more successful than conifers.

The winter distribution of rain is of extreme importance, especially
to those trees that hold their leaves, for KusANo'! has shown that
evergreen leaves transpire considerably even at temperatures below
freezing. While there are no data to show whether the bare twigs
of deciduous trees transpire more or less than those of conifers, yet
a prior it is very likely that they give off less moisture, for they have
less surface. If this be the case, conifers are more in danger of
desiccation during the winter months than deciduous trees; for even
though the amount of evaporated moisture is slight, it must be remem-
bered that the ground may be cold or even frozen, and that absorp-
tion is thus checked. However, if the rainfall is sufficient and the
relative humidity of the atmosphere is high, this danger is less. The
figures will show that the winter rainfall of the Puget Sound region
is excessive, while that of the Flathead valley and the northern
peninsula of Michigan is equal in quantity to the summer rainfall.
Conifers exist in all three situations. The extensive rainfall in the
Winter months at Seattle must be coupled with the fact that the
temperature conditions during these and the early spring months are
exceedingly favorable to certain physiological processes and account
for the almost tropical luxuriance of the trees of that region.

RELATIVE HUMIDITY.

The relative humidity of the atmosphere is another factor closely
associated with the amount of rainfall. The drier the atmosphere
the greater the transpiration. Other things being equal, the nearer
relative humidity is to absolute humidity, the less water will the trees
give off, and the less danger will there be of their desiccation. The
greater the saturation deficit, the greater the danger of losing water.
The following table will show the mean monthly deficits of the three
stations to be considered. Unfortunately there are no data from
Columbia Falls.

The high averages of the saturation deficit for July and August for
Kalispell, together with the small rainfall, probably account for the

™t Kusano, S., Transpiration of evergreen trees in winter. Jour. Coll. Sci. Tokyo
00: 313-366. 1902.

118 BOTANICAL GAZETTE [FEBRUARY

TABLE IIT.
Stations Jan. | Feb. | Mar./April | May | June} July | Aug. | Sept. Oct. | Nov.] Dec.
Kalispell 16.4] 21.2] 21.5] 30.3] 23.1] 26.1] 44.0] 41.2] 22.1] 25.0] 17.6] 14.6
Marquette 16.5] 16.4] 19.7] 23.7| 27.0] 27.0] 27.6] 23.3] 22.3] 10.8] 17.5] 17.4
Seattle 16.0] 20.7] 25.0] 29.2] 28.9] 30.9| 32.6] 29.6) 24.5] 18.4] 15.6] 15.5

prairie condition there and on the west side of the Flathead valley.
It must be emphasized, however, that the saturation deficit is of
great importance only in connection with the amount of water avail-
able in the soil. Ifthe ratio of the water obtained by the tree from
the soil to that given off is 1: —1, then the tree is in no danger of
desiccation; if, however, the ratio is —1:1 or 1: 1, the tree is in danger
of desiccation. From this it will be seen that if there is in the
soil plenty of water that the tree can obtain, there will be tree growth
though less luxuriant, even though the saturation deficit is high.
This accounts for the existence of trees along streams even in prairie
regions.

A comparison of the atmospheric deficits of Marquette and Seattle
reinforces what was said concerning the rainfall of these two regions.
It will be seen that both have a fairly low atmospheric deficit during
the winter months, thus decreasing the possibility of transpiration
at a time when the ground is cold. In the Puget Sound region the
comparatively high temperature renders greater transpiration more
likely than in the Marquette region, but this is offset by the fact that
the temperature of the soil is no doubt warm, thus rendering avail-
able for absorption some of the great amount of water that reaches
the soil in the form of rain during these months.

Again, a comparison of the atmospheric deficit data for the five
growing months of the year shows a uniformly higher deficit for
Seattle than for Marquette. This coupled with the fact that Mar-
quette has a rainfall of 414™™ during these months against 2027”
for Seattle makes deciduous forests possible in the former region,
but not in the latter.

VELOCITY OF WIND.

Another climatic factor that is likely to play a part in the dis-

tribution of the forest is the velocity of the wind. Perhaps the great-

1905] WHITPORD—FORESTS OF FLATHEAD VALLEY 119

est influence the wind has on trees is to increase the transpiration,
and they more than any other form of vegetation are subject to the
drying effects of winds. Indeed so important does SCHIMPER (pp.
542-555) think this factor that he considers it influential in bringing
about the prairie condition. Below is given the mean monthly and
annual wind velocities for the three stations. Again, there are no
data for Columbia Falls, but the amount of wind is probably not
very different from that at Kalispell. The figures are in kilometers
per hour.
TABLE Iv.

Jan. | Feb. | Mar.|April | May | June | July | Aug. |} Sept.) Oct. | Nov.) Dec. |Ann’]

Kaiepell, ... 500d 4.4) 7.7| 9-5] 10.4] Io.0] 10.1] 10.0] 9.6{ 9.0; 8.2] 7.2] 7.2) 8.8
Marquette............] 17.3] 16.1] 16.7] 15.4] 16.2] 12.6] 13.8] 14.2] 16.2] 18.0] 17.7] 18.0] 15.9
Seattle I10.3| f1.%| 16.6) TOcd|) OS] Geol 7:6f 6-6) 7.4) Svcl id. 1) 20.7; 6-39

As shown from these figures, one of the remarkable facts about
the Flathead valley is the absence of excessive wind. It is believed
that this, coupled with the rather low mean monthly temperature for
the summer months, is what makes possible so luxuriant a forest,
with a rainfall of about 500™™. In passing from the east side of
the Rocky Mountains into the Flathead valley, the decrease in the
amount of wind is very noticeable. Thus, Kipp on the east side of
the Rocky Mountains has a normal rainfall of 512™™, yet it has no
tree growth. Although there are no data to show that there is more
wind here than in Flathead valley, the fact is very apparent to one
who has been in the two regions. It is believed that the excessive
winds prevent the growth of trees, in spite of the fact that there is a
rainfall of about 517™™, or nearly as much as at Columbia Falls.

Compared with Marquette the wind velocity of Seattle is low.
This would again favor a more luxuriant vegetation in the latter
region than in the former.

SUNSHINE.

The more light, other things being equal, the more work trees
can do. On clear days more food is manufactured than on cloudy
days. This process can go on, as has been shown, during the non-
growing season when the temperature is not too low, though the

120 BOTANICAL GAZETTE [FEBRUARY

amount of food manufactured then is much less than at higher tem-
peratures. Light is the least variable of all the climatic elements.
Possibly for a given altitude and a given latitude the variability is
not great enough to have much influence on the kind of vegetation.
However, the sunshine data may prove of importance in comparison
with other regions. In the table below the mean possible hours of
sunshine, which would be the same for a given latitude, and the
mean actual hours are given. The observations cover a period of
short duration.

TABLE V.
JANUARY FEBRUARY Marcu APRIL
Poss. Act. Poss. Act. Poss. Act. Poss. Act.
Mahseell. 25ers 276.2 88.2 286.8 111.8 | 370.1 180.7 410.4 | 240.2
Escanaba 283.1 85.3 290.4 120.5 370.3 129.8 407.0 178.4
Seattle 276.2 66 286.8 | 103.8 | 370.1 168.6 | 410.4 | 205.2
May JUNE Juiy AuGUST
Poss. Act. Poss. Act. Poss. Act. Poss. Act-
pe! eee eee 471.3 | 231.4 | 479.8 | 279.6] 483.2 364.4 | 442.5 | 269.6
De oP na eee | 464.3 | 163.8 | 473.7 |] 238.7 | 475.7 | 211.5 | 437-6 | 230-5
Seattle 471.3 | 226.9 | 479.8 | 248.5 | 483.2 | 301.7 | 442.5 | 257-5

——

SEPTEMBER OcTOBER NoveMBER | DECEMBER ANNUAL

Poss. | Act. | Poss. | Act. | Poss. | Act. | Poss. | Act. | Poss. | Act

Kalispell — 202.5] 335.8| 174.6] 278.0] 73 | 262.1] 49 |4473-7|2262-9
PMMA, cial) ptel eae Bele al 376.1] 149.1) 338.5] 148.8] 284.1] 52.9] 269.6) 61.4) ..-- |1727-7
Seattle. 377-5| 186 335.8] 110 278 | 40.6] 262.1] 45 |4473-7 1959-6

ae

In his classification of climatic formations, SCHIMPER (ppP- 550-
565) does not recognize a distinct formation for the coniferous forest,
placing it in what he calls the summer-green forest (deciduous).
There is little doubt that he is right so far as the eastern part of the
United States south of the latitude of Lake Superior is concerned.
Here, as shown by Cow tes (/. c.) and the writer (J. c.), the conifer-

1905] WHITFORD—FORESTS OF FLATHEAD VALLEY 12!

ous forests occur as edaphic formations in the summer-green climatic
formation. In this region they are xerophytic formations (societies)
forming a stage in the progression toward the climax mesophytic
deciduous forest formation. With the wearing down of the xero-
phytic hills, the filling up of swamps, and the accumulation of humus
on the sandy plains and hills, the coniferous societies which now
prevail there will give way to the climax deciduous forest, which is
the highest expression of a climate similar to that found in the east-
ern United States.*?

The Pacific coast district north of San Francisco, and including
the moister mountainous regions inland, presents a climate entirely
different from that of the eastern United States. Here, as compared
with the eastern deciduous forest region, there is a more equable dis-
tribution of temperature throughout the year, with winter rains and
excessively dry summers. The forests here are coniferous, with the
deciduous element occupying only edaphic situations along water
courses. It is my belief that in some such climate as that found in
the Puget Sound region the coniferous forest is at its best, the
deciduous type being unable to compete with it because of the dry
summers.

The climate and character of the vegetation of the Pacific coast
region corresponds more nearly with ScHIMPER’s sclerophyllous wood-
land (pp. 464-469, 507-540) than they do with his summer-green cli-
mate and vegetation. Indeed, elements of the sclerophyllous vegeta-
tion are found in the Puget Sound district, for here such trees as Arbu-
tus Menziesii show a type of leaf decidedly like that found in climatic
districts which ScummpeER has so aptly called sclerophyllous woodlands.
The sclerophyllous formations are in a climate with winter rain and
comparatively high and equable temperature. Likewise the Pacific
coniferous district has winter rains and a comparatively equable
temperature, that is rather warm winters and cool summers. How-
ever, the mean average monthly temperature is much lower than
that of the sclerophyllous districts of the warm temperature belt.
This is no doubt influential in bringing about the narrow type of
evergreen leaf rather than the broad type found in the warmer climate.

*2 See ScHIMPER (p. 545) for table showing rainfall of Atlantic forest district and
Pacific coast

E22 BOTANICAL GAZETTE [FEBRUARY

For reasons given above, it is my belief that ScHIMPER’s summer-
green climatic formation should not include the coniferous forests
of the Pacific coast of North America, but that these should be
separated from it. In order to show its relations to the broad-
leaved sclerophyllous formation, I would suggest that it be called
the needle-leaved sclerophyllous formation. The limits of this forma-
tion are not clear. While no doubt it reaches its best development
in the Puget Sound region, this does not prevent its spreading into
more northerly regions with cooler but still damp winters. Whether
this type is the climax forest formation in the region north of the
Lake Superior district, as it is on the Pacific coast, or only edaphic
formations in the summer-green climatic formation of the United
States, our present knowledge cannot determine.

The forests of the Flathead valley clearly belong to the needle-
leaved sclerophyllous formation, but since they are on the border of
a prairie climate they are not so good an expression of it as is that
of the district farther west.

[To be continued]

STUDIES IN THE GRAMINEAE.
VIII. MUNROA SQUARROSA (Nutt.) Torr.’
THEO. Horm.

(WITH TWELVE FIGURES)

Ir often happens that plants which are common and easily avail-
able are the least studied. Being frequently collected and repre-
sented in herbaria by almost numberless specimens, such plants
become so familiar “by name” that botanists as a rule do not take
the pains to examine them with much interest or care.

The steady increase in the publication of cheap systematic works
in this country is hardly beneficial to the study of taxonomic botany.
The danger lies in the fact that authors of such works seldom take
the time to reexamine the plants or to compare them with diagnoses
of earlier date. It is no surprise, therefore, to discover points of
great importance mentioned in earlier diagnoses which have been
totally overlooked or ignored in works of a more recent date. As a
matter of fact, the older botanists were more careful in examining
and describing their plants, and their diagnoses, brief as they may
be, often allude to certain characters which are well worthy of
notice. A case in point may be illustrated by Munroa, and in select-
ing this genus as the subject of the present paper, it is with the inten-
tion of showing that the plant was actually better understood formerly
than it is now, and that it possesses certain morphological and histo-
logical characters which have not been observed hitherto and which
may be of some importance to future research.

Munroa was first described as a species of Crypsis—C. squarrosa—
by Nurrarz, but Torrey corrected the mistake and established the
genus.? At that time it was considered as belonging to the HORDEAE,
while it is now generally placed among the Festucear. As described
by Torrey, the plant is readily recognized, and both the singular

«Earlier papers were published in this journal as follows: I, June 1891; I,
August, 1891; III, October 1891; IV, November 1892; V, August 1895; VI, June
1896; VII, November 1896.

2 Pacific R. R. Rep. 4:158. 1856.

1905] 123

124 BOTANICAL GAZETTE [FEBRUARY

habit and the inflorescence are exceedingly well characterized.
There is one point especially which has escaped the attention of
later botanists, namely, that the spikelets are dimorphic in respect
to the shape of the glumes. It is true that Torrey did not express
this as plainly as he might have done, but his diagnosis certainly
shows that he did observe the different structure of the spikelets, as
shown by the following statement: “In the uppermost spikelet, and
often in the middle one, these nerves are bearded with long white
hairs towards the base; but the flowers of the lowest spikelet are
usually quite naked.” That BENTHAM and Hooker considered this
character of some importance is evident from their description of
the flowering glumes as being “‘longe ciliatis v. in eodem fasciculo
glabris.”’3 But none of the subsequent authors, not even the agros-
tologists, appear to have noticed this point, and it so happens that
the genus has been received as a plant of rather ordinary habit, dis-
tinct from Monanthochloe by “Spikelets usually in threes, terminal
in the axils of stiff spinescent leaves which project far beyond them.”

Since Torrey called attention to this slight modification in struc-
ture of the flowering glumes, it seemed natural to investigate the
matter further, as it might lead to the discovery of additional charac
ters sufficient to attribute a certain degree of dimorphism to the
spikelets. A careful examination of a large number of specimens,
studied in the field, revealed the fact that Munroa does exhibit such
dimorphism constantly, and to a greater extent than hitherto observed.
The deviation is not only due to the presence of long white hairs at
the base of the flowering glume in the uppermost spikelet in contrast
with the others, as first described, but the complete structure of these
spikelets is quite prominently distinct. As a rule, each inflorescence
(fig. 1) in Munroa consists mostly of three spikelets, one median and
two lateral; and when only two are developed, one of them shows
always the structure of a median and the other that of a lateral
spikelet. Of the two spikelets here figured, fig. 2 represents the
median and fig. 3 one of the two lateral. It is readily seen from
the drawings that the structure of these spikelets is distinct. The
empty glumes are very unequal (figs. 2 and 4) in the median, but

3 Genera plantarum 3:1180. 1883.

4 ScRIBNER and SouTHWoRTH, The true grasses. New York. 137. 1890-

1905] HOLM—MUNROA SQUARROSA 125

almost equal in the lateral spikelets (fg. 3); the flowering glume is
pubescent in both, but in the median there are three distinct nerves
with a tuft of long hairs at the base (figs. 2, 5, and 6), while in the
lateral the nerves are not visible, and no long hairs are developed
(figs. 3 and 7); moreover, the apex of the flowering glume is long-
awned and distinctly bidentate in the median (jigs. 5 and 6), but merely
emarginate and short-awned in the lateral spikelets (jig. 7). Some
distinction in regard to texture is also observable, the flowering glume

1G. 1.—Munroa squarrosa.—The inflorescence, magnified. Fic. 2.—A terminal
spikelet. Frc. 3-—A lateral spikelet. Frc. 4.—The empty glumes of a terminal
sR Fic. 5.—Flowering glume of same, dorsal view. Fic. 6.—Same, side view.

i ee Fiaaigaiaiag glume of lateral spikelet, side view. Fic. 8.—The prophyllon,
sit
and the palet being membranaceous in the median, but quite thick
and coriaceous in the lateral spikelets. It may seem justifiable,
therefore, to consider the spikelets of Munroa as being dimorphic,
even if we have not been able to observe any deviation in regard to
the distribution of the sexes within them.

While Torrey’s description of the spikelets contains a good deal
more than is recorded by subsequent writers, except BENTHAM and
Hooker, there are some other points which may be added to the
diagnosis of this singular grass. The mode of growth is very well

126 BOTANICAL GAZETTE [FEBRUARY

characterized by Torrey, but there are some peculiarities which
have been omitted, and these consist especially in the structure of
the shoots, the vegetative and the floral; furthermore, the histology
has only been touched upon very briefly.

The peculiar bushy habit of the plant depends upon the fact that
the culms are not directly terminated by the inflorescence. Usually
in the Gramineae the long internode of the culm passes immediately
over into the inflorescence, but in Munroa the spikelets are preceded
by two or three green leaves which are borne upon the culm near the
apex, and whose internodes are exceedingly short, often being barely
visible. These green leaves all subtend shoots, which develop into
floral or vegetative branches, and quite often into secondary culms
from the apex of which the same development of shoots is repeated.
Beneath the terminal inflorescence there is thus a small cluster of
shoots, developed in the axils of green leaves, a ramification that
calls to mind the well-known condition known in teratology as ‘“‘pro-
lification,’’ but which is normal and constant in Munroa. To sepa-
rate these shoots and to define their proper order might at first glance
seem difficult, not only on account of their relatively large number,
but also because the internodes are so very short. However, if we
examine the shoots carefully, it is easily seen that the’ first leaf of
several of them represents a fore-leaf (prophyllon) of most character
istic and striking shape.’ The presence of a fore-leaf actually gives
the clue to the composition of these “fascicles of leaves and shoots,”
since it always signifies a lateral ramification, and in the Gramineae
is nearly always situated on the branch with its dorsal surface toward
the mother axis. In Munroa this prophyllon has attained a high
development, and is not only represented by an open sheath-like
body, but the two nerves are extended into very long, stiff, divergent
setae (jig. 8); the sheath-like portion is membranaceous, ciliate, and
minutely pubescent on the dorsal face. Altogether it is a structure
so remarkable and characteristic that it may well deserve to be men
tioned in the diagnosis of the genus.

The composition of the shoots in Munroa is illustrated by the two
accompanying diagrams (jigs. g and 10), which represent two fre-

5 Some new anatomical characters for certain Gramineae. Beihefte Bot. Cent.
11:28. 1901.

‘
ae
LE eT

1905] HOLM—MUNROA SQUARROSA 127

quent instances of ramification. In these diagrams L', L?, etc., all
represent assimilating leaves with sheaths and blades developed as
ordinary grass leaves, and they differ in no respect from the others
which are situated at the base of the culms, where they form a
dense rosette. On the other hand, P*, P?, etc., are prophylla or

CS: O and 10.—Musvros squarrosa. Diagrams of two fascicles of shoots from
the apex of two culms: I, the green leaves; p, the fore-leaves; s, the spikelets; for
further explanation see text.

fore-leaves, and their form as indicated is almost like their cross-
Section. Thus their actual position may be readily understood,
the concave side (dorsal) turning toward the mother axis. S*, S?,
etc., are the spikelets, of which only the empty glumes and one of
the flowering have been drawn, in order to show the mutual position
of the spikelets—median and lateral—and of their respective glumes.

In fig. 9 the two leaves L' and L? are developed a very short dis-

128 BOTANICAL GAZETTE [FEBRUARY

tance below the inflorescence (51), which terminates the culm of the
main stem; they are exactly alternate, and so are the glumes of the
median spikelet (S‘); in the two lateral the same position is notice-
able, but these have been turned so as to form an angle of 90° with
the median, the terminal spikelet. In the axil of L' a complex of
shoots may be seen, beginning with a prophyllon (P*) with its back
toward the main axis, but alternating with the leaf Lt. The subse-
quent leaves (L3-L°) of this axillary shoot, on the other hand, are
turned go° to the side of the leaf Z', but are otherwise alternate.
The first of these leaves (L3) supports a bud with a prophyllon and
two small leaves, and similar buds are also observable in the axils
of the other leaves (L4—L°). The apex of the whole shoot terminates
in a single spikelet (S?), the arrangement of whose glumes, alternat-
ing with the green leaves of the same axis, shows that it is a median
and not one of the lateral spikelets; these have become suppressed
entirely in this instance.

There is still another shoot observable, however, namely the one
whose prophyllon is marked P?, and which has only two green leaves.
This little shoot comes from the axil of the large prophyllon P’,
Munroa thus illustrating a case, not unusual among Gramineae, in
which prophylla may subtend shoots. It is furthermore noticeable
that this prophyllon (P?) and the leaves of its shoot lie distinctly in
the same plane as the first leaves (L' and L?) and the first prophyllon
(P). In this way the spikelet S* terminates the main culm, while the
other (S?) terminates the shoot developed from the axil of the leaf L.*

The succession of these various leaves (L and P) seems invariably
the same, inasmuch as no instance was observed where the inflores-
cence was directly preceded by a prophyllon alone. But the number
of vegetative shoots in Munroa is sometimes much larger than that
of the floral, and the reason seems to be, at least judging from this
particular instance (jig. 9), that the shoots were so crowded within
the leaf axil that the development of flowers became arrested. This
explanation may seem justifiable when we examine the other diagra™
(fig. 10), which shows a more balanced growth of the whole syste™
of axes, instead of the one-sided development of shoots shown in the
former diagram (fig. 9), where no axillary branch was developed from
the leaf L?, but only from L’.

1905] HOLM—MUNROA SQUARROSA 129

In jig. ro there are represented four inflorescences and five vege-
tative shoots, but much more equally distributed than before, since
the main culm bears here a larger number of leaves—four instead
of two—three of which (L'-L3) subtend shoots. These leaves
(Z'-L4) are also exactly alternate and lie in one plane, and
the uppermost (Z4) precedes an inflorescence (S*) which termi-
nates the main culm. Similar to the specimen illustrated in
jig. 9, the first leaf (Z") subtends a shoot with a large prophyllon
(P*), and this shoot is also turned go° to the side of the supporting
leaf and the prophyllon; but it bears only two leaves (L‘ and L°),
one of which (Z5) has an axillary shoot consisting of a prophyllon
and a vegetative apex. The other leaf (Z°) has no shoot and precedes
the terminal inflorescence (S?) of three spikelets, arranged in con-
formity with the leaves (L5 and L°). Another shoot is observable
in the axil of the large prophyllon P', which shows the same develop-
ment as the one in jig. 9, but with the exception that an axillary shoot
is visible within the prophyllon (P); thus two prophylla of the shoot-
complex pertaining to leaf Lt support buds.

Examining the other shoots, we find one in the axil of leaf LZ? with
a prophyllon (P?), whose leaves and inflorescence are likewise turned
go° to the side, with exception of the prophyllon. This shoot con-
sists of a terminal portion with two leaves and a two-flowered inflor-
escence (S3; one median and one lateral spikelet), besides a sec-
ondary, purely vegetative bud which is developed in the axil of the
prophyllon, but although axillary its prophyllon has become sup-
pressed. The position of the floral shoot, however, shows clearly
that it is terminal, and that the other much smaller one must belong
to the prophyllon. A similar construction of two shoots pertaining
to one system is to be found in the axil of the leaf L3, where we observe
the same position of the prophyllon (P3) and of the green leaves as
before. The shoot bears two leaves and is terminated by a single
spikelet (S+), which shows the arrangement of the glumes as that of
a median rather than of a lateral spikelet, since the glumes alternate
with the leaves. It is now interesting to see that the prophyllon (P)
here also subtends a bud, and a purely vegetative one. The fourth
leaf (L+) has no bud and precedes directly the inflorescence (S‘),
which terminates the main culm; thus the median spikelet represents

130 BOTANICAL GAZETTE [FEBRUARY

the uppermost terminus, and its glumes are situated so as to alter-
nate with the four leaves (Z'-L4) borne on the culm. -

Such is the usual composition of the fasciculate shoots in Munroa
squarrosa, but there are of course many variations in proportion to
the number of shoots developed. However, the succession of leaves
appears constant, and the modification by suppression of some of the
spikelets or of the vegetative shoots does not influence the principal
structure as described above. It is a structure which no doubt is
uncommon among the Gramineae, even if the components of the
shoots may be identical. Somewhat similar and equally complicated
cases of ramification may be observed in Coix, in Andropogon, and
especially in Jouvea. North American Gramineae exhibit altogether
a number of types in regard to inflorescence and ramification of axes
which are very little known and which deserve attention. In Munroa
the most striking peculiarity depends, as stated above, upon the
presence of short internodes with green leaves above the longest
internode of the culm, preceding the terminal inflorescence. Fur-
thermore, it is characterized by the profuse development of shoots,
floral and vegetative, from the axils of these leaves; and also by the
elongation of some of these into secondary culms, at the apex of
which the same development of fasciculate shoots occurs.

From a morphological aspect, therefore, Munroa squarrosa oflers
several important characters, but it exhibits also some points of
histological interest, which may be mentioned in this connection.
As an inhabitant of the arid plains, it would be expected that our
plant would possess a xerophytic structure, which it does; but such
structure is far from common among its associates. Ecological
studies are very seldom extended beyond the mere fact that such and
such plants constitute associations; whether the external and internal
structure of such xerophytes, for instance, is in real harmony with
the surroundings is very seldom touched upon. The xerophytic
characters have been very excellently defined by several prominent
authors, by WARMING and ScHimpER for instance; but whether
these characters are to be observed in all the members of such
xerophilous associations remains to be seen. It would require a?
immense amount of work, and nothing more or less than a complete
investigation of the internal structure of desert plants. Very few

1905] HOLM--MUNROA SQUARROSA 131

of our plants have been studied from this particular point of view,
and the following notes on the histology of Munroa squarrosa may
be regarded, therefore, as a small contribution towards the knowl-
edge of this interesting plant association. With this object in view,
the writer has examined the structure of the roots, the culm, the
proper leaves, and the prophyllon, and finally the flowering glumes
of both forms of spikelets.

THE ROoTS.—The roots are thin, hairy, not very strong, rela-
tively short, and but little ramified. In a mature root the epidermis
and the moderately thick-walled hypoderm persist, but the cortex,

Fic. 11.—Munroa squarrosa.—Transverse section of a mature root: e, endoder-
mis; p, pericambium; /, leptome. X 360. Fic. 12.—Transverse section of a capil-
lary lateral root: h, proto-hadrome; the other letters as above. X 560
which consists of only three or four layers, collapses radially. The
inner walls of the endodermis (jig. rz, e) are considerably thickened
with distinct layers and pores; and it surrounds a pericambium
which is thickened throughout. Therefore the location of the proto-
hadrome vessels could not be clearly ascertained, and the inner por-
tion of the central cylinder is occupied by a mass of thick-walled
conjunctive tissue with about ten broad rays of hadrome. The lep-
tome constitutes very small groups, when viewed in transverse sec-
tions (fig. rz, 1). A younger stage of the root shows these same ele-
ments much less thick-walled. Thus it was possible to distinguish the
pericambium cells from the proto-hadrome vessels, and it was noticed
that the latter had penetrated the pericambium in all places. If
one of the capillary lateral roots (fig. r2) is examined the same struct-
ure is observed, but the center of the root is here occupied by a large
vessel, and the hadromatic rays are merely represented by four proto-
hadrome vessels (fig. 12, h), all of which are located inside the peri-

132 BOTANICAL GAZETTE [FEBRUARY

cambium. We do not attach much importance to this varied position
of the proto-hadrome, however, since we have ascertained that it is
not constant, a fact that has been recorded in some previously pub-
lished papers.° A characteristic of the root of Munroa seems to be
the poorly developed cortex in contrast with the prominent and
thick-walled inner tissues of the central cylinder.

THE cutM—The long internode is semicylindric, smooth, and
almost glabrous at the base; but cylindric, deeply furrowed, and
scabrous above. The smooth and quite thick cuticle covers a very
small-celled epidermis, whose outer walls are prominently thickened.
Stomata are frequently near the furrows of the upper part of the culm,
and their guard cells are level with the epidermis. Unicellular,
pointed projections abound along the elevated ridges, but decrease in
number toward the base, where the surface of the culm is smooth.
The cortical parenchyma is well differentiated in the upper part,
where it occurs as a single layer of palisade cells, filled with chloro-
phyll, and bordering on the parenchyma sheath of the peripheral
mestome bundles; the palisades are very regularly arranged, radi-
ating from the parenchyma sheath. Another tissue of about five
layers, but composed of colorless, more roundish cells, occupies the
space between the peripheral mestome bundles with their covering
of palisades, and this tissue belongs also to the cortex. It is developed
to a somewhat larger extent at the base of the culm, where no pali-
sades were observed; thus the cortex consists here of a continuous
ring of colorless tissue surrounding the mestome bundles.

The mechanical tissue as stereome is well represented and forms
sub-epidermal groups in the ridges, bordering on the palisades,
besides occurring as a solid, continuous zone all around the inner
mestome bundles. Toward the base of the culm the stereome does
not occur as sub-epidermal layers, but only as a heavy ring inside
the cortex and surrounding both the outer and inner circle of mestome
bundles. Throughout the culm there are thus two almost cot
centric circles of mestome bundles, the peripheral ones being the
smaller, orbicular in transverse section, and containing mostly lep-
tome alone; while the larger inner ones are oval and contain both

6 Compare: Eriocaulon decangulare L. Bot. GAZETTE 31:17. 1901; and Studies
in the Cyperaceae. XIV. Amer. Jour. Sci. 10:278. 1

1905] HOLM—MUNROA SQUARROSA 13%

leptome and hadrome. Common to both is a thick-walled mestome
sheath, but the parenchyma sheath is developed only on the outer
face of the peripheral bundles, thus bordering on the palisades, being
quite large-celled and containing chlorophyll of a somewhat bluish-
green color. This parenchyma sheath is thus restricted to this
particular portion of the peripheral mestome bundles in the upper
part of the culm, where palisades occur, and is entirely absent in all
the others. The inner portion of the culm is occupied by a thin-
walled pith, which is broken in the center.

Characteristic features of the culm, therefore, are the small develop-
ment of chlorophyll-bearing parenchyma in the shape of palisades
and as an incomplete parenchyma sheath, and the predominating
chlorophyll-less cortical parenchyma which evidently has the function
of storing water. The stereome is well represented, being thick-
walled and constituting heavy layers both in the ridges and inside
the cortex.

THE LEAF.—The leaf blade is furrowed on both faces, deepest
on the ventral, and covered with small obtuse papillae and somewhat
larger prickle-like projections like those observed on the culm. The
midrib is hardly larger than the other nerves, the blade thus being
nearly flat and of the same thickness throughout. Stomata occur
on both faces of the blade along the sides of the furrows, their guard
cells being level with epidermis. While the cells of epidermis are
small and quite thick-walled on the dorsal face of the leaf, those of
the ventral are larger and developed as bulliform cells in the furrows
between the mestome bundles.

The parenchymatic tissues of the leaf consist of a colorless water-
Storage tissue, which is located in the furrows beneath the bulliform
cells, and consisting of three or four strata. There is also a chloro-
phyll-bearing tissue, developed as one layer of palisades surrounding
the mestome bundles and radiating from them. Similar palisade
cells were also noticeable in the upper portion of the culm, and
bordering on the incomplete parenchyma sheath.

The mestome bundles are all orbicular in transverse section, and
Possess a complete green parenchyma sheath besides a mestome
sheath, which is somewhat thick-walled in the larger bundles. The
leptome and hadrome are well differentiated in all the nerves, espe-

134 BOTANICAL GAZETTE [FEBRUARY

cially in the larger ones. In regard to the mechanical tissue, a stere-
ome occurs in large groups, one in each margin of the blade, besides
smaller groups accompanying the nerves, and located above and
below them between epidermis and palisades.

Characteristic of Munroa and several of its associates among the
Gramineae are the nearly orbicular mestome bundles surrounded by
the single layer of straight-walled palisades, radiating from them and
bordering on a rather large-celled parenchyma sheath, inside of which
a typical mestome sheath occurs; and also the deeply furrowed blade
with bands of bulliform cells covering strata of colorless tissue, the
function of which is evidently to store water. The numerous epi-
dermal projections in the shape of papillae or long-pointed spines
partly surround and cover the stomata.

THE PROPHYLLON.—This very characteristic leaf, as stated above,
consists of a sheath-like, bicarinate, membranaceous body and two
long setae, the nerves being extended beyond the apex of the leaf.
The structure of the sheath-like portion is very simple, since the
margins and the central part possess only an epidermis (the dorsal)
with a few stereomatic cells scattered here and there. The two keels, in
which two mestome bundles are located, show a more solid structure,
since each keel has a support of quite thick-walled stereome, which
borders on a large-celled, colorless parenchyma. Each mestome
bundle has a closed parenchyma sheath and a moderately thickened
mestome sheath. The awns contain chlorophyll, located in a stratum
of palisades and in the parenchyma sheath, and stomata occur on
the upper surface. A cross-section of one of these awns is sharply
triangular, with the two lateral angles occupied by stereome, while
the keel contains a large mass of colorless parenchyma. The mestome
bundle is located in the center, but is very small and consists only of
leptome.

THE FLOWERING GLUMES AND THE PALETS.—As stated above, the
glumes of the terminal spikelet are somewhat different from those of
the lateral in regard to shape and texture, and therefore the spikelets
may be designated as “dimorphic.” This distinction, howeve!,
seems much more pronounced when the internal structure is com
pared. The empty glumes of both spikelets possess the same struc
ture, even if their form be distinct; but the flowering glumes and the

aaa ree RS ee

1905] HOLM—MUNROA SQUARROSA 135

palets exhibit a structure so distinct that when detached the spikelets
may be easily assorted into terminal or lateral by their tissues alone.
In the terminal spikelet the flowering glume and the palet are thin
and membranaceous, as in the majority of Gramineae. The flower-
ing glume has four prominent ribs on the dorsal face supported by
heavy groups of stereome, and small scattered groups of this same
tissue are observable also on the ventral face of the glume, but only
between the two innermost mestome bundles. Otherwise the glume
consists only of a single layer throughout, represented by the dorsal
epidermis. The palet of this same spikelet—the terminal—is two-
nerved, but very slightly bicarinate, and the structure is even more
delicate than that of the flowering glume.

In the flowering glume of the lateral spikelets the structure is
very different, a large mass of very thick-walled stereome covering
nearly the whole ventral face of the glume, and a colorless parenchyma
occupying the larger inner portion. The four mestome bundles are
imbedded in this tissue, thus forming no prominent ribs, as is the
case in the terminal spikelet. A similar broad stereomatic tissue is
observable in the palet on the ventral face, besides a colorless tissue
nearer the dorsal. The two nerves are located in this tissue, which
extends into two sharp keels with numerous epidermal projections.

So great a diversity in structure observable in the glumes of
spikelets belonging to the same inflorescence is evidently very rare
within the order. It seems the more peculiar since the spikelets are
otherwise identical, the number of glumes being the same, and the
flowers perfect in both the terminal and lateral spikelets. The very
firm structure, described above, which is especially marked by the
unusual development of stereome, is also known in other genera.
We might mention, for instance, the coriaceous or cartilaginous
empty glumes in MaypEar and ZoystEAE; the cartilaginous flower-
ing glume and palet in PANICEAE and OryzEAE; finally the promi-
nent structural deviation noticeable in the unisexual spikelets of
Buchloe and Amphicarpum. But none of these cases are really
comparable with that of Munroa, where the only distinction between
the spikelets depends upon position, terminal or lateral, and not upon
the distribution of sexes, and still exhibiting such marked modification
in structure.

136 BOTANICAL GAZETTE [FEBRUARY

The systematic position of Munroa seems naturally to be within
FESTUCEAE, but it is difficult to place it near any of the genera so
as to demonstrate its nearest affinity. Most frequently Munroa is
placed next to Monanthochloe, a genus to which it certainly shows
no affinities whatsoever. The peculiarity of the genus seems to
depend upon the dimorphism of the spikelets, and to some extent
upon the profuse development of shoots, especially vegetative, in the
axils of leaves near the apex of the culm, a structure which is exceed-
ingly common in teratological cases, but which is normal and con-
stant in Munroa squarrosa.

There may exist some analogous cases, and normally so, among
the other members of the order in this country, but so far as known
to the writer no investigations have been published dealing with such
morphological peculiarities.

BROOKLAND, D. C.

ALTERNATION OF GENERATIONS IN ANIMALS FROM
A BOTANICAL STANDPOINT.?
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY.

CHARLES J. CHAMBERLAIN
(WITH TWO FIGURES)

SINCE zoologists do not recognize in animals an alternation of
generations comparable with the alternation of gametophytic and
sporophytic generations in plants, it may seem presumptuous for a
botanist to propose any theories. Nevertheless, after hesitating for
several years I have decided to publish my belief that animals exhibit
an alternation of generations comparable with the alternation so
well known in plants.

In short, the theory is this: the egg with the three polar bodies
constitutes a generation comparable with the female gametophyte in
plants; similarly, the primary spermatocyte with the four sperma-
tozoa constitute a generation comparable with the male gametophyte
in plants. All other cells of the animal constitute a generation com-
parable with the sporophytic generation in plants, the fertilized egg
being the first cell of this series.

In support of this theory I shall present two lines of evidence:
(1) the gradual reduction of the gametophyte in plants, with the
constantly diminishing interval between the reduction of chromosomes
and the process of fertilization; and (2) the phenomena of chromatin
reduction in both animals and plants.

1. EVIDENCE FROM THE REDUCTION OF THE GAMETOPHYTE IN
PLANTS.—For convenience, the female and the male gametophyte
will be considered separately. Further, since dioecism in gameto-
phytes is regarded as a more specialized condition than monoecism,
and since the tendency is toward dioecism, all cases in which the
gametophytes have not attained the dioecious condition may be
disregarded.

* Presented before Section G of the A. A. A. S., at the Philadelphia meeting,
December 30, 1904.

905] 137

138 BOTANICAL GAZETTE [FEBRUARY

(1) The female gametophyte—In the liverworts the gametophyte
is a green, independent plant. It is the conspicuous generation,
the sporophyte being comparatively small and never entirely inde-
pendent. Here the gametophyte generation, beginning with the
spore mother-cell and extending to the fertilization of the egg, is a
complex plant, of which the egg, the culmination of the gametophyte,
constitutes only a very small part. In the mosses the disparity
between the gametophytic and sporophytic generations is not so
marked, but the gametophyte is still predominant and independent,
while the sporophyte never becomes entirely independent. In the
ferns the sporophyte has become the conspicuous, independent gen-
eration, and the gametophyte is much reduced. In most cases the
gametophyte is still a green independent plant, although it is so small
that it is likely to be overlooked by the layman. . In the heterospor-
ous forms the gametophyte is still smaller, develops little or no
chlorophyll, and shows but little differentiation. Compared with
the liverworts or mosses, or even with the homosporous ferns, the
interval between the reduction of chromosomes and the fertilization
of the egg has been immensely reduced.

In the gymnosperms the gametophyte is entirely dependent, being
parasitic in the sporophyte at all stages of its development. Although
there is a more or less prolonged period of nuclear division before
cell walls are formed, the ancestors of these gametophytes were doubt-
less cellular from the beginning of their development. In Gnelum
Gnemon, cell walls are formed only in the lower portion of the gamet-
ophyte, in the upper portion the nuclei lying free in a common layer
of cytoplasm. Any one of the free nuclei of the upper portion seems
capable of functioning as an egg nucleus. In other species of the
same genus no walls are formed even in the lower portion, all the
nuclei lying free in the general cytoplasm. In the angiosperms,
both the conditions shown by the genus Gnetum are found, but
reduction has proceeded much farther. A considerable tissue 8
formed in the lower portion of the gametophyte in Sparganium, 1D
the grasses, and to a less extent in some Compositae and other forms;
the tendency, however, is toward the free nuclear condition already
reached by the latter type of Gnetum. The gametophyte of Peper
mia, with its sixteen free nuclei, one of which functions as an €88

E
;
5
|

1905] CHAMBERLAIN—ALTERNATION OF GENERATIONS 139

nucleus, does not differ very markedly from the free nuclear type
of gametophyte seen in some species of Gnetum. Four mitoses of
the simultaneous type give rise to these sixteen nuclei, one of which
becomes the egg nucleus. Between the reduction of chromosomes
and the fertilization of the egg there are only four mitoses. In
many angiosperms, as in Lilium, the interval is still shorter, the
gametophyte containing only eight nuclei, resulting from three mitoses.
This is the most reduced female gametophyte yet discovered in plants,
and it shows only one more mitosis than does the gamete-producing
generation in animals, which regularly shows two mitoses.

Up to this point I have made no mention of the tetrad of spores,
a feature almost universal in plants above the thallophytes. Where
the tetrad is present, the reduction of chromosomes takes place dur-
ing the two mitoses by which the four spores are formed from the
mother-cell. It is only in some heterosporous forms that the tetrad
is incomplete or fails to appear. In such cases the reduction takes
place during the first two mitoses in the mother-cells, so that whether
a tetrad is formed or not, the reduction takes place during the first
two mitoses in the mother-cell. To suggest that the gametophyte
of the Lilium type represents a reduced tetrad might only cause
confusion, and since the question does not affect essentially the
theory proposed, I shall not discuss it at this time, but shall depend
for comparison upon forms in which a tetrad is present. With a
few well-known exceptions, one member of the tetrad develops,
while the other three are evanescent and have no obvious function.
It is only rarely that any nuclear division occurs in the three evanes-
cent spores of the tetrad. In the functional spore of the tetrad three
mitoses occur, giving rise to eight free nuclei, one of which becomes
the nucleus of the egg. :

I believe that this latter condition—the most prevalent one in
angiosperms— corresponds closely with the egg and three polar
bodies so characteristic of animals. In my opinion, the egg with
its three polar bodies constitutes a generation directly comparable
with the gametophytic generation in plants. The accompanying
diagram (fig. 7) illustrates the comparison. Animals do not furnish
any example more directly comparable with the condition shown in
D of the diagram. In D the cell which we have compared with the

140

BOTANICAL GAZETTE

[FEBRUARY

animal egg has undergone some development so that it contains eight

nuclei.

This is no serious objection, however, when we remember

that in the pines the cell directly homologous with the larger cell in

D contains

oO
& co ro io)

. 1.—A-D, successive

ee
3

reduced; the shaded nucleus
in D is the egg nucleus.

thousands of nuclei, and in Gnetum contains too large

a number to be counted accurately, and
within the angiosperms shows a series
ranging from more than a hundred nuclei
down to sixteen and then to eight. Com-
paratively few plants have been investi-
gated, and it is not improbable that plants
will yet be found which will complete the
reduction series by showing only four nuclei,
or only two, or even only one. The latter
condition, with the megaspore functioning
directly as the egg, would be illustrated by
C of the diagram.

To me the comparison seems so obvious
that I can explain the previous absence of
a theory of alternation of generations in anl-
mals only by the fact that the gamete
bearing generation is’ extremely reduced
and is not approached by any gradual
series as in plants. Had observations in
plants been confined to angiosperms, there
would doubtless be no theory of alternation
of generations in plants.

(2) The male gametophyte—It is not
necessary to trace in detail the reduction
of the male gametophyte. Originally 4
conspicuous, independent plant, it becomes
reduced, loses its independence, acquires’
the parasitic habit, and then undergoes
a progressive reduction until it becomes 4
simple, microscopic, parasitic structure
which no one would think of homologizing
with the gametophyte of a liverwort, were
it not for the close series leading to the

Lal

1905) CHAMBERLAIN—ALTERNATION OF GENERATIONS 14

reduced condition. In many gymnosperms the vegetative tissue has
become reduced to two evanescent ‘‘prothallial” cells, and even these
are lacking in mostforms. In the angiosperms it is only rarely that a
vestige of the vegetative tissue in the form of a prothallial cell is found.

The formation of a tetrad of spores
seems to be universal. The interval
between the reduction of chromosomes

and the formation of the sperm is gradu- , a
ally reduced, until in the angiosperms

there are only four mitoses in the game- (|e) 2)
tophytic generation.

I believe that the condition which pre- B B
vails in angiosperms is directly compar-
able with that found in animals. This
comparison is illustrated in the accom-
panying diagram (fig. 2). Here again .
the plant shows a slightly more extended () (0) (0) (o)
development, the condition shown in E CO} (o}(2)
of the diagram having no parallel in ani-
mals. But, as in the case of the female
gametophyte, the objection is not serious, :
for the male gametophyte has been gradu- ae
ally reduced from a conspicuous, inde- ad
pendent plant to a microscopic, parasitic Fic. 2.—A-E, spermato-
structure with only three nuclei. Itis not genesis in an angiosperm. 4,
at all improbable that instances will yet
be found in which the last two mitoses nuclei in Erepresent the sperm
have been suppressed, the microspore nuclei, the unshaded nucleus
functioning as a sperm. This condition
could then be illustrated by E in the ie B’-D’, succeeding stages;
diagram. the reduction of chromosomes

The sporogenous tissue which pre- ee eae me soa
cedes the formation of megaspore mother- =
cells and microscope mother-cells is comparable with the oogonia and
Spermatogonia of animals. Preceding the formation of megaspore
mother-cells this tissue becomes much reduced in the higher plants
and in many cases is altogether lacking.

142 BOTANICAL GAZETTE [FEBRUARY

2. EVIDENCE FROM THE REDUCTION OF CHROMOSOMES.—The
reduction of chromosomes is a phenomenon so complex and so unique,
and at the same time so essentially identical in plants and in animals,
that any adequate theory must be applicable to both kingdoms.
In plants the reduction always occurs during the first two divisions
of the spore mother-cell, whether the plant be homosporous or
heterosporous. In animals the reduction occurs during the first two
divisions of the primary oocyte and primary spermatocyte. The
structural character of the mitoses are too well known to need any
review at this time. According to STRASBURGER? the reduction of ©
chromosomes indicates a return to the more primitive gametophytic
generation, and this view is quite generally accepted by botanists.
I am aware that zoologists distinguish somatic and germ plasm,
and that the application of this theory of alternation with its terms
gametophyte and sporophyte (or rather similar terms, since “ phyte”
is obviously unsuitable) would cut the line of germ plasm, leaving
part in one generation and part in the other. The sporogenous tissue
of plants is quite analogous to the germ plasm of animals, and yet
most botanists have no hesitation in assigning it to the sporophyte
and in regarding the mother-cell as the first term of the gametophytic
generation. The gametophytic generation—whether long as in the
liverworts or short as in the angiosperms—is characterized by the
reduced number of chromosomes; the sporophyte, except possibly
in cases of parthenogenesis, contains double the number of chromo-
somes found in the gametophyte. We should recognize two gener
ations in animals, characterized respectively by the reduced and the
double number of chromosomes.

COROLLARY.

It is obvious that this theory, if well founded, affects not only
the application of sexual terms in plants, but also our notions in regard
to the nature of sexuality. It is hardly necessary to say that most
terms relating to sexuality in plants have been borrowed from zoolo-
gists. Many botanists, especially the more rigid morphologists,
have attempted to confine sexual terms to the gametophytic genet

2 STRASBURGER, E., The periodic reduction of chromosomes in living organisms:
Ann. Botany 8:281-316. 1894.

.

1905] CHAMBERLAIN—ALTERNATION OF GENERATIONS 143

ation, which is spoken of as the sexual generation, while the sporo-
phyte constitutes the asexual generation. They have ceased to speak
of male trees and female trees, or male flowers and female flowers,
and have ceased to regard the stamens and carpels as sexual organs.3

Notions as to what constitutes a male individual or a female indi-
vidual among animals, or as to what constitutes a male organ or
female organ, are firmly established. If the method which the
rigid botanical morphologist applies to plants should be applied to
animals, the result would be interesting, for there would be no male
or female individuals, nor would there be any male or female organs.
All sexuality would be confined to the microscopic egg with its three
polar bodies, and to the spermatocyte with its four sperms. The
gametophytes of bryophytes, pteridophytes, and most gymnosperms
bear well-developed archegonia, which are correctly designated sex
organs, but by a gradual reduction the archegonium in the angiosperms
becomes reduced to its essential structure, the egg. It is only through
respect to its ancestry that this egg may be termed an organ. The
antheridium is only less reduced. In animals there remain only
these essential elements, which by similar courtesy may be called
organs.

I prefer to apply the terminology as it is applied by zoologists,
and consequently should regard dioecious sporophytes as male and
female individuals. In the gametophytic generation the dioecious
condition is universal in heterosporous plants. Stamens and car-
pels, which contain the male and female gametophytes, may be
termed male and female organs as properly as may the reproductive
organs of animals. It is strictly correct to speak of male and female
gametophytes in plants; but in my opinion to designate the sporo-
phyte as an asexual generation is a mistake. The sporophyte is
male or female as truly as is the gametophyte; and, like the indi-
vidual in animals, it is characterized by male or female reproductive
organs which produce the male or female gametophytes.

I do not claim any acquaintance with zoological literature further
than a reading of the latest edition of W1tson’s “The cell in develop-
ment and inheritance.’ Were there any theories as to alternation
ny oe discussion of monoecism and hermaphroditism would neither rein-
en the argument these subjects are omitted altogether.

144 BOTANICAL GAZETTE [FEBRUARY
of generations in animals, doubtless they would have been thoroughly
discussed in that book.

It is hardly necessary to state that the theory has no bearing what-
ever upon relationships or interrelationships. It merely adds another
to the already long list of parallel processes, and suggests that the
theory of alternation of generations in plants is applicable to animals.

THE UNIVERSITY OF CHICAGO.

BRIEFER 2 UICEES.

NEW PRECISION-APPLIANCES FOR USE IN PLANT
PHYSIOLOGY. It
(WITH FOUR FIGURES)

IN the first of these articles I gave the reasons which have led to the
development of some new pieces of apparatus for educational use in plant
physiology, and I described two of these pieces. In brief, believing that
improvised apparatus has been brought to its fullest practicable, if not to
an actually harmful, degree of development, I am trying to devise for each
of the leading physiological topics appropriate normal apparatus, viz.,
appliances which, manufactured expressly for the specific work, will yield
accurate quantitative results, will be applicable to their work with economy
of time and effort, and will be obtainable by purchase at any time from
the stock of a supply company. In the foregoing article I described the
new clinostat and the new portable clamp-stand; below are accounts of
three additional appliances, and similar descriptions of other pieces in
advanced preparation will follow later. They all are now, or soon will be,
for sale by the Bausch & Lomb Optical Company, of Rochester, N. Y.

III. AUTOGRAPHIC TRANSPIROMETER.

A practicable and obtainable form of autographic (self-registering)
transpirometer, suitable for use both in educational demonstration and also
in certain lines of investigation, is one of the first desiderata of plant physi-
ology. Several forms have been devised, of which the best known are
the “évaporométre” of Richard Fréres, and the registering balances of
Woods and of Anderson. The first of these, while obtainable, has serious
limitations both in practice and in principle; while the two others, though
admirable in their accuracy, must be made to order at large cost and with
much delay, and they are somewhat elaborate withal. My new instru-
ment, illustrated in the accompanying jig. I, is constructed as follows:
A cylinder, shown in the upper part of the figure, contains on a spiral
track between its outer and an inner wall some 2 50 spherical gram weights.
These weights are steel bicycle balls of one-fourth inch diameter, the same as
Anderson used in his balance; they weigh almost exactly one gram each.

* Continued from 37:307. April 1904.

1905] 145

146

BOTANICAL GAZETTE

Fic. 1.

2

Fe ee

1905] BRIEFER ARTICLES 147

and vary not over one-thousandth of this weight from one another. These
feed by gravity, one at a time, into a simple releasing valve, so arranged
that when acted on by an electro-magnet a slide rises and allows one ball
to drop through a tube into a scale pan, a new ball immediately taking its
place in the releaser-slide. Attached to the releaser-slide is a bar carrving
a pen, so adjusted that every time the slide moves, that is, every time a
ball is dropped, the pen makes a vertical fine line with chronographic ink
upon a record paper attached to a revolving drum. In use the plant to
be studied is prepared in the manner usual for transpiration studies, and
is balanced on a scale pan of any good balance, while the transpirometer
is adjusted beside it. As the plant loses water this pan rises, and at the

fT
I

ETE eT rT =
LE ?
— ew[v—_w_€€_wo§_i aml v
cc AAA t
ce AE
oO E_wc_cEoSE Tek RMA
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Fic. 2.—One-third the true size.

top of its swing is made to touch a wire, thus closing an electric circuit;
this excites the electro-magnet, which then raises the releaser-slide, drop-
ping a ball into the scale pan (which is immediately depressed, breaking
the circuit), and making a mark on the record paper. This operation is
repeated thereafter every time the plant has lost a gram of water. The
drum revolves once in twenty-four hours, and the record paper, a reduced
copy of which is presented in the accompanying jig. 2, is divided into
numbered spaces corresponding to the hours; these spaces are subdivided
into twelve parts, each therefore representing five minutes,? and these in
turn can easily be read by estimation to fifths, or one-minute intervals.
It is possible, therefore, to read off from the drum directly the number of
minutes it takes the plant to lose one gram of water, data which are readily
transformable into other terms. The record paper is divided into seven
horizontal spaces, marked by initial letters, one for each day of the week.
The pen slides on the bar, which contains seven notches; and each day,
when the plant is watered and the clockwork is wound, the pen is slipped
along the bar one notch. Each record paper therefore contains a complete
att five-minute spaces are exactly 1mm broad; hence millimeter cross-section
Paper may be used instead of the record paper. This happened to be the case at the
time the photograph (fig. 1) was made.

148 BOTANICAL GAZETTE [FEBRUARY

record for a week. The tripod stand is adjustable for height, and can be
leveled, while the mechanism in use is protected by a glass bell-jar.

Such is the instrument in its present form.3 For some special purposes,
such as for use out of doors, it would be better to have the weight-cylinder
and the recording drum separated, so that the latter may be removed to
any desired place in laboratory, lecture-room, or elsewhere. Probably
this form. will later be obtainable. For greenhouse and general demonstra-
tion use, however, the arrangement figured will be found most convenient.
The releaser and reservoir are arranged for gram weights, which are the
only ones likely to be needed in educational work. For special investi-
gation purposes lighter or heavier balls could of course be used after appro-
priate alterations in the size of reservoir and releaser. The instrument
may be used with any balance sensitive to a gram; but a special balance,
adapted expressly for transpiration work, and provided with a mechanism
to prevent oscillation under action of the wind, is in preparation, and will
later be described.

IV. ADJUSTABLE LEAF-CLASP.

In several phases of the study of the physiology of leaves it is necessary
to apply some object or special device to two exactly corresponding areas
of the two surfaces of a leaf. The most familiar instance of this occurs in
the application of Stahl’s cobalt chloride method to the study of transpira-
tion, and there are other cases nearly as important. For these purposes
some simple devices are improvised from clamps, watch-crystals, mica,
etc., but there is at present no obtainable appliance by which this end can
be accomplished with certainty, celerity, and convenience. The new
leaf-clasp, designed to meet this need, is illustrated in the accompanying
fig. 3, and is constructed as follows. Two similar brass rings, ‘‘chambet-
rings,” each 3o™™ in diameter and 5™™ in depth, are attached at the
ends of parallel flexible-elastic bars, so arranged that they hold the rings
firmly and exactly edge to edge, while allowing of their separation, by
means of a screw, to any desired extent.4 For each ring there are provided
two accessory rings. One of these is right-angled in section and holds 4
removable cover-glass, so that when pushed over the exposed edge of the
chamber-ring, it converts the latter into a glass-topped chamber, as show?
in the figure. If disks of filter paper treated with cobalt chloride (and

3 While this description is in press it has been decided, in order to give the inst™
ment greater compactness, to place the weight-cylinder and recording drum side by
side, instead of one above the other.

4 In the final form of the instrument, a second screw, not shown in the figure,
has been added, permitting the rings to be brought still more tightly together if needful.

1905] BRIEFER ARTICLES 149

preferably cut slightly larger than the chamber-rings so they will cling
half-way up the latter) be placed in the chambers, which are then applied
to the leaf, the change of color in transpiration may be observed with the
greatest clearness and facility. Incidentally, the tightness of the chambers
(when not on the leaf) permits the papers to retain their dryness and red
color for a considerable time, so that there is no need of haste in applying
them to the plant. The second accessory ring is broken, and is intended
to hold tinfoil or any fabric tightly to the chamber-ring. Thus if projecting

FiG. 3.

veins prevent a connection of chamber withleaf "sufficiently tight for
Some special purposes, a band of thin rubber may be held by the broken
ring in such a way that it will project against the leaf, filling the spaces
between the veins. But there is a more important use for the broken
ting, which is to attach tinfoil patterns or screens to the chamber-rings
for the demonstration of the necessity of light in photosynthesis. The
common method of demonstrating this important physiological fact by
vias of corks or other material attached to both faces of a leaf is fallacious
(since the absence of starch thus shown is due as much to absence of
carbon dioxid as to absence of light), and a logical proof involves use of

150 BOTANICAL GAZETTE [FEBRUARY

a method which cuts off light without cutting off the carbon dioxid. Most
leaves have all, or most, of their stomata on the under surface, so that if this
is left free, the upper surface may be covered by screens as closely as desired.
By means of the broken ring a tinfoil screen, with a pattern cut therein,
(preferably backed by a thin glass), or a photographic film or similar device
may be attached to the inner edge of one chamber, this is then applied to
the upper face of a leaf, with the lower chamber-ring pressing the leaf

mly against it. The access of light is now shut off from the lower sur-
face by a simple accessory diaphragm arrangement (not shown in the
figure), which does not impede gas passage; and such an arrangement
gives a perfectly logical, and beautifully clear and conclusive demonstra-
tion of the necessity of light for starch formation. Of course, this end may
be attained by simpler make-shift devices; the virtue of this instrument
consists simply in the facility and certainty with which the end may be
reached. As shown by the figure, a universal joint and a screw-joint
fitting over any upright support permit the apparatus to be adjusted at any
desired height, plane, or angle.

V. LEAF-AREA CUTTER.

The most striking and conclusive way of demonstrating the funda-
mentally important fact of increase of organic substance through photo-
synthesis is by Sacus’s method of comparing the morning and evening dry
weights of equal areas of similar green tissue; but it is rarely employed
because of the inconvenience of the manipulation. The leaf-area cutter
here figured (fig. 4), and described below, is designed to permit all parts
of this valuable experiment to be performed with exactness and facility.
The cutter works on the principle of a punch; the steel dies, operated by
proper handles, cut disks cleanly from a leaf between them, the disks then
dropping into a perforated aluminum cup screwed below the lower die.
The diameter of the punch-dies is as nearly as possible 1.128°™"s and
hence every disk is nearly 1°™? in area. In use the arms of the punch
are slipped above and below the leaf, when the disks may be cut very
conveniently and rapidly, in any desired number, care being taken t0
avoid the larger veins. The larger the number taken the better, since
local variations in thickness, etc., may thus be compensated, and 100 is a
fair number. The cup is then unscrewed and covered by its own screW
cap, which projects sufficiently to allow the cup to hang near the top of
an ordinary test tube, as shown in the figure. Water in the bottom of the
test tube is then boiled in the gas flame (a convenient holder for the test
tube being shown in the figure), and the steam enters the perforations of

1905] BRIEFER ARTICLES 151

the cup, killing the living cells of the disks, thus preventing any loss of
weight by respiration. The cup with its contents is then placed in the
drying oven. In the evening, using the other cup, an equal number of
disks is cut from corresponding positions in similar leaves, or, better, is
cut from the second halves of leaves, from the first halves of which the morn-

FIG. 4.

ing disks were taken. These are treated as for the first
series and the cup is placed beside the first in the drying
oven. When thoroughly dry both cups are carefully
weighed, the weights of the cups (stamped upon them,
together with the letters M and N respectively to distinguish
the morning and night cups) are subtracted, when the remainder gives the dry
Weight, which is always greater for the evening than for the morning
set. If steps be taken (by use of the methods described by SACHS) to
eliminate transfer of material into the stem, the difference in the weights
shows approximately the amount of substance formed through photosyn-
thesis in the particular plant studied in so many hours per so many square
centimeters of surface. In practice the performance of the instrument
1S very satisfactory.

In addition to the above-mentioned educational demonstration use, the
instrument should also serve in those phases of ecological investigation
where it is desirable to determine the relative photosynthetic powers of

152 BOTANICAL GAZETTE [FEBRUARY
various sun and shade plants, or of the same plants under sun and shade
conditions. It should also permit of the determination of the photosyn-
thetic constants of many ecologically important plants, as a part of their
physiological life-histories, without a knowledge of which their real relations
to their physical environment and to other competing plants cannot be
understood.—W. F. Ganonc, Northampton, Mass.

CURRENT LITERATURE.
BOOK REVIEWS.
The teaching of biology.

THE SIxTH volume of the American Teachers’ Series is devoted to a discussion
of the teaching of biology, by F. E. Ltoyp and M. A. BrcEtow of the Teachers
College, Columbia University.° The book is in reality two books in one cover,
‘the first by Professor Lioyp, devoted to the teaching of botany, and the second,
by Professor BicELow, to the teaching of zodlogy. With the latter we are not here
concerned.

Dr. Luovp considers two general topics which lie somewhat outside of his
title, namely, the value of science (particularly of biology) in education, and nature
study. The remaining chapters treat of the value of botany, the principles
which should determine the content of the botanical course, the various types of
botanical courses that have been proposed, and the botanical principles to be
emphasized. These topics are followed by detailed discussion of a course in
botany for high schools, and of the equipment and materials for the laboratory.
The final chapter consists of a list of books most useful for teachers.

It is hardly necessary to say that such a book as this, from the hand of a suc-
cessful teacher and a skilled botanist like Dr. Lioyp, will be of great service to
teachers, especially to those whose training is somewhat inadequate and who
therefore are familiar neither with the literature of the subject nor with those
general principles and outlines which ought to be presented in secondary schools.
The number of such teachers is still unfortunately large. Nor is it to be doubted
that even the more experienced teachers and those better trained will profit by
many of the suggestions and discussions, for Dr. Lioyp has clear, definite con-
ceptions, and a pedagogical rather than an academic point of view. His chapter
on nature study is commended to the serious attention of those who are advocates
of this sort of work in the primary schools. The chapter on the ‘“‘method of
thought” in teaching as against the “‘method of discovery” is admirable in many

iculars; but it must not be forgotten that even this method depends on the
skilful teacher, and such an one may even “introduce a system of lenses between
the eye and the object” without confusing his pupils.

___ The part of the book for which the author naturally expects the most criticism
is that which discusses the types of botanical courses and marks out one which
appeals most strongly to himself. ‘There is no gainsaying the value of this course,
and no one more strenuously than the author would insist that there are other
eas

*Lioyp, F. E., and BicEtow, M. A., The teaching of biology in the secondary

School. T2mo, pp. viiit+491. New York: Longmans, Green & Co. 1904.
1905] 153

154 BOTANICAL GAZETTE [FEBRUARY

courses equally valuable. It cannot be too often repeated that only the incom-
petent slavishly follow courses prescribed by other teachers. Each really good
teacher is obliged to make his own in details, though he may follow another's
general plan.

The book certainly does not overshoot its audience. One who is familiar
with the teaching of botany in schools of the upper Mississippi valley and west-
ward is tempted to think that Professor Lioyp’s ideal has already been realized
and perhaps even surpassed in a considerable number of the better high schools;
and he wonders whether the book is not better adapted to the teachers and schools
of the North Atlantic states, which, in the teaching of science, have certainly
lagged behind the less conservative western ones.

It would be of no service to attempt to summarize the author’s discussions,
which will well repay serious consideration, and should be read both by the college
man and the secondary teacher.—C. R. B

A monograph on transpiration.

‘THE ABSOLUTE necessity that the investigator be thoroughly cognizant of
what others have done and thought in his field, together with the ever-increasing
hopelessness of coping with this task, offer probably the most serious discourage-
ment to the scientific man. This difficulty, however, can be very much lessened
by the publication from time to time of exhaustive monographs in restricted fields.

the whole subject of transpiration,” a subject whose literature has been extraor-
dinarily difficult to compass. The book is a natural growth from the author's
three previous bibliographical and critical papers on this subject, Materialien 2u
einer Monographie betreffend die Erscheinungen der Transpiration der Pflanzen,
which appeared from 1887 to 1901, but it is much more than a bringing of these up
to date; it is not a mere critical bibliography but a thorough treatise on the various
phases of the subject. New experimental data are scattered through the work,
being presented in connection with the discussion of the several questions involved.
Besides true transpiration, we have also discussed here the allied phenomeno?,
guttation, in a chapter which is altogether the best treatment that this subject has
yet received. A very complete bibliography occupies thirty-three pages at
end of the volume, a Niigata in which reviews are uniformly noted as well
as the articles themselves

The subject-matter is , divided logically into thirty chapters, each of which
deals with a certain part of the general field. The headings of some of these at
as follows: Methods of investigation, Light in general, Light rays of certain
refrangibility, Atmospheric carbon dioxid, Air temperature, Humidity, Air move
ments, Chemicals, Periodicity, Guttation, etc. The language is clear and concise

? BURGERSTEIN, ALFRED, Die Se apeig der Pflanzen. Imp. 8vo. pp- x +283,
jigs. 24. Jena: Gustav Fischer. 1904. M 7.

1905] CURRENT LITERATURE 155

and the matter so well arranged that the reader has little trouble in finding the
discussion of any particular point in which he may be interested. Different
readers will doubtless differ as to the relative importance of the several subjects,
but it seems on the whole that the balance is well preserved. All workers in plant
physiology will find the work practically indispensable. The only source of
regret is that so many minor a aa errors should have escaped the proof
reader.—B. E. LivincstTon.

NOTES FOR STUDENTS

AMONG THE MOST important ecological investigations of late years are those
that have been carried on in New Zealand by CocKAyNE, which have
recently brought him the degree of Ph.D., causa honoris, from the University of
Munich. His most recent publication is an ecological presentation of the flora
of the famous “‘southern islands.”’3 CocKAYyNE had the privileges of a govern-
ment steamer, and was able to visit Auckland, Campbell, Antipodes, and Bounty
Islands. Although it was necessary to make his studies with ‘‘feverish haste,”
his was the first ecological trip ever made to these islands, and the first botanical
trip in winter. These islands lie between lat. 47° 43’ and 54° 44’ S., yet in spite
of the high latitude, the larger islands are clothed with a luxuriant rain-forest.
Auckland Island (50° 45’) rarely has snow for over three days at a time on the
lowlands. Although the rainfall is not excessive, almost every day is rainy
or at least cloudy and the evaporation is slight. These features, together with
the mild winters, make an ideal rain-forest (hygrophytic) climate. However
the winds are constant and violent, a feature which commonly accompanies a
xerophytic climate. The resultant vegetation reflects the peculiar climate to a
most extraordinary degree. The forest trees are short (not exceeding five meters
in height), and present the gnarled aspect so familiar in mountain regions and
near the sea. The lateral branches grow to such length as to make the forest a
true jungle. Within the forest, where the air and the forest floor are always
moist, the wind does not enter; hence the vegetation is amazingly luxuriant,
reminding one of the tropical forests; mosses, liverworts, and filmy ferns grow in
wild profusion, making a soft carpet, while the trunks of the trees are covered
with epiphytes. Even a tree fern (Hemitelia) was discovered, extending by some

escribed; Campbell Island has merely a scrub; while Macquarie Island is without
either forest or scrub. All of these islands have essentially similar climates, but
the smaller islands are too exposed for trees, having instead the tussock formation.
The dominant tree of these hygrophytic rata forests is the myrtaceous M: etrosideros
lucida. The much more local forests, in which Olearia Lyallii dominates, have
4 somewhat different aspect. An interesting formation, found also in the Falk-
es

3 Cockayne, L., A botanical excursion during midwinter to the southern islands
of New Sei Trans. N. Z. Inst. 36:225-333. 1904.

156 BOTANICAL GAZETTE [FEBRUARY
land Islands and Fuegia, is known as the tussock meadow; the physiognomy here
is chiefly determined by grasses or sedges, which grow on mounds of their own
formation, such as are sometimes seen in low sedgy pastures in North America.
The tussock formation is scanty on Auckland Island, more extensive on Campbell
Island, and almost excludes other formations on the smaller islands. It seems
therefore that the forest and tussock form a beautiful instance of vicarious forma.
tions. Among the character tussock plants are Danthonia bromoides and a species
of Poa, the former especially in subalpine tussocks. On Antipodes Island, where
the wind is very severe, the tussock often grades off into a xerophytic heath-like
formation, where the grasses are less dominant, being replaced by stunted Cop-
rosma bushes, lycopods, and lichens. Some space is devoted to a discussion of
the destructive influence of animals, which is especially marked in the sheep
pastures of Campbell Island. Cockayne is a firm believer in the presence of a
former land connection between New Zealand and these islands, because it is
impossible otherwise to account for the presence of entire New Zealand formations.
Certain species might be transported hundreds of miles, but scarcely the con-
stituents of an entire formation.

In an earlier paper, CocKAYNE has given an admirable picture of the vegeta-
tion of Chatham Island, 450 miles to the east of New Zealand. The climate
resembles that of the southern islands, but is of course much milder. As woul
be expected, the climatic formation of the island is the hygrophytic forest, which
presents a somewhat xerophytic physiognomy, though much less so than on
Auckland Island. The trees commonly come close to the sea. Palms are fre-
quent, and tree ferns are among the dominant elements, especially species Of
Dicksonia and Cyathea. Perhaps the dominant tree is the celastrineous Coryno-
carpus. COCKAYNE made a genetic study of the dune and bog floras. On the
beach the dominant plant was once the handsome Myosotidium nobile, the only

ndemic genus of the island, now almost extinct in its native state. The dunes,

4 CocKayng, L., A short account of the plant-covering of Chatham Island. Trans.
N. Z. Inst. 34: 243-325. 1902.

1905] CURRENT LITERATURE 157

of the characteristic bog plants are just as characteristic of dry as of wet xerophytic
situations; this is true of Olearia, and notably so of Phormium tenax, the

aland flax. This plant, once so abundant, is being exterminated by fires
and grazing animals, and is now almost confined to the seemingly opposite habitats
rocks and lake margins. Fires have wrought great destruction to the native flora
and as a result there are vast areas dominated by the typical fireweed, Pferis
esculenta; the exotic Rubus fruticosus also covers large areas.

DENDYS has also written concerning the flora of Chatham Island. He thinks
that the absence of many characteristic New Zealand types is due to the absence
of many typical New Zealand habitats, such as alpine and other xerophytic situa-
tions, as well as to the long time that has elapsed since the islands were connected
with New Zealand; even when there was a land connection, it is likely that much
of it was a desert, and hence a barrier to many forms.

An earlier paper by Cockayne® deals with the vegetation of the New Zealand
mainland, in the neighborhood of the Waimakariri River. In this paper there
are brought out the characteristic features of the eastern and western climatic
regions, and their various edaphic formations.—H. C. CowLrs

THE PREMATURE decease of Nicoras ALBOFF has been much mourned by
plant geographers. His studies in Fuegia have been edited and issued by EUGENE

UTRAN,’ who has published as a preface an appreciative biographical sketch,
together with a bibliography and an excellent portrait. In the historical sum-
mary, especial praise is given to the work of Hooker more than half a century
ago. There are two dominant formations, the forest and the moor. ‘The forests
are extremely dense and luxuriant, and bear witness to the humidity and uniform-
ity of the climate. The forest floor has a wealth of bryophytes and filmy ferns.
Extreme floristic poverty characterizes these forests, only two tree species being
present: Fagus antarctica and F. betuloides. Moors occur where the forest can-
not exist, either through exposure or soil moisture. Kerguelen Island, 140°
distant, represents, from a floristic standpoint, the farthest point reached by this
Fuegian flora. The balsam bogs or dry moors, so characteristic of the Falkland
Islands, are also found in Fuegia. Fifty-three per cent. of the species are endemic.
The most interesting elements in the flora are the neozelandian and boreal.

OFF agrees with most authors in holding to an ancient antarctic continent or
archipelago, as accounting for the similar floras throughout antarctic regions.
The long-known and most perplexingly large boreal element has been a stumbling

’ Denby, A., The Chatham Islands; a ole in biology. Manchester Memoirs
46: 1-29. r902 See Bot. Cent ay 9. 19

Cockayne, L., A eels of the plant ineeste of the Waimakariri River Basin,
Considered chiefly from an oecological point of view. Trans. N. Z. Inst. 32: 95-136.
goo

si Apo, Nicos, Essai de flore raisonnée de la Terre de Feu. Anales del
Museo de La Plata. Seccién Botanica. I. pp: vit+85 +xxili. With portrait. La
=a

158 BOTANICAL GAZETTE [FEBRUARY

block to students of floristics. Apart from species that are more or less cosmo-
politan, the author holds that alternating glacial climates, advocated by CRott,
afford the best explanation for the similarity of austral and boreal floras. Since
glacialists generally reject CROLL’s hypothesis, it is seen how slender a support
Avzorr’s theory has. ‘To the reviewer, it seems that here, if anywhere, we shall
be forced to consider, at least as a possibility, Brrqurr’s polytopic (polygenetic)
theory.

In connection with the work of ALBorr, brief mention may be made of
Dusén’s admirable studies in the same region. Dusfn’s earlier work has been
noted in these pages.? One of his papers? was for the most part a floristic and
taxonomic account of the flora. A later paper'® presented a more detailed eco-
logical account of the vegetation, along the line of the short earlier articles. His
latest paper™® gives an excellent account of the ecological and floristic features
of western Patagonia. There are three great regions: the evergreen forest, the
deciduous forest, and the steppe. The evergreen forest resembles that described
by Atporr, and for New Zealand by Cockayne. Bryophytes, especially liver-
worts, form a forest floor, sometimes five or six feet thick. Epiphytes reach a
high degree of development. One or more species of beech everywhere page
although Drimys Winteri and Libocedrus tetragona are often abundant,
aig especially in moory soil. Of extraordinary interest is the deciduous eo

e only one yet found in the southern temperate zone. The dominant tree is
F agus (Nothofagus) antarctica, the very tree that dominates so much of the ever-
green forest. Very few cases are known, at least on the lowlands, where one tre¢

species dominates in two radically different climatic forest types. The whole
forest aspect differs, being more parklike, and without the luxuriant undergrowth
of the evergreen forests. The rich bryophyte carpet of the latter is wholly miss-
ing. The steppes call for no special mention. Through the work of ALBOFF
and Dus&n, it is clearly to be seen that the Fuegian and Patagonian vegetation
is of almost equal interest to the ecologist as the vegetation of New Zealand.—
H. C. Cow1es.

ITEMS OF TAXONOMIC INTEREST are as follows: J. Rick (Ann. Mycologici
2:407. 1904) has described Psendohydnum as a new genus of fleshy fungi from
South America—A new part of Komarov’s Flora Manshuriae (Acta Hort.

8 See Bor. Gaz. 24:135. 1897. Also Engl. Bot. Jahrb. 24:179-196. 1898:
USEN, P., Die Gefasspflanzen der Magellanslander, nebst einem Beitrage

zur nea der Ostkiiste von Patagonien. Svenska Expeditionen till Magellanslandern®-
T9000. See Bot. Cent. 85:47-49. 1901

to DusEN, P., Die A UetaegeTaRe der Magellanslander, nebst einem Beitrag 2UT
Oekologie der nas iisaeliog Vegetation. Svenska Expeditionen till Magellans-
landerna 3:351-523. 1903. See ne Cent. 96: 468-469.-1904. Also Engl. Bot.
Jahrb. 33:litt.28-29. 1903.

«« Dusén, P., The vegetation of western Patagonia. Reports of the Princeto®
University Expeditions to Patagonia, 1896-1899. Part I. Princeton. 1903-

1905] CURRENT LITERATURE 159

Petrop. 22:453-787. pls. 1-17. 1904) begins with Rosaceae and ends with
Balsaminaceae, bringing the serial numbers of species to 1058.—P. A. RYDBERG
(Bull. Torr. Bot. Club 31:555-575. 1904), in his 12th age entitled “Studies on
the Rocky mountain flora,” has described new species of Draba, Smelowskia,
Sophia (2), Arabis (2), Erysimum (2), Opulaster (2), lane. items,
Rosa (2), Astragalus (2), Homalobus (3), Ceanothus, Sphaeralcea (2), T
(2), Acrolasia (2), Epilobium (4), Gayophytum, Anogra (2), saa ©,
aura, Suida, Aletes, Phellopterus, and Pseudocymopterus.— . GLEAS
10 Nat. 5:214. 1904) has published a new Helianthus nee THinois. a
HM (Hedwigia 44: 1- 13. pl. rt. 1904) has published Trichophyma (Myrian-
a and Stictoclypeolum (Mollisiaceae) as new South American genera of fungi
from the Ule collection.—H. Curist (Bull. Herb. Boiss. II. 4:10 9-1104. 1904)
has described new species of Costa Rican ferns under Asplenium (2), Lomaria
(3), Adiantum (3), Gymnogramme (2), Saccoloma, and Polypodium (3).—G.
ERONYMUs (Bot. Jahrb. 34:417-560. 1904) has published Lehmann’s pteri-
dophytes from Guatemala, Colombia, and Ecuador, enumerating 315 numbers,
and describing new species in Trichomanes (2), Hymenophyllum (2), Loxsom-
opsis, Cyathea, Nephrodium (6), prerani, Polystichum, Diplazium (2), Blech-
num, Gymnogramme (5), Adiantum, Polypodium (12), and Elaphoglossum (7).—
E. L. GREENE (Leaflets 1: 65-81. 1904) has described 5 new species of Ceano-
thus; has separated from Gentiana the genus Pneumonanthe, to include the
“closed gentians” and their allies, and has transferred to it nearly 30 species;
and has described from middle California new species under Lupinus (4), Lotus,
Sidalcea (2), Silene, Aquilegia, Delphinium, Bistorta, Eriogonum, Swertia,
Castilleia (2), Pentstemon, Apocynum, Cryptanthe, Galium, and Chrysothamnus
(2).—R. M. Harper (Torreya 4:161-164. 1904) has recognized a new species
of Ludwigia (L. maritima) among the forms commonly referred to L. virgata.—
C.F. raed ge 172) has published a new species of Euphorbia from
the Bahamas.—W. H. BLANCHARD (Rhodora 6: sen es 1904) has described
eines Rubus Gilackberey) from New England.— House (idem. 226 pl. 59-)
has described a new Viola from New England. a - ed

WIELAND” has secured transverse seed-sections of one of the Bennettitales
showing tissue filling the archegonium, which he interprets as the proembryo.
If he is correct, this is an interesting confirmation of the current morphological
view that the Ginkgo type of proembryo is the most primitive among living gymno-
sperms. The Previously known mature seeds of Bennettitales are singular among
8yMnosperms in the entire absence of endosperm; and now even in this reported
Proembryonic stage WIELAND finds no trace of endosperm.—J. M. C
SS

8. * WIELAND, G. R., The proembryo of the Bennettiteae. Am. Jour. Sci. IV.
19:445-447. pl. 20. 1904.

NEWS.

Dr. Oscar BrEFELD, professor of botany at Breslau, has retired from acti
service.

Dr. JOHANN GoROSCHANKIN, professor of botany in the Imperial University
and director of the Botanical Gardens at Moscow, died recently at the age 0
sixty years. ;

ProFEssorR GEORGE J. Perrce has returned from working at the Zodlogical
Station at Naples and resumed his duties at Leland Stanford University at the
beginning of the year. .

THE ForRESTRY QUARTERLY, whose chief editor is Professor B. E. FERNOW, a4
has increased its subscription price to $2, as the amount of material for publica:
tion demands an issue of three to four hundred pages a year.

Henri T. A. Hus, M.S., a graduate of the University of California and
student with Huco pr Vrtss, has been appointed to take charge of the herbarium =
work at the Missouri Botanic Gardens, St. Louis. Mr. Hus has given much -~
attention to fasciation and to mutations in plants. ‘

4

i

Py
ee

iS

Notice.—Owing to the increasing pressure upon our space, and in pares
to numerous contributors, the Editors of the BoTANICAL GAZETTE are constrained
to announce that with the beginning of volume XL (July 1905) no paper exceeding
thirty pages in length (about 11,000 words) can be accepted for publication,
unless the author is willing to pay for additional pages; in which case the
of the number will be correspondingly increased. Longer papers that —
already been received and accepted will be published in due time.

Attention is also directed to the more liberal terms for separates an
in the advertising pages.

nounced

to the rapidly accumulating collections that are making Washington a great eee
for taxonomic work. Captain Sarru will retain for the present the custody ee
library and the greater part of his collection, but some 25,000 specimens
already been sent to Washington.

BUFFALO
LITHIA WATER

No Remedy of Ordinary Merit Could Ever
Have Received Indorsations from
Men Like These.

Sam uel O. L. Potter, A. M., M. D., M.R.C.P., Z ondon,
Peatror of the Principles "and Practice of "Medicine and Clinical
ayes ime in the wen ro Physicians and Surgeons, San Francisco,

q Sepa r. Wm. H. Dru ond, Professor Medical Jurisprudence,
s Disease Bistop’ Ss srtisca Montreal, Canada. :
Cyrus Edson, A. M., M. D., G fi ith Commissioner Nez
York City and GH ke, President Board of Phar Sigrid New York
City, Biaaeaie. Physician Corporation Council, é
John V. Shoediunee M.D., LL. D Profess
mean and Ther Medico-Chirurgical C College, Pudladelphia,

Dr. eee eorge e Ben. Johnston, Richmond, Va., Ex-President
Southern Surgical and Gyne ological Association, Ex-Presi com
Medical Society of Va., aa Professor r of Gynecology and Abdomina
Surgery, "Medical College of Va.

Dr. A. Gabrief Pouchet, Proj seins = cient omits shred
Materia Medica of the Faculty of Me
Dr. J. T. LeBlanchard, 707 Macun Sen See N.,V. U.
Professor Clinical Medicine
a et Asante Ho west Pare Post Ceaduaie pe Medical School.
. Louis C. Horn, M. D., Ph. D., hi iopal Diseases of Chile
dren and Dermatology, pelos i) oe sity.

wer: J. Allison Hodges, nd Professor Nervous and
ental Diseases, University De iivueee of ‘Medicine, Richmond, V be!

” lia, Robert Bartholow, M

Medica and srs 7; herapeutics, Je ee Medical College, Phila.
New Artes City, incase 8 Professor Diseases of
and Sur, SeOns, and in en Sims :
;D.. Ex-President American ‘ i
Sinaoce, Lae Clinte ical eght ins

L. D., Professor Materia ake

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March, 1905 |

Editors |
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COULTER and CHARLES R .
LES R. BARNES ©

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" a

The Botanical Gazette

A Montbly Fournal Embracing all Departments of Botanical Science

Edited by JoHN M. CouLrer and CuHar.es R. BARNES, with the asistanc of other members of the
botanical staff of the University of Chica

Vo. U Socux. No. 3° Issued March 20, 1905.

CONTENTS

SAM Side Set AND EMBRYO OF TORREYA TAXIFOLIA. ContrisvTIons FROM
oe, OTANICAL PARTIES: LXIX Signs PLATES A, I, I, uf): ane am
, Cone oe cea W. J. G. Land 161
T FETE PRINCIPLES OF PHYTOGEOGRAPHIC NOMENCLATURE. Pehr Olsson-Seffer 179

? FORESTS OF THE FLATHEAD each: MONTANA. CONTRIBUTIONS FROM THE

Hvuit Botanican Pye ade tages T (wir MAP AND TWENTY-THREE ps comin ds
Harry N. Whitjord 194
RIEFER i
TERMINOLOGY OF THE SPORE-STRUCTURES IN THE UREDINALES. J.C. Arthur - o~ aig

; NT oo ee

. - - - - - - - - 93g

eer Sec PLANT ORGANIZATION.
MINOR NOTICES - - - - - - - - - - 225
_ NOTES FOR STUDENTS | - - - - - - : - - 226
N EWS 2 - - - - - = - - - - - 240
Communications for the Editors should be addressed to them at the University of Chicago, Chicago, Ill.
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Copyright, 190 Seah University of Chicago.

| q4

VOLUME XXXIX NUMBER 3

BOTANICAL -G4ze ee

MARCH, 1905
GAMETOPHYTES AND EMBRYO OF TORREYA TAXI-
FOLIA. —

CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY.

JoHN M. Covutrer and W. J. G. LAND.
(WITH PLATES A, I, II, 111)

Torreya taxijolia Arnott! occurs in a narrow belt along the eastern
side of the Apalachicola River, extending from the southern boundary
of Georgia for about thirty miles southward (1). In April 1904 this
region was visited by H. C. Cowes of this laboratory, among whose
_ Notes the following are of interest in this connection:

My visit was to the northernmost colony, west of Chattahoochee village, and
close to the Georgia line. The distribution lines on CHAPMAN’s map (1) would
lead one to suppose that the tree is xerophytic and frequents the steep and dry
castern bluffs. I was much surprised to find that it was confined (in the Chatta-
hoochee station at least) to the extremely mesophytic slopes of ravines, growing
exclusively in the shade of trees, and in places that are continually moist, prefer-
ably on slopes facing north. The northern and southern known limits of the
tree are only about thirty miles apart, and the east-west range is much less.

urthermore, on account of the great economic value of the wood, and the
familiarity of the tree to all the inhabitants of the region, the likelihood of finding
other areas is very slight.

It is associated with a remarkable and somewhat extensive group of northern
mesophytic plants, and the conclusion is irresistible that Torreya is a northern
plant of the most pronounced mesophytic tendencies, and to be associated with

* Unfortunately, Torreya was used at least three times as a generic name, in as
many families, before 1838, the date of the publication of Torreya Arnott. Hence
ARNOTT’s genus has been replaced by T'umion Raf., and our species becomes Tumion
taxifolia Greene. In the present paper, however, the more familiar name is used for

onvenience.
161

162 : BOTANICAL GAZETTE [MARCH

such forms as the beech-maple-hemlock forms of our northern woods, our most
mesophytic type of association. Probably it never becomes a large tree, although
farmers always cut the trees as soon as they become at all usable. Rarely were
any found over 30°™ in diameter or more than 9 to 12™ high. It has remark-
able capacity for vegetative reproduction, almost equaling the redwood in this
respect. Many suckers issue from cut stumps, and even from fallen trunks;
even rotten stumps show vigorous suckers, and it seems to be as tenacious of life
as the poplar. Staminate trees appear to be the more numerous; however,
when in blossom they are far more conspicuous than are the pistillate trees, and
the conclusion as to proportion may not hold.

Cow Les secured collections of staminate and ovulate material
April 4 and 5, and arranged for subsequent collections to be sent to
the laboratory. These collections were made at intervals of approxi-
mately’ two weeks up to October 21. The strobilus-bearing twigs
were packed in damp cotton in tightly closed tin buckets and reached
the laboratory in good condition in about five days. The length of
the intervals between collections and between collecting and killing
has prevented so close a series as is desirable at certain stages, but
the gymnosperm periods are so long that a fair outline of morpho-
logical events was secured.

SPERMATOGENESIS.

Srropii.—The staminate strobili occur in the axils of leaves of
young shoots. Their appearance in October, on a shoot of the
same season, is shown in fig. r. At this time they are small and
ovoid, solitary in the axil of each leaf, and by a curving of the short
peduncles are displayed along the under surface of the shoot. The
strobilus consists of a series of closely overlapping sterile bracts, 2
four vertical rows, completely enveloping the tip of the axis bearing
numerous stamens. It is thus distinctly sterile below and fertile
above, the sporophylls continuing the spiral succession developed
by the bracts (figs. 6 and 7).

The young strobili were first observed in July, and at that time
no primordia of sporophylls were evident; but in August these wer’
beginning to appear. Fig. 7 is from a longitudinal section throug
a strobilus at this period (August 12), showing the overlapping sterile
bracts and the beginning of stamineal primordia. A remarkable
development of the pith region of the axis below the staminate Por

1905] COULTER & LAND—TORREVA TAXIFOLIA 163

tion of the strobilus, with its investing vascular cylinder, may be
observed, apparently an important storage region for the strobilus.

STAMENS.—At the stage of the stamineal primordia observed
August 12, there is no evident differentiation of tissues (fig. 8), but
early in September the young sporangia become distinct, the sporo-
genous tissue being represented in each case by a single primary
sporogenous cell. At this stage the further development of the
young sporangia differs. Seven sporangia are seen in the primary
sporogenous cell stage, three adaxial and four abaxial (jig. ro),
radially arranged about the central axis as are the five or more spor-
angia in the allied Taxus. The four abaxial sporangia develop in
the usual way, and become the pendent abaxial sporangia character-
istic of Torreya. The three adaxial sporangia, however, do not
develop further, the primary sporogenous cell not dividing, and its
nucleus showing signs of disorganization (fig. 9). The disorganiza-
tion continues until it involves all the cells separating the three adaxial
sporangia, which are thus replaced by a single large flattened cavity,
which becomes a resin cavity. Intermediate stages were found in
which the three sporogenous regions were still distinct (fig. 10). In
jig. 11 (October 21) is shown a median longitudinal section through
a stamen, in which the large resin cavity is seen above and the nor-
mally developing sporogenous tissue below. As a result, the mature
stamen of Torreya is characterized by four abaxial sporangia and a
large adaxial resin cavity.

In one instance the two lateral sporangia were smaller than the
middle ones (fig. ro), suggesting a tendency to still greater reduction
in the number of sporangia. Accordingly Pinus Laricio was exam-
ined, and two resin cavities were found related to the two sporangia
exactly as the two lateral sporangia of T ‘orreya are to the middle
ones; and in early stages the tissue on the sites of these two resin cavi-
ties of Pinus resembles that of sporangia. It is interesting to note
that in T. californica Miss ROBERTSON (8) found occasional stamens
bearing six or seven mature sporangia, indicating that in these cases
some or all of the usually abortive sporangia reached maturity.

There is evident a tendency to reduce the number of functioning
sporangia by abortion, a reduction that has proceeded farther in
Pinus than in Torreya; and in the latter farther than in Taxus. That

164 BOTANICAL GAZETTE [MARCH

the cavities formed by the breaking down of young sporogenous and
adjacent tissue should become resin cavities in conifers is to be
expected.

It was only in the abortive sporangia of Torreya that stages were
- found indicating the history of the sporangium up to the primary
sporogenous cell stage. For example, from jig. g it seems evident
that there was a single hypodermal archesporial cell, which at this
stage had given rise to two layers of wall cells and a single primary
sporogenous cell. In the mature sporangium there are three or four
wall layers. The epidermis contains numerous stomata, and its
cells become prominent and have thickened walls. The sporangia
are bluntly four-angled from pressure, and the epidermal cells at
the angles are much larger than elsewhere.

MALE GAMETOPHYTE.—We secured no stages between the very
young sporangium and the shedding of the pollen, which occurs late
in March and early in April. Miss Roperrson (8) has observed
the mother-cell stage and the formation of tetrads in T. californica.

In germination the microspore of 7. taxifolia cuts off no prothallial
cell, a feature common to all the Coniferales except Podocarpeae
and Abieteae. After the first division the generative and tube cells
are distinct and separated by a very delicate wall or membrane.
The tube nucleus is sometimes spherical, but more often amoeboid
(fig. 12).

Early in April the binucleate pollen grains, rich in starch, wert
found resting on the nucellus (fig. 19); and at the end of June micro-
pyles were found full of pollen grains, most of which had sent out no
tubes, but were still full of starch, nucleate, and apparently alive.

As fertilization occurs about the middle of August, active pollen
tubes occupy the sterile cap of the nucellus about four months, and

their behavior is exceedingly variable. They may advance towards
the embryo sac rapidly, reaching it at a very early stage, a5 early as
June 21, when the endosperm consists of only sixteen to sixty-four
parietally placed free nuclei (figs. 14 and 15); or they may advance
slowly, being found at all stages of progress at the same date. They
may advance so directly that the tube resembles a straight cleft
through the nucellar tissue to the sac, or they may pursue a remar*
ably devious course. In one case a tube passed directly half-way

See sae, eee

PR te ee

Te a Oe ey eT EE TS ce

PRS ene

oh al

ie i a a a la col a ae aa a a

1925] ' COULTER & LAND—TORREYA TAXIFOLIA 165

down the nucellar cap, then advanced spirally downward and out-
ward until it reached the peripheral cells of the nucellus, and after
destroying several of these it turned abruptly inward, penetrated the
nucellus at the level of the archegonium, crossed the top of the endo-
sperm, and discharged its contents into the archegonium on the farther
side. In another instance, such a pollen tube had entered the inner
integument and destroyed some of its cells before turning back into
the nucellus. It is very common for the pollen tube to push into the
sac at its free nuclear stage, making a deep invagination, often to the
middle; from this pocket the tube turns back again into the nucellus.
It is of interest to note in this connection that the usually solitary
archegonium is never centrally placed in the endosperm, and the
pollen tube enters the sac to one side of it (fig. 16).

In all the wanderings of the pollen tube, the body cell, stalk
nucleus, and tube nucleus are conspicuous. The division of the
generative cell was not observed, but that it occurs very early in
the development of the tube is evident. The body cell is relatively
very large, with a conspicuous nucleus and investing cytoplasm, and
was always found consorting closely with the stalk and tube nuclei
(figs. 13 and 14).

e division of the body cell just before fertilization results in
unequal male cells (fig. 23), almost exactly resembling those of Taxus.
It is not a case of the extrusion of one nucleus from the common
cytoplasm, as observed by CoKER (2) in Podocarpus, but the cyto-
plasm is unequally divided, so that there are two distinct and naked
male cells very different in size. The whole cavity of the pollen
tube surrounding the cells and nuclei is rich in starch and other food
materials (fig. 13).

A consideration of the time involved in these various events shows
that a period of about fifteen months elapses between the first appear-
ance of the microsporangiate strobilus and fertilization, divided as
follows: June, first appearance of strobilus; August, first appearance
of stamineal primordia; September, distinct differentiation of spor-
angia; April, shedding of spores; August, fertilization. The stro-
bilus was first observed in July, but in such a condition that it must
have been evident in June, if not earlier. In comparing this schedule
with that given by Miss ROBERTSON (8) for T. californica, it is evident

166 BOTANICAL GAZETTE [MARCH

that the sporangium in both species passes the winter in the mother-
cell stage, but that the subsequent stages appear earlier in T. tawijolia
growing in its natural habitat than in 7. calijornica growing in Eng-
land. For example, in the latter case the reduction division takes
place about the time that 7’. taxifolia sheds its pollen, and pollination
in T. californica does not seem to occur until late in May or early in
June. |
OOGENESIS.

STROBILI.—The ovulate strobili are borne in the axils of the lower
leaves of short young shoots (fig. 2). They usually occur in pairs
upon a very short axillary branch, usually one pair, frequently two
pairs, very rarely three pairs appearing upon a single shoot. A
cluster of two to six strobili, therefore, appears near the base of the
strobilus-bearing shoot, the upper pairs never maturing, usually
one and sometimes both strobili of the lowest pair producing the
large plum-like seeds (fig. 3). The strobilus is a very simple one,
consisting of four enveloping bracts and a single terminal ovule with
two integuments (figs. 17 and 78), the outer one often called an arillus
because it ripens fleshy. The whole structure resembles a simple
ovulate flower with a perianth of four bracts, which perhaps deserve
to be called a perianth as much as the so-called perianth of Gnetales;
but it is none the less evident that they are the sterile bracts of a much
reduced strobilus.

The strobili were first seen July 26, at which time the growing
point, enclosed by the bracts, was composed of entirely homogeneous
tissue, showing no differentiation as an ovule (fig. 17). No subse
quent stages were found until April 7 (fig. 18), when the mother-cells
were in synapsis (fig. 20); and it seems probable that the winter was
passed in the mother-cell stage. At this time the integuments 47
entirely free from the nucellus, appearing to arise from the base of
the ovule. Soon, however, extensive intercalary growth below the
mother-cell becomes evident, the ovule being greatly elongated and
broadened below, the original free nucellus with its integumen
forming only the tip. This growth continues throughout the season
of fertilization and the following one; and Oxiver (6), who has
described this intercalary growth in T. nucijera, estimates that ™
that species in the maturing seed the original nucellus with its free

1905] COULTER & LAND—TORREYA TAXIFOLIA 167

integuments represents only one-twentieth of the entire length of the
seed. -The characteristic structures of the integuments are con-
tinued as two distinct peripheral layers of this large mass of additional
tissue, and the tissue within these layers may be taken to represent
in a similar way the downward extension of the nucellus. It is this
additional nucellar tissue that the endosperm chiefly invades, giving
rise to the phenomenon of “rumination,” to be described later.
OLIVER (7) has also described fully the course of the vascular strands
in the ovule of certain species of Torreya, a description which seems
to serve as well for T. taxijolia.

FEMALE GAMETOPHYTE.—The mother-cell is solitary and with
no differentiation of a nutritive mechanism about it, such as appears
in connection with the “spongy tissue” of Pinus, and in all the
Pinaceae investigated. In Torreya it is directly in contact with the
cells that are resorbed, without any intervening digestive layer (jig.
20). Since this is true also of Podocarpus (2) and Taxus (10), it
suggests the possibility that Taxaceae in general may be character-
ized by the absence of a special digestive layer about the mother-cell.

The reduction division was not seen, but a more or less complete
tetrad is formed, as observed by Miss RoBERTSON (8) also in T.
californica.

The germination of the megaspore begins ick the usual free
nuclear division, the nuclei being in the parictal position when only
sixteen to thirty-two in number (figs. 14 and 15). The interior of
the sac contains cytoplasmic material, much less dense than the
cytoplasm of the parietal layer, and also some reserve food. In this
early few-nucleate stage of the endosperm there is always an appear-
ance suggesting that the sac has sent a beak-like projection, contain-
ing a nucleus, upwards into the nucellar tissue (fig. 15). After a
careful comparison of the position of this apparent projection in
reference to the surrounding parts with that of the megaspore, it
seems that the “projection” is the original site of the megaspore,
and that the appearance of a projection is due to the fact that the sac
has encroached almost exclusively upon the chalazal tissue. This
conspicuous beak, containing one of the parietal nuclei, often appears
close to the tip of an advancing pollen tube, and suggests a possible
explanation of the peculiar behavior ascribed to the archegonium

168. BOTANICAL GAZETTE [MARCH

initial of Tumboa. Inasmuch as the archegonium initial of Torreya
often occupies this beak, the suggestion becomes still more pertinent
(fig. 21).

The formation of walls in the endosperm was not observed before
July 1, and in several instances they did not appear for a month later.
When wall-formation began, repeated countings showed 256 free
nuclei, which seems to be a very common limit of free nuclear division
among gymnosperms.

Since fertilization was observed August 12, it is evident that the
archegonium is developed very early in the history of the gameto-
phyte. In fact, as soon as the very small sac is filled with extremely
delicate tissue the archegonium initial becomes evident. It does not
seem possible for archegonia to appear any earlier, for the initials
are organized as soon as the free nuclear stage has passed. In
Torreya, therefore, nearly all of the endosperm, which becomes an
extensive tissue, develops after fertilization.

The single archegonium initial is always to one side of the central
axis (fig. 16), often occupying the “beak” referred to above, and so
projecting above the endosperm (jig. 2z). A neck cell is cut off
and divides anticlinally, forming a two-celled neck (fig. 22), the
usual limit of neck-formation among gymnosperms. In fact, it is
only among Podocarpeae and Abieteae that a more extensive neck
is usually formed, consisting of more than one tier of cells, unless the
somewhat anomalous neck of Ephedra be included. The fact that
there is variation in the number of neck cells in the same form (two to
twenty-five in Podocarpus), and that as a rule necks are destroyed
as soon as formed by the growth of the central cell and pollen tube
(fig. 23), suggests that their extent depends somewhat upon the
approach of the pollen tube, which usually checks neck formation
early, but sometimes permits it to become more extensive. In T.
californica Miss ROBERTSON (g) has found that the archegonia are
usually three in number, ranging from two to five, and that the necks
consist of four or six cells.

The central cell enlarges rapidly, no jacket-layer being evident
until after fertilization, and even then it is weakly organized (fig- 25).
The nucleus is spherical and lies near but not against the neck cells,
more nearly resembling an egg nucleus than in any gymnosper™

1905] COULTER & LAND—TORREYA TAXIFOLIA 169

we have observed (jig. 22). We could not detect the formation of a
ventral canal cell or nucleus, or anything that stood for such a struct-
ure at later stages. A ventral nucleus was expected, for a distinct
ventral canal cell among Coniferales seems to be restricted to the
Abieteae and does not always occur in them, and there seemed to be
no excuse in our preparations for missing it. We are fully aware
that all previous negative evidence as to the occurrence of at least a
ventral nucleus in archegonium-forming gymnosperms has proved
to be deceptive, but a study of the behavior of the central cell of
Torreya, from the formation of the neck cell to fertilization, not only
failed to show any indication of division but suggested that it may
not occur. In 7. californica a spindle seen twice in the central cell
was interpreted by Miss ROBERTSON (g) as representing the ‘cutting
off” of a ventral nucleus, but no other traces of it could be found.

In the single case in which two archegonia were observed, they
were at opposite sides of the gametophyte, with the tip of a pollen
tube between them.

At the time of fertilization the gametophyte contains 400-800
cells, with extremely thin walls and scanty cytoplasm (fig. 23). The
only differentiation observable is the abundant accumulation of
reserve food in the peripheral cells of the antipodal region. The
whole mass of endosperm at this period usually measures 20 by 30#;
while in the mature seed the endosperm mass is ordinarily about
20o™™ long by 14™™ at its widest part, and all of it surcharged with
starch and other food materials. The food material is particularly
conspicuous in a broad central band extending from the advancing
tip of the embryo and widening to the antipodal end of the sac. Fig.
5 shows the longitudinal extent of the band, and fig. 4 its cross-section.
The peculiar behavior of the endosperm after fertilization will be
considered under the discussion of the maturing of the seed.

FERTILIZATION. :
The forcible discharge of the contents of the tube may be inferred
from the vacuole-like appearance in the center of the egg, produced
by the inrush (figs. 24 and 25). The male nucleus in its cytoplasmic
sheath passes through the cytoplasm of the egg and comes in contact
with the egg nucleus. The male cytoplasm becomes closely appressed

170 BOTANICAL GAZETTE [MARCH

to the surface of the female nucleus, slips from its own nucleus, and
was observed extending over fully two-thirds of the female nucleus
(fig. 24). This behavior of the male cytoplasm has been observed
by CoKER (3) in Taxodium and by Miss RoBERTSON (g) in Torreya
californica. The male cytoplasm of Torreya taxijolia 1s sharply
differentiated by staining from the cytoplasm of the egg, and undoubt-
edly completely invests the fusion nucleus. The appearance of a
similar mass of cytoplasm investing the free nuclei of the first division
(fig. 25), and continued in the second division in connection with
wall-formation (fig. 26) suggests the possibility that the male cyto-
plasm may remain differentiated through more than one cell genera-
tion. Near the neck end of the archegonium nuclei are evident,
which seem to be the stalk and tube nuclei and the other male nucleus
with its investing cytoplasm (fig. 24).

EMBRYO.

Soon after fertilization the first division of the egg nucleus was
observed (fig. 25), and almost immediately the second division fol-
lows, giving rise to four large free nuclei almost filling the egg (7g:
26), one nucleus in the base of the egg, the other three in a plane
above. At this time wall-formation occurs, the cytoplasmic radiations
which precede it being very evident (fig. 26). Two weeks later the
egg is completely filled with a proembryo consisting of twelve to
eighteen cells (fig. 27). This complete filling of the egg by the pro-
embryo is remarkable among Coniferales, having recently been
observed also by Lawson (4) in Sequoia, but as yet not recorded
in other genera. In Torreya, at least, this fact seems to be related
tothe relatively small size of the egg, the very large nuclei, and the
early appearance of walls.

The cells of the proembryo at this early stage are distinctly in three
tiers; that nearest the neck of the archegonium comprising five or SIX
cells and forming the primary suspensor tier; the middle tier, Co™
prising five or six cells and forming the secondary suspensor tier; and
the lowest consisting of a single cell which ultimately contributes to
suspensor-formation and forms the embryo. The inequality in the
number of cells entering into the tiers seems to be characteristic of
Taxaceae. In Podocarpus CoKER (2) found the three tiers made up

1905] COULTER & LAND—TORREYA TAXIFOLIA ry A

of eleven cells in each of the upper two and one in the lowest; and a
similar but less striking inequality is to be observed in Taxus. Occa-
sionally in Torreya other divisions may occur, giving rise to approxi-
mately four tiers and a proembryo of about eighteen cells; but no
other division occurs until the following spring, the winter condition
being a proembryo of three or four tiers of cells as described above.

In the following season the suspensor develops and the embryo
is formed, along with the characteristic “‘rumination” of the endo-
sperm and the development of the testa. The first indication of
change from the winter proembryonic condition is the elongation of
the primary suspensor cells (fig. 29); a little later this is shared by
the secondary suspensor cells; and this is followed by the elongation
of the third tier, if four tiers are formed. In the meantime the termi-
nal cell has begun a series of rapid divisions, resulting in a cylindrical
mass of meristematic tissue, much as Lyon (s) has described in the
case of the embryo of Ginkgo. This meristematic cylinder advances
gradually into the endosperm, its basal tiers of cells successively
contributing to the suspensor elongation. Thus in the formation
of the suspensor there seems to be developed what may be called a
wave of elongation, beginning with the uppermost tier of the pro-
embryo and extending gradually downward, tier after tier, until it
includes the upper region of the meristematic cylinder formed by
the terminal cell. This same phenomenon is very evident also in
Thuja.

After the meristematic cylinder has advanced into the endosperm
and has become prominent, the growing points are organized; the
two cotyledons presently becoming beautifully crescentic in outline
and completely surrounding the stem-tip; and the root-tip being
organized deep within the meristematic cylinder.

In several instances a number of small embryos were observed
imbedded in the endosperm about the suspensor region of the normal
embryo. Our material did not permit any determination of their
origin, but they resemble the proembryo of the normal embryo, and
are developed while the latter is in its second season’s growth. After
the pollen tube has reached the archegonium, the endosperm. grows
up around it (figs. 23 and 24), so that the tube lies in a cup-shaped
depression. After fertilization, the rim of this endosperm cup con-

172 BOTANICAL GAZETTE [MARCH

tinues growth and gradually incloses the fertilized egg, in most cases
forming quite an elongated beak above the embryo. Later many
of these cells round off, forming a loose tissue, and it is among these
rounded-off cells that the feeble accessory embryos are produced.
Whether these have been developed apogamously from the endosperm
cells, or have budded from the suspensor cells can only be conjectured.
In any event, they might develop further if there was any failure in
the development of the normal embryo.

The time involved in the series described above, that is from the
first appearance of the megasporangiate strobilus to the maturity of
the seed, is about thirty months, distributed as follows: June (*);
first appearance of the strobilus; April, mother-cells in synapsis;
August, fertilization; October, proembryo of 12 to 18 cells (winter
condition) ; following season, development of embryo, “rumination”
of endosperm, and development of testa; October, fall of seed.

MATURING OF SEED.

The outer integument and its histological continuation about the
ovule develops a thick fleshy coat containing very numerous large
resin cavities, and completely inclosing the structures within (figs.
4and 5). This fleshy coat gives to the mature seed the appearance
of a plum (fig. 3), as in the seeds of Cycads and Ginkgo. Within
the broad band of resin cavities, near the inner limit of the integu-
ment, two conspicuous vascular strands occur, directly opposite one
another (fig. 4). These are the main strands of the very charactet-
istic vascular system of the ovule described by OLIVER (7)-

The inner integument early differentiates into two distinct layers,
a differentiation just as evident in Cycads and Ginkgo. The outer
layer forms the stony coat, and the transformation from soft to very
hard tissue begins after the embryo and endosperm have completed
their development. The hardening begins at the apex of the ovule,
and on account of resistance to stains appears under low power as 4
clear band (black in fig. 5). The hardening band gradually extends
downwards through the relatively very short integument, and differ-
entiating as a distinct layer in the much larger mass of tissue below
completely invests the ovule within the fleshy coat. Protoplasmic
connections between the cells of the stony coat and striations in the
‘ cell walls are unusually clear.

1905] COULTER & LAND—TORREYA TAXIFOLIA 173

The inner layer of the inner integument comprises several layers
of thin-walled cells, but beyond the integuments it is not histologically
differentiated as a layer distinct from the tissue within. Accordingly,
through the great bulk of the seed this layer may be neglected, and
the whole mass of tissue within the stony layer and outside of the
embryo sac will be spoken of as nuclear tissue or perisperm.

The behavior of the endosperm is peculiar, resulting in what is
called “‘ruminated endosperm,” a phenomenon peculiar to Torreya
among gymnosperms, and commonly illustrated by the nutmeg.
‘“Rumination” of endosperm proves to be a misnomer, for the endo-
sperm is always the successfully aggressive tissue in developing this
condition. The perisperm continues to grow throughout the matur--
ing of the seed, and the final condition results from what might be
called the struggle of two growing tissues that have been abutting upon
one another through their whole period of growth.

In ordinary seeds the endosperm invades the surrounding tissue
more or less uniformly; in the case of Cycads and Ginkgo, for exam-
ple, obliterating most of the perisperm. In Torreya, on the other
hand, the invasion by the endosperm is irregular in the extreme.
It is in the season after the proembryo has been formed that the active
invasion of the perisperm begins. The extension of the endosperm
into the tip of the nucellus above the sac proceeds in the usual way,
obliterating all of it except a few peripheral layers of cells. This uni-
form invasion seems to be due to the fact that in this apical region
(the original nucellus) the perisperm is not growing actively if at all.
Below this small region at the tip, however, the perisperm is very active
and evidently resists disintegration much more at certain points than
at others. As a consequence, the perisperm becomes eroded by the
irregularly advancing endosperm, and is left in the condition of a much
dissected coast-line (figs. 4 and 5). To the casual observer this results
in an appearance suggesting that the endosperm is being invaded by
plates of perisperm, but this is no more true than that the promontories
of a dissected coast-line are advancing into the sea. The suggestion of
an invading perisperm is further strengthened by the fact that within
the perisperm bordering the endosperm a dark brown and finally
black band of cells is developed, due to abundant food storage (jigs.
4; 30, 31), but this really recedes as the endosperm advances.

174 BOTANICAL GAZETTE [MARCH

A cross-section of the mature seed always shows a definite and deep
constriction of the endosperm in the center, exactly opposite the two
opposed vascular strands that run up on each side of the seed through
the inner part of the outer integument (jig. 4). This constriction is
the cross-section of two opposite and deep longitudinal furrows in the
endosperm, and it means that in this longitudinal plane the endosperm
encounters the greatest resistance in invading the perisperm. This
most resistant perisperm certainly seems to hold a very definite topo-
graphical relation to the principal vascular strands, and this relation
may explain the resistance.

_ That endosperm is the aggressive tissue at every point, even at
.the region of most resistant perisperm, is evident for several reasons.
In no case were the peripheral cells of the endosperm broken down;
and in no case did there fail to appear one or two layers of disorgan-
ized cells of the perisperm in contact with the endosperm (jigs. 30
and 3r). In every case, also, the peripheral cells of the endosperm
appeared active and very vigorous, and their different appearance in
regions of more and less active encroachment is striking. In regions
of active invasion the endosperm cells are radially elongated, and
many of them are binucleate (jig. 30); while in regions of less active
invasion the cells are more nearly isodiametric and rarely binucleate
(fig. 31).

Another proof that endosperm is the encroaching tissue may be
obtained from comparative measurements. At the times of fertili-
zation the gametophyte usually measures 20 by 30#. In the mature
seed the ordinary length of the gametophyte is 20™™, the greatest
width being 14™™, and the least (at the deep constriction opposite
the vascular strands)) 1.5™™. At this most resistant region of the
perisperm, therefore, where it is hard to escape the conviction that
the perisperm plate has cut the endosperm nearly in two, the endo-
sperm has increased its diameter against the perisperm seventy-five
times.

The best reason, however, for concluding that the endosperm .
the invading tissue in this case is that this is always the behavior of
endosperm; and it is singular that the old explanation of “ rumination’ ;
was ever suggested. An examination of the nutmeg, the classic
illustration of “ruminated endosperm,” and of Asimina triloba,

1905] COULTER & LAND~—TORREYA TAXIFOLIA 175

showed that precisely the same explanation applies to them that we
have given in the case of Torreya.

SUMMARY.

The staminate strobilus consists of a series of closely overlapping
sterile bracts, in four vertical rows, completely enveloping the tip of
the axis bearing numerous stamens. The large adaxial resin cavity
that occurs in the stamen occupies the site of three abortive sporangia.

The male gametophyte has no prothallial cell, and the male cells
are very unequal, resembling those of Taxus. The pollen tube is
exceedingly variable in the rate and direction of its advance through
the nucellar cap, sometimes pushing in the embryo sac while it is
in an early free-nucleate stage.

The ovulate strobilus consists of four enveloping bracts and a
single terminal ovule with two integuments. Extensive intercalary
growth below the mother-cell forms the bulk of the mature ovule
and seed. There is no organization of a special digestive layer
around the mother-cell.

The solitary archegonium initial appears as soon as walls are
formed, is always at one side of the central axis of the gametophyte,
and forms a two-celled neck. The nucleus of the central cell was
not observed to divide, nor could any trace of a ventral nucleus be
found.

In fertilization the male cytoplasm invests the fusion nucleus,
and seems to remain distinct until wall-formation at the four-
nucleate stage of the proembryo.

In the development of the proembryo, four free nuclei appear
before wall-formation, and the proembryo completely fills the egg,
having no “open cells.” A proembryo of twelve to eighteen cells
is the winter stage. In the spring the suspensor is formed by what
may be called a wave of elongation, beginning with the uppermost
tier of the proembryo, and extending gradually downward, tier after
tier, until it includes the upper region of the meristematic cylinder
formed by the terminal cell.

Small embryos are formed during the second season in the sus-
pensor region of the normal embryo; but whether they arise from
Prothallial or suspensor cells was not determined.

176 BOTANICAL GAZETTE [MARCH

The “rumination” of endosperm, peculiar to Torreya among
gymnosperms, arises from the extremely irregular encroachment of
the endosperm upon the perisperm, the endosperm being resisted
much more at certain points than at others. The same was found
to be true of other “ruminated”’ seeds.

THE UNIVERSITY OF CHICAGO.

LITERATURE CITED.

1. CHapMAN, A. W., Pash oe Amott; a reminiscence. Bor. GAZETTE
10:251-254. map. 1
Coker, W. C., Notes on ee sspeiet and embryo of Podocarpus.
Bor. Gazerre 33:89-107. pls. 5-7.
, On the gatos and hime it Taxodium. Bot. GAzkTTE
36:1-27, 114-140. pls. 1-11. 1903.
4. Lawson, A. A., The ees archegonia, fertilization, and embryo of
Sequoia aguas. Ann. Botany 18:1-28. pls. 1-4. 1904
5. Lyon, H. L., The embryogeny of Ginkgo: Minn. Bot. Studies. 3:275- 29°
pls. 29-43. 1904.

N

Y

6. Oxiver, F. W., On some points of apparent resemblance in certain fossil
and recent gymnospermous seeds. New Phytologist 1:145-154- 19°?

ve , The ovules of the older gymnosperms. Ann. Botany 17: 451-479.
pl. 24. 1903.

8. sclera AGNES, Spore Sal in Torreya californica. New Phytolo-
gist 3:133-148. pls. 3-4.

9g. ———.,, Studies in the a aS of Torreya californica Torr. II. The

sexual organs and fertilization. New Phytologist 3:205- 216. pls. 7-9:
1904.

10. STRASBURGER, E., Anlage des Embryosackes und Prothalliumbildung bet
der Eibe nebst anschliessenden Erérterungen. Festschrift Ernst Haeckel.
Jena. 1904.

EXPLANATION OF PLATES LIV.

With the exception of Plate I, all figures were drawn with the aid of an Abbé
camera lucida and reduced one-half in reproduction. Abbreviations are ad
follows: br, bract; a, primordium of stamen; oi, outer integument; ii, inner
in

cells; im, tube nucleus; g, primary spermatogenous cell; sé, nucleus of stalk
cell; b, body cell; m, functional male cell; m, functionless male cell; fr, haus
torial region of pie eee nc, nucleus of neck cell; 0, e883 end, eD
sperm, per, peris 2 peris,

1905] COULTER & LAND—TORREYA TAXIFOLIA 77

PLATE A:

Fic. 1. Staminate branch; October 21, 1904. X1I.

Fic. 2. Ovulate branch; April 7, 1904. X1.

Fic. 3. Mature seed; October 21, 1904. XT.

Fic. 4. Cross-section of seed showing ruminated endosperm with storage
region in center, seed pe stored food in the perisperm, and resin ducts. X 3.

Fic. 5. Longitudinal median section through seed showing embryo, rumi-
nated endosperm, and seed coats. 3.

PLATE J,

Fic. 6. Transverse section through a staminate strobilus showing bracts and
primordia of stamens; gust 12, 1904 4.

Fic. 7. Longitudinal section through a staminate strobilus showing envelop-
ing bracts, primordia of stamens, and storage ges the position of the bundles
is shown by the dotted lines; August 12, 1904.

Fic. 8. Primordium of a stamen; August 12, ai

Fic. 9. Median section through an adaxial sporangium sieice the disor-
ganizing primary sporogenous cell and two wall cells; October 21, 1904. X 460.

F

resin cavity above and a functioning sporangium below; October 21, 1904. X 255.

Fic. 12. Pollen grain showing primary spermatogenous cell and tube cell;
April 7, 1904. X 1250.

Fic. 13. Tip of a pollen tube which has penetrated about half way through
the nucellus; June 10, 1904. 460.

Fic. 14. Tip of pollen tube in contact with the embryo sac; the female gamet-
ophyte is in the free nuclear stage and the nuclei are placed parietally; June 21,
1904. X 460.

Fic. 15. Embryo sac in free nuclear stage showing one of the nuclei occupy-
ing ae oS of the megaspore mother-cell; June 10, 1904. 185.

- 16. Pollen tube in contact with the endosperm; an archegonium is at
m i of the tube; July 26, r904. x25.

PLATE II.

Fic. 17. Young ovulate strobilus showing enveloping bracts, inner integu-
ment, ind a gpeiies July 26, 1904. X47.

Fic. 18. Ovulate nobihes showing bracts, integuments, nucellus with rudi-
any ss chamber, and megaspore mother-cell; April 7, 1904. X24.

Fic. 19. Tip of nucellus with pollen chamber containing a microspore;
April 8, 1904. X 220.

IG. 20. Megaspore mother-cell in synapsis; April 7, 1904. X 460.
Fic. 21. Micropylar end of female gametophyte with oe initial

Projecting into the space formerly occupied by the megaspore mother-cell; July
26, 1904. X 460.

178 BOTANICAL GAZETTE (MARCH

Fic. 22. Archegonium consisting of two neck cells and central cell; the
nucleus of the central cell has rounded out, and is passing downward to the center
of the cell and taking on the appearance of an egg nucleus; July 26, 1904. X 460.

Fic. 23. Median longitudinal section of female gametophyte showing egg,
remains of a neck cell, and antipodal haustorial cells; the tip of the pollen tube
in contact with the egg contains stalk and tube nuclei, functional and functionless
male cells; the upward growth of the endosperm cells forms a sheath around
the pollen tube; August 12, 1904. X 460.

PLATE III.
1G. 24. Fertilization; the male nucleus is in contact with the egg nucleus;
the cytoplasm of the male cell is closely applied to the egg nucleus; the function-
less male cell, and tube and stalk nuclei are in the upper part of the egg cytoplasm;
the cavity in the egg cytoplasm is caused by the inrush of the contents of the
pollen tube; August 12, 1904. X 460.

Fic. 25. Two-celled proembryo; the dense cytoplasmic mass surrounding
the ey is — in greater part derived from the male cytoplasm; August
27, 1904.

Fic. - , eee proembryo; walls coming in; August 12, 1904. 40.

Fic. 27. Proembryo shortly after walls are laid down; the proembryo passes

X 460.
Fic. 28. Cross-section through suspensor cells; September 12, 1904- X 460.
Fic. 29. Proembryo showing elongation of suspensor cells; April 7, 1994

Fic. 30. Endosperm cells encroaching on perisperm; August 12, 1904- ee
Fic. 31. Endosperm cells which have ceased to encroach on perispe!™
August 12, 1904. X460.

—

PLATE

BOTANICAL GAZETTE, XXXIX

COULTER & LAND on TORREYA.

BOTANICAL GAZETTE, XX XIX PLATE 1

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BOTANICAL GAZETTE, XX XIX PLATE Jf

PLATE (Ee

BOTANICAL GAZETTE, XX XIX

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THE PRINCIPLES OF PHYTOGEOGRAPHIC NOMEN-
CLATURE.'
PEHR OLSSON-SEFFER.

THE confusion prevailing within the nomenclature of phyto-
geography has of late been repeatedly brought up for discussion.
A unanimous opinion exists as to the disadvantages resulting from
the present chaotic terminology. Such a diversity of ideas prevails
that every writer is obliged to explain in what sense he has used a
technical term, and if he omits an explanation we are often left in
doubt as to the interpretation of the expression used. An agreement
has to be arrived at, sooner or later, and the sooner there is an end
to the present disorder, the better for the progress of phytogeography
and all concerned.

This science is still in its infancy and very few of its doctrines are
settled beyond doubt. We meet with contradictory views at every
step. Sweeping generalizations, often based upon very imperfect
observations, and described in vague and uncertain terms, full of
ambiguities, threaten the development of this important science.
Its advancement and success depend upon well-settled methods of
investigation and description. The necessity of adopting a nomen-
clature similar to that of other descriptive sciences is obvious. The
prevalent fault of which we complain is not the absence of names
and technical terms, but of the defective definition of the terms
now in use. It has always been much easier to offer censure than
to correct mistakes, easier to state evils than to relieve them. My
object in this paper is not to give any proposals as to the detailed
atrangement of phytogeographical nomenclature, but to discuss some
of the general principles which ought to prevail in any attempt to
revise the nomenclature so as to meet the present demands of the
science. I have elsewhere, in papers appearing simultaneously
with this, drawn attention to the terminology of certain phytogeo-
graphical phenomena.

Read before Section G of the American Association for the Advancement of
Science, Philadelphia meeting, December 30, 1904.
Igo5 179

180 BOTANICAL GAZETTE [MARCH

Since the appearance of the papers of WARBURG, FLAHAULT,
NILsson, and CLEMENTS on this vexed question of nomenclature,
it could have been expected that some discussion would arise, but
instead it seems as if phytogeographers were content with the opinions
brought forward, although these are by no means harmonious. The
object of plant geography is the study of the distribution of plants,
and of the laws that govern this distribution, not wrestling with words
nor philological hair-splitting. But it cannot be denied, as I have
just said, and as is generally conceded, that this science needs uni-
formity of expression; how to obtain this is the problem.

The next international congress, to be assembled at Vienna in
June 190, is to take this matter under consideration. Some of
the foremost phytogeographers of the world constitute a committee
which is to make a report and submit certain proposals that may oF
may not lead to a solution of the problem. It is only the general
principles, however, that can be laid down by any one botanist or
any joint committee, because it is quite beyond the power of any
one except the individual monographer to decide what expression
may be necessary in his individual case. But so long as each one
is allowed to do what he thinks is right, there will be no end of the
present trouble. What we need is a good, clear system of wholesome
general rules. The practical and gradual application of these must
be left to the discrimination of the individual writers, who have
describe conditions essentially different in different cases. The
reader is then the judge whether the writer has succeeded in his
application of the rules, whether his terminology is correct oT not.
No permanent international committee is needed for the purpos’
of acting as a guardian oracourt. Ifa good code of rules for ia
clature is laid down, it will be followed spontaneously by all writers
of any consequence, without the fear of a court of judges. If the
code is a bad one, it is not worth following, and will not be accepted
in spite of any supervising committee, however great its authority:
It must be remembered that no matter from what association
individual such a law of nomenclature has emanated, it is and
always be temporary. It is impossible to determine upon any rules
that will stand for all time, because what meets the present needs
will most likely not satisfy the next generation. How long, them

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1905] OLSSON-SEFFER—PH YTOGEOGRAPHIC NOMENCLATURE 181

shall a writer consider himself bound to follow these rules? They
will certainly have to be changed from time to time. It is especially
impossible to decide upon the fixity of special terms, because a name
may be founded on ideas, which, in the course of time, owing to the
progress of science, will be shown to be incorrect. No such rules
as are applicable to the nomenclature of taxonomy can be brought
into effect in regard to a system of terminology.

The law of priority, which is the first principle in the nomen-
clature of systematic biology, cannot be strictly adhered to in this
connection. It has been proposed by CLEMENTS? that “priority of
term and of application is to be regarded as the fundamental principle
of phytogeographical nomenclature.” ENGLER, who must be con-
sidered as better qualified than most men to judge in such a matter,
in a footnote to CLEMENTS’ article expresses strong objection to the
introduction of a law of priority in phytogeography. The acceptance
of such a law would lead to the retaining of names which are neither
expressive of the idea they represent nor suitable in other ways. If
an absolute rule of priority is maintained, how are we to arrange for
the retaining of names that originally expressed ideas now considered
as false? Every terminology shows traces of such names. What
conception in that case shall represent the type of the systematist
and bear the old name? It can be seen at a glance that the
tule of priority is not practicable here as it may be in taxonomic
nomenclature.

The question whether to retain an old term which is not good,
or to abolish it and substitute a new name, will always be difficult.
If a free hand is given, phytogeography will have a heavy load of
useless synonyms that always will act as a drag on true science, and
create much more confusion than exists now. If on the other hand
any restriction can be brought about, it must be to the effect that
Priority should be conceded to such a name only as has been properly
defined in a work accessible to scientists. To impose new names
needlessly upon previously named conceptions will always be con-
sidered bad form, and a general consensus of opinion prevails as to
this habit. Suddenly introducing a large number of new terms into

* CLEMENTS, F, E., A system of nomenclature for phytogeography. Beiblatt
Bot. Jahrb. 70: — 20. 1g02.

182 BOTANICAL GAZETTE [MARCH

a new science is the best way to stifle it. It is with great hesitation
that a new term should be coined, and it needs in each case a spe-
cialist to decide as to the necessity for such an action. Good and
forcible reason should always be shown why an old term is not sufl-
cient to indicate the conception that is to be described. It should
never be forgotten in rejecting already established names that there
is a possibility that the new term may meet a similar fate from a
later writer. If that were always borne in mind, perhaps writers
would think twice before entering the arena as name-makers. The
true test of the quality of a term is generally the time it is able to
exist, provided the conception as such remains unchanged. Not
infrequently, however, names are introduced into a science—phyto-
geography not excepted—for speculative theories and ideas, upheld
and supported by facts which are consciously or unconsciously mis-
interpreted to fit preconceived notions. In other cases new terms
are proposed for already named conceptions, because of ignorance
on the part of the writer. We may here remember what DECANDOLLE
says in his Phytographie: ‘“‘the perfectly honest and right-minded
botanist may sometimes have failings. He may neglect to cite his
predecessors, or cite them inexactly, either from negligence or from
want of literary resources. The latter case may be deemed a mis-
fortune and no fault.” “But,” continues DECANDOLLE, “if he
has not the necessary books within his reach, why not go where
they are and consult them? Or if unable to do that, why need he
publish?” Where a writer may have enriched the nomenclature
with a new term for which no need existed, the application by sub-
sequent writers of the rule of priority is to be recommended. If the
term in question be introduced by a writer who enjoys real or affected
authority, and his term is accepted upon such motives by a thought-
less multitude, it will naturally sooner or later be suppressed oF
ignored and finally disappear.

When we undertake to revise nomenclature and find terms the
meaning of which is doubtful, the only proper way out of the dilemmé
seems to be to ascertain the conception given to such terms by the
original proposer. If the term cannot be used in that sense it should
be discarded. We have a good illustration of this in the much ™*
treated term “‘phytogeographic formation.” It was originally pro

bis Teas

1905] OLSSON-SEFFER—PHYTOGEOGRAPHIC NOMENCLATURE 183

by GRISEBACH (1838) to designate an aggregation of plants char-
acterized by a dominant species. But, as his examples show, GRISE-
BACH considered his formations as having a certain physiognomic
aspect, and when in 1872 he moderated his conception of the term
formation to an association characterized by a physiognomic type
instead of by a dominant species, he only more pronouncedly brought
forward the conception of a physiognomic unit. Leaving aside all
the various uses of this term by later writers, we have to consider
whether the original conception of GrIsEBACH of a general plant-
topographic or physiognomic unit, such as forest, steppe, tundra, or
prairie, can be retained in the light of modern investigations. If
that is the case the name stands, if not it falls. I have endeavored
to show, in a paper now in print, that we need the term formation
in the sense of GrisEBACH. Did our limits allow, we might call
attention to many other instances where we could clear up the
muddy stream of phytogeographical names. One more example will
suffice to show how this rule would work. The word zone is now used
as a technical term in phytogeography to designate at least the fol-
lowing conceptions: the successive belts of vegetation on a mountain
side, the horizontal climatic zones of the surface of the earth, the
belts of vegetation surrounding a pool or succeeding each other on
a shore, the submerged belts especially of marine algae, the layers
in a fossil-containing deposit; in many local descriptions it is adapted
for designating any convenient floral area delimited from others;
and finally in anatomy the term “zone” is applied to any area dis-
tinguished in structure from its surroundings. That this multiform
interpretation of the word zone needs adjustment is manifest. In
the technical language of a science a term should have only one
meaning. Nothing but confusion will come from the admission
of enigmatic terms, and the clarifying process is therefore the one
we expect the nomenclature of phytogeography will shortly be
exposed to. If in regard to the term zone we follow the principle given
above, we have to ascertain what author first introduced it as a tech-
nical word, and in what meaning it was used by him. In 1839
Bolssrer designated with 'this term the vertical belts of plants in
Mountains. In a corresponding sense it had been used for a long
time previously in topography, as also FLAHAULT? mentions. The
$ Projet de nomenclature phytogéographique. 1890.

184 BOTANICAL GAZETTE [MARCH

term zone in plant-geography, therefore, should signify the successive
stages of vegetation from the base to the summit of a mountain, and
nothing else. It is true that WAHLENBERG4 in 1812 designated these
belts with the term regio, and if we followed the rule of absolute
priority, this would be the correct term. Regio, however, was used
long before WAHLENBERG’s time by botanical writers in the general
sense of a geographical area of more or less definite extension, and
so was zone. In this case we are confronted with two synonymous
terms, and the only principle on which a decision can be based seems
to be that of general usage. It must be admitted that it is an
extremely difficult matter to lay down any rules that would take us
out of dilemmas such as this. If it were possible to canvass the
various authors to ascertain which term has been used more than
the other to designate this special feature of mountain belts we are
considering, it is very doubtful at what result we would arrive.
Region, however, since Martius used it in 1831 for a certain phyto-
geographical area, has generally been understood and adopted by
the best writers for that purpose. FLAwAuLT has made the relation
and usage of these two terms, zone and region, clear by adopting
them in the sense advocated above. If that is universally done, we
have to find other appropriate terms for the various conceptions
that often have been called zones. Nor need we take refuge to the
method of making new terms in this case, for we only have to make
a selection from the multitude of expressions already used, and in
selecting we can make a choice that will serve in other languages
as well as our own, and thus to some extent satisfy the call for an
international nomenclature.

The first and most essential principle of nomenclature is clearness.
To obtain this result, all the expressions used in technical terminology,
whether they be old established names or newly coined ones, must
be definite, concise, perfectly distinctive, and easily intelligible.

All names and terms are for the sake of convenience. In order
to insure mutual intelligibility, greater precision, and clearness,
is imperative to avoid names that will create error or ambigully-
It is not inconceivable that the need of short compendious namé
and terms to denote phytogeographical facts or processes can be met

4 Flora Lapponica. Berlin. 1812.

Pe eee ee Tee

1905] OLSSON-SEFFER—PH YTOGEOGRAPHIC NOMENCLATURE 185

by individual monographers following certain general principles,
which should be applied with all possible generality. Festina lente
is the maxim into which we might condense the prevalent, but per-
haps not yet outspoken condemnation of the tendency to drown our
science in a torrent of unpronounceable so-called international terms,
which cannot but embarrass the student, render the subject less
accessible and more difficult to handle, and be exasperating to lovers
of a clear, consistent, and uniform nomenclature.

Emphasis must also be laid upon the manner in which a term
comprises the idea it is to convey to the reader. In systematic
biology it is now held that a name need not contain any reference to
the subject it represents, and may be wholly meaningless. This

would hardly be convenient, however, in a system of terminology.

The limitation of human memory makes it important that the term
or name employed in a descriptive science should not be merely a
name, but also associate in one form or another our thought with the
subject we are discussing. In making new terms or in discriminating
between already existing ones we should bear this in mind. It is
just as easy to coin a name of this kind as it is to make a meaningless
one. There might be a tendency to attribute too much importance
to the meaning of a name, but all things considered it seems easier
to remember a term that at once conveys to our mind the conception
it stands for.

It has been recommended by several writers that we ought to
avoid having names that are already used in geology or some other
science nearly related to phytogeography. It stands to reason that
such a course is neither absolutely necessary nor very advisable.
Although we speak of stratification in connection with sedimentary
processes in dynamic geology, it does not follow that the term strati-
fication could not be used in plant geography to designate the division
of a plant community into strata, without implying any ambiguity
or causing any confusion. Objection has been made in regard to

‘certain terms such as formation and province, the former word being

used in geology, the latter in a political sense. If we were consistent,
a great number of names which have been used for a long time in
botany ought to be ruled out, because they are also used in zoology,
Or vice versa, as anatomy, cytology, heliotropism, parasitism. Any

186 BOTANICAL GAZETTE [MARCH

small inconvenience that may result from this principle of ignoring
the fact that a term is already established in another science would
be counterbalanced by the appropriate use of similar terms to desig-
nate related phenomena in related sciences.

It is of importance in applying technical terms or inventing new
names to take a broad view of the subject, and not use geographical
terms, that generally refer to large areas, to signify local phenomena.
We might mention as an example the use of the expression Austral
zone for a phytogeographic area of North America. It has always
been understood, however, that the term austral refers to the southern
hemisphere, and it is as wrong to use the word in the way mentioned
as it would be to apply its counterpart boreal to the northern part of
Brazil or Australia. It would be greatly misleading if a botanist,
say in Australia, would designate for instance the eastern coast
region of that continent as the Oriental region.

When Linnaeus brought about the reformation of systematic
nomenclature he freed the names from the cumbrous descriptive
phrases by assigning to each object a generic and a specific name.
Similarly a concise mode of expression in phytogeography ought to
be agreed upon, so as to save a great deal of verbosity which at pres-
ent naturally must accompany an exact and complete phytogeogt aphi-
cal description. The difficulty of presenting the results in a compact
form would not be very much bettered, however, by adopting sud-
denly a number of new terms, because most likely the remedy would
in that case be worse than the disease. It may be safely said that
by instituting a uniform method of applying necessary terms, and by
bringing such an agreement into universal practice a long step would
have been taken towards establishing order.

The question of obtaining a nomenclature of an international
character has been discussed to some extent. There can be no doubt
as to the beneficial results that would follow the adoption of such
names. The practicability of the application of any rule to that

effect, however, seems somewhat doubtful. Be that as it may, W°

are justified, I think, in looking forward to some kind of tacit agree:
ment in this case between plant-geographers of different countries:

Another question is how such international terms should be
formed. Warpurc and CLeMeEnts think that only Greek and Laun

Se ey ee Oe a

—

1905] OLSSON-SEFFER—PHYTOGEOGRAPHIC NOMENCLATURE 187

can be used. FLAHAULT, NILSSON, WARMING, and ENGLER, among
others, are of the opinion that vernacular names ought not to be
excluded. While it seems absolutely imperative that in referring to
plants we should use only the scientific name, it does not appear to
be so overwhelmingly important to change all those terms of ver-
nacular origin, which already are established in phytogeography,
into quasi-international substitutes drawn from Greek or Latin. In
the former case we have thousands of names, and consequently we
use the accepted scientific names, which can be easily identified,
instead of the vernacular, as the latter would surely give rise to con-
fusion. On the other hand, the number of technical terms in phyto-
geography is fortunately not yet so great as to make the list a very
voluminous one. Even if vernacular names are retained, or intro-
duced to designate certain facts, this would not militate against.
a uniform nomenclature so long as the names are clear and do not
give rise to any doubt as to their significance.

One fatal objection to a change into Greek-Latin of such com-
monly understood and accepted names as tundra, prairie, chaparral,
scrub, savannah, and others, is that in spite of the adaptability of
these ancient languages, it is impossible to translate these terms
adequately, since the ancients did not have any conception of the
ideas or facts these names represent, and any attempt to fabricate a
modern name from the ancient languages to signify, for example, the
formation known as the patana of Ceylon would be a failure, so long
as we want the term to suggest to our mind the peculiar conditions
that characterize this particular formation. If we consider the chief
object of nomenclature to be to serve our convenience, I fail to see
why the name patana would not be acceptable in any language, and
thus be international. As a matter of fact, it is so already, and in
all probability very few persons would approve of a Greek-Latin
equivalent coined according to the principle of constructive naming.
CLEMENTS, who is the principal advocate of the latter method, and
who has augmented the labor of those who are endeavoring to find
a4 way in the labyrinth of phytogeographical terms by proposing at
least 500 new names, has given the name psilium to a prairie forma-
tion, deriving this “international” term from WeAeiou or yura, bare,
naked. Now this new name does not convey any idea of a prairie

188 BOTANICAL GAZETTE [MARCH

to the reader because it can be applied as well to any treeless forma-
tion. Prairie, on the other hand, is a term well-known to every school-
boy the world over, and there is no need whatever to overload mem-
ory with a new name that serves no useful purpose. For the term
“bad lands” of Nebraska CLEMENTS suggests hydrotribium, and
for plants on that formation hydrotribophyta. Terms such as these
are very expressive in a way, but they are certainly not an improve-
ment. The English “bad lands” applied to this particular forma-
tion is widely known, it causes no ambiguity, and there would be no
objection raised against its acceptance in French, German, or any
language, but most people would certainly protest against hydro-
tribium. And still this last term can be pronounced, but what about
ptenothalophyta, rhoium, ammochthophilus, proodophytia, mesoch-
thonophilus, chosen at random from CLEMENTS’ catalogue ? The
terms rolling prairie, rolling foothill, rolling downs, and so on, can
readily be adapted in any language, and be just as characteristic as
if we translated them into some more or less high-sounding name
derived from the Greek word for rolling, or wheel, or ball, or some
similar expression. In geology many characteristic words have been
borrowed from the vernacular for technical use, as fjord, atoll, and
canyon. Would physiography have been better off in regard to clear-
ness and brevity of expression if names of mixed Greek and Latin
origin had been invented for these conceptions, which had no equiva-
lent in the language of the ancients? ENGLER, in the footnote to
CLEMENTS’ article already referred to, gives the following categorical
judgment in this matter: .“‘dass es sicht nicht empfielt, die volkstiim-
lichen Bezeichnungen von Pflanzenformationen aus der pflanzen-
geographischen Literatur zu verbannen.”

What has here been said may suffice to show that new names
cannot be invented to advantage for features that already have well
established and characteristic designations in the vernacular language
of the country where they constitute a salient feature. Good common
sense in this as in many other cases must be the guide in choosing
technical expressions.

If a suggestion were offered as to the first step necessary in order
to obtain uniformity, it would be that we have to decide about the
various kinds of floristic, topographic, and ecologic units that cam be

a li i a a

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1905] OLSSON-SEFFER—PH YTOGEOGRAPHIC NOMENCLATURE 189

and are necessary to distinguish. It has been suggested’ that the
division of the vegetation into formations must be founded upon the
concept of habitats. This principle is a good one, the only difficulty
seeming to lie in the practical working of the rule. Any one conversant
with the great variety of forms of habitat is aware that such a classi-
fication is no easy undertaking. In all attempts made the authors
have decided for one or another environmental factor that has influ-
enced the development of the formation, and consequently the classi-
fication has been more or less artificial. The task of identifying,
classifying, and naming plant aggregations, or features of the vege-
tation, is extremely difficult because of the comprehensive data
necessary to illustrate the complex factors influencing distributional
phenomena. The use of one class of names that refer to habitat,
however, are inevitable and absolutely necessary. For such terms
we can turn to the vernacular language, which often possesses very
expressive names that combine in one word the main features of
environment. Together with the term formation as representing
the large topographical units of vegetation such vernacular terms are
very adequate. Let me give a few examples to illustrate this. The
chaparral formation of California and southwestern United States
generally is one of the most: peculiar anywhere. There can be no
doubt about the meaning and scope of the term, when the formation
has once been clearly defined, because it has no counterpart in any
other region of the world, although it certainly is paralleled in many
places by related formations. The expression chaparral formation
gives not only a general idea of the component plants, but it also
includes a conception of the topographic aspect of the country where
the formation occurs, and whose physiognomy it assists to mold.
Still the term is strictly devoid of any reference to the dominant
species concerned in the aggregation of plants in any part of the
formation. Formational names should always be so. A formation
can be subclassified into associations, and these into communities.
The latter can be designated adequately by adding the suffix etwm to
the scientific name of the dominant plant after the method first sug-
gested by Hutr.® The limitations of this article do not allow me

5 CLEMENTs, be G.

® Férs6k till en analytisk behandling af vixtformationerna. Medd. Soc. F. Fl. F.
8:1-155. 188; :

.

Igo BOTANICAL GAZETTE [MARCH

to enter into details with regard to this method of classification,
which I have found works equally well in the arctic parts of Europe
and in the Mediterranean countries; in the primitive tropical forests of
Ceylon, northern Australia, and Polynesia, and in the semi-arid regions
of New South Wales and Western Australia; in New Zealand and
in California. I may say that in my work in these countries I have
not found it necessary to introduce any new system of naming, but
merely to coordinate and classify previously existing conceptions.
And I firmly believe that this working out of the synonymy will be
the only method by which a final agreement can be reached.

We have further to classify all the units of one kind or another
into groups according to relationship, and to give names to them,
availing ourselves as much as possible of terms already existing, the
synonymy of which must be cleared up thoroughly. After laying
down the general principles of nomenclature in the code to be recom-
mended, we have to leave the application of these canons to the
individual writers.

This should be the program for a permanent committee to be
appointed by the next international congress. Let this body then
publish the results of their work and submit them for leisurely con-
sideration and discussion. When conflicting opinions, if there are
any, have been expressed, and direct or circumstantial evidence
has been brought in to illustrate and reinforce the various principles,
it is time to settle the matter by adopting a general code of nomen-
clature. There can be no hope of getting any substantial improve
ment in existing conditions through any immediate action of the
botanical congress, because of the great diversity of opinion and
practice that prevails, and because no definite proposal based on
facts and logical arguments has yet been held forth which could be
made subject to a detailed criticism. Any proposition that presents
principles or terms without proper and clear definitions can naturally
not be considered. -

The question may arise as to what constitutes a definite descrip”
tion. The degree of exactness and clearness of expression W!
naturally vary with the different authors, but so long as there remains
any doubt as to the feature meant by the writer his term can hardly
expect to be generally accepted.

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a ee eee eee

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1905] OLSSON-SEFFER—PHYTOGEOGRAPHIC NOMENCLATURE 1g1t

Nomina nuda in the sense of systematic biologists appear not
infrequently in phytogeographic literature, and it must always be
considered as insufficient. definition to supply merely a translation
of a formational name, without giving a description sufficiently clear
to remove every trace of doubt as to what the writer has described.

Whenever coining of pseudo-classical names is resorted to, it is
to be expected that the author would at least take into consideration
some degree of linguistic purity, besides the matter of precise mean-
ing, because it is important that terms which are also to be spoken
should be euphonious and in some harmony with the pronunciation.

In regard to the rules of coining new terms from Greek and Latin
it has been claimed that the classical languages only should be con-
sidered. Rules of that kind are difficult to follow. In the real
classical Latin, for example, the words are used in so many different
ways that it will often be difficult to bring them into conformity
with the primary rule of technical nomenclature that each idea
should be represented by a single term only, and each name should
have only one meaning. The classical Latin of botanists has been
the Latin of Linnaeus, and it will most likely always remain so,
because of its definiteness and conciseness. Whatever language
is chosen, in the forming of new words we must follow the rules of
the language; but we must also remember that by driving the sys-
tematizing too far we will only increase the difficulties, and by a too
sedulous adherence to preconceived notions we might arrive at results
which are not in unison with the true progress of science.

One important point is that in forming new terms for phyto-
8eography we must avoid terms which already, in one form or
another, are used in botany. To call an orchard formation dendrium
will lead to difficulties, because that name is already used to designate
@ genus of plants. The same objection can be made to eremia,
amathia, lophia, petria, xylia, and scores of other terms proposed
by CLEMENTS

The rule suggested by that author “that a term to be valid must
be Proposed by a botanist” is incongruous. In hazarding this crit-
1cism I must confess that it appears to be a hard rule that forbids
any one who has facts to present concerning the vegetation or flora
from doing so, provided he is able to express himself correctly, no

192 BOTANICAL GAZETTE . farce
a

matter whether he claims the title of botanist or not. Or is it the
coining of names only that should be restricted to botanists? Who
is a botanist? Where shall the limit be drawn? Does knowledge
of a certain number of plants entitle a man to this privilege? Or is
publication of a certain number of pages on some botanical subject
a sufficient qualification? If such a rule were accepted and applied,
we should have to reject, for instance, the term “ecology,” because
it was first introduced by Ernst HAECKEL, who never claimed the
name of a botanist, although he knew more about the subject than
most “botanists.”

Enough has been said upon this matter. We should take up a
conservative position in this question of nomenclature, but at the
same time insist upon the adoption of a code announcing certain
principles, the application of which will prevent such a plentiful
harvest of confusion as we have now, and assist in bringing about a
reform resulting in a nomenclature better adapted to the needs of
scientific workers. ‘Prove all things; hold fast that which is good”
is the very essence of such a code.

To summarize the previous discussion:

Clearness and conciseness are the main requisites for a system
of terminology.

Each technical term should have only one meaning.

In case of doubtful terms consult the proposer of the name. If
the conception it represents is not absolutely clear, the name has no
status In nomenclature.

If a term has been commonly used and understood in another
sense than the original author proposed, it should be retained, but
only in case there can be no doubt as to its interpretation.

If a conception has already received a name and there is no obvious
reason to discard that name, an author has no right to propose @
new term.

A law of priority is practicable, we think, only so far as the prin-
ciples laid down in the previous pages of this article will admit.

A name, the conception of which has been materially changed 10
the course of time, naturally has no standing.

A technical term should be associated in our mind with the idea
it represents.

1905] OLSSON-SEFFER—PH YTOGEOGRAPHIC NOMENCLATURE 193

A technical term should be defined clearly, so as to leave no doubt
as to its significance. Authors should desist therefore from pro-
posing new terms in mere catalogues.

A new term should be published in some work accessible to scien-
tific workers.

Vernacular names should not be excluded from phytogeographical
nomenclature, but they must in every case be definite and give rise
to no ambiguity.

An international committee of phytogeographers should be
appointed by the Vienna congress, to continue the work on a pro-
posed code of rules.

This committee should consider what kinds of technical terms
are needed; how they should be classified, for example with regard
to distribution, abundance, elevation, phenological phenomena, etc.

The result of the work of the committee already existing, and of
the succeeding one, should be published at an early date, so as to
give the public ample time to discuss the various phases of the ques-
tion, before the following congress assembles.

STANFORD UNIVERSITY, CALIFORNIA.

THE FORESTS OF THE FLATHEAD VALLEY, MONTANA.
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY.
XVII.
Harry N. WHITFORD.
(WITH MAP AND TWENTY-THREE FIGURES)
[Continued from p. 122.]
II. EDAPHIC FORMATIONS IN FLATHEAD VALLEY.

In the previous pages an attempt was made to show why the west
side of the valley contains a prairie formation, and the east side a
forest formation, and why the coniferous type of forest prevails
rather than a broad-leaved deciduous type. A plot will now be
presented in detail to show why in the forest formation there are
some areas altogether without trees. In order to reach definite con-
clusions, a limited area was selected to represent the whole forest area
of the valley. This area, comprising about 250 square kilometers, was
mapped, and the conditions for forest growth were determined (map).
The relation of this area to the whole valley is shown in fig. 3- It
lies at the northeast end of Flathead Lake, and includes a portion
of the main valley on the east side of the Flathead River at the head
of the lake, the low end of the Mission Range of mountains, and a
portion of Swan valley. In this area are found many types of topo-
graphic diversity, from Flathead Lake (890™) to the highest point
(about 1372™) in that part of the Mission Range which is included
in the map. No detailed study was made in the higher Swan Range
east of the valley, though some conditions of tree growth were noted
in these mountains. As shown by the map, five distinct edaphic
formations were found: meadow (hydrophytic), Engelmann spruce
(meso-hydrophytic), western larch-Douglas spruce (meso phytic),
Douglas spruce-bull pine (meso-xero phytic), and prairie (xero phytic):

THE MEADOW FORMATIONS (HYDROPHYTIC).

As already mentioned, Swan valley was formerly a lake, the pres-
ent condition being only a stage in its recession. The drainage
consists of Swan Lake, Big Fork River, and a number of sma

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1905] WHITFORD—FORESTS OF FLATHEAD VALLEY 195

streams; also one large pond known as Ross (Mud) Lake, which,
with its outlet Ross Creek and the numerous small inlets, comprise
the drainage of the extreme north end of the valley.

The most noteworthy fact about the hydrophytic vegetation is
its resemblance to that found in similar edaphic situations in the
eastern United States, both types of swamps being found here.
Near the head of Ross Lake is a sphagnum meadow, in which Meny-
anthes trijoliata, Drosera rotundifolia, Comarum palustre, Erioph-
orum polystachyon, and Betula pumila are characteristic. At least
two other large meadows show this combination. One of these is
along Little Bear Creek, and is known as the School Meadow;
another is found along Flathead River, and will be discussed in con-
nection with the spruce forest which lies close to it. Ross Lake
(fig. 9) is a pond which shows very well the zonal distribution of
plants, being shallow and with a very muddy bottom. In its center
and in streams bordering it are found the following characteristic
species: Nymphaea advena, Brasenia purpurea, four species of Pota-
mogeton, Myriophyllum, and Hippuris. The sedge zone bordering
the water of the shallow ponds and streams contains the following
species: Bromus Richardsonii pallida, Muhlenbergia racemosa, Carex
utriculata, C. viridula, C. hystricina, Calamagrostis caespiiosa,
Phalaris arundinacea, Juncus Regelii, Scirpus lacustris occidentalis,
Lobelia sp., Cicuta maculata, Solidago sp., and Dodecatheon sp.

The meadows are usually submerged during the spring and early
summer months, when the melting snow of the mountains to the east
swells the streams, and brings the water level slightly above the sur-
face. In the latter part of the summer and fall the underground
water level is a little below the surface; and it is this condition of
submergence and emergence that determines the meadows and
excludes trees. Of course some of these meadows are never entirely
submerged, but are always wet, with few exceptions being associated
with streams. The largest meadows are found near the head of Ross
Lake, at its foot on the east side, near the mouth of Wolf Creek, at
the foot of Swan Range along Little Bear Creek, and in the large
bend of the river just east of Swan River cafion.

The meadows are given brief mention here in connection with
this forest study because they are genetically related to the spruce

196 BOTANICAL GAZETTE [MARCH

forests that usually border them. It is obvious that trees do not
grow in them because the amount of moisture in the soil is too great.
Some of the meadows have been artificially drained, others have
been planted to timothy, and one in particular, in the northwestern
corner of the area (map), has been drained and cultivated for a num-
ber of years, so that its former nature was discovered with difficulty.
What has been done artificially is being done naturally, but more
slowly. The detritus that is washed into the ponds and the vegeta-
tion that dies down from year to year are filling up these bodies of
water, and are making the meadow condition possible. Again, the
streams are cutting their channels deeper, thus lowering the under-
ground water level at their borders, and this changes conditions that
favor meadows and introduces those that make the forest possible.
It is very probable that many areas now occupied by spruce and other
forests were formerly occupied by meadows. Indeed, all stages of
development towards this forest condition were noted, and will be
discussed under the next head.

THE SPRUCE FORMATIONS (MESO-HYDROPHYTIC).

In many places species of Salix and Alnus incana encroach upon
the meadows, sometimes forming dense breastworks around the
grass areas, so that it is difficult for other trees to get a start. This
is the case when the zones of underground water level are sharply
and narrowly marked off. Usually, however, when the underground
water level is at a fairly uniform distance below the surface for 4
rather broad area, as it is where the small streams spread out over @
level extent of land, the willow-alder breastwork is absent. These
situations are favorable for the advance of Picea Engelmanni into
the meadows. The youngest trees are found on the hummocks 10
the meadows; nearer its’ borders are the older ones, though they
are still scattered; these grade imperceptibly sometimes into dense
forests of spruce, many of the trees reaching huge dimensions. since
this formation is found best developed on situations where the under-
ground water level is a little farther below the surface than it 5
the meadows, it may be called the meso-hydrophytic formation.

It must not be supposed that there are no other trees with the
spruce; indeed there are only small tracts where pure spruce woods

1905] WHITFORD—FORESTS OF FLATHEAD VALLEY 197

are found. The most common companion trees are Populus angusti-
jolia, Populus tremuloides, and Betula papyrijera. These species
nearly always form a small per cent. of the spruce forests, and in the
meso-hydrophytic area along the Flathead River they occupy nearly
the entire space. In Swan valley the soil is a mixture of clay and
humus; along Flathead River, especially in the delta region, the
subsoil is sand, overlaid by a fine alluvial deposit. In the dense
woods the undergrowth is scanty, sometimes only one or two species
being present, the most common plant associated with the pure
spruce growths being Rhamnus alnijolia, although Cornus stolonijera
is often found. In the narrow strip of this forest along the streams
nearer the mountain Echinopanax horridum and Veratrum calijor-
nicum are present. Since the spruce woods are transitional between
the meadow and the mesophytic woods, the undergrowth of these
associations is often present. Thus, near the meadow side of the
forest, grass plots are frequently present, and on hummocks between
anastomosing channels Lysichiton kamtschatcensis is frequent. On
the other hand, toward the mesophytic side of the forest the elements
of the undergrowth of the western larch-Douglas spruce formation
are to be found.

An area of spruce is found in connection with a meadow in the
northwestern corner of the plot not far from the Flathead River.
Here on the border of the prairie formation is an area that has almost
identically the same plants as the spruce forest in the Swan valley.
Along the east side of this swamp there are numerous springs, some
large enough to give rise to small streams, and others hardly notice-
able. The source of the water supply is thought to be Echo Lake,
which, with a number of smaller ones, lies in the depressions of a
moraine that extends beyond the limits of the plot to the northeast.
Save a small stream, there is no visible inlet to this lake, and its
source of water supply is probably by underground seepage from the
mountains that lie northeast. There is also no outlet above ground,
and since the drainage is toward the Flathead River it is thought
that the springs mentioned are fed by underground seepage from
these lakes and ponds. This underground water level, as shown
by the wells in the neighborhood, is in places nearly a hundred feet
below the surface, and extends underneath the larch-Douglas spruce

198 BOTANICAL GAZETTE [MARCH

and bull pine forests and the prairie to a low ridge near the Flathead
River, where it issues in the form of springs, making conditions
favorable for an Engelmann spruce formation. Thus a bit of forest
is projected, as it were, into the prairie formation. Where the water
level comes nearer the surface the meadow already mentioned is
associated with the forest.

Another meso-hydrophytic forest in which the spruce is almost
entirely wanting needs to be mentioned. Around the head of Flat-
head Lake, including the delta at the mouth of the river (figs. 1 and 4),
and extending up the river from its mouth, is a region that is annually
submerged." This region is occupied by a dense growth of willows,
cottonwoods, birch, aspen, and other trees. On each side of the
mouth of the river are long, low peninsulas of sand, the ends of
which are entirely barren of vegetation. These are above water
only a short time during the growing season, and this fact no doubt
prevents plants from getting a start. A short distance back from
the end of the delta on a little higher land is Salix fluviatilis, in some
places so thick as to be almost impenetrable. Underground stems
are sent out from this center, and thus new territory is gradually
conquered, so that as fast as the delta is built out it is occupied by
this pioneer willow, which prepares the way for changes that make
it possible for higher forms to exist. When the water is high its
contact with the willows causes a deposit of fine silt, so that the delta
is gradually built higher and higher, making conditions favorable
for the growth of such plants as the red dogwood, other species of
willow, narrow-leaved cottonwood, paper birch, aspen, choke cherry,
two species of hawthorn, and service berry.

Around the head of the lake there is a low terrace which is also
submerged in the spring and early summer months. Instead of
Salix fluviatilis there is a thicket of other willows; and on the lake
side of this there are a few scattered specimens of S. fluviatilis and a
amygdaloides. On the land side of the willow thicket are the birch,

sive floods do great damage to the country lying at the head of the lake. Since nearly
= oe in the valley have their sources in the mountains to the east, the pooner
tion of this region would greatly increase these spring floods and greater damage woul
be the result.

a

1905] WHITFORD—FORESTS OF FLATHEAD VALLEY , 199

cottonwood, and aspen; and here and there are found small patches
or single specimens of the western larch, Engelmann spruce, Douglas
spruce, and bull pine. None of these were noticed on the delta
proper, and with the exception of the Engelmann spruce they seldom
occupy the areas that are often submerged. Of course in the lowest
places the meadow type of vegetation prevails, and elements of it
are scattered more or less through the area occupied by the trees.
As already shown, the lake formerly extended over the valley north
of it, and as its waters receded it is very probable that vegetation
similar to that found around its shores today occupied similar areas
around the shore line of the former lake. Of course as the gradual
lowering of the underground water level brings about unfavorable
conditions for those plants that exist around the shores, these are
replaced by those needing other conditions. On that portion of the
shore line that borders the area where the rainfall is insufficient to
support trees, the swampy areas will pass through the meadow, the
cottonwood-birch-aspen type, to the prairie. However, if the shore
line be in a region that has sufficient rainfall to support a forest, the
first two, stages will be followed by a western larch-Douglas spruce-
bull pine stage.

Since the Engelmann spruce is the most prevalent and the most
important tree in the formation just described, its sylvicultural habits
will be summarized. It has been shown that it does best in soils
where the underground water level is not far below the surface, that
is, for the plot under consideration, in low-lying lands bordering
lakes, ponds, or streams. Thus it occurs not only in Swan valley,
but also along the banks of Flathead River, where it is somewhat
sparingly mixed with cottonwood-birch-aspen groups. It is the first
tree to advance into the meadows. However, the Engelmann spruce
is not confined to the more or less swampy region, for it is found
scattered through the mesophytic areas in Swan valley. In the
stands that lie adjacent to the spruce forests it is almost always pres-
ent, and in some places it extends well out into the drier parts of the
western larch-Douglas spruce formation, being found beyond the
limits of the silver pine and lowland fir, and having a wider life range
than either of these two species. It was often observed in the bull
pine-Douglas spruce association where depressions in the topog-

200 BOTANICAL GAZETTE [MARCH

raphy brought the surface of the soil near the underground water
level, and along streams. However, trees in these situations seldom
reach large dimensions.

The Engelmann spruce can tolerate shade fairly well; in this
respect it is different from western larch and Douglas spruce and
is classed with silver pine and lowland fir. The fact that it can endure
shade probably accounts for the presence of its seedlings in the forests
that occupy the drier areas in the plot, for the forest canopy in these
situations brings about a local climate that makes it possible for trees
to exist that could not otherwise do so. The Engelmann spruce has
the power of germinating in the open if there is sufficient soil mois-
ture, and like the other shade-enduring species it does better in these
situations than in the shade.

It will be seen from the foregoing that outside the semi-hydrophytic
conditions Engelmann spruce does not flourish in the plot under
discussion, being confined to the level stretches of land bordering
on the meadows and open bodies of water. There is little doubt
that as the waters of the former Swan Lake withdrew, meadows first
occupied these areas and then were replaced by spruce stands. Just
as the meadow conditions may be considered the first stage in the
development toward the climax forest of the region, so the spruce
forest may be considered the second stage. As the channel of Swan
River is cut deeper the water level of the valley will be lowered, thus
bringing about a condition favorable to the development of other
types of forest which will replace the spruce association.

In the above account the distribution of the Engelmann spruce }§
discussed only in the area mapped. In the mountains bordering
the area on the east it shows a wide altitudinal range, but is always
associated with good moisture conditions, for it is found best devel-
oped in the damp cafions and on damp slopes where it is invariably
associated with Abies lasiocarpa. In the high basins it does not do
so well as the fir, but in the lower valleys it flourishes where the alpine
fir does not.

THE WESTERN LARCH-DOUGLAS SPRUCE FORMATIONS (MESOPHYTIC)-

In the discussion of the Engelmann spruce formation it was shown
that on its borders and on the dry hummocks in stands of this tree
other trees obtained a foothold. Just as there is a gradation from

:
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4

1905) § WHITFORD—FORESTS OF FLATHEAD VALLEY 201

the meadows into the meso-hydrophytic forest in which the Engel-
mann spruce predominates, so there is a similar change from these
into a forest in which the western larch and Douglas spruce are the
principal trees. Again, this difference is due to a change in the under-
ground water level; in the meadows it is near the surface; in the
Engelmann spruce stands it is slightly below the surface; and in the
western larch-Douglas spruce combinations it is still farther below.
Indeed, in this association the water level is so far below the surface
in places that the roots of the trees do not reach it, but rather depend
on the water in the surface layers that comes from the rains or by
capillarity from the underground water level. Since there is a medium
condition of water in the soil, the forests occupying these areas are
known as the mesophytic formations (fig. 8). Other trees found
with Larix occidentalis and Pseudotsuga taxifolia are Pinus Murrayana,
Abies grandis, Pinus monticola, Picea Engelmanni, Thuja _plicata,
and occasionally Abies lasiocarpa, Pinus ponderosa, and Tsuga
heterophylla; also certain deciduous trees.

Associated closely with the spruce and almost invariably sur-
rounding stands of it, except where the lodgepole pine is present, is
found a forest in which the silver pine and lowland fir are at their
best. Thus on the west side of the bay-like area of meso-hydrophytic
forest west of Ross Lake, the spruce is gradually replaced by a mag-
nificent growth of lowland fir, silver pine, western larch, and Douglas
spruce. The lodgepole pine occupies a good deal of the spruce
stands that would otherwise have developed into a forest like that
just mentioned, were it not for the influence of fires. In various
other areas are found forests in which silver pine and lowland ‘fir
form a more or less conspicuous element, especially in the area near
the mouth of Meadow Creek. They are never unaccompanied by
other species and seldom become the dominant trees, probably
because the climate is not so congenial to them here as to some of
the other dominant species.

The silver pine closely resembles its relative the eastern white
pine. As already stated, in Flathead valley it does its best in the
Conditions surrounding the spruce stands, or where the water level
1S not far below the surface. However, it spreads over nearly all the
mesophytic region in Swan River valley and ascends the east slope

202 BOTANICAL GAZETTE [MARCH

of the Mission Mountains in moist situations. In the swampy soils
silver pine has a shallow root system and is easily blown over by
winds, many large trees being observed lying prostrate with nearly
the entire root system exposed. Where the soil has sufficient moisture,
this tree can germinate in the open with no shade whatever, but it
can pass the seedling stage in rather shaded conditions. The only
two trees that have a shade too dense for it are the giant arborvitae
and the western hemlock. While it can exist in the shade around
the drier edges of the mesophytic forest, it fails utterly to maintain
a stand in the mature growth of these areas, and except in extremely
favored places the silver pine cannot be considered a successful tree
for the region under consideration. It succumbs easily to fires,
many instances being noted where fires had swept through the forests
destroying the silver pine, while such trees as western larch and
Douglas spruce were only slightly damaged.

The distribution of lowland fir is very much like that of silver
pine, both reaching their climax growth in about the same situations.
Where the water level is rather far from the surface it is more suc-
cessful than silver pine in the low altitudes; but its altitudinal range
is less than that of the latter. It is able to develop in open places,
in soil so dry that silver pine is excluded. Where the moisture was
sufficient it was not an uncommon thing to find stands of small (3
to 5™ high) trees so dense that absolutely all vegetation was lacking
beneath them. It is more tolerant of the shade than is silver pine, and
because it can grow in drier soils it extends as undergrowth into the
borders of the meso-xerophytic regions, where silver pine is seldom
if ever found. Like silver pine and Engelmann spruce, the range
of this tree is extended because of the protection of the forest canopy;
but this is not the region of its best development. The largest tree
seen were less than a meter in diameter and not more than 38" 7
height. SaRcEnr reports that it attains its best development along
the Pacific coast in Washington, Oregon, and northern California,
where it is frequently 75-90™ in height.

By far the most successful tree in Flathead valley is wester? larch.
It is a tree that closely resembles its eastern relative (Larix lari na)
in general appearance, although its habits are decidedly different
(fig. 11). As already shown, it is one of the first trees to gain a foot-

203

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hold in the spruce association, where it is found on the ridges and
hillocks, and it forms an important element in the neighboring
forests. There are extensive areas in Swan valley where this is
decidedly the predominating species, and on the west slope of the
Mission Range in some places there are almost pure stands. It
extends out into the prairie regions along the river courses and on
protected slopes. It does not grow in as dry soils as Douglas spruce
and bull pine, and hence is more restricted in its distribution in
regions where the rainfall is slight. The young trees, unlike those
of silver pine and lowland fir, cannot endure shade. In no instance
was a seedling observed growing in the shade of a forest. Other
things being equal, it does best in broad open places, although it
can do fairly well in slight openings where the sunlight enters at
least during the middle of the day. The western larch then does not
tolerate shade and cannot reproduce itself unless an opening be
made first in the forest. This may be done in the mature primeval
forest by the dropping out of old trees, but more frequently accidents
such as fires and winds bring about open places suitable for its repro-
duction, provided of course seed-bearing trees are left to stock the
soil with seeds. In contrast with silver pine and lowland fir, it may
_be called a fire-resisting tree. Its thick bark enables it to withstand
fires that would kill silver pine and lowland fir, old trees often being
found still living in the midst of burned areas. Sometimes isolated
trees stand towering over the young lodgepole pine growth that has
come in after fires (fig. 12), and the only young trees competing with
lodgepole pines are western larch.

Observations on the rooting habits of trees are in any case difficult
to obtain, unless windfalls are frequent. Many trees of weste™
larch were found standing in windfalls where silver pine, lowland
fir, and Engelmann spruce had been blown over. From this on¢
would judge that its roots penetrate much deeper than the roots of
these trees; indeed this may be one of the reasons why western larch
is able to exist in drier areas than can some other trees, for 4 deep
root system will enable it to reach nearer to the underground water
level or at least below the superficial dry surface layers where 4 tree
with shallow roots could not get sufficient moisture.

Since Douglas spruce is a successful tree in the regions bordering

1905] WHITFORD—FORESTS OF FLATHEAD VALLEY 205

on the prairie, a discussion of its sylvicultural habits will be deferred ~
until the meso-xerophytic forest is described. It is almost invariably

; - —Three generations of forests in a “burn” near Ross Lake; first genera-
tion represented by isolated old specimens of western larch; second generation con-
sists of a forest of lodgepole pines mixed with a few individuals of western larch; in
the foreground is the third generation, which consists almost entirely of young trees

of lodgepole pines.—From photograph by PRAEGER.

4 companion of western larch in the mesophytic area, hence the name

Western larch-Douglas spruce association.

206 BOTANICAL GAZETTE [MARCH

The giant arborvitae is another tree that needs brief mention here.
In the plot under discussion, along the base of the Swan Range
there are three isolated places where the arborvitae was noted. It is
usually found in moist places and probably ought to be considered
as a meso-hydrophytic rather than a mesophytic tree (fig. 13). West
of the Cascade Mountains, where it reaches its greatest development,
it is distinctly a tree of the moister regions.

Another tree found in Swan valley is alpine fir, mentioned before
as associated with Engelmann spruce in the moist cafions of the
mountains. From here it spreads to the subalpine regions where

it occupies a prominent place in the basins where snow lies all the

year. In places it also finds its way into the lower altitudes in rather
moist situations, as in Swan valley. The birch maintains a place
throughout the mesophytic area; especially is it frequent in the
moister situations and mixed with the lodgepole forests.

The soil in the mesophytic area is variable in nature, the character
of the vegetation changing to a certain extent with the variations.
It is probably not the chemical composition that determines the
kind of forest, but rather the physical composition, the capacity of
the soil for holding water being the controlling factor. If the under-
ground water level is not far below the surface, it makes little differ-
ence whether the soil is sand, clay, or gravel, so long as there is
sufficient humus to furnish the needed nitrates. However, back
from the low-lying lands, where the water level is too far below the
surface to be reached by the roots of the trees, the physical character
of the soil plays an important role. As a rule the soil of the valley
is made up of clay, probably derived from the decay of the dolomitic
shale, the principal rock of the surrounding mountains. This has
been washed in and deposited in the bed of the former more extended
Swan Lake, and is mixed more or less with silt. Humus has acct
mulated in places, and this is more abundant where fires have nom
been so prevalent; for not only do fires destroy the humus, but 10
the open places left after the destruction of the forest it dries out
rapidly. In contrast with the rich beech and maple woods of the
eastern United States, the humus content of the soil is considerably
less in these coniferous forests. This is of course due to the fact that
the needle leaf of the conifers is not so good a humus producer as the

207

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208 BOTANICAL GAZETTE [MARCH

broad leaves of the maple, beech, and other deciduous trees. In
places, however, considerable quantities of humus are present, espe-
cially where the trunks of fallen trees have been allowed to decay
and have not been destroyed by fire. A very good indication of a
soil rich in humus is the number of species of fungi and mycotrophic
seed plants found in it; Monotropa uniflora, Pterospora Andromeda,
and Corallorhiza multiflora being scattered rather thickly throughout
the forest. However, one misses the rich array of fleshy fungi found
so abundantly in the leaf mold of the deciduous forests. Except in
those places where arborvitae, hemlock, and lowland fir are prominent,
the shade is not very dense; and under the rather thin canopy of the
other conifers there is enough light to permit the development of a
number of semi-shade species.

A woody plant that gives a decided tone to the undergrowth
throughout the mesophytic area and to some extent in the meso-
xerophytic area is the dwarf maple (Acer glabrum), the only repre
sentative of the maples found in Flathead valley. It can hardly be
called a tree, for it seldom becomes more than 3™ in height. Other
characteristic shrubby plants are Salix sp., Philadelphus Lewisti,
Holodiscus ariaefolia, Amelanchier alnifolia, and Sym phoricar pos
sp. In some places, usually more mesophytic, Mensiesta urceolaris
and Taxus brevifolia are found. Some of the forms, like snowber'y,
service berry, and mock orange, become more frequent as the forest
becomes more open. Among the prominent more lowly forms were
noted the following: Berberis aquifolium, Aralia nudicaults, Cornus
canadensis, Chimaphila wmbellata, Pyrola rotundijolia, Linnaca
borealis, Rubus parviflorus, Clintonia uniflora, Adenocaulon bicolor,
Tiarella unijoliata, Lycopodium sp., and Disporum sp.

At the south end of Swan Lake, on the west side of Big F
River, is an area with a soil decidedly pebbly, and probably of glacial
origin (map), its inability to hold water being indicated by the chat
acter of the forest it supports. The Douglas spruce is about the only
mesophytic tree found here, and in places there are groves of bull
pine. On the east side of the river, so persistent is the absence ’

ork

other species than Douglas spruce and bull pine, that the area W#*
mapped as meso-xerophytic forest rather than mesophytc- ie

rather open, and the vegetation in it becomes somewhat prairie-

1905] WHITFORD—FORESTS OF FLATHEAD VALLEY 209

indeed everywhere that the mesophytic forest grades into the Douglas
spruce-bull pine combination the forest floor has prairie plants rather
than forest forms.

It has been shown that the western larch-Douglas spruce associa-
tion is found in those areas where the amount of water in the soil
is less than in the areas that support the spruce formation. Again,
attention must be called to the fact that in the former extension of the
lake many parts of the valley now having mesophytic forest were
moist enough to support spruce forests, and as the water of the lake
receded the amount of water in the soil became less and less until
the encroaching mesophytic forests crowded out the spruce forests.
As there is a relation in origin, therefore, between the meadows and
the spruce forests, so there is also a genetic relation between the
spruce forests and the western larch-Douglas spruce forests. In
places where the hills end precipitously near the lake shore, as on
the west side of Swan Lake for instance (map), this relation is not
So apparent, for the mesophytic forests reach in many cases to the
water’s edge. Yet even here narrow, flat areas at the foot of the
Mountains on the borders of the lake have small meadows with scat-
tered Engelmann spruce and cottonwood in them. These will give
Way, of course, to the encroaching mesophytic element when the
waters of the lake have further withdrawn.

In conclusion it may be pointed out that the climax forest of the
plot under consideration is the western larch-Douglas spruce for-
mation. This association occupies a greater portion of the com-
paratively level stretches of Swan River valley (fig. 8). It also is
found on the protected slopes and foothills of the low Swan Range.
No attempt was made to determine its range outside of the region
mapped. The trees that are characteristic of this forest are western
larch, Douglas spruce, silver pine, lowland fir, Engelmann spruce,
and lodgepole pine.

THE DOUGLAS SPRUCE-BULL PINE (MESO-XEROPHYTIC) AND THE
PRAIRIE (XEROPHYTIC) FORMATIONS.
Since the Douglas spruce-bull pine associations and the prairie
are more or less connected, they will be treated together. A refer-
ence to the map will show that a rectangular plot of ground known

210 BOTANICAL GAZETTE [MARCH

as Nigger Prairie lies just north of Swan Hill, at present used for
farming purposes. It is bordered on the south, east, and west by

narrow strips of pure bull pine forests, and on the north a forest of

‘fh
3

poten kane tae tan
«

? org | a a

in the center is 4 /aT8

las spruce.
— i

Fic. 14.—View of bull pine forest south of Echo Lake;

bull pine, the younger growth around consists of bull pine with some Doug

—From photograph by PRAEGER.

a ae ade as - ine
the same species extends to Echo Lake ( fig. 14). This bull ps 3
See oo toe Si eR
stand is surrounded on all sides by a mesophytic forest 1n wht
Tecteany ¢ ~} _ * F ‘ = ne.
western larch and Douglas spruce almost entirely replace bull p!

1995] WHITFORD—FORESTS OF FLATHEAD VALLEY att

The question arises, what is the cause of this difference in the char-
acter of the vegetation in so limited an area? So far as could be
ascertained, trees have been absent from Nigger Prairie for at least
twenty years, and it is very probable that it has never been forested.

The character of the soil in both the prairie and the bull pine zone
surrounding it is decidedly different from that of the country im-
mediately adjacent. That to the south and west is a residual clay
from the dolomite shale of Swan Hill; that to the east is a clay rich
in humus; and that to the northwest, north, and northeast is a pebbly
clay of glacial origin. On the other hand, the soil of the entire
prairie and bull pine region is composed of sand mixed in places
with a considerable amount of humus. The limits of this sandy
area extend a little beyond the limits of the bull pine forest. It
seems very probable that the absence of any other trees than bull
pine is due chiefly to the physical character of the soil. It must be
remembered that the climate of the area under discussion is such as
barely to favor a forest, and a slight difference in the amount of
water which the soil can hold may lead to the exclusion of certain
trees, and in places of all trees. Since a sandy soil lets water drain
through it readily, it is very likely that this area is too dry to support
a mesophytic forest. As shown by wells, the underground water
level in Nigger Prairie is some distance below the surface, so that
those trees which have shallow root systems would not be so likely
to succeed as those whose root systems are extensive and deep. The
bull pine is one of the latter kind, and it is very likely that this is one
reason why it has the habit of growing in drier situations than some
of the other trees.

If the surface layers of a sandy soil can be made to retain water,
the objectionable feature of such a soil will be partly overcome, and
this can be brought about by an accumulation of humus. Lower
forms of vegetation, even grass, by their decay will add to the humus
content of the soil. In very dry countries this accretion would be
exceedingly slow, for the oxidation of the organic débris would be
great; but even in dry situations by the accumulation of a slight
amount of humus the water-holding capacity of the soil might be
increased sufficiently to tide a few trees over the driest months of
the year. Trees once established in this way would help accumulate

212 BOTANICAL GAZETTE [MARCH

humus more rapidly, for by fall of the dead limbs and leaves more
organic material would be added than from the decay of lower forms
of vegetation. The shade of the trees thus established would check
the rapid oxidation of the organic material that takes place in the
open. The ultimate effect would be the establishment of a forest
in a place that was formerly occupied by a prairie. It is very probable
that the whole yellow pine area just described was at one time
occupied by a prairie vegetation. There is evidence that the present
prairie island in this forest would become covered with forest trees
if not interfered with by man, for along the roadside groups of young
growth of bull pine seedlings are not uncommon. All around the
border of the prairie young trees are present, and even in a timothy
field there were noted seedlings that had escaped being cut by the
mowing machine. Before the prairie island was settled it is very
probable that surface fires swept it occasionally and thus checked
the conquering of the prairie by the forest. Another prairie area
similar to the one just described lies directly west of Echo Lake
(map). Again, west of Nigger Prairie, just across the low range of
Mission Mountains, is another ‘‘sand pocket” that has a prairie
vegetation. These however border on the prairie climate.

The question arises whether the presence of bull pine will in any
way change the conditions sufficiently to make them capable of sup-
porting other species of trees. It was shown that the humus content
of the soil increases more rapidly when trees are present than when
they are absent. With the gradual increment of organic decay,
it is very possible that the water-holding capacity of the soil may
be increased to such an extent that it will be rendered capable of
supporting other species of trees.

A bull pine seedling does not tolerate shade; the young tree must
have broad open places where the sun strikes the soil a greater por
tion of the day. An opening in the forest made by the removal -
one or two trees does not allow sufficient light and heat to enter;
thus young stands are excluded from anything like a tolerably thick
growth of these trees. This is not the case with western larch and
Douglas spruce; it has been shown that the former does not tolerate
shade, but grows in slight openings in the forest; the latter 1 iti
similar in this respect to western larch. If humus has accumulate

1905]

W HITFORD—FORESTS OF FLATHEAD VALLEY

the

a few years ago this w

Lake, showing submerged cabin and timber land;

‘ of Echo
trees in the foreground are western larch and Douglas spruce.

arm

as dry land;

214 BOTANICAL GAZETTE [MARCH

so as to give the soil a sufficient water-holding capacity, open places
in the bull pine forest may permit seedlings of both the species named
to thrive. Numerous instances of this were noted in the bull pine
forest under consideration. In places where bull pine had been
removed there were clumps of young trees of Douglas spruce and
western larch. If the place is sufficiently open, bull pine seedlings
are found with the others. It will be seen readily that such a condition
will lead to a gradual replacement of the bull pine forest by the
Douglas spruce-western larch combination. There is some evidence
that such a succession has already taken place, for all around the
borders of the “sandy pockets” western larch and Douglas spruce
are more numerous in places than bull pine. There is also reason
to believe that in the pebbly clay morainic soil around Echo Lake,
especially to the south and northwest, a more or less pure bull pine
stand preceded the present stand consisting of a mixture of the three
species. The bull pines are found in groups, or solitary full-grown
specimens overtop the younger growths of the next generation of
Douglas spruce and western larch, these isolated specimens doubtless
representing all that is left of the former pure bull pine stand. Of
course this succession has been a gradual one, occupying many gel
erations, the more successful trees elbowing their way, as it were,
among the less successful until the spread of the latter is limited by
their incapacity to adapt themselves to the new environment. All
stages in this succession are present today in the forest under dis
cussion, and it was the study of these stages that led to the conclusion
that has been advanced.

Out towards Flathead River the zone between the prairie and the
forest contains a few scattered specimens of bull pine; next to this
there is a more dense stand of this species with a few scattered trees
of Douglas spruce; back farther still from the prairie is a zone
which bull pine is less and Douglas spruce more frequent; and
finally another condition is found in which western larch joins these
two species, bull pine being the least conspicuous element of the
three. Such an arrangement is approximated just west of Echo
Lake. This region is decidedly morainic in its character, and wher?
depressions approach the underground water level western Jar
becomes more numerous, and in places near Echo Lake the surface

i

eee ee

a

or ee en a ae oe oe

SS Ee NE ed

Oe eT ee

q
Fs.
:
a
; "i

1905] WHITFORD—FORESTS OF FLATHEAD VALLEY 215

of these depressions comes near enough to the water level to permit
the growth of scattered Engelmann spruce. This level of under-
ground water varies, and this change is shown in the rise of the lake
itself and the ponds around it (fig. 15).

Mention has already been made of the western larch-Douglas
spruce combinations on the protected slopes of the Mission Moun-

16.—View on the top of the Mission Mountains showing area on which the
Douglas spruce-bull pine forest has been destroyed by fire; the outcrop of dolomitic
Shale has been scoured smooth by the action of glacial ice; Ceanothus bushes on the
right, lichens of various sorts on the rocks, and prairie grasses where the soil is suffi-
ciently deep.—From photograph by PRAEGER. .

tains. Toward the top of this slope western larch becomes less and
Douglas spruce more prominent; in places the latter is mixed with
lodgepole pine. On the west side of the range Douglas spruce and
bull pine are the principal species; hence this area has been mapped
4S Meso-xerophytic, although western larch is present in some
situations.

216 BOTANICAL GAZETTE [MARCH

The altitude of the Mission Range within the limits of the plot
is nowhere more than 1372™ above the level of the surrounding
country. The core of the mountains is composed of a dolomite
shale, which outcrops in many places. Especially is the rock near
the surface on the steep northwest fault face and where the soil has
been removed by the action of glaciers (fig. 16). South of the river
the forest on the western slope has been partially destroyed by fire
(figs. 16 and 17), but enough of the original stand remains to give
a clue to the conditions of the whole slope. A discussion of the burned
area will be taken up in another connection.

Wherever the rock outcrops there is little or no vegetation (jig.
16), due to the fact that there is no soil; the pioneer plants on the
rocks here, as elsewhere, are the lichens. The crustaceous lichens
appear where there is absolutely no disintegrated rock; sometimes
associated with these are the foliaceous forms, but these are more
common where a little soil has accumulated; with these are found
mosses adapted to the dry conditions. After more soil has accumu
lated, conditions are favorable for plants that require more moisture;
among these are the fruitcose lichens, Cladonia rangijerina sylvatica
being a good example. Associated with these lichens are Selaginella
densa, Sedum Douglasii, Heuchera parvijolia, the last being usually
found rooted in cracks. These plants prepare the way by their decay
for the higher forms that can get a foothold as soon as sufficient soil
is present. The inorganic material made by the disintegration of
the rocks is blown or washed in from the surrounding region and
reinforces the organic accretion. In the meantime the rock crevices
have been filling up with soil, thus furnishing favorable places ae
shrubs like Prunus demissa, Symphoricar pos sp., Amelanchier alm
folia, and Juniperus sibirica; and Campanula rotundifolia, Arc
siaphylos Uva-ursi, and grasses are early pioneers in these places.
If the outcrop is near a large body of water like Flathead Lake,
Juniperus scopulorum is one of the first trees. This tree is found
distributed around the shores of the lake and along water courses
in the prairie region in the valley at the foot of the lake, and with ™
are found Douglas spruce and bull pine. These of course g4!? 1g
foothold by sending their roots into the rock crevices and may play
an important réle in the weathering processes by prying the ate

a4

VALLE

FLATHEAD

OF

WHITFORD—FORESTS

ay
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189] |

I “You JO S}so10j

218 BOTANICAL GAZETTE [MARCH

apart in the growth of their roots. However, on the rocks there is
never more than a scattered growth of trees. Where the soil is less
shallow, more trees are present and a more closed stand is developed.
In the ravines and other situations where a deep soil has accumu-
lated, a heavy growth often occurs; especially is this true in the
protected ravines and on the sheltered slopes. Along the front of
Flathead Lake there is a series of ridges which has an open growth
of Douglas spruce and bull pine on the side facing the lake; and
protected from winds, the slopes on the east side of these ridges and
the valley between them and the higher ridges to the east have in
addition western larch, which in places is the dominant tree. Again,
where the surface of this valley is not far above water level, Engel-
mann spruce and its associates are found, though these places are
infrequent. In many protected areas on the west side of the Mission
Range western larch is present, though as contrasted with the eastern
slope the exposed western face of the mountains is not so favorable
a habitat for the mesophytic trees.

The temperature conditions on top of these mountains and at the
foot are not known. No doubt the climate is considerably cooler
at an altitude of 1372™, the highest point of the range in the plot,
than it is at 8g90™, the level of the lake. However, this difference 15
not enough to exclude the lowland species and introduce those found
on the higher Swan Mountains to the east. The presence of a large
body of water like Flathead Lake no doubt influences the tempera
ture conditions along its shores, for early fall frosts are less frequent
here than at a short distance inland. Whether or not this affects
the forest growth is not known.

[To be concluded.]}

BRIEFER ARTICLES.

TERMINOLOGY OF THE SPORE-STRUCTURES IN THE
UREDINALES.:

SOME time ago I began to study groups of species among the rusts in
a more methodical way than usually pursued, and as part of the method
undertook to prepare uniform descriptions in which like parts of each
structure should receive the same kind of treatment in every case. I soon
met with an obstacle that had not been anticipated. The terminology in
use was found to be clumsy, ambiguous, poorly correlated, and quite inade-
quate to show homologies properly.

I will illustrate a few of the difficulties which confronted me. When
an aecidium is mentioned, there is a tolerably clear conception of a cup-
like structure filled with spores; but when one reaches the next stage of
the rust, what is the meaning conveyed by the word uredo? I think it is
most generally employed as a collective term, referring to the second stage
of the rust. We can properly say that the uredo occurs on leaves of a
certain host, but if we wish to specify a single uredinial structure correspond-
ing to an aecidium, it is necessary to say uredosorus; and yet it is not per-
missible to say aecidiosorus. Passing to the third stage of the rust there
appears to be neither a specific nor a collective name with which to desig-
nate it. The nearest approach to a specific term analogous to aecidium is
teleutosorus, and to a collective term analogous to uredo is teleutostage.
We can Say properly that aecidia appeared on a leaf, after a time they were
followed by the uredo, and finally the teleutospore stage was developed;
in which statement we have used three different methods to express the
same idea relative to similar spore-structures.

Taking up the first stage again, there appears to be nothing inappro-
priate in calling the single spore-structure an aecidium so long as we have
in mind the first stage of species belonging to the genus Puccinia or Uro-
myces; but in the genus Phragmidium there is no peridium present, and in

ymnoconia not only is the peridium absent but the spore-layer is extended
and indefinite, and the term aecidium now seems far less applicable. For
such cases it is customary to call the structure a caeoma, instead of an

* Read before the American Mycological Society at Philadelphia, December 30,
04. '

1905] 219

220 BOTANICAL GAZETTE [MARCH

aecidium, adopting the generic name established by TULASNE in 1854,
who applied it to forms without a peridium and having the spores in chains.
But in the genus Coleosporium it is the uredostage, and not the aecidium,
that possesses the caeoma structure. Moreover, care must be exercised
not to confound TULASNE’S caeoma with Linx’s caeoma. For in 1809
Lryk established a genus Caeoma to cover pulverulent forms of rust,
whether the spores were borne singly or in chains, which was extensively
used by mycologists of the time and subsequently, including SCHWEINITZ
and others in America. But the terminology for the first stage of a rust
is not yet exhausted. If the stage occurs on a pomaceous host, one is
usually safe in calling it a roestelia; and if on a coniferous host, it is usually
styled a peridermium. But there are plenty of instances among the less
common forms of rust where none of these many terms seems accurate.

Turning to the second stage of rusts fortunately we do not find so many
terms in use. The spores are usually borne singly on pedicels, and are
generally called uredospores, or when borne in chains they are caeomo-
spores. If the sorus happens to be surrounded or filled with prominent
paraphyses, the spores become epiteospores, although this term is derived
from a generic name first applied to an aecidial form. It has seemed to
me providential that when the uredosorus is surrounded by a peridium, as
in species of Pucciniastrum and Cronartium, we are not required to have
some other sort of spore, and are permitted to say simply uredospore.

I have only cursorily mentioned some of the most obtrusive difficulties
encountered, when I undertook seriously to record comparative studies of
the rusts. I should probably never have publicly rebelled against this
confusion of superfluous and yet inadequate terms, had I not after a time
become interested in generic nomenclature. So long as there seemed to be
no impropriety in allowing a generic name to wear out and be cast aside,
or to be subjected to the menial duty of designating a form of spore, It
seemed unreasonable to make complaint. But when the conviction
became firmly established that a uredineous genus Was entitled to as much
dignity of position as the genus in any other group of plants; and when It
became evident, moreover, that in the application of the rule of priority
some of these degraded genera might suddenly assume the place usurped
by long accepted names, as for instance, Kuntze would have us believe
that Roestelia should take the place of Gymnosporangium, then I thought
* ae - look into the possibility of emending, changing, or in some "ey
improving the terminology of the spore-forms. Pursuing my generic
studies further, it appeared little short of absurd that one of the earliest
genera, and until recently one of the largest, which has embodied the mos .

Ne ae ee eee Pe ee

ae ae

1905] BRIEFER ARTICLES 221

prominent idea of the order both in the scientific mind and in the popular
mind, and which supplied the accepted name by which the order is now
known, I mean the genus Uredo, should be wholly discredited and aban-
doned, and according to many able mycologists even lose its rights in synon-
ymy. When I came to believe that the names Uredo, Aecidium, Roes-
telia, Peridermium, Epitea, and Caeoma were entitled to rank as accept-
able genera, provided they were not antedated, it became certain that these
names should no longer be made to do duty as spore-names.

For the several reasons which I have now partially illustrated, I was
impelled to seek for more serviceable, accurate, and concise names for
uredineal spore-structures. More than a year ago I presented the first
installment of my problem to Dr. FREDERIC E, CLEMENTS, of the Uni-
versity of Nebraska, and it is to his continued and hearty cooperation and
suggestion that I am able to present at this time a well considered series
of terms to designate the several spore-structures of the Uredinales. I
think the claim can be justly made that these new terms, all constructed
by Dr. CLements, will do away with ambiguity, have a definite and easily
recognized application, permit of uniformity of statement for the several
Spore-stages, and promote clear conceptions of homologies.

The terms I have to propose apply to the sorus. By sorus is meant
the structure which arises from a single, fertile hyphal mass, or hymenium,
either with ot without a peridium, now usually called spermogonium,
aecidium, uredosorus, teleutosorus, or kindred names. A compound sorus
is produced by a number of sori standing close together, separated and sur-
rounded by modified hyphae, which form a stroma, erroneously termed
paraphyses, e. g., the teleutosori in Puccinia rubigovera and Puccinia
Virgaureae. The stroma is not considered a part of the sorus, but an
adjunct structure. A simple sorus is surrounded by the tissues of the
host, or by unmodified mycelium, and includes the peridium, and all true
paraphyses, whether peripheral or discal.

The new terms consist of four words, with their derivatives, one for each
of the four stages of uredineal fungi. For the sorus of the initial stage,
usually designated by a cipher, and called spermogonium, pycnidium,
etc., I propose pycnium (mvxviov); derivatives pycnial, pycnios pores, etc.
For the sorus of the first spore-stage, usually designated by the Roman
numeral I, and called aecidium, roestelia, peridermium, etc., I propose
aecium (aixiov); derivatives aecial, aeciospore, etc. For the sorus of
the second spore-stage, usually designated by the Roman numeral II,
and called uredosorus, etc., I propose uredinium (uredo); derivatives
wredinial, urediniospore or if preferred uredospore, etc. For the sorus 0

222 BOTANICAL GAZETTE [MARCH

the third spore-stage, usually designated by the Roman numeral III, and
called teleutosorus, I propose felium (rédiov); derivatives telial, telio-
Spore, etc.

These four words and their derivatives have sufficient resemblance to
corresponding terms now in use to suggest their meaning, and thus facil-
itate their introduction; and yet they are all new words and are now brought
before the scientific public for the first time. All the words are shorter
than the corresponding ones now in use, except urediniospore, which is
longer by three letters than uredospore, but no more difficult to pronounce,
and for this the word at present in use, uredospore, may well be retained.
No permissible shortening of the stem of this term has seemed feasible,
and yet have the uniformity and classicism of the series preserved.

I may be permitted to illustrate the use of these words by a similar
sentence to that employed at the beginning of my paper. In the case of
wheat rust, aecia appear on the leaves of the barberry in spring, preceded
by pycnia; the aeciospores give rise in due course of time to uredinia on
the wheat plant, followed by telia. A similar statement could be made
for a Coleosporium, Phragmidium, or any other rust. One might explain
the difference between the telial stage and the uredinial, or discuss the
status of pycniospores as compared with teliospores or other sorts, oF he
might point out that the sorus of a uredinium has the same essential
structure whether without peridium and stylosporic, as in Puccinia; with
peridium and stylosporic, as in Pucciniastrum, or without peridium and
the spores in chains, as in Coleosporium; and so on.

Finally, the acceptance of these terms would require that the following
terminology for the receptacles, spores, etc., of the rusts should be sub-
stituted for the series recently given by Saccarpo in his NV omenclatura
mycologica, viz., sorus, pycnium, aecium, uredinium, telium, peridium /
stroma. Mesospore, amphispore, stylospore, etc., are special descriptive
terms, which may be extended indefinitely as need arises, but do not form
part of the general category. Caeomospore, epiteospore, aecidiospores
and other terms compounded from generic names should be wholly dis-
continued. Peridium is to be preferred to pseudoperidium, in accord
with the usage of DECANDOLLE, LEVEILLE, and many recent mycologists-
Brevity, directness, and accuracy are excellent qualities to keep in mind
- when deciding upon terminology.—J. C. ARTHUR.

UE UNIVERSITY,
Lafayette, Ind.

CURRENT LITERATURE.
BOOK REVIEWS.
Plant chemistry.

Many BooKs have been written on physiological chemistry, but for the most
part these deal specifically with substances which are of chief interest to the
animal physiologist; and generally they undertake to treat the characteristics and
modes of reaction of a very great number of individual substances more or less
independently of their biological relations. The book before us has a definite
ai

of departure for further researches. The field of plant chemistry has been so
neglected, comparatively, that scarcely more than ordinary industry suffices to
make notable advances in knowledge.

Such a book, of course, will be of value to many other than plant physiol-
ogists. The service which CzapexK has done is likely, therefore, to have wide
and hearty appreciation. The book is not intended for the beginner in bio-
chemics, but assumes a general knowledge of both botany and chemistry, being
intended specifically as a reference book and repertory of literature for the help
of investigators of special problems.

The book opens with a brief historical introduction, in which the author
touches lightly yet clearly the service of chemists of the Middle Ages and of those
who labored after the transformation which chemistry underwent through the
genius of Lavorsrer. The work then is divided into two parts, general and
Special. The general part occupies something over ninety pages, and is of
decided interest to the reader who is familiar enough with modern chemistry
to appreciate this brief and illuminating discussion. The author considers the
nature of the protoplasm and the substances of which it is composed; the prop-

: erties of colloids, the structure of the protoplasm and its biochemical signifi-
cance; the conditions and rate of chemical reactions, and especially the ionic
reactions in living cells. The last three sections, in which are discussed catalysis
and catalysers, the general chemistry of enzymes, and the cytotoxines and similar
compounds, deserve particular notice.

The special part of the present volume is confined to a consideration of fats,
lecithines, phytosterines, waxes, and the carbohydrates. To the last named much
t vg larger part of the volume is devoted. Each of these classes of materials is
discussed in its relations to various groups of plants, or to the various plant parts
1 which it is found. In this feature we have the most radical difference between

* Czarex, F., Biochemie der Pflanzen. Bd. I. Imp. 8vo. pp. xv+584. Jena:
Gustay Fischer. 1904. M 14.

1905] 223

224 BOTANICAL GAZEITE [MARCH

this book and most of the previous works on physiological chemistry, and even
of the few that have been written with special reference to plant chemics.

A second volume is already in press and may be expected ere long. We
suppose that a detailed treatment of the proteids will occupy the greater part
of it. It will be accompanied by full indexes.—C. R. B.

Plant organization.

In 1899 we noticed the first part of a work by BERTHOLD on ~_ “—
zation,? of which the first half of the second part, promised “very soon,” has
recently appeared.s The author announced his intention of Pander in this
second part in a comprehensive way the investigations made by himself and others.
This promise rests in large part unfulfilled.

nd an “introduction” of twenty-one pages (which seems queerly out
of place in the middle of a book) setting forth the aims and methods of this “Phys-
iologie der pflanzlichen Gestaltung.”” BrRTHOLD assumes that the protoplast is
a mechanism, made of different individual parts, even though they cannot be
distinguished, each having its particular function. This assumption is developed
at length. Of course, therefore, he objects to the idea of adaptation in connection
with anatomy, especially as developed by HaBERLANDT. He rejects the ideas
‘of gro enzymes and of Anlagen as indicating useful directions for research,
and minimizes the value of studies on regeneration in the problem of plant
organization.

In the special part (pp. 22-257) BERTHOLD discusses in four chapters the
morphology of the typical (sic) shoot, the pith, the primary cortex, the course of
development in the pith and cortex, and entitles the fifth “a comprehensive synopsis
of the development and rhythm of the shoot.’ These chapters make almost
no reference to the “mechanism” on which he lays so much stress in the in
duction, and certainly do nothing toward revealing it. Profuse details encum cumber
the pages; indeed, in 1 sain it oe seems as though the author had published
his notebook. The “synopsis” covers 136 pages, a formidable number for 8

“glance” to take in. It is so beclouded with details and so unorganized -
one cannot readily discover what the author is driving at. Great stress is laid
throughout upon the cell-contents as determining factors in the final configuration
of tissues, yet there is no indication of the nature of the relation, and su
categories as “‘starch,” “sugar,” and “tannin” are treated as though the

Picturing clearly either the anatomical or physiological features of plant —
zation. Perhaps the — instalment will sum up his views in @ definite a”
precise fashion.—C. R

? Bor. Gaz. 27:146. 1899.

n.
3 BERTHOLD, G., Untersuchungen zur beige der pflanzlichen Org rigs
8vo. pp. iv+257. Leipzig; Wilhelm Engelmann. 19

CURRENT LITERATURE 225

MINOR NOTICES.

THE TENTH AND ELEVENTH parts of RorH’s+ Die européischen Laubmoose
complete the second volume and the work. Supplements to both volumes and
indexes to accepted names and synonyms in the second volume make up the
larger portion of the eleventh part. The four indexes (two in the first volume)
should have been combined into one for convenience, though we are thankful
for full indexing of any kind. The author and publisher deserve special com-
mendation for the promptness with which the work has been issued. The book
will be almost as useful to American as to European bryologists, and the total
price, 44 marks, for a work with 62 plates and nearly 1400 pages is cen
very reasonable.—C. R. B.

Major Squier,’ of the Signal Corps, U. S. A., in experimenting with field
Wireless telegraphy, has incidentally paid some attention to the conductivity
and electromotive force of live trees. Being, naturally enough, unfamiliar with
the literature, he has rediscovered a number of well-known phenomena and raises
Some questions which have already been investigated. The discharge of a
powerful induction coil and Leyden jars through live-plant electrodes, and the
so-called ‘“‘floral spectra” with lines of carbon, nitrogen, sodium, etc., are more
novel than valuable.—C. R. B

Die Pflanzenjamilien drags on, the sections on mosses and lichens not yet
being complete. The former are being elaborated by BRoTHERUS.® The last
two parts to appear (219, 220) treat the remainder of the Bryaceae, the mono-
typic Leptostomaceae, Mniaceae (72 spp.) Rhizogoniaceae (35 spp.), Aulacom-
niaceae (11 spp.), Meesiaceae (11 spp.), Catoscopiaceae (monotypic), Bartram-
iaceae (400 spp.), Timmiaceae (10 spp.), Weberaceae (13 spp.), Buxbaumiaceae
(2 spp.), eansng ( : spp.), and Georgiaceae (5 spp.). The Polytrichaceae
are just begun.—C.

Brat? has ae what is really a third edition of the list of vascular
plants that bears the title “Michigan Flora.” The first edition (1880) was pre-
pared by C. F. WHEELER and Erwin F. Situ; the second (1892) by C. F.
WHEELER and W. J. Beat; and this third edition bears only the name of Pro-
fessor Brat. Besides the incorporation of additional plants, the most notable
————

4RortH, G., Die europdischen Laubmoose. 2 Band, Lief. ro, 11. Imp. 8vo
a8 et rk = 41-62. Leipzig: Wilhelm Engelmann. 1904. Each M 4. Parts
not sold
5 SQUIER, gee O., On the absorption of electromagnetic waves by living
vegetable organisms. 8vo. pp. 32. Reprinted from Maj. Gen. Arthur MacArthur’s
Report to War De ept. on Military Maneuvers in the Pacific Division, 1904.
° ENGLER and PRANTL, Die natiirlichen Pflanzenfamilien. Lief. 219, 220. pp.
377-672. ay! ba in 37. Leipzig: Wilhelm Engelmann. 1904. M 6 or 12.
7 BEAL V. J., Michigan Flora. A list of the fern and seed plants growing
Without ee Reprinted from 5th Report of the Mich. Acad. Sci. 1904.

226 BOTANICAL GAZETTE [MARCH

change in the present edition is the use of the Engler and Prantl sequence as
presented in Britton’s Manual.—J. M. C.

THE SECOND FASCICLE of L&VEILLE’s? monograph of Oenothera has appeared,
the first fascicle having been noticed in this journal for April 1903 (p. 296).
Naturally it is of great interest to American taxonomists, and contains a profusion
of illustrations. The genera Eulobus and Gayophytum have been merged under
Oenothera, and O. bistorta Nutt. has been replaced in great part by O. chieranthi-
jolia Hornem.—J. M. C.

THE FOREST WEALTH OF OREGON is the title of a small pamphlet prepared
by E. P. SHELDON? setting forth the forest resources of the state, with a list 0
forest trees and larger shrubs. Of these thirty-eight are gymnosperms.—C. R. B.

In conTINUING his revision of Eucalyptus, MAIDEN’? presents E. stellulata,
E. coriacea, and E. coccifera, under each species giving the description, synonyms,
range, and affinity.—J. M. C.

THE EIGHTH VOLUME of the fourth series of HooKEr’s Icones Plantarum
closes with the publication of the fourth part January 1905. The plates of this
part are 2776-2800.—J. M. C.

NOTES FOR STUDENTS.

HESSELMAN™ has published a paper on Swedish meadows, which should
be carefully read by all who are engaged in ecological research, since no recent
paper has gone more fundamentally or successfully into the real problems of
ecology. The studies have been carried on for nearly a decade, and in a rather

imited area in the neighborhood of Stockholm, and with especial detail on the
little island of Skabbholmen. The “Laubwiesen” are meadows in which there
are scattered deciduous trees, the general aspect being park-like. They might
perhaps be regarded as edaphic savannas. ‘These formations are rich in herbs
and ses, and are essentially without low shrubs. At an earlier time the
“‘Laubwiesen” covered extensive areas, but they are now restricted to what may
be called new terranes, especially near the coast and about inland lakes. They
seem to be particularly favored by maritime climates and calcareous soils. lor-
istically the vegetation is closely related to that of oak or beech woods on calcareous
soil rich in humus. The dominant trees are ash, oak, linden, elm, and beech,

8 L&verLié, H., Monographie du genre Oenothera. Le Mans. 1905- 10° fr.

9 SHELDON, E. P., The forest wealth of Oregon. _12mo. pp. 32- Pls. 4- Portland,
Ore. Printed by direction of the Lewis and Clark Exposition Commission. 1994-

t° Marpen, J. H., A critical revision of the genus Eucalyptus. Part v. pls. 4:
ae by the authority of the Government of the State of New South Wales. 1994+
2s.
aubwiesen-

Zur Kenntnis des Pflanzenlebens schwedischer L
- Bot. Cent.

1 HESSELMAN, H.,
Eine physiologisch-biologische und pflanzengeographische Studie. Beih.
17: 311-460. pis. 4-8. 1904.

1905] CURRENT LITERATURE 227

while the hazel is the chief shrub. The herbage is so dense as to exclude most
mosses. The formation is often developed from salt marshes and HEssELMAN
gives an interesting account of the developmental stages. The alder is commonly
the pioneer ligneous form, and is followed by the ash. The author’s most impor-
tant contribution is experimental, and in this field he has clearly pushed forward
the frontiers of our knowledge. His results deal chiefly with the ecology of starch
formation, respiration, and transpiration. An important feature of his results
is that they are rigidly quantitative. In the study of starch formation, close
correlation was made with the light intensity, in which WrESNER’s methods were
employed. The “Lichtgenuss” varied from about 1 in open meadows to #5 in
dense hazel thickets. Much formation of starch was observed under the trees in
early spring, and all through the summer as well in open meadows. The very same
species that were found rich in starch in the woods in spring were found nearly
Without it in the summer shade. In the ash woods, however, starch is formed in
summer by all herbs, except a few which are always poor in starch. This is corre-
lated with the greater light intensity of ash woods, as compared with other kinds.
Anatomical studies are also correlated with the above, and it is found that those
plants which develop their leaves late in the forest shade have a weak develop-
ment of palisade cells, while the plants which develop their leaves in the intenser
light of early spring have more palisades. Trees with high light requirement
were found to have but one leaf type, namely a sun leaf; while shade-enduring
trees have both sun or shade leaves, depending on their growth conditions. Trees
of the first class, such as the ash or birch, show starch formation in all leaves,
wherever placed. Plants of the second type, such as oak or hazel, do not form
Starch in the innermost leaves. In equal light shade leaves form more starch
than do sun leaves of the same species. Starch formation decreases from spring
to summer in the woods more in sun plants than in shade plants. Shade plants
Tespire much less than do sun plants. Shade plants transpire much less in the
shade than do sun plants in the sun, especially on hot or dry days. The leaf
surface being equal, plants transpire more as they have a greater development
of palisade cells, though many have thought that palisade cells in some way chec
transpiration, since they commonly occur where such protection seems necessary.

The great detail with which HessELMAN’s studies have been pursued, and
the tedious quantitative experiments, may well furnish a model for many workers
in all lands. The paper is illustrated by several reproductions of photographs,
and a number of text figures showing anatomical details —H. C. Cow Les.

_ THE AcHRoMATIC figure in Pellia epiphylla has been investigated again, this
time by GREGorrE and BERGHS,™ who have devoted their attention to the intra-
Sporal mitoses of the germinating spore, and in some degree to the first mitosis in
the spore mother-cell. The figures show some improvement over those of their
Predecessors and indicate a technique of the highest order. The researches of
Senses

*? GRrecorre,V., and BERGHS, J., La figure achromatique dans le Pellia epiphylla.
La Cellule a1: 193-233. pls. I-2. 1904.

228 BOTANICAL GAZETTE [MARCH

Davis and of the reviewer are frequently discussed through the work. In the
opinion of the reviewer, our technique was also good and enabled us to see the
structures described in the present paper, so that differences in conclusions are
due in large measure to differences in interpretation of structures which other
investigators have seen. The present writers support the reviewer’s conclusions
that the two poles of the chromatic figure arise in succession and independently,
and also that the nuclear membrane belongs to the category of the H autschicht.
Without details, some of the principal conclusions are the following: In the
segmentation of the spore (beginning with the third mitosis) the asters are con-
tinuous from pole to pole and the achromatic figure surrounds both the nucleus
and the “‘polar vesicles” (‘‘caps’’). The achromatic figure results simply from
a rearrangement of the general cytoplasmic reticulum. Neither the nucleus nor
the vesicles (caps) contribute to the formation of the spindle. The vesicles are
nothing but nuclear sap diffused into the cytoplasm. After mitosis the achro-
matic structures are transformed into the general cytoplasm of the cell. At no
stage is there a genuine centrosphere or centrosome. The first mitoses in the
germinating spore and the mitoses in the spore mother-cell show neither asters
nor polar vesicles (caps), but in other respects the evolution of the achromatic
figure is the same as in other mitoses. The authors hold that there is no distinc’
tion between kinoplasm and trophoplasm. No importance is attached to the
structure described by others as a centrosphere. These researches on Pellia
contradict many theories in regard to the mechanism of mitosis—C. J. CHAM-
BERLAIN.

PHILLPs's has contsibuted some ‘Startling discoveries to an already confused
subject. Particularl t to the structure of the cen-
tral body of the Cyanophyceae, which he regards as a nucleus, and the occur-
rence of sexual fusions in the formation of the spores. In unsectioned filaments,
since “‘sections revealed very little that could not be seen equally as well in the
uncut object,” the author has discovered that the chromatin of the central body
is aggregated in hollow vesicles in the resting cell. ‘This vesicular appearance
disappears in the dividing cell and the chromatin granules become arrange
loose network. In this condition, he finds the minute granules themselves multi-
plying by transverse division. Nuclear division follows, which may be by one of
two modes, both occurring even in the same species. One, according to the
author, evidently corresponds to a direct division, while the other is 4 primitive
mode of karyokinesis. The latter resembles closely the method of mitosis

13 PHILLips, O. P., A comparative study of the cytology and movements : cpa
phyceae. Contrib. Bot. Lab. Univ. Pennsylvania 2:237-335- pls. 23-25- 19°

14 Kout, F. G., Ueber die Organisation und Physiologie der Cyanophyce
und die mitotische Theilung ihres Kernes

at

1905] CURRENT LITERATURE 229

(one to four) into one, and after certain adjoining cells, called by him “nurse
cells,” have gradually disintegrated and given up their chromatin to the forming
spore. The first fusion is regarded as probably a sexual act, but he does not
appear to attach this importance to the giving up of the chromatin by the “nurse
cell.” His figures of the spores of Oscillatoria remind one of small hormogonia,
while his ‘“‘nurse cells” evidently correspond to the cells adjacent to the hor-
mogonia which have been heretofore regarded as collapsed, dead cells.

The author makes a third distinct contribution, which is also far from con-
vincing, in that he finds that the movements of the Cyanophyceae are caused by
delicate cilia, distributed along the sides of the filament—Epcar W. OLIve.

VULE of seven genera of cycads, Zamia, Macrozamia, Ceratozamia,

10)
Encephalartos, Bowenia, and Dioon, have been investigated by Miss Stoprs.'s

The work was done in GorBEr’s laboratory and the standpoint is that of phylo-
genetic anatomy.

e integument in all the forms studied is differentiated into three layers,
the outer and inner being fleshy, while the middle is the well-known stony layer.
The inner system of bundles does not lie in the nucellus (which in all living cycads
and their fossil relatives is absolutely without vascular bundles), or between the
nucellus and integument, as is commonly supposed, but belongs to the inner
fleshy layer of the integument. These bundles, which in all the seeds examined
are similar and simple, sometimes extend beyond the nucellus, reaching almost
to the micropyle. The larger ones are collateral and the phloem is not strongly
developed. It is often hard to determine where the protoxylem lies, but, in general,
the bundles are endarch. In the simplest case, these bundles come from the
branching of a single concentric or nearly concentric bundle.

The bundles in the outer flesh, with few exceptions, are collateral, with outer
phloem and inner mesarch xylem. The central strand, from which these bundles
come, is usually concentric.

It is certain that the fossil seed, Lagenostoma, is related to living cycads.
Evidence indicating this relationship is presented in detail and is illustrated by

agrams. Miss Stopes believes that Lagenostoma lies near the common origin
of the two groups. The single integument of the cycads is thought to correspond
to the two separate integuments of Lagenostoma.

It is particularly interesting to note that Cycas itself, generally thought to be
the most primitive of the cycads, has the most highly developed seed to be found
in the group.—C. J. CHAMBERLAIN.

ZACHARIAS® adds another article to his long series of writings on the structure
of the cyanophyceous cell, which leaves the subject in deeper uncertainty than
ever. This latest addition is in the main a sharp criticism of the methods and
_

*S STOPES, Marie C., Beitrige zur Kenntniss der Fortpflanzungsorgane der
Cycadeen. Flora 93:435-482. figs. 37- 1904.

*© ZacHarias, E., Ueber die Cyanophyceen. Jahrb. Hamburg Wiss. Anat. 21: 49-
89. pl. 1. 1903.

230 BOTANICAL GAZETTE [MARCH
conclusions of Kout and a reiteration of the principal results of his paper of 1890.

kérner,” or slime globules. While ZacHarias denies that it has been proved
that the central body contains any granules other than the “Zentralk6rner,” yet
he leaves the question still open as to the possibility of the existence of chromatin
in this enigmatical body. He asserts strongly his belief, however, after an exam-
ination of the preparations of both Kout and HEGLER, that there are no structures
present which can be called chromosomes, as maintained by Kout and BiTscHtt.

ZACHARIAS adds in the present paper some new observations on the nature
of the cyanophycin granules. After repeating his earlier experiments on the
effects of dilute HCl, he comes to a conclusion somewhat at variance with his
earlier results, namely, that the granules in certain cases are not actually dis-
solved by the acid, but that nevertheless they undergo some change. He con-
cludes, further, that there is no difference in the behavior of the cyanophycin
toward dilute HCl and toward pepsin, thus disputing the results of HEGLER and
Kout. The author as well cannot agree with these investigators in their conclu-
sions as to the albuminous and crystalloidal nature of the cyanophycin granules,
although he adds nothing in support of his earlier supposition of their carbohy-
drate nature.

In regard to the duration of cyanophycin in filaments grown for a long time
in the dark, Zacwartas differs from HEGLER and Kout in asserting that there
is no complete disappearance of cyanophycin and slime.—Epcar W. OLIVE.

KoERNICKE reviews critically the present condition of researches upon the
plant cell,*? confining his attention to the morphological aspect now commonly
spoken of as cytology. A little more than one half the work is devoted to the
cytoplasm, the rest dealing with the nucleus.

That modern preparations do not show a series of artifacts but give a reliable
view of structures, is proved by observations upon living material. The cytoplasm
is regarded as consisting of two distinct substances, the kinoplasm and the tro-
phoplasm. Spindle formation is described in all groups and the various views
are clearly presented. The centrosome problem is discussed at some length.

impartial summary of the literature dealing with the blepharoplast is given,
but the author expresses no opinion as to its homology. The nuclear membrane,
cell plate, and Hauéschicht are regarded as kinoplasmic structures. The pat
graphs on protoplasmic continuity are especially valuable because the important
Investigations are so recent. Less attention is paid to the cell wall.

In treating the nucleus, the author deals principally with the works which
have appeared since 1896, the date of Zimmerman’s book on the nucleus, sinc
when many investigators have studied the nuclei of the lower plants, especially
the schizophytes and yeasts. Of course, the discussion of the chromatin occuples
most of the space devoted to the nucleus. It is rather surprising that the Chara-

*7 KOERNICKE, M., Der heutige Stand der pflanzlichen Zellforschung- esas
Deutsch. Bot. Gesells. 21 : Generalsammlungs-Heft 66-134. 1904-

1905] CURRENT LITERATURE 231

ceae, Liliaceae, and Amaryllidaceae should be credited with the largest nuclei,
for their nuclei are small when compared with those of the eggs of some gymno-
sperms.

o figures are given, but the references are very numerous and authority is
cited for every statement.—C. J. CHAMBERLAIN.

R REVIEWING several of the more important publications regarding the
statolith theory, TrscHLER* states that adequate evidence to establish a correla-
tion between movable starch grains and the perception of the gravitation stimulus
is not available; and it is his opinion that such evidence cannot, with present
methods, become available, for the simple reason that while the reaction ma
optically observed the perception is hidden and a constancy of quantitative rela-
tion between the two cannot be depended upon. For similar reasons, the author
believes a demonstration of the statolith theory cannot come from the study of
organs which are little or not at all reactive; hence we must regard his contribution
as consisting of merely circumstantial evidence. The general trend of the paper
is that in the various types of roots slightly or not at all geotropic, geotropism and
starch migration are coincident and associated phenomena. He finds that in
adventitious roots which are constantly ageotropic these starch grains are either
not present in the cap or if present are irregularly distributed. In temporarily
ageotropic roots the starch grains are either absent or if present are irregularly
distributed during the ageotropic period. With incipient geotropism, however,
the starch grains appear if previously absent, and commence to collect and func-
tion as statoliths. When geotropic response is prevented by the application of
stronger stimuli, the starch grains which would otherwise change position to
function as statoliths do not do so. Some of the orchids having aerial roots
slightly geotropic do not have starch grains, and the author calls in chloroplasts of
the root-cap to act as statoliths —Raymonp H. Ponp.

PALEOBOTANICAL NOTES.—STENZEL”? has just published a very elaborate and
valuable monograph describing all the known forms of fossil palm-wood.—
NATHORST?° briefly summarizes our knowledge of Antarctic fossil floras. A
rather rich deposit of Jurassic age on Louis-Philippe Land yields numerous remains
of Cladophlebis, besides representatives of Brachyphyllum, Elatides, Palissya,

axites, Araucarites, Otozamites, Pterophyllum, Equisetum, Sagenopteris,
Thinnfeldia, Scleropteris, and Stachypteris; all hati cteristic Jurassic forms,
emphasizing the uniform cosmopolitan character of the Jurassic flora even in
lat. 63° 30'S. Seymour Island in lat. 64°S. yields fragments of a dicotyledon, of
Araucarites, and of a conifer resembling Sequoia. The Malouine Islands furni

ER, G., Ueber das Vorkommen von statolithen bei wenig oder gar nicht
grotropischen Wer Flora 94:1-68. figs. 3I. 1905.
"9 STENZEL, K. G., Fossile Palmenhdélzer. Beitr. Palaeont. u. Geol. Oesterr.
Ungarn. fol. pp. wa pls. 22. 1904.
*° NatHorst, A. G., Compt. Rend. 138:1447-1450- 1904.

232 BOTANICAL GAZETTE [MARCH

fragments of an Asterocalamites indicative of Upper Devonian or Lower Carbonif-
erous age.—Maury?! reports Hicoria minor, Castanea vulgaris, Fagus pliocenica,
Tlex aquifolium, and Bambusa lugdunensis from a new Pliocene locality at Capelle
in the Department of Cantal, south central France.—Hartz?? records characteris-
tic spikes of the common Dulichium arundinaceum (L.) Britt. of eastern North
America, along with the remains of Picea, Brasenia, Hydrocharis, Carpinus, and
Betula, from the interglacial of Brérup in southern Jutland, Denmark.—EpwarD
W. BERRY

Hume? calls attention to the anthracnose of the pomelo in Florida caused by
Colletotrichum gloeosporoides. He reported in 1g00 an injury to pomelo leaves
caused by this fungus, a disease that was then referred to as leaf spot. HUME
first noticed the same fungus on pomelo fruits in 1901 and now reports it upon
twigs of the same plant. It is also known to cause a disease of lemon and lime
fruits and is frequently seen on the leaves and twigs of the sweet orange and rarely
on its fruit. Trees whose vitality has been reduced by improper soil conditions
or by mechanical injury or through injuries inflicted by winds, frost, insects, oT
other diseases are more subject to this anthracnose than trees in perfect condition.
It is recommended that all diseased fruits and branches be removed and burne
and the trees sprayed with the usual Bordeaux mixture early in the season from
May to July. Spray treatment should particularly be applied to trees whose
branches show any evidence of the anthracnose. It is also recommended that
the fruit be washed, before packing, with water to which has been added some
ammoniacal copper carbonate solution or potassium sulfid. The later treatment
will prevent the serious injury to the fruit that frequently occurs on the way to
market.—E. Mrap WItcox. -

By MEASURING the distance between oppositely located branches of small
trees and shrubs, GANoNG?4 has established that with autumnal defoliation the ©
branches commence an inward movement toward the main axis. This inwat
movement continues after defoliation, reaching a limit with the full winter condi-
tion in January. With the swelling of the buds in April a reverse movement
commences which continues through vernal foliation to the full summer condition
in June. The movement accompanying defoliation and foliation is attribute
merely to decreasing and increasing weight of branches, by loss and gain of leaves,
but the intervening movement is regarded as indirectly thermometric because
from experimental data (quite limited and so regarded by the author) it seems
the movement is correlated with variations in the amount of water in the stem

that
and

2t Maury, P., Le Monde des Plantes, Nov. 1903, PP- 54-55-

22 Hartz, N., Meddel. Dansk. Geol. Foren. 10:13-22. figs. 5. 1904:

23 Hume, H. Haroxp, Anthracnose of the pomelo. Bull. Florida Exp. Stat
74:157-172. pl. I-4. 1904.

24 GANONG, W. F., An undescribed thermometric movement of the branc
shrubs and trees. Ann. Botany 18: 631-644. figs. 6. 1904.

hes in

Pe! Tipe gM

- authors (DEtTTo?7 for

1905] CURRENT LITERATURE 233

that fluctuations in temperature induce such variations in water content. The
author regards the movement as without ecological significance. The practical
absence of other literature on this subject gives the paper a quality of uniqueness,
and although negative results appear to equal the positive in number, the high
ideals of the accurate and unprejudiced worker are everywhere suggested.—
Raymonp H. Ponp

Ganonc has published in an earlier paper hiss views concerning some of the
underlying principles of ecology, and to these he has added others?° dealing espe-
cially with adaptation. After criticising the idea that ecology is synonymous
with ecological geography, and deploring the lack of any significant advance in
ecological methods in recent years, he plunges into a discussion of the philosophy
of adaptation. Five principles are laid down; the first, as to the reality of adapta-
tion, and the last, as to the inevitable imperfection of all adaptation, must meet
with general acceptance. The second principle, dealing with the evolutionary
phylogeny of adaptation, is taken up from a Lamarckian standpoint. GANONG
thinks that the concomitance of diverse adaptations in a single species is to be
accounted for best along Lamarckian lines. While this is doubtless true, there
are as many objections wan 05 OD we as 2 oe suena hk poe

of all. The third and fractile principles are concerned ith adaptation as a race
Process rather than an individual process, and with the metamorphic nature of
its origin—H. C. Cow es.

TANSLEy’s?8 address before the British Association on The problems of ecology
is very suggestive, and indicates the rapid progress which is being made along
ecological lines in the British Isles. ANSLEY proposes to narrow the definition
of ecology to include only its geographic aspects. For instance, he would include

“the geographic distribution of pollination mechanisms, but would exclude the

study of pollination mechanisms themselves. The reviewer can well imagine
what a shock this will give to those who imagine that ecology includes nothing
but the study of seed dispersal and pollination! TANSLEY uses as a synonym
« sores | wperpnice Papaalngy, eur is practically identical with what the
gy. It does not seem likely that TANSLEY’s
view as to the limits of ecology will meet general approval, since the broader term
has been so universally regarded as highly desirable and much needed. The
distinctions between descriptive and experimental ecology are well brought out,
and proper emphasis is laid upon the ultimate aim of ecology, the discovery of
causes.—H. C. Cow es.
a,

*5 See Bor. Gaz. 36: 447-453. 1903.
© GANONG, W. F., The cardinal principles of ecology. Science 19: 493-498. 1904.
*7 See Bor. Gaz. 38: 385-386. 1904.
28 TANSLEY, A. G., The problems of ecology. New Phytologist 3: 191-200. 1904.

234 BOTANICAL GAZETTE [MARCH

PARTHENOGENESIS in Taraxacum and Hieracium is being investigated by
MvrBECK.?° Two species of Taraxacum were studied, T. vulgare (Lam.) Raunk.,
which produces abundant, but imperfect pollen, and 7. speciosum Raunk., which
produces no pollen at all. e embryo-sacs present nothing unusual in their
appearance. The egg cell increases in size, its nucleus divides, and the embryo
is formed just as if normal fertilization had occurred. The polar nuclei fuse and
then divide, giving rise to the endosperm.

Three species of Hieracium were examined, H. grandidens Dahlst., H. ser-
ratijrons Almqu., and H. colophyllum leiopogon Gren. In general, the embryo-
sac resembles that of Taraxacum. The embryo is developed from the egg, and
embryos are formed in eighty per cent. or ninety per cent. of the flowers, so that
it is probable that these species are always parthenogenetic. A full account with
illustrations is to follow.—C. J. CHAMBERLAIN.

Miss Cootrys° has given an account of the ecology of the tamarack, which
has been based upon a considerable amount of field study in various states.
Attention is called to the unusally wide range of the tree, which occurs in five of
MeERRIAM’s zones. The tree displays great vigor even at the extreme northern
and southern limits. The range of topographic habitat is also wide; although
commonly thought of as a swamp tree, it occurs in almost every habitat, even to
the most xerophytic. Among the characteristics which may serve to give this
tree its great success, Miss Cootey mentions the abundance of seeds, the fre-
quency with which the tree produces seeds, the power of the seedlings to endure
light, indifference to soil, and rapid growth. A great disadvantage is the inability
of the tree to endure shade, or to compete with other forest trees. In a Maine
area a tamarack swamp has succeeded a hardwood forest.—H. C. COWLES.

Apocamy in Hieracium is described by OSTENFELD,3? who in 1903 in a paper
written conjointly with RAUNKIAER reported that twenty-two species of the genus
had been found to produce seeds even after pollination had been prevented.
OsTENFELD now finds that many of the seeds thus produced germinate normally.
In the botanical garden at Copenhagen two species of Hieracium were found
which produce only female flowers, a careful examination showing not 4 single

was not determined. Attempts to germinate the pollen of pollen-bearing a2
failed. This would indicate that the great number of forms in the genus, US" :
attributed to hybridization, are due rather to mutation. All attempts to er
ybrids were unsuccessful—C. J. CHAMBERLAIN.

29 MURBECK, Sv., Parthenogenese bei den Gattungen Taraxacum und Hieracium-
Bot. Notiser 1904: 285-296.

3° CooLEy, Grace E., Silvicultural features of Larix americana. F
Quarterly 2: 148-160. 1904.

3t OsTENFELD, C. H., Zur Kenntniss der Apogamie in der Gattung Hiera
Ber. Deutsch. Bot. Gesells. 22: 376-381. 1904.

orestry

cium.

1905] CURRENT LITERATURE 235

THE NUTRITION of the chromatin thread and the chromosomes by nucleolar
material is well shown, according to VON DERSHAU,3? in the active nuclei of the
parietal endosperm of Fritillaria imperialis. The increasing staining capacity
of the chromatin thread is correlated with the dissolution of the nucleoli; besides,
slender connections unite the nucleoli and the thread and indicate a streaming
of material. Resting nucleoli are spherical or ellipsoidal, while those in the active
condition are more or less irregular or star-shaped. A study of karyokinesis
showed a direct relation between the nucleoli and spindle formation. An investi-
gation of the development of the peristome of Funaria hygrometrica and Bryum
argenteum showed that the nucleoli play an active part in the local thickenings of
the cell wall. Von DersHav draws the conclusion that the nucleoli consist of
reserve substance.—C. J. CHAMBERLAIN.

BISPORANGIATE STROBILI have not hitherto been observed in Juniperus, and
only a single case has been reported for the Cupressineae. An interesting case is
described by RENNER33, who discovered a plant which bears such strobili almost
exclusively. Only on a few twigs, and here only toward the tips, was there a
transition to purely ovulate strobili. In the base of the bisporangiate strobilus
there are a few sterile leaves, above these are two or three whorls of staminate
sporophylls, and at the tip are the ovules. There may be a whorl of sterile leaves
between the staminate sporophylls and the ovules. The pollen develops nor-
mally, but so much later than the ovules that there is no possibility of self-
pollination. RENNER sees in these bisporangiate strobili a suggestion of the
method by which the bisporangiate flowers of angiosperms may have arisen.—

J. CHAMBERLAIN.

Waires+ has published the first American contribution to our knowledge of
those paleozoic fern-like plants which had acquired the seed-habit, a subject
under active investigation abroad of late. He records abundant gymnospermous
fruits in the fern-genus Aneimites from the lower Pottsville of West Virginia.
They are usually borne singly at the apices of the slightly dilated terminal exten-
sions of the outer pinnae, and show some slight indications of a pollen-chamber.
They have been found only as impressions and therefore cannot be compared
Properly with the beautifully preserved foreign specimens showing internal
structure. The author is of the opinion that the American form is more closely
related to Lagenostoma than to any other cycadofilicinean type at present recog-
hized as seed-bearing —EDWARD W. BERRY.

3? DeRSHAU, MAx vON, Wanderung nukleolarer Substanz wahrend der Kary-
okinese und in lokal sich verdickenden Zellen. Ber. Deutsch. Bot. Gesells. 22: 400-
411. pl. 21. 1904.

33 RENNER., Orro, Ueber Zwitterbliithen bei Junip mmunis. Flora 93:297-
300. 7ig. 3. 1904.

: 4 WauitE, D., Smithsonian Miscellaneous Collections 47:322-331- pls. 47-48.
904.

236 BOTANICAL GAZETTE [MARCH

PARTHENOGENESIS in Wikstroemia indica, one of the Thymelaeaceae, is an-
nounced in a preliminary paper by WINKLER.35_ The pollen develops imperfectly
or not at all, and attempts to germinate the pollen by the usual methods were unsuc-
cessful. Although pollination was prevented, embryos developed, and sections
showed that the embryos arise from unfertilized eggs. The behavior of the
chromatin will be studied later. Up to date the angiosperms in which partheno-
genesis has been shown to exist are Antennaria alpina, Alchemilla (several species),
Thalictrum purpurascens, Taraxacum officinale, Hieractum, and Wikstroemia
indica. Gnetum Ula is the only gymnosperm in which parthenogenesis has yet
been reported.—CHarLes J. CHAMBERLAIN.

LSSON-SEFFER?* has written a short defense of LiNNAEUS, 2 propos of
the recent Swedish biography by T. M. Frres. The particular occasion
for OLSSON-SEFFER’s paper is a choleric and most unjust attack upon LINNAEUS
by KatiscHer and Hansen. Considerable attention is also paid to the
unfair treatment which is accorded to LinnaEus in Sacus’s History of Botany.
There can be no doubt that LrnNAEus has a secure place among the greatest
botanists of all time, and the imputations that have been cast upon him are certain
to be less harmful to - renown of LinNAEvs than to the reputation of his
accusers.—H. C. Cow

RCELLULAR PROTOPLASM has been further investigated by Kny.7_ The
principal material was cotyledons of Lupinus albus and L. angustifolius. That
the protoplasm in the intercellular spaces is in direct communication with that
of the neighboring cells is probable from the fact that the spaces contain starch
grains. If there is protoplasmic continuity between the intercellular spaces and
the neighboring cells the absence of nuclei in the spaces would not be strange.
Perforations and connecting threads could not be identified positively. How
the protoplasm en) into the spaces the author is not yet ready to explain. J.
CHAMBERLAIN.

THE PHYTON THEORY is supported by Cor in a lengthy papers® which deals
mainly with abnormally placed vascular bundles in dicotyledons. Starting with
the Campanulaceae the author has extended his observations to a large number
of families, and Teaches the conclusion that the bundles in question are normal

tly for only a short distance. The chief argu-
ments advanced in favor of the ‘phyton theory are (1) the bundles coming from
eaves diminish in volume as they descend through the stem; (2) practically

KLER, HANs, Ueber ae bei Wikstroemia indica (L.) C. A- =
Ber. stacerage Bot. Gesells. 22:573- 1905.

36 OLSSON-SEFFER, P., The a of Linnaeus in the history of botany. Jour $
Bot. 42:262-269. 1904.

. “

37 Kny, L., Studien iiber intercellulares protoplasma II. Ber. Deutsch. Bo
Gesells. 22: 347-355. 1904. d

38 Co, A., Recherches sur la disposition des faisceaux dans la tige et les feuilles “©
quelques dicotylédoien. Ann. Sci. Nat. Bot. 8: 20: 1-288. 1904.

1905] CURRENT LITERATURE 237

all of the eee formations of the stem supply oo organs.—M. A.
CHRYSLE

tae has published further additions to the Cretaceous flora of Long
Island, treating of plants from the Northport Clays and from the vicinity of
Hempstead Harbor, Oyster Bay, and Montauk Point. Nine new species, includ-
ing an interesting form of Marsilia, are described. The supposed palm Serenop-
sis is reconsidered in the light of better material and is definitely referred to Ne-
Jumbo. This leaves a species of palm (about to be published by the undersigned
in Torreya) coming from the mid-Cretaceous of Delaware and Maryland, as the
earliest known plant of this type from American strata.—EDWARD W. BERRY.

NrwcomBE,*° in his presidential address before the Michigan Academy of
Science, contrasts the old and new types of biological surveys, and makes a strong
appeal for the new progressive and definite survey of the ecological type, accom-
panied by detailed maps, such as has been carried on by FLAHAULT in France,
by the SairHs in Britain, and begun in Michigan by Livincston. Concise
statements are made concerning the uses of such a survey, both scientific and
practical. It is much to be hoped that the plan here advocated will be carried
out, not only in Michigan but in many other states as well.—H. C. Co

LAWRENCE gives the results of his investigation of the apple scab in Wash-
ington. 4 He was able, as were CLINTON and others, to confirm the discovery made
by ADERHOLD, that Fusicladium dendriticum, found as a parasite on the fruit
and-leaves of the apple during the summer, is but a stage in the development of
Venturia inaequalis, the perfect form being found on the dead leaves as a sapro-
phyte during the winter. He recommends that the fallen leaves should be gath-
ered and destroyed and that the usual highs with Bordeaux mixture be con-
ducted in the spring —E. Mrap Witc

CHEMOTAXIS of the sperms of .. toward a number of albumins,
ea nucleo-albumins, proteids, and enzymes, is the subject of a paper by
by Lrprorss.4? Out of nineteen substances tested only two gave a negative
sce et the others attract the sperms when in the right concentration. The
author suggests that proteid substances are probably the main factor in the attrac-
tion of such sperms to the archegonia in nature, as well as in the tropism of pollen
ay There is some evidence that Marchantia sperms are positively aerotropic.
—B. E. Livincston.

39 HoLicx, A., Additions to the paleobotany of the cretaceous formation on Long
Island. II. Bull. N. Y. Bot. Gard. 3:403-418. pls. 70-79. 1904.

4° Newcomsr, F. C., A natural history survey for Michigan. Annual Report
Mich. Acad. Sci. 6: 28-36. 1904.

4 LAWRENCE, W. H., The apple scab in western Washington. Bull. Wash. Exp.
Stat. 64: 1~24. Sis I-5. pl. I-2. 1904.

“ Liprorss, B., Ueber die Reizbewegungen der Marchantia-Spermatozoiden.
Jahr. Wiss. fie 41: oe he. 1904.

238 BOTANICAL GAZETTE [MARCH

sity has published an account of observations and experiments regarding
five serious diseases of the Irish potato that are known to occur in Florida:
late blight, caused by Phytophthora infestans; early blight, caused by Alternaria
solani; scab, caused by Oospora scabies; rhizoctonia blight, caused by a species
of etclanctunig and bacterial blight, caused by Bacillus solanacearum. A brief
discussion of the nature and effects of each disease is accompanied by statements
as to the proper methods of control to be employed in each case.—E. MEAD
WILCox.

From A consideration of the various causes assigned by different authors as
inducing flower-formation, Lozw4+ believes that such causes have one common
physiological factor, namely, an increased concentration of sugar. He advances
the theory that flower-formation is a phenomenon of irritability; that a certain
concentration of sugar in the plant stimulates the embryonic substance to differ-
entiate male and female nuclei. nian Si the evidence offered is very
limited and not at all conclusive-—RAYMOND ND.

NATHANSOHN’S last work on permeability of the plasmatic membranes of
plants,45 which was reviewed in these pages some time since,*® includes in SS ~
cussion the results of a large series of experiments with the tissue of Dahlia tu
The paper which embodied the details of this work, though actually ere
before the one already reviewed, has but recently come to hand.47_ T hose inter-
ested in this subject will need to read the two papers together —B. E. Livinc-
STON.

THE FIRST NUMBER of the second series of KaRSTEN and ScHENck’s*® Vege
tationsbilder comes from the hand of Ute, the well-known student of the
South American tropics. The number is devoted to the illustration of some
characteristic epiphytes of the Amazon district, and the Standard of the preceding
numbers is fully maintained. Among the epiphytes figured are Nidularium,
Platycerium, Polypodium, Cereus, and Anthurium.—H. C. CowLes.

THE INFLUENCE of weak aqueous solutions of ether upon the growth of
Spirogyra has been studied by Grrasstmmow.49 In these cultures many of the

UME, H. Haroxp, Potato Diseases. Bull. Florida Exp. Stat. 75 77-199

pl. I-4 and frontispiece. 1904.

44 Loew, Oskar, Zur Theorie der bliitenbildenden Stoffe. Flora gg: 124-128
1905.

45 NATHANSOHN, A., Weitere ce at iiber die Regulation der sic
nahme. Jahrb. Wiss. B Bot. 40:

46 Bot. GAZETTE = 477- 1904. al

47 NATHANSOHN, A., Ueber die Regulation der Aufnahme anorganischer —
durch die Knollen von gett aa ahrb. Wiss. Bot. 39: 607-644. 1904-

48 Karsten, G., und ScHenck, H., Vegetationsbilder. Zweite Reihe, oe
E. Ure, Die Epiphyten des date Tafel 1-6. Jena: Gustav .
1904. Single parts M 4; to subscribers M 2.50. s

49 GERAssimow, J. J., Aetherkulturen von Spirogyra. Flora on

1905] CURRENT LITERATURE 239

cells swell laterally and thus become barrel-shaped. Certain cells without nuclei

increase in length but never swell in this way. Thus the author concludes that

the swelling is brought about in some manner through the action of the ether upon
e nuclear mechanism.—B. E. Livincston.

Tscuircus° finds that many dicotyledonous plants possess two types of root
in the same individual, one sort having a nutritive function and the other serving
to fasten the plant to the substratum. The so-called “fastening roots” are dis-
tinguished by the presence of mechanical fibers and generally by the absence of
pith and by the larger size of the central cylinder—M. A. CHRYSLER.

R. M. Harpers? has continued his valuable ecological and taxonomic obser-
vations on the flora of Georgia, a state whose botany has been peculiarly unknown
until these and previously noteds? studies were made. In addition to critical
taxonomic notes, these papers contain accounts of the author’s itinerary, together
with landscape figures and ecological remarks.—H. C. CowLeEs

EPIPHEGUS VIRGINIANA has been studied by Cooke and ScHIVELY,53 who
find that the parasitic haustoria arise from the roots of the host, the huni and
not from the parasite. As is usual in such plants, there is a sperm
and the embryo is rudimentary. Bicollateral bundles are found both in nthe
tuber and in the aerial shoot—M. A. CHRYSLER.

Wooprvrre-Pracocks4 has published some rather peculiar suggestions on
the study of rock-soil floras. An immense number of habitat types are proposed,
but no attempt is made at analysis. It is obvious that the author is not conversant
with modern ss literature —H. C. CowLes

£ TURE and sprouting of the seed, and the structure of the mature
plant of Cisivahi filijormis are described by Miss Borwic.5s This plant, though
a parasite resembling Cuscuta, has functional chlorophyll tissue, especially when
growing in the shade.—M. A. CHRYSLER.

TANSLEYS® gives an account of the methods which he has employed in eco-
logical surveying with his classes on the coast of Brittany. The scheme makes
Possible accurate and detailed mapping.—H. C. Cow Les.

5° Tscuircu, A., Ueber die Heterorhizie bei Dikotylen. Flora 94:68-78. 1905.
5t Harper, R. M., Botanical explorations in Georgia during the summer of rgor
Bull. Torr. Bot. Club 30: nn pats 319-342. 1903. SSE HNC in the coastal hai
of Georgia during the season of 1902. Bull. Torr. Bot. Club 31:9-27. 1904.
5? See Bor. Gaz. 34: 386. 1902.
OOKE, E., and Scuive ty, A. F., ti n the structure =i ae
of Eephecu virginiana. Contrib. Bot. Lab. Univ. Penn. 2: 352-398.
54 WooDRUFFE-PEACOCK, E. A., How to make notes for a ee hh Rural
Studies Series, No. 5. Louth, 1904
53 BoEwIc, Harriet, The histology and development of Cassytha_ filiformis.
Contrib. Bot. Lab. Univ. Penn. 2: 399-416. 1904.
8° TANSLEY. A, G., A second experiment in ecological surveying. New Phytol-
ogist 3:200-204. 1904.

NEWS.

Dr. Ernst HALuier, the mycologist, died at Dachau, near Munich, Decem-
ber 20, 1904, at the age of seventy-three.

Dr. A. Ernst, formerly privatdocent at the University of Ziirich, has been
- appointed professor and director of the botanical Jaboratories.

Miss Ciara E. Cummines, of Wellesley College, went to Jamaica in Feb-
ruary, to spend several months in a study of the flora, especially the lichens.

A. S. Hrrcucock, of the Division of Agrostology, Department of Agriculture,
has purchased the grass herbarium of F. LAMSoN-SCRIBNER, containing many
of his types.

Dr. Ernst ABBE, of Jena, whose name is so well known in connection with
the optical apparatus of the Karl Zeiss establishment, died January 16 at the age
of sixty-four years.

Str JosePH Hooker retired from the editorship of the Botanical Magazine
at the close of 1904, after forty years of service. He is succeeded by Sir W.
THISELTON-DYER. ;

AT THE MEETING of the Linnean Society on December 15, 1904, fifteen ladies
were elected Fellows. Among those thus honored were Miss MARGARET BENSON
and Miss ETHEL SARGANT.

AT THE ANNUAL MEETING of the Torrey Botanical Club, held on January *
H. H. Russy was elected president in the place of ADDIsoN Brown, who resigned
after fifteen years of service in that office.

JouN F. CrowE Lt, director of the Buffalo Botanical Garden, will visit Panama
during February and March as an agent of the New York Botanical Garden,
which will undertake botanical explorations in co-operation with the engineers
of the Panama Canal Commission.— Science.

Proressor N. L. Brirron, Mrs. E. G. Britton, Dr. C. F. MriisPAUGH,
and Mr. M. A. Hows sailed in January to New Providence for a further botan!
exploration of the Bahaman archipelago. Mrs. Britton will collect on New
Providence, while the rest of the party will explore the Berry and Great Bahama
groups. :

Tue Macuiiian Co. announces for the spring a revised and enlarged —
tion with many new illustrations of CAMPBELL’s Mosses and ferns. They wil ich
publish a book by Maup G. PETERSON entitled How to know wild fruits, wh of
classifies wild plants by their fruits. They are also to publish a translatio?
Connnem’s Chemistry of the proteids, by Dr. Gustav Mann. The a
be of interest not only to chemists, but especially to biologists. The advance :
the knowledge of proteids will be fully set forth. [ a

240

BUFFALO
__ LITHIA WATER

No Remedy of Ordinary Merit Could Ever

j Have Received Indorsations from
j Men Like These.
mul ©. L Fotter, A. M., M. D., M.R.C.P., London,
jr be of the Principles d Practice of "Medicine and Clinical

ae in the College oo Physician s and Surgeons, San Francisco.
Dr. . H. Dru ond, iid oad Medical Jurisprudence,
Bishop’ Ss blk Monreal, Cana
Cyrus Edson, A. M., M. D., an ith Commissioner New
Yor z City and State, President Board of Pharmacy, New York
City, sam agp.) Hos opts Corporation igi ar ete,

John V. aker, M. D., vofessor Materia
Pregnancy Medica and Taree Mules Cnaeie vical Die ge, Philadelphia.
: Ben. Johnston, Richmond, Va., Ex-President
~President

Dr. George
Southern verse Soene and Gynecolog ical anole Ex-Prest
Medical Society of Va., and Professor of Gynecology and Abdominal
Surgery, Madscat College of Va.

PTS > ae a ae Sey eee Seats an Se eS ae Se ee Ln rE SF rs
Be goatee aien) as se tee en ae eee

. A Pouchet, ssor of Pharmacology and

Materia’ inne, nih the Faculty of fsa sem Plots =

Dr. J. T. LeBlanchard, Prof Montreal Clinic, SM.,SN..V.T.
Jas. M. Crook, A. M., M. D., Professor Clinical Medicine

and Clinical Diagnosis, New York Post Graduate Medical Schoot.
Louis C. Horn, M. D., Ph. D., Professor Diseases of Chil-

dren and Dermatology, Baltimore Lely ity.

Pees a Allison Hod President and Professor Nervous and
ental Diseases, Unssorsty College af Matician Richmond, Va.

"Dr. Robert Bartholow, M.A., LL. D., Professor Materia
Medica - General eels mare Je (ihimag Medical Coliege, Phila.

i i fove, Z New York City, Former Professor Diseases of

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Che Botanical Gazette

a {Monthly Journal Embracing all Departments of Botanical Science

3 - Edited by JoHN M. CouLtrer and CHartes R. Barnes, with the — of other members of the
botanical staff of the University of Chica

; Vol. XXXIX, No. 4 Issued April 18, 1905

CONTENTS

CONTRIBUTIONS FROM THE CRYPTOGAMIC LABORATORY OF HARVARD
UNIV gpd i >, Gia. di kyies AMERICAN 2 ee OF a ER PLATES IV AND x

Roland Thaxt ra 241
(ON THE DEVELOPMENT OF HAMAMELIS VIRGINIANA. ConrTRIBUTIONS FROM THE
AL LABORATORY OF derbi eeise poh anni No. -3 sale PLATES VI AND

VII). 2. N. Shoemaker 248
EXUAL REPRODUCTION IN THE RUSTS (WITH PLATE vitt). A. H. Christman - - 267
THE FORESTS OF THE FLATHEAD VALLEY, MONTANA. ConrtRIBUTIONS FROM THE

Hurt BotanicaL ‘ibaa hr ara LXVII (wr ITH MAP AND TWENTY-THREE FIGURES).

Harry N. Whitjord — - RE Be A dy en RES

_BRIEFER ARTICLES. ©
OTES ON THE PHYSIOLOGY OF STIGEOCLONIUM. CONTRIBUTIONS FROM THE HULL
Boranicat LaBoratory. LXX (wITH THREE FIGURES). Burton Edward Livingston - 207
FERTILIZATION IN THE SAPROLEGNIALES. . A. Trow . - - - - - goo
URRENT LI fast cue

BOOK RE 2 EB y ‘ j ss : 2. ee
* THE le OF NORTH AMERICA. ORGANIC EVOLUTION.
_. MINOR NOTICES : ‘ . 3 ‘ fl a <* 308

NOTES FOR STUDENTS - : - - - - - : - 302
EWS : 3 i ¥ ‘ - " a - : - 319

Contin one for the Edit } ee aA pao , Chica: cago, Tl.
are requested to write ea - proper names yi Wieclior ca care and in citations
wn in the pages of the Bor AZETTE.

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1904, by the University of Chicag

VOLUME XXXIX NUMBER 4

BOTANICAL GAZETTE

“APRIL, 1905

CONTRIBUTIONS FROM THE CRYPTOGAMIC LABORA-
TORY OF HARVARD UNIVERSITY. LX.
A NEW AMERICAN SPECIES OF WYNNEA.
ROLAND THAXTER.
(WITH PLATES IV AND V)

In the third volume of Hooxer’s Journal of Botany and Kew
Garden Miscellany (1851) BERKELEY published a description without
figures of a large and striking discomycetous fungus, distinguished
by the possession of long ear- or spoon-shaped apothecia, arising in a
fasciculate fashion from a well-developed common stem. This form,
which was said to have been found abundantly on rotten wood near
Darjeeling, India, he placed in‘ the then very comprehensive genus
Peziza, comparing it to Peziza onotica and P. leporina, and designating
it as P. macrotis. Some years later, however, having received from
Dr. Curtis a closely related North American species collected by
Botrert near Orizaba, Mexico, he was led to create the new genus
Wynnea for the reception of these two forms (Jour. Linn. Soc.
9:1866), the Mexican species being used as the type under the name
Wynnea gigantea. Both species were subsequently illustrated by
Cooke in his M icrographia, the colors being no doubt guessed at
from BERKELEY’s descriptions and from the dried specimens. W.
macrotis is here said by CooKE to occur also in Mexico, but no author-
~ ity for this statement is mentioned. During the forty years that have
elapsed since the collection of W. gigantea by BotTert, there seems
to have been no further mention of the occurrerice of these or of other
species of Wynnea, and in more recent years the genus has been
consigned to the limbo of synonymy by Saccarpo in his Sylloge,
where both species are included in the genus Midotis.

241

242 BOTANICAL GAZETTE [APRIL

’ In the summer of 1888, a portion of which was spent by the writer
in collecting fungi among the mountains of Tennessee and North
Carolina, a species of Wynnea was found near Burbank, Tennessee,
growing on the ground.in rich woods, in a single locality; where
several clusters of its long bluntly pointed, rabbit-ear-shaped dark
brown apothecia were scattered in a limited space, each cluster borne
on a well-defined stout stem emerging directly from the humus. The
resemblance of this plant to CooKe’s figure of Wynnea macrotis, to
which it closely corresponds in form and color, was so striking that it
was assumed to be that species, despite certain differences in the size
and appearance of the spores when fully mature, and in the absence
of any authentic material of the Indian species for comparison it was
so referred.

A second visit was made to the same region in 1896, and the Wynnea
was again encountered, both at Burbank and at Cranberry, North
Carolina; one of the specimens from the last-named locality being
parasitized by a fine species of Syncephalis described in a former
number of the GAzETTE (24:1. 1897) as S. Wynneae, the host being
here recorded as W. macrotis. Having noticed, while gathering this
material, that the stems appeared to have been broken from some
attachment in every case, and not to have arisen like most humus
Pezizae from an indefinite mycelium, a more careful examination
was made in subsequent gatherings, and a little digging about the base
of the stem showed that it originated in every case from a large,
irregularly lobed, brown, firm, tuber-like body buried a few inches
deep in the humus. This body, which was somewhat cartilaginous
in consistency, showed, when cut, a chambered structure (figs. 5 and
6), the interior-being traversed by light and dark more or less con
trasting winding areas, closely resembling those characteristic of
many Tuberaceae or Hymenogastreae; and at first sight it seemed not
impossible that the Wynnea might actually be parasitic on some
hypogaeous fungus. A microscopic examination of sections cut from
this tuber, however, showed no signs of any structures which could
by any possibility be considered to represent modified hymenia-_ Te
chambered interior, as is shown by the accompanying figures, 1 vet
rounded by an external layer or cortex of large, empty; thin-walled,
brownish cells, those on the surface showing signs of degeneration

1905] THAXTER—NEW SPECIES OF WYNNEA 243

and wearing, which is rather abruptly succeeded by a peripheral
region lying beneath it, composed of densely woven brownish fila-
ments. As these filaments pass toward the interior, they become
more loosely woven, the walls are much thickened, and the living
protoplasmic contents are evident; the brownish tinge is lost as they
pass into the white or colorless more or less gelatinous areas by which
the adjacent chambers are separated. The chambers themselves
show the greatest irregularity in contour, and are clearly marked off
by a layer of dark cells which line them and contrast rather abruptly
with the intermediate colorless areas above mentioned. ‘These dark
hymenium-like layers closely resemble the general external layer, and
are made up of dark-walled, rounded, empty cells of somewhat
smaller size, packed irregularly four or five deep; the somewhat
ragged appearance of the superficial ones suggesting the lysigenous
origin of the cavities which they line. Two such layers lying one on
each side of an intermediate colorless region thus form the wall
separating the cavities of two adjacent chambers (fig. 6). The early
condition of this tuber-like body could not be determined, since all
the specimens examined were well matured, and showed no signs of
developing chambers. The body, however, appears to be in the
nature of a sclerotium, which from its spongy structure may possibly
serve the double purpose of supplying moisture as well as nutriment
to the developing apothecia.

An examination of the specimens of Wynnea gigantea in the Curtis
Herbarium shows that this species is characterized by a general habit
closely resembling that of the Carolina form; and the main stem,
Where present in these specimens, is evidently broken from some
attachment which may safely be assumed to have been a sclerotioid
body similar to that above described. The presence of such a body
might offer an additional reason for retaining BERKELEY’S genus, but
its characters seem otherwise quite sufficient to remove it from Mi-
dotis or other known discomycetous genera. In regard to Midotis
it may further be said that no one appears to have any definite knowl-
edge of this genus at the present time, and it is altogether doubtful
what the nature of this generic type really is. The name is first
mentioned by Fries (Syst. Orb. Veg. 363), without further allusion
'0 a specific form than the remark “ Species unica pleuropus;”’ while

~

244 BOTANICAL GAZETTE [APRIL

in the Elenchus (2:29), published five years later (1830), the first men-
tion of the type species, Midotis lingua, appears. This fungus,
which is described as growing ‘“‘ad basos truncorum” and as pos-
sessing an inferior hymenium, has not been found again, so far as is
known; and it seems quite uncertain what it may have been, or even
if it were anything more than some well-known form growing under
abnormal conditions. The only discomycete known to the writer
which might be referred to Midotis, and which possesses a truly
inferior hymenium, is a plant found growing on dead logs in Septem-
ber, 1889, near New Haven, and again at Burbank; the material in
the former locality being rather abundant. The apothecia are very
thin, broadly spathulate, proliferous, and fasciculate, having a habit
of growth very like that of some basidiomycete; the spores small and
insignificant, and the inferior hymenium very characteristic and by
no chance accidental. This plant, which has been provisionally
referred to M. plicata Cke. & Hark., may perhaps really be a Midotis
in the Friesian sense, on account of the unusual position of its hyme-
nium; but that the other species included in the genus by various
writers really belong here seems doubtful. The only species among
these for the most part tropical forms, that the writer has examined,
is the Cuban M. verruculosa B. & C., in which it is not possible to
determine whether the hymenium is inferior or not. The status of
Midotis itself being thus decidedly uncertain, it seems doubly unde-
sirable to include in it the members of what at least appears to be a
well-marked and peculiar genus.

Within the past year Mr. MAsseE has been so kind as to send to
the writer a small fragment of the type material of Wynnea macrotis
and W. gigantea, and, as has already been mentioned, he has eel
ined the specimens of the latter species in the Curtis Herbarium,
which form a part of the original gathering of Borrert, and include
one large and very well-developed example. None of these, how-
ever, correspond very closely to the figures given by BERKELEY and
Cooke, the apothecia, though somewhat more blunt than in the
Carolina form, hardly presenting the thin, broadly spathulate, and
freely proliferous habit represented in these drawings. Mr. MASSE
informs me that in his opinion the Berkeleyan species are not spe-
cifically distinct, although the color and habit of the two are s° very
differently represented in the Micrographia.

ahh age

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1905] THAXTER—NEW SPECIES OF WYNNEA 245

A comparative examination of the spores in all three species
shows that when fully mature all are characterized by the presence
of longitudinal markings, apparently corresponding to slight depres-
sions and somewhat roughened intervening elevated regions which
extend the whole length of the spores. Although these markings
are conspicuous in the Carolina species, they are hardly visible until
the numerous oily globules with which the spores are originally filled
(jig. 4, upper spore) have been obliterated by treatment with glycerine
or otherwise (fig. 4, two lower spores). While they are much less striking
in W. gigantea, they are nevertheless readily seen (fig. 8), but in the
material of W.macrotis received from Mr. MasseEE, they are exceedingly
faint, possibly owing to the fact that the spores may not have been
thoroughly matured. In all three species the spores are charac-
teristically inequilateral, being more strongly curved on one side
than on the other, apparent deviations from this rule being probably
in all cases due to differences in the point of view. While in W.
macrotis and W. gigantea they are bluntly rounded, they are more
or less conspicuously apiculate or papillate at each extremity in
the Carolina species, and the spore as a whole is distinctly larger.
The spores of W. macrotis sent by Mr. MassEE (fig. 7) are slightly
smaller than those of W. gigantea; but the difference is insignificant,
and could not serve to distinguish the species in the absence of other
distinctive characters. Nevertheless it seems more desirable to
retain both the Berkeleyan species until further data may be obtained
by a re-examination of fresh material.

For convenience of reference the original descriptions of BERKE-
LEY are quoted below. The genus may be characterized as
follows, the sclerotium of the Carolina form being assumed to occur
in the other species as well.

WYNNEA Berkeley & Curtis, Jour. Linn. Soc. 9:124. 1866.

Apothecia thick, firm, subcartilaginous, tough and subcoriaceous
on drying, erect, elongate, ear-shaped, simple or subproliferous,
several- to many-clustered on a common stalk arising from a sclerotium
buried in the substratum. Paraphyses cylindrical, simple or branched.
Asci cylindrical, tapering to an elongated base penetrating the sub-
hymenium with the filaments of which it is continuous, without
articulation. Spores large, inequilateral.

246 BOTANICAL GAZETTE [APRIL

WYNNEA GIGANTEA B. & C., Jour. Linn. Soc. g:124.. pl. 17.
fig. 31. Midotis gigantea Sacc. Sylloge 8:547.

‘Common stem three inches high, three-quarters of an inch thick,
deeply rugose and cracked, so that the surface resembles that of
Opegraphae; above divided repeatedly, with subdivisions elongated
into ear-shaped cups, which are smooth externally, but wrinkled,
though not cracked like the stem when dry; the cups are from two
and one-half to three inches long, with incurved margins variously
divided and sometimes proliferous. Asci cylindrical, containing
eight subcymbiform spores .o00g5 inch long, and more or less obtuse
at either end. When steeped in water, the inside of the stem acquires
a slight foxy tinge. The substance is so totally different from Peziza,
though this curious fungus is closely allied to P. leporina and P.
onotica, that it cannot be placed in the same genus. Peziza macrotis
Berk., a species found abundantly at Darjeeling at 7,500 feet, is clearly
congeneric and may be characterized Wynnea macrotis Berk., etc.”

Collected near Orizaba, Mexico, by Borrert, without note as to substratum.

WYNNEA MACRoTIS Berk.

Peziza macrotis Berk., Hook. Jour. Bot. & Kew Gard. Misc.
3:203. 1851. Wynnea macrotis Berk., Jour. Linn. Soc. 9:124- 1866.
Midotis macrotis Sacc., Sylloge 8:547.

“Inodorous, dry, firm, leathery, subcartilaginous, varying in size,
sometimes five inches long; erect, tufted, connate below and thence
branched; cups elongated, oblique auriform, of a bright liver color,
smooth externally; margin subinvolute. Hymenium even, purplish.
Sporidia oblong-elliptic, with one side in general more conve:
Nucleus single, in dried specimens.”

On rotten wood Darjeeling, India, 7500*, June-July.

Wynnea americana, nov. sp.

Sclerotium tough, subgelatinous, coriaceous on drying, irreg-
ularly lobed, variable in size, 50x 40™™ more or less, brown. Main
axis becoming variously divided above almost immediately after
emerging from the ground, the short divisions giving rise at once
clusters of apothecia of variable size and number. Apothecia sever al
to about twenty-five on a single plant, typically simple, rarely —
liferous, erect, elongate ear-shaped, very variable in size; the longest
seen 130X60™™, the average about 80™™, the margins somewhat

PLATE IV

THAXTER on WYNNEA

+» KRIEGER, PINX.

1. €

BOTANICAL GAZETTE, XXXIX

ee ae ee aati = eee aie - ies a Se las Fe eae Sere

PLATE 'V.

BOTANICAL GAZETTE, XXXIX

g
;
Z
:

1905] THAXTER—NEW SPECIES OF WYNNEA 247

involute on drying, the external surface rich blackish-brown, finely
verruculose, the roughness due to projecting groups of irregular
parenchyma-like cells becoming somewhat hair or chain-like toward
the margins. Hymenium even, dark dull purplish-red, or brown.
Asci elongate, about 500-540 u long, the sporiferous part cylindrical,
about 18 # in diameter. Paraphyses septate, simple or irregularly
branched, clavate, the slightly brownish terminal enlargement more
abruptly and conspicuously distinguished in older specimens. Spores
eight, subcymbiform, 32-40 X 15-16 #, the extremities apiculate or
papillate; marked when mature by about eight roughened areas
running longitudinally and separated by a corresponding number
of smooth slightly depressed areas, the spore when fresh completely
filled by numerous small round refractive oily masses.

Growing on the ground in rich woods, Burbank, East Tennessee, and Cran-
berry, North Carolina.

Professor DuRAND has kindly allowed me to examine a specimen
collected by E. WitxKrnson at Mansfield, Ohio, and communicated
by Professor KELLERMAN, which corresponds in all respects with
the Carolina plant. The spores of BERKELEY’s species differ in
shape and in their smaller size; those of W. gigantea measuring
25-30 X12 m, and those of W. macrotis received from Mr. MASSEE
20-25 X 11-12 #. Theapothecia of W. gigantea seem also to be some-
what smaller, more rounded distally, more numerous in a single
plant, and distinguished by their much paler color. .

HARVARD UNIVERSITY.

EXPLANATION OF PLATES IV AND V.

Fics. 1-6.—Wynnea americana Thaxter.
1G. 1. General habit of a well-developed plant. For the original of this
- figure, done in color by Mr. Krrecer from a formalin specimen and colored
sketches of my own from fresh material, I am indebted to Professor FARLOW.
Fic. 2. Paraphyses.
1G. 3. An ascus showing connection with subhymenial filament.
Fic. 4. Three spores, the upper in optical section.
Fic. 5. Portion of a section of the subterranean sclerotium showing chambers
and cortex. Leitz obj. C+oc. 1. |
Fic. 6. Portion of walls between two chambers. Leitz obj. 1+0¢. 1.
1G. 7. W. macrotis Berk. Three spores, two in optical section.
Fic. 8. W. gigantea Berk. & Curt. Three spores, one in optical section.
The spores were drawn with Zeiss apochr. 1.25™™ and 4 ocular.

ON THE DEVELOPMENT OF HAMAMELIS VIRGINIANA.
CONTRIBUTIONS FROM THE BOTANICAL LABORATORY OF
JOHNS HOPKINS UNIVERSITY, No. 3.

D. N. SHOEMAKER.

(WITH PLATES VI AND VII)

INTRODUCTION.

Tuts work on Hamamelis was undertaken on account of its
peculiar habit of flowering. It is one of the few angiosperms that
flowers in the fall and matures fruit the following year. This
peculiarity made it seem worth while to investigate its entire embryo-
logical history, and especially the behavior of the pollen tube and the
time and manner of fertilization. The subject was suggested by
Dr. D. S. JoHNson, to whom I wish to express my gratitude for
sympathetic guidance and instruction during its prosecution.

The literature of the family is not extensive, and none of it has
to do with the embryology of any of its forms. The most complete
working out of the anatomy and affinities is by REmNscH (12). Batt-
LON (3) carefully described the organogeny of the flower in Ham-
amelis virginiana and Fothergilla Gardeni. VAN ‘TIEGHEM (14)
worked on the secretory canals of Liquidambar and Altingia. THOU-
VENIN (13) described the structure of the root, stem, and leaves of
various members of the family. The account given by NIEDENZU in
Engler and Prantl’s Natiirlichen Pflanzenfamilien is the most com
plete I have seen.

In some American oaks, which require two years to mature seed,
it has been found that fertilization takes place about a year after
pollination. The statement is made by GOEBEL (15, P- 392) that 4
period of rest occurs after the pollen tube has reached the embryo
sac in Ulmus, Quercus, Fagus, Juglans, Citrus, Aesculus, Acer,
Cornus, and Robinia. As Miss BENSON (4) points out, this sane
ment is erroneous in the case of British Amentiferae. It is not true
of Hamameli

In Colchicum autumnale, according to HOFMEISTER (7), the pollen

248 [APRIL

1905] SHOEMAKER—HAMAMELIS VIRGINIANA 249

tube reaches the embryo sac at the latest by the beginning of Novem-
ber, and it is not until May of the next year that the embryo begins
to form. This plant and the autumn-flowering species of Crocus,
like Hamamelis virginiana, bloom and shed their pollen in the fall.
HOFMEISTER’s account of Colchicum shows that it differs essentially
from Hamamelis in the behavior of the pollen tube and the time of
fertilization. The other genera named as having a longer or shorter
period between pollination and the beginning of embryonic growth
are all pollinated in the spring, so that the resting period for most
of them does not extend through the winter.

The family of the Hamamelidaceae comprises some fifty species,
in eighteen genera, of which eight are monotypic. North America
has three representative genera: Hamamelis, Liquidambar, and
Fothergilla. The first is found nearly always in such sheltered places
as harbor Polystichum acrostichoides, from Labrador to Florida and
west to the Mississippi River. The second is found on the coastal
plain and on bottom lands from New York to Florida, thence to
Central America, and up the Mississippi River to southern Indiana.
The third is found from Virginia to Florida, east of the Appalachian
Mountains. Hamamelis has two moré species, one in southern
China and one in Japan; Liquidambar is also represented by two
species which occur in southern Asia from Asia Minor to Formosa;
and Fothergilla is represented by one other species in Persia. The
remaining genera are confined to southern and eastern Asia and
Malaysia, with the exception of three genera found in Madagascar
and southern Africa. Thus the whole family, with the exception of
three genera, is confined to the eastern and southern parts of North
America and Asia. This peculiar and as yet unexplained distribution
occurs in a large number of genera, but this family is one of the most
Pronounced cases found. It should also be said that Hamamelites
and Parrotia are found in the Dakota group of the Cretaceous (9),
and that Hamamelis (5) and Hamamelidanthium are found in Eocene
strata in northern Europe, so that the family had formerly a much
more extended range than at present.

I have usually found Hamamelis on rather steep hillsides with a
horthern exposure, or more rarely on low ground along streams.
It is said to grow abundantly on mountain tops in Pennsylvania

250 BOTANICAL GAZETTE {APRIL

and western Maryland, and CowLes (6) reports it as growing on sand
dunes near Chicago. From his description, however, it is doubtful
if the seed had germinated in these dry localities. It has thus a very
restricted range, and seems to grow only in positions in which moisture
is abundant and transpiration slow. Liquidambar is confined to
alluvial soils and moist situations, and so follows river valleys and coast
lines; its northern limit is the valley of the Hudson River in New
York. Fothergilla grows in moist places.

The material of Hamamelis virginiana and Liquidambar for the
present work was obtained from the .region about Baltimore, from
northern Virginia, from Long Island, and from southwestern Ohio.
I am indebted for exotic forms to the kindness of Mr. Gro. V. NASH,
of the New York Botanical Gardens, and to Mr. J. G. Jack of the
Arnold Arboretum, Jamaica Plain, Mass. The long period of
development made it necessary to collect nearly every week of the
_ year for Hamamelis. Material of Fothergilla was collected in
central South Carolina.

Killing and fixing was mostly done with a sublimate-acetic mixture
made by adding 5 per cent. glacial acetic acid to a saturated aqueous
solution of corrosive sublimate; this was often used hot. The
material was cleared in xylol and imbedded in paraffin; and sections
were cut 5 to 15 in thickness. For staining, a combination of
haem-alum and Bismarck brown was tried, but Flemming’s triple
stain was found more satisfactory and was used almost exclusively.
It was necessary before the young carpels could be sectioned '
paraffin to remove carefully and laboriously the hairs from their
bases, as on account of their thick walls the hairs could not be cut
in paraffin, invariably tearing the sections. To avoid this process
celloidin imbedding was used in a few instances, but it was impossible
to get the sections thin enough for most purposes.

ORGANOGENY OF THE FLOWER.
The flower buds arise from axils of the leaves of the current yeah
or from latent buds of the two preceding years, and appear early
May. They, as well as the leaf buds and the young twigs, are covered
by a dense coating of tufted hairs. Each hair develops from a single
epidermal cell, which protrudes from the surface and is cut by antl-

1905] SHOEMAKER—HAMAMELIS VIRGINIANA 251

clinal walls into four to twenty cells, each of which sends out a
long process, making a many-rayed star. They are often raised on
a low multicellular papilla.

Each flower bud produces a head of two to four flowers, and
there are often as many as three buds from an axil. At first the
tip of each bud is protected by three or four alternate bracts, which
are soon left below on the stem of the flower head and finally fall off.
The first floral organ to appear is the outside bract of each flower.
As the buds grow the other two bracts arise successively, one on each
side of the flower. The sepals appear in pairs, the first pair being
anterior and posterior. The petals then arise in one cycle of four
rudiments, inside which two successive alternating cycles of four
rudiments each develop. The outer cycle, opposite to the lobes of
the calyx, becomes stamens, the inner sterile staminodes.

The torus by this time, through unequal growth, has become
concave, and on its floor are developed two horseshoe-shaped ridges,
one anterior and one posterior. These grow together on the median
line, and this line of fusion is carried upward by growth, so that there
is a solid wall between the cavities of the carpels for a short distance.
The carpels have separate styles and stigmas, but are united through-
out their hollow portions. In each ovary there is found one ovule,
which is suspended from the margin of the carpel. This develop-
ment of the flower is essentially as described by BATLLON (3), except
that he describes the ovary as originally having two ovules, one of
Which nearly always atrophies. Le Maour and DEcaIsNE (16)
also figure a cross-section of the fruit showing two mature seeds in
each carpel. Although I have examined several hundred carpels I
have found but a single one with two ovules: BarLLon (17) also
speaks of Hamamelis as being polygamous, but of this I have seen no
evidence in my material. It is possible that these conditions may
occur more frequently in places from which I have no material, or
In other surroundings, yet the form has been very constant from all
my collecting points.

n the mature flower the temporary parts, the stamens, petals, and
nectaries, are smooth. The outside of the sepals and bracts, and
the bases of the carpels are thickly covered with hairs. Figs. 8 and
9 show how the rudiments of the growing flower fit together, and

252 BOTANICAL GAZETTE [APRIL

how the bud is protected by its hairy covering. Most of these hairs
have a double function. While young they act as slime cells in
keeping the growing point and growing tissues moist, this function
being best performed by the hairs on the tips of the sepals and bracts,
and the bases of the carpels. These young hairs are long and tortu-
ous, and wind among the growing rudiments (fig. 9). Their cavities
are full of sap, and their nuclei take a very dark stain; they remain
in this active stage longer than hairs on other parts of the plant. As
they grow older all the hairs acquire thick cell walls, and tend to
straighten. In their mature state they act as a protection against
moisture. This function they perform by means of a waxy covering
which repels water, so that it is very difficult to moisten a young
flower bud, or a growing twig, or fruit; but if these hair-covered parts
be soaked a short time in strong alcohol, and allowed to become dry
again, they may be very readily moistened.

POLLEN SACS AND POLLEN.

Each fertile stamen rudiment begins early to form two microspo-
rangia. There is apparently no evidence of the presence, at any stage;
of the other two microsporangia that are usually found in angio-
sperms. No indication of pollen-formation is ever seen in the sterile
rudiments.

The first evidence of the formation of the archesporium is found
about the middle of June. The subepidermal layer divides by pet!
clinal walls at the place where the microsporangia are to be formed
(fig. 1). The exact derivation of layers is hard to trace, but it is quite
certain that it is from the inner layer thus formed that all the arche-
sporial tissue comes. By the middle of July the archesporium is well
blocked out, and shortly after the spore mother-cells are formed. At
this stage there is about them a moderately well-defined tapetu™
and the outside wall of the microsporangium is three or four ce
layers in thickness (fig. 2). Here the pollen mother-cells are only
noticeable by their slightly larger nuclei and more deeply staining
contents.

The further growth of the microsporangium is brought about by
the increase in size of its cells, both sporogenous and tapetal.
Before the tetrad division, the nuclei of the tapetum divide without

;

1905] SHOEMAKER—HAMAMELIS VIRGINIANA 253

the formation of cell walls, and the tapetal cells have two or three
nuclei each; the nucleoli also increase notably, and the contents of
the cell become more largely vacuolate (fig. 3). The pollen mother-
cells thicken their walls, and soon float freely in the cavity of the
anther ; increasing further in size, with a much larger nucleus
and nucleolus (fig. 29). The two tetrad divisions occur almost
simultaneously, although stages are found showing two nuclei
(fig. 30). In the tetrad division the nuclear processes do not show
at all clearly, though there seem to be fibers connecting the nuclei in
one stage observed (fig. 31). The pollen grains have cell walls and
but one nucleus when the mother-cell wall is broken down (fig. 32)
and the grains released into the cavity of the microsporangium. The
mature grain was described by Von Mout, very briefly, in 1835. It
has the shape of an oblate spheroid, with three meridional furrows
(fig. 33) ; between these furrows the surface is covered by a fine
reticulation. An equatorial section shows that the intine is strongly
developed under the furrows, which gives the section of the interior
a decidedly three-lobed appearance (figs. 33 and 34).

Soon after the pollen grain is freed from the mother-cell, its nucleus
divides, and the smaller nucleus, probably the generative, retires into
the extremity of one of the lobes. Here it becomes closely applied
to the intine and is cut off from the larger cell by a very noticeable
wall, which is probably of cellulose (fig. 33). Shortly before the pollen
is shed this wall disappears, and the two nuclei then lie free in the
Cavity of the grain. The larger of the two, the tube nucleus, is loosely
vesicular, while the structure of the generative nucleus is dense and
deeply Staining (fig. 34). ;

Of the four layers in the microsporangium wall only the subepider-
mal layer has any part in the opening of the anther, becoming the
fibrous layer and covering the inner and lateral faces of the anther
(figs. 5 and 6). The first and for a long time the only evidence of the
differentiation of this layer is the radial lengthening of the cells. At
this stage it is possibly not inappropriate to mention an instance of
tegeneration observed in this fibrous layer. From some unknown
Cause, the first subepidermal layer had been destroyed over a small
area. The remaining part of that layer had developed normally.
Into this gap tissue had grown from both sides, but that which came

254 . BOTANICAL GAZETTE _ [APRIL

from below was slight in amount, and had retained its original char-
acter; while the epidermal cells had greatly elongated and had divided
into epidermal and secondary fibrous layer cells (fig. 4). These sec-
ondary fibrous layer cells resembled very closely at this stage those
of the primary fibrous layer, and I sec no reason to suppose that they
would not have developed fibers at the proper time. Here then the
epidermis seems to have been more plastic under the injured spot
than the deeper lying tissues.

The fibers are developed in this layer shortly before the time the
anther is to open, appearing on the side and bottom of each cell.
Around the top, posterior, and bottom of the microsporangium there
is a groove (figs. 5 and 7), and in the bottom of it the cells have no
fibers and are quite thin-walled, so that they readily break, thus form-
ing the line of dehiscence of the anther. As LECLERC DU SABLON (8)
has shown for anthers in general, the opening is due to unequal
shrinkage of the two walls of the fibrous layer. The outer wall being
of cellulose, shrinks more on drying than the inner wall, which is
strengthened by its lignified fibers. By this means the whole outer
covering is bent upon itself and points directly toward the carpels
(fig. 7); nearly all of the pollen adheres to these wings, and so is placed
in the way of any insect that comes to visit the nectaries. The
stigmas are ripe for pollination at the same time that the anthers open
Any insect visiting many flowers in succession must scatter pollen
promiscuously, so that there is sufficient adaptation to insure cTOss
pollination, but no well-developed mechanism to prevent self-pollina-
tion. Pollen is ripe and begins to be shed in the latter part of October,
and is shed from that time on into the winter, as the flowers keep
opening with each return of warm weather, even as late as January.

OVULES AND EMBRYO SAC.

The ovules show specialization of archesporial tissue at the tip of
the nucellus before the integuments have begun to develop, but #t
very difficult to distinguish archesporial cells, although there a
to be several of these, each one cutting off a tapetal cell which divides
repeatedly. The cells of this tissue then elongate in the direction of
the long axis of the nucellus, and bury the mother-cells by some eight
layers of cells in the nucellus.

™ SE
wy

1905] SHOEMAKER—HAMAMELIS VIRGINIANA 255

It is also probable that this part of the nucellus forms the extremity
of the conducting tissue for the pollen tube. In early stages
three to five macrospores can be seen, but only one germinates. This
goes through the usual stages, the resulting nuclei being arranged in
the order most often found among angiosperms (fig. 10). The antip-
odals very early disappear, so that they are hard to find at the time
of fertilization. The endosperm nucleus is found about the middle
of the sac.

The tissue surrounding the mature embryo sac is much disinte-
grated (jig.r0). Around the chalazal end of the sac the tissue always
stains deeply, and there is a quite evident strand of conductive tissue
from the tip of the fibrovascular bundle at the chalaza to the lower
end of the embryo sac (fig. 10). The base of the nucellus shows by its
smaller cells that it is the most rapidly growing part. The develop-
ment which the ovule has attained at the beginning of winter is shown
in fig. rz. The integuments up to the early part of April are still
behind the nucellus in growth; in spring their growth is hastened and
they soon project beyond the nucellus, leaving a wide open micropyle
(fig.21). This is finally closed to a slit-like fissure between the edges of
the outer integument.

The outer integument is now quite thick, formed of about eight
layers of cells, and is uniform in structure throughout. The inner
integument is made up of three layers of cells which are very much
alike. The epidermal layer of the nucellus has already become
slightly different from the underlying tissue.

POLLEN TUBES ARTIFICIALLY GROWN.

Pollen taken from open anthers was very readily sprouted in a 16
ber cent. sugar solution made with tap water, in which 1.5 per cent.
gelatin was dissolved. The pollen grains first became spherical by
, € filling out of the furrows, thus using the masses of intine on the
tner sides of the furrows. Tubes sprout from the pollen grains when
Placed in the nutrient gelatin in one to three hours, and always
arise from the smooth bands. The cultures were kept at room tem-
Perature and growth was more luxuriant in the dark. The growth
of molds, etc., usually disturbed the cultures at the end of a few days.

The behavior of the nuclei was not readily observed. Methyl-
Steen-acetic-acid was used to kill the pollen tubes, and no more

256 BOTANICAL GAZETTE [APRIL

than two nuclei were ever found in a tube. The tubes showed a
marked tendency toward the formation of cellulose plugs (jig. 12).
It rarely occurred that part of the contents of the tube was in this way
shut off, as the spaces walled off by plugs were mostly empty.

In the course of about three days’ growth the pollen tube frequently
‘“encysted” (fig. 35), that is a spherical swelling developed at the tip
or near the tip of the tube into which nearly all the contents of the
tube were withdrawn, including one or both nuclei. A wall was then
formed completely closing off the swelling, which was often as large
as the original grain. I have not determined the exact conditions
which called forth this action, neither have I found such things in the
style in normally grown pollen tubes. Miss BENSON (4) reports a case
of somewhat similar character as occurring in Carpinus, though the
spherical character and the separating wall were not nearly so pro-
nounced. She suggests that this may be of use in the short resting
period of this form, but was not able to find such appearances in the
style.

The pollen showed ability to sprout at room temperature whenever
the flowers opened. A collection made in Ohio early in January;
after the unopened buds had endured a week of very cold weather,

~when the temperature had been as cold as —15° F., sprouted ina
seemingly normal way, though not so vigorously as earlier. I am
inclined to think, however, that pollen shed so late never functions.

CONDUCTIVE TISSUE OF THE STYLE AND OVARY.

By the folding of the carpels each style has a groove fo
which leads from the stigma to the ovary. It is open at the top, and
where its sides are slightly separated and its inner surface thus
exposed, it bears the loosely arranged papillose cells of the stigma.
The epidermis of the groove and two subepidermal layers continue
this stigmatic tissue down to the base of the funiculus, the strand 0
conducting cells getting gradually deeper and deeper in the tissue
of the style. The cell walls of the strand are thickened, partly
ized, and the contents of the cells are dense. Fig. 13 shows its appea
-ance and position about midway in the height of the style, ae
course in longitudinal section is shown in fig. 18. When the omg
first opens, the epidermis of the funiculus is not yet differentiate”

rmed,

ae
gelatl
pear-

1905] SHOEMAKER—HAMAMELIS VIRGINIANA 257

but as winter approaches the base of the funiculus becomes glandular.
The epidermal cells enlarge, the walls thicken, and the contents
become vacuolated. In the spring this process is carried still farther,
and as the ovule occupies more and more of the cavity of the ovary,
these cells secrete a mucus which fills the small remaining cavity
around the base of the funiculus (jig. 10).

THE DESCENT OF THE POLLEN TUBE.

The pollen grains begin growth very shortly after falling on the
stigma, and the growth is at first comparatively rapid. Its course
is readily traced in the conducting tissue of the style, which is greatly
disorganized by the growth of the large number of tubes usually
present. The course is between the cells rather than through them.
By the time that winter sets in the living part of one or more tubes is
to be found in the neighborhood of the base of the funiculus (fig.
11). There are usually several living tubes at varying heights in
the style at this time ( figs. 17 and 18), and evidence of many more
which have. been stranded above. The unprotected tip of the style
is dead and withered, while that part which is clothed with hairs is
alive. It is in this protected part of the carpels that the pollen tubes
hibernate. Soon after pollination the flower head twists on its stalk
so as to invert the blossoms. The inverted calyx then very effectually
Protects the carpels from rain, sleet, and snow.

The pollen tubes found at this time are usually of greater diameter
than at the beginning of growth. Tubes grown in sugar-gelatin
solutions are 5 to 8 w in diameter, and those which sprout on the
stigma are at first about the same size; but those found during the
resting stage are 12 to 15 # in cross-section, and the wall is thicker
than. at first (fig. 14). The nuclei found did not exceed two in any
tube.

When growth is renewed in the spring, the area of conducting
tissue in the funiculus being increased, the pollen tube is soon seen
In the cavity of the ovary. More than one tube may reach the ovary.
At first the ovule is by no means ready for fertilization, and the
integuments have not yet closed up the micropyle (fig. 21). The
tubes do not appear at this time to have any definite direction of
Stowth, but grow down beside the ovule or into the.wide open

258 BOTANICAL GAZETTE [APRIL

micropyle, or between the integuments. The course to the egg cell
is through the micropyle and into the tip of the nucellus through the
tissue derived from the tapetal cells, which stains deeply at the time
and is probably conductive tissue. Thus it is seen that the tube
grows just about as fast as the conductive tissue is prepared for it,
and stops in the fall when it reaches the end of the mature conductive
strand. The transference of the male nucleus has not been observed,
but fertilization takes place about the middle of May, which is from
five to seven months after pollination.

ENDOSPERM.

The antipodals very early disappear. The first result of fer-
tilization is apparent in the action of the endosperm nucleus, which
immediately begins to divide. The stage of free endosperm nuclei
is very short, as cell walls have appeared in the twelve-nucleate
stage. These walls first arise in the bottom of the embryo sac.
Both endosperm and nucellus grow rapidly from this time forward.
The endosperm early disintegrates the neighboring nucellar tissue
except at two points, the tapetal strand of tissue leading down from
the micropyle and bearing the fertilized egg at its lower end, and the
pit at the chalazal end of the embryo sac which earlier held the
antipodals. This and the deeply staining tissue surrounding it
resist the action of the endosperm for some time, and by the growth
of the base of the nucellus are pushed into a position on the side of
the growing endosperm (fig. 23). This antipodal pit is finally
absorbed.

The nucellus keeps pace with the growing endosperm,
layer being changed to make part of the inner seed coat.
entiation of this layer is shown mainly by the larger size of its
especially at the tip of the nucellus, and by the crowded cell contents
which take up blue stains very readily. This layer is the only part
of the nucellus that permanently resists the action of the endosperm
and it is completed across the region of the chalaza so that it entirely
surrounds the endosperm. Its nuclei are usually applied to the outer
cell wall, and help doubtless in making the clear membrane whic
surrounds the nucellus in the mature seed. It is thrown into folds
shortly before the ripening of the seed (fig. 20). The endosperm

its epidermal
The differ-
cells,

1905] SHOEMAKER—HAMAMELIS VIRGINIANA 259

is finally stored with food in the form of proteid grains, which shortly
before ripening show numerous globoids (fig. 22) that disappear later.
The ripe endosperm contains in its cells much oil along with proteid
material, and in the cell walls there are imbedded numerous crystals
of calcium oxalate (fig. 15).

THE EMBRYO.

The embryo begins growth comparatively late, so that the endo-
sperm has already acquired some size before the first division of the
egg occurs. The egg after fertilization becomes slightly imbedded
in the tissue of the tapetal strand, and enlarges greatly. The first
division is transverse and cuts off a small cell below and a large one
above. By this continued cross division the suspensor may have
five or six cells (fig. 23). The first division of the embryo is longi-
tudinal. The embryo dissolves the endosperm in much the same
Way as the endosperm dissolves the nucellus, and lies free in a cavity
formed by the disintegration of the endosperm. At maturity there
isa Straight axial embryo which extends from end to end of the seed
and is richly stored with oil and proteid material in all its cells.
The upper side of the cotyledons has already a well-developed palisade
layer (fig. 24).

INTEGUMENTS.

The inner integument is at the beginning three cell layers in
_ thickness, the innermost layer taking part in the formation of the
inner seed coat. It early becomes filled with dense contents that
stain blue readily (fig. 20), and finally shrink and become applied
to the inner cell wall. The remaining layers are crumpled up so
that they can only be made out with difficulty.

The outer integument thickens greatly, and its cells elongate,
taking a curved oblong shape. The cell walls then begin to thicken
and the whole integument forms the outer seed coat which is moder-
ately hard, black, and very resistant to water. The outer integument
1S very smooth over the whole surface, except at the place of attach-
ment of the funiculus, where there is a white saddle-shaped scar.
The seed is ovoid, but very decidedly pointed at the lower end.
The shape of the seed is a very important part of the discharging
mechanism,

260 BOTANICAL GAZETTE [APRIL

THE CARPELS.

At blooming time the carpels are very slightly imbedded in the
tissue of the torus (fig. 17). There is a very short calyx tube, how-
ever, shown in the figure below the attachment of the anther. As
the fruit matures, the calyx tube lengthens proportionately more
than the carpels, and this gives the fruit the appearance of being half
buried in the torus (fig. 38). A longitudinal section (fig. 39) shows
that this is only apparent and that the fruit is only slightly buried.

The substance of the carpels develops into two tissues, the outer
one becoming fleshy with numerous roundish stone cells, the inner one
forming part of the mechanism for expelling the seed. In eac
carpel this inner Jayer is formed in two halves, which are not closed
at the top and are higher toward the posterior of each carpel, as
shown in side view in fig. 42. These halves are never closely joined
on the inner sides of the carpels, and there is provision for a split
along the dorsal line also. The cells are developed into fibers diag-
onally from the inner edge to the dorsal line of each carpel, parallel
to the top of the layer. In opening, this layer splits down the midrib
of the carpel and in front. It then opens at the top and each half
below begins to contract in a transverse direction. The cros*
section of the opening layer is shown before contraction in fig. 49%
after the opening of the fruit in fig. gz. The pressure exerted comes
gradually on the seed, and it is thrown out, not by a sudden move-
ment of the capsule, as in many such contrivances, but by being
pinched on the smooth pointed lower end. The great smoothness
of the seed and of the inside of the capsule assists greatly in the proces
The seeds are often thrown to a distance of twenty feet. This move-
ment is caused by drying, as can be proved by placing an opened
capsule in water, when after some hours it will close entirely, and
will open again on being dried.

, GERMINATION.

The seed, thus distributed lies on the ground for two —
according to Bartey (x), sprouting the second year. Under —
which fruited abundantly in the fall of rgor, but where the crop
a failure in 1900, it was not possible to find young seedlings ™ —
1902, though many seeds were found. Under trees which fruite

1905] SHOEMAKER—HAMAMELIS VIRGINIANA 261

in 1900 it was easy to get young seedlings in various stages. The
cotyledons remain in the seed coats until they have absorbed the
stored up nourishment of the endosperm (jig. 16); they are then
freed and exposed as green assimilative leaves. Attempts at sprout-
ing the seed in damp sphagnum were made in the laboratory. The
seed was planted in September of 1900, and by May of 1902 had just
begun to protrude the tips of the radicles. They had been in the
temperature of an unheated room constantly, but had not been
subject to frost, and had never been allowed to dry out.

_HAMAMELIS ARBOREA. |

I procured one stage of the Japanese species H. arborea inthe
latter end of October. This differs from H. virginiana in its time
of flowering, which is in very early spring. “A variety, H. arborea
succariniana, flowers as early as February, and thus approaches
the flowering time of the American species. The flowers are in
about the same condition in October as those of H. virginiana,
except that the stamens are rather backward. The pollen grains
are free and have each two free nuclei, and evidently pass the winter
in that stage (fig. 36). As the pollen is shed in March at the latest,
it probably must rest about two months before fertilization occurs.
The petals are coiled involutely in the bud as in H. virginiana, but
instead of being entirely smooth have a tuft of hairs on the tips. In
other respects the two genera are much alike in their development
So far as studied.

FOTHERGILLA GARDENI.

An incomplete series of stages of this species was studied. Its
flowers appear in the spring along with the leaves. It lacks a corolla,
and its calyx tube is much longer than that of Hamamelis, having
five to seven very small teeth. The development of the stamens
's described by Barton (3). They arise first as five single rudi-
ments, which are followed by other rudiments on each side, so that
there are finally five groups of five or six stamens, those of each group
being of different ages and different heights. They pass the winter
In the pollen mother-cell stage. At the time of flowering the ovules
are not yet ready for fertilization, so that the pollen must have a resting
Period of nearly a week. The anthers have four microsporangia and

262 BOTANICAL GAZETTE [APRIL

open by two valves instead of one, very much as is common in most
angiosperm stamens which open by slits. The structure of the seed
and fruit is like that of Hamamelis, except that the seed is smaller.

CORYLOPSIS PAUCIFLORA.

Only one stage of this species was examined. It was obtained
in the spring before the flowers open,which occurs before the leaves
appear. They are borne in drooping racemes, with many bracts,
which are smooth outside, but covered by silky hairs within. The
structure of these hairs is much like that in Hamamelis, but they are
not rigid. The stamens, which have four microsporangia, pass the
winter containing nearly mature pollen grains (fig. 37), with two free
nuclei. The ovule passes the winter in the same stage as does that
of Hamamelis (fig. 25). It is difficult to determine whether there
is present a definitive macrospore or a macrospore mother-cell; how-
ever, there is no evidence of the presence of more than one macrospore
mother-cell. There must also be some time here between pollination
and fertilization.

LIQUIDAMBAR STYRACIFLUA.

Liquidambar is not so closely related to Hamamelis as the other
genera studied. The buds in this species pass the winter with the
merest rudiments of the floral organs present. The stamens are only
small protuberances which do not show any archesporium. The
mature anthers have four microsporangia and open by slits (AS.
26). The fibrous layer is very slightly developed as compared with
Hamamelis. The flowers are imperfect, with rudiments of stamens
appearing as nectaries among the flowers in the female heads. These
were formerly thought to be both petals and stamens; abortive
pollen is sometimes developed in them, which is evidence of their
staminal nature (fig. 27). The carpels, which occur in pair a
in Hamamelis, are collected into large heads containing aid
five to fifty flowers each. Each carpel has a double row of ovules
developed on marginal placentae, and a broadly expanded stigmatic
surface. With very rare exceptions, only one of these many ovules
is fertilized, and this one is near or at the bottom of the cavity- han
is a week or ten days between pollination and fertilization in this
The developing seed shows the same resistant tissue at the antipoda

3
a

1905] SHOEMAKER—HAMAMELIS VIRGINIANA 263

end of the embryo sac as is found in Hamamelis, but here it persists
into the ripe seed. The epidermal layer of the integument is not
used up, and around the chalaza there is a small fragment of the
nucellus left in the ripe séd. This is never stored with food mate-
rials, and so cannot be called perisperm (fig. 28). The macrospore
is buried about as deeply as in Hamamelis. It germinates only in
the lower ovules, the upper ones never showing typical embryo sacs
and being less developed progressively toward the top of the ovary.
In the sterile ovules the cells of the outer integument become very
much enlarged and at last empty. The substance of the nucellus
is absorbed, and the ovules become polygonal bodies, resembling saw-
dust, which fill the upper part of the ovary. The outer integument
of the fertile ovule (fig. 28) grows into a wing. The embryo is
Straight, and in bulk bears about the same relation to the endo-
sperm as in Hamamelis.
SUMMARY AND CONCLUSION.

1. In Hamamelis the anthers have two microsporangia from the
beginning.

2. The generative cell in the pollen grain has a cell wall developed
Which is afterward dissolved.

3- The pollen tube when grown artificially shows a marked ten-
dency to form cellulose plugs, and also forms spheres into which the
Contents of the tube are withdrawn. Thus far these phenomena

ave not been observed in normal growth.

4. The development of the pollen tube in the style may be divided
into three periods: first period of growth, hibernation, and second
Period of growth. During hibernation the walls are thickened and
the diameter of the tube enlarged, in the next stage having a
smaller size and thinner walls.

5 There are several macrospores developed, only one of which
becomes functional. It is deeply buried in the nucellus by the growth
of tapetal tissue.

6. The germinating macrospore is nourished through a strand of
conductive tissue from the chalaza.

7- The antipodals are sunk in the tapering lower end of the
embryo sac. This tip is surrounded by deeply staining tissue which
for a time resists the dissolving action of the endosperm.

264 BOTANICAL GAZETTE {aprit

8. The epidermis of the nucellus is the only part not used up by
the endosperm. Its walls are thickened and it helps to form the
inner seed coat.

g. Fertilization takes place in May, five to. seven months after
pollination.

1o. The embryo is slow to begin growth and has a short sus-
pensor.

11. The seeds sprout normally after lying on the ground for two
winters.

12. The hairs serve a twofold function while young, to keep the
growing tissues moist, and when mature to keep off moisture.

13. One case of the regeneration of the fibrous layer of the anther
wall by the epidermal layer was observed.

14. The other investigated genera of the family all have a resting
stage of the pollen, although it is much shorter.

15. The other genera of the family all have anthers with four
microsporangia.

In comparing Hamamelis virginiana with its relatives, it seems
certain that ii was once a spring-flowering plant, whose blossom-
ing has worked backward through the winter. It differs from H.
arborea essentially in the way its pollen passes the winter, for the
development of each is much the same in October.

Most of the plants showing long resting periods in pollen growth
belong low in the system, in the Amentiferae; but with the exception
of some oaks Hamamelis has the longest resting period known. It
seems probable, therefore, that this resting period can not be regarded
as a primitive character in any case.

U. S. Bureau or PLant INDUSTRY,
Xas.

- LITERATURE CITED.
1. Barey, L. H., “Hamamelis” in Cyclopedia of American Horticulture. ae
2. BarLton, H., Observations sur les Saxifragées. Adansonia 5:297- 1864-1 ‘ 5:
3: , Nouvelles notes sur les Hamamélidées. Adansonia 10:120- 1871
1873.
4. BENSON, Marcaret, Contributions to the embryology of the Amentiferae-
. Part I Trans. Linn. Soc. Bot. II. 3: 409-427. 1894-
5. Conwentz, H. W., Die Flora des Bernsteins. 2:93. 1886.

BOTANICAL GAZETTE, XXXIX PLATE VI.

: Sa
a

%
i

; Af

=
:*

Si

AK}

Sy
acs bef WI
JE is ay 4) Minne
DS SCRE “408 .
BT Nat é
4 u . .
f te ‘a = sy ys = Oo .
COL OES
© CY 6
my, ;
\ SAS ;
ej
“ = ; a

, —_ Ay Se
‘ me cons sr) Carr: = ae

pee tthe

we = eh <a 2

Cores “tie ee
pee
ae

Paes

— = a i er)
i Sat i cease nee ,
a os BENE a

SN eee

ee nace

ay MY
i il

at Sa

‘\

(Iinieaee
iy

in

: \ HAVE

N® ees
au as
WY ‘i ede
\ \\ NX) ifn SED Tg a, ul
iN ANY % Hegrllte U
Nw seh
AN eee
AS Ss
SS \
oN NC!
SINAN
STR
Sot ¢ Sz
et
QR

SSS =
LO Se
“ee
SA Smee

Li
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ie

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ane
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oe
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4°,
74

vw

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PLATE Vii.

SHOEMAKER on HAMAMELIS

~

1905] * SHOEMAKER—HAMAMELIS VIRGINIANA 265

6. Cowtes, H. C., The ecological relations of the doen on the sand dunes
of Lake Michigan. Bor. GAZETTE 27:95. 1899.

7. Hormetster, W., Neue Beitrage zur Sone der Embryobildung der
facies: Abhandl. KGnigl. Sachs. Gesell. Wiss. vols. 6 and

8. LECLERC DU SaBLon, Recherches sur la structure et la déhiscence des anthéres.
Ann. Sci. Nat. Bot. VII. 1:97-134. 1885.

9. LEsQUEREUX, LEO, The flora of the Dakota group. Monographs of U. S.
Geological Survey 17. 1892. pp. 139-14I.

10. Mout, H. von, Sur la structure et les formes des grains de pollen. Ann.
Sci. Nat. Bot. II. 3:325. 5-

tI. es ice Notes on poanne and apr momine Trans. Linn.

:457. pl. 48. 1862

12. "ara me Ueber die wnstniiacle Verhiltnisse der Hamamelidaceae mit

Riicksicht auf ihre systematische Gruppirung. Bot. Jahrb. 11: 347-395.
88

~ 2, 1BBp.

13. THOUVENIN, Maurice, Recherches sur la structure des Saxifragées. Ann.
Sci. Nat. Bot. VII. 12:135-147. 1890.

14. VAN TIEGHEM, Px., Seconde mémoire sur les canaux sécréteurs des plantes.

n. Sci. Nat. Bot. VII. 1:1-96. 1885.

15. GOEBEL, K., Outlines of classification and, special morphology. 1887.

16. LE Maout Et DECAISNE, Traité général de botanique. 1865.

17. Battton, H., Histoire des plantes 3:389-465. 1872.

18. NIEEnzv, Fr. , Hamamelidaceae, rere and Prantl’s Natiirlichen Pflanzen-
familien 324: 115-130.

EXPLANATION OF PLATES VI AND VIL.

All figures were drawn with a Sale camera lucida, and are of Hamamelis
virginiana unless otherwise stated. The magnifications indicated are those of
i reduced plates; those of Plate VI having been reduced one-half, and those -

f Plate VIT one-fifth. The abbreviations are as follows. ar, archesporium;
is, br?, br3, first, second, and third bracts; c, carpels; ccs, chalazal conducting
strand; ct, condos tissue; em, embryo; en, endosperm; ep, epidermis; ¢s,
: embryo sac; j/, funiculus; ff, fibrous layer; gw, generative nucleus; 7, inner
integument; m, Ciera BN. n, nectaries; mu, nucellus; of, outer intégu-
ment; ov, ovules; p, petals; pm, pollen mother-cells; pf, pollen tube; s, sepals;
St, stamens; 1, peen: ae vascular bundle; vm, vegetative nucleus.

Fic Growing stamen; longitudinal section. * 550.

Lal

2. Growing stamen; trans section. X 550
Fic. 3. Tapetum and pollen mother-cells. < 550
Fic. 4. Anther wall showing a eration. X 220.
Fic. 5. Mature anther; matic transverse section. X 35-
Fic. 6. Mature anther eae longitudinal section. 35
Fic. 7. Mature anther, open; diagrammatic transverse section. X 35.

266

Fic.
Fic.

32h

BOTANICAL GAZETTE [apRIL

8. Growing flower bud; transverse section in plane ab in fig. 9. X 22.
9. Growing flower bud; long'tudinal section along line wy in fig. 8

. 10. Ovule; longitudinal section. * 97.

. 11. Ovule at beginning of winter. x 220.

. 12. Pollen tube with cellulose te grown in gelatin. X 330.
. 13. Style; transverse section. x 2

. 14. Hibernating pollen tube. X 550.

. 15. Endosperm cell walls with crystals. X 550.

16. Seedling in seed coats. X 1.
17. Flower in winter; longitudinal section. < 9

. 18. Carpels in winter; diagrammatic longitudinal section. * 20.

. 19. Ovule; longitudinal sections. x 35.

. 20. Seed coats. X 175.

. 21. Ovule in spring; longitudinal section. x 80.

. 22. Endosperm cell stored with food. * 550.

. 23. Embryo and suspensor. X 550.

. 24. Cotyledon in seed. X 220.

. 25. Nucellus of Corylopsis pauciflora in spring. * 55°.

. 26. Mature anther of .Liguidambar styraciflua; transverse section.

. 27. Nectary of the same; transverse section. * 35.

. 28. Seed of the same, nearly mature. X 5.

. 29. Pollen mother-cell. * 1745.

. 30. First division of pollen mother-cell. X 1745.

. 31. Second division of pollen mother-cells. 1745.

. 32. Pollen grain, in pollen mother-cell. < 1745-

. 33- Pollen grain. 1745.

. 34. Pollen grain. < 1745.

. 35- Pollen grain sprouted in gelatin. < 1745.

. 36. Pollen grain, winter condition; Hamamelis arborea. X 1745.
- 37. Pollen grain, winter condition; Corylopsis pauciflora, X 1745:
. 38. Ripe fruit. x 1

. 39. Fruit; Lspaitisdined section. X 1.6.

. 40. Fruit; transverse section. x 1.6.

. 41. Fruit, after discharge of seed; transverse section. % 1- 6.

- 42. Opening layer of capsule; side view. x 1.6.

pti i, a ape bore. ak i

SEXUAL REPRODUCTION IN THE RUSTS.
A. H. CHRISTMAN.

(WITH PLATE VII1)

THE nature of the aecidium has remained one of the most vexed
questions in mycology. BLACKMAN’s' recent discovery goes far
toward clearing up the most important points and my own results,
described below, confirm his general conclusions, though showing a
widely different method of conjugation. The literature relating to
sexuality and the general cell processes in the rusts has been so thor-
oughly and justly reviewed and critically estimated by BLACKMAN
that I need hardly devote further space to it here.

My studies on the winter conditions of the rusts and the nature
of the aecidium led me to the preparation of material for cytological
study, and in the spring of 1904 Professor J. C. ARTHUR proposed
to Professor HARPER to furnish material from his cultures for cyto-
logieal investigation. This material was turned over to me. It
consisted of a quantity of carefully identified aecidia of several rusts,
and since our knowledge of the method of origin of the binucleated
condition found in the aecidiospores was at that time entirely lacking
as to the details of the behavior of both cells and nuclei, my study
was directed largely toward that point.

It was soon found that aecidia of the caeoma type, with unlimited
Stowth, were by far the most favorable material. This type of
development apparently admits of less crowding of the hyphae, and
as a result the cells and their relations can be made out more easily.
Two forms were further found especially favorable because of the
large size of their cells and nuclei. These were Caeoma nitens S.
Stowing on Rubus cult., and Phragmidium speciosum Fr. on Rosa
humilis,

The best fixation was secured by the use of Flemming’s fixing solu-
tions; both the strong and weak were very satisfactory. Fixing
Solutions were sent to Professor ARTHUR and the specimens were

I

BLackman, V. H., On the fertilization, alternation of generations, and general
Cytology of the Uredineae. Annals of Botany 18:323-373- pls. 21-24. 1904.

“ 1995] 267

BO See td Ree ee et ee ee
oe st .

268 BOTANICAL GAZETTE [APRIL

put into them by him at Lafayette, Indiana, and mailed at once to
Madison. The usual time for fixation was from two to three days.
Flemming’s triple stain gave the best results. Good preparations
were also obtained by the use of Heidenhain’s haematoxylin.

In the case of Phragmidium, material was found of all ages,
so that the entire history from the uninucleated mycelial cells to the
fully formed binucleated spores could be easily followed. It was
found that in the young pustule the hyphae form a layer just beneath
the epidermal cells (fig. 1). This layer, in which the direction of
the individual hyphae is almost lost, is usually from one to three
cells thick. The mycelial cells here are much thicker and shorter
than those found in the vegetative hyphae. Each cell has a very
dense, finely granular cytoplasm and a single large nucleus. The
nucleus occupies a central position in the cell. There is a well-defined
nuclear membrane and one large nucleole., The chromatin is always
-more or less massed at this stage, making a ragged net with the
strands very irregular in thickness. Thus a great deal of clear space
is left in the nucleus, which is filled with nuclear sap. The chromatin
is regularly stained by the violet of the triple stain and the nucleole
stains a deep red.

Certain cells now become elongated in a direction perpendicular
to the epidermis, forming a continuous series and raising the epl
dermis (fig. 2). The exact origin of the cells of this series is hard to
determine; many of them can be seen to be end cells of short hyphal
branches. In thick sections, long branches may often be traced
between the subepidermal cells of the host, which terminate in one
of these cells. In the cacoma the cells are not crowded; there 's
even considerable intercellular space. Very often it may be seen
that cells standing side by side arise from different hyphal branches,
or at least from distant parts of the same hypha (jig. 2; 4 and 6).

The single nucleus of the cell now apparently divides and the cell
elongates into a rather narrower upper part which is cut off as 4
small distal cell from the larger basal cell. The cell division 1s very
unequal. The end cell as a result is only about one-third ee
of the cell beneath it (fig. 3). The protoplasm of the end cell is at
first dense and appears quite normal. It soon becomes very eee
however, and finally appears quite clear, with only a few strands 0 |

1905] 1 CHRISTMAN—SEXUAL REPRODUCTION IN RUSTS 269

granular material, and the whole cell dwindles in size. The chromatin
of the nucleus seems never to have passed into the resting condition,
but lies in dense masses, apparently as it was left on reaching the poles
of the spindle in the nuclear division preceding. The whole nucleus
is very small compared with the nucleus of the larger cell beneath
and contains little space between the chromatin masses. The
material of the nucleus begins to disorganize and soon becomes a
homogeneous mass, which stains a hazy red with the triple stain.

The cell beneath this terminal cell, on the other hand, begins to
enlarge, bulging at the middle, and becomes irregularly barrel-
shaped. The cytoplasm is dense with small vacuoles. The nucleus
enlarges greatly and the chromatin matter is distributed through
its interior, forming a ragged network. There is a large, well-defined
nucleole.

We have thus a series of oblong cells standing vertically side by
side but not much crowded. If we study a series from the margin
of a sorus to its center, we find gradually more advanced stages, so
that very many conditions of development may often be found in a
single section.

Up to this time in the history, the writer’s observations agree very
closely with those of BLACKMAN. The subsequent behavior of the
larger cells, however, which BLACKMAN terms the fertile cells, is very
different. Many times two can be found which incline toward each
other, coming in contact in a region on their adjacent walls. At this
period the remains of the degenerating sterile cells may often still be
seen (fig. 4). At the point of contact an opening is formed by solu-
tion of the cell wall and thus the protoplasts are brought into contact.
The pore is small at first, leaving the bases of the gametes quite inde- ©
pendent, and often the two tips of the gametes are also separate, as is
indicated by the notch at the apex of the conjugated cells (fig. 5).
By gradual enlargement of the pore the upper halves of the proto-
Plasts of the gametes unite to form a continuous cell mass which still
shows plainly the two distinct bases (fig. 6). The nuclei, which
before fusion occupied a central position in the fertile cells, now come
to lie in their upper portions, consequently the two nuclei are brought
side by side in the conjugated region. Simultaneous division now
takes place. . Two spindles are formed which lie side by side in about

270 BOTANICAL GAZETTE [Apri

the position in which the pore first appeared (fig. 7). The spindles
in this division are more distinct and not quite so parallel as in sub-
sequent conjugate divisions. Each shows a few short polar rays and
to each pole are drawn several (certainly more than two, though the
number was not determined) distinct chromosomes. The two
nucleoli are at this time lying in the cytoplasm outside the spindle
figures. They are less dense than at earlier stages and appear to be
disintegrating. Two of the daughter nuclei, one from each spindle,
wander back into the bases of their respective cells. The other two
remain lying side by side and move into the distal end of the fused
portion of the gametes, which now enlarges and elongates (fig. 8).
A cell wall now cuts off the distal portion of this” region, which
contains the upper pair of nuclei, and thus the first aecidiospore
mother-cell is formed (fig. g). This cell, as has been described fre-
quently, at once divides into two unequal cells, the aecidiospore and
the small intercalary cell.

After one spore has been formed, the nuclei in the bases of the
fused gametes again move upward into the conjugated portion,
which has meanwhile grown in length, and the process of division is
repeated. In this way a single row of aecidiospores is formed from
each pair of the gametes (fig. 10). During these processes the cyto-
plasm retains the same granular appearance. The nuclei, excepting
in figures showing division, always have the same general structure.

From the fact that the bases of the conjugating cells diverge
widely, it is suggested that the conditions shown in jig. 6 might be
produced by the bending of a single hypha, as is the case in the
development of an ascus. The earlier development of the gametes,
however, shows conclusively that nothing of the sort occurs here,
and that the cells which fuse may belong to distinct hyphal branches.

We find thus a typical case of the fusion of gametes at the base of ,
each row of aecidiospores, with the difference that the nuclei do not »
fuse and the cell produced by the fusion germinates at once. The
- subsequent history of Phragmidium has been well worked out and I
shall not take it up here. ~ ;

My material of Cacoma nitens did not afford so complete 4 sc
of stages, but I have been able to trace with perfect certainty the sil
tory as far as the formation of the fertile and sterile cells. As in

1905] CHRISTMAN—SEXUAL REPRODUCTION IN RUSTS 271

Phragmidium, hyphae made up of large dense cells collect in layers

, one to four cells deep just beneath the epidermis of the host (jig. 11).
From these hyphae a series of large elongated cells is formed which
raise the epidermis. A cell wall cuts each of these cells into a sterile
cell and the gamete (fig. 12).

The remainder of my material of Caeoma was so far advanced
in development that each pair of gametes had already formed a con-
siderable series of spores. Here, as in Phragmidium, a most con-
spicuous structure through all the later stages of the development of
the aecidium is the basal remnant of the walls which separate the
two gametes (see jigs. 6-10). There may be more or less of it accord-
ing to the completeness with which the adjacent walls were dissolved
away when conjugation occurred, but in all cases it and the two dis-
tinct bases remain throughout as an evidence of the double origin of
each row of aecidiospores (jig. 10).

Modifications of figs. 6-10 sometimes occur when two gametes
lie, before fusion, with the entire length of their adjacent walls in
contact. A very complete destruction of the walls in the formation
of the conjugation pore now may cause the two separate bases of the
gametes almost to disappear. A section of jig. 8, showing a lateral
or edge view, would appear as a single large cell containing two
nuclei. Either of these conditions gives us a figure which might
readily be mistaken for a binucleated basidium.

In Uromyces Caladii Pers. a further interesting modification was
observed. In this form the fused portion of the gametes elongates
Steatly and the two nuclei come to lie in the upper part of this region.
Much of the base of this region, and the two basal parts of the gametes,
are occupied by a large vacuole (fig. 13). By normal conjugate
division of the nuclei followed by cell division, an aecidiospore
mother-cell is formed. The fused portion of the cell is so long and
the base is so difficult to trace on account of the vacuole that this also
may be readily mistaken for an ordinary binucleated cell.

BLACKMAN considers the process described by him for Phrag-
midium violaceum as a ‘“vegetative fertilization,” in which an egg—
the fertile cell—is fertilized by a vegetative nucleus, and his conclu-
sions seem justified by the nature of the process in Phragmidium
violaceum. In P. speciosum and Caeoma nitens, however, the cells

272 BOTANICAL GAZETTE [APRIL

which fuse are apparently equal gametes. If we consider the matter
simply from this standpoint, the whole sorus must be considered as
a collection of such pairs of equal gametes. The fusion of each pair
should form a zygospore like that of the lower molds. There are,
however, two important differences. The product of the fusion in
the aecidium is a temporary structure which germinates at once, and
further, in germinating it produces a considerable number of spores,
the aecidiospores, while in the mold normally only one zygospore
is produced. If it is found that the same method of fusion occurs in
Micropuccinia this form might be considered an intermediate stage
between the molds and Eupuccinia.

Further, it seems plain that each pair of gametes with the spores
produced by them constitutes an individual structure. The aecidium
cup, which has so often been considered the individual fruit body on
account of its likeness to the cystocarp or pyrenocarp, is shown, by
the discoveries described above, as well as by those of BLACKMAN, to
be a complex of coordinate units rather than itself a morphological
unit. An ascocarp is the product of the fusion of a single pair of
gametes, while the aecidium arises from a collection of many such
fusing pairs. Further, the peridium of an aecidium cup is not to be
compared to the perithecial wall of the ascocarp, since it is composed
of rows of cells which are morphologically equivalent to the aecidio-
spores. The tissue protecting the asci, on the other hand, is pro-
duced by the massing together of vegetative filaments which do not
arise from the fertilized egg. :

I am convinced that the method of conjugation above described
will be found in the forms with the limited cup as well as in the more
indefinite aecidial sori of Caeoma. Considering the aecidium a
collection of individuals, it would seem that those forms having @
peridium are to be regarded as the more highly specialized, while
the caeoma is the more primitive type. ;

Against these views, which seem to me the obvious conclusions
be drawn from my discoveries, it may be urged that the spermogom
with their spermatia, showing as BLACKMAN points out very man}
resemblances to the male cells of the lichens and the red algae, 4°
still left unexplained, and it must be admitted that there is ground for
BLACKMAN’S suggestion of the origin of the rusts from the red algae.

1905] CHRISTMAN—SEXUAL REPRODUCTION IN RUSTS 273

RicHARDs’? observations of acarpogonial branch in the young aecid-
ium of U. Caladii and other forms favor this view. It is quite
possible that the gametes in U. Caladii are borne on branches arising
from such a carpogonium. The final settlement of these questions
of affinity must await the study of further forms of the rusts and
related fungi. It is especially important, as suggested above, that
the nuclear phenomena in species of rust with an abridged life
cycle be determined. There can be no doubt that the proof
given above of the existence of a sexual fusion of gametes in the
aecidium finally disposes of the conception of DANGEARD’ and
APPIN-TRourry* that the teleutospore is an egg, and the nuclear
fusion which occurs in it the equivalent of the sexual fusions else-
where among the algae and fungi. On the other hand, these discov-
eries also confirm and give‘new significance to the conceptions, so
clearly set forth by ArTHuR,> that the aecidium represents the stage
of sexual rejuvenescence. There can be no question that BLACKMAN
is in general correct in the contention that the nuclear phenomena in
the teleutospore are concerned with chromosome reduction and mark
the close of a sporophyte generation which began with the cell fusion
in the aecidium. Here again; we need further facts as to the
Tusts with reduced life cycle before the doctrine of an alternation of
generations, such as BLACKMAN maintains, can be regarded as finally
established.

Matre’s® conception that*the nuclear fusion in the teleutospore
is a mixie, was developed on the basis of the belief that no real cell
fusion occurs in the life cycle of the rusts. It is at least a fair pre-
sumption that while no nuclear fusion occurs in the aecidium, the
fusion of gamete cells described above presents all the essential
* Ricuarps, H. M., On some points in the development of the aecidia. Proc.
Amer. Acad. a descien. $2 ho 1806.

poy PA Tie siesbet sexuelle des champignons. Le Botaniste,
7:89~1 30.
7 “Sarr Taovre M., Recherches histologiques sur la famille des Urédinées.

Otaniste 5: 59-244. figs. 69. 1896-97.

a a . C., The aecidium as a device to restore vigor to the fungus.
on Agric. Sci., 23d annual meeting.

6 ‘ sei
. Marre, R., Recherches cytologiques et taxonomiques sur les Basidiomycetes.
go2.

Proc.

274 BOTANICAL GAZETTE [APRIL

features of sexual conjugations as found in other plants and animals.
Superficially considered, RAcIBoRSKI’s’ conception that the sexual
union may be regarded as consisting of two phases, cell fusion and
nuclear fusion, might seem to fit the conditions found in the rusts.
I am inclined, however, to accept BLACKMAN’s conclusion that the
fusion in the teleutospore has wholly to do with the reduction of the
number of chromosomes.

The difference between the method of conjugation in P. violaceum
as described by BLACKMAN and P. speciosum as I have described it
above indicates the necessity for a comparative study of a large
number of aecidia to determine the nature of the sexual process in
the group as a whole. There can be no question, however, that with
the discoveries already made the existence of true sexual cell fusions
in the rusts is finally established. ,

The writer feels greatly indebted to Professor ARTHUR for the
careful selection and fixation of the material used in this work, and
to Professor Harper for valuable suggestions and kindly assistance
in many ways.

THE UNIVERSITY OF WISCONSIN,

Madison, Wis.

EXPLANATION OF PLATE VIII.

The figures were drawn with the camera lucida. Figs. 1, 2, 10, 17 » and : .
were drawn using the Bausch & Lomb r-inch eyepiece and +(x oil objective; al
others were drawn with }-inch eyepiece and jy objective, the tube length in either
case being 185™™.

Phragmidium speciosum.

Fic. 1. Hyphae massing beneath the epidermis in the early stages of 1

ma. ;

Fic. 2. The series of large cells formed preparatory to division into wee
cell and gamete; a and 4, cells lying close together, which appear to be parts
different hyphal branches. oss: toe, Cee Te

Fic. 3. A cell of such a series as is shown in fig. 2 after the division into
smaller sterile cell and the large gamete.

Fic. 4. Two gametes about to fuse; sterile cells remain though
reduced in size.

Fic. 5. Fusing cells with the small conjugation pore just formed.

7 Ractsorsxr M., Uber den Einfluss dusserer Bedingungen auf die Wachstums”
weise des Basidiobolus ranarum. Flora 82:107-132. figs. II- 1896. :

very much

TE Vill

PLA

BOTANICAL GAZETTE, XXXIX

|
|
|

eo ee ee ae eee

1905] CHRISTMAN—SEXUAL REPRODUCTION IN RUSTS 275
Fic. 6. Same as jig. 5, slightly more advanced; the nuclei lie nearer the distal

end than in earlier stages.

Fic. 7. Simultaneous nuclear division following the fusion.

Fic. 8. A stage following nuclear division in which the fused mass contains

four nuclei, two side by side in the distal end and one in each of the bases.

Fic. 9..The aecidiospore mother-cell formed by a cell wall cutting off the
upper portion of the protoplast.

Fic. ro. A figure taken from a thick section showing several aecidiospores,
intercalary cells, and the two gametes at the base; the remnant of the basal part
of the dividing wall being evident between the two gametes.

Caeoma nitens.

Fic. 11. Hyphae of rust massing beneath the epidermis in very young sorus.
Fic. 12. Series of large upright cells; @, a nuclear division preceding the
throwing off of the sterile cell.

Uromyces Caladit.

Fic. 13. A pair of gametes fused, showing the nuclei in the upper portion
the protoplast, the large vacuole in the lower part, and the two less conspicuous
ases,

THE FORESTS OF THE FLATHEAD VALLEY, MONTANA.
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY.
LXVII.

; HARRY N. WHITFORD.
(WITH MAP AND TWENTY-THREE FIGURES)
[Concluded from p. 218]

THE undergrowth of the Douglas spruce-bull pine combination is
decidedly heath- or prairie-like. In the more open places grass is
predominant, and associated with the grasses are Balsamorrhiza
sagittata, Monarda scabra, Lupinus ornatus, and Clarkia pulchella.
The most common bushes are Prunus demissa, Amelanchier alni-
jolia, Opulaster pauciflorus, and Symphoricarpos sp. In places
where the rock is near the surface a heath-like appearance is given
to the undergrowth by the presence of Cladonia sp., Arctostaphylos
Uva-ursi, Campanula rotundifolia, Selaginclla densa, Lepargyraea
argentea, and Pteridium aquilinum; other forms noted were Galium
- boreale, Achillea millejolium, Holodiscus ariaefolia, and Populus
tremuloides; and along the pebbly shores of the lake Crataegus is
very characteristic.

On the southern slope of Swan Hill the vegetation conditions
found on the exposed slopes of Mission Mountains are approximated,
though this area is slightly more mesophytic; and the same may be
said for the east side of Swan Lake. Here the shores end abruptly
in a range of hills on whose west slope Douglas spruce and bull pine
are characteristic, though here and there are scattered groups of
western larch. Another meso-xerophytic area on the east side of
Big Fork River, not far from the foot of the lake (map), has already
been mentioned.

The country lying to the south and southwest of Flathead Lake
is semi-arid (fig. 5), but in favorable situations there is some woody
vegetation. A fringe of trees around the borders of the lake yond
tains principally Rocky Mountain juniper, hawthorn, and bull pine;
and the same condition is found along Pend d’Oreille River where

276 [sven

|

1905] WHITFORD—FORESTS OF FLATHEAD VALLEY 277

a low terrace has been formed. Dry ravines extending back from
this lake also have a woody vegetation. The difficulties which trees
meet in getting a start in a prairie region should be recognized. A
very large majority of the seeds never germinate, and many that do
germinate dry out before they have made a good start; and the sod
also in many instances prevents the seeds from reaching the soil.
The head of a ravine works back during freshets, undermines the
prairie sod, and thus disturbs the vegetative equilibrium that had
been established, offering a place more or less free from competition.
If seeds fall on one of these bare places and do not dry out before the
next rain which disturbs more of the soil, they may be buried by the
moving soil; the seeds thus having the advantage of being planted

- and supplied with moisture at the same time. If the conditions

remain stable long enough for the plant thus started to establish
itself, it may hold its own even though there is considerable movement
of the soil in the erosive process. Of course only a few plants out of
the number thus established can exist long in the severe conditions

_ of drouth that prevail in these regions.

The large moraine at the foot of the lake is almost destitute of
trees (jig. 5), but isolated trees of Douglas spruce and bull pine were
observed on its northern slope, and around them were a large number
of seedlings, nearly all of which, in spite of their needle-like leaves,
Were nipped off by the cattle. It is very possible that if it were not
for the cattle more of the trees would reach maturity, though it is
doubtful if the increase would be very appreciable. However, if
a slope was slightly more protected from the drying winds and the
cattle did not interfere, open stands of both Douglas spruce and bull
pine might exist. The hillside near the region shown in fig. 6 illus-
trates the point; it slopes to the north, and the part of it under dis-
cussion is too high up the cafion of the Pend d’Oreille River to be
influenced by proximity to the river, though probably trees near the
"ver are the source of supply for the seeds that stocked the hillsides
higher up. The bank is too steep for cattle to maintain a foothold
easily, and except along paths young trees are not eaten by them.

As one approaches the forest formation to the north and east
from the south end of Flathead Lake, trees become more prevalent,
& number of hilly islands (fig. 5) in the lake showing an interesting

278 BOTANICAL GAZETTE [APRIL

series of gradations. First there are hills with bull pine and a few
Douglas spruce on the north and east slopes and few or none on the
exposed slopes; then there are hills where a considerable number of
bull pines find favorable conditions for development on the exposed
sides of the hill (fig. 18), and with these there is some Douglas spruce.
On the protected slopes the Douglas spruce becomes more abundant
and may even destroy new growths of bull pine by their shade.
Another set of hills nearer the forest formations show the south slopes
fairly well covered with bull pine and Douglas spruce, while the bull
pine has become less abundant on the protected slopes where the west-
ern larch element has been introduced. Thus there are all gradations
from hills with few or no trees to those that have mesophytic elements
on the north slopes. Of course there are many variations; for
instance, hills a little to the north of west of the town of Big Fork
have their southern slopes almost destitute of trees, while the pro-
tected slopes have a stand of Douglas spruce, western larch, and
some bull pines. The series may be carried still further. It has
been shown that the west slope of Mission Mountains has a forest
of Douglas spruce and bull pine, while on the protected slope western
larch is the most prevalent, and with it trees like lowland fir, silver
pine, and giant arborvitae, which require still more moisture, are found.
If the rainfall be sufficient, a mesophytic forest may be found on the
exposed as well as on the protected slopes.

It may be well to analyze the conditions that make the so-called
protected slopes more desirable for trees. Of course the great
factor that prevents tree growth is the lack of moisture. If an annual
rainfall of 4oo™™ falls upon a hill similar to those just described,
and in gentle showers so that there will be little or no run-off, the page
that leaves the soil directly must do so by evaporation, the remainder
soaking in and becoming available for absorption. It is obvious
that the slopes receiving the strongest insolation and exposed to dry-
ing winds will lose the most water by evaporation, and these are the
south and west slopes; hence the north and east slopes will have
more moisture in the soil. Again, the plants growing on these pro-
tected slopes do not receive so much heat and are not exposed ae
drying winds, hence do not lose so much water by transpiration;
they not only get more moisture but do not need so much to supply

1905] WHITFORD—FORESTS OF FLATHEAD VALLEY 279

the transpiration streams, therefore trees that require better moisture
conditions can exist in these situations.

The sylvicultural habits of Douglas spruce and bull pine remain
to be summarized. Douglas spruce has the widest life range of any
of the species found in Flathead valley, being found at higher alti-
tudes than any other of the lowland species, with the exception of

2 pe 7% - - A f Fl: De ake; a
18.—View of a portion of an island near the south end of Flathead Lake;

Fre.
park-like growth of bull pine and Douglas spruce is present.

Engelmann spruce. It is found accompanying western larch all
through the mesophytic forests of Swan valley, and forms even a
greater percentage of the trees in the meso-xerophytic regions, and
advances into the prairie sometimes as far as bull pine. However,
this power of adaptation is not without an effect upon its form, for
in high altitudes and on the border of the prairie region it is dwarfed
and sometimes fasciated. In mesophytic conditions it reaches the
dimensions of the trees with which it is associated, and even these

230 BOTANICAL GAZETTE [APRIL

trees, though larger than most of the conifers of the eastern states,
are small compared with the vigorous trees found in the hemlock-
arborvitae-Douglas spruce forests west 6f the Cascade Mountains.
It is in this region of greater rainfall and warmer winter months that
Douglas spruce makes its best growth. With its great power of adapt-
ing itself to adverse conditions, it has spread through all the Rocky
Mountain region as far south as Mexico. While Douglas spruce
can adapt itself to varying conditions of moisture, it is very intoler-
ant of shade, in this respect being like western larch, and even requir-
ing slightly more open places in the forest to gain a foothold. Thus
the tree will be reproduced only in open places in the forest.

The distribution of bull pine in Flathead valley is more limited
than that of either Douglas spruce or western larch. It occurs in
open park-like growths on the borders of the prairie formation, with
patches of the prairie between (fig. 18). In the forest formations
in the vicinity of Nigger Prairie there is a close stand of this species
(jig. 14). Indeed so thick are the trees in certain situations that
there is not sufficient light under them for the reproduction of either
Douglas spruce or western larch, though young trees of Engelmann
spruce and lowland fir can endure the shade. Isolated groups of
bull pine are scattered through Swan valley, usually along streams
where abundant light can reach them, and in pebbly soils, where
other trees have difficulty in maintaining a stand, bull pine is found.
Wherever a single tree is found surrounded by other trees it usually
overtops them, showing that it probably started before they were
present, for it is exceedingly intolerant of shade; it must have open
places in which to pass its young stages, and this perhaps accounts
for its scarcity in the mesophytic area in deep rich soils. Where
found in Swan valley, it is a very healthy tree and very likely would
do well in pure stands there if given a chance. In other words, bull
pine does not grow in the dry soils on the border of the prairie because
it prefers the moisture conditions found there, for it does better in
the soils where there is more moisture; it is almost entirely forced
out of the latter soils in the struggle for existence with the more suc-
cessful trees. It undoubtedly demands a greater amount of heat
than the other species, with the possible exception of lowland fir,
for its altitudinal range is more limited. The highest point at which
it was observed on the surrounding mountains was 1375*-

1905] WHITFORD—FORESTS OF FLATHEAD VALLEY 281

A summary of the relation of the prairie to the forest in this region
is as follows: (1) there is less moisture in the prairie soil than in the
forest soil; (2) this is due primarily to the smaller amount of rainfall;
(3) in the prairie formation forests may exist in certain topographic
situations, as along streams and other bodies of water, and on protected
hill-sides; (4) in the forest formation prairies exist where the charac-
ter of the soil is such that it will not easily hold water; by the gradual
addition of humus such soils may be changed sufficiently in their’
water-holding capacity to permit more mesophytic conditions and
in some instances a climax western larch-Douglas spruce combination.

Objection may be made to the use of the term mesophyte for plants
with xerophytic leaves. WARMING classifies all conifers as xero-
phytes because they grow in dry soils. In the eastern United States
in contrast with the broad-leaved deciduous trees the conifers are
undoubtedly xerophytes, comparatively speaking. That is, during
the summer months the deciduous tree requires more moisture than
the conifers, therefore the deciduous tree is excluded from the dry
soils. In the northwestern United States the coniferous forests occupy
the mesophytic soils almost to the entire exclusion of the deciduous
clement. As shown in the discussion on the climatic formations,
this is due to a peculiar climate in which the summers are compara-
tively cool and dry, and the winters comparatively warm and exces-
sively wet. In such a climate the deciduous tree is lacking, except
in edaphic situations, on account of the cool dry summers, because
with its broad transpiring surface it requires more moisture than it
is able to get. The narrow-leaved sclerophyllous trees, on the other
hand, while they do not necessarily thrive during the dry summer
months, because of their reduced transpiring surface they are per-
mitted to exist, while the broad-leaved deciduous trees cannot.
During the winter the reverse is the case, for deciduous trees with
their bare twigs are better able to endure severe conditions than are
the conifers. On the other hand, if the climate is moist and warm,
the conifer is able to do a considerable amount of photosynthetic
work. Thus the deciduous tree requires more moisture during the
summer months and is consequently more mesophytic at that season;
on the other hand, the coniferous tree requires more moisture during
the winter months and consequently is more mesophytic at that

282 BOTANICAL GAZETTE [APRIL

time than the deciduous tree. Because of the equable distribution
of moisture throughout the year in sufficient quantity in the eastern
United States both can exist, although the deciduous element is able
to occupy the mesophytic areas almost to exclusion of the coniferous
element. In the northwestern United States the conifers, because
the climate in which they grow is more suited to them, are able to
occupy the mesophytic areas, and hence, so far as that. climate is
concerned, are mesophytes. Although they may have xerophytic
leaves, the structure of the tree as a whole is more mesophytic during
the non-growing season than is that of the deciduous tree. Taking
the entire year into consideration, for the reasons given above |
think that I am entirely justified in speaking of conifers as meso-
phytes.

III. THE INFLUENCE OF FIRES ON THE PRESENT COMPOSITION OF

THE FORESTS OF FLATHEAD VALLEY.

In the discussion of the forest conditions up to this point little
attention has been given to the influence of fires. There is scarcely
a section of land in the area investigated that has not been more or
less burned over. In some places mere surface fires have run through
the woods, scorching the trunks of the trees sufficiently to scar them.
In other situations the fires have burned vigorously through small
areas killing many of the trees. Still other fires have destroyed com-
pletely large areas, leaving many acres with not a single tree. Such
is the case on the west slope of the Mission Mountains (figs. 4, 16, i ).
There are many small clearings made by settlers, who after proving
up their claims have deserted the cabins erected upon them (fis ‘ 9).

By marshaling the facts collected by a study of the conditions
of reforestation in these fire clearings, nearly all stages in the ag e
lishment of new mature forests were determined. Studies in similar
regions outside of the area plotted have proved very helpful in the
interpretation of these conditions. Some important principles must
be kept in mind in explaining what plants will first get @ foothold in
the open places made by fires. These are as follows:

1. The subterranean parts of some plants that are able to sprout
from roots or underground stems may not be destroyed by ~
The sprouts of these species will give the burn a decided aspect
a short time.

1905] WHITFORD—FORESTS OF FLATHEAD VALLEY 283

2. Other things being equal, the plants whose seeds are in the burn
first will gain the earliest foothold.

3. Those plants that have seeds there early after fires will be
those that have seeds’ well adapted for distribution.

4. Of the plants that have their seeds equally well adapted for
distribution, those with seed-bearing representatives standing nearest
the burned area will have the advantage.

5. Again, other things being equal, of those species that have
their seeds equally well adapted for distribution and have seed-
bearers equally near the clearing, the species that produce seeds most
abundantly will be apt to win out in the struggle.

6. The species that can resist fires the best are likely to have left
standing in or near the area itself seed-bearing parent plants.

7. The conditions of the soil must be such that it will permit the
germination of the seeds that fall upon it. If the soil is too moist,
too dry, too poor, or too much shaded, no matter how many seeds
fall upon it, none will germinate. :

These well-known principles will aid in the determination of the
causes of the many complex conditions of forest growth after fires.
Before the actual conditions of the clearings are considered, however,
the ecological habits of another tree, the lodgepole pine, must be
known.

The lodgepole pine covers large areas in Swan valley, sometimes
forming almost pure stands. It is able to exist and thrive in those
moist areas where it has to compete only with the spruce and its
associates. It is found mixed with all the other species in the meso-

‘ phytic portion of the valley. Toward the borders of the meso-xero-

phytic areas it is not so prevalent, though signs of it were noted west
of Echo Lake and in the mesophytic portions of the west slope of the
Mission Range. It does not advance into the prairie formation,
however, so far as the bull pine and the Douglas spruce. Its alti-
tudinal range was not investigated. It is difficult to tell whether
or not this tree would maintain a stand in the Flathead valley if it
Were not for the influence of fires. It is intolerant of shade, in which
Tespect it may be ranked with western larch and Douglas spruce,
Teproducing in open places only. No young trees were noted in

€ shade, save an isolated poorly developed specimen now and then

284 BOTANICAL GAZETTE [APRIL

in the slight shade of a mature stand of the same species. Probably
it is a little less exacting in its light requirements than western larch
and Douglas spruce. ~ .

The lodgepole pine is a prolific seeder, beginning to bear fruit
early in life. Out of twenty trees varying in age from five to twenty
years, many had cones. The youngest tree noted with cones was six
years of age, one at this age having seven cones. On one tree nine
years old fifty cones were counted. It was a common thing to find
clumps of trees 3 to 4™ high fruiting abundantly. As will be shown
below, this habit of fruiting early in life is of very great advantage
to this species. Another thing of very great importance is the fact
that the cones remain closed in some instances a number of years,
thus preserving the seeds. The heat of a fire will open them and
liberate the seeds, many of which will escape injury and germinate
at once. The lodgepole: pine, during its early stages at least,
grows rapidly in height, and this gives it some advantage over its
competitors. From the measurements of twenty-five specimens each
of lodgepole pine, western larch, and Douglas spruce, the average

rapidity of growth in height per year is shown to be as follows: lodge-

pole pine 52°™, western larch 27°", Douglas spruce 20°™. Although
these averages are from rather meager data, they are sufficient to
show that the lodgepole pine has by far the most rapid growth.

In contrast with western larch, Douglas spruce, and bull pine,
lodgepole pine has poor fire-resisting qualities. Except in old trees
the bark at the base is comparatively thin; the cambium layer 1s
thus easily scorched and killed. In this way many whole forests
of trees are destroyed by fires that are not intense enough to consume
the trunks. It is not an uncommon thing to see acres of dead stand-
ing poles of this species that have thus been swept by fire. In showing
how successful lodgepole pine has been in obtaining a foothold in
the forests of Swan valley, the principles mentioned above must be
kept in mind. The réle that the plants other than conifers play '
the reforestation stages will be treated in another connection. In
order that the conditions may be understood more clearly, hypothet-
ical cases will be assumed, and when these hypothetical cases =
realized attention will be called to them. Suppose a limited area ©
burned in the midst of a forest in which western larch and Douglas

*

1905] WHITFORD—FORESTS OF FLATHEAD VALLEY 285

spruce are the dominating trees, and that these are mixed with lodge-
pole pine, silver pine, and lowland fir. Suppose that the trees surround-
ing this area all bear cones, and all have their seeds equally well
adapted for wind distribution. Since not one of the conifers found
in the region is able to sprout from the roots that would be protected
from the fires, all would have to start from seed. Let us suppose

east slopes of

1G. 19.—A clearing in a lodgepole pine forest in Swan valley;
Mission Mountains in the background; these slopes have a mesophytic forest of
Western larch and Douglas spruce.—Photograph by PRAEG

that the seeds of all fall in equally favorable places, and that the
seedlings that spring up are numerically proportional to the parent
trees in the undestroyed stand surrounding the burn. Each species
in the forest adjoining the new growth will thus be represented in
the burn in the same proportion as it is in the mature stand. If all
the young trees grow with equal rapidity, and the natural thinning
Out is proportionately distributed among the species, the new stand
will be just like the old. There are some small burns where this

286 BOTANICAL GAZETTE [APRIL

condition is approximated; of course the proportions are not exactly
the same in the old and new stands (fig. 20). Granted that any one
of the species in the young forest grows more rapidly in its youth
than the others, other things being equal it would gain an ascendancy
over its neighbors, and in the forest approaching maturity it would
have more representatives than in the old forest. As already shown,
lodgepole pine bears exactly that relation to Douglas spruce and

a clearing Sur-

Fic. 20.—Young growth of lodgepole pine and western larch in ; i
rounded by older trees of the same species, both of which have seed-bearing trees;
other plants are dwarf maple and a willow, both of which sprout from old stumps;
fireweed is also a characteristic plant.—Photograph by PRAEGER.

western larch at least, and probably to the other species, though no
measurements were taken for them. In the new forest thus estab-
lished, lodgepole pine has made a gain on the other species. How-
ever, it reaches maturity sooner than the others, and in the old a
it is the first to drop out in the struggle for existence, So that while in
the middle-aged forest it may have had some slight advantage, 1
loses this and often is entirely eliminated from the mature stand.

|

1905] WHITFORD—FORESTS OF FLATHEAD VALLEY 287

Many instances were noted, especially in the region outside the plot
mapped, in which lodgepole pine was thus being driven out of forests
where doubtless it was more prominent in the young stages of devel-
opment.

Referring again to the original case, it will be seen to be highly
improbable that the species in the stand around the hypothetical burn

Pic. 21.—Young growth of lodgepole pine and Engelmann spruce in a clearing
of the same species.—Photograph by PRAEGER.

Would produce seeds equally well. Some species might not have
any seeds at all, and they would form no part in the new forest unless
a few seeds are blown into the area from regions where there are
Seed-bearing trees. The lowland fir during the past two years has
Produced few if any seeds in Swan River valley. Any burn made
in the valley then within two years could not be stocked with seeds
of this Species. If for oe Douglas spruce or western larch
Produce seeds more abundantly than any of the other trees, they are
likely, other things being equal, to have more representatives in the

288 BOTANICAL GAZETTE [APRIL

new forest than in the old. So if any other tree, lodgepole pine for
example, has more seeds than the others, it will increase in numbers
in the new stand that is formed. So far as its relation to restocking
a burn is concerned, a tree that does not produce seeds might just
as well not exist at all.

This leads naturally to another modification of the hypothetical
case, in which the forests that surround the burn have only one or
two species instead of anumber. The result will be that these species
are likely to be the only ones found in the burn. Such a condition
is seen in fig. 20, where the forest that surrounds the clearing is com-
posed mostly of western larch and lodgepole pine, and these species
are the predominating ones in the new growth. Fig. 21 shows a
clearing surrounded principally by lodgepole pine and Engelmann
spruce, which are almost the only trees present in the new stand.
Again, fig. 22 shows that bull pine, western larch, and Douglas spruce
are present in an opening in a mature forest of these species. Another
explanation for this, however, is that the area is situated on the border
of the meso-xerophytic region where probably no other species could
exist, even if their seeds were present.

This leads to still another modification of the hypothetical case.
Assuming that the seeds of all the species are present, it is highly
improbable that they would find equally favorable places for germina-
tion. The case just cited (fig. 22) is an instance of where all except
western larch, Douglas spruce, and bull pine are ruled out. If the fire
burnt out an Engelmann spruce stand surrounded by a less swampy
region in which Engelmann spruce, lowland fir, silver pine, Douglas
spruce, western larch, and lodgepole pine were present, only Engel-
mann spruce and lodgepole pine would be able to restock it, for
these are the only species that could grow in the swampy situations.
An instance of such restocking was noted near the head of the bay-
like area of meso-hydrophytic forest southwest of Ross Lake.

Still other conditions of reforestation remain to be expla
Taking the hypothetical area that has been restocked with the seed-
lings of all the trees in the surrounding forest, and assuming that
the young growth is approximately fifteen years of age, the only
species of this young growth that would have fruit is lodgepole pine:
The cones on this pine would be more or less abundant, and on

ined.

he et i?

1905] WHITFORD—FORESTS OF FLATHEAD VALLEY 289

the others the cones would be absent. If a fire should sweep through
this young growth, destroy it, and at the same time consume some
of the original forest around it, thus extending the limits of the
original burn, what would be the result? The extent of the burned
area being greater, the seeds of the trees bordering it could not so
readily be carried to the center of the burn. But this portion of the
clearing would be restocked with the seeds of young lodgepole pines
that were destroyed by fire, but whose seeds would be more or less
protected from the fire by the cones. Some of the cones would be
cracked open by the heat of the fire and the seeds would be liberated.
The result would be that the ‘next forest would contain more repre-
sentatives of lodgepole pine than the former forest, and that they
would be more numerous in the center. Indeed this center might
contain a pure growth of lodgepole pines. Another such fire in the
course of fifteen or twenty years or less would enlarge this area at
the expense of the other species. Thus almost if not quite pure
forests of lodgepole pines of considerable extent would be established.
It is very probable that the mature lodgepole pine forest found in
Swan River valley was established in this way (fig. 23). The evidence
for this is as follows:

I. In no case was astand of this species found in which there were
not noted dead and charred trunks of western larch and Douglas
spruce, mostly the former; these because of the thick bark would be
the last to yield to the fire. In some instances mature live trees of
Western larch were observed towering above the younger lodgepole
Pine forest. In these cases there were isolated specimens of young
"estern larch of about equal age growing with lodgepole pines (fig. 12).

2. In nearly all cases these forests grade imperceptibly into more
Mature forests in which the lodgepole pine element is entirely or nearly
Wanting. The mature forests of western larch, Douglas spruce,
silver pine, lowland fir, and lodgepole pine are in many instances
Stowing in soil that is similar in moisture content. Thus it cannot
be said that the difference in the two stands is due to the character of
Soil conditions.

3- In the meso-hydrophytic situations there is often a gradation
from a spruce forest to a mixture of Engelmann spruce and lodgepole
Pine, the latter stand being much younger than the former.

ES ee tO ee ee a ae

ee

290 BOTANICAL GAZETTE [APRIL

4. In stands that contain little or no signs of fires, the lodgepole
pine element is absent or nearly so.

5. Nearly all stages in the development toward this condition were
noted. Thus, fig. 20 shows a comparatively recent clearing in a young
lodgepole pine forest, which is in turn surrounded by a mature forest of
western larch and Douglas spruce in which the lodgepole pine element
is inconspicuous. Fig. 12 shows an almost pure growth of this young
forest with mature trees of western larch in it. In another place there
is a more mature growth in which only charred trunks of larch tell
the tale.

For the reasons given above, it is believed that the explanation for
the lodgepole pine in the area plotted is the correct one. As before
stated, observations were not limited to this immediate region, and
many instances might be cited where burned areas similar to those
described have become reforested with lodgepole pine. It will not
be out of place to repeat that the advantage which it has over the
other species in the region is due to its capacity to produce seeds
early in life, and its habit of retaining the seeds in its cones for a
number of years, thus preserving them for a greater or less length of
time. In forests partially destroyed by fire the trees of western larch
and Douglas spruce, because of the capacity of their trunks to resist
rather severe burnings, will restock the burned areas. Fires of this
nature, repeated sufficiently often to prevent any young lodgepoles
from becoming old enough to produce cones, will militate against
the latter, while Douglas spruce and western larch will have cone-
bearing trees on the ground so long as the fires are not too intense.
As soon as these are destroyed, then the seed supply is cut off and
restocking from that source at least will discontinue.

_ It will be seen readily that if the fires that have made the lodgepole
pine condition possible are repeated every five years, for instance, the
young growth forests of that tree will not be permitted to follow each
other in procession, for the five-year interval between fires will be too
short a time to permit lodgepole pine to produce seeds. Then 8
course all forest growth will be completely destroyed and the area wil
not become clothed with trees until restocked with seeds from the
neighboring undestroyed forests. The further these are away, the
longer it will take for seeds to reach the devastated area. Howevel;

1905] WHITFORD—FORESTS OF FLATHEAD VALLEY 291

if the interval between the fires is sufficiently long to give lodgepole
pine time to produce seeds, after each fire a forest of lodgepole pines
will spring up to replace the old. Such fires enable lodgepole pine

to perpetuate itself- so
long as the soil is able
to furnish the requisite
amount of nourish-
ment, but a checking of
fires will in time bring
about the normal con-
ditions. This has been
done about as follows:
The fires being
absent, the lodgepole
pine stand is permitted
to reach maturity. As
it does so, seeds of forest
trees in the neighbor-
ing undestroyed or par-
tially destroyed stands
have had time to spread
into the lodgepole pine
forests. Seedlings of
those trees that can
tolerate the shade will
et a start at once, if
the other conditions are
favorable. Thus in
Many instances silver
Pine, lowland fir, and
Engelmann spruce were
hoted growing under
lodgepole pine stands
in Swan valley. Also in
ae Shae Lake r region

: ea
10f western larch, Douglas

FIG 22.—Young growt : h :
S “ol i -learing I the same
= a clea nf Oo
spruce, and bull pine in ~

i : , PRAEGER.
species.—Photograph by PRA!

Ss, alth ug € hey | S c re a to W hich

292 BOTANICAL GAZETTE [APRIL

these two species are confined, their occurrence under lodgepole pine
stands is not common. As soon as openings are made in the forest of
mature lodgepole pines, due to causes other than fires, this undergrowth
may spring at once into prominence and may come to occupy a place
in the mature forests. At the same time the openings have made it
possible for the existence of seedlings of such trees as western larch
and Douglas spruce which demand light for germination. Of course
lodgepole pine seedlings can germinate here also, and in the first
generation or two they will still predominate; but each new generation
will have fewer specimens of the latter species, for since it is not a
long-lived tree, a canopy of the mature forest of the other species will
in the long run crowd it out,by density of shade, or reduce its occur-
rence to isolated trees here and there in the forests. In this way the
forests will revert to the normal conditions. The time it will take to
do this depends upon the nature of the conditions that the last fire
left. If the lodgepole pine conditions had been established for a wide
area, the time would be long, perhaps covering many generations of
growth. If on the other hand the fire only partially destroyed the
original forest, one or a few generations would suffice to permit the
re-establishment of a forest similar to the original. Between these
two extremes are all stages, some of which have been described.
Indeed the present forest formations are a complex expression of the
influence of fires upon them. | ;
The general aspect of a lodgepole pine forest approaching matunty
is somewhat different from that of other stands. It has already been
shown that the growth in height is rapid. In dense stands of young
growth the poles are long and spindling, hence the name lodgepole
pine. The small diameter of the bole of the tree is very characteristic.
A forest in which the average age of the trees is about one hundre
years is seen in fig. 23. In this forest, where the trees are over @
hundred years old, the average diameter is probably 20°™, many —
being only 15°™ in diameter.
Compared with the other forests growing in like situations, the
canopy that a lodgepole forest forms is not very heavy. This per-
mits more light to reach the forest floor, hence there is 4 greater
development of undergrowth. It has already been shown that seed-
lings of silver pine, Engelmann spruce, and lowland fir can endure

1905] WHITFORD—FORESTS OF FLATHEAD VALLEY 203

the shade of these forests, and that except in open places Douglas
spruce, western larch, bull pine, and lodgepole pine are ruled out
altogether. The birch is found scattered throughout the limits of
the forest. Indeed, as will be shown, it is often a conspicuous tree

5 bet ao ae, f RD. ase 7 <j
PRR aa eG a eA eA
rc ‘ : “~—F.

5
:

; Fic. 23.—Mature lodgepole pine forest in Swan valley, showing a dense stand in
ie the trunks are 15 to 20cm jn diameter; the undergrowth consists of Acer glabrum,
Vv m Pose 3 - is
yrola, Pteridium, Linnaea, etc.—Photograph by McCaLium.

in the burns, and grows up with lodgepole pines, though as the forest
Matures it drops out early. The undergrowth consists of Acer
slabrum, Lepargyraea canadensis, Symphoricarpos sp., Rosa sp.,
Lycopodium sp., Pyrola secunda, Pteridium aquilinum, Linnaea
borealis, Chimaphila umbellata, Clintonia borealis, and Aralia nudi-

7

294 BOTANICAL GAZETTE [APRIL

caulis. Scattered specimens of juniper are found in places.. In
other respects the forest is like the other stands in the mesophytic
regions.

In what has been said concerning fires, no mention has been made
of species other than conifers. They may often play an important
role in the first stages in the natural process of reforestation, but their
importance diminishes as the life history becomes complete. One of
the first plants to give a decided aspect to the forest after fires is the
fireweed (Chamaenerion angustijolium). The birch and aspen, by
virtue of their light seeds, gain an early place in the burns. Indeed
sometimes their stands become quite dense and they check the advent
of coniferous species. Many species that were on the forest floor of
the mature stand may have survived the fires and spring up even
more vigorously than before, because the shade has been removed.
Especially is this true of those forms that can send shoots from their
roots or from underground stems. In the burn on the east slope of
the Mission Range south of Big Fork, there are shrubby growths of
Salix spp., Ceanothus sanguineus, Opulaster pauciflorus, Acer
glabrum, and Holodiscus ariaejolia, many of which probably were
present as underbrush in the stand that existed previous to the burn.
These shrubs will retard the reforestation of the area in some instances
to a marked degree. If the forest that is destroyed be on the border
of the prairie, plants from that association will form an important
element in the growth that follows. This is of course detrimental to
forest growth. Indeed if the fires be repeated often enough, the
forest plants will gradually diminish in quantity, and the prairie
element will become more and more prominent, until finally a prairie
will come to replace a forest. If the fires be checked, however, for
any length of time, the forest will gradually reconquer the territory
thus lost. It is probable that some of the prairie region in Flathead
valley has been won from the forest in this manner. It must be
remembered that in the area bordering on the prairie, in Flathead
valley at least, lodgepole pine is not successful and is thus ruled out
from playing any important réle in these places after fires.

In the discussion of the causes for the Nigger Prairi
mention was made of the importance of the accumulation of hum
in rendering the sandy soil more capable of holding moisture.

e region,
us
It

1905] WHITFORD—FORESTS OF FLATHEAD VALLEY 295

can be seen that fires will tend to reduce the humus content of the
soil. This is of extreme importance, for that which would otherwise
add to the richness of the soil and increase its water-holding capacity
is destroyed. Especially is it of importance in those regions that
border on the prairie. There is evidence of many such surface fires
in the open woods around Nigger Prairie, and it is very probable
that these fires have played an important réle in keeping the prairie
vegetation from being encroached upon by the forests that surround
it. Even in the mesophytic conditions, fires influence the capacity
of the soil to reforest itself quickly, by partially or totally destroying
the humus. However, with the reclothing of the burned area, the
floor of the new forest will gradually resume its normal condition.
From the above it will be seen that forest fires play an important
part in determining the composition of the forest. That forest fires
prevailed in this region before the advent of civilized man is a logical
inference. In the lodgepole pine forest in the Swan River there are
unmistakable signs of fires before the present forest, which is now
about one hundred years old, was started. How these fires started
can only be conjectured, and it is not within the province of this paper
to discuss their origin. It is also very evident that the fires are more
numerous since the settling of the country by civilized man than
ore.

- SUMMARY.
_1. Fires play an important part in determining the present compo-
sition of the forest.

2. The lodgepole pine is the “fire tree” of the region.
~ 3- It is favored after fires principally because it has the capacity
to produce seeds early in its life.

4. Many complex conditions of growth are introduced after fire.
The species that have seed-bearing trees near the burn will generally
be represented in the new forest. e

5- Repeated burnings at intervals of ten to thirty years will
establish a lodgepole pine forest where formerly there existed a normal
mesophytic forest.

6. Repeated burnings at intervals of five years or less will destroy
all forest growth.

296 BOTANICAL GAZETTE [APRIL

7. After the lodgepole forest is once established and the fires are
checked, it will slowly be replaced by the species that exist in the
normal forests.

8. The lodgepole pine is not successful in the bull pine belt.

I wish to thank Dr. H. C. Cowrrs for many valuable suggestions
made in the preparation of this paper; also Professor M. T. ELRop,
through whose kindness I was permitted to make the Montana
Biological Station headquarters while collecting data; also Professor
Exrop, W. E. Prarcer, and W. B. McCativm for the photographs
from which the illustrations were made. The taxonomic nomen-
clature used in this paper is that found in RypBERG’s Catalogue oj
the flora of Montana and the Yellowstone National Park, and Svv-
WORTH’S Check list oj the jorest trees of the United States.

LABORATORIES,
Manila, i.

BRIEPER ARTICLES,

NOTES ON THE PHYSIOLOGY OF STIGEOCLONIUM.
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY. LxXX.
(WITH THREE FIGURES)

THE experiments with which this paper has to deal fall into two groups,
those with low temperatures and those with sea water. The subject for
experimentation was the polymorphic form of Stigeoclonium with which
I have been concerned in several preceding papers.*. As has been already
shown, this alga takes either of two very distinct forms according to the
nature of the medium in which it is grown. In solutions of relatively high
Osmotic pressure and in those of low pressure to which stimulating metallic
salts have been added, the palmella form is assumed. This consists of
nearly spherical cells lying singly or in irregular groups. In unpoisoned
solutions of low osmotic pressure the alga grows as branching filaments
composed of cylindrical cells. When such filaments are placed in a poisoned
solution or in one of high pressure, they become transformed to the other
form by the simple rounding off of their individual cells. In general the
Production of zoospores is checked where the palmella form is produced,

ut there are a number of exceptions to this among the metallic poisons.?
In such cases this process may be accelerated even where the germination
and growth of the zoospores are inhibited.

I. Low tem peratures.—Since high pressure of the medium acts to pre-
vent water absorption, and since low temperatures are known to cause the
extrusion of water by both plant and animal cells,3 it occurred to me that

__? Livinesron, B. E., (1) On the nature of the stimulus which causes the change
n form of polymorphic green algae. Bor. GAZETTE 30:289-317- 1900.

~_—+ (2) Further notes on the physiology of polymorphism in green algae.
Bor. Gazerre 32:292—302. 1901.
~~» (3) The réle of diffusion and osmotic pressure in plants. Chicago. 1903
Part II, Chapter III. This chapter was reprinted as “The effect of the osmotic pres
“ure of the medium upon the growth and reproduction of organisms. Chicago. 1993.
>» (4) Chemical stimulation of a green alga, Bull. Torr. Bot. Club 32:1-34.
figs. bi 1905.

? Loc. cit. (4).
5 Loe. cit. (3), pp: 75 and 141, and the references there given.

298 BOTANICAL GAZETTE [APRIL

possibly low temperatures might also cause filaments of this alga to take
the palmella form. Experiments were devised to test this point, and their
results are here given.

Cultures of the filamentous form were made in small glass dishes with
loosely fitting covers. The medium employed was the modification of
Knop’s solution previously described, and had an osmotic pressure of
60™™ of mercury. The culture dishes were placed in weighted beakers
which floated about three-fourths immersed in ice-water contained in a
galvanized iron tank. Ice was added from time to time as melting took
place, and the superfluous water was drawn off. The tank was covered
with glass and stood in the conservatory, so that the plants were supplied
with the necessary light for growth. The cultures were shaded from direct
sunlight.

In the medium employed the alga grows rapidly as filaments at labora-
tory and conservatory temperatures. Zoospores are produced and germi-
nate to form new filaments. Such normal filaments are shown in jig. 2-
The figures are all from camera drawings and are magnified about 300
diameters. In the cold cultures, whose temperature rarely rose above
6° C.,5 the growth of the original filaments was checked, but the production
of zoospores continued at about the normal rate. At the end of fifteen
days the old filaments had completely changed to the palmella form, in the
manner already described for solutions of high osmotic pressure. Z0o-
spores fail to germinate normally in the cold; most of them simply lie quies-
cent on the bottom of the dish, having assumed the spherical form, while
a few enlarge slowly and divide into new palmella cells. Growth of the
palmella form is comparatively very slow at ordinary temperatures. It Ls
still more so in the cold, and this retardation is here even more marked in
resting zoospores than in cells produced by the breaking up of the original
filaments. Palmella cells from one of the cold cultures are shown in 7g: 7-
They are seen to be exactly similar to those produced by high pressure =
toxic cations. Several resting zoospores and empty sporangia are also
figured. That the plant was not permanently injured by the low tempera-
ture was shown conclusively by continuing the cultures in the conservatory
after they had been taken from the cold bath. They all responded to te
return to normal temperature, by producing typical filaments in from win
to fourteen days. A portion of one of these cultures after the filamentov
form had been assumed again is shown in jig. 2. Germinating ee
and empty sporangia are also shown.

4 Loc. cit. (4), p. 4. ;
During the period of the experiment, 20 days, the temperature was u

nwittingly
allowed to approach ro° C. several times, for periods of a few hours only.

1905] BRIEFER ARTICLES 299

Five cultures made from different stock material, and always compared
with controls maintained at the temperature of the conservatory, all agreed
perfectly in the results. Thus it seems safe to conclude that low tem pera-
tures act upon the vegetative growth of this alga with the same result as do
high osmotic pressure and poison cations. No acceleration of zoospore
formation, a phenomenon often exhibited in poisoned solutions, has been
observed here. It appears that in low temperature we have another
method of withholding water from the plant, and that the uniform
response to such withholding is the production of the palmella form.®

Fic. 1.—Palmella from filaments like fig. 2 in culture at low temperature.

Fic. 2.—Normal filaments and germinating zoospores from cold palmella culture
returned to normal temperature.

Fic. 3.—Palmella in sea water from normal filaments like fig. 2.

Il. Sea water.—During my residence at the New York Botanical
Garden I was able to determine the behavior of this Stigeoclonium in sea
water. Natural water was collected from the surf at Far Rockaway, L. L,
and was brought in bottles to the laboratory, where it was used in making
the cultures. Filaments of this plant placed in undiluted sea water take
the typical palmella form, as in other solutions of high pressure. Zoospores
are not produced, nor do those produced previously germinate. Such a
culture is shown in fig. 3. The water used had a pressure of about 25,000"™
of mercury.7_ In sea water diluted to one-tenth and one-hundredth of its
natural concentration respectively—water redistilled in glass being used
In the dilution—the response is the same, although the change is not so
Tapid. In the latter dilution it seems impossible that osmotic pressure
Is the main stimulating factor, for here is a pressure of only about 250"™

° See the remarks on this subject in the paper on chemical stimulation, loc. cit-
(4), p. 21 et seq.

7 The calculation was made by the method of the depression of the freezing-point,
loc. cit. (3), Pp. 37.

300 BOTANICAL GAZETTE [APRIL

of mercury, while a pressure of over 15,000™™ is necessary to bring about a
marked response in the alga.2 Perhaps there is a stimulating chemical
in sea water which aids the somewhat high osmotic pressure to bring about
the result. At a dilution of one-thousandth the filaments grow normally
as in weak nutrient solutions.

The result of this test suggests how a fresh-water form coming into sea
water may be influenced to change its character and still live and thrive.
This may have been a factor in the evolution of certain algal forms.—
Burton Epwarp Livincston, The University of Chicago.

FERTILIZATION IN THE SAPROLEGNIALES.

Proressor B. M. Davis, in his criticism of my views ‘‘On fertiliza-
tion in the Saprolegnieae,” makes use of the term ‘“‘ovocentrum” in a
sense very different from that which I gave to it. His use of the term,
moreover, is obviously based on a misconception of my meaning. He
says ‘‘Trow calls the egg-asters ovocentra.” I neither do this, nor do
I approve of it being done. Such an innovation would be worse than
useless, as it would increase the confusion which already exists. At present,
unfortunately, structures of more than one kind have apparently been
grouped together by giving them a common name—‘“coenocentrum.” I
suggested the term “ovocentrum” as suitable for use in describing the
dense mass of protoplasm found at the center of the eggs of the Sapro-
legnieae. Imbedded in the ovocentrum of an egg I find a single nucleus
accompanied by a single centrosome and its astrosphere. It would have
been correct, I think, for Davis to have said that the “coenocentra” dis-
covered by him in Saprolegnia were interpreted by me—rightly or wrongly
—as consisting of centrosomes with their astrospheres. The “ovocentra
of the Saprolegnieae may be the equivalents of the coenocentra of
Peronosporeae. They are altogether different from centrosomes and as
spheres. I cannot, therefore, accept DAvIs’s statement that I call “the ese-
asters ovocentra.’’ I do not propose to discuss the many other points of
interest raised by Professor Davis, for new facts are required now, and
these can only be obtained by patient and prolonged investigation 19 a
laboratory.—A. H. Trow, Cardiff, Wales.

8 Loc. cit. (1) and (2). 9 Bor. GAZETTE 39:61. 1995.

tro-

CURRENT LITERATURE.
BOOK REVIEWS.
The trees of North America

PROFESSOR SARGENT’ has brought into a convenient volume the information
concerning the trees of North America that is much more elaborated in his Silva.
The sequence is that of ENGLER and Pranri’s Die natiirlichen Pilanzenfamilien.
Especial attention has been given to the construction of simple analytical keys,
so that a species may be determined with the minimum of trouble. For example,
the key to the families is based on the arrangement and character of the leaves;
and in the same way genera and species are reached by the important and easily
discovered contrasting characters. Each of the 630 or more species is further
made clear by the admirable illustrations of Mr. C. E. Faxon, showing the leaves,
flowers, and fruits. There is no reason why this manual should not become at
once extensively used by all those interested in trees, a constituency that extends

far beyond the boundary of professional botanists.—J. M. C.

Organic evolution

A RECENT BOOK by Professor METCALF? presents in a clear and simple style
the fundamental principles of organic evolution in a form very well adapted to
the needs of the general reader and to those who wish an outline of the theory of
arwinism, The standpoint is clearly that of the Neo-Darwinist, and the main
topics the familiar ones of adherents of this school. Perhaps the most striking
feature of the book is the wealth of clear and very well selected illustrations, a
large proportion of which are presented for the first time in a general text of this
character. These give to the work a character quite its own, and in themselves
Justify a book which in its general outlines follows very closely the subject-matter
and method of presentation in several popular accounts of evolution, including
those of WaLLace and RomANEs. The subject-matter is chiefly zoological, and
the book would hardly claim an extensive treatment of the principles of plant
evolution.—B. M. Davis

MINOR NOTICES.

THE REPORT for 1904 of the Chief of the Bureau of Plant Industry, Dr.
GaLLoway, is commended to the perusal of all botanists that they keep in touch
With the botanical work in progress under this Bureau of the Department of

S
of Mexico). Imp. 8vo. pp. xxiii +826. figs. 640. Boston: Houghton, Mifflin and
Company, 1905. $6.00,

MErcacr, M. M., Anoutline of the theory of organic evolution. Imp. 8vo. pp.
204. pls. rer, figs. 46. New York: The Macmillan Co. 1904.

1905] 301

302 BOTANICAL GAZETTE [APRIL

Agriculture. It is impossible here to summarize the report, itself a summary.
Certainly every American who understands the past success and future plans of
this bureau must feel proud that our government is thus leading the world in
turning applied botany, based upon researches by a capable staff, to the produc-
tion of wealth and comfort for the people. Asa mere matter of dollars and cents,
any one of a dozen or more discoveries made or practically applied by this bureau
recently will add to the annual income from our fields and gardens more than
the whole bureau has cost from its beginning. The report is a concise and
clear showing that will interest all and will surprise those who have not followed
closely the recent developments.—C. R. B.

Tue Butrerin fromthe laboratories of natural history of the State Univer-
sity of Iowa contains in its last issue (Vol. V, no. 4) two botanical papers on the
local flora, viz., The flora of the St. Peter sandstone in Winnesheik Co., Iowa, by
Professor B. SHimEK: and The Discomycetes of eastern Iowa, by FRED J. SEAVER.
In the latter, out of nearly one hundred species collected in the state, about fifty
are described and illustrated in twenty-five plates, the remainder being reserved
for further study. No new species are described.—C. R

THE SECOND ANNUAL ISSUE of the volume on Botany of the International
Catalogue of Scientific Literature3 was published in December last, the manuscript
having been completed in March 1904. ‘There is no occasion to explain the
scope and quality of this publication, for it has already been extensively rev iewed,*
and the opinions of botanists concerning it have become well settle d.—J. M.

J. Perkrns’ has issued the third fascicle of contributions to the flora of the
Philippines. The collaborators are as follows: C. DE CANDOLLE, Piperaceae,
J. Perkins, Rutaceae; O. WARBURG, Ulmaceae, Moraceae, Urticaceae, Balano-
phoraceae, Artciodochintess: Magnoliaceae, Thymelaeaceae, Ericaceae, and Ficus;
E. B. Copetanp, Ferns (38 n. spp. and Christopteris, n. gen.).—J. M- ©-

NOTES FOR STUDENTS.
RECENT STUDIES IN HYBRIDIZATION.—The literature of Mendelian inherit-
ance has been enriched by the appearance of several important recent papers:
SCHERMAK © gives an account of further studies in the hybridization of peas,
stocks (Matthiola), beans, etc., which have given him so many noteworthy results
reported in earlier contributions. The great amount of interesting detail makes
= review impossible in a short note. ‘Kreuzungsnova,” which have

3 International Catalogue of Scientific Literature. M. Botany. 2d annual issve
8vo. pp. 1111. London: Harrison & Sons, 45 St. Martin’s Lane. 1904- sad

4 Bot. GAZETTE 34:455.

5 S Je b saices ae ae Philippinae. Fasc. Til. pp. 155-7 -*
Leipzig: poe Borntraeger. 1905. M5.

© TscHERMAK, E., Weitere Kicusanennedien an Erbsen, Leukojen und Bohnen
Zeits. Landw. Versuchsw. 7: 533-638. 1904.

1905] CURRENT LITERATURE 303

usually been considered uncommon, he has found to result regularly in seventeen
of the hybrids that he reports. Races which show the possession of such latent
characters that are externalized as a result of crossing he has designated in another
recent paper? as “‘cryptomeric” races, and the process of externalization of
latent characters as ‘‘cryptohybridism.” In many cases the new characters are
recognizable as atavistic, in other cases they appear to be retrogressive or degres-
sive modifications. The processes by which these arise he calls ‘“‘hybrid atavism”
and “hybrid mutation,” and he looks upon the latter as an important source of
new forms, especially of ‘‘defect-races.”’

In nearly all cases the new characters as well as the parental characters
behaved in a Mendelian way. Thus in a cross between-two races of peas, white-
flowered X rose-colored gave complete dominance of red (atavistic) in the first gen-
eration (F;) splitting in F, to red: rose: white =9: 3: 4, the red dominating the rose,
and red+rose the white, in the simple ratio 3:1. In a case of this kind the red
is called ““dominant” and rose “co-dominant.” In other cases “co-recessives”
occurred, giving four forms in the second generation, the ratio being 9:3:3:1
A still further complication was found in a cross between white-flowered Matthiola
glabra and red-flowered M. incana, in which pure violet dominated in F, and five
forms appeared in F,, giving approximately the proportions 27:9:9:3:16.
This last result was reported by TSCHERMAK two years ago, and he explained it
by assuming that each of the four colors (the fifth class being the white reces-
Sives) is a compound, a+b, a+d, b+c, and c+d, and on this assumption he
predicted then what would be the product of each of these groups in F; when self-
fertilized. The third generation is here presented, showing that in every instance
his prediction was confirmed.

Lock" obtained very similar results in crosses of various cultivated peas with
Species native at Peradeniya, Ceylon. Thus with respect to seed-color, self-
colored x white gave in F, self-colored : purple spotted : white=9: 3:4, and
another case precisely resembling that of TSCHERMAK’s Matthiolas gave
Categories of seed-coloration in the second generation nearly in the ratios 27:9:
9:3:16. The explanation offered by Locx is essentially the same as TSCHER-
MAK’S, though differently expressed. He assumes that the allelomorphs ABC
in the colored pea are matched by corresponding recessives abc in the white, but
that neither B nor C can reach external expression except in the presence of A.
he chance combinations of gametes containing these allelomorphs then result
In the observed ratios

; Lock’s paper gives an admirable general treatment of Mendelism, illustrated
With new examples taken from his own studies, and presents briefly but fairly
the results of other workers. It can be recommended to any one who wishes to
at =,

| PSCHERMAK, E., Die Theorie der Kryptomerie und des Kryptohybridismus.
Beihefte Bot. Centralbl. 16: 11-35. 1904.
P tennan R. H., Studies in plant breeding in the tropics. Ann. Roy. Bot. Gard.
erade

niya 2: 209-356. 1904.

304 BOTANICAL GAZETTE [APRIL

orient himself with the least possible expenditure of time and effort in the impor-
tant fields which have been opened up by MENDEL and his belated successors.

EMERSON® gives a continuation of his studies in bean hybrids, in which he
now treats his results statistically, instead of qualitatively as in his preliminary
report two years ago. e numbers are mostly not large, but considering the
smallness their agreement with the theoretical ratios is fairly close. The chief
interest in EMERSON’s results is the class of characters investigated, often making
classification difficult, and considerable error probably being introduced in this
way. A few of these characters with the dominant member of the pair given first
are as follows: running habit (flowers axillary), bush habit (flowers terminal);
pods tender, pods tough; pods green, pods yellow. In the case of stringless vs.
stringy pods, about half the crosses showed the former dominant, the other half
being intermediate; though all the progeny from any given cross behaved con-
sistently. EMERSON shows that no prediction can be made regarding the hered-
itary behavior of seed-color from knowing the relations obtaining in other hybrids
having the same seed-color, a fact also noted by TsCHERMAK in the first paper
mentioned above.

Correns’? has reported the results of further observations on hybrids of
Hyoscyamus niger, H. pallidus, and H. major. The former crossed with its var.
annuus shows complete dominance of the biennial habit with typical splitting in
F, and later generations. H. pallidus x H. niger, which was reported in a pre-
vious paper as giving intermediate flower color in F,, isnow shown to split typically
in later generations, regardless of the annual or biennial habit of the offspring
and independently of evironmental conditions. :

ORRENS" has also investigated the hereditary relations of gynodioecism,
dealing chiefly with Satureja hortensis and Silene inflata. In the former species
bisporangiate flowers crossed together showed a predominance of bisporangiate
plants in F,, but when stigmas of the ovulate plants were pollinated with pollen
from bisporangiate plants the ofispring were almost without exception ovulate.
In Silene the results were very similar, ¢X¥% giving almost entirely °, and #X%
resulting in a predominance of ¢. This is quite contrary to Mendelian —
tion, and the author does not believe that sexuality can be aligned under ordinary
laws governing characters of hybrids, which it will be recalled has been attempted
by CasTLe*? with reference to animals.—G. H. SHULL.

9 EMERson, R. A., Heredity in bean hybrids. Ann. Rept. Agr: Exp. Sta.
Nebraska 17: 33-68. 1904.

t° CorRENS, C., Ein typisch spaltender Bastard zwischen einer einjahrigen SIppe
des Hyoscyamus niger. Ber. Deutsch. Bot. Gesells. 22:517-524- 1994:

: LG f.
™ CoRRENS, C., Experimentelle Untersuchungen iiber die Gynodioecie. sa

Deutsch. Bot. Gesells. 22: 506-517. 1904.
7-218.

12 CASTLE, W. E., The heredity of sex. Bull. Mus. Comp. Zool. 40:18
1903.

1905] CURRENT LITERATURE 305

PENHALLOW’S has recently published the first and general part of an extensive
treatise on the anatomical determination of the North American Coniferales
as well as certain species from Japan and Australia. The memoir represents
the results of a quarter of a century’s work in this field andis of great importance
from the paleobotanical and phylogenetic standpoints. Although the author
expresses his appreciation of the value of a general anatomical study of the group,
he limits himself to the structural features of the ligneous cylinder, for the reason
that coniferous remains are ordinarily best preserved as fragments of wood.
Wood structure is discussed under the following headings: spiral tracheids;
bordered pits, distribution and structure; medullary rays; resinous tracheids
and resin cells; and resin Passages.

Under the caption of bordered pits, the author points out that multiseriate
and crowded pits deforméd by mutual pressure are characteristic of the older
8ymnosperms, the Cycadofilices, Cordaitales, etc. Bordered pits are in general
confined to the radial walls of the tracheids, exceptions to this mode of distri-
bution occurring only in the primary wood and in the autumnal tracheids of the
annual rings. Under the heading of medullary rays, the various types of struct-
ure which are of diagnostic importance are clearly and definitely described, and
in this respect the present work marks a long advance on its predecessors. Two
types of ray are distinguished, namely the linear ray made up of a single series
of cells, and the more complex fusiform ray, which is so broad as to contain a
horizontal resin canal. Resinous tracheids are described as diagnostic for the
Araucarineae, but also occur sporadically in certain species of Abies. Resin
cells, according to the author, appear in the more primitive conifers (exclusive
of the yew-like Taxineae) and are characterized by their scattering, zoned, or
Segregated distribution, the latter condition being considered the most specialized.
Where the resin-cells are highly segregated they may give rise schizogenously
to resin cysts of limited extent, with walls constricted at intervals. Resinous
cysts are found characteristically in the abietineous genera Tsuga and Abies,
but also occur in living and extinct Sequoias. In the abietineous genera Larix,
Pseudotsuga, and Picea, the resin spaces form a continuous system of vertical
and horizontal passages, which according to the author betray their derivation
from resin cysts by the constricted character of their walls. In Pinus the resin
Passages no longer show signs of constriction and are moreover lined entirely
with thin-walled epithelium.

The author, as the result of plotting frequency curves based on the occurrence
of the various characters of the wood, reaches certain conclusions as to phylogeny
which may be briefly stated as follows: The Taxaceae are the oldest of the Con-
iferales, and from their general plexus have branched off on the one hand the
Podocarpeae, and on the other the common trunk, which gave rise to the Taxo-
ee

"3 PENHALLow, D. P., The anatomy of the North American Coniferales together
With certain exotic species from Japan and Australia. Part I. American Nat. 38:
743-273, 331-359, 523-554, 691-723. 1904.

306 BOTANICAL GAZETTE [APRIL

dineae, Cupressineae, and Abietineae in approximately ascending order. The
Araucarineae, contrary to the usual opinion, have a separate and direct origin
from the Cordaitales. The conclusions reached by the author from the study of
wood harmonize on the whole very well with those adopted in general taxonomic
works such as Engler and Prantl’s Die natiirliche Pflanzenfamilien, and are based
on the assumption that greater complexity of structure is necessarily charac-
teristic of more modern forms. There is of course room for difference of opinion
on this subject, in view of the evidence of general paleobotany, comparative
anatomy, and development, which seems to show that the Coniferales form a
‘series of recession rather than of progression. There can be no doubt, however,
as to the very important character of the facts brought out by the investigations
of Professor PENHALLOw, and the second part of the memoir, which is to contain
the specific determinations of the conifers from the structure of their wood, will
be awaited with interest —E. C. JEFFREY.

JEFFREY" has published the second of his contributions to our knowledge
of the anatomy of the Coniferales, in so far as it bears upon relationships and phy-
logeny. The immense service that anatomy of this kind is rendering to morphol-
ogy that concerns itself with phylogeny cannot be overestimated. The student
of gymnosperms is particularly indebted to JEFFREY for the studies he is just
now prosecuting, for this new method of attack could not have been directed
more usefully than upon Coniferales. In the preceding number of the series
the author has reached the conclusion that Sequoia shows an abietineous origin,
and it was natural that a presentation of the Abietineae should follow. No
better summary of his results could be given than that prepared by the author
himself; and since the subject is one of much general interest and importance
no apology is needed for reproducing it here.

1. The Abietineae are divisible, on the evidence supplied by a study of their
vegetative and reproductive organs, into two distinct subfamilies, viz., the Pineae
and the Abieteae.

Resin canals are present in the outer margin of the primary wood of the root.
scales of the female cone are not deciduous. Pinus, Picea, Larix, Pseudotsuga- :
3- The Abieteae ordinarily do not possess resin canals in the secondary W
root and shoot. Resin canals, however, are sometimes found in the wood of the
female reproductive axis, and in the first annual ring of vigorous sh
trees. Resin canals occur in the secondary wood in tangential rows,
of injury. Resin canals are invariably found in the center of the pri rimary wood ane
root. The scales of the female cone are generally deciduous. Abies, " Pseudolarix,
Cedrus, Tsuga.
The evidence derived from anatomy and experimental morphol
that the presence of resin ducts in the woody tissues and in the cortex 0

ogy goes to show
f the Abietineae

* Y Oe
14 JEFFREY, Epwarp C., The comparative anatomy of the Coniferales. Part
The Abietineae. Memoirs Boston Soc. Nat. Hist. 6:1-37. p/s. I-7- 194+

1905] CURRENT LITERATURE 307

is primitive for the group. The resin canals persist longest in the reproductive axis,
the leaf, and the first annual ring of root and shoot. In the more specialized genera the
resin canals of the wood are replaced by resin cells, but in the latter condition of the
wood, resin canals may always be recalled as a result of injury. e di f
resin canals and their replacement by resin cells is probably for the sake of economy
of carbohydrate material. In Pseudolarix and Tsuga even the cortical resin canals
disappear from all organs except the female reproductive axis, together with its append-
ages, and the vegetative leaf.

root, shoot, and leaf. It is confirmed by the examination of the female reproductive
organs and of the pollen. It is further in conformity with paleontological evidence.
6.

daitales, the Ginkgoales, and the Cycadales. This feature serves to separate them
from the Cupressineae in the larger sense, and to unite them with the Cordaitales,
which they resemble in’ other important particulars, described in the body of the
memoir,

7. The Abietineae must be regarded on comparative anatomical and morpho-
logical grounds as a very ancient order of the Coniferales, and may even be the oldest
living Tepresentatives of this group.—J. M. C.

SKOTTSBERG,'S botanist of the Swedish Antarctic Expedition of 1901-1903,
gives a preliminary report on the phytogeographical conditions in the Antarctic
Tegions. He proposes for the lands south of 40-50° S. the following names: Aws-
‘ral zone, including Terra del Fuego, with the Isla de los Estados, the Falkland
islands, South Georgia, and no doubt the South Sandwich islands; and Antarctic
zone, including South Orkney and ‘South Shetland islands, and Graham Land.

objection must be raised against the use of the terms Antarctic and Austral
for local geographical areas such as the author is speaking of. The reviewer had
tecently (in a paper read before the Philadelphia meeting of A. A. A. S. 1904)
occasion to mention in passing that the name of Austral zone for a certain phyto-
8eographical area of North America was incorrect, and he holds a similar opinion
in regard to SKorrsBERG’s use of the term. What is to be called the Austral
Zone ? Certainly not a limited area in South America with a few neighboring
Oceanic islands. We have at present an almost endless number of phytogeograph-
‘cal divisions, and this is not the place to enter upon a discussion of the relative
nents of these, but it seems proper here to point out that if we are ever to get a
“onsistent nomenclature in phytogeography it will not do to apply to local or minor
areas Names generally used to designate larger divisions. The Arctic region Is
recognized by DRuDE and ENGLER, for example, as a subdivision of what the
former calls the Northern realm and the latter the North extratropical realm.
ae es to this Northern realm the name Boreal region. The reviewer

% SKorTSBERG, C., On the zonal distribution of South Atlantic and Antarctic
Vegetation, Geographical Journal 655-664. 1904.

308 BOTANICAL GAZETTE [APRIL

would call it the Boreal realm, of which the Arctic region constitutes a part. In
the southern hemisphere we have a corresponding Austral realm, of which the
Antarctic region is a subdivision. What SKOTTSBERG now calls the Austral zone
is a region that may be designated as Patagonian, or Fuegian, or Magellanian,
or by some other local name, and which is a division of equal rank to the Ant-
arctic, subordinating this and several other regions under the term Austral
realm. Our criticism here refers to the use of the terms only, not to the author's
definition or limitations of the areas under consideration. In regard to the flora
of the South Atlantic ocean, the author considers that the division into Austral
and Antarctic is just as evident as in the land flora. Until the detailed results
of the expedition are made known and until this question is thoroughly discussed,
final judgment in the case must be reserved.—OLssON-SEFFER.

IN A PAPER READ before the B. A. A. S. last August, CZAPEK presents’ @
fuller account of his researches on the anti-ferment reaction which occurs in
tropistic movements, especially of roots. In geotropically stimulated roots, for
instance, homogentisinic acid, a product of decomposition of tyrosin by tyrosinase,
accumulates slightly, instead of being oxidized at once, because an anti-enzyme is
produced by the stimulation, which inhibits the action of the oxidase. The quant
tative determination of the homogentisinic acid is not practicable as a measure of
the reaction, but a solution of homogentisinic acid may be used as a reagent for
determining by titration the retardation of its oxidation by the anti-oxidase. The
roots to be tested are ground in water with powdered glass to a thin paste, and a
standard solution of homogentisinic acid added; the initial reducing power of the
wash is determined. and repeatedly at intervals of five days, by titration with
n/to AgNO3. Czapex finds that only tropisms produce the anti-enzyme;
narcosis, poisoning, mechanical hindrance of growth, and wounding having failed
by themselves to produce the reaction. Six minutes was determined to be the
limit for roots placed horizontal at 17° C. to show the anti-enzyme reaction, which
persists for about four hours. Though not observably intensified by prolonged
stimulation of the root, it persists for a much longer time, even up to thirty Liesl
after fifty minutes stimulation. The reaction becomes certain for thirty minutes
induction at an angle of 7° from the normal and at 10° is maximal, remaining con-
stant up to 170°; decreasing very much at 176°, and not showing at all at 180°.
By reducing the time of induction it can be shown that the reaction is stronger
at 135°-150° than at horizontal, which assists in settling a disputed point. Roots
rotated on the klinostat show the reaction, clearly supporting the view that a
tropic stimulation occurs under such conditions though curvature does not resv a
This reaction also confirms CzAPEK’s experiments with glass slipper, esis
that more than the root cap with its statolith cells is involved in the | uaeeilll
phenomena. He lists these phenomena (assuming them to be consecutive) thus:
x6 CzZAPEK, F., The anti-ferment reaction in the tropistic movements of plants
Ann. Botany 19:75-98. 1905.

1905] CURRENT LITERATURE 309
(1) statolith effect; (2) anti-ferment reaction; (3) the processes hindered by shock;
(4) transmission of stimulus; (5) curvature.—C. R. B

A NOTABLE FEATURE of the recent meetings of the affiliated scientific organi-
zations at Philadelphia was the annual discussion before the American Society
of Naturalists, which this year had for its subject the mutation theory. Two
of the seven addresses were botanical, D. T. MacDovuGat opening the discussion
from the standpoint of plant-breeding, and L. H. Barrey presenting the taxo-
nomic bearing of the theory. In the opening address MacDoucat” stated the
main thesis of the mutation theory thus: “the saltatory movement of characters
Tegardless of the taxonomic value of the resultant forms,” its principal corollary
being “‘that the saltations in question do result in the constitution of new species
and varieties.” He believes that many current misconceptions regarding species
are due to the failure to discriminate between elementary species and the com-
posite or “group” species of the systematist; and that there is accumulating
evidence that the supposed deleterious effects of close-fertilization are groundless.

arning is given of the confusion which must arise through the unguarded use
of the terms “mutation” and ‘‘variation” to designate phenomena of segrega-
tion and alternative inheritance in hybrid strains. It will be recalled in this
connection that in a previous paper MacDoucat'® has discussed the difference

tween fluctuating and mutative variations, and has included as mutative
Variations only “newly arisen and transmissible qualities,” emphasizing the
fact that it is a pure presumption to designate any aberrant condition a mutation
until its hereditary character has been demonstrated. Regarding the causes
of mutations little is known, but the results of the mutation cultures both at
Amsterdam and at New York are held to indicate that they arise in greater
numbers under environmental conditions which are especially favorable for
vegetative development and seed-production. Some mutations occur much more
frequently than others, and the highest total number of mutants found in any
Progeny of Onagra Lamarckiana in New York was over six per cent., as com-
Pared with the five per cent. maximum observed by DE Vries in Amsterdam.

“port on other mutating species is promised for the near future—G. H. SHULL.

‘7 MacDoveat, D. T., Discontinuous variation and the origin of species. Tor-
Teya 5:1-6. 1905,

ie MacDoveat, D. T., Mutation in plants. Amer. Nat. 37:737-779- 1993-

*9 Jorpan, D. S., Some experiments of Luther Burbank. Pop. Sci. Monthly
*201-225, figs. 22, 1905.

66

310 BOTANICAL GAZETTE [APRIL

authoritative character of the account. The presentation is given in the form of
quoted paragraphs from conversations with BURBANK, and the assurance is given
that these have been referred to him for verification. This method certainly
has advantages which to some degree counterbalance the disconnectedness and
the frequent repetition of ideas which necessarily result from it. There is
nothing new or unusual in the methods employed by BURBANK, namely, ‘‘selec-
tion, crossing, hybridization, and mutation,” and it is apparent that his success
is dependent upon his great activity rather than upon the methods used. This
paper is of great scientific interest, but of little scientific value, and emphasizes
with unusual force the difference between the economic and scientific ideals.
It will be a source of information to scientists, but certainly of a great deal of
misinformation to the general public, with whom the source of the statements
made will carry a weight wholly incommensurate with their scientific value. It
appears to the reviewer that the compiler of these paragraphs from BURBANK’S
conversations owed it to his readers to explain two features of the technique
employed which would lead to a correct valuation of the statements made.
“Strawberry-raspberry hybrids” cause less surprise when it is known that
insufficient precautions were taken to prevent the entrance of foreign pollen of
unknown origin; similarly, when apple-trees grow from blackberry seeds it is
well to bear in mind that the seeds were sown in unsterilized soil. The dis-
cussions of the mutation theory and other subjects bearing upon evolution will
prove amusing.—G. H. SHULL.

ITEMS OF TAXONOMIC INTEREST are as follows: EpitH M. Farr (Contrib.
Bot. Lab. Univ. Penn. 2: 417-425. 1904) has described a new species of Pachy-
stima from the Selkirks of British Columbia.—W. A. MvrriLt (Bull. Torr. Bot.
Club 31:593-610. 1904), in his ninth paper on the Polyporaceae, has described
Laetiporus, Trichaptum, and Pogonomyces as new genera.—J. A. SHAFER (Torreya
4:177-181. 1904) has separated a new species of Cassia (C. Medsgeri) from C.
marilandica.— In continuing their account of the flora of western Australia,
L. Drets and E. Pritzer (Bot. Jahrb. 35:161-528. 1904) describe numerous
new species and also a new genus (Psammomoya) of Celastraceae—EDWARD L.
GREENE (Leaflets 1:81-96. 1904) has segregated from Streptanthus many of ie
Californian plants that have been referred to it, and has established the following
genera to include them: Euclisia (14 Spp-, 3 new), Pleiocardia (9 spp-, 2 new),
Mitophyllum, Microsemia, and Mesoreanthus (3 spp., 2 new); has repl
Chlorogalum Kunth by the older Laothoe Raf.; has recognized Aloitis Rat as
represented by Gentiana quinqueflora occidentalis and adds two new species: an
has described new species of Batrachium (2) and Sophia (2).—B. F. Bus# sea
Acad. Sci. St. Louis 14:181-193. 1904) has monographed the Texan species .
Tradescantia, recognizing 18 species, 10 of which are new—P. A. coat
(Bull. Torr. Bot. Club 31:631-655. 1904), in his 13th ‘Studies on the ear
Mountain flora,” has described new species in Dodecatheon, Gentianella, > '
(6), Polemonium (2), Lappula (2), Oreocarya (2), Mertensia (6), Stachys, Mon

See

1905] CURRENT LITERATURE 311

ardella, Solanum, Pentstemon (3), Castilleia (3), Valeriana (2), Coleosanthus,
Grindelia (2), Gutierrezia (2), Chrysopsis (4), Solidago (6), Oligoneuron, Chrys-
othamnus (2), and Aster (5).—J. M. C.

Lyon’? has stated more in detail his views as to the phylogeny of the cotyle-
dons of angiosperms. The monophyletic origin of angiosperms is argued with
considerable fullness; a view which recent anatomical studies have helped to estab-
lish, and to which there is probably little dissent at present. The cotyledons are
regarded as primarily haustorial organs, related phylogenetically to the so-called
foot of bryophytes and pteridophytes; a view which seems to be well taken and
cogently argued. The monocotyledonous condition is claimed to be the primitive
one among angiosperms, the dicotyledonous condition being derived from it
through the “bifurcation” of the originally single cotyledon. This last view is
probably the only one that will meet serious opposition, since the recent studies
of the comparative anatomy of the vascular systems of the two groups have con-
tributed great strength to the view that monocotyledons have been derived from
dicotyledons. However, this detail does not affect the general claim as to the
nature of cotyledons. Beginning with the sporophyte of bryophytes, in which
the body is differentiated into two regions called “sporophore” and “haustrum,”
the latter is traced through into the angiosperms and shown to include cotyledon, _
hypocotyl, and primary root. In fact, the hypocotyl is regarded as a new “haus-
tral” structure that is differentiated between the root and the sporophore. The
term “protocorm” is used instead of ‘“proembryo” for the undifferentiated embryo,
and “metacorm” for the “plant body after the differentiation of its permanent
members.” The author announces that a paper is in preparation in which he
“will endeavor to demonstrate the validity of his hypothesis concerning the phy-
logeny of the cotyledon.” —J. M. C

with large mobile starch grains. Roots with 1.5™™ removed remain straight,
having no such new cells. The regeneration of the perceptive complex is not
always complete before the root becomes geotropic, certain older cells often
acquiring motile starch grains and perceptive sensitiveness. also ad —
‘urther evidence from the behavior of inverted root-tips and replies to objections
based upon the results with glass-capped roots, where thigmotropic curvatures
(this in the face of NEwcomBE’s results) may enter. Against CzAPEK’s objec-
tion, that the chemical differences between stimulated and unstimulated roots are
observed whether the statocysts are present or not, it is suggested that there
a,
*° Lyon, HAROLD L., The embrvo of the angiosperms. Amer. Naturalist 39:
13-35. 1905.
NEmec, B., Einiges iiber den Geotropismus der Wurzeln. Beiheft

it : e Bot.
in
Centribl. 17:45-60. 1904

3x2 BOTANICAL GAZETTE [APRIL

may be two kinds of perception, of which that by heavy starch grains alone leads
to curvature, the other (e. g., by radial pressure) leads to the chemical changes
—a strained answer. After defending his ideas against some criticisms by NOLL
and by Mrrue, he pays his compliments to WresNER’s objections based on Clivia
nobilis and C. miniata, the former being geotropic, while the latter is not. But
NeéEmec finds both confirm the statolith theory, inasmuch as in C. nobilis there
is abundant motile starch in the perigonial leaves, and none in C. miniata. Stato-
lith starch is found widely distributed in mosses. It is wanting in certain geotropic
liverworts (Metzgeria furcata), abundant in the strongly it ie T richocolea
tomentella, and scarce in the weakly geotropic Plagiochila —C. R. B.

STRASBURGER”? gives a popular but strictly scientific Aaa of the genus
Alchemilla as it occurs on the Grand Saléve at Geneva. At least thirty-one
species of this genus are found on this one mountain where Lrnn£é recognized
only three. No wide gaps separate these species, but their distinctive characters
are fully constant in inheritance. The author contrasts the Linnean conception
of species as an abstraction with the present conception in which a species is &
real entity which he defines as a group of individuals that agree among them-
selves, and are separated from other species by definite characters that they
inherit, however slight these characters may be. The species of Alchemilla are
believed to be the result of a rather recent period of mutation, the evidence of its
recency being found: (t) in the want of marked gaps between the species, which
would result through the extinction of some species by natural selection; (6)
the almost perfect development of stamens rendered functionless by the apog-
amous habit of nearly all species; (c) the marked development of nectaries that
are likewise functionless and secrete no nectar. The species are regarded as
apogamous rather than parthenogenetic since no reduction of the chromosomes

curs. No case of parthenogenesis in this sense is yet known among higher
plants. Alchemilla is paralleled with two other noteworthy apogamous genera,
Taraxacum and Hieracium, both of which likewise show numerous elementary
forms, and it is suggested that the apogamous habit of these genera may have
resulted through excessive mutation that introduced continuous hybridizations
with attendant deterioration of the sexual processes.—G. H. SHUL

IN AN ADDRESS before section K of the British Association, WARD’ has given
an extremely interesting review of the growth of our knowledge of parasitism
among the fungi and bacteria, from the researches of DEBARY to the sel
time. The first part of the paper deals with the more important earlier dis-
coveries which have marked distinct phases or epochs in the advancement of this
subject, notably those of A. BRauN, DEBARY, TULASNE, ConN, Kocx, and ene

a1 SrrasBuncer. E., Unserer lichen Frauen Mantel. Eine phylogenetisch¢
Studie. Naturw. Wochenschr. 20 : 49-56. 1905.

23 Warp, H. MARSHALL, Recent researches on the parasitism of fungi. An?-
of Botany 19:1~54. 1905.

1905] CURRENT LITERATURE 313

The latter part of the paper is occupied principally with questions brought out
by recent discoveries and theories relating to the rusts. It is shown that the
uredo stage is probably responsible for the wintering of many rust fungi whose
distribution and annual recurrence is not easily explained by the regular alter-
nation of teleuto- and aecidiospores. Specialized races, immunity, susceptibility,
and related subjects are discussed. The mycoplasm theory of Eriksson is con-
sidered to be without foundation and absolutely erroneous in the light of the
speaker’s investigations. Students will find this a valuable paper for refer-
ence, outlining briefly the main questions that have occupied the minds of
investigators at different periods in the development of mycology, while the
bibliography of 209 titles is a means of easy access to the literature of the subject.

ERIksson *4 also read a brief paper on the vegetative life of the Uredineae.
This paper is a review of his mycoplasm theory, mainly as set forth in his former
papers.—H. HASSELBRING.

CAMPBELLS discusses in detail and combats Bower’s theory that the spike
of Ophioglossum is the equivalent of a single sporangium of Lycopodium and
that all pteridophytes are reducible to a common strobiloid type. He repeats his
belief, hitherto published, that the progenitor of the large-leaved ferns may Have
Sprung from some bryophyte type, and that the Ophioglossaceae may have arisen
directly from an Anthoceros-like prototype. He regards a species of Ophio-
glossum from Sumatra described by Bower (Ann. Bot. 18:205. 1904) as nearly
realizing the hypothetical form suggested by him as the fern ancestor. The
close relationship of the Ophioglossaceae and the Marattiaceae is pressed upon
the grounds of structural resemblances in both gametophyte and sporophyte,
and the conclusion is drawn that there is “no valid reason for removing the
Ophioglossaceae from their association with the ferns.” He also reviews the
evidence offered by recent writers in regard to the likeness existing between the
Marsiliaceae and the Schizaeaceae. He thinks there is a marked resemblance
between the sporocarp of certain of the former group and the fertile leaf segment
of Schizaea, both as regards structure and the origin of the sporangium. se
features combined with “the remarkable correspondence in the structure of the
Sporangia”’ he considers sufficient evidence to ally the Schizaeaceae and the Marsi-
liaceae not remotely—FLORENCE Lyon.

IN continvatIon of his experiments with the Erysiphaceae SaLmon?° has
established that i injury to plants in various ways makes such plants susceptible
to biologic forms of Erysiphe to which they are normally immune. The fungus
I a ia

os gt J., On the vegetative life of some Uredineae. Ann. of Botany
Ig: 55-60.

*5 CAMPBELL, D. H., The affinities of the Ophioglossaceae and Marsiliaceae.
American Nati es 168-715 figs. 9. 1904.
- S., Further cultural sa with “biologic forms” of the

20 SALMON,
Erysiphaceae. pe of pina 19:125-148. 1905

314 BOTANICAL GAZETTE [APRIL

used was the biologic form of E. graminis on wheat. This form will not infect
tye, but by subjecting rye leaves to various treatments it was found that the
fungus could be made to infect this plant also. When leaves of rye were injured
either by cutting or bruising or by slugs, the cells around the injured parts could
be infected with conidia or ascospores from E. graminis on wheat. Other
experiments showed that rye leaves subjected to the influence of alcohol, ether, or
heat become susceptible to infection from conidia taken from wheat. In some
of these cases the fungus produced vigorous growth nearly covering the leaves.
Conidia grown on barley in this way retained the power of infecting their original
host, but were unable to infect normal untreated leaves of barley. These experi-
ments are especially interesting, since they contribute an experimental proof
to the generally current notion that plants of ‘weakened vitality” are more easily
attacked by fungi than vigorous plants. The experiments show that at least
in certain cases plants may become more susceptible as a result of injurious
agencies affecting the general health of the plant.—H. HassELBRING.

THE COMPLETED FORM of CLINTON’s??7 monograph of the North American Usti-
lagineae has recently appeared. This monograph is the result of studies carried
on for a series of years at the University of Illinois and completed at Harvard
University. The descriptions are based on the author’s personal examination of
type material so far as this was possible. To the technical descriptions are
appended notes of interest giving the distribution of the species, and special
characters of aid in separating closely related species. Complete synonomy and
references to exsiccati are given, while a bibliography of over 200 titles is found
at the end of the volume. A feature of the book is the tabulated list giving the
distribution by continents of the species treated in the work. This volume forms
an excellent handbook for the study and determination of the smuts of North
America. It will prove extremely useful to students and station workers by whom
the lack of monographs of groups of parasitic fungi has been long felt. It would
have added to the utility of the work to have had the index of hosts arranged
alphabetically, with page references for the parasites; instead of classifying the
hosts according to orders, which is a time-robbing arrangement.—H. HASSEL-
BRING.

Wa Inn shoots

GOEBEL”* has previously shown that Bryophyll

from the growing region at the margin of the leaves if the midrib be severed, OF
if all the buds be removed from the shoot. Further studies’? show that the 6
result follows if the buds be left intact, but inhibited from growing. Ale

J ist.
27 CLINTON, G. P., North American Ustilagineae. Proc. Boston Soc. Nat. His
31: 329-529. I904.
28 GOEBEL, K., Ueber Regeneration im Pflanzenreich. Biol. Ce
1902.
29 GOEBEL, K., Morphologische und biologische Bemerkungen. 1
Studien tiber Regeneration. Flora 92:132-146. figs. 6. 1903-

ntralbl. 22:418-

4. We jtere

ae ee

1905] CURRENT LITERATURE 315

growing points of several shoots were encased in plaster,and after four weeks’
time buds appeared along the margin*of the leaves. Upon removing the plaster
the buds of the shoot continued growing. In this plant the leaf buds arise from
already existing growing points. In Begonia, however, none of these are present
on the leaf, and here also continued removal of the growing point of the shoot
resulted in the development of buds on the leaves.

In Streptocarpus Wendlandi when the inflorescence is removed a number of
the adventitious shoots develop. The interesting feature about these is that they
present seedling characters, viz., each has a short shoot-like axis and one large
leaf corresponding to the large cotyledon, the opposite leaf being almost invisible
or entirely suppressed.—W. B. McCatium.

LECLERC Du SABLON3° finds that the carbohydrate and nitrogenous reserve
foods of trees reach their maxima in the autumn, and their minimain May or June
after the formation of the leaves and shoots of the season. For carbohydrates
the roots are more marked reservoirs than the stems, and the leaves do not func-
tion as a storage place. Nitrogenous reserves, however, are more abundant in
leaves than in stems or roots, diminishing from spring to autumn, at first very
rapidly, then slowly. The stem and roots give up to leaves in process of forma-
tion most of their nitrogenous food, and their store is replenished little by little
during the season. The fats are scarce in stems or roots; they are more abundant
in leaves, in which they increase from spring to autumn. Water is at a maximum
in early spring and a minimum in autumn. Autumn, when the water is at a
minimum and the reserve food at a maximum, is the most favorable time for the
transplantation of trees. SABLON seems to have overlooked some important
German work on reserve food in trees, especially that of MUELLER.—C. R. B.

BERTRAND! obtained from pure cultures of various orchids the same fungus,
and found that the seeds of these species, sowed with this fungus (an Oospora ?)
to the exclusion of other micro-organisms, gave normal plants, regularly infested.
Cypripedium seeds grown aseptically do not develop; Cattleya embryos grow
to the spherule stage and no further unless then infected; Bletia hyacinthina
develops a stem and some leaves, but does not pass this stage unless infected,
when it pursues a normal course. The action of the fungus is to incite growth
In cells other than those attacked. This knowledge will be useful to growers.
In a discussion of tuberization BERTRAND expresses the view that though tuberi-
zation is an anomaly of growth (from the morphological point of view) due to an
abnormal concentration of the sap, in nature this concentration is brought about
not by any of the methods of the laboratory (which have been urged as disproving
this theory of tuberization), but by the attacks of endophytic fungi—C. R. B.
ae ee

3° SABLON, LECLERC DU, Recherches physiologiques sur les matiéres de réserves

des arbres. Rev. Gén. Bot. 16: 341-368, 386-401. 1904.
ERTRAND, N., Recherches expérimentales sur les Orchidées. Rev. Gén.
Bot. 16: 458-476, pls. IS-19. 1904.

316 BOTANICAL GAZETTE [APRIL

It APPEARS from Poropko’s work on the growth of bacteria and molds under
different pressures of oxygen,3? that the maximum pressure of this gas which can
be borne varies, in twenty-five organisms studied, from 0.676 to g.38 atmospheres.
A single nutrient medium was used throughout, and the author points out that
with other media perhaps the maximal pressures might have been different.
The upper limit of oxygen pressure for optimal growth seems not to be related
to the maximal pressure. Furthermore, every aerobe has its specific minimal
pressure of the gas. This point lies considerably higher for the molds than for
bacteria, being about 0.6 atmospheres for Aspergillus, Penicillium, and Mucor,
while it is 0.00016 for Bacillus subtilis. Of course, in case of facultative anaerobes
there isno minimum. The lower limit for most obligate aerobes lies low enough
to allow the growth of obligate anaerobes.—B. E. LivincsTon.

Miss ROBERTSON,33 in continuing her study of Torreya californica, has
described the sexual structures and fertilization. Two to four archegonia are
produced, the neck consisting of a single tier of four or six cells. The division of
the central cell was observed, and an ephemeral ventral nucleus inferred. The
two male nuclei were observed in a common cytoplasmic sheath, but no inequality
in the male cells was noted. In fertilization the cytoplasm of the male cell was
observed investing the fusion nucleus. In the development of the proembryo
wall-formation occurs after four free nuclei have appeared. A count of chromo-
somes indicated that the reduction number is eight. The general conclusion is
reached that the morphological evidence does not bear out the suggestion of se
dean resemblances obtained from the anatomy of the seed and seedling. —J.M.

Grécorre+ has investigated the reduction of chromosomes in the pollen
mother-cells of Lilium speciosum and Allium fistulosum. The immediate occa-
sion for the investigation was that STRASBURGER’S recent conclusions** conflict
with those of BERGHS,3° a pupil of Grécorre. The present investigation con-
vinces Grécorre that his pupil is correct in his interpretations, and that STRAS-
BURGER’S recent interpretation of mitosis in Thalictrum purpurascens will not
apply to Lilium, Allium, and Convallaria. The most interesting mgm
in the present paper concerns alternations of generation. GREGO RE claims
that since reduction is not complete until the four spores are cam anion the

32 PoropKo, THEODOR, Studien iiber den Einfluss der Sauerstofispannuns auf
pflanzliche Mikroorganismen. Jahrb. Wiss. Bot. 41:1-64. 1904

33 ROBERTSON, AGNES, Studies in the morphology of Torreya cali wii

sexual organs and fertilization. New Phytol. 3:205-216. pls. 7-9. 1904

ia, 1. ‘The

34 GREGOIRE, Victor, La réduction numérique des chromosomes et les cine
de maturation. La Cellule 21: 297-314. 1904.
: - XK, Preus.
35 STRASBURGER, Epuarp, Ueber Reduktionsteilung. Sitzungsb. K. K. Pr

Akad. Wiss. 18:—: [1-28.] 1904. See Bor. GazeTre 38:397-398- 1994:
TO-
3° BERGHS, JULES, La formation des chromosomes hétérotypiques oe a tak
géntse végétale I. La Cellule 21:173-189. 1904. See Bor. GAZETTE es

1905] CURRENT LITERATURE’ 317

mother-cell, the gametophytic generation begins with the spore rather than with
the spore mother-cell—CHaries J. CHAMBERLAIN.

BoRrGESEN and JENSEN37 have given a floristic description of a portion of
heath in Jutland, Denmark. On the suggestion of WARMING, a part of the estate
now meadow has been reserved for experimental purposes, the object being to
trace the developmental phases from cultivated soil to heath. Another portion
of the heath was burned over in order to trace the progressive stages from no
vegetation until the tract is again converted into the Calluna heath. Another
object in view is to trace the succession of plants on areas which now are or are
to be planted with conifers. The data of the present condition seem to be thor-
oughly detailed and classified. Hence it would seem that future observations
on this tract will furnish interesting and valuable material for the ecologist —
G. H. Jensen. ,

lactic acid is one of the intermediate products in alcoholic fermentation (with
which StToKLAsA absolutely identified anaerobic respiration), in the light of the
researches of Ner. At the close of his paper StoxtasA hints darkly at results
which indicate that chlorophyll-free plant cells form CHOH as in photosynthesis,
oe being reduced by nascent H to formaldehyde, water being again formed.
ok. B.

Livincston finds3° that the cations of many metallic nitrates and sulfates
act in the same way and at the same concentration upon Stigeoclonium, producing
death at certain concentrations, causing the filamentous form to become palmel-
loid at lower concentrations, and sometimes at the latter concentrations, but
oftener at a still lower one, accelerating in a marked degree the formation of
Zoospores. The palmella effect is identical with that produced by withdrawal
vd Water, but the zoospore effect is exactly the opposite. The degree of tox-
‘city of the metals found agrees in a general way with that found by other observers,
though there are many unexplained discrepancies.—C. R. B.

N THE BASIS of comparative researches with the quartz spectrograph,
TScurrctt comes to the conclusion? that the yellow pigments of flowers and

ei Bor SEN, —, and Jenson, —, En floristisk Underségelse af et Stykke Hede
: Vestjylland. (A floristic research on a portion of heath in West Jutland.) Bot.
ldsskrift 26 :—___. 1904
38 STOKLASA, .. Ueber das Enzym Lactolase, welches die Milchsaiurebildung in
der Pflanzenzelle verursacht. Ber. Deutsch. Bot. Gesell : 460-46 On:
39 Livingston, B. E., Chemical stimulati f a green alga.
32°1-34. figs. 17, 1905. ,
IRCH, A., Vergleichend-spektralanalytische Untersuchungen, etc. Ber
Deutsch. Bot. Gesells. 22: 414-439. 1904.

Bull. Torr. Bot. Club

318 BOTANICAL GAZETTE [APRIL

fruits betray no close relationship with artificial yellow pigments, a large
number of both being investigated. Rather the plant pigments show alliances
with the phytosterins. Carotin, he adds, probably is allied to THIELE’s /ulvin,
and its chromatophore radicles contain a quinate ring of which at least three
C-atoms have double bon

VuILLEMrn*! has published a paper on the development and structure of the
membrane of the zygospore in the mucors. The main contribution of the paper
is the recognition of the fact that the membrane is more complex than has usually
been supposed. The author distinguished five distinct layers which differentiate
successively and differ from each other in thickness, appearance, and in their
behavior toward reagents. The aay includes Sporodinia, Spinellus, Zygorhyn-
chus, and Mucor.—H. HAsSELBRIN

AN INVESTIGATION under the sae of PALLADIN by L. PETRASCHEVSKY*?*
shows that when Chlorothecium saccharophilum is fed with raffinose its respiratory
quotient in anaerobic life is raised much above 1 (in aerobic life it is 0.74—0-89);
while with maltose it falls below the normal. Acids are probably produced as
oo aga in the former case and alcohol-like substances in the
latter.—

Parla in a greenhouse modifies notably the external form, the habit,
and the structure of plants, according to the experimental studies of BEDELIAN.*$
This he ascribes to the humidity, nearly uniform temperature, and the weaker
diffuse light. In general differentiation is more or less arrested, and there are
manifested various adaptations to the new conditions.—C. R. B.

FRIEDEL finds the effect of insufficient oxygen on the anatomy of certain
plants quite parallel with that of darkness. The relative thickness of the cortex
is increased, that of the pericyclic region is diminished, and lignification is incom-
plete.44 The experiments, however, cannot be continued long enough to make
these effects as marked as in etiolation.—C. R. B

KeLticorr‘s finds in roots of Allium and Podnplepthain two maxima (c. rand
_ 1 P.M.) and two minima (c. 7 A. M. and 3 P.M.) in cell division, which are not

affected by slight variations in temperature, but are affected by solutes in ne
water. Elongation maxima and minima in Allium coincide respectively
minima and maxima in cell division. —C.

4t VUILLEMIN, P., Recherches morphologiques et morphogéniques Sur la mem-
brane des zygospores. Ann. Mycol. 2:483-506. pls. 8-II. 1904-

42 PETRASCHEVSKY, L., Ueber Atmungscoefficienten der einzelligen Alge ail
thecium sacchara sibs. Ber. Deutsch. Bot. Gesells. 22:323-327- 1994

43 BEDELIAN, J., Influence de la culture en serre sur quelques plantes ee
de Paris. Rev. Gén. Bot. 16: 318-336. pls. I0-13. 1904-

44 FRIEDEL, JEAN, Influence d’une faible pression d’oxygen
anatomique des plantes. Rev. Gén. Bot. 16:305-317- 1904-

48 Ketiicott, Wm. E., The daily periodicity of cell division and
the root of Allium. Bull. Torr. Bot. Club 31:529-550. figs. 8. 1904-

e sur la structure

of elongation "

NEWS.

Dr. G. Bitrer, formerly of the University of Miinster, has been appointed
director of the recently established Botanical Garden at Bremen.

Dr. W. Micuta, formerly at the Technical School at Carlsruhe, has been
appointed professor of botany at the School of Forestry at Eisenbach.

Prorrssor CHARLES R. BARNES, University of Chicago, sailed for Europe
April 8, to be absent seven months. He will attend the International Botanical
Congress at Vienna.

Tar Cepar Pornr LAKE Laporatory of the Ohio State University offers
instruction in general botany and ecology from June 26 to August 4. e
instructor is Otto E. JENNINGS, Carnegie Museum.

THE THIRD ANNUAL MEETING of the Central Branch of American Society
of Naturalists and Affiliated Societies, held at Chicago March 31 and April x,
the annual address of the chairman, PROFESSOR JoHN M. Courter, University
of Chicago, was upon ‘‘Public interest in research.”

HE MARINE BroLocicaL LABORATORY at Woods Hole, Mass., offers instruc-:
tion in Thallophytes and Cell studies from June 28 to August 9, 1905. The
lnstructors are BRaDLEY Moore Davis, University of Chicago; James J. WOLFE,
Trinity College, N. C., and Ropert B. Wyu1e, Morningside College, Iowa.

Tue Borantcat Socrety of the Netherlands has begun the publication of
Its Archives in two parts. The title of one is Nederlandsch Kruidkundig
Archie}, dealing with the Dutch flora and printed in- Dutch ; that of the other is
Recueil des travaux botaniques Néerlandais, dealing with subjects of more gen-
eral interest.

Tur Bermupa BrotocicaL STATION FOR RESEARCH will be open in 1905
under the direction of Professor E. L. Mark, of Harvard University, and Pro-
fessor C. L. Bristor, of New York University. Botanists who wish to take
advantage of the station must apply not later than May 1. The date of sailing
from New York is July 1 and six weeks will be spent at the station.

Wirt rrs January number the Indian Forester commences its thirty-first
volume with a great improvement in appearance. In typography, size, and —
tents the subscription price (12s 6d) is exceedingly low. While the journal
devotes considerable space to the discussion of forest problems that confront the
Forest Service in India, it discusses problems of much wider scope and is of
8eneral interest to all students of forestry.

T9035] 319

320 BOTANICAL GAZETTE [APRIL

Tue AMERICAN Myco.ocicar Society met in affiliation with the A. A. A.S.
at Philadelphia, December 28 to 31. The following officers’ were elected:
President, CHARLES H. Peck; Vice-President, F. S. EARLE; Secretary-treasurer,
C. L. SHear; and the following committee on organization and relation to the
‘other societies was appointed: C. L. SHEar, S. M. Tracy, and ROLAND
THAXTER. The following program was presented: Suggestions for the study
of dairy fungi, CHARLES THom; A new disease of the cultivated agave, GEORGE
G. Hepccock; A study of North American Coleosporiaceae, and The termi-
nology of the spore-structures in the Uredinales, J. C. ARTHUR; Classification of
the Geoglossaceae, E. J. DuRAND; Generic characters of North American The-
lephoraceae, E. A. Burt; Cultures of wood-inhabiting fungi, PERLEY SPAULDING;
Two fungous parasites on mushrooms, and The genus Balansia in the United
States, G. F. ATKINSON.

Tue “Boranists of the Central States” met at the University of Chicago,
March 31 and April x, in connection with the Central Branch of the American
Society of Naturalists, with STANLEY CoutTeR, of Purdue University, as chair-
man, and Cuas. F. Mritspavcu, of the Field Columbian Museum, as secretary.
The following papers were read and discussed:

C. F. Hortes, University of Illinois, The effect of starvation on the cells of
the root tip; W. B. McCatium, University of Chicago, Regeneration and polarity
in the higher plants; R. H. Ponp, Northwestern University, The endosperm
enzyme of Phoenix dactylijera; E. W. Outve, University of Wisconsin, Nuclear
‘and cell division in certain lower plants; E. O. JORDAN, University of
Chicago. A rosette-forming micro-organism; F. M. Lyon, University of Chicago,
Development of the megaspore coats of Selaginella; R. A. HARPER, University
of Wisconsin, The nuclear fusion in the ascus, and alternation of generations ™
the Ascomycetes; A. H. Cuerstuan, University of Wisconsin, Sexual reproduc:
tion in the rusts; B. M. Davis, University of Chicago, The sexual organs ot the
Rhodophyceae and Uredinales; C. J. CHAMBERLAIN, University of Chicago,
Apogamy and apospory in ferns; E. B. Simons, University of Chicago, The
morphology and development of the conceptacle in Sargassum; J. B, Oven
University of Wisconsin, The permanence of the chromosomes and their behavior
in synapsis; C. F. MirtspaucH, Field Columbian Museum, Recent field-work
in the Bahamas.

A permanent organization of the society was effected, with Wi.
LEASE, Missouri Botanic Garden, as president, and H. C. Cowles,
of Chicago, as secretary-treasurer.

pram TRE-
University

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City, Examining Physician Corporation Comers. ete. aoe
John V. Shoemaker, M. D., LL. D., Professor Materia
Medica and Therapeutics, a College, Philadelphia.
Dr. Geo earns e Ben. Johnston, Va., Fi iano
Souther: na nar ane Cynccological prmoncarciss Ex-President
Medical Societ nd Professor of of Gynecology end Aicpectea
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and Clinical yes doe New York Post Graduate Medical School.

Louis C. Horn, M. D., Ph. D., Prof
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ol. XXXIX, No.5 : Issued May 20, 1905

ree

CONTENTS
eeOUY TIC: ENZYMES, I: Arthw L.\Dean - “*- SSR SS a ten = = ‘gal

O relly OF SPORANGIUM IN BOTRYCHIUM. Conrrrutions FROM THE
ULL BoTanicaL LABoratory. LXXI (WwiTH PLATE Ix). Ira D. Cardi, iff 340°

LOGICAL PROPERTIES OF BOG WATER. Cowtrisuti0ns FROM THE Hoi
Borantcat Laporatory. LXXII (WITH THREE.FIGURES). Burton Edward Livingston . 348

S FOR OBSERVING ee TRANSPIRATION STREAM, (wit TWO
Geten:), Otto V. Darbishire ef 356

POLYEMBRYONY IN SPHAGNUM (WITH THREE FIGURES). Harold L.Lyon -- -® = 365
ADENODERRIS, A VALID GENUS OF FERNS (WITH TWO FIGURES). William R. Maxon 366 ~

5 - hs - ~ “ - ~~ 37
RUSTS OF SWITZERLAND
4NOR NOTICES - 2 : 8 '§ . is * w on FI
TES FOR STUDENTS - - - - : - : : - 374
a ee es oe ee a ee
ications for the Edit } $22.4. 4] 4 ech cago, Til

for AZETTE.
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Copyright, 1904, y the University of Chicago

VOLUME XXXIX NUMBER 5

HOTANICAL (GAZETTE
MAY, 1905

ON PROTEOLYTIC ENZYMES. I.
ARTHUR L. DEAN.

NUMEROUS investigators have studied the proteolytic enzymes
in plants and their work appears to have been carried on from two
different points of view. The physiological chemists working with
vegetable proteases have made careful studies of the products and
nature of the action of enzymes of marked activitity. Of such
work, the investigations carried on by CHITTENDEN and his pupils
on bromelin are excellent examples. In studies of this kind the
enzyme is the important thing, the réle of the enzyme in the physi-
ology of the plant is secondary. On the other hand, the studies
of plant physiologists have been directed less to working out the
nature of the enzymes and more to locating them and to discovering
their function in the life of the plant. Of such researches the inten-
sive studies of BUTKEWITSCH on germinating seeds, and the extensive
ones of VINEs on all sorts of plant tissues, are good illustrations.

Looking at the subject from the point of view of plant physiology,
we may conceive of proteolytic enzymes as being digestive, 7. ¢.,
acting on stores of proteid, as in germinating seeds; or metabolic,
* €., acting on the proteids in the life of the vegetable cells. The
idea of enzymes digesting the reserve proteids was the first to occur
to physiologists. Acting on the hypothesis of the existence of such
snzymes, numerous investigators have examined germinating seeds,
"sing a variety of methods, and obtaining results which have not
always been in harmony with the hypothesis or with each other.

The view that wherever we find proteids in an organism, there
we should always find a proteolytic enzyme, is not a new one; the
€xperimental facts to support such a conception have been slow

321

322 BOTANICAL GAZETTE [MAY

in coming. Manifestly the first step in proving this assumption
is to show that proteolytic enzymes are present in all parts of all
living plants, or to demonstrate their presence in a sufficient number
of places to warrant the conclusion that they are universally present.
Such extensive researches have not been undertaken until the last
few years.

I shall briefly review the results of the various investigations,
taking first the studies on the digestive enzymes, as I have called
them, and then those on the metabolic enzymes.

Gorup-BESANEZ' appears to have been the first to examine
seeds for proteolytic enzymes. Using the Wittich glycerin method
he obtained protease preparations from the resting seeds of hemp,

ax, and vetch, and from the germinating barley; he failed to find
enzymes in ungerminated barley, lupin seeds, Secale cornulum,
pine, and corn, and in seedlings of almonds and beans.

VAN DER Harst? found that a glycerin extract of the cotyledons
of germinating beans would digest fibrin.

The first careful study of a seed proteolytic enzyme was that of
J. REYNOLDS GREEN? who investigated the germinating seeds of
Lupinus hirsutus. GREEN’s results showed conclusively that these
plants contain an active enzyme, capable of attacking fibrin, as well
as the proteids of the seeds themselves. The enzyme was not present
in the resting seeds. Later GREEN discovered a similar enzyme
in the seedlings of the castor oil bean.*

NEUMEISTER’ undertook the study of a number of seeds and
seedlings, using a method different from that of any previous investi-
gator in this field. Pieces of fibrin were soaked in the watery extracts
from the tissues to be tested; subsequently the fibrin was trans
ferred to a weakly acid solution and allowed to digest. In case @
proteolytic enzyme was present it was taken up by the fibrin, and
in the acid medium proteolysis of the fibrin took place. The value

« Gorup-BESANez, Ber. Deutsch. Chem. Gesells. '7:1478. 1874; 8:1510- 1875} “
9:673. 1876.

2 VAN DER Harst, Moonblad voor Naturwetensch. 1867, Sept. 29-
3 GREEN, J. REYNOLDS, Phil. Trans. 1'78:(B) 39. 1887.

4 GREEN, J. REYNOLDs, Proc. Roy. Soc. 48:370. 1890.

5 NEUMEISTER, Zeitschr. fiir Biologie 30:447. 1894.

1905] DEAN—PROTEOLYTIC ENZYMES 323

of the negative results by this method has been called in question
by FRANKFURT.° NEUMEISTER’S results may be summarized as
follows:

POSITIVE RESULTS NEGATIVE RESULTS
Barley seedlings—sprouts 3°™ long. Barley seeds.
Barley seedlings—sprouts 5 °™ long. Barley seedlings—sprouts 0.5 to 1°™.
Barley seedlings—sprouts 15-20°™ long.
Poppy seedlings. Poppy seeds.
Beet seedlings. Beet seeds.
Corn seedlings—sprouts 10 °™ long. Corn seeds.
Corn seedlings—sprouts 16°™ long. . Lupin seeds.

Wheat seedlings—sprouts 17°™ long. Lupin seedlings—sprouts 3°™.
Lupin seedlings—sprouts 12°™.
‘ Lupin seedlings—sprouts 20°™.
Vicia see
Vicia seedlings.
ea seeds.

Oat seedlings.

_ A number of different investigators have worked on malt; the
evidence shows that the germinating barley contains a proteolytic
*nzyme. WErss’ thinks there are two, a peptic and a tryptic enzyme,
. view which is supported by FERNBACH and Husert,® but ques-
tioned by Vings.

In the course of the extensive studies of Fermi and BuscaLIont®
on proteases in plants, they tested a number of seeds and found
that the ripe seeds of several plants contain an enzyme capable
of liquefying gelatin. They likewise found that the germinating
Seeds of some plants contained such enzymes, but in other cases
no evidence of their presence could be found.

BurKEwirscu'® made a careful study of the germinating seeds
of two lupins, Z. angustijolius and L. luteus, and a somewhat shorter

° FraNxkrurr, Landw. Versuchsstat. 47:466. 1896.

7 WEIss, Zeitschr. fiir Physiol. Chem. 31:79.

8 FERNBACH and HuBeErt, Compt. Rend. 130:1783.-

® FER and Buscatront, Centralbl. Bact., etc. II. 5:24. 1899-

*° BUTKEwrrtscu, Zeitschr. Physiol.-Chem. 32:1. 1got.

324 BOTANICAL GAZETTE [MAY

examination of Ricinus major and Vicia Faba. His experiments
show that the proteids in the sprouting seeds of all these species
will undergo autolysis, giving rise to products not precipitable by
phosphotungstic acid. From L. /uteus he made a proteid prepara-
tion by the Wittich method (solution in glycerin and precipitation
by alcohol) which acted on “conglutin” with the formation of leucin
and tyrosin, but not of asparagin.

Vines"! found some difficulty in getting a formation of tryptophan
in the autolysis of germinating seeds of Vicia Faba; the reaction
was readily obtained, however, if Witte peptone was added to the
mixture and digestion allowed to proceed. He found that the embryo
of the wheat contains an enzyme which readily attacks Witte peptone
and apparently has some slight action on some of the proteids of
the embryo.

As stated above, the first step in demonstrating that proteases
are universally at work where proteids occur, is to show that all
living plant tissues contain proteolytic enzymes. Investigations
of this sort have been undertaken but recently. The older researches,
which did not have this purpose, did tend to show that active pro-
teases occur in places other than germinating seeds. The papain
of the pawpaw fruit, the bromelin of the pineapple juice, the cradein .
of the latex and juice of the fruit of the fig, and the protease of the
Kachree gourd and of Anagallis arvensis have been known for some
time, but these plants were looked upon as exceptional.

Fermtand Buscationr? were the first to make a systematic examl-
nation of a large number of plant tissues. Their method consisted
in placing the objects to be tested on gelatin containing phenol as an
antiseptic. If the gelatin was liquefied by the tissue or tissue extract,
a protease was judged to be present. The positive results of this
method are valuable; negative results cannot be taken as conclusive
evidence that no proteolytic enzyme is present, since all proteases
do not attack gelatin, and phenol is said to have an inhibitory influ-
ence on the action of some enzymes. These investigators succeeded
in showing that proteases occur in a great diversity of plants and
of plant parts. Their negative results are sufficiently numerous

™ VinEs, Annals of Botany 16:12. 1902.

12 FERMI and BuscaLiont, Centralbl. Bact., etc., II. 5:24. 1899-

1905] DEAN—PROTEOLYTIC ENZYMES 325

to cause some doubt in respect to the universal occurrence of pro-
teases in living tissues, and the positive results were obtained with
such a diversity of tissues that they do not lead to any definite hypothe-
sis of the physiological réle of these enzymes. In view of the evidence
of VINES’ researches, it appears probable that a more thorough
study of the tissues examined by Fermi and Buscatront would
show that proteases are present there which cannot act under the
conditions of their experiments. Their positive results support
: the theory of the universal distribution of proteolytic enzymes,
their negative results do not disprove it.

Finally we come to a consideration of the results of VinEs."3
He believes that all vegetable proteases are proteolytic in distinction
from peptonizing, i. ¢., that they change the proteids which they
attack into their ultimate cleavage products, amido-acids, hexon
bases, etc., and do not stop, like pepsin, with the albumoses and
peptones in preponderance. Acting on this well-justified assump-
tion, he expects to find tryptophan as the result of the action of
vegetable proteases, and uses as a test for proteolysis the progressive
formation of tryptophan as shown by the color reaction with chlorine
or bromine water. He uses the method of autolysis, allowing the
€nzymes of a tissue to digest the proteids of that tissue and from
time to time tests samples of the mixture for tryptophan. He fre-
quently adds other proteids to the digestion, especially Witte peptone.
The only objection to the method is that some proteids do not contain
| the tryptophan group, or only a small amount of it. A negative
. tesult by Vines’ method with a tissue containing such a proteid would
7 hot mean much. This probably explains the differences between
the results obtained by Vines and BuTKEWwITscH in their exam-
inations of the germinating seeds of Vicia Faba, since OSBORNE
and Harris‘ have shown that the proteids of this seed give but
4 faint Adamkiewicz reaction, a reaction which Hopkins and
CoLe"s have shown to be due to the presence of the tryptophan
Stoup in the proteid molecule. By using Witte peptone, VINES
obtains evidence of proteolysis in cases where the tissue contains

a a

13 VINEs, Annals of Botany 16:1. 1902; 1'7:237 and 597. 1903; and 18:289. 1904.
“4 OSBORNE and Harris, Jour. Am. Chem. Soc. 25:853- 1903-
*S Hopkins and Cote, Jour. Physiology 27:419- 1901.

326 BOTANICAL GAZETTE [MAY

no peptonizing enzyme, but does have an enzyme which, like erepsin
of the animal body, attacks albumoses and peptones. In cases
where all the trials were made with the addition of Witte peptone,
it is impossible to say whether there is an enzyme present of the
trypsin type (tryptase), the erepsin type (ereptase), or both. In
cases where a positive result was obtained both with or without
the addition of Witte peptone, a tryptase is present if, as is probable,
the tissue contains no marked quantity of albumoses or peptones;
an ereptase may also be present and in the table its presence is indi-
cated as questionable. When no tryptophan reaction is given on
autolysis, except after the addition of a proteose, I have assumed
that no tryptase was present, an assumption which, as pointed out
above, is not entirely justified because certain proteids give no reaction
for the tryptophan radical (see table).

This table records but three failures to obtain evidence of a pro-
tease, the pulp of the apple, the juice of the orange, and the sap
of the bleeding birch tree (Betula alba), which last is not a cell extract.
The two failures, the apple and the orange juice, are scarcely suffi
cient to prevent the acceptance of these results as evidence of the
‘universal occurrence of proteases in living plant tissues.

No mention has been made of the proteolytic enzymes of the
insectivorous plants and the bacteria and lower fungi. Some of
the latter are doubtless endo-enzymes and belong to the group which
I have termed metabolic enzymes. The others are apparently
secretions designed to render food materials available and should
be classed with the digestive enzymes.

The results of Vines tend to show that there are enzymes of the
erepsin type widely distributed in plants, yet the evidence is not
wholly conclusive and he has made no attempt to isolate such enzymes
DELEZENNE and Mouton’? made extracts from several fleshy fungi,
Amanita muscaria, Amanita citrina, Hypholoma fasciculare, and
Psalliota (Agaricus) campestris, which were judged to contain an
ereptase since they digested proteoses but not fibrin. VINES finds,
however, that Agaricus campestris, which DELEZENNE and MOUTON
found to yield an especially active ereptase, is capable of autolysis
and will digest fibrin if the fibrin is not boiled. The unboiled fibrin

76 DELEZENNE and Mouton, Compt. Rend., April 9, 1903-

= DEAN—PROTEOLYTIC ENZYMES

327
Erep-
Tryp- | Erep- | tase or
Tissue — = Pla
both
ee Or MONEE 4 i fe ks Eas Hen Roa Rd Lae ees - = a es
Ae AGSrISis CONDOMS. «6.6.0. vos jens deena gedendenmen a et :
Mpa Seeds OL Pisin Salvi. 53 o0.0 0 oc od eke eee + ce o-
SO MERWE WEE 5 6. ciess sas bs Bias ove aR Oxi eee Le ee a oa i“
RI Cr CMONINES SORTOUS oo 6s once accede g cede ee a Hd
Fruit of Cucurbita pepo-ovifera........0. 0c eae cnceeceeeeees se a
Fruit of Echallium Eloterium.......000cc0ccecceneseseveeons = Le ;
RIE Ol Musa sa pientwi.. 6 5 occ. va nin v1 gedeee rns toes + 77
Fruit of Lycoperiscum esculentum...........2.2-+0200e eee eees ai ‘ =
emcee Fs wieted (ENED)... . ow nseiare nen Dw nth Sead 2 R
eee ee 2 WORS MeAbee (Sheen). 60.25 ac sao dsc saem ae cae a -
Fruit of Pyrus communis............ Salo Rhe Met enToe eee og ae
Fruit of Citrus aurantium ( JRICE) Soke tenth aee ete res é -
Fruit of Citrus aurantium (peel). ..........020c0eeeeeeeeeees
PRO Vr TAR MOIE .. s sven Caecum emadans Soran ae \ om
Of Se eternal) aeowe: Ur tet ot 2 oe sboe a
Rumen da Creme MeaeiNned. cis vu ce avcd-deaveoesvenes eS ae
BRA Nachos FB ccc eta. ihe aoe 4. be lek Cane ft etek: Remeatecmrea meet rye bi 7 i
Latex of Euphorbia Characias. ........0cccensegeccneveccers pee zi
OE LAAN SONNE. figo sv ok vena vaansctiounic cee eenters bab Fees es
S RUPE ANN GOT AD IE ood cag sews ho eae Oe ie : a eee
Ste f Cucurbita pepo-ovifera..... i... cece anodes deusvente ; . > -
CPIM MMUMNS Sula. «5 9 co ook de inn ces v ees 4
Stems of Helianthus tuberosus...cccccccccucecncucecceeeetes t i o
RPO MMMNN fo fo oc as os wh nh du nated Cala me . ia owe
Leaves of Spinacia oleracea. 0.05. 6iccccseseureanerecersbes id 4 es
Leaves of Dahlia variabilis..... 00.02. ccccuccucceeceueeeues ‘ ee ere
Leaves of Tropacolum majus..........2..cccceeeeeeeeeceees id eS Be
Leaves of six other species of plants. .......- a Sie phehlge heels .
ves of Holcus mollis and three other species of plants. ..... Pe res fe
Bulbs of Twlipa and two other plabieccs Puget eee aS ad eae
Tubers of Solanum tuberosum. .......02..cc0cceceeeeeeeeees . + ?
Tubers of Helianthus tuberosus. .......0.0.0ccceeeceeeeeces : ot
Roots of Dahlia variabilis...........00ceeeene vee eeeeeeeres - i
Roots of Brassica Rapa and seven other plants.......----+++-

which VinEs used would not digest itself and he believes its solution
Was not due to any enzyme which the fibrin itself contained. VINES
agrees with DeELEzENNE and Movron in believing that this fungus

Contains an enzyme of the erepsin type.

EXPERIMENTAL DATA

In undertaking the study of the proteolytic enzymes I have had
two ends in view. In the first place it was thought desirable .
test a number of different plant tissues in order to confirm, if apes
the results of Vines. The second and more important part of the
investigation was to make a careful study of the enzymes In some

328 BOTANICAL GAZETTE [MAY

one or two plants, tracing their occurrence and studying their nature
throughout the life history of the plant. In this way it was hoped
that some results might be obtained which would throw light on
the nature and functions of the proteases in the life of plants. For
this study the common field bean, Phaseolus vulgaris, which has a
large proteid content in its seeds and shows marked vigor and rapidity
in its growth, was selected. The work on the enzymes of Phaseolus
has proved so absorbing that scant attention has been paid to the
testing of various plants. The few positive results which have been
obtained will be set down at this time.

SPINACIA OLERACEA.—Fresh leaves of spinach were carefully
picked over to remove all imperfect tissue and then ground up and
the juice removed by a screw press. The residue was rubbed up
with a little water and the fluid again pressed out. The turbid
dark green extract was examined as follows:

Flask no. 1—g50°¢ extract—boiled.

Flask no. 2—s5o0°° extract—unboiled.

Flask no. 3—50° extract +.5 8™ Witte peptone—boiled.

Flask no. 4—50°¢ extract +.5 ®™ Witte peptone—unboiled.

Flask no. s—so°¢ extract +.5 8™ Witte peptone +0.25 ®™ citric acid—boiled.

Flask no. 6—50°* extract +.5£™ Witte peptone + 0.25 ®™ citric acid—unboiled.

The antiseptic was chloroform, the length of digestion 30 hours.
Ten cubic centimeters of each digestion were tested by acidifying
with acetic acid, drying, taking up in alcohol and filtering off the
residue, drying the alcoholic extract, taking up the residue in water,
and applying the tryptophan test with chlorine water. The results
were negative except in the case of no. 4, which showed a fair trypto-
phan reaction. This result shows that the leaves of spinach contain
an enzyme capable of attacking Witte peptone, but not acting on
the proteids of the tissue, or if it does these proteids do not yield
tryptophan on hydrolysis.

BRASSICA OLERACEA.—The leaves of a half-matured cabbage,
some of which were green and some etiolated, were run through
a chopping machine and the pulp macerated with water. The
fluid was removed by a screw press and the pale green turbid
extract filtered. This extract was faintly acid to litmus, but not
to lacmoid, and yielded a coagulum on boiling, the filtrate from

—

1905] DEAN—PROTEOLYTIC ENZYMES 329

which gave a faint tryptophan reaction. The extract was examined
in the following way:

Flask no. 1—s5o0°¢ extract.

Flask no. 2—so°¢ extract—boiled.

Flask no. 3—50°¢ extract +1°° 4 per cent. HCN

Flask no. 4—50°¢ extract +1°° 4 per cent. HCN—boiled.

Flask no. 5—5o°¢ extract +1°° 4 per cent. HCN + 0.5 &™ Witte peptone.

Flask no. 6—so0°° extract + 1°° 4 percent. HCN +0.5 &™ Witte peptone—boiled.

Flask no. 7—50°° extract +0.25 Na, CO;.

Flask no. 8—so°¢ extract +0.25 Na, CO;—boiled.

Ten drops of toluol were added to nos. 1, 2, 7, and 8. The flasks
were corked and placed in the incubator. After twenty hours’ diges-
tion 10° was removed from each flask, boiled, filtered, and 1°
of acetic acid added to each filtrate. The tryptophan test with
chlorine water showed that proteolysis had taken place in all the
unboiled digestions except no. 3; that in no. 1 was feeble compared
with the action in nos. 5 and 7. These results show that although
the enzymes of the cabbage leaves act on the proteids of the tissue,
yet that action is slight compared with that exerted upon Witte
peptone. The tissue residue from which the extract had been
obtained was shaken up with water and toluol, a small portion
temoved and boiled, and Witte peptone added to both parts. After
two days’ digestion a sample of the fluid in each flask was tested for
tryptophan; the unboiled digestion gave a strong reaction, the
boiled one a very faint reaction. The fluid was removed as far as
Possible from the unboiled digestion mixture, concentrated to a
Sytup, treated with several volumes of alcohol to remove the pro-
teoses and peptones, and the filtrate concentrated to a thick syrup.
After standing, very characteristic crystals of both leucin and tyrosin
Separated.

Daucus Carota.—Blossoms of the wild carrot were ground
in a mortar with water. The mashed tissue and water were divided
into four parts and placed in test tubes.

Tube no. 1—tissue + H,O + toluol + uncoagulated egg albumen.
Tube no. 2—tissue + H,O + toluol + uncoagulated egg albumen—boiled.
Tube no. 3—tissue + H,O + Witte peptone + toluol.
Tube no. 4—tissue +H,O + Witte peptone + toluol—boiled.
The egg albumen was added because a test of the tissues with

330 BOTANICAL GAZETTE [May

Adamkiewicz reaction, using glyoxylic acid and concentrated sul-
phuric acid, gave no well-defined positive result. After two days,
digestion no. 3 was found to give a weak tryptophan reaction, whereas
none of the others gave a positive result. An experiment carried
out in the same way with the developing seeds gave the same result
except that the unboiled test with Witte peptone gave a much stronger
reaction for tryptophan, showing that the proteolysis had been
more vigorous. The evidence indicates that the enzyme present
is an ereptase.

CASTANEA SATIVA AMERICANA.—-The leaves of this tree were
ground with sand in a mortar, the pulp mixed with water, divided
between four test tubes, and examined as follows:

No. 1—toluol + uncoagulated egg albumen.

No. 2—toluol + uncoagulated egg albumen—boiled

0. 3—toluol+ Witte peptone
0. 4—toluol+ Witte peptone—boiled.
After two days in the incubator no. 3 gave a faint reaction for tryp-
tophan, the others none. The original pulp from the leaves gave
no Adamkiewicz reaction. The unripe seeds of the chestnut were
ground to a paste and mixed with water. No Adamkiewicz reaction
was given by this mixture, which was divided between four test
tubes and examined in precisely the same way as the leaves. The
result was qualitatively the same, but the tryptophan reaction in
the unboiled trial with Witte peptone was very strong, showing the
presence of an active protease, probably an ereptase.

PHasEoLUs Munco.—Three weeks old etiolated seedlings of
this bean were rubbed up in a mortar and covered with glycerin
and a little toluol. After extracting for twenty-two hours, the gly-
cerin was somewhat diluted, strained through gauze, and filtered
through absorbent cotton. This extract was tested as follows:

No. 1—10° extract + fibrin.

No. 2—10°¢ extract boiled + fibrin.

No. 3—10° extract + Witte peptone.

No. 4—10° extract + Witte peptone—boiled.

Examined after three days’ digestion no. 1 showed no digestion
of the fibrin and gave no test for tryptophan, no. 3 gave a mark
tryptophan reaction, and no. 4 none.

1905] DEAN—PROTEOLYTIC ENZYMES 331

CucuRBITA MAxIMA.—The seeds of this plant and the young
seedlings with radicles an inch and a half long were found to contain
an ereptase of considerable power as shown by the action of their
tissues on Witte peptone. Under conditions for autolysis none
seemed to take place, since no tryptophan reaction was obtainable,
notwithstanding the fact that the proteids of this seed give a strong
Adamkiewicz reaction. When the seedlings were thirteen days
old, however, they were found capable of autolysis with the formation
of tryptophan. The results show that the seeds contain an ereptase
and that after germination has proceeded for some time a tryptase
also makes its appearance.

Cucursira Prepo.—The seeds of the pumpkin give the same
evidence of the presence of an ereptase as do those of the Hubbard
squash. Two attempts were made to obtain preparations of this
€nzyme. A quantity of ground seeds of the pumpkin were whipped
up in water, and the shells, which floated on the surface, were skim-
med off. After extracting for several hours, the mixture was strained
through gauze and an attempt made to filter it clear. The fluid
appeared to have a quantity of emulsified fat in it, and even the
repeated filtration through pulp filters failed to make any impression
on it. The mixture was allowed to stand two days with toluol
to prevent bacterial action and was then readily filtered, yielding a
perfectly clear filtrate, which however possessed no proteolytic
Power. In the second attempt 452™ of ground seeds were extracted
with sufficient water to make 275°° of milky looking extract which
was found to have a marked action on Witte peptone. As soon
as this extract was strained from the ground seeds it was mixed
with an equal volume of saturated ammonium sulphate solution
and allowed to stand over night. The precipitate was filtered off
the next day, and when tested with Witte peptone was found to be
very active, showing that the enzyme is either precipitated by half
Saturation of its solution with ammonium sulphate, or else it sticks
to the proteid precipitated by the salt. The precipitate was whipped
up in water, transferred to a parchment paper bag, and dialyzed
for two days in running water. When taken from the dialyzer and
filtered clear, the solution was found to be inactive on Witte peptone.
These results show that this enzyme is very sensitive and that some

332 BOTANICAL GAZETTE [way

other means must be resorted to in order to obtain preparations
of it free from the bulk of proteids, etc., in the tissue extracts.

STUDY OF THE PROTEASES OF PHASEOLUS VULGARIS

The beans used in the experiments to be described were of two
sorts, the red kidney bean and the white medium field bean; there
has been nothing to suggest that the proteases in these varieties are
different, and in the records of experiments no mention will be made
of which kind was used.

Resting seeds.—The resting seeds were tested a number of times
for proteases; the results always showed that the seeds contained an
enzyme capable of hydrolyzing Witte peptone with the formation of
tryptophan, but incapable of digesting the phaseolin of the seed.
Later in this paper will be found a more detailed description of this
ereptase. The record of a typical experiment is given here.

Seeds of Phaseolus vulgaris which were known to germinate readily
were ground to a fine meal, one gram of this powder placed in each
of four test tubes, and further additions made as follows:

No. 1—15°¢ H,O +toluol.

No. 2—15°¢ H,O +toluol—boiled.

No. 3—15°° H,0+toluol+o.38™ Witte peptone.

No. 4—15°° H,0+toluol+o.38™ Witte peptone—boiled.

All were corked, shaken, and kept in the incubator at 37-39° C.
for forty-one hours. At the expiration of that time the contents of
each tube were boiled, acidified with acetic acid, and filtered. The
filtrates were then tested. :

No. 1—no biuret, Millon’s, nor tryptophan reactions.
No. 2—not tested.

No. 3—strong tryptophan reaction.

No. 4—no tryptophan reaction.

Seeds soaked in water for twenty-four hours.—Seeds from the
same lot used in the above experiment were soaked in distilled water
for twenty-four hours, the hulls removed, and the cotyledons ground
in a mortar and 1.5®™ of the paste tested in the same way a5 the
dry bean meal, and with the same results.

Seeds soaked jor forty-five hours.—The beans not used in the last
experiment were soaked twenty-one hours longer, peeled, and the
tests repeated, with the same results.

1905] DEAN—PROTEOLYTIC ENZYMES 333

Seeds soaked twenty-jour hours and germinated in sand three days.
—At this stage the hypocotyls were about one inch long. The
cotyledons were removed, ground in a mortar, and the paste tested
as in the previous experiments. The result was the same, the Witte
peptone was attacked and the seed proteids were not.

Seeds soaked twenty-jour hours and germinated six days.—The
cotyledons from the young plants with hypocotyls two to three inches
long were removed and ground to a paste. One gram of this was
placed in each of six test tubes and other additions made as follows:

No. 1—10°¢ H,O + toluol.
No. 2—10°¢ H,O + toluol—boiled.
No. 3—10° H,O+0.1 per cent. of citric acid +toluol.
No. 4—10°¢ H,O+0.1 per cent. of Na,CO, + toluol.
No. 5—10°¢ H,O0+0.2™ Witte peptone +toluol.
No. 6—10°¢ H,O+0.22™ Witte peptone—boiled + toluol.
After nineteen hours in the incubator the contents of each tube were
iled, acidified, filtered, and the filtrates tested with these results:
No. 1—no tryptophan nor biuret reactions, Millon’s ?
No. 2—not tested.
No. 3—no tryptophan reaction, biuret ? Millon’s?
No. 4—no tryptophan reaction, biuret ? Millon’s?
No. 5—strong tryptophan reaction.

No. 6—no tryptophan reaction.

Seeds soaked twenty-four hours and germinated ten days in the dark.
—The cotyledons at this age of the plant were comminuted and tested
in tubes as follows:

No. 1~ground cotyledons + H,0 + toluol.

No. 2—ground cotyledons +H, Ce cies egg albumen.
No. 3—ground cotyledons + H,O + toluol + edestin.

No. 4—ground cotyledons + H,O +toluol+ Witte peptone.

After eighteen hours in the incubator the tests showed that only the
Witte peptone had undergone proteolysis.

Cotyledons of eleven-day old seedlings grown in light.— Beans
allowed to germinate in washed sand, with a good supply of light,
for eleven days had attained a height of five to six inches and had
two well developed leaves besides the shriveled cotyledons. The
cotyledons were found still to contain an active ereptase.

334 BOTANICAL GAZETTE [way

Cotyledons of etiolated seedlings thirteen days old.—These cotyledons
gave convincing evidence of the presence of ereptase, but no tryptase.

Besides testing the cotyledons of germinating beans by the method
of autodigestion, extracts were also prepared with glycerin and 20
per cent. alcohol. The tests applied to these extracts showed that
they were incapable of digesting the bean proteids or boiled fibrin.

It should be noted that OsporNE and Harris‘? have found that
phaseolin, the globulin which makes up the main proteid store of
the resting bean, gives but a faint Adamkiewicz reaction. Therefore
the absence of a tryptophan reaction in the autodigestion experiments
with the bean cotyledons would not be conclusive evidence of the lack
of proteolysis. The use of the Millon’s reaction for tyrosin and
the biuret reaction for albumoses and peptones would have shown
the presence of the products of peptonization or proteolysis had they
been present. Still more conclusive evidence that the enzyme in
question does not act on the unaltered proteids of the cotyledons is
afforded by the more thorough study of the enzyme.

Beans were allowed to germinate for four days and the cotyledons
separated and ground in a mortar to a paste. Forty grams of this
pulp were extracted with 200° of water and 5° of toluol. The
extract was removed from the tissue residue by straining through
fine muslin, and, since phaseolin is soluble in water and the salts
of the seed, this milky looking fluid was rich in proteid matter. The
200°° of extract were divided between two flasks and one of the por
tions boiled. After digesting for seventeen and one-half hours,
a comparative quantitative estimation of the amount of nitrogen in the
proteid decomposition products was made in the following manner.
Fifteen cubic centimeters of each digestion fluid were mixed ina dry
beaker with 1 5°° of tannic acid reagent (7 per cent. of tannic acid
dissolved in 2 per cent. acetic acid) and the precipitates removed by
filtration. Nitrogen determinations were made in duplicate by the
Kjeldahl method on 5° of each filtrate. The amount of standard
acid required to neutralize the ammonia formed by the sulphuric
acid treatment was, in the unboiled digestion, 0.7°°; in the boil
digestion 0.5°°. The digestions were allowed to proceed for two

17 OSBORNE and Harris, Joc. cit.

7
a
|

1905] DEAN—PROTEOLYTIC ENZYMES 335

days longer and the tests repeated. This time the unboiled digestion
required 0.5°° of acid; the boiled digestion required 0.5°°.

This experiment was repeated, using different conditions of
reaction to make sure that such conditions played no part in the
results previously obtained. Fifty grams of cotyledons from seven-
day etiolated bean seedlings were comminuted and an extract pre-
pared as before. This was divided between five flasks as follows:

No. 1— 50°¢ extract.

No. 2— 50°¢ extract—boiled.

aaa. eee
No. 3—50°¢ extract +o. 14®™ citric acid (= ms acid).

N
No. 4— 50° extract +o. 1068™ Na,CO, (= jee
No. 5— 50° extract + 28™ Witte peptone.

Toluol was added in equal amount to all the flasks. The quantitative
estimations were made in the same manner as in the last experiment
except that 10° of each filtrate were used for an analysis instead of
5°. In place of a boiled control for no. 5, a determination was
made at the start of the experiment, when it appeared that 1.4°° of
standard acid were required to neutralize the ammonia formed from
the nitrogen compounds in 10° of the filtrate from the tannic acid
Precipitate. After three days’ digestion quantitative determinations
were made on all.

No. I—required 1 .3°¢ of standard acid.

No. 2—required 1.0°¢ of standard acid.

No. 3—Tequired 1.5°¢ of standard acid.

o. 4—
No. oe analyses lost by breaking of the apparatus.

| Three days later the tests were repeated.

0. I—required 1. 4°¢ standard acid.

No. 5—required 6.5°° standard acid.
The digestion fluid of no. 5 at the close of the experiment gave a

Strong tryptophan reaction. The variations among the digestions,
xcept no. 5, are within o. 3°° of standard acid; in such a method

336 BOTANICAL GAZETTE [way

these differences cannot be held to show the presence of a trypsin-
like enzyme, especially when we see the change in the digestion
containing Witte peptone which at first required 1.4°° of acid for
10°° of filtrate and at the close 6.5°°.

Being convinced by the experiments tried that the resting and
germinating bean contained an ereptase, I attempted to separate
it from the mass of other materials in the seed extracts.

One hundred and fifty grams of bean meal were extracted for
five hours with 375°° of water with the addition of toluol and chloro-
form. The fluid was strained and pressed out through cheese cloth,
the residue mixed with 150°° of water and this likewise removed.
The total extract, amounting to 450°°, was allowed to settle over night
to get rid of the starch, etc. by sedimentation. The next morning
the supernatant fluid was siphoned off and filtered three times through
pulp filters, yielding 365°° of an opalescent fluid. This was half
saturated with ammonium sulphate by adding an equal volume of
saturated ammonium sulphate solution. The flocculent precipitate
A was removed by filtration, and to the filtrate saturated ammonium
sulphate solution was added to make two-thirds saturation. Pre-
cipitate B was likewise removed and ammonium sulphate added to
three-quarters saturation. With this concentration of the salt nothing
but a turbidity resulted; accordingly ammonium sulphate in substance
was added to saturation, yielding the precipitate C. A small portion
of each precipitate was placed in a test tube and mixed with Witte
peptone and water; after standing over night in the incubator the
fluids were tested for tryptophan. The results showed that A con-
tained sufficient ereptase to cause a strong tryptophan reaction, B
a weak one, and C none at all. Each precipitate was dissolved in
water so far as possible, and the turbid solution placed in dialyzing
bags and dialyzed until practically free from sulphates. In A a
considerable amount of phaseolin was precipitated on the sides of the
‘dialysis; this was filtered off, washed with water, and both the clear
filtrate and the globulin tested with Witte peptone. It was found
that the ereptase had remained in solution and had not been carried
down with the precipitated proteid. The solutions of the precipitates
Band C, after dialysis, were found to have respectively a weak action
on Witte peptone and no action at all.

1905] DEAN—PROTEOLYTIC ENZYMES 337

The ereptase solution obtained from A gave a weak biuret reaction
and a faint coagulum on heating. Attempts to precipitate the enzyme
by means of uranium acetate according to Jacoby’s method failed;
no precipitate was obtained except in acid solutions in which the
enzyme was destroyed. The ereptase was tested with crystallized
edestin from hemp seed, crystallized excelsin from the Brazil nut,
phaseolin from the bean, boiled fibrin, and Witte peptone. The
native proteids were all unattacked and the Witte peptone was
actively digested.

An ereptase solution was prepared from 15008" of bean meal by
extraction with water, filtering the extract, half saturating the solution
with ammonium sulphate, and dialyzing the solution obtained from
this half saturation precipitate. A perfectly clear solution resulted
after filtering off the phaseolin precipitated by dialysis, and this
was dried at 4o to 50° C., yielding 2.98™ of a golden yellow powder.
This powder was found to be active in digesting Witte peptone,
although it seemed that the purification and drying had considerably
diminished the power of the enzyme. This protease preparation was
tested with some of the fractions of Witte peptone which had been
separated by Picx’s'® method and with a preparation of phaseolin.
The first experiment was carried out using the secondary protease
preparations without the removal of the ammonium sulphate left
adherent by the method of separation.

One and one-half grams of the enzyme preparation mentioned
above were dissolved as completely as possible in 300°° of water,
35°° of this solution measured into each of five flasks, and 2°° of toluol
added. Proteids were added as follows:

No. 1—18™ of phaseolin.

No. 2—18™ of proto-proteose.

No. 3—18™ of hetero-proteose.

No. 4—18™ of secondary albumose A.
No. 5—18™ of secondary albumose B.

After shaking, 1 5°° were removed from each flask, 15°° of tannic
acid reagent added, and the precipitates filtered off. Analyses of
to** of each filtrate were made in duplicate:

No. 1—required 0.7°¢ of standard acid.
No. 2—required 0.6° of standard acid.
*® Pick, Zeitschr. Physiol.-Chem. 24:246. 1898.

338 BOTANICAL GAZETTE [aay

No. 3—required 0.8°° of standard acid.

No. 4—required 10.3°¢ of standard acid.

No. 5—required 7 .6°° of standard acid.'9
After six days in the incubator the quantitative estimations were
repeated.

No. 1—required 0.6°° standard acid=gain of 0.0°°.

No. 2—required 1.9°° standard acid =gain of 1.3°°.
0. 3—Tequired 1.3°° standard acid=gain of 0.5°°.
No. 4—required 10.9°° standard acid = gain of 0.6°°.
0. 5—required g.2°° standard acid =gain of 1.6°°.

The results in the case of the phaseolin, the proto-proteose, and
the deutero-albumose B are well defined; those with the hetero-
albumose and the deutero-albumose A were not sufficiently marked
to lead to a definite conclusion.

A second experiment was tried with the four proteose preparations
after the ammonium sulphate had been removed from the two deu-
tero-proteoses. In these trials, instead of using the ereptase prepara-
tion used in the previous experiment, an extract was made from some
bean meal and filtered several times through pulp filters until it was
nearly clear. Of each of the four proteoses 0.125%" was dissolved
in 25°° of water. Ten cubic centimeters of the bean extract were
placed in each of eight test tubes and four of the portions boiled.
Ten cubic centimeters of each proteose solution were added to a boiled,
and 10° to an unboiled, portion of the extract. Nine drops of toluol
were added to each tube, the tubes corked, shaken, and kept in the
incubator for sixty-five hours. At the expiration of that time the
contents of all the test tubes were boiled and filtered. Of each
filtrate 10° were removed to a clean test tube, 5°° of a ro per cent.
sodium hydroxide solution added, and then dilute copper sulphate
to the maximum biuret reaction. After standing for several minutes
the strengths of the biuret reactions from the boiled and unboiled
digestions of the different proteoses were compared. The reactions
showed that all of the proteoses were attacked by the enzyme. boas
portions of the filtrates not used in the comparative biuret reactions
were employed to test for tryptophan. The unboiled digestions
with proto-proteose, hetero-proteose, and deutero-proteose A gave

9 The ammonium sulphate in the last two proteose preparations accounts for the
high nitrogen in the filtrates from the tannic acid precipitates.

Z

Z

Ne ee os ee RE en ee Pe ES SY a eT ee er he eee

To ee ee ee eT eee ere ee ee

1905] DEAN—PROTEOLYTIC ENZYMES 339

distinct tryptophan tests, that with deutero-proteose B was very faint;
none of the boiled digestions showed any trace of tryptophan.

In summing up the results of the experiments described above,
we may note that evidence of the presence of a protease has been
obtained in the leaves of the spinach and cabbage, the blossoming
heads of Daucus Carota and the developing seeds of that plant, in
the leaves and unripe seeds of the chestnut, the etiolated seedlings
of Phaseolus Mungo, the seeds and seedlings of Cucurbita maxima,
and the seeds of Cucurbita Pepo. In the case of Phaseolus vulgaris,
experiments with all stages of the germination of the seed have shown
that the cotyledons always contain an enzyme of the erepsin group,
and at no time can any evidence of the presence of an enzyme
capable of attacking the proteids of the seed be obtained. This
€reptase may be removed from the bean extracts by half saturation
of the solution with ammonium sulphate. On dissolving this precipi-
tate in water and dialyzing the solution free from sulphates, the
enzyme is obtained in a solution which is perfectly clear and gives
but feeble reactions for proteids. A preparation of the enzyme may
be obtained by drying this solution at a temperature below 50° C.
The enzyme acts on the proto-proteose, the hetero-proteose, and the
deutero-proteoses, separated from Witte peptone; it is quite inactive
on phaseolin of the bean, excelsin of the Brazil nut, edestin of the
hemp seed, and boiled fibrin.

There can be no doubt but that the large proteid store in bean
seeds is utilized in germination, and in all probability this utilization
is preceded by cleavage to the amido-acids, hexon bases, etc. What
effects this cleavage, and what part, if any, the ereptase plays, are
Unsolved problems. It is likewise unknown whether or not this
®nzyme occurs in other parts of the plant or whether enzymes of the
trypsin type are present in certain tissues.

The investigation of which this is a partial report was carried
Cut with the assistance of a grant from the Carnegie Institution.

LaBoratory oF PLANT PHysIoLocy,

SHEFFIELD SCIENTIFIC SCHOOL,
YALE UNnrversity

DEVELOPMENT OF SPORANGIUM IN BOTRYCHIUM.
CONTRIBUTIONS FROM THE HULL BOTANICAL
LABORATORY. LXXI.

Ira D. CARDIFF
(WITH PLATE IX)

THE present work upon Botrychium was taken up with the pur-
pose of investigating what was thought by some to be septation
caused by sterilization of mother-cells in the sporangium of B. ternaium.
Both B. ternatum and B. virginianum were examined, and the two
species were found to be essentially the same in their sporangial
development. The material of B. virginianum was collected near
Woodville, Indiana, March 14, 1904; that of B. ternatum was col-
lected near Sullivan, Ohio.

The early stages in the development of the Botrychium sporangium
were studied by BowER (1), CAMPBELL (2), and GOEBEL (5), but
apparently none of them studied the interesting later stages. In
the present work, though a number of preparations were examined,
no special attention was given to the early development further than
to confirm the studies of the above mentioned investigators, namely,
that the sporogenous mass originates from a single hypodermal
archesporial cell. CAMPBELL (2) says that the later divisions in the
archesporium do not follow any definite rule, but take place irregt-
larly. This does not accord with my observations, the divisions in
B. ternatum and B. virginianum taking place with great regularity j
CAMPBELL implies that his fig. 129, C, which shows no regularity 1n
cell-arrangement, is in the mother-cell stage. If this is true, it indicates
an unusually small output for a Botrychium. Horrzman (6) made
some observations upon the later development of the sporangium of
Botrychium, but Bower (1) thinks that the sequence of segmenta
tions, as shown in Horzman’s figs. 3-6, is not sufficiently intelli
gible, and should be investigated afresh. He thinks that Holt?
MAN’S description suggests a mode of segmentation more clearly
analogous to that in leptosporangiate ferns.

[May
340

* 1905] CARDIFF—SPORANGIUM IN BOTRYCHIUM 341

In the sporangium of Lycopodium, the sporogenous mass _ is
divided into blocks of cells, each block apparently being the descend-
ant of a single archesporial cell. The blocking here is evidently
related to a multicellular archesporium. In the Botrychium spor-
angium there occurs a blocking of the sporogenous mass that must
have a different cause, as the archesporium is unicellular. The single
hypodermal archesporial cell divides usually anticlinally, then peri-
clinally (fig. r). Regular divisions then follow, as shown by jig. 2.
At the stage shown in fig. 2 the wall is six to eight cells in
thickness, including the tapetum; and the extremely glandular char-
acter of the two inner layers of cells indicates that they are definitely
set apart as tapetum. In the tapetum at this stage walls are formed
both periclinally and anticlinally. The divisions in the sporogenous
Mass occur with remarkable regularity. The position of the original
walls, from stages shown in jigs. r and 2, are still perfectly apparent
up to the spore mother-cell stage (jig. 3). At this stage the sporog-
€nous mass has somewhat the appearance of the spermatogenous
mass in a bryophytic antheridium, except for the size of the cells and
the character of the nuclei. By the time the mother-cell stage is
reached, the tapetum has become four or five cells thick, is quite
glandular in appearance, and there is no further evidence of mitotic
division. The sporangium wall-cells adjacent to the tapetum have
commenced to collapse as a result of the drain upon them by the
tapetum.

Probably the most common and most primitive method of nour-
ishment of mother-cells is by abortion and absorption of a portion of
the mother-cells to form a diffuse tapetum, as in Equisetum. Another
method is by the formation of sterilized tracts from potentially sporog-
€nous tissue through which material may be conducted to the interior
of the sporangium, as the trabeculae of Isoetes or the septations in
the microsporangia of Lemna minor.

In the sporangium of Botrychium, no sterilization of either kind
was found, every mother-cell functioning. So far as I was able to
determine, division up to the mother-cell stage is simultaneous
throughout the sporogenous mass, yet the original blocks (jig. 3)
of sporogenous cells still remain perfectly distinct. As the mother-
cell enters upon the synapsis stage, the original walls or wall separating

342 BOTANICAL GAZETTE [MAY

the blocks is apparently thicker, and by the time the spirem is
formed the blocks have begun to separate. This separation takes
place in the order in which the original walls were laid down in the
archesporium and young sporogenous mass (jigs. 3, 4, 6, 7). At
about prophase the mass has separated into at least sixteen (in section)
distinct and separated masses (fig. 6). ‘‘About” prophase is used,
since with the separation of the blocks differences in stages of division
begin to appear, so that the cells throughout the entire sporogenous
mass are not in the same stage, though those in the same block are
always in the same stage.

The progressive separation into smaller blocks continues in the
same manner in which the first blocks were formed. It takes place
along the same lines and in the same order in which the earlier walls
were laid down. Whatever stimulus caused the simultaneity of divis-
ion during the early life of the sporangium seems to have been
interfered with here by the cleavage into disjoined blocks,
for in the same sporangium the blocks are in different stages of
division, it being quite common to find four or five stages. For
example, in three adjacent blocks, the cells of one were in early meta-
phase, of another in telophase, and of the third in anaphase. In
another case blocks were found in the same sporangium varying from
metaphase of first to metaphase of second division; and it is very
common to find them varying from metaphase of first to prophase of
second division. One sporangium was found in which all the cells
of one-half the entire sporogenous mass were in metaphase, while
those of the other half were in telophase. All of the sporogenous
tissue throughout its entire development appears in a vigorous and
perfectly normal condition.

What may be the cause of this retention of their individuality by
the developing sporogenous cells is difficult to say. In regard to bryo-
phytic antheridia this phenomenon has been explained as due to the
independent development of the original spermatogenous cells; and
there is no further blocking of the spermatogenous mass after the
periclinal walls which cut off this mass from the antheridial wall
layer have been formed. The difference in rate of development of
these blocks has been attributed to differences in food supply, oF '°
some purely physiological cause. It seems that as much might be

SARE ne aN ge em ee ae

1905] CARDIFF—SPORANGIUM IN BOTRYCHIUM 343

said of the blocking in a Botrychium sporangium; though each divis-
ion, from the very first almost to the formation of the mother-cell,
separates masses which retain their individuality throughout their
further development. The rate of growth, however, and apparently
the food fupply are absolutely the same for all the blocks up to the
mother-cell stage. It would seem that in such large sporangia some
blocks or cells would be more favorably located than others with
reference to food supply or conditions of growth, and there would
be greater growth on the part of some regions than others, thus
causing irregularities in the arrangement of cells, such as is found in
most sporangia, instead of the very regular arrangement in those of
Botrychium. Therefore, the above mentioned physiological explana-
tion in regard to bryophytic antheridia is not entirely satisfactory.

What may be the exact cause of this progressive separation of the
sporogenous mass along lines where the earlier walls were laid down
is impossible at present to say. To me, the most reasonable explana-
tion which can be offered is that the middle lamellae of the walls are
acted upon by an enzyme at a time when the sporogenous mass is
greatly in need of food. As the lamellae grow older their composi-
tion may change, so that they are more easily digested than those
more recently formed, thus effecting a progressive separation of the
sporogenous tissue. As the blocks become separated, and wholly
or partially surrounded by the tapetum, some blocks will of necessity
be under slightly different conditions of osmotic pressure or chemical
stimulation than others, thus bringing about differences in the rate
of their development.

In Botrychium the tapetum is derived from the wall, and absolutely
no contribution is made to it from the sporogenous tissue. By the
time the sporogenous mass has reached as much as sixty-four cells,
the tapetum is clearly delimited from the sporangial wall and is two
cells in thickness. From this time on periclinal divisions take place
rapidly until the spore mother-cell stage is reached, when the tapetum
is four or five cells in thickness and very glandular in appearance.
At the first separation of the blocks (figs. 3 and 4) in the sporogenous
mass—about the synapsis stage—the cell walls of the inner layer of
tapetal cells begin to disintegrate. Some of these inner cells are at
this time binucleate, while the nuclei of the next one or two layers

344 BOTANICAL GAZETTE [MAY

of cells are in process of division amitotically, and can be found in
all stages. All the tapetal nuclei have now increased much in size,
those of the young tapetal cells being 8# in diameter, and those of
the later stage 15 to 20H. As the blocks of sporogenous cells con-
tinue to be formed and more widely separated, the tapetum com-
mences to grow rapidly. The number of nuclei increases greatly,
as well as the volume of the cytoplasmic mass in which they float.
This pushes inward between the blocks with quite a regular outline
(fig. 6). At prophase of the first division of the mother-cell, a section
of the sporangium shows that these tapetal plates have extended almost
across the sporogenous mass between the first formed blocks, and have
commenced to grow inward between the blocks of later formation
(fig. 6). This rapid tapetal growth continues pushing thinner and
thinner plates between the smaller blocks as they are formed (jigs.
7,8, 9), making a network which finally invests the individual tetrads,
or groups of two or four tetrads (fig. g); and at last the spores separate
and float in this tapetal mass. The original thicker plates of tapetum
may often be found after the spores are completely formed.

As this excessive tapetal growth takes place, the cell walls of the
original tapetum break down successively from the inner layers
outward, until at metaphase of the mother-cell the walls of only the
outermost layer of tapetal cells remain. By the time anaphase of
second division (fig. 8) is reached, the last walls of the tapetal cells
have entirely disappeared, and the inner cells of the sporangial wall
begin to take on tapetal characters; especially is this true of the nuclei
which resemble tapetal nuclei very closely (jig. 9). The cell walls,
while they collapse, have not been found to disintegrate entirely, as
in the case of the true tapetum. In fact, there is no reason why these
inner layers of sporangial wall might not be called tapetum.

Probably the most interesting feature of the development of the
Botrychium sporangium is the unusual growth of the tapetum.
before mentioned, it increases greatly in volume and in number of
nuclei, yet not a wall is formed anywhere, though it was stained
especially for walls. The nuclei are found in all stages of amitotic
division (figs. 5, 8, 9), but no evidence of mitosis is found. They
take stains strongly, are exceedingly large, as mentioned above, and
have an unusually thick nuclear membrane. That the nucleus

1905] CARDIFF—SPORANGIUM IN BOTRYCHIUM 345

bears an important relation to-the metabolic processes of the cell
is too well recognized to need discussion, and. this enlargement of
nuclei, or increase in nuclear surface, is undoubtedly in response to
the increased demand for nourishment on the part of the sporogenous
tissue at this stage.

It has been noted frequently that the ovarian follicle cells of
arthropods have many large nuclei which divide amitotically, and
this has been explained by FLEmminc (4) and Cun (3) as being a
means of securing more rapid metabolic processes between cytoplasm
and nucleus through the use of a larger nuclear surface. Our reliable
knowledge of the process of amitosis is so meager that one can at
this time venture only a tentative explanation of the nuclear behavior
in the Botrychium tapetum. Whether amitosis can take place more
rapidly than mitosis is unknown, but from the mechanics of the two
processes, it would seem that the former would be much the more
rapid. WILsoN (9) considers that all nuclear division is the response
to particular stimuli, and is probably incited by local chemical
changes, an idea which is confirmed by Prerrer (8) and NATHAN-
SOHN (7), who were able to produce mitosis or amitosis at will in
Spirogyra orbicularis. May we not then look upon amitosis in the
tapetum as simply an acquired character due to the unusual demand
upon it by the fertile tissue for nourishment at a particular period
in its development? Walls being unnecessary, the energy of the
Organism would not be used in forming them.

As the spores commence to separate in the tetrad, the tapetal
cytoplasm has entirely filled the sporangium and many of the nuclei
have begun to disorganize, though they seem unusually persistent
and many are found after the tetrad is fully formed. Later, when
the spores are entirely separated and mature, the tapetum disappears.

Thus we have here worked out the problem of nourishment in
a large sporangium by a method entirely different from the two
formerly. mentioned, namely by the preservation of the individuality
of sporogenous cells, thus enabling the mother-cell mass to separate
easily into regular blocks, and leaving straight open passageways
through which a non-sporogenous tapetum grows rapidly, furnishing
the required nourishment for the developing spores. As a general
tule there are more spores provided for in a sporangium than nutri-

|
|

346 BOTANICAL GAZETTE [May

tive conditions will allow, and there are usually two methods by
which this difficulty is overcome: by abortion of mother-cells, and by
arrangement for a succession of sporangia according to nutritive
supply. In Botrychium, however, there is no abortion of mother-
cells and very little difference in the stages of development of the
different sporangia in a spike. The nutritive supply is equal to the
demand of all mother-cells, probably owing, in part, to the slow
growth of the plant, and also to the large amount of food material
stored in the stem.
SUMMARY.

1. The sporogenous tissue develops from a single hypodermal
archesporial cell.

2. As the successive sporogenous cells are formed, each retains
its individuality throughout the development of the sporogenous tissue.

3. Divisions in the sporogenous tissue are simultaneous up to
the mother-cell stage.

4. Beginning with the mother-cell stage, the sporogenous mass
separates successively into blocks of cells in the same order in which
the earlier cells were formed. :

5. The blocks develop independently, and at a different rate in
the same sporangium, though all cells of one block develop at the
same rate. Whatever stimulus caused the simultaneity of division
in early sporogenous tissue, is interfered with by separation of the
cells into disjoined groups.

6. The progressive separation of thc sporogenous mass is probably
caused by the digestion of the middle lamellae.

7- All mother-cells produce spores.

8. The tapetum is of non-sporogenous origin.

9. With the separation of mother-cell groups, the tapetum grows
rapidly between them without the formation of walls; the nucle!
increasing greatly in size, and dividing amitotically.

10. The problem of nourishment in large sporangia may thus be
solved by individual development of sporogenous cells, by their later
separation into regular groups, and the rapid growth of tapetum
between them.

11. The nuclei of the old tapetum are four times the size of those
in younger stages of its development.

Pee ee

PLATE 1X

_ BOTANICAL GAZETTE, XXXIX

a.
=
~
P a
&
=
°
a
z
tu
5
4
3

1905] CARDIFF—SPORANGIUM IN BOTRYCHIUM 347

12. Amitotic division and increase in size of nuclei are both
devices for rapidly increasing nuclear surface, thus effecting a larger
increase of metabolic products.

I am indebted to Professor Joun M. Courter and Dr. CHARLES
J. CHAMBERLAIN for criticism and advice.
CoLumBra UNIvERsIry.
LITERATURE CITED.
1. Bower, F. O. Studies in the morphology of spore-producing members. II.
P. 37. 1896.
- CAMPBELL, D. H., Mosses and ferns 250. 1895.
3. Can, C., Uber he Bedeutung der direkten Zelltheilung. Sitzb. Schr. Physik-
Okon. Ges. Kénigsberg. 1890
4. Fremuine, W., Entwicklung und porte der Kenntnisse iiber Amitose. Merkel
und Bonnet’s Ergebnisse. IT.
- GoEBEL, K. , Beitrige zur rE Entwickelungsgeschichte der Sporan-
jen Jot. Zeit. 38:545. 1880.
; Hortzman, C. L., On the apical growth of the stem and the eea RS of
— nj Botrychium virginianum. Bot. GAZETTE 1'7:21
7. NATHANSO Physiologische Untersuchungen tiber cies Kern-
hctane. pu b. Wiss. Bot. 35: 48-79. 1900.
PFEFFER, W., Ueber die Erzeugung und die Stee cab Bedeutung der
Amitose. Verhandl. Kénigl. Siichs Akad. Wiss. 53:4-12. 1899.
9. Witson, E. B., The cell in development and inheritance 391. 1902.
EXPLANATION OF PLATE Ix
All figures were made — Zeiss nia - oculars and Bausch Lomb
camera lucida. The origina half in reproduction.
IG. 1. Young spovangiam. x (en:
IG. 2. Sporogenous tissue of thirty-two cells; gas two cells thick and
visions taking place both periclinally and anticlinally. % goo.
G. 3. Sporogenous tissue in mother-cell stage showing comes’ in cell
rangement the two original walls, a and 6, beginning to separate. X 400.
lata showing sporogenous tissue separating re “blocks along
me pee walls. X 150.
Fic. . Portion e fig. 4; mother-cells in synapsis; commencement of tapetal
X goo.

tS

a MK

9

gtowth.
Fic. 6. Sporangium showing sporogenous tissue separating along original
ood shown i in fig. 2; plates of tapetum extending between earlier formed blocks.

7 7. Sporangium showing cosine tissue separated into blocks in
each of which the cell division is simultaneo

Fic. 8. Portion of sporangium showing ane : the forming tetrads,
also the large increase in the number of tapetal nuclei. ;

Fic. 9. Later stage than shown in fig. 8. X goo.

PHYSIOLOGICAL PROPERTIES OF BOG WATER.
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY.

BurRTON EDWARD LIVINGSTON.
(WITH THREE FIGURES)

ALTHOUGH it has been shown (1) that water from the bogs of the
northern United States contains solutes to such a very small amount
that its osmotic pressure is generally not appreciably above that of
the river swamps and lakes of the same region, still the generally
observed xerophilous character of bog vegetation may be due to small
amounts of dissolved substances of such nature that they affect the
plants chemically through toxic stimulation. Having found that
metallic ions affect the vegetative growth of the polymorphic
Stigeoclonium with which the author has been experimenting for
some time (2), and that the effect thus produced is identical with
- the response of the alga to high osmotic pressures, it was suggested
that this alga might be used as an indicator in a study of the physi-
ological properties of bog waters. In accordance with this suggestion,
natural waters of a number of different types were collected in bottles,
filtered through filter paper, and tested as culture media for the
alga. The result of these tests is, briefly, that many bog waters act
upon the plant like poisoned solutions. Details of the work are
given in the following pages.

The form of Stigeoclonium here used has already been shown
(2-6) to take either of two forms according to the medium in which
it is grown. In solutions of low osmotic pressure at ordinary tem-
peratures it assumes the form of branching filaments composed of
cylindrical cells. In the same solutions at a temperature slightly
above the freezing point of water, and at ordinary temperatures 1)
solutions similar to these but poisoned with certain metallic salts, as
well as at ordinary temperatures in solutions of high osmotic pressure;
the plant takes the palmella form, in which the cells are spherical or
nearly so and lie in the medium singly or in irregular groups- If

348 [may

1905] LIVINGSTON—PROPERTIES OF BOG WATER 349

filaments are taken from the conditions which favor their growth
and are placed in those favoring the palmella form, their cells round
off and partially or completely separate, they begin to divide slowly
in all planes, and the result is the typical form for these conditions.
A return to the conditions for filaments is followed by the resumption
of that form, partly by the growth of filaments directly from the pal-
-mella masses, and partly by the production of zoospores which germi-
nate to form new filaments. Zoospores are not produced in solutions
of high pressure, and they fail to germinate in solutions which produce
the palmella form, though they are produced in the cold and in
Poisoned solutions.

The waters tested in these experiments are in large part the same
as were used in the determination of the osmotic pressure (1). The
work was carried on partly at the New York Botanical Garden
and partly at the University of Chicago, and extended from June
1902 to January 1905. The cultures were made in the manner
described for water cultures of this plant in the author’s earlier
papers.

Since a number of authors have loosely attributed SCHIMPER’S

“physiological dryness” (7) of bogs to acidity, titrations of most of
the natural waters here employed were made with n/100 KOH
solution, using phenolphthalein as indicator.”

Data from the experiments are given in the following table. In
the column of responses, F—F denotes that the filamentous form
persisted as such when placed in this particular water. F—P denotes
that the original filaments became palmella and that no new filaments
were produced. F—4P denotes that filaments persisted, but that
there was also a marked production of palmella. A number of cul-
tures were made with each of the waters, and the result is the general
one for all. The cultures were continued for two to four weeks.
The acidity data are given in terms of normal acid and the pressure
in terms of millimeters of mercury at 25°C.

From the table it is seen at once that in some of the waters the
palmella form was produced; in others it was produced in some
— but filaments persisted; and in still others the filamentous

the indicators at hand this was the best, although it is —— not per-
fect ‘3 oh acids as are probably present in these waters.

350 BOTANICAL GAZETTE

DATA FROM THE EXPERIMENTS

[MAY

Response Osmotic Acidity of
Source of water and nature of vegetation of pressure water
Stigeo- of water, in terms of
clonium | mm.of Hg| normal acid
Drained swamps of Hackensack River, N. J. (river swamps)—
Average of six samples F—4P 50 0.00192
Maximum of six samples.........-+-+-seeeeeeeeeeeeeeeeee| teers os 0.0042
Ce or gt i) 2 a en eee earn rr aoe ua 0.001
New York City supply, Croton and Bronx Rivers | ge 50 0.0005
Chicago City supply, Lake Michigan. ...... 205 <sic0-eccweorence F—F 100 Always alkaline,
: (see fig. 1) about 0.0015
Grand River; Grand Rapide; Michi ic. od cca cee ceccnesieg nor EF—F 100 As last, or more
ne
Aetna, Ind. (Rhus vernix, ra)—

Average of three samples pressed from black peat.......-...- F—4P feo, =| *hssmrenee
Miller me , Rhus vernix)—

Average of two samples pressed from black pea F—4P 50 0.0003
Oconomowoc, Wis.* (typical Larix swamp), ditch............. F—P 200 0.0004
Stewart ~— e, Ill., average of three samples pressed from black ce

— Iso 0.0002
Ann Arbor, Mich} ;

First Sister Lake bog (Larix, Chamaedaphne, Sphagnum)—

Sample A F—P 50 0.0033
72.2
ised e aoe. Chamaedaphne, Potentilla palus-
Sample A PP 100 0.0026
Sample P—P 150 0.0024
1 R: , (see fg. 3)
Tom’s River, N, J. (Chamaecyparis, Sphagnum, Chamaedaphne,
Sarracenia, Oxycoccus)—

Sample \, pressed from Sphagnum F—F 170 0.0004
Sample B, beneath Sphagnum ep 140 0.00048
Sample C, pressed from black peat rp go 0.0003
Sample D, margin of pond Fop 40 0.0003

Ree ee Say eee

, , N.Y. (Alnus, Eri , Spha,

Sample A, edge of pond ( eee, Semekeae:) F—P 0.0015
Sample B, decayed leaves F—}P 110 0.0022
Sample C, ditch F—F 90 °
Sample D, pressed from Sphagnum F— 100 -OO1

* This water was obtained for me by Dr. H. C. Cow es of this laboratory.
+ The samples from Ann Arbor weré obtained for me by Dr. H. N. TRaNsEAv, of Alma College, Mich.

form persisted without the production of palmella at all. The char-
acteristic forms of this plant in three different waters are shown 1n
jigs. 1,2, and 3, which are to be compared with previously published

figures, and are self-explanatory.

As has been shown in the case of many inorganic poisons (5)»
the production of the palmella form is sometimes accompanied in
these waters by a stimulation of zoospore production. Usually,
however, the swamp water acts more like low temperatures, producing
the vegetative response without either accelerating or retarding the

reproductive activity.

The palmella response in certain of these waters may

be due,

1905] LIVINGSTON—PROPERTIES OF BOG WATER 351

a priori, to either of two sets of factors, the osmotic pressure of
the solution or the chemical nature of the solutes. Since the experi-

ments were carried on at room temperature, it is unnecessary to con-

sider low temperature as a possible stimulus.

In the work on the influence of osmotic pressure upon this plant

it was found (2, 3) that there is no tendency to form palmella till a

pressure of about 1618.6™™ of mercury has been attained. Filaments

still persist at a pressure of 3237.1™™, but have practically all dis-

appeared at a pressure of 6474.2™™. But no swamp water studied

has a pressure at all approaching the lower limit for even the incipient

Fic. 1. Stigeoclonium filaments in Lake Michigan water.

Fic. 2. Stigeoclonium, palmella form from filaments, in water from West Lake,
Ann Arbor, Michigan, sample A.

Fic. 3. As fig. 2, but in water from First Sister Lake, Ann Arbor, sample B.
formation of palmella. Therefore, we are forced to the conclusion
that the palmella response in these bog waters is due to the chemical
nature of the solutes. The plant grows well as filaments for a long
time in distilled water, so that it is impossible to relate the reasons .
to absence of inorganic salts.

Definite knowledge of the chemistry of bog water is beyond our
reach at present. It appears that all bogs are acid to some degree,
and there has been a tendency for many authors, e. ., SCHIMPER
(7, Pp. 4, 8, 657, etc.), to attribute the peculiarities of bog plants to
this Property. In the author’s study of chemical stimulation (5) it
was found that nitric and sulfuric acids produce palmella at concen-
trations of from o.coorm to o.c0006n. The natural swamp waters are
uniformly more acid than this; therefore, were the acid property
of the latter due to either of these mineral acids, we should expect
all of these waters to produce the palmella response. This is obviously

352 BOTANICAL GAZETTE [way

not the case, and so it seems highly probable at least that the stimu-
lating factor of bog waters is not the hydrogen ion.

Further, a comparison of the acidity figures with the response of
the plant in the different waters shows clearly that the former data
could not be used as a criterion for the prediction of the latter. This
is clearly brought out in the following list, in which the acidity figures
are arranged in order of their magnitude, with the responses occurring
in the corresponding water placed after each. Alkalinity 0.0015”, F;
acidity 0.0002”, F; 0.0003n, F, $P,P; o.cco4n, F,P; 0.00048”,
P; 0.0005”, F; 0.0008n, F; 0.0o1m, F; 0.0015”, P; o.co1gn, $P;
0.0022n, 4P,P; 0.0026n, P; 0.00332, P; 0.0038”, P. The lower
acidities appear to produce both filaments and palmella, the

igher ones only palmella. This would seem to indicate that, while

high acidity is always accompanied by the presence of the stimulating

substances, these substances are not necessarily accompanied by
high acidity.

Boiling the stimulating waters for five or ten minutes and then
rediluting to the original volume with redistilled water decreases their
acidity from 30 to 50 per cent., but appears not to alter their stim-
ulating power. Apparently the active substances are not volatile
at 100° C. Diluting the Ann Arbor samples, and also those from
Tom’s River numbered 2, 3, and 4, with distilled water or with weak
nutrient solution, decreases the toxic effect, and this effect practically
disappears when the water has been diluted to twice its volume.
This is evidence that the stimulating substances are present in
extremely small amount.

The relation of the source of these waters and the type of vege-
tation growing therein to the physiological properties exhibited
toward Stigeoclonium throws some light on the general problem of
the xerophilous character of bog plants. The drained swamps of
the Hackensack valley are not in any sense bogs. In many places,
however, are found spots where Sphagnum has taken a foothold in
small pools. Eriophorum, Typha, and some other semi-xerophilous
plants are also found here. The water samples studied were taken
from such places, and the experiments show that they possess the
toxic property to a considerable degree. The data for this broad
area of swamps are averaged from a large number of samples taken
near Englewood and Closter, N. J., as well as from the western slope

SE ER Te ited iin ee

a a a A a a a a a a At

‘

1905] LIVINGSTON—PROPERTIES OF BOG WATER 353

above the palisades of the Hudson, where the water is held in the
irregular rock basins.

The New York City water, although quite acid, is not active upon
the alga. This water is from streams with well-drained river swamp
margins. Lake Michigan water and the water of Grand River
appear to be identical. Both are somewhat alkaline; neither has
any physiological effect upon Stigeoclonium.

The Aetna and Miller swamps are practically alike; both are
composed of black peat with no Sphagnum. The general nature of
these Swamps suggests an intermediate condition between bog and
river swamp, leaning toward the former, and the water shows a
marked tendency to produce palmella.

The bog near Oconomowoc has Larix, Sphagnum, Vaccinium
corymbosum, Oxycoccus, Chamaedaphne, Sarracenia, etc. Its water
has a very marked action upon the alga.

The Stewart Ridge swamp is a peat deposit but not a true bog.
A few patches of Sphagnum seem to show a tendency in this direction,
but the samples tested showed no effect upon the indicator plant.

The swamps of Ann Arbor are as typical and characteristic bogs
as the author has seen. The character of their vegetation, consisting
of Larix, Drosera, Sarracenia, Andromeda, Chamaedaphne, Arethusa,
Calopogon, etc., agrees well with the fact that the water is markedly
toxic toward the alga.

The vegetation upon all the vast stretches of lowland about Tom’s
River is practically alike; these are dense Chamaecyparis swamps,
abounding in Oxycoccus, Sphagnum, Sarracenia, Chamaedaphne,
etc. All the samples from here, with the exception of one pressed
from living moss, produced the palmella response. It appears that
the active bodies are more plentiful in the mass of dead material
beneath the Sphagnum than in the moss itself.

The Richmond swamp is not a true bog, and yet it contains
considerable amounts of loosely-growing Sphagnum, together with
Eriophorum, Typha, etc. Here there appears a disagreement between
the physiological properties of water samples from different parts.
That from the ditch should be the most dilute, and shows no action
Upon the indicator. That from moss is again harmless; while that
Pressed from decayed Jeaves near by shows a marked toxic effect.
The pond sample was taken up within a centimeter or two of sub-

354 BOTANICAL GAZETTE [May

merged powdery peat, and should be saturated or nearly so with any
slightly soluble substances contained therein. Its active property is
very marked.

From the last eight paragraphs it seems clear that the stimulating
substances with which we have been dealing are present in swamp
waters to an extent roughly proportional to the xerophilous character
exhibited by the swamp vegetation. It is possible that the factor in
such bogs which prevents the growth of plants other than xerophilous
ones may be these unknown toxic bodies. They act upon Stigeo-
clonium in much the same manner as do drying media. Perhaps
ordinary plants are affected by these substances with the same end
result as though they were in a truly dry soil. If this be true it becomes
easy to see how plants whose protoplasm is naturally adapted to dry
situations may alone be able to thrive in these bogs.

The behavior of this alga toward dryness, cold, and bog water
are quite parallel with results obtained by TRANSEAU (8) with Rumex
acetosella. This author found that in dry mineral soil, Rumex pro-
duces thickened leaves reduced in size and with revolute margins,
while the palisade tissue is very much increased in amount, and the
epidermal cells are reduced in size and have thick, cuticularized
outer walls. These changes give the plant, which in moist conditions
is anything but a xerophyte, a very characteristic xerophilous struct-
ure. The same responses are exhibited, to a somewhat less degree,
when the roots are kept at a low temperature, and also when the
substratum is a bog soil. But when both the last named conditions
are allowed to act together the response is the same in direction and
amount as in dry mineral soils. The changes occurring in these
leaves are very similar in their nature to those just described for
Stigeoclonium. From an ecological standpoint, the palmella form of
the alga is extremely xerophilous in character, while the filamentous
form lies at the other extreme. It appears that we have here vee
very widely different plants, both of which respond to these various
conditions in the same way.

SUMMARY.

The results of this series of experiments are as follows:

1. There are chemical substances, in at least some bog waters,
which affect Stigeoclonium as do poisoned solutions and solutions
of high osmotic pressure.

1905] LIVINGSTON—PROPERTIES OF BOG WATER 355

2. The responses of this alga to bog water and to cold are as nearly
identical with those obtained by TraNnsEAu with Rumex, for the
same conditions, as the nature of the two plants would permit.

3. The active Se neeauEte are not directly related to the acidity
of the water.

4. Boiling the water decreases its acidity but does not appreciably
affect its action as a stimulating agent.

5- The stimulating substances are most markedly present in water
from those swamps whose vegetation is most definitely of the bog
type. They are absent from river swamps and large lakes; in water
from swamps whose vegetation is of a character intermediate between
those of the river swamp and the bog, they are present to some degree,
their amount being roughly proportional to the extent of the xerophi-
lous character of the vegetation.

6. The stimulating substances here demonstrated may play an
important réle in the inhibition from bogs of plants other than those
of xerophilous habit.

THE UNIversity oF CHICAGO.

LITERATURE CITED.

I. eee B. E., Physical properties of bog water. Bot. GAZETTE 37: 383-
385. 1904.

, On the nature of the mua which causes the change of form in

polymorphic green algae. Bor. GAZETTE 30:289-317. 1900

, Further notes on the elbeii of polymorphism in green algae.

Bor. GAZETTE 32:292- gol.

, The réle of diffusion we osmotic pressure in plants. Chicago. 1903.
The last chapter of this monograph was reprinted as “The effect of the
osmotic pressure of the medium upon the growth and reproduction of
organisms.” Chica

5- ———,, Chemical euitae of green algae. Bull. Torr. Bot. Club 32:1-34.
figs. 17. 1995.

, Notes on the physiology of Stigeoclonium. Bor. GAZETTE 39:297-
300. figs. 3. 1905.

7. Scurprr, A. F. W. Plant geography upon a physiological basis. Translated
by W. R. FisHEer. Oxford. 1903.

8. TransEav, E. N., On the development of palisade tissue and resinous deposits
in leaves. Scienan Bt S. 19:866-867. 1904.

AN APPARATUS FOR OBSERVING THE TRANSPIRATION
STREAM.

Otto V. DARBISHIRE,
(WITH TWO FIGURES)

THERE are, roughly speaking, two sets of experiments by means
of which we can investigate the presence of the transpiration stream
in small plants. One of these is concerned with the root of the plant
and with the demonstration of the phenomenon known as root pres-
sure. The other set deals with the shoot and the suction exerted
by it on the water which is being absorbed by the root.

We all know what happens when we fix some kind of root pressure
apparatus to the stump of a small plant, like a fuchsia for example.
Water will soon be pressed out of the stump with sufficient force to
raise a column of mercury to a considerable height. We can at the
same time attach to the shoot of the very plant used for the root
pressure experiment some form of potometer. We will observe
suction and a column of mercury can be raised to a considerable
height thereby. Do these two separate experiments really—even
approximately—show what was going on in the plant at the time
the experiment was set up? I think the answer must be no. When
the plant in question was cut across, the stump exhibited the phenom-
enon of pressure, but the shoot that of suction at the same level.
It is obviously impossible that pressure and suction should be exhib-
ited by an intact plant at the same level. It must be mentioned here,
however, that if the two experiments are set up very quickly, we may
find the stump exhibiting suction for a short time. This changes
to pressure when the stump is saturated with water. The root may
of course become saturated in a few minutes by simply being exposed
to water during the setting up of the experiment. We do not
therefore get a clear idea of what is going on in the plant, when we
isolate the shoot from the root, by attaching to each one a separate
apparatus. r

In order to learn more about the relation between shoot suction
and root pressure it is obviously necessary not to separate the cea

356 .

1905] DARBISHIRE—APPARATUS FOR TRANSPIRATION STREAM 357

portions of the plant entirely, but rather to keep them connected
by some continuous bit of apparatus, so that, although the plant
be cut in two, the shoot suction may still act on the root or the root
pressure on the shoot. I use for this purpose a simple bit of glass
apparatus, which I have called a pinometer. It will be necessary
first to describe the apparatus; secondly, the way it is set up; and
finally, to refer to a few of the experiments and the results
obtained.

The pinometer consists of a straight bit of glass tubing (jig. 1, b-d),
to which is obliquely attached by annealing on one side another short
bit of glass tubing (c-/). On the opposite side a U-tube with an
oblique connection is annealed on (a-e.) We have therefore four
Openings to this part of the apparatus (a, 6, c, d). The bore of the
glass tubing used for the pinometer depends entirely on the thick-
ness of the stem of the plant used. The diameter of the glass tubes
should about equal the diameter of the latter. A very convenient
plant to use for demonstration purposes is a healthy fuchsia plant,
not exceeding two feet in height.

The experiment should not be set up till all the various parts
and tools are quite ready. The glass tubing should be cleaned to
Temove any greasiness, which is often the cause of introducing minute
air bubbles into the system of tubes. The air should also be care-
fully removed from the rubber tubing. The more quickly every-
thing is set up, the more nearly will the results obtained show what
Was going on in the plant at the time the experiment was made.

When everything is ready, the plant in its flowerpot is put into
a bucket of water, so that it is immersed to a few inches above the
point at which it is to be cut across. The leaves should not be mois-
tened more than is absoluely necessary. The stem of the plant is
Now cut across under water in such a way that there is about one
inch of stem, above and below the cut, devoid of buds or branches.
Tf the stem has already a complete wood cylinder, the cortex may
be removed with a sharp knife for about half an inch above the cut
on the shoot, and below on the root.

The lower end of the shoot, without removal from the water,
is fixed by rubber tubing to the opening a, jig. I, and the portion a-e
of the pinometer remains full of water, even when removed from the

358 BOTANICAL GAZETTE [MAY

water, and can be temporarily held by a clamp (7) on a retort stand."
Next some rubber tubing is slipped over the upper end of the root
stump. This bit of tubing also fills with water and the whole
flowerpot can now be removed. The end 6 of the pinometer is now
quickly fixed to the rubber tubing over the stump. A gauge is
securely attached to c, and water is poured into d from a small
reservoir (jig. 3, r) till the whole system of tubes is full of water.
Mercury is then poured into the outer limb of the gauge, and this
causes water to pour out at d, to which some pressure tubing has
been firmly fixed. When there is enough mercury in the gauge to
give the columns sufficient play to rise and fall, the opening at d
should be closed by a pinch-cock and the experiment is set up. Retort
stands and clamps are used for keeping the various parts in position,
and a millimeter scale (k) can be attached to one or both limbs of
the gauge. |

Should air make its appearance, it will collect under d, if it comes
from any part of the plant except the lower end of the shoot. It
can be removed from d by opening the pinch-cock and allowing water
to run in from the reservoir. Should it accumulate under a, the
shoot must be removed from the rubber tubing attaching it to the
glass, and water allowed to enter the pinometer at d rather slowly.
It will then be running out slowly at a, and the shoot is again fixed,
the current of water preventing any air getting in.

In any case the removal of air means the opening of the pinch-
cock at d, and this causes the mercury to go back to its starting point.
This can be obviated by inserting a stop-cock between the oblique
bit j-c (fig. 1) and the gauge. I do not however consider this necessary
for the apparatus, which as described here is intended chiefly for
qualitative and not quantitative observations.

The results obtained with the pinometer depend very much on
the point at which we fix it in the plant. I will refer therefore to 4
few actual experiments, which I hope will show the value, however
small, of the simple bit of apparatus just described.

In the first experiment to be described, a pinometer was fixed to

t For fixing the plant to the glass tube it is best, if possible, to employ some kind
of pressure tubing. ‘The latter can be made secure by tying with string or by employ-
ing some kind of clamp. The use of wire is to be deprecated.

irate oc

1905] DARBISHIRE—A PPARATUS FOR TRANSPIRATION STREAM 359

the main shoot by cutting the stem a short distance above the lowest
lateral shoot (fig. r). After a very short time the mercury in the
limb (g) of the gauge nearest the plant was seen to rise, as the latter
was withdrawing water from the pinometer. As the mercury rises,
the pull on the lower end of the shoot and on the upper end of the

m) | WHMIS EL

Fic, 1.—A fuchsia plant with one pinometer inserted just above the lowest
lateral shoot: a, the attachment of the upper shoot; }, that of the root stump; c, the
connection leading to the mercury gauge; d, the top opening of the pinometer, closed by
rubber tubing and pinch-cock; g, the level of mercury in inner, /, that in outer limb of
gauge; k, millimeter scale; the latter indicates shoot ti d the shoot wilting;

the stump at b, under pressure from the root, is flourishing.

Toot-end of the plant gets stronger, and gradually the leaves of the
Upper shoot wither. It will be noticed that the leaves of the lowest
lateral shoot are quite fresh. Before the upper shoot dies altogether
its axillary buds will generally develop and form small leaves, but
€ven these succumb in the end.

360 BOTANICAL GAZETTE [MAY

In this experiment, therefore, and at the height in question, our
plant exhibited shoot suction when tested by the pinometer. A
modified barograph recording cylinder, made by NEGRETTI & ZAMBRA,
was arranged to record the rise of the mercury in this particular
experiment. A burette-float was placed on the mercury in the open
end of the gauge (/). It was suspended by a fine thread, which ran
over a pulley and was attached on the other side to the free end of a
lever, the other end of which carried a pen which wrote on the revolv-
ing cylinder. The records taken during the first week showed that
the water taken in from midday to midnight and that taken in from
midnight to midday was in the proportion of three to two. Never
did any pressure from the root, during night or day, cause the
amount of water taken in to fall below nought.

In a second experiment a pinometer was fixed into a fuchsia
plant about one inch above the soil and quite below the lowest lateral
shoot. Root pressure manifested itself very soon and the mercury
was forced out of the inner limb of the gauge, rising of course as
rapidly in the outer one. In this case there was obviously pressure
on the cut surfaces of shoot and root. The leaves of the shoot kept
fresh as long as the pressure lasted, which was for sixteen days.
On the sixteenth day there was a difference in the level of the mer-
cury in the two limbs of the gauge of 20™™. Allowance however
must be made for the column of water resting on the mercury in the
inner limb of the gauge. After the sixteenth day the pressure was
gradually reduced, very probably owing to the root becoming exhaus-
ted, its supply of organic food from the green leaves being cut off.
The leaves began to wither as the pressure of the root decreased.

In a third experiment two pinometers were employed (/ig- 2 ).
One was fixed to the fuchsia plant just above the soil and the other
just above the lowest lateral shoot. The plant was therefore cut into
three parts, the lowest one of which, the stump, was devoid of any
lateral shoots. The gauge attached to the lower pinometer (P;) Very
soon indicated pressure from the root, that of the upper pinometer
(P,) suction from the two portions of the shoot. We have therefore
in this experiment an arrangement by which root pressure and shoot
suction can be observed at the same time. The difference in the
appearance of the leaves on the two portions of the shoot is very

ggg

1905] DARBISHIRE—APPARATUS FOR TRANSPIRATION STREAM 361

striking. The leaves of the upper shoot are dead, there being a strong
pull on the lower end. The leaves of the middle shoot are flourishing,
there being a push on its lower end, although the lower pinometer
is separated from the upper one by about two inches of stem only

LTA,

Fic. 2.—A fuchsia plant with two pinometers: the lettering as in fig. 1; the
lower Pinometer (P,) shows root pressure and its shoot (az) is flourishing i the upper
one (P,) exhibits shoot suction, and its shoot (a2) is wilting; r, reservoir of water,
5; its spout.

(between a, and b,). A fortnight after being set up, a reading for
this experiment showed a difference in level of the two columns of
Mercury of 18™™ in the lower pinometer, indicating pressure from
the root, and a difference of 20™™ in the upper pinometer, indicating
Suction from the shoot.

362 BOTANICAL GAZETTE [May

An experiment was also made with three pinometers inserted into
the stem of a fuchsia plant. The following readings were taken
after the experiment had been going on for some time. The lower
pinometer showed root pressure with a difference in the level of the
mercury columns of 31™™, the middle pinometer showed suction
with a difference in level of 85™™, and the upper. pinometer suction
with a difference of 63.5"™. The figures for the next day in milli-
meters were 39 (an increase of 8), 127.2 (42.2), and 128 (64.5)
respectively. The two lower pinometers were below the lowest
branches. In this case, therefore, root pressure could not be observed
even up to the lowest lateral shoot. As in all the previous experi-
ments, suction, where observed at all, was maintained day and
night, till, owing to the pull on the cut surface of the plant, air made
its appearance and the mercury returned to its original level.

The pinometer, as described here, is meant to be of use chiefly
for class and demonstration purposes. I wish now to point out what
the results obtained by employing it.are, that is to say what obser-
vations can be made on the transpiration stream.

We can readily see that in our plants, at least, it isnevera function
of the root to press water up into the leaves. By day and by night
the phenomenon of root pressure can be observed only below the
lowest shoot. But root pressure is an extremely useful if not a
necessary process by which the rise of the water is started. Referring
to those of our experiments in which the lower cut surface of the shoot
was exposed to pressure, we see that the plants do not suffer much by
the stem being cut across. It appears to be necessary that water should
be forced into the lowest end of the vascular system; or at least
there should be no pull on it. Root pressure, therefore, is not only a
symptom of the avidity with which the roots are absorbing wateT,
but it is of importance as assisting in the starting of the transpiration
current.

It must be mentioned here, however, that the insertion of a pino-
meter into a fuchsia plant is in any case a serious thing for the latter-
The phenomena of exudation and bleeding seem to cease entirely
when the shoot has been severed from its connection with the root.
This statement holds good whether the gauge of the pinometer shows
pressure or suction. The fact that at different heights in the plant

Oe EN ae ee ee ee a ee

1905] DARBISHIRE—APPARATUS FOR TRANSPIRATION STREAM 363

we get readings which at least show that the pressure may vary very
much in even two or three inches of stem, is a proof, I think, that the
“atmospheric pressure” theory of the rise of water is not correct.

When we get away from the root we find the phenomenon of shoot
suction manifesting itself. In fact, as already pointed out, shoot
suction generally seems to be stronger than the pressure from the
root at any point except just above the root. That is to say, the
shoot is able at any time to take in more water than can be supplied
by the root. Numerous experiments show that this is the case in
winter and summer, day and night, in the plants I experimented upon.

Which part of the plant is exerting this suction? I have spoken

here of shoot suction, but it is possible to split up this part of the
transpiration stream into two distinct processes, namely the leaf
suction and the wood lift. There is not much mystery about the
ormer. We can understand how the water is removed from the
finest endings of the vascular system. If, however, we remove the
leaves from a shoot which has been attached to a pinometer or even
cut off the upper part bearing leaves, leaving only a short leafless
shoot stump, we still get water rising in the wood and exerting a pull
on the gauge of the pinometer. The activity of the leaves simply
removes the-water from the top of the vascular system, and _ this
water is replaced by a process going on in the wood.

I am mentioning this simply to show that in the experiments with
the pinometer the taking in of water by the shoot is not a phenomenon
of leaf suction but one of wood lift. The force of the wood lift is very
great; it is generally greater than the root push. But, as our experi-
ments show, it cannot act efficiently for any length of time if the
lowest end of the vascular system is exposed toa pull from below. It
is as yet not known how the wood lift acts. It is therefore all the more
necessary to make as many observations as possible on this process.
I think the pinometer does make it possible to observe at least one
property of this water current. It is continually in a state of what is
known as ‘negative pressure.” A natural result of this is that the
air, which the water absorbed by the plant contains under ordinary
atmospheric pressure conditions, escapes from the water when inside
the plant. Over and over again my experiments have been brought -
to an end by air collecting, generally under the shoot. Air-bubbles

364 BOTANICAL GAZETTE [MAY

are bad for the conducting of the experiments, and the more they are ~

kept out the more water is absorbed by the plant when attached to a
pinometer. It is obvious that the plant must also guard against the
accumulation of air in the column of water which fills the vascular

system. No doubt this will in part account for the peculiar structure |

of the woodelements. Do the latter bear any relation in their struct-
ure to the nature of the water generally found in the localities in
which the plant grows ?

The form of pinometer mentioned in this paper is intended, as
already mentioned, essentially for demonstration purposes. It is
possible by its use to observe, more clearly than hitherto, the relation
between shoot suction and root pressure. Owing to the preliminary
nature of this paper I have refrained from giving any lengthy readings
taken during the experiments. Their value in any case would not
be very great, being taken with the simple form of pinometer here
described. A more elaborate bit of apparatus is therefore in course
of construction, by which any air making its appearance in any
part of the pinometer is removed automatically at regular intervals
without altering the conditions of pressure inside the system of tubes.
Furthermore, it is connected with an automatic recording instrument,
in which the difference in the height of the two mercury columns will
be reduced as much as possible.

The nature of this paper, I hope, will excuse my having made no
reference to any literature. It was not my intention to discuss the
old or bring forward a new theory with regard to the rise of water.
We still have the suction by the leaves, the pressure by the root, and
the as yet little understood lifting of the water by the wood. In
writing this paper, it was my object to give an account of a very
simple bit of apparatus. by means of which various phenomena con-
nected with the transpiration stream in small plants could be readily
observed. I hope later on to be able to publish some more detailed
observations on the properties of the wood lift, taken by more
elaborate instruments.

Diagrams illustrating the working of the pinometer were shown
at the last Cambridge meeting of the British Association.

UNIVERSITY OF MANCHESTER, ENGLAND.

Spas at's Te Se aa a ee er ee

eT at ee ee ee ee

BRIEFER ARTICLES.

POLYEMBRYONY IN SPHAGNUM
(WITH THREE FIGURES)

DuRING July 1904, while collecting sporophytes of Sphagnum for class
use, the writer discovered the peculiar specimens described in this note.
The particular species of Sphagnum has not been determined, but it is a
large variety which grows abundantly
in the swamps around Tower, Minne-
sota

Among the thousands of sporo-
phytes handled, five cases were found
Where in two archegonia at the end of
the same branch the oospheres had
been fertilized and each oosperm had
developed into a normal sporophyte.
In these specimens each capsule was
enclosed by its separate calyptra and
the haustra of each two sister sporo-
phytes were inserted side by side in
the bulbous end of the pseudopodium
(ig. 2).

Two cases were also found where two sporophytes had developed in
the venter of one archegonium (jig. 2). They were more or less distorted

|
|
|

ts £2

and modified by mutual pressure, but each presented the essential structure
of a normal sporophyte (fig. 3). When studied in sections, the tissue of
wa. 365

366 BOTANICAL GAZETTE [MAY

each sporophyte appeared to be entirely independent of that of its twin
except in the loose tissue forming the base of the haustra, where the line
of separation was not clearly recognizable in all sections. In the region of
contact, the epidermis, while quite distinct, was less strongly developed
than on the free faces. Since the sporophytes were nearly mature at the
time of collection, it was impossible to determine the conditions which led
to the production of twins. It is probable that two functional oospheres
were produced in the venter of each archegonium, although it is not impos-
sible that twinning took place through an early branching of the young
protocorm.—HaAroip L. Lyon, University of Minnesota.

ADENODERRIS, A VALID GENUS OF FERNS.
(WITH TWO FIGURES)

THE genus Polystichum,! typified by the well-known P. Lonchitis
constitutes a natural and well-defined group of ferns and has commonly
had general recognition, recently even by American writers.? Its species
are characterized, briefly, by their rigidity and erect habit, smooth firm
dryish texture, mainly auriculate and spinulose or mucronate divisions,
abundant chaff, free venation, and ordinarily by the orbicular centrally
peltate indusia. The genus was divided by Jon SmirH into two sections:
the one, typified by P. aculeatum and P. Lonchitis, with fasciculate generally
erect acaulose vernation; the other, typified by P. coriacewm, with uniserial
sarmentose vernation. In view of that writer’s natural bent for generic
Segregation along the very lines on which the species of this genus were
placed in two sections, it would not have been surprising if he had accorded
to each section generic rank. That such a disposition—in view of the two
radically different types of vernation—was not made is significant, for it
emphasizes Smiru’s belief that in other respects the species are in too close
agreement to warrant their division into two genera. There is, however,
variation in one other particular; in certain species of both groups the
indusium is caducous, or, as in P. tenue,3 apparently quite wanting; but
even in these cases the appearance of the plants is so unmistakably that
of Poly stichum and their agreement in the main particulars so essential
that a rational course seems to necessitate their recognition as true mem
bers of the genus. We have thus a group of species which, though offering

? Rorg, Teut. Fl. Germ. 3:69. 1800.

2 GILBERT, in papers presented at the Boston Meeting (1898), 22. 1899. UNDER
woop, Our native ferns, ed. 6, 11 5. 1900. DAVENPORT, Rhodora 4:9. 190?) etc.

3 GILBERT, Fern Bull. 8:63. rg00.

Ne ee Dg RS CE ee, pay eS ee ee a cee sa

el eee

Se

1905] BRIEFER ARTICLES 367

variation in two lines, is nevertheless, in the characters enumerated at the
beginning of this paper, one of the most unified to be found among the
ferns. Indeed, as GiLBeRT has remarked, the general characters of the
genus are so distinctive and obvious that the descriptive term polystichoid
has come into rather common use.

The historic treatment of Polystichum, although of much interest,
need not be here discussed at length. The genus has been made often to
include diverse groups not properly associable with its typical members;
and in several cases it has been an author readiest to admit numerous small
genera who has failed to realize the essential unity of the group and to
refuse to admit unlike forms. us Moore retained Cyclopeltis J. Sm.
under Polystichum, though it constitutes a most distinct natural genus.

d more recently, Drets,4 while properly removing Cyclopeltis, never-
theless allows Phanerophlebia, Cyrtomium, and Adenoderris to remain «
within the gerius. Cyrtomium and Phanerophlebia have recently been
held distinct from each other and from Polystichum by UNDERWoOD,5
and it now appears certain that Adenoderris, long ago founded by JoHN
SMITH upon the anomalous Aspidium glandulosum of Hooker and Greville,
Tepresents a perfectly valid genus. The discovery of a second species of
Adenoderris with very different venation, from Guatemala, is of interest
and has seemed to render desirable the present notice of the genus and
its relationship.

ADENODERRIS J. Sm. Hist. Ferns 222. 1875.6—Small plants of lax
habit, distinct from Polystichum by their herbaceous texture, aspinulose
margins, and dense glandular-pilose covering —Type, Aspidium glandu-
locum Hook. and Grev. from Jamaica. Species two.

Adenoderris viscidula (Mett.).— Aspidium glandulosum Hook. and
Grev. Icon. Fil, 2: pl. 140. 1831; not Aspidium glandulosum Blume,
Enum. Pl. Javae 2: 144. 1828. Adenoderris glandulosa J. Sm. Hist. Ferns
223.1875. Aspidium viscidulum Mett. Abhand. Senck. Nat. Gesells. 2: lal
1858.—The species was very fully characterized upon specimens received
from several collectors in Jamaica and is the subject of an excellent illus-
tration by Hooker and GREVILLE. Lately it has been well described by
JenMan,? SMITH, basing his genus upon the single species, published the

4 DIELs, in Engler and Prantl, Die natiirlichen Pflanzenfamilien 14: 183-189. 1899.
_ _5 UNDERWoop, Bull. Torr. Bot. Club 29:121-136. 1902. See also UNDERWOOD
in Bull. Torr. Bot. Club 26: 205-216. pls. 359-360. 1899.

’ ° The genus seems to be published here for the first time, although the author
cites the date 1852.
7 Bull. Bot. Dept. Jamaica 2: 197-198. 1895.

368 BOTANICAL GAZETTE [MAY

following characters: ‘‘Vernation fasciculate, erect, acaulose. Fronds 6
to 8 inches in length, oblong, lanceolate, pinnatifid, densely covered with
pilose glands, decurrently attenuated to a short stipe. Veins pinnately
forked. Receptacles punctiform, medial. Sori round. Indusium orbicular,
occasionally reniform.”8 In habit the plant was said to be “totally at
variance” with any species of Polystichum, and
this opinion was formed entirely from her-
barium specimens. An examination of mate-
rial in the field indicates even more plainly
how inappropriate has been the usual sys-
tematic association of this peculiar plant with
true members of Polystichum. The fresh
plants are spongy, very lax, and intensely viscid,
and except for the peltate indusia have nothing
1.—Adenoderris to suggest a close relationship with the stiff
smooth spinulose Polystichums. The venation
of A. viscidula, which was not indicated by
Hooker and GrevILLE, is shown in fig. 1, which represents the middle
portion of a Jamaican specimen (Clute no. 333; U. S. National Herbarium,
no. 349588). This feature and the position of the sori are discussed under
the next species.

A. viscidula is known only from Jamaica and Cuba. JENMAN states
that in Jamaica it occurs upon ‘“‘rocky banks and skirts of forests 1500~
3000 't altitude; plentiful in one place at least between Gordontown and
Guava Ridge. There is, however, but one sheet in the Jenman Herbarium
at New York. Other Jamaican specimens are: Clute no. 333; collected
above Gordontown, March 12, 1900, at an altitude of 4507; Underwood
no. 2498, collected near the Green River (below Cinchona), April 22, 1903,
at an altitude of 750™; and specimens collected by D. E. Watt at or ‘cee
the last locality in May 1903. The Cuban record? is based upon C. Wright
no. 1052. Specimens of this number in the D. C. Eaton herbarium ‘gabe
identical with the Jamaican plant; to them is attached Wright’s original
label stating that they were collected in rocky ravines on mountain sides
near Josephine, October 25 (1859).

Adenoderris sp. nov.—A delicate plant of small size, the fronds
glandular throughout. Rhizome slight, erect, having Jong fibrous rootlets
rather thickly clothed with delicate bright brown chaff: fronds 8°™ long,
short-stipitate, spreading, oblong-lanceolate, deeply pinnatifid, with about

8 Only orbicular indusia have been observed by the writer.

9 Hooker, Sp. Fil. 4:6. 1842.

Fic.
viscidula (Mett.) Maxon;
natural size.

1905] BRIEFER ARTICLES 369

ten pairs of usually subopposite to alternate pinnae; pinnae ovate to deltoid,
the middle ones to-13™™ long, the lowermost scarcely reduced, all deeply
divided into about three blunt lobes upon both upper and lower margins,
each lobe usually once soriferous near the outer side at the base, the sorus
being borne at the extremity of a spur given off from the otherwise usually
simple single veinlet; venation terminating well within
the lobe; indusium minutely glandular, orbicular,
centrally peltate.

“Founded upon no. 868 of John Donnell Smith’s
Guatemalan plants; said to have been collected by
von Tiirckheim at Sesisp, Department of Alta Vera
Paz, altitude 1200™, March 1886, and distributed as
As pidium glandulosum. -The most perfect material
of this number the writer has seen is that preserved
in the D. C. Eaton herbarium at Yale University,
and this, having served for the accompanying illus- 0”, 0. sp.; natural
tration (fig. 2), may stand as the type, though the
specimens in the United States National Herbarium and the herbarium
of the New York Botanical Garden are of the same collection. Captain
SMITH has stated (in litt.) that duplicates were presented also to the
W. M. Canby, Philadelphia Academy, Kew, Berlin, Paris, and DeCandolle
herbaria.

Adenoderris sororia is distinct in all states from A. viscidula, though in
its lax habit, slight texture, aspinulose margins, and glandular covering
it shows an undoubted generic alliance with that species. It differs spe-
cifically in its less size, bipinnatifid condition throughout (A. viscidula
though larger is only deeply once-pinnatifid), more sparse glandular cover-
ing, and in its spreading simpler included venation, the sori being borne
terminally at the apices of the veinlets. In A. viscidula the veinlets are
Pinnately forked and excurrent to the suberose margins, the sori being
borne dorsally, i. é., upon the veinlets and at some distance from the margin.
These differences, while very marked, appear to be no more than specific,
and SuitH’s original generic diagnosis quoted above must therefore be
amended as regards venation. :

Both drawings, which are by Mr. H. D. Howse, are natural size.
That of A. sororia represents parts of the third, fourth, fifth, sixth, and
seventh pairs of pinnae of a frond of the type specimen—WruaM R.
Maxon,

CURRENT LITERATURE.
BOOK REVIEWS.
Rusts of Switzerland.

THREE NOTABLE WORKS on plant rusts have appeared within a year: KiE-
BAHN’s Wirthswechselnde Rostpilze, the first volume of Sypow’s Monographia
Uredinearum, and FiscHer’s Die Uredineen der Schweiz. Each of these works
covers a distinct field, and in its own way marks advance in the elucidation of the
world’s rust flora, and in an understanding of the problems connected therewith.
The last named work,? although embracing a limited region, is conceived upon
such a broad plan, and carried out with so much completeness, that it serves as
the best model yet produced for a uredineous manual.

A large number of the species found in Switzerland are cosmopolitan. Every
species is described in detail, and with few unavoidable exceptions direct from
specimens, and is also illustrated with outline drawings. The descriptions are
especially full, embracing not only the usual characters, but those derived from the
pycnidia, the peridial cells, and the germ pores. The illustrations are drawn to a
uniform scale, and are skilfully made. Often a dozen or more teleutospores are
shown. Usually uredospores are included, and always drawn in a norm
upright position, a most commendable innovation. In most cases a transverse
section of two or more peridial cells shows their varying thickness of wall and
sculpturing, another helpful innovation.

The notes which follow the diagnoses, briefly stating how much is known of
the life history of the species, are helpful and suggestive. The list of Swiss stations
for each species is more especially of local value to collectors.

The systematic arrangement is essentially that of DreTeL in ENGLER and
PRAnTL’s Pflanzenfamilien. Under the genera the species are distributed by
hosts and morphological characters, with a view to showing relationship. A
general key on the same basis is provided, together with excellent indexes, and a
full modern bibliography.

chapters are of unusual interest to the general mycologist. In one the

distribution of the Uredineae in Switzerland is analyzed and discussed, taking

into account the ecological factors controlling the hosts. In another the general

classification and the grouping of species within the genera are considered in the

light of the most probable hypotheses regarding the phylogenetic descent —

rusts. And in the third the difficult questions regarding the value of various
Sy =e, ee ee, AN 5 1: ee es ie } Pp

recente =
LCo

1 FiscHer, Ep., Die Uredineen der Schweiz. Beitr. z. Kryptogamenflora one
Schweiz. II? imp. 8vo. pp. xciv+590. figs. 342. Bern: J. K. Wyss. 1904- 20 4 Z
MAY

379

1905] CURRENT LITERATURE 371

This work by Dr. FiscHer, for which he has been preparing for more than a
dozen years, is replete with new matter of great value and is altogether admirable.
—J. C. ARTHUR.

MINOR NOTICES.

THE FOSSIL FRUITS from the lignites of Brandon, Vermont, are the subject
of an important contribution by PeRKins.? In 1861 LEsQUEREUX described
twenty-one species from this locality, which he regarded as of approximately the
same age as the (Eningen stage of the Swiss Miocene. In 1902 KNowLTon pub-
lished a brief paper of forms from this locality which represents all we knew of
this interesting flora up to the time of publication of the present paper. The
deposits have been much obscured and inaccessible for a half century, until the
coal famine of 1902, when the lignite came into demand locally as a substitute for
coal. The state geologist, GEORGE H. PERKrNs, was enabled to secure a mag-
nificent collection of the fossil fruits during the mining operations, and the present
paper contains the result of preliminary study of these collections. One hundred
and eighteen species are recorded, and many new forms of more or less doubtful
botanical affinities are described. These are largely included in the following
new genera: Monocarpellites (11 spp.), Hicoroides (5), Bicarpellites (5), Bran-
donia, Rubioides, Sapindoides (6), and Prunoides. The illustrations consist
chiefly of photographs of type specimens, which are perhaps less satisfactory in
showing details than careful drawings. The flora is unique in the abundance
and variety of its fruits, and it is to be hoped that future study by Dr. PERKINS
will demonstrate with more precision the exact age of the formation containing
them. The accompanying clays should be searched for leaf remains.—EDWARD
W. Berry.

PopuLaR ACcouNTs of soil inoculation for legumes devised by Moore have
attracted wide attention, so that it is of special interest to receive his own account.
The nitrogen is fixed by the tubercle-forming bacteria within their bodies. This
was determined by cultures in flasks containing nutrient solutions without nitro-
gen. There was no increase of nitrogen in the solution, but a marked increase
in the organisms themselves. In its biology the organism is therefore considered

* PERKINS, Geo. H., Description of species found in the Tertiary Lignite of
Brandon, Vermont. Rept. State Geologist, Vt. 1903-1904. pp- 174-212. pls. 75-SI.
1904.
> Moorg, G. T., Soil inoculation for legumes, etc. _U. S. Dept. of Agric., Bureau
of Pl. Industry, Bull. 71, pp. 72. pls. 10. 1905.

372 BOTANICAL GAZETTE [MAY

a method of putting up for distribution pure cultures of Pseudomonas radicicola,

own in nitrogen-free media and dried on cotton immersed in the culture. These
cultures are sent out by the U. S. Department of Agriculture together with pack-
ages of nutrient salts to multiply the organism. e mass-culture thus obtained
is used to inoculate the seed or the soil. Numerous reports from farmers of all
states indicate that this method will prove successful and practicable——H. Has-
SELBRING.

Tue Orrice of Experiment Stations of the U. S. Department of Agriculture
has recently issued a valuable syllabus of the diseases of the Irish potato in the
United States by Stewart and Eustace of the New York (Geneva) Experiment
Station. This is the second of a new series of publications designed primarily
to assist farmers’ institute lecturers in the presentation of various important sub-
jects to farmers. Each lecture is accompanied by a set of lantern slides, there
being in this case forty-seven slides in the set. Arrangements have been made
to loan these slides to lecturers, or they may be purchased through the Department
of Agriculture at a very moderate cost. Both the syllabus and the accompanying
set of slides should prove of great value to teachers of plant pathology in presenting
the diseases of the potato. A brief bibliography is appended to the syllabus,
which contains references to the most important American literature of this sub-
ject.—E. Mrap WILLcox.

AN UNDATED EDITION of ANDREWS’s Botany all the year round has been issued,
which differs from the preceding one’ only in being bound into one volume with
a flora by W. Nevin Geppes.° This flora brings together compact and simple
descriptions of the more common flowering plants, not only native and naturalized,
but also cultivated, and includes about 1,250 species. The sequence of families
is an interesting reminiscence of the old time classification, and it is quite a penne

0 see gymnosperms once more as a subclass of “Exogens or Dicotyledons.”
The nomenclature, too, is frankly “conservative and eclectic;” but the book does
as well as any other to enable a student to find names for plants, which seems to
be regarded as a very valuable exercise.—J. M. C.

THE FOURTH PART of SARGENT’s Trees and shrubs? contains plates and
descriptive text of species of Acer (13), Parthenocissus (3), Malus, Oroxy lum,
Phellodendron (3), Arctostaphylos (2), Dracaena, and Pinus (2). There are

4 Stewart, F. C. and Eustace, H. J., Syllabus of illustrated sgn on Lage
diseases and their treatment. Office of Exp. Stat. U. S. Dept. Agr. Farm —
Lecture 2:1-30. 1904.

5 See Bot. GAZETTE 35: 439. 1903.

6 GeppEs, W. Nevtn, A brief flora of the eastern United States. Ne
Cincinnati, Chicago: American Book Company, with no date.

7 SARGENT, C. S., Trees and shrubs. Illustrations of new or little known ligneous
St — chiefly from material at the Arnold Arboretum of Harvard Unt iversity-

IV. pp. 151-217. a 76-100. Boston and New York: Houghton, Miffin &

Cages 1905.

Ww York,

;
3
;
4
7
:

ea i NSS a ak ity ese a is A

|
|

1905] CURRENT LITERATURE 373

two new Chinese maples, a conspectus of the maples of eastern continental Asia,
two new American species of Parthenocissus, and one new Chinese Oroxylum,
by ALFRED REHDER; a new Phellodendron, by CHARLES S. SARGENT; two new
Californian species of Arctostaphylos, by ALICE Eastwoop; a new Central
American Dracaena, by JoHN DoNNELL-SMITH; and two new Mexican pines by
G.R.SHaw. This a closes the first volume, and contains the title-page, index,
and corrections.—J.

HERBERT hiss we brought together a most interesting collection of
descriptions and excellent photomicrographs of commercial timbers. Being
able to draw upon the British colonies for material, the list is unusually full.
An introductory chapter gives a general account of the varying structure of wood
from the standpoint of one who has a practical acquaintance with timber; and
this is followed by practical hints as to its identification. The list of timbers
includes 247 numbers, each species with a very full description, all the genera
being illustrated by photomicrographs.—J. M. C.

E PARTS of an illustrated handbook of the angiospermous woody plants
native or hardy in middle Europe . K. ScHNEER have appeared.2 The
illustrations are numerous and excellent, floral dissections, fruits, and leaves
being given, and in many cases fine half tones of characteristic bark. The
ENGLER and PRANTL sequence is used. The first part includes Salicacae to
Fagaceae, with 95 illustrations; the second part, Fagaceae (Quercus) to Ber-
beridaceae, with 102 illustrations; the third part, Berberidaceae to Spiraceae,
with go illustrations.—J. M. C

A SECOND EDITION of Grout’s Mosses with a hand lens has been published,'°
including the common liverworts. The review of the first edition’: stated the
general scope and purpose of the book. The demand for a new edition has
justified the undertaking, and with the addition of much new material the author
believes that he has reached the limit of the hand lens. The addition of the
common liverworts represents a very natural extension of the author’s method,
and introduces beginners to a group they must have encountered repeatedly in
collecting mosses.—J. M. C

8 STONE, HERBERT, The timbers of commerce and their identification. Illus-
trated with 186 oo . by Arthur Deane. Imp. 8vo. pp. XXXViii + 311
London: William Rider & S 1904.

9 SCHNEIDER, CAMILLO pee Illustriertes Handbuch der Laubholzkunde.
Charakteristik pa in a. npr und im Freien angep ten a
Spermem Geholz-Arten und Formen mit Ausschluss der Bambuseen und Kakteen.
Jena: Gustav Fischer. r904. Each part M4.

Illustrated by Mary V. Tuayer. Second edition, revised, enlarged, and including
the hepatics. 8yo. pp. xvit+208. pls. 39. figs. 118. New York: The Author, 360
Lenox Road, Flatbush. 1903.

™ Bor. GAZETTE 31:132. 1900.

374 BOTANICAL GAZETTE [way

NuMBERS 221 and 222 of ENGLER and Pranti’s Pflanzenjamilien have
appeared, the former containing a continuation of the lichens by A. ZAHLBRUCK-
NER, the latter a continuation of the mosses by V. F. BrorHerus.—J. M. C.

THE SEVENTH FASCICLE of DALLA Torre and Harms’s Genera Siphono-
gamarum*? has just appeared, including wou oe Acanth (7938. Echina-
canthus) to Compositae (9371. Liabum).—J. M

NOTES FOR STUDENTS.

PALEOBOTANICAL NOTES.—BRABENEC'? records thirteen species of plants
from a new locality, Holedeé, in the Saaz basin of northern Bohemia, describing
new species in Carya, Acacia, Paliurus, and Porana.—BrErry™ describes addi-
tional material from the Cretaceous near Cliffwood, N. J., consisting of rather
fragmentary remains of nine additions to that flora, making the total number of
known forms ninety-five —The same author describes*s a new palm, represented
by abundant but fragmentary remains of leaves, from the mid-Cretaceous of the
coastal plain in Delaware and Maryland.—In notes on Tertiary plants PEN-
HALLOW’® describes wood of Taxodium and Thuja from the basal Eocene of
Alberta, and of Cupressoxylon from the Cretaceous of Assiniboia. Wood of Pseu-
dotsuga Douglasii is identified from near Bozeman, Montana, and the charac-
teristic Pleistocene Larix americana (Larix laricina) is identified from Georgia.
The latter find is most interesting, showing as it does that the larch had pushed
about 480 miles farther south than its present southern limit during the Scar-
borough period of the Glacial. leat of Pinus rigida is recorded from the
Pleistocene of New York; and Juniperus californica and Pseudotsuga macrocarpa
are recorded from the Seiad a at Orleans, Humboldt county, California.
The paper concludes with an enumeration of a number of species, based on leaves
and wood, from the well-known Don River Pleistocene of Canada.—WaiTE”’ has
a paper on “Fossil plants of the group Cycadofilices,” in which the chief points
of interest in this most interesting group are enumerated. He concludes, with
WILLIAMSON, that secondary wood originated, polyphyletically and polychron-
ously, as an engineering feature. It is suggested that the heterosporous filicean
ancestors of the group Pteridospermae will be found in the lowest Carboniferous
if not in the Upper Devonian. Should the author’s Kalymma grandis prove ”

12 Datta Torre, C. G. DE, and Haras, H., Genera Siphonogamarum ad systema
Englerianum conscripta. Fasc. 7. pp. 481-560. Leipzig: Wilhelm Engelmann. 19°5-
M6.

13 BRAB , F., Ueber einen neuen fundort v. egees Pfi., etc. v- Saazer
Schichten Bull 1 ar “iel Sci. Bohéme, 1904. pp. 8

14 Berry, E. W., Bull. ee Bot. Club 32: 43-48. a I-2. 1905.

15 BERRY, E. W., Torreya 5: 30-33. figs. 2. 1905.

x6 PENHALLOW, D. P., Trans. Roy. Soc. Can. II. 10:57-76. 190

17 WuIte, Davi, U. S. Geol. Surv. Professional Paper No. 35: PP- ee pls. 2-0-
1905.

ies

1905] * CURRENT LITERATURE 375

be pteridospermic, as is expected, it will serve to emphasize the evidence of the
fossil wood (Dadoxylon) as to the occurrence of gymnospermous-like plants in
the middle Devonian (Genesee), a time nearly as remote as that furnishing the
oldest known ferns of undoubted authenticity, and would seem to render the search
for ancestral forms of these comparatively highly organized pteridospermic types
futile—The same author gives'® a systematic account of the Devonian flora of
the Perry basin, Maine, listing twenty-nine species, not including nine more or
less doubtful species which Dawson reported from this locality. Several new
generic and specific types are described, and numerous nomenclatorial points are
cleared up. The paper is fully illustrated —Epwarp W. BERRY.

CLAUSSEN” gives an account of the life history of the ascomycete Boudiera.
The sexual organs consist of short filaments which are developed in clusters and
become very much twisted together, presenting spirally wound structures some-
what resembling the sexual organs of Gymnoascus. These filaments are differ-
entiated into antheridia and ascogonia which fuse. After fertilization a cluster

ascogonium, containing several nuclei, which bears a terminal sterile cell, perhaps

0. The nuclei in the sterile cell break down, and it finally becomes the medium
(conjugation tube or trichogyne) through which the protoplasm from the anther-
idium enters the ascogonium. The antheridial filament is thinner and also con-
tains several nuclei, which after the fusion with the sterile terminal cell pass into
the ascogonium. The fertilized ascogonium then contains some ten or twelve
nuclei and the antheridial filament and sterile terminal cell are empty. The
nuclei in the fertilized ascogonium fuse in pairs. The empty antheridial filament
and sterile cell collapse. The fertilized ascogonium increases in size, the fusion
nuclei become conspicuous, and it finally divides into several cells by cross walls.
The ascogenic hyphae are very short and the tips take at once the form of a hook
and contain four nuclei. One nucleus lies in the tip, another in the stalk, and the
remaining two in the bent portion of the hook. The four- nucleate condition is
derived from a two- nucleate, but the preceding history is not known. e pair
of nuclei in the bent portion of the hook become separated by walls from the tip
and base of the filament and shortly coalesce; this cell becoming then a young
ascus containing the single fusion nucleus. Thus each ascogenic hypha develops
a single ascus. The terminal cell remains indefinitely, but there is no development
from it. Several asci are formed from one ascogonium, but not simultaneously.
Their number is believed to agree with the number of fusion nuclei in the ascogo-
nium. The process of aor formation appears to follow HARPER’s account, but
LL

8 WuiTE, Davip, Smithsonian Miscellaneous Coll. 47: 377-39°- pls. 53-55: 1995-
*9 CLaussEN, P., Zur Entwickelungsgeschichte der Ascomyceten Boudiera. Bot.
Zeit. 631: 1-28. pls. I-3. 190

376 BOTANICAL GAZETTE [MAY

details are not given. The paper ends with a long historical discussion of the
disputed sexuality of the ascomycetes and supports HARPER’s conclusions.—
B. M. Davis

ITEMS OF TAXONOMIC INTEREST are as follows: T. MAKINO (Bot. Mag.
Tokyo 18:156. 1904) has separated the Japanese form referred to Croomia
pauciflora T. & G. as a distinct new species (C. kiusiana).—W. B. HEMSLEY
(Hooker’s Icon. Plant. IV. 8:1. 2781. 1905) has named the type of a new
West Australian genus of Compositae (Helichryseae) Thiseltonia Dyeri.—Pu.
VAN TIEGHEM (Jour. Botanique 19: 45-58. 1905) has made the genus Octocnema
(provisionally placed under Olacaceae) the type of a distinct family (Octocnema-
ceae).—A. A. Hetter (Muhlenbergia 1:111-118. 1905) has described new
Californian species of Ribes, Trifolium, and Stachys.—J. Carport (Bull. Herb.
Boiss. 5:208. 1905) has described a new genus (Alophosia) of mosses from San
Miguel —L. RADLKOFER (idem 222) has described a new genus (Sisyrolepis) of
Sapindaceae from Siam.—Cart Merz (idem 232) has described a new Venezuelan
genus (Glomeropitcairnia) of Bromeliaceae with two species.—J. C. ARTHUR
(Annales Mycol. 3:18. 1905) has described a new genus (Baeodromus) of Uredi-
neae with two species.—E. L. GREENE (Ottawa Nat. 18:215-216. 1905) has
described new Canadian species of Malus and Fragaria (2).—W. FAwceETT and
A. B. RENDLE (Trans. Linn. Soc. London II Bot. '7:1-13. pls. 1-2. 1904), in a
monograph of the Jamaican species of Lepanthes (Orchidaceae), recognize 12 spe-
cies, describing 5 as new.—N. L. Brirron (Bull. N. Y. Bot. Garden 3: 441-453-
1905), in his first ‘Contributions to the flora of the Bahama Islands,” has pub-
lished 15 new species and a new genus (Bracea) of Apocynaceae.—J. J. SMITH
‘(Recueil Tray. Bot. Neerl. 1:49-51. pl. 2. 1904) has described a new genus
(Gynoglottis) of Orchidaceae from Sumatra.—M. L. FERNALD (Rhodora 7: 47-49-
1905) has described a new species of Comandra that ranges from Quebec to Assini-
boia and Kansas.—F. STrpHANI (Hedwigia 44:74. 1905) has described two new
Himalayan genera (Gollaniella and Massalongoa) of Hepaticae——L. DIELs and
E. Pritzet (Bot. Jahrb. 35:564. 1905) have published a new Australian genus
(Pentaptilon) of Goodeniaceae—G. E. Davenport (Fern Bulletin 13:18-21-
1905) has published a new Mexican Aneimia, with plates.—A. and E. S. GEPP
(Jour. Botany 43:108. pl. 470. 1905) have ra a new genus (Leptosarca)
of the red algae from the South Orkneys.—J.

BERGH’s second paper?° on the formation ? ire chromosomes deals
with the behavior of the chromatin from the telophase of the last division in the
sporogenous tissue up to the strepsinema stage of the first mitosis in the micro-
spore mother-cell. In a very early prophase of the first mitosis in the mother-
cell, slender filaments are formed from the chromosomes of the sporogenous
cell. Synapsis then takes place and at its close the filaments appear a5

20 BERGHS, JULES, La formation des chromosomes hétérotypiques dans la spear

génése végétale. II. Depuis la sporogonie jusqu’au spiréme définitif dans la mic-
rosporogénése de l’Allium fistulosum. La Cellule 21:383-397- pl. I- 1904-

ee Se

ee ee i eS ee ee

1905] CURRENT LITERATURE : 377

“thick spirem.” During synapsis the dual nature of the filaments is evident.
It is due to a longitudinal approximation of two filaments and leads to the forma-
tion of the thick spirem. The so-called longitudinal division is merely the
reappearance of the line of approximation. Consequently the reduction in the
number of chromosomes seen in the prophase of the heterotypic division is only
an apparent reduction, due to the approximation of two somatic chromosomes,
which in the mature chromosomes play the réle of daughter chromosomes. The
reduction is really accomplished by the first mitosis, which separates these daughter
chromosomes

The third paper?! in the series deals with microsporogenesis in Convallaria
majalis. The results confirm the conclusions already drawn from the author’s
study of Allium fistulosum and Lilium speciosum. The two daughter chromo-
somes which constitute the mature heterotypic chromosome represent two longi-
tudinal parts of one segment of the thick spirem. This thick spirem is formed
by the longitudinal approximation of slender oe ey Seite by ais
This approximation takes place during th
takes place in the strepsinema stage, resuietere fe in an apparent sper Si division.
The heterotypic division is a reduction division, which distributes to the two
poles complete somatic chromosomes.—C. J. CHAMBERLAIN

THE RESEARCHES of SCHWARZ, ELFVING, VoCHTING, RAcrBoRSKI, and others
on the influence of gravitation upon the rate of growth of erect and inverted organs

(vertical) is not affected by gravitation, such organs when inverted manifest a
checking of the rate of growth in length. Improvement upon ELFvinc’s tech-
nique was secured by using artificial light of constant intensity instead of sun-
light for keeping the organs parallel with gravitation; also by substituting a non-
shrivelling substratum for enue crumbs for the growing of fungi. For such
organs as could not be t 1 in a vertical position by = ee of light, recourse
was taken to the use of pulleys and weights. The expe terial employed
includes both positively and negatively geotropic organs as found in members of
various genera of fungi, of apie s, and of dicotyledons, as well as the
branches of weeping trees. While nodes of grasses constitute an viet HO the
uniformity of behavior revealed by numerous other forms is so great to permit
a formulation of this general statement, namely, parallelotropic ses as 2
consequence of inversion suffer a reduction in the rate of growth in length. In the
case of the sporangiophores of Phycomyces prolonged inversion is followed by
-_-—

« BeRGHS, JULES, La formation des chromosomes hétérotypiques dans la sporo-
wha végétale. III. La microsporogénése de Convallaria majalis. La Cellule
53: pl. 4.

22 HerInc, GeorG, Untersuchungen iiber das Wachsthum inversgesteller Pflanz-
€norgane. Jahrb. Wiss. Bot. 40:499-562. 1904.

373. BOTANICAL GAZETET [MAY

accelerated growth, although a temporary inversion (one hour) is followed by
diminished growth. In the case of the descending branches of weeping trees,
gravitation not only causes a reduced rate of- growth, but also determines the
locus of origin of new shoots. Growth correlations similar to those observed by
RAcIBoRSKI in tropical lianas were obtained.—Raymonp H. Ponp.

LAWRENCE? has published a bulletin containing a summary of our present
knowledge and the results of his own investigations of a serious apple disease
found thus far only in Oregon and Washington, that was first described by
Corpiry*+ under the common name “apple tree anthracnose.” The disease is
also known under the following common names: Oregon canker, black canker,
blackspot apple canker, blackspot, deadspot, and sour sap disease. The present
writer proposes that the disease be known as the “blackspot canker.” The
fungus causing it was described by Corpiry?5 under the name Gloeosporium
malicorticis, and has been described also by PECK?® under the name Macrophoma
curvispora. This fungus is known to injure apple trees seriously in the Pacific
northwest by the production of cankers upon the smaller twigs and branches.
Similar cankers were produced by inoculation from apple cankers upon the
cherry, plum, prune, and peach by Lawrence, and he is inclined to the belief
that the same fungus is responsible for cankers upon all of the hosts named. As
a rule cankers do not appear upon the larger branches and the trunk, owing
apparently to the resistance offered by the corky outer layers of the bark of such
regions to the penetration of the fungus. LAWRENCE here reports for the first
time that the same fungus causes also a dry rot of stored apples that may be
rather serious under certain conditions. The cankers may appear within a week
from the time of infection; ordinarily appearing from November to February,
rapidly completing their development after renewal of growth on the part of the

ost, an oming mature by the next June or July. No definite statement can
yet be made regarding either treatment of the disease or as to possible resistant
varieties. —E. Mrap WILcox.

A MUCH NEEDED INVESTIGATION of the réle of latex in the metabolism of
plants has been commenced by Knrep,?7 and if continued may lead to valuable
results. The paper here cited is presented in four sections. The introduction
carefully reviews the literature, showing that theories are not supported by facts
and that experimental results do not agree. The second section contains experl-

23 LAWRENCE, W. H., Blackspot canker. Bull. Wash. Exp. Sta. 66. pp- 35-
pls. 13. 1904.

24 CORDLEY, A. B., Apple tree anthracnose: a new fungus disease. Bull. Oregon
Exp. Sta. 60. pp. 3-8. pls. I-3. 1900.

25 CORDLEY, A. B., Some observations on apple tree anthracnose. BOT. GAZETTE
30: 48-58. figs. I-12. 1900.

26 Peck, C. H., New species of fungus. Bull. Torr. Bot. Club 27:21- 1g00-

27 Knrtep, Hans, Ueber die Bedeutung des Milchsafts der Pflanzen. Flora
94:129-205. 1905.

ee ee ee oe

1905] CURRENT LITERATURE 379

mental evidence obtained by the author, which establishes that the latex partici
pates very little, if at all, in the translocation of nutrient substances. Seedlings
subjected to conditions intended to force an exhaustion of storage products showed
no difference as to quantity or form of starch grains in the latex as compared with
controls. The author does not regard the latex starch of the euphorbias with
which he experimented as a typical storage product. ScHIMPER’s results th
the latex tubes of the euphorbias do not participate in the depletion of carbo-
hydrates is verified. The third section comprises an anatomical study of thirty-
six species distributed in seven families of laticiferous plants. DrBary’s view
that latex tubes and sieve tubes are reciprocal tissues is rejected. The opinion
of HABERLANDT that latex tubes function as translocation parenchyma is also
discredited. The final section presents an ecological view of the problem, and

exudation of latex, and protection against animals secured by presence of poison-
ous, acrid, or ill-smelling substances. That the problem remains unsolved, and
that the chief value of his paper consists in formulating the present status of
investigation as well as in seit, promising lines of research, is fully realized
by the author.—Raymonp H.

Mitosis in the salamander (Salamandra maculata) has been investigated by
Kowatski’8 under the direction of GREGOIRE. The object of the investigation
was to compare typical animal mitoses with the mitoses in the roots of Trillium
as described by GREGOIRE and WycaeErtTs (see Bot. GAZETTE 38:396. 1904).
Mitosis was studied in various tissues, but principally in the epithelial cells. In
general it may be said that even in minute details the structures and the sequence
of events are essentially the same as in Trillium. In the telophase the action of
the nuclear sap outside the chromosome results in the production of anastomoses

tween the neighboring chromosomes and between parts of the same chromo-
some; while the action of the sap within the chromosome brings about an alveolar
and reticular structure. Each chromosome forms a network of lamellae and
ents of various shapes and sizes which cross each other in every direction.
The entire chromatic network of the nucleus is made up of the individual networks.
The behavior of the chromatin in the prophases is strong evidence in favor of the
individuality of the chromosome. ‘The longitudinal splitting of the chromosome
is not due to the bipartition of a series of chromatic disks imbedded in a sub-
stratum, but consists simply in the longitudinal cleavage of a chromatic ribbon.
No disk spirum is formed at the telophase, nor is any spirem formed at the pro-
Phase. The nuclear membrane is formed by the condensation of the cytoplasm
bordering upon the nuclear vacuole, and at its formation no cytoplasm is included
within the nuclear cavity.—C. J. CHAMBERLAIN.
ee

28 KOWALSKI , J-. Reconstitution du noyau et formation des chromosomes dans
les cinéses somatiques de la larve de Salamandre. La Cellule 21:349-377- pls. I-2
1904.

380 BOTANICAL GAZETTE [way

SEVERAL RECENT INVESTIGATIONS have brought to light a curious condition
in certain plankton diatoms which is believed to be a method of spore formation.
The contents of certain cells break up into a number of small bodies which are
termed “‘microspores.”” These structures have been recently investigated by

STEN?? in a species of Corethron. Their development begins in this form
with a remarkable increase in the number of nuclei in a single diatom, until as
many as 128 are present. The nuclei divide mitotically and the protoplasm succes-
sively until this large number of bodies are found, which however often remain con-
nected with one another by delicate fibrils of protoplasm. KarsTEN believes that
“microspores” which escape from the mother diatom are gametes and unite with
one another in pairs to form a zygospore. Further, there is evidence that the
zygospore gives rise to two new diatoms and that each of these contains at first
two nuclei, one of which afterwards breaks down. If this should be established,
we should have presented in the diatoms a method of sexual reproduction followed
by a type of germination with two successive mitoses apparently of the same
character as in the desmids. It is fair to say, however, that KARSTEN seems to
draw his conclusions from a rather imperfect series of stages.—B. M. DAvIs. °

PELETRIsoT’? has published an extended study of the development of the
seeds of Ericaceae, including in his examination twenty-six genera and sixty-two
species. Such a survey can only mean a glimpse of a general situation, which
subsequent research will develop properly. Aside from anatomical details of
tissues, which can hardly be related as yet, the general morphological situation
seems to be about as follows. The most striking feature of the embryo sac is
the development of a large haustorial pouch at each end. This haustorium varies
much in size in different genera, reaching its maximum in Arbutus; and also in
its degree of separation from the central cavity, sometimes being separated only
by a slight constriction, and in other cases by a narrow, more or less prolonged
neck. In many of the genera there is also a well-developed jacket-layer investing
the central cavity of the sac. The synergids are often beaked and extended into
the micropyle, and the antipodals are ephemeral.

ide from the extent of the survey and the prolix presentation, the most
notable feature of the paper is the absence of all knowledge of any English or

erican publications dealing with similar haustorial developments.—J. M. C

THE PosITION of the monocotyledons is discussed in a recent paper by
Fritscu.3t Since the geological history is so fragmentary, we must rely upon
the evidence of comparative morphology. After discussing this evidence,

29 KarstTEN, G., Die sogenannten “Mikrosporen” der Plankton-Diatomeen und
ihre weitere Entwickelung beobachtet an Corethron Valdiviae. Ber. Deutsch. Bot-
Gesells. 22:544-554. pl. 23. 1904. Ewe
3° PELETRIsOT, C.-N., Développement et structure de la graine chez les Ericacees-
Jour. Botanique 18: 309-367, 386-402. figs. 173. 1904-
3* FritscH, K., Die Stellung der Monokotylen im Pflanzensystem.
Bot. Jahrb. Beiblatt No. 79. pp. 22-40. 1905.

Engler’s

.

1905] CURRENT LITERATURE 381

including recent investigations in anatomy, the conclusion is reached that the
Ranales and Helobiales are related groups; not, however, that any particular
family in one has come from a family in the other, but that both have been differ-
entiated from a common ancestry. HALLIER’s theory that the entire group of
the angiosperms is monophyletic and that it has come from a Ranales-like type,
is, according to FrrtscH, a broader conclusion than is warranted by the present
condition of our knowledge. Since the dicotyledons show some characters more
nearly resembling conditions found in gymnosperms, the dicotyledons should be
sep before the monocotyledons in any scheme of classification. —C. J. CHam-

SHIBATAS? has investigated the chemotaxis of the sperms of Isoetes. He finds
the stimulation limit to be m/20000 malic acid. Of seventy other substances
tested in concentrations of m/10 to M/ roo, only neutral salts of succinic, fuma-
ric, and d-tartaric acids, which are in structure closely allied to malic acid, were
attractive, the lower limit being m/1oo to 5m/1000. WEBER’S law was found
applicable. The relative chemotactic effect of H-ions was found equal in various
mineral acids and salts; similarly, OH-ions from equivalent solutions of alkalies
acted alike. Other interesting results bear on negative chemotaxis of anions,
Se and the nature of the reactions. The notice is a preliminary one.—

oe Eh,

KoERNICKES3 has reviewed several recent cytological papers, thus bringing
his work on “The present condition of cell studies” up to date. The only impor-
tant change of opinion noted is in regard to the occurrence of a reduction division
in plants. The evidence given in the previous paper was against such a division,
but in light of the more recent researches, especially those of FARMER and Moore,
KoernIckeE believes that the evidence in favor of reduction division is gradually
increasing. The work of Witurams, GreGorY, Lotsy, ROSENBERG, BovERI,
Grécorre and Wycaerts, and HACKER are considered in this summary.—
Carters J. eae ROR

WIESNER,3+ who has lready pointed out two cases of the dropping of leaves
which he calls Sokmnossal and Treiblaubjall, adds another, due to high
temperature combined with drouth, which he calls Hitzelaubfall. The death
of the leaves is due to excessive transpiration not covered by the available water.
Naturally the peripheral leaves and especially those reached most directly by the
sun are most affected. Observations are adduced from the author’s extended tour
in the United States during the months of August and September.—C. R. B

3? Surpara, K., Studien iiber die Chemotaxis von Isoetes-Spermatozoiden. Ber.

tu
Deutsch. Bot. Gesells. era 1904.
33 KOERNICKE, M., Die neueren Arbeit
anzenreich und daran pie ethos karyokinetische Probleme.
621: 305-314. 1 990
34 WIESNER, J., Ueber den Hitzelaubfall. Ber. Deutsch. Bot. Gesells. 22: 501-505.
904.

en iiber die Chromosomreduktion im
Bot. Zeitung

382 BOTANICAL GAZETTE [MAY

STEMS OF CALAMITES with attached roots are rare, but several specimens
have been described by MAsten,35 whose account clears up several doubtful
points. The roots in question are mainly adventitious and spring from the
nodes of the aerial as well as the subterranean stems. These roots may be dis-
tinguished from branches among other ways by their point of origin, which how-
ever is different from that of the roots of Equisetum. The writer finds no evi-
dence for connecting the roots with the infranodal organs described by WILLIAM-
son.—M. A. CHRYSLER

GuIGNARD?° has presented further interesting details in connection with
double fertilization in the Malvaceae, using Althaea rosea and Hibiscus Trionum.
Each large and very characteristic pollen grain puts out several pollen tubes,
which advance through the tissues of the style and the nucellus in a peculiar way.
In Hibiscus, when the embryo sac is reached, the tube branches irregularly.
The formation of the generative nucleus and male cells, and the occurrence of
starch are described in detail—J. M. C

Miss SARGANT and Miss ROBERTSONS7 call attention to the fact that the
furrows on the dorsal face of the scutellum in Zea Mais should be regarded as
glands which increase the extent of secreting surface. These glands are situated
in contact with the floury part of the endosperm, and so secure a rapid transfer
of food to the seedling while it as yet has no assimilating organs. An interesting
observation is the presence of amphivasal strands and of transfusion tissue in
the scutellum.—M. A. CHRYSLER.

DRABBLE’s studies on the roots of palms** show that the polystelic condition
found in Areca is not exceptional, but that it commonly occurs in the proxi ximal
region of secondary roots. These separate steles usually fuse in the distal region
of the root to form a single cylinder; certain vascular strands, however, may
remain unfused and constitute the well-known medullary strands. The mono-
stelic central cylinder of palm roots has thus by no means a simple origin.—
M. A. CHRYSLER.

WALLER finds3 that practically no sea plants give blaze currents following
an induction shock, while practically no land plants fail to do so. Resist-
ance of young tissues is very high, and is lowered much by one shock, which

35 MASLEN, - ee The relation of root to stem in Calamites. Ann. Botany
19:61-74. pls. I-2. 1905.

- 36 Guienarp, L., La double fécondation chez les Malvacées. Jour. Botanique
18: 296-308. figs. 16. 1904

37 SARGANT, E., and RoBERTSON, A., The anatomy of the scutellum in Zea
Ann. Botany 19:115-124. pl. 5. 1905.

_ 8 DRapsie, Eric, On the anatomy of the roots of palms. Trans. Linn. Soc.
London. II. Bot. 6: 427-490. pls. 48-51. 1904.

39 WALLER, A. D., On the blaze currents of vegetable tissues. Jour- fot

Bot. 37:32-50. 1904.

Mais

1905] CURRENT LITERATURE 383

probably means that the lacking electrolytes are produced by dissociation upon
passing the induction current. Killed tissues show no blaze currents.— B.

TREBOUX*° in a preliminary paper announces that for chlorophyllous plants
the source of N par excellence is the ammonia radicle, rather than amido-acids.
The significance whereof lies in its relation to a general pate of proteid syn-
thesis to which the author has been led by his experiments.—C. R. B.

Von Spiess shows‘! that the so-called green aleurone does not exist, such
grains being really more or less degenerate chloroplasts. Bluish aleurone grains
occur in maize, the color being due to anthocyan.—C. R. B.

Gatin*? has described cases of polyembryony in two palms, Phoenix canarien-
sis and Pinanga patula, but gives no account of the origin of the embryos.—J. M. C.

BONNIER 43 has compared in detail the embryo sac of gymnosperms and
angiosperms, reaching the conclusion that the most pate tee difference
between the two groups is that in gymnosperms all the eggs formed in an embryo
sac are alike; while in angiosperms the two eggs usually produced are unlike, one
forming the embryo, the development from the other stopping at the proembryo
Stage (endosperm). This is an effort to include double fertilization in con-
trasting gymnosperms and angiosperms, but does not explain anything.—J. M. C.

Miss RIDDLE 44 has studied the Fei of Staphylea trifoliata from the
megasporangiate archesporial cell of the embryo. The account presents no
unusual features, but it is a matter of interest to note that double fertilization was
observed. A is account of some of the forms of Sapindales was muc
needed.—J. M

4° TrEBouX, O., Zur Stickstoff-Ernahrung der griinen Pflanzen. Ber. Restart.
Bot. Gesells. 22: 570-372. 1905.

41 Von ee Kart, Ueber die Farbstoffe des Aleurone. Oesterr. Bot. Zeits.
54:440-446. 1904

42 Gatin, C. L., Ee a cas de polyembryonie chez plusieurs espéces de palmiers.
Rey. Gén. Botanique 17:60

43 BONNIER, GASTON, halen sur la comparaison entre les ai aa
les gymnospermes. Rev. Gén. Botanique 17:97-108. figs. 6. 1905.

44 RippLe Lumrna C., Development of the embryo sac iia embryo of Staphylea

drijoliata. Ohio Naturalist 5: 320-325. pls..19-20. 19095-

NEWS.

Proressor F. W. Oxtver, University College, London, has been elected a
Fellow of the Royal Society.

Dr. E. Prrrzer, University of Heidelberg, has been elected associate member
of the Royal Botanical Society of Belgium.

ALFRED Ernst, professor of botany at Ziirich, has received from the govern-
ment $1000 for botanical studies at Buitenzorg.

Dr. GEORGE BITTER, docent in botany at Miinster, has been appointed direc-
tor of the Botanical Garden at Bremen.—Science.

ProFEssor Huco DeVries, University of Amsterdam, has been elected asso-
ciate member of the Royal Academy of Belgium.

W. A. KELLERMAN has returned from a three months’ exploration of Guatemala
with a large amount of material, especially parasitic fungi.

Proressor E. C. Jerrrey, Harvard University, has received a grant of $200
from the Elizabeth Thompson Science Fund ‘‘for the study of cupressineous
conifers.”

A. S. Hircucocx, U. S. Department of Agriculture, has been transferred to
the Office of Botanical Investigations and Experiments, his position being styled
“systematic agrostologist.”

PROFESSOR THEOPHILE DURAND, director of the Botanical Garden at Brussels,
and Professor JEAN Massart, University of Brussels, have been elected corre-
sponding members of the Royal Academy of Belgium.

F. E. Luoyp, Teachers College, Columbia University, has received a grant
of $500 from the Carnegie Institution to aid him in continuing his studies on
stomatal action and transpiration in desert plants. He spends three months
at the Desert Botanical Laboratory for this purpose.—Science.

Ir Is ANNOUNCED by Science that Professor Douctas H. CAMPBELL ee
spend next year in an extensive trip through Europe, Africa, and Asia. He w
visit the regions about Victoria and Zambesi Falls; the botanical gardens
at Paradeniya and Buitenzorg; and will then return by way of the Philippine
Islands

LITHIA Wi

Remedy of Ordinary Merit Could Ever
Have Received Indorsations from |
Men Like These.

Ss 1 o. F otter, = - 7s R.C.P.,
Professor of ie Principe ee reo P. London, I
Zeons, Son Francie, es

m. H. D: Souttnisuns Prof Medical Juri.
hn Bright's Disease Buoy fa Poe Papers Cas ewe ah J tim.

se and C. Edson, A. M., M. D., Health Comm clistemnd Wout
Albuminuria York City and State, President Board of | Pharmacy, New York
4 City, Examining Physician Corporation
BRS apohn V. Shoemaker, M. Day LL, De Prefor Nalria
, 5 y Dr. George Ben. Johnston, Pikssiad: ae Re Pretied

Southern Surgical and Gynecological Association, Ex-President .
Eeteonl Sasoy ay Tes ra of Caigy od dado
oul Bis Medical Ca. BF OEY Ge

Materia Medica of the Faculty of Medicin gpg =
Dr. J.T. t Bla ca ee Prof. Me i? oh te CMF ESN VO. :

i = Bg}, Cc. Horn, M. Di, Ph Ph. DB.» Pr Projessor Diseases of ae
: én @ } /

: 4 E phat vod Prt Neng an

Dr. J. Allison Hedges,” — of ied I Vee < :

Dr. I N. Love, 1¥ New York City, ‘ormer Profes
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The Botanical Gazette

A Monthly Journal Embracing all Departments of Botanical Science

Edited by JonN M. CouLrer ane CHARLES R. BarRNEs, with the oom of other members of the
. botanical staff of the University of Chic

Jol. XXXIX, No. 6 Issued June 22, 1905

CONTENTS

Becummous RUSTS FROM MEXICO, J..€. Apia. (2 6 ees ea ge

ON THE WATER-CONDUCTING SYSTEMS OF SOME DESERT PLANTS (WITH
TEN FicuRES). W. A. Cannon - - = SOF

1 #E EFFECTS OF TOXIC AGENTS UPON THE ACTION OF BROMELIN. Conrrt-

; BUTIONS FROM THE Hurt BoranicaL Laporatory. LXXIII. Joseph Stuart Caldwell - 409
BRIEFER ARTICLES.
THe Earty History or ANciosperMs. Ethel Sargant - - - - - - 420

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VOLUME XXXIX NUMBER 6

DOTANICAL-. GAZEI Te
JUNE, 1905

LEGUMINOUS RUSTS FROM MEXICO.
J. C. ARprwuR.

THE following thirty-seven species of Uredinales, represented by
128 numbered collections, are, with two exceptions, part of the rich
fruits of three vacation trips into Mexico, made by Mr. E. W. D.
HOLWAY in 1808, 1899, and 1903. Some of the material secured
on these excursions has already been reported upon by Mr. Hotway
in an article in this journal (31: 326-338. 1901), which included
descriptions of thirty-one new species; a still earlier trip, made in
1896, was also reported by Mr. Hotway in this journal (24:23-38.
1897), in which forty-five new species of Uredineae were described,
the descriptions being drawn by Dr. P. DreTeEt, and with two species
of ascomycetous fungi, one being the type of a new genus. Much
material remains to be studied, which will undoubtedly yield many
more new species.

The amount of excellent material secured by Mr. Hotway within
the limited periods at his command shows him to be a collector
of unusual activity and acumen. His specimens are ample, and
as a rule contain the various spore stages, or all that could reasonably
be expected to occur at one season of the year. Another most com-
mendable feature in making these collections is the care taken to
Secure the exact determination of the host whenever feasible. Not
only are bits of the inflorescence and fruit included, but whenever
the host is in any way unfamiliar, specimens suitable for the phanero-
gamic herbarium are taken illustrating the foliage, flowers, and fruit
of the plants as well as possible. These are given the same numbers
as used for the fungi found on them. They are subsequently sent
to specialists for determination, by which means accuracy of naming

385

386 BOTANICAL GAZETTE [JUNE

is insured; incidentally also the leading herbaria of the country have
been enriched with many valuable specimens, while not a few new
species of flowering plants have been brought to light.

The collections made by Mr. Hotway show that Mexico has an
abundant rust flora, and especially so in the groups represented by
the genera Ravenelia and Uropyxis. If collections made during
September, October, and November, and over a comparatively
small part of that great country have proved so interesting and val-
uable, more extended search is likely to be rewarded with almost
or quite equal results for some time to come.

The following enumeration includes the portion of Mr. Hotway’s
uredineous collections in Mexico, not heretofore published, possessing
leguminous hosts, 7. ¢., the rusts occurring upon members of the
families Fabaceae, Cassiaceae, and Mimosaceae.

1. Uromyces Triroti (A. & S.) Levy.—On Trifolium amabole
HBK., Jalapa, Oct. 3, 1898, no. 3091: Trifolium sp., Pachuca,
Oct. 27, 1903, no. 5245; City of Mexico, Oct. 28, 1899, no. 3746.

2. Uromyces MEDICAGINIS-FALCATAE Wint.—On Medicago
denticulata Willd., Toluca, Sept. 19, 1898, no. 3129.

3. Uromyces rugosa, n. sp.—Uredosori hypophyllous, cinnamon-
brown; uredospores globoid, 20-24 by 21-27; wall light yellow,
medium thick, 2-2.5, minutely and sparsely verrucose, pores 8,
scattered: teleutosori hypophyllous, small, round, 1™™ or less across,
scattered or somewhat gregarious, soon naked, pulverulent, dull
chocolate-brown; teleutospores broadly oval or globoid, 18-22 by
21-27, wall dark chestnut-brown, medium thick, 2-3, rugose Or
irregularly and closely verrucose, apex usually bearing a very low,
hyaline umbo; pedicel colorless, short, fragile, mostly deciduous.
—On Lupinus sp., Amecameca, Oct. 21, 1903, no. 5208.

The very rough and opaque spores easily separate this species from all others
reported from North America on Lupinus. The uredo stage was not well shown
on the specimens studied, and is consequently incompletely described.

4. Uromyces montanus, n. sp.—Teleutosori hypophyllous, small,
round, crowded in circinating groups, 5-10™™ across, centripetal
in development, early naked, cinnamon-brown or darker, usually
cinereous from germination; . teleutospores oval or obovate, pale,
18-24 by 30-40, wall very pale brownish, smooth, thin, 1-1-5":

1905] ARTHUR—RUSTS FROM MEXICO 387

thickened at apex, 5-7, pedicel colorless, thick, as long as the spore
or shorter— On Lupinus mexicanus HBK., Nevada de Toluca,
10,400 f* alt., Oct. 16, 1903.

This collection also showed well-developed aecidia of Uromyces Lupini B. & C.
upon the same leaves. The teleutospores germinate in the sorus upon maturity.

5. Uromyces Lupint B. & C.—On Lupinus elegans HBK.,
Toluca, Sept. 19, 1898, no. 3177: Lupinus sp., Pachuca, Oct. 5,
1899, no. 3575; Zapotlan, Oct. 9, 1903, no. 5140; Nevada de Toluca,
Oct. 15, 1903, no. 5154; Amecameca, Oct. 21, 1903, no. 5209.

6. Uromyces Cologaniae, n. sp.—Uredosori hypophyllous, scat-
tered, round, small, o.25™™ or less across, soon naked, pulverulent,
pale cinnamon-brown, ruptured epidermis barely noticeable; ure-
dospores globoid, 16-23 by 20-274, wall pale cinnamon-brown,
thin, 1.5-24, evenly and minutely echinulate, pores 3-4, equatorial:
teleutosori hypophyllous, scattered, round, small, 0.25 ™™ across,
soon naked, pulverulent, cinnamon-brown, ruptured epidermis
barely noticeable; teleutospores oval or globoid, 16-20 by 18-26,
rounded at both ends, wall cinnamon-brown, thin, 1.5-2#, closely
and finely verrucose, apex a little thicker or with a very low, pale
umbo, 2-4#; pedicel colorless, short, fragile; mostly deciduous.—
On Cologania pulchella HBK., Patzcuaro, Oct. 20, 1898, no. 3192
(type); Toluca, Sept. 20, 1898, no. 3179; Uruapam, Oct. 11, 1899,
no. 3615: C. congesta Rose, Toluca, Sept. 20, 1898, no. 3179: C.
ajinis Mart. & Gal., Jalapa, Oct. 3, 1898, no. 3193: Cologania sp.,
Oaxaca, Oct. 21, 1899, no. 3703.

The determination of the several hosts was made by Dr. J. N. Rose of the
U. S. National Museum, all with some slight question, as the material was far
from complete.

7. UROMYCES APPENDICULATUS (Pers.) Unger—On Phaseolus
vulgaris L., City of Mexico, Oct. 15, 1898, no. 3049; Atequiza, State
of Jalisco, Oct. 6, 1903, no. 5120: P. retusus Benth., near Tula,
Sept. 20, 1898, no. 3161: P. anisotrichus Scheele, Rio Hondo, near
City of Mexico, Sept. 22, 1898, no. 3155: P. disophyllus Benth.,
Chapala, Sept. 18, 1899, no. 3455: P. coccineus Jacq., City of Mexico,
Oct. 9, 1898, no. 3037; Dos Rios, near City of Mexico, Sept. 22,
1898, no. 3094; Uruapam, Oct. 12, 1899, no. 3625: P. obvallatus
Scheele, Santa Fé, near City of Mexico, Oct. 18, 1903, nO. 5179;

388 BOTANICAL GAZETTE [JUNE

Amecameca, Oct. 20, 1903, no. 5188: P. atropurpureus Moc., Iguala,
State of Guerrero, Nov. 4, 1903: Phaseolus sp., Cuernavaca, Sept.
24, 1898, no. 3021; Cuautla, Oct. 12, 1898, no. 3070, and Oct. 22,
1903, no. 5217; near Tula, Sept. 21, 1898, no. 3187; Etla, State of
Oaxaca, Oct. 23, 1899, no. 3729; Guadalajara, Sept. 22, 1903, no.
5021, Iguala; Nov. 4, 1903, no. 5336.

The Mexican collections of this species of rust do not vary in any noticeable
way from those made in the United States. Uromyces obscurus D. & H. was
founded on a Phaseolus rust having what was taken to be a peculiar form of
Aecidium, but which proves to be a species of Synchytrium. This name is there-
fore a synonym of U. appendiculatus.

8. Uromyces Fasas (Pers.) DeB.—On Faba vulgaris L., Toluca,
Sept. 17, 1898, no. 3189; Patzcuaro, Oct. 13, 1899, no. 3628; and
Oaxaca, Oct. 24, 1899, no. 3733.

g. UromycEs TENuISTIPES D. & H.—On Meibomia sp., Dos
Rios, near City of Mexico, Sept. 22, 1898, no. 3095; Toluca, Sept.
19, 1898, no. 3097; Cardenas, Oct. 22, 1898, no. 3148; Atequiza,
State of Jalisco, Sept. 26, 1899, no. 3503.

The teleutospores are minutely rugose, although they appear smooth when
wet. The apex of the spore has a very low, semihyaline umbo, which often
disappears, leaving a shallow depression, as mentioned in the original description
(Bor. Gaz. 24:25. 1897).

10. URoMYCES MExIcANUS D. & H.—On Meibomia sp., Cuer-
navaca, Sept. 24, 1898, no. 3019, and Sept. 28, 1898, no. 3124; near
Tula, State of Hidalgo, Sept. 21, 1898, no. 3195; Cardenas, State of
San Luis Potosi, Oct. 22, 1898, no. 3150; Rio Hondo, near City of
Mexico, Sept. 22, 1898, no. 3154; Chapala, Sept. 18, 1899, no. 3447-

11. Uromyces HEpySARI-PANICULATI (Schw.) Farl.—On Meibo-
mia amplijolia (Hemsl.) Kuntze, Patzcuaro, Oct. 20, 1898, no. 3141:
M. elegans (Desv.) Kuntze, Rio Hondo, near City of Mexico, Oct.
4, 1899, no. 3569: M. strobilacea (Schl.) Kuntze, Guadalajara,
Oct. 12, 1903, no. 5146: Meibomia sp., City of Mexico, Oct. 14
1898, no. 3048; Cuernavaca, Sept. 29, 1899, no. 3518 with uredospores
only; Morelia, Oct. 14, 1899, no. 3633; Oaxaca, Oct. 18, 1899, NOS:
3658, 3659, Oct. 23, 1899, nos. 3716, 3723, Nov. 11, 1903, no. 53853
Santa Fé, near City of Mexico, Oct. 18, 1903, no. 5165; Tlalpam,
near City of Mexico, Oct. 19, 1903, no. 5180.

1905] ARTHUR—RUSTS FROM MEXICO 389

12. Uromyces Clitoriae, n. sp.—Uredosori hypophyllous; uredo-
spores globoid, about 23-27# in diameter, wall thick, 3, fuscous,
evenly and sparsely echinulate, pores apparently 2: teleutosori hypo-
phyllous, small, round, scattered, dark cinnamon-brown, soon naked;
teleutospores ellipsoid, 20-24 by 22-30#, wall chestnut-brown,
evenly thick, 33.5, minutely and evenly verrucose; pedicel delicate,
colorless, less than the length of the spore, caducous.—On Clitoria
mexicana Link, Jalapa, Oct. 5, 1898, no. 3088.

Only a few uredospores were seen, which were found in- parasitized sori.
The sculpturing of the teleutospores is minute but very distinctly shown when

13. Uromyces bauhiniicola, n. sp.—Spermogonia epiphyllous,
gregarious, punctiform, fuscous, subepidermal, seen in section
globose, 60-130 in diameter: teleutosori at first hypophyllous,
becoming amphigenous, small, round, about 1™™ in diameter,
often confluent, early naked, blackish; teleutospores broadly oval
or globoid, 15-21 by 20-24; wall dark chestnut-brown, semi-
opaque, thick, 2.5-—3.5, prominently rugose, thicker and paler at
apex, 5-7#; pedicel colorless, firm, as long as the spore——On Bau-
hinia Pringlei Wats., Guadalajara, Sept. 28, 1903, no. 5060 (type):
Bauhinia sp., Iguala, Nov. 4, 1903, no. 5334-

Readily distinguished from the four or five species occurring upon Bauhinia
in South America by the very dark, rugose spores, unaccompanied by uredo-
spores. The spores germinate somewhat readily in the sori, not long after
maturity.

14. PHRAGMOPYXIS DEGLUBENS (B. & C.) Diet.—On Brit-
tonamra Edwardsii (A. Gr.) Kuntze (Cracca Edwardsii A. Gray),
Cuernavaca, Oct. 30, 1903, no. 5278.

This is the first time the host of this rare and interesting species has been
definitely determined. It has only twice been reported before, once in Ecuador
on an undetermined species of Coursetia, a genus closely related to Brittonamra.
The type collection is recorded on “leaves of some leguminous plant,” and is
said to come from Texas (Grev. 3:55), which latter, however, must be an error.
The specimen in the cryptogamic herbarium of Harvard University says, “ North-
ern Sonora,” and bears the collector's number, the same as recorded by BERKELEY
in Grevillea. A specimen in the herbarium of the Department of Agriculture at
Washington, D. C., says “N. Mexico,” but does not bear the collector's number.
Both of these specimens are undoubtedly part of the one original collection made
by C. Wright. The Washington specimen consists of two leaflets, 5 by 12™™,

390 BOTANICAL GAZETTE [JUNE

elliptical, entire, nearly smooth above, slightly hirsute below with scattered hairs,
mucronate, short petioluled, texture firm, bearing a few, well-formed, amphi-
genous teleutosori. The specimen was compared, under the guidance of Dr.
J. N. Rose, with specimens in the phanerogamic herbarium of the U. S. National
Museum, but could not be matched.

The collection by Mr. Hotway is furthermore noteworthy in possessing
aecidia, not heretofore known, which are rather pale and inconspicuous. ey
may be described as follows: Aecidia hypophyllous, or somewhat amphigenous,
crowded in small groups, pale yellow, without peridium, but sparingly encircled
by incurved paraphyses; aecidiospores catenulate, globoid or globose-oblong,
13-17 by 15-19, wall very pale yellow, thin, 1, finely and closely verrucose,
paraphyses small, cylindrical, 5-7 by 24—30#, more or less contorted, wall usually
thick, sometimes nearly obliterating the lumen, smooth, colorless: spermogonia
epiphyllous, few in groups opposite the aecidia, brownish-yellow, punctiform,
inconspicuous, subcuticular, conical, small, 60-80u broad and half as high, ostiolar
filaments 20-24» long, free.

CALLIOSPORA, nov. gen.—Teleutosori arising from beneath
the epidermis, soon naked; teleutospores 2-celled by transverse
partition, wall colored, with an external layer which swells in water,
germ pores 2 in each cell, lateral. Aecidium and uredo wanting.
Spermogonia arising from beneath the cuticle, conical.

15. Calliospora Holwayi, n. sp.—Spermogonia epiphyllous, numer-
ous over areas 2-7™™ across, punctiform, golden yellow, becoming
brown, subcuticular, conical, 80-125 broad: teleutosori epiphyllous,
scattered, sometimes confluent, small, round, blackish-brown; teleuto-
spores elliptical, 26-34 by 40-514, rounded at both ends, slightly
or not constricted at the septum, inner wall chocolate-brown, thick,
3-4#, pores two in each cell, outer wall colorless, 2-3 thick in water,
smooth; pedicel colorless, as long as the spore, swelling in water to
the diameter of the spore and bursting—On Eysenhardtia amor-
phoides HBK., Guadalajara, Sept. 28, 1903, no. 5059 (type), Sept-
29, 1903, no. 5068; Oaxaca, Oct. 25, 1899, no. 3737: E. orthocar pa
Wats., Etla, State of Oaxaca, Nov. 13, 1903, nos. 5404 and 5495.

This species resembles Uropyxis Eysenhardtiae (D. & H.) Magn., but differs
not only in the absence of the uredo,but in the position of the sori, the measure-
ments of the teleutospores, and their smooth, colorless, outer wall. The collec-
tions have been examined by Dr. J. N. Rosr, who says that the species of .
Eysenhardtia have not been sufficiently studied, and some doubt must attach
to the determination of the several numbers.

1905] ARTHUR—RUSTS FROM MEXICO | 391

10. Calliospora Farlowii, n. sp.—Spermogonia caulicolous, and
sparingly on midrib of leaf, small, punctiform, yellowish-brown,
conical, 80-100 broad, ostiolar filaments 25-40u long: teleutosori
caulicolous, large, confluent, early naked, pulverulent, cinnamon-
brown; teleutospores elliptical, 18-24 by 29-42, rounded at both
ends, slightly or not constricted at the septum, inner wall cinnamon-
brown, 2-3 thick, pores 2 in each cell, outer wall colorless, barely
noticeable when swollen in water, minutely verrucose; pedicel color-
less, about 6# thick, short and mostly deciduous——On Parosela
domingensis (DC.) Heller ae domingensis DC.), Orizaba, Feb.,
1888 (W. G. Farlow).

I am indebted to the collector for excellent material of this interesting species.
In the selection of the specific name I take the opportunity of showing my appre-
ciation of courtesies in sending me material, and also my high regard for the
eminent services rendered by Dr. FARLow to botanical science.

17. Calliospora Diphysae, n. sp.—Spermogonia amphigenous,
crowded in dendritic groups, or disposed in lines, pale brown, bullate
or hemispherical, subcuticular, too-175# broad and less than half
as high: teleutosori amphigenous and caulicolous, round or elongated,
I~5™™ long, early naked, nearly black; teleutospores elliptical,
30-33 by 45~50#, rounded at both ends, not constricted at septum,
inner wall dark chestnut-brown, 3-4 thick, pores 2 in each cell,
outer gelatinous, pale amber-color, 3-4 thick in water, sparsely and
evenly verrucose; pedicel half length of spore or longer, colorless,
firm above, at base slightly bulbous and swelling in water to bursting.
—On Diphysa suberosa Wats., Rio Blanco, near Guadalajara,
Sept. 30, 1903, no. 5082.

18. Uropyxis Nissourar (D. &. H.) Magn.—On WNissolia
laxior Rose, Guadalajara, Sept. 25, 1903, no. 5039: N. hirsuta DC.,
Sayula, State of Jalisco, Oct. 8, 1903, no. 5127; Cuautla, State of
Morelos, Oct. 2 3, 1903, no. 5230: Nissolia sp., Cuernavaca, Nov. I,
1903, NO. 5309.

These collections, except the one made in November, show a few ibileonpoces
among the teleutospores. The uredospores are globoid or ellipsoid, pale yellow
or nearly colorless, 12-16 in diameter, wall very thin, 0.75, finely verrucose,
pores very indistinct, few. The paraphyses seen are capitate, thin- walled, and
colorless.

392 BOTANICAL GAZETTE [JUNE

19. Uropyxis DALEAE (D. & H.) Magn.—On Parosela Dalea
(L.) Britt. (Dalea alopecuroides Willd.), Guadalajara, Sept. 26, 1903,
no. 5048: P. mutabilis (Willd.) Rose, Toluca, Sept. 19, 1898, no.
3188; Amecameca, Oct. 20, 1903, no. 5196, Nov. 20, 1903, no. 5426:
P. acutijolia (DC.) Rose, Cuautla, State of Morelos, Oct. 23, 1903,
no. 5227: P. Holwayi Rose, Iguala, State of Guerrero, Nov. 3, 1903,
no. 5319: P. trifoliata Rose, near Tacubaya, City of Mexico, Oct. 7,
1896, without number; Aguascalientes, Oct. 9, 1903, no. 7705 (Rose
& Painter).

20. Uredo Aschynomenis, n. sp.—Sori mostly hypophyllous,
grouped on small reddish spots, or scattered, subepidermal, small,
globose, with pseudoperidium formed of imbricated paraphyses by
the union of the slender stipes, the free capitate ends lining the inner
wall, dehiscent by a central orifice; uredospores stylate, broadly
elliptical or globoid, 14-18 by 16-23, wall pale yellow, thin, 14,
very minutely verrucose, pores obscure, about 6, scattered.—On
Aischynomene americana L.., Cuautla, State of Morelos, Oct. 22,
1903, NO. 5220.

The spores of this collection are much like those of Uropyxis Nissoliae,
the hosts of the two species being closely related. But no genus of leguminous
rusts known to the writer contains species with a pseudoperidium for the uredo.
Until the teleutospores are discovered, it will be impossible to state the affinities
of the species.

21. Ravenelia Lysilomae, n. sp.—Spermogonia epiphyllous,
crowded in small groups, punctiform, rather large, pale brownish-
yellow, subcuticular, flattened hemispherical, 80-130” wide, one-
fourth as high: teleutosori epiphyllous on pale spots, becoming
amphigenous, small, o.25-0.5™™ across, chestnut-brown, subepi-
dermal, numerous hyphoid paraphyses intermixed with the spores,
inconspicuous; teleutospore-heads chestnut-brown, 7-9 cells across,
80-120 in diameter, smooth, cysts appressed beneath the head,
extending from periphery to pedicel, united laterally; pedicel color-
less, short, deciduous; paraphyses cylindrical, 7-10 by 30-49A,
light golden-brown, wall thin or rarely thick—On Lysiloma ter
gemina Benth., Iguala, State of Guerrero, Nov. 3, 1903, NO. 5317-

22. RAVENELIA VERRUCOSA Cke. & Ell—On Leucaena micro-
phylla Benth., Iguala, State of Guerrero, Nov. 3, 1903, 00. 5314-

1905] ARTHUR—RUSTS FROM MEXICO 393

This collection differs slightly from the type in havi ing heads almost or quite
smooth, and with paraphyses fewer and paler.

23. RAVENELIA LEUCAENAE Long.—On Leucaena diversijolia
Benth., Etla, State of Oaxaca, Nov. 13, 1903, no. 5408: L. esculenta
DC., Etla, State of Oaxaca, Nov. 1 3, 1903, no. 5413; Iguala, State
of Guerrero, Nov. 3, 1903, no. 5311: Leucaena sp., Guadalajara,
Sept. 25, 1903, no. 5044; and Tehuacan, State of Puebla, Nov. 7,
1903, NO. 5340.

There appears to be considerable uncertainty in the exact determination of
the species of Leucaena. Although these numbers have been submitted to com-
petent i the correctness of the specific names employed here cannot
be vouched for.

24. RAVENELIA EXPANSA Diet. & Holw.—On Acacia filiculoides
(Cav.) Trel. (A. filicina Willd.), Iguala, State of Guerrero, Nov. 3,
1903, no. 5312: A. cochliacantha Humb. & Bonp., Iguala, State of
Guerrero, Nov. 3, 1903, no. 5315; Tehuacan, State of Puebla, Nov.
8, 1903, no. 5353: Acacia sp., Yautepec, Oct. 24, 1903, no. 5237.

25. RAVENELIA sILIQUAE Long.—On Acacia pennatula Benth.,
Etla, State of Oaxaca, Nov. 14, 1903, NO. 5395.

The type was seen by Mr. Lone only on the fruit, but the present collection,
Which does not differ in any perceptible way from the type, only shows rust on
the leaves. No teleutospores could be found.

26. Ravenelia gracilis, n. sp—Spermogonia amphigenous, puncti-
form, crowded in groups, prominent, subcuticular, hemispherical,
60-100# broad and half as high: uredosori epiphyllous, scattered,
less than o.25™™ in diameter, nearly round, mamillose, dehiscent
by central, irregular rupture, the encircling epidermis forming an
aecidioid cup, paraphyses peripheral, or also intermixed with the
Spores; uredospores elliptical or obovate-oblong, 16-21 by 30-404,
wall rather thin, 1.5-2.5, golden-brown, strongly and evenly echinu-
late, pores 4-6, equatorial; paraphyses hyphoid, 7-10 by 40-604,
Somewhat contorted, smooth, thin-walled, nearly or quite colorless:
teleutosori like uredosori, but without paraphyses; teleutospore-

eads chestnut-brown, 5—7 cells across, 75-100” in diameter, each
Spore bearing 4~7 slender, nearly colorless tubercles, 3-4 high;
cysts delicate, appressed beneath the head, extending from periphery
to pedicel, united laterally; pedicel short, colorless, deciduous.—

304 BOTANICAL GAZETTE [JUNE

On an undetermined species of Mimosaceae, Cardenas, State of
San Luis Potosi, Oct. 22, 1898, no. 31444.

This rust has many characters in common with that on Pithecolobium,
although the hosts do not appear to be closely related. The host is a thorny
shrub or tree, with evenly twice pinnate leaves; the leaflets oblong, nearly glab-
rous, entire, about 3 by 7™™. A cupulate gland usually occurs on the main
rachis at the insertion of each pair of petioles.

27. Ravenelia Pithecolobii, n. sp——Uredospores in the teleutosori
elliptical or broadly oval, 15-18 by 24-30, wall golden-yellow,
medium thick, 2-3, thicker at apex, 3-5“, evenly verrucose, pores
4-6, equatorial: teleutosori amphigenous, small, round, at first
bullate, scattered, subepidermal, chestnut-brown; teleutospore-heads
chestnut-brown, 6-8 cells across, 70-go# in diameter, each spore
bearing 2-3 slightly curved tubercles, 5-7 long, acute, pale brownish;
cysts appressed beneath the head, extending from periphery to
pedicel, united laterally ; pedicel colorless, short, deciduous ; paraphyses
none.—On Pithecolobium dulce (Roxb.) Benth., Guadalajara, Sept.
27, 1903, NO. 5051.

28. RAVENELIA MIMOSAE-SENSITIVAE Henn.—On Mimosa stipt-
tata Rob., Cuautla, Morelos, Oct. 28, 1903, no. 5228; and Iguala,
Nov. 4, 1903, no. 5326: M. caerulea Rose, Cuernavaca, Oct. 30, 1903,
no. 5290: M. Galeottii Benth., Cuernavaca, Oct. 31, 1903, no. 5303:
M. polyanthoides Rob., Iguala, Nov. 3, 1903, no. 5324: M. albida
H. & B., Cuernavaca, Sept. 24, 1898, no. 3125, Oct. 29, 1903, nO.
5265; and Cuautla, Oct. 20, 1903, no. 5213: M. alba floribunda
Rob., Oaxaca, Nov. 10, 1903, no. 5368; Etzatlan, State of Jalisco,
Oct. 2, 1903, no. 5086: Mimosa sp., Etla, State of Oaxaca, Nov. 16,
1903, no. A.

This species appears to be remarkably uniform on all the hosts so far reported
for it. Uredo sensitiva Speg. (An. Mus. Nac. Buenos Aires 6: 236. 1899) 1s
probably a synonym, judging from the description. All the numbers under
this species were submitted to Dr. B. L. Roprnson of the Gray Herbarium, who
determined the hosts.

29. RAVENELIA CASSIAECOLA Atk.—On Cassia sp., Oaxaca,
Nov. 10, 1903, no. 5359.

This species has not before been reported from Mexico, having been known
only from the southeastern United States.

1905] ARTHUR—RUSTS FROM MEXICO 3905

30. Ravenelia inconspicua, n. sp.—Uredosori hypophyllous,
small, o.5™™ or less across, subcuticular, soon naked, pulverulent,
ruptured cuticle inconspicuous, paraphyses abundantly intermixed
with the spores; uredospores globose or broadly elliptical, 15-20 by
15-22, wall medium thick, 2-2.5, golden-yellow, closely verrucose,
pores about 10, scattered, paraphyses spatulate, smooth, cinnamon-
brown, paler below, walls thick, 3-6#: teleutosori hypophyllous,
very small, scattered, blackish, shining, subcuticular; teleutospore-
heads chestnut-brown, 6-8 cells across, 60-100 in diameter, each
cell bearing 4-6 cylindrical tubercles, 3# wide by 5-7» long, slightly
colored; cysts appressed to the under side of the head, 6-10, some-
what united to one another; paraphyses none.—On Cassia (or
Caesalpinia) sp., Zapotlan, State of Jalisco, Oct. 9, 1903, no. 5135-

31. RAVENELIA SPINULOSA Diet. & Holw.—On Cassia Holwayana
Rose (C. multiflora Mart. & Gal.), Cuautla, State of Morelos, Oct-
23, 1903, no. 5226; Oaxaca, Nov. 11, 1903, no. 5390: C. Galeottiana
Mart., Tehuacan, Nov. 7, 1903, no. 5348: Cassia sp., Oaxaca,
Noy. 10, 1903, no. 5372; Cuernavaca, Nov. 1, 1903, no. 5310.

32. Ravenelia pulcherrima, n. sp.—Uredosori amphigenous, in
loose groups or scattered, round, small, 0.25-0.5™™ across, soon
naked, subcuticular, cinnamon-brown, paraphyses numerous, inter-
mixed with the spores; uredospores ellipsoid or oblong-globoid,
15-18 by 17-24, wall thin, 1.5-2#, pale golden yellow, finely and
closely echinulate-verrucose, pores 6-8, scattered, paraphyses large,
capitate or spatulate, 12-18 by 35-55, smooth, walls of the stipe
thin and nearly colorless, walls of the head much thickened above,
3-10, chestnut-brown: teleutosori amphigenous, small, scattered
or confluent, subcuticular, blackish-brown; teleutospore-heads choco-
late-brown, 6-7 cells across, 75-120 in diameter, each spore bearing
4-5 inconspicuous papillae, semihyaline, 1-2# high; cysts appressed
to the underside the head, extending from periphery to pedicel;
paraphyses none; pedicel short, colorless, deciduous.—On Poinciana
pulcherrima L. (Caesalpinia pulcherrima Sw.), Yautepec, State of
Morelos, Oct. 24, 1903, no. 5236 (type); Iguala, State of Guerrero,
Noy. 4, 1903, no.

33. RAVENELIA LAEVIS Diet. & Holw.—Or Indigojera jaliscen-
sis Rose, Chapala, Oct. 5, 1903, no. 5108: I. densiflora Mart. & Gal.,

396 BOTANICAL GAZETTE [JUNE

Cuernavaca, Sept. 26, 1898, no. 3225, Oct. 30, 1903, no. 5292; Oax-
aca, Nov. 11, 1903, no. 5380; Amecameca, Oct. 20, 1903, no. 5203;
Santa Fé, near City of Mexico, Oct. 18, 1903, no. 5167: Indigofera
sp., Etzatlan, State of Jalisco, Oct. 2, 1903, no. 5091.

34. RAVENELIA INDIGOFERAE Tranz.—On Indigojera cuernava-
cana Rose, Cuernavaca, Oct. 30, 1903, no. 5296: I. Conzaitii Rose,
Mt. Alban, State of Oaxaca, Nov. 12, 1903, no. 5392.

35. RAVENELIA BRONGNIARTIAE Diet. & Holw.—On_ Brong-
niartia podalyrioides HBK., Iguala, State of Guerrero, Nov. 3, 1993,
no. 5310.

36. RAVENELIA SIMILIS (Long).—On Brongniartia podalyrioides
HBK., Guadalajara, Sept. 30, 1903, no. 5181.

37. RAVENELIA TALPA (Long).—On Cracca Talpa Rose (Teph-
rosia Talpa Wats.), Guadalajara, Oct. 12, 1903, no. 5151: C.
macrantha (Wats.) Rose (Tephrosia macrantha Wats.), Etzatlan,
State of Jalisco, Oct. 2, 1903, no. 5100.

The collection on C. macrantha shows some deviation from the type, the
tubercles on the teleutospore-heads being larger, and the uredospores being
slightly smaller, with germ pores more often scattered.

PURDUE UNIVERSITY,
Lafayette, Ind.

SSS ean ete

ON THE WATER-CONDUCTING SYSTEMS OF SOME
DESERT PLANTS.*
W. A. CANNON.
(WITH TEN FIGURES)

A STATEMENT by VOLKENS® to the effect that the ducts of the
stems of the shrubs and trees of the Egyptian-Arabian deserts are
relatively poorly developed led to an examination of these elements
in the shrubs and trees which are growing within reach of the Desert
Botanical Laboratory. It appeared at once that our native plants
are not poor in water-conducting tissue; on the contrary they are
very well provided for in this regard, and I cast about for some means
of expressing this condition of affairs from a comparative standpoint.
It was especially desired to compare these plants with those of the
deserts of Egypt and Arabia, but since the direct comparison was
impracticable, because VOLKENS does not give exact data, I have
sought a way out of the difficulty by using irrigated desert plants
of the species studied as controls. I assume that the desert plants
if provided with an abundance of water are for practical purposes
mesophytes, and if VoLKENs had in mind the plants of middle Europe,
Which might well have been the case, we have in this roundabout
Manner a means of instituting the desired comparison. Aside from
this, the great difference in the development of the conductive tissue
between desert forms that have been irrigated and those that have
not is of considerable interest in itself.

THE PLANTS STUDIED.

This study includes as many of the native desert trees and shrubs
as are readily obtained, and also such as have been irrigated in various
places in whatever manner or degree. These are the hackberry
(Celtis pallida), the creosote bush (Covillea tridentata), the candle
bush or ocotillo (Fouquieria splendens), and a relative of the cruci-
*Papers from the Desert Botanical Laboratory of the Carnegie Institution.
No. 12,

* VOLKENS, G., Flora der iigyptisch-arabischen Wiiste, p. 82.

905] 307

398 BOTANICAL GAZETTE [JUNE

fixion thorn (Zizyphus lycioides), representing the shrubs; and the
cat claw (Acacia Greggii), the mesquite (Prosopis velutina), and two
species of palo verde (Parkinsonia microphylla and P. Torreyana),
representing the trees.

These plants grow in habitats which are very unlike in appearance,
and which presumably are dissimilar also in such fundamentals
as soil characteristics and water supply. Speaking now of the vicin-
ity of the Laboratory only, for a radius of about ten miles the habitats
may be classed as (1) mesa, (2) river bottoms, (3) the rocky slopes
of the lower mountains, and (4) the “draws” of these mountains.
As regards the amount of water in the soil available to the plants,
the river bottoms should be classed first, after which should be placed
the draws; the rocky slopes and the mesa are frequently very dry.
The mesa is peculiarly desertic, because a subsoil, the so-called
“‘calliche,” which is practically impervious to water, reaches within
a few inches of the surface of the ground. The top soil, the adobe,
is a clay. In the river bottoms the top soil is several feet in depth.

The mesquite and the cat claw are found especially on the bottom
lands, where they may attain the size of forest trees, especially the
former. Both of these trees adapt themselves to the drier habitats,
however; but in such places they are of much smaller size and are
gnarled and much changed in outward appearance. Celtis and
Zizyphus also grow on the river bottoms; Fouquieria seems to be
confined to the rocky slopes. Parkinsonia microphylla also is mostly
found on rocky or well-drained slopes, while the other species of palo
verde (P. Torreyana) is to be found in the draws at a lower level.
Finally, the creosote bush, which is the characteristic shrub of the
mesa, grows on the bottoms as well as on the mesa, and is there
large and most vigorous. The natural distribution of these plants,
and also the modification in form which they assume in the various
habitats, point to the conclusion that the optimum conditions for
them are attained where, other conditions being equal, water is
to be had in abundance.?

That the greater vigor of the plants on the river bottom, as opposed
to those on the mesa, for instance, is not due mainly, if at all, to

2 Compare “The creosote bush (Covillea tridentata) in its relation to water supply,”
V. M. SPALDING, Bot. GazETTE 38:124. 1904.

1905] CANNON—WATER-CONDUCTING SYSTEMS 399

differences in the character of the soil, is shown by the manner in
which they react to an abundance of water when they are well irri-
gated, the substratum being otherwise the same. The creosote
bush and both species of palo verde thrive equally well by the side
of the same irrigating ditch, and the same is true of the other plants
mentioned above. That is to say, each of these plants, whatever
may be their natural habitat, is most vigorous where the water supply
is adequate. The suggestion cannot be avoided, therefore, that
the distribution of these plants in this vicinity is in some way indicative
of their specific reaction to the minimum rather than to the optimum
supply of water. The extent to which this phase of the reaction of
desert plants to water may influence their present distribution in
this vicinity will be the subject of future experimental investigation
at the Laboratory.

As has been suggested above, the irrigated desert plants are the
more vigorous in every way. The leaves are larger and are functional
for a longer period, and the plants attain also much larger proportions
than those that are not irrigated, and they also grow faster. To give
but a single instance of the latter, a palo verde well-irrigated four
years from the seed is now about eight feet high, while there are many
of these trees in the neighborhood of the Laboratory, without doubt
Many times as old, that are not so high.

With the larger leaf surface of the irrigated plants one would
look for a faster rate as well as for a larger total transpiration. What-
ever may be the absolute transpiration of the irrigated forms, the
Structure of the stems of the irrigated as opposed to the non-irrigated
plants indicates that the latter have the faster rate. That is, the non-
irrigated stems have larger ducts and more of them per equivalent
area of cross section than the irrigated stems, which can be taken
as indicating the more rapid transpiration of the former.’

To anticipate the general conclusions to be drawn from this
Paper, we may therefore be justified in thinking that the desert
plants of this locality, being probably better provided with conductive
elements than those of the Egyptian-Arabian deserts, may also have
a faster rate of transpiration.

3 inci Physiology of plants 1: p. 216; HABERLANDT, Pflanzenanatomie
(3d ed.), p.

409 BOTANICAL GAZETTE [JUNE

THE METHODS AND THE RESULTS.

The number of ducts of the woody cylinder was determined in
the following manner and under the following conditions: The
branches selected for study were as nearly of the same size as possible,
namely 0.6°™ in diameter; it did not appear feasible to get them
all of the same age. All of the sections were made midway between
the nodes. In counting the ducts an arbitrary area as a standard
of comparison was taken as follows. A circle 14°™ in diameter was
drawn on metric paper and cctants were struck off. By means of
an Abbé camera lucida an image of the section was so thrown on the
metric paper that the periphery of the woody cylinder and the two
points delimiting any octant coincided. The standard octant had
an area of 19.2499™ (14? Xo0.7845 +8). Since the magnification
employed was 66 diameters, the area actually examined was 0.29% ™,
or 29%4™™,

The method of studying the ducts thus chosen gives the greatest
prominence to the latest growth. This was desired for the reasons
that the most active transfer of water probably takes place in the
more peripheral ducts, and because I could be most sure of the recent
history of the plants as regards irrigation.

The term “irrigated” as used in this paper needs a word of
explanation. I have been obliged to take whatever plants I might
find that had received any water in addition to the rains. In certain
instances, to be detailed below, the plants were growing by the side
of an irrigating ditch and without question had water in abundance
every day of the year. In other cases they were a greater or less
distance from the ditch; and in yet other instances the ditch was
filled with water for a portion only of the time, say once a month,
during the dry season. Some of the forms received little water
artificially, as, for example, Zizyphus, which is in the cactus garden
at the University of Arizona and is not irrigated oftener than once
or twice in a year. When the plants received a small amount of
water I have called them “‘semi-irrigated.” The distinction between
irrigated and semi-irrigated forms is thus purely arbitrary.

The accompanying table presents a summary of the study. The
numbers refer to the number of ducts in 29°¢™™ of stem cross section
made under the conditions described above.

T905] CANNON—WATER-CONDUCTING SYSTEMS 401

|
Not irrigated | Irrigated Semi-irrigated

Fic. 1.—Acacia Greggii: left, not irrigated; right, irrigated.

The following outline indicates some of the conditions of growth
of the plants studied.

Acacia Grecct (fig. r).—The non-irrigated specimen was taken
by the roadside between an old and at present unused irrigating
ditch and the road ditch. Water stands in the latter for a consider-
able period after each heavy rain, and thus the effects of the rain are
conside ‘rably prolonged. The irrigated form was growing about
6™ from a ditch in which the water flows the most or perhaps all of
the year. The older portion of the silent cain’ stem does not

4 The ducts of the irrigated plants were very small; see figs. 5, 0, 7:

402 BOTANICAL GAZETTE [JUNE

have as many ducts as the more recent, an indication according to
the present view that this plant might formerly have been irrigated.
The owner of the field in which the non-irrigated specimen is found
tells me that such, indeed, is the case.

Fic. 2.—Celtis pallida: left, not irrigated; right, irrigated.

ha
mountain on which the Desert Laboratory is situated. The irrigated

CELTIS PALLIDA (fig. 2).—The non-irrigated plant grows on the

Fic. 3.—Covillea tridentata: left. not irrigated; right, irrigated.
plant is on the grounds of the University of Arizona, and has been
watered occasionally.
CoVILLEA TRIDENTATA (jig. 3).—The non-irrigated plant is on
the mesa at the northern base of the laboratory mountain. The

1905] CANNON—WATER-CONDUCTING SYSTEMS 403

irrigated plant is growing by an irrigating ditch. Its roots are
without question in a soil which is saturated with water all of the year.
A third specimen, not figured, was originally chosen for study as
being representative of the irrigated form. It was placed near an
irrigating station and was supposed, therefore, to have received
considerable water. When I examined the conductive system of
this plant I found it to be essentially the same as the non-irrigated
specimen. It was learned on in-
quiry that the plant had not been

Fic. 4.—Fouquieria splendens: left, not irrigated; right, irrigated.

irrigated at all, a curious confirmation of the view advanced in this
paper.

FOUQUIERIA SPLENDENS (fig. 4)—The non-irrigated specimen
is growing on the mountain near the Laboratory. The irrigated is
on the grounds of the Indian Industrial School, Tucson. It has
received a variable amount of water, more during the past two years
than formerly.

PARKINSONIA MICROPHYLLA (fig. 5).—The plant that is not
irrigated is on the mesa at the northern base of the Laboratory
mountain. The irrigated specimen is on the campus of the University
of Arizona and has been irrigated frequently.

PARKINSONIA TORREYANA (fig. 6).—The non-irrigated palo verde
is in the draw at the western base of the Laboratory mountain.

404 BOTANICAL GAZETTE [JUNE

The irrigated plant is on the campus of the University of Arizona
and has been irrigated in the same manner as the other species of

palo verde given above.

Fic. 5.—Parkinsonia microphylla: left, not irrigated; right, irrigated.
PROSOPIS VELUTINA (fig. 7)—The non-irrigated specimen of
mesquite 1s growing near the Laboratory. It is about 2™ high,

Fic. 6.—Parkinsonia Torreyana; left, not irrigated; right, irrigated.

gnarled, and evidently in need of a better supply of water. The
irrigated form of mesquite is growing near an irrigating ditch in

1905] CANNON—WATER-CONDUCTING SYSTEMS 405

which there is water several times during the year; not continuously

as in the other ditch mentioned above.

at ~

“ater Pe RS

10t irrigated; right, irrigatec

Fic. 8.—Zizyphus lycioides: left,
1e non-irrigated plant is on the

ZIZYPHUS LYCIOIDES (fig. 8).—T!
he campus

mesa near Tucson. The irrigated plant is growing on t

406 BOTANICAL GAZETTE [JUNE

of the University of Arizona in the cactus plantation. It receives
water by direct application about twice a year. The nearest irrigating
ditch is across the drive, about 13™ distant. It should be explained
that the drive is so constructed with reference to the underlying
calliche that presumably no water seeps across.

GENERAL CONCLUSIONS.

Although, much to my regret, the amount of water given to the
irrigated plants is not known with any degree of exactness in any case,
there can be no mistaking the fact that branches of irrigated plants
(even if “‘semi-irrigated” only) are poorer in conductive tissue than
branches of the same diameter of non-irrigated plants. This is an
unexpected condition and of especial interest in view of the small
development of the water-conducting elements in the non-irrigated
forms of the Egyptian-Arabian deserts as given by VoLKENs. It
merits further investigation.

Without doubt the irrigated plants have a greater absolute trans-
piration and organize each year a larger amount of wood than the
non-irrigated plants of the same age; but the composition of the
wood is different in the two instances. The irrigated plants con-
struct a relatively large amount of non-conductive tissue each year
(as compared with the amount of the other wood elements formed),
while the reverse is true of the non-irrigated plants. So that it
happens, in stems of equal diameter but not of the same age, that
the non-irrigated and older stems have more vessels than the irri-
gated and younger. One other characteristic of non-irrigated stems
was also noted, namely, their ducts were usually or frequently of
greater diameter.

We find, therefore, a quantitative and a qualitative difference in
the structure of irrigated and non-irrigated stems, and from what
has been stated it appears that any adequate account of the causes
which induce these structural peculiarities must separate the pro-
cesses which may be associated with the formation of the woody
cylinder as a whole from those which may be connected with the
organization of that part of the wood that is primarily to conduct
water.

I shall not presume at present to offer any hypothesis to explain

1905] CANNON—WATER-CONDUCTING SYSTEMS 407

the phenomena noted. It seems, however, on the surface of things,
that such explanation must take into consideration the differences
in the length and the character of the growing season of the two
classes of plants. The irrigated plants probably do not cease growing
during any month of the year and are most active when the tem-
perature is also the highest.
The curve of the growth of rae {|
these plants probably is | | |
very nearly parallel to that Anes
of the temperature for the |
year. The non-irrigated Pa |
plants are subject to the |
peculiar climatic conditions |
of the desert. The summer |
|
|

is the season of the maxi-

mum temperature. At

\

Tucson there are each
year two rainy seasons: a L\ |
that of winter, January- ae Bad | |
February; and that of sum- ee) bd il | L L
mer, July-Au gust. The Fic. 9.—Mean yearly rainfall and maxi-
period of the greatest vege- mum temperature at bs segeuibobiars Metin
tative activity once ar the temperature (rem =10°F.); solid line is rainfall
i ‘ in inches (1° =o.2').
time of the midsummer
rains (it is not known how the winter rains affect the growth
of these forms), and all growth ceases very soon after the
rains are over. The curve of growth of the non-irrigated plants
would be very similar, no doubt, to that for the yearly rainfall.
Fig. 9 expresses ina graphic manner the annual rainfall and tempera-
ture at Tucson. It may be taken also to indicate the rate of growth
of the non-irrigated and the irrigated desert plants. ;

Whatever may be the effect in detail, it must be admitted, I think,
that the growth of the non-irrigated plants, and this may also be true
of their transpiration, is directly associated with the distribution of
the annual rains. Further, the intensity of this growth, as sipeesabai
by the structure of these plants and the general vegetal conditions
of the desert, is probably directly connected with the fact that the

408 BOTANICAL GAZETTE [JUNE

heaviest rainfall occurs in summer. These facts encourage the view
that the peculiar structure of the plants of the Egyptian-Arabian
deserts, as regards the points under consideration, may likewise be
due wholly or in part to analogous causes.
The climate of these foreign deserts is quite different from that
of the desert about Tucson
| | LT | in the distribution, as well
j / % as the amount, of the
\ annual rainfall; and conse-
/ \ quently in the relation of
rf \ the rainfall to the period
/ i of the highest temperature.
i \ The most rain occurs in
f the deserts of northwest-
4 ern Africa and southwestern
4 Asia in winter, but the
\

maximum temperature is
in summer.

The period of the
greatest vegetative activity
in these deserts is in win-
ter; in summer the plants

Fic. 10.—Rainfall and temperature at 4!€ dormant. Inthe Egyp-
Cairo, Egypt, in 1887; broken line is temper- tian-Arabian deserts, there-

ature (1¢™=2°C.); solid line is rainfall ‘ th
(rem = mm), fore, the renewal of growt

N

occurs at the time of the
year when the rainfall is also greatest, but not at the season of
the highest temperatures. The curve of growth of the plants of
this region probably coincides very accurately with the curve of the
yearly rainfall. I have no means at hand of telling whether, on the
other hand, if such plants were irrigated the period of their greatest
growth would fall insummer. Such is apparently the case with some
of the cultivated forms’ in deserts farther west.

Fig. 10 represents the relation of the yearly rainfall to the tem-
perature at Cairo, for the year 1887.

5 SWINGLE, W. T., The date palm. U.S. — Agric. 1904.

DESERT BOTANICAL LABORATORY.

THE EFFECTS OF TOXIC AGENTS UPON THE ACTION
OF BROMELIN.
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY.
LXXIII.

JOSEPH STUART CALDWELL.
HISTORICAL.

ATTENTION was first directed to the existence of a proteolytic
enzyme in the juice of the pineapple (Ananas sativus) by MaRcano
(13). It wascalled bromelin by CHITTENDEN, who, with his pupils,
examined it in some detail (3, 4). They prepared it from the expressed
and filtered juice, after neutralization, by saturation with crystals of
ammonium or magnesium sulfate or sodium chlorid, any of which
precipitate the enzyme together with some proteid matter. The
precipitate was dialyzed free from salt in running water, collected
upona filter, and dried at 40° C. upon a water-bath. The flaky, whit-
ish residue thus obtained consisted of the enzyme with some associ-
ated proteid. It was soluble in water and active in acid or neutral
solutions, very slightly or not at all in an alkaline medium. The
sodium chlorid preparation was more active and greater in amount
than the others, and was used throughout the experiments. Curt-
TENDEN’S study was directed principally to a determination of the
products of digestion of blood-fibrin, myosin, and coagulated egg
albumen. These were found to be hemi- and anti-peptones, proto-,
hetero-, and deutero-proteose, leucin, and tyrosin. The enzyme was
thus determined to be tryptic in nature, and akin to other vegetable
trypsins in that it acted best in an acid medium.

PURPOSE.

The experiments described in this paper were undertaken with
a view to ascertaining whether a similarity existed between the
effects of poisonous metals upon the action of an enzyme and those
observed in experiments upon living organisms. While an immense
amount of work has been done with plants and animals, the results
are at variance, not even agreeing as to the most poisonous metal.
1905] 409

410 BOTANICAL GAZETTE [JUNE

It was hoped that the limits of toxicity might be more clearly defined
in the case of enzymes.

Very little work has been done in this field. In a general way it
is known that slight amounts of metallic salts affect enzymic action.
Cotes (6) found that cations and hydroxyl ions depress and anions
accelerate the action of ptyalin and invertin. Results with acids
and alkalis vary with the enzyme used (7, 8, 10). So far as I am
able to ascertain, the only extended work with poisonous metals has
been that of McGuican (16) upon diastase. His results confirm
the theory advanced by MATHEWws (10, 11) that the physiological
action of any atom or ion depends primarily upon its affinity for its
electrical charge. The work described in this paper was undertaken
with a view to determining whether this theory would hold for a
tryptic enzyme, and was completed before MGGuican’s results were
known to me.

Bromelin was chosen as a typical vegetable trypsin, rapid in its
action, easily prepared, and hitherto studied only with reference to
its digestion-products and its action in the presence of acids and
alkalis.

: PREPARATION OF THE ENZYME.

It was desirable to prepare the enzyme in as pure a state as pos-
sible, since it was early found that the presence of associated proteid
matter obscured the results of my experiments. Even very slight
amounts apparently shielded the enzyme from the action of poisons.
Furthermore, it was shown by some preliminary experiments, sum-
marized elsewhere in this paper, that autodigestion, with the forma-
tion of peptones, leucin, and tyrosin, occurred in solutions of the
impure enzyme when kept for some time at temperatures between
25 and 65° C.

After several trials, a preparation was obtained which contained
only very slight traces of associated proteid, and which was not at
all autodigestive. The dialyzed NaCl-precipitate was dissolved in
a little water, reprecipitated by the addition of 95 per cent. alcohol,
again dissolved in water, and again precipitated by adding crystals
of (NH,),SO,. A little of the associated proteid is left behind at
every precipitation, and five repetitions of the process gave a prepa-
ration containing only slight traces of proteid. Precipitation was
much hastened by placing the vessel in the refrigerator at about 4° C-

1905] CALDWELL—ACTION OF BROMELIN 41L

After being dried upon the water-bath at 40° C., the preparation
was wholly soluble in water and was not autodigestive in any medium
even after prolonged standing. It was active in faintly acid or
alkaline solutions, though not at all in neutral media. This prepara-
tion was used throughout the experiments.

In order that the results might not be rendered inaccurate through
the use of preparations of unequal purity, all the enzyme used was
prepared at one time and thoroughly mixed after drying. From 46
pineapples of average size, 16.4** of filtered juice were obtained,
which yielded 14.88™ of the enzyme dried at 40° C. This was kept
in a calcium chlorid desiccator and from time to time compared
with fresh preparations, but no deterioration occurred in the period
covered by the experiments.

The enzyme was most active in solutions of alkalinity equaling
m/30-m/40 NaOH, and was considerably less active in acid solu-
tions irrespective of strength. Its action was wholly inhibited by
m/1to HNO, or H,SO,, m/15 HCl, NaOH, or KOH, or m/20 NH,OH.

The fact that the enzyme had been found most active in acid and
neutral solutions by CHITTENDEN, while my own preparations were
markedly more active in alkaline than in acid media and showed
only the slightest trace of activity in neutral solutions, led me to a
very careful examination of this point. CHITTENDEN’S statement
of his method was followed in making a number of preparations.
Pineapples of varying ripeness were chosen, upon the hypothesis
that a change in the nature of the enzyme might occur with the ripen-
ing of the fruit, but all of my preparations agreed perfectly in requiring
an alkaline medium for optimum action, very much less action cccur-
ring in acid media. One preparation from very immature fruits
showed faint activity in a neutral solution. This difference from
the results of CHITTENDEN’s examination seems explicable only on
the ground that I have misunderstood his statement of his method
of preparation; yet the method is simple and the statement is appar-
ently lucid and exact enough to be followed without difficulty.

ISOLATION OF THE ALKALINE ENZYME.

Some of the facts observed in the preparation of the enzyme
suggested that it might be in reality a mixture of two enzymes separ-

412 BOTANICAL GAZETTE [JUNE

able by proper treatment. Some experiments with this purpose in
view met with partial success.

A neutral solution of one gram of the dry enzyme in 100° water
was carefully heated to 65° C. Upon gradual addition of NaCl
crystals a slight precipitate settled out, which when collected by
filtering, and dried, weighed 0.31%". The filtered liquid when
freed from NaCl showed no activity upon egg-albumen in acid
solutions, regardless of strength, but its activity in alkaline solutions
was undiminished and accorded in all respects with that of ordinary
preparations of the enzyme in alkaline media, the products of diges-
tion with the two being identical. The heat precipitate was imper-
fectly soluble in water and showed no action upon egg-albumen in
any medium.

I believe that these facts, with the differences, elsewhere stated,
in the action of poisons upon acid and alkaline solutions of the enzyme,
justify the conclusion that there are present in the preparation two
enzymes, one active in acid, the other in alkaline solutions. J am
aware that full proof of this statement demands isolation of the acid-
enzyme without destroying it in the process, but this I am as yet
unable to do. | ’

That the presence of NaCl is a necessary condition for the pre-
cipitation of the acid-enzyme is shown by the fact that no precipitation
occurs in unsalted solutions until heated to 76° C., activity in both
acid and alkaline media being retained until complete precipitation
occurs at 87-90%.

EXPERIMENTS WITH POISONS.

In the experiments with poisons, solutions containing o.006*™

of the enzyme per cubic centimeter were made up with distilled
water previously made acid or alkaline by the addition of HCl or
NaOH. An m/32 acid or alkali was adopted as a medium for all
digestions, since this strength allowed optimum activity in each case.
Thymol was used to prevent bacterial infection and was found per-
fectly satisfactory. Of this solution 5°°, containing 0.03%™ of the
enzyme, were transferred to each of the test-tubes to be used. To
each was then added 5°¢ of the poison to be used, of twice the desired
strength, 7. e., 5°° enzyme solution plus 5°° m/so000 Ba(NO,), gives

1905] CALDWELL—ACTION OF BROMELIN 413

1o°* m/10000 Ba(NO,),. To each tube was then added 1%™ egg-
albumen previously boiled and granulated, and the tubes were then
placed in the water-bath at 40° C. for 24 hours. They were then
removed, filtered to remove undissolved albumen, and tested.

Tests were made for peptones, leucin, and tyrosin by all the
standard tests. The biuret reaction after precipitation of the albu-
moses, and the tryptophan reaction described by ViNEs (25, @) were

OPTIMUM ACID MEDIUM m/32 HCl

OPTIMUM ALKALINE MEDIUM
m/ OH

32 Na
SALT | Dilution of molecular solution | Diluti f molecular solution
| Inhibiting Allowing | Inhibiting | Allowing
ea Si
| eee 110,000 130,000 80,000 100,000
PO Ni race ccd a's a wey 80,000 110,000 80,000 96,000
ie ae 60,000 90,000 45,000 60,000
fe ne 8 000 70,000 40,000 54,000
EDs erica ¢ 55 Ks 75,000 90,000 64,000 80,000
3) 65,000 85,000 45,000 60,000
te 40,000 60,000 36,000 56,000
pee ee ae 37,500 56,000 0,000 56,000
lt (0) oe 28,000 32,000 16,000 22,000
oS ae ne 16,750 23,000 14,000 17,000
nl in 10,000 13,000 ,000 8,
Se ae 8,000 12,000 6,000 (i Shee
CAN Oa snd ca ,0 ,600 5,000 7,000
RN eek nck ew 24 4,800 6,650 4,750 5,650
Od 045 Ge ss 4,800 6,400 3,000
Sheree sk era es 500 5500 O00 3,000
LiNO, at itera ete cp ga : 1,200
PAPO viveacc. ... 750 1,050 °
3 2 Se 512 1,024 500
is Shas a eae 1,600 2,000 1,100 1,675
oa goo 1,200 72 6
a a een 750 1,100 650 1,050
a eee 800 1,000 600
MalN Os). ces os 650 | 1,000 3 512
oS NO oe 00 | 1,000 375 512
ae 100 | 160 60 120
RG ee Sits a wes 100 200 125 575
Ais 256 512 256 432
PaO... a. 400 | 512 256 432
15 ie Sa a 320 | 480 275 420

found to be most delicate, HorrMANN’s and SCHERER’S tests being
used as confirmatory. The unequal delicacy of the reactions employed
made it necessary to adopt a rule to secure uniformity in results.
Hence only such concentrations were considered inhibiting as gave
ho results with any test, while the only strengths tabulated as allowing

414 BOTANICAL GAZETTE [JUNE

action were those that gave unmistakable results with a majority of
the tests. SCHERER’s test—the isolation of leucin and tyrosin—
was of course conclusive. The application of this rule resulted in
some cases in comparatively wide gaps betiveen the toxic and non-
toxic strengths given in the table; e. g., m/650 Mg(NO,), is given
as inhibiting, since it gave none of the tests; m/1000 as allowing
action, since it gave all; while m/850 gave faint tryptophan reaction
but no SCHERER’S test, and was therefore disregarded.

In beginning to experiment with a particular salt, a stock solution
was first made; a portion of this was diluted to give the greatest
strengths desired, and greater dilutions were made from this by
successive additions of distilled water. For example, in my first
experiment with CuSO, a series of tubes with m/5000, m/ 10000, up
to m/400000 were made up in duplicate. The results indicated
where the critical points—between m/50000 and m/1000co—were
to be expected, the duplicates preventing error. Closer series were
then made and repeated until the limits of toxicity and non-toxicity
were clearly determined. Finally the whole table was twice checked
over in duplicate, so that the figures given represent in every case
from eight to twelve concordant results.

The results obtained may be shown by arranging the metals used
in the order of toxicity, beginning with the most poisonous. The
results obtained by Marnews, working with eggs of Fundulus
heteroclitus, and those of McGurcan with diastase are given for
purposes of comparison.

MATHEWS (14) McGuican (16) CALDWELL
Ag Ag

Ag
Hg Hg Hg
Cu Cu Cu
Cd Cd Pb
Pb Co Zn
Zn Zn Ba
Co Pb Cd
Li Sr Co
Sr Ba Na
Na Mg Li
Ba is Sr
Mg Na Mg
NH, NH,

Fi at a i

1905] CALDWELL—ACTION OF BROMELIN 415

While there are a number of exceptions, these results show a general
agreement which is more striking in view of the total lack of harmony
in the earlier work with poisons, whether with plants or animals.
Silver, mercury, and copper stand at the head of the three lists. In
my experiments, cadmium stands near zinc, nearly in the place it
should occupy in accordance with the theory of MATHEWs, who
found it in his work only less poisonous than copper. MATHEWS
and McGuiGan have found sodium less poisonous than lithium or
strontium, using the chlorids. I find the three sulfates practically
equal in toxicity, while sodium nitrate is much more poisonous than
the other nitrates. My results agree more nearly with those of
Martuews in the places given lithium, lead, and cobalt. The great
toxicity of barium in my experiments cannot be attributed to impuri-
ties in the salts used or to inaccuracy in performing the experiments,
since several standard preparations gave markedly uniform results,
and the series has been worked over so many times that the possibility
of mistake is excluded.

The nitrates uniformly inhibit the action of the enzyme in some-
what greater dilution than the corresponding sulfates and chlorids,
which agree very closely.

The action of the enzyme is markedly weaker and is uniformly
inhibited by poisonous solutions of less concentration than when
acting in alkaline media. This confirms the evidence already given
for the existence of two enzymes.

EXPERIMENTS UPON AUTODIGESTION.

The experiments upon autodigestion were suggested by the dis-
covery that while none of the tests for peptones, leucin, or tyrosin
were given by freshly prepared aqueous solutions of the impure
enzyme, all these were present in such solutions after standing for
a little time at 25-60° C. CHITTENDEN (loc. cit.) has stated that the
proteids of the juice are exceedingly resistant to the action of the
enzymes, while VINES (25, ¢) has mentioned the occurrence of auto-
digestion in the expressed juice. My own observations led to the
supposition that the associated proteids were acted upon by the
enzyme at suitable temperatures. This was found to be true of acid
and alkaline solutions, not of neutral ones. The associated proteid

416 BOTANICAL GAZETTE [yUNE

was first rapidly digested, the breaking down of the enzyme itself
proceeding more slowly and continuing until complete.

Solutions containing 12™ each of the impure enzyme were placed
in the water-bath and kept at 40° C. for 12 days, the growth of bacteria
being prevented by the addition of thymol. At short intervals,
portions were removed and examined for digestion products by all »
the methods employed in other experiments. Traces of peptones,
leucin, and tyrosin were found after 14 hours, rapidly increased in
amount for 10-14 hours, then very slowly increased for 9-10 days.
During the first day, portions were hourly removed, filtered, and
tested as to their activity upon egg-albumen, no perceptible decrease
in power being shown. After the first day, portions were removed
once or twice daily and tested, and there was found a decrease in
power to digest egg-albumen proceeding pari passu with the increase
of digestion products in the stock solution, until after the eighth day
no perceptible action upon egg-albumen occurred, even after pro-
longed standing. Contrary to my expectation, the process of auto-
digestion was not checked or inhibited by the accumulation of its
products, since it was equally rapid in the stock solution and in
another from which the digestion products were daily removed.
Furthermore, the two enzymes break down with equal rapidity,
since activity in acid and alkaline media decreased equally until the
eighth day, after which no action occurred. :

No such decrease in activity occurred, nor were digestive products
present, in purer preparations of the enzyme kept for a much longer
period (five weeks) under similar conditions, nor were such solutions
affected by light.

It was hoped that a preparation purer than I had otherwise been
able to make might be obtained by allowing the digestion to proceed
until the associated proteid had been broken down, separating the
products of digestion from the enzyme and drying the latter. To
my surprise, digestion immediately began in aqueous solutions
made from such a preparation, and would occur perceptibly in the
moist precipitate while drying in a calcium chlorid desiccator.

Autodigestion does not begin so long as there are present proteids,
either egg-albumen, fibrin, or that present in the juice, but does begin
as soon as these have been broken up.

1905] CALDWELL—ACTION OF BROMELIN 417

Autodigestion occurs most rapidly in alkaline solutions equaling
m/16~m/32 KOH or NaO, less rapidly in m/10-m/24 HCl, H,SO,,
or HNO,. Action was inhibited by greater concentrations and
occurred more slowly in weaker ones. It occurs with poisons in
strengths very much greater than those inhibiting the action of purer

. preparations upon egg-albumen, as shown by the following table:

AUTODIGESTION IN m/24 HC] AT 40° C.

DILUTION OF MOLECULAR SOLUTION
SALT a a i ee oe oe
Inhibited Allowed
CNB ISOg es aeeoe oe 72 120
NH,NO, Fi AiNe coemenrer sss sehen fe 2 48 4 64
EOIN Os Vac Saat also dies 128 256
rot 0) Fea irae fo eee ae 100 I
CUS Oped ces eoprenomcns 6,000 8,650
UNE OD ieee one teld tee lene 16,000 22,750
FEU) DR APURRRY Oa teas 10,000 16,000
EIN dlp ecsin oh reiterate fo) fete)
CON Oa airs ctreneccntte 12 68
lyr Corey areiy eee aes ahs 1,600 2,400
INAS CE oars sa Sard siael Ses 150

In an alkaline medium (m/32 NaOH), inhibiting and allowing
strengths were uniformly about one-third greater with the few salts
tested.

The table just given also represents pretty closely the conditions
for the digestion of egg-albumen by such impure preparations, in
so far as this was studied. For example, digestion would just occur
in m/14000 ZnSO, m/7200 CuSO,, or m/1800 Co(NO,),, as deter-
mined by decrease in the amount of albumen subjected to the test.
Hence the degree of susceptibility to poisons is determined largely
by the purity of the preparation employed, apparently because the
enzyme is shielded by the associated proteid from the action of
poisons. The results obtained with preparations of varying purity
are parallel, 7. e¢., the toxic strengths of different metals bear approxi-
mately the same relation to each other irrespective of the preparation
used,

SUMMARY.

1. Impure preparations of bromelin are strongly autodigestive
in acid or alkaline media, such digestion beginning when the breaking
up of proteid impurities has been completed and proceeding to total
destruction of the enzyme.

418 BOTANICAL GAZETTE [JUNE

2. The effects of poisons vary with the purity of the preparation
used, slight amounts of proteid impurities rendering necessary an
enormous increase of concentration in order to inhibit action.

3. The toxic strengths of the salts used maintain a constant
relationship irrespective of the purity of the enzyme used, 7. e., silver
is always most poisonous, copper third, zinc sixth, and so on.

4. Bromelin, when prepared in a relatively pure condition is
not at all autodigestive, the presence of some proteid of the juice
apparently being a prerequisite for such action.

5. Such preparations appear to be in reality a mixture of two
enzymes, one active in alkaline solutions, slightly more resistant
to poisons, and twice as great in amount as the other, which is active
in acid media and is destroyed by heating to 65° C. in saline solution.

6. The limits of toxicity and non-toxicity are somewhat. more
clearly defined than has been the case in experiments upon living
organisms (5, II, 12, 18, 19, 2

7. The results obtained agree in general with MATHEWS’S arrange-
ment of the metals upon the theory that ‘‘the affinity of the atom
or ion for its electrical charge is the main factor determining its
physiological action.’”” Cadmium in my experiments occupies the
position it should hold in accordance with this theory, while barium
is far out of place.*

My sincere thanks are due to Dr. B. E. Lrvincston, at whose
suggestion the work was begun, for much help in its inception, and
to Dr. C. R. Barnes for assistance in its progress and completion.

* Experiments undertaken in order to determine more conclusively the place to

barium are still in progress. I have found it only relatively less poisonous

than zinc in every case, whether bromelin,: papain, or animal trypsin were the
enzyme u
THF UNIVE RSITY OF CHICAGO.
LITERATURE CITED.
I. Bourque or, Les fermentations. Paris. 1
2. CHITTENDEN, MENDEL, and McDermort, Papiin proteolysis. Am. Jour.
Physiol. 1:255. 1808.
3- CHITTENDEN, R. H., The enzyme of the pineapple. Trans. Conn. Acad.
8:36. 1801.

1894.

, The enzyme of the pineapple. Jour. Physiol. 15:290-

q
:

4

.
i,
4

i

1905]

25.

26.

CALDWELL—ACTION OF BROMELIN 419

CrarK, J. B., On the toxic effects of deleterious agents. Bot. GAZETTE
28: 289. 1899

Cotes, S. W., Contributions to our knowledge of enzyme action. I. Influence
of electrolytes on the action of amylolytic ferments. Jour. Physiol. 30:
202. 1903. II. Influence on invertin zbid. 30: 281. 1903.
UCLAUX, Traité de biologie. Paris. 1896.

Errront, L., Les enzymes et leurs applications. Paris. 1899.

GREEN, %. The soluble ferments and fermentation. 1899.

Gus, Archiv. Gesam. Physiol. 91:145. 1902.

KAHLENBERG and True, On the toxic action of dissolved salts. Bor.

. 1896.
Minos. M., Studien iiber die oo einiger Metallsalze auf das
Wachee héherer Pflanzen. Jour. Coll. Sci. Imp. Univ. Tokyo 19:

Art. 13. pp. 37. pl. 1

Marcano, Bull. of Ponies 5:77.

MaruHews, A. P., Relation between ce tension, atomic volume, and
physiological sation of the elements. Am. Jour. Physiol. 10:290. 1904.
—————_—__———, Nature of chemical and electrical stimulation. ibid.11:
455. 1904.

McGuican, Am. Jour. Physiol. 10: 444. 1924.

Nertson, C. H., and Brown, O. H., Effect of ions on the hydrolysis of
butyric ether ties watery extracts of the pancreas. Am. Jour. Physiol.
10: 335.

Ono, N., eee die Wachsthumsbeschleunigung einiger Algen und Pilze
durch cuennache Reize. Jour. Coll. Sci. Imp. Univ. Tokyo 13:141-156.
pl. 13. 1900

RICHARDs, H. M., Die Beeinflussung des Wachsthums einiger Pilze durch
chemische Reise. Jahrb. Wiss. Bot. 30:665-688. 1897.

Ripeat, O. S., Papiiin digestion. Pharm. Jour. IIL. 24:845. 1894.

Suarp, Gorpon, Papiin digestion. Pharm. Jour. II. 24:633. 1893.

ScHUTZENBERGER, Les fermentations. Paris. 1896

Stevens, F. L., The effect of aqueous solutions upon the germination
of fungus spores. Bor. GAZETTE 26:377. 18

Vines, S. H., Proteolytic enzymes of Nepenthes. (a) Ann. Botany 10: 292.
1896; (6) 11:563. 1897; (c) 12:548. 1898; (d) 15:1. 1902; (e) 15:

563. 1902.
Vernon, H. M., Conditions for the action of pancreatic ferments. Jour.
Physiol. roe rgor et seq.

BRIEFER ARTICLES.

THE EARLY HISTORY OF ANGIOSPERMS.

THE paper lately published by Mr. Harotp Lyon on the embryo of
angiosperms' gives a clear account of existing views on the race-history
of monocotyledons and dicotyledons. In the course of his argument
Mr. Lyon has referred to my own work on the subject in very generous
terms, while giving in greater detail than before the arguments which had
already led him to the opposed conclusion. We are, as before, in complete
agreement as to the field of battle. We both hold that monocotyledons
and dicotyledons come from a common stock, and that the single cotyledon
of the one is strictly homologous with both cotyledons of the other. But
on the question of the comparative antiquity of these two classes, we start
from opposite ends of the lists. I do not propose to discuss the points on
which we differ. The evidence on both sides has been published, and also
our respective interpretations. Those interested in the question are in a
position to form their own judgment on it. The issue will be determined
by the results of future research. But Mr. Lyon’s lucid statement of the
case has shown-me that my own is obscure in two points, and I wish to
take the first opportunity of restating them.

First I wish definitely to disown the suggestion that the fusion of two
ancestral cotyledons might have taken place within the seed where they
were acting as sucking organs. This appeared in my first sketch of the
whole hypothesis, published in May, 1902.2, In later papers the fusion
of the cotyledons is attributed to the reduction of assimilating organs
characteristic of geophilous seedlings in their first season, a second sug-
gestion, inconsistent—as Mr. Lyon has pointed out—with the first, which
is implicitly abandoned. It would have been clearer to state definitely,
as I do here, that—if we consider monocotyledons as derived from a dicoty-
lous stock by adaptation to a geophilous habit—the fusion of two cotyledons

Lyon, Harotp L., The embryo of the angiosperms. American Naturalist
39:13-25- 1905.

2 The origin of the seed-leaf in monocotyledons. New Phytologist 1:107-113-
pls. 2. 1902.

3A theory of the origin of monocotyledons. Ann. Botany 17:1-92- pls. I-7-
1903. The evolution of monocotyledons. Bot. GAZETTE 37:325-345- 1904-

JUNE] 420

1905] BRIEFER ARTICLES 421

into one is simply and sufficiently explained by the necessity of reduction
in the green parts of the seedling, and that this explanation of course
requires that the ancestral cotyledons should be acting as green assimilating
organs above ground during the period of fusion, and excludes my first
suggestion that they fused as sucking organs within the

at my argument was obscure in a more important respect I infer
from Mr. Lyon’s statement that the evidence on which it is based could as
well be read backwards as forwards; in short that the facts on which I
rely might be used to prove the derivation of dicotyledons from monocot-
yledons. My statement of the argument must have been very defective
if it admitted of any doubt on this head. No observer dealing with that
evidence at first hand could hold such an opinion. He would find the
evidence incomplete; he might rate its value much lower than I do; but
so far as it goes he must allow that it points in one direction—the derivation
of the single from the double cotyledon. To make this clear, the nature
of the evidence must be taken into account. It is of three kinds: anatom-
ical, embryological, general.

The anatomical argument is due to Professors QuEvA and JEFFREY.4
Both have pointed out that the vascular structure of the young stem in
monocotyledons is of the exogenous or dictoyledonous type. They both
conclude that monocotyledons are derived from dicotyledons. No botanist,
I believe, has denied the facts, which I could confirm if necessary. Their
theoretical value may be discounted, but the most ingenious critic could
hardly use them to prove the descent of dicotyledons from monocotyledons.

The embryological evidence rests primarily on my conclusion that a
certain type of vascular structure in the cotyledon and hypocoty] of liliaceous
seedlings is primitive, and that various other types of seedling structure
in the same family are genetically connected with it. Where we find a
single line of related structures there is nothing—in the absence of inde-
pendent evidence—to show which of the extreme forms is the more primi-
tive. But where—as in this case—there are several distinct lines of descent
ending in the same vascular type, it is fair to assume that type as the primi-
tive structure. For a common ancestor naturally gives rise to divergent
stocks, but it would be an extraordinary series of coincidences which
should lead several distinct types of structure to produce remote descend-
ants of a single ty

It is true that uniform conditions of life do lead to great superficial
resemblance between organs which are morphologically distinct. But

4 Quéva, C., Contributions & l’'anatomie des Monocotyledonées, p. 147. 1899
Jerrrey, E. C., in Courter and CHAMBERLAIN’s Morphology of Angiosperms,
P- 316. 1903

422 BOTANICAL GAZETTE [JUNE

the resemblance in such cases is mainly external; the metamorphosis is
revealed by differences in internal anatomy. Now in this study of lilia-
ceous seedlings we are dealing with anatomical details which are often
uniform, or very nearly so, among allied genera of the most varied external
form. Thus, among the Tulipeae some species of Lilium have hypogaeic,
other species epigaeic cotyledons; the cotyledon of Fritillaria is of the
common green rush-like form; that of Tulipa resembles Fritillaria, but
the whole structure of the seedling is transformed by the conversion of the
stem bud into a dropper. Yet the same vascular symmetry is found in the,
cotyledon and hypocotyl of all these divergent forms. On the other hand
seedlings belonging to almost every tribe of the family possess the green
rush-like cotyledon, but it masks a great variety of vascular structure.
Thus the reappearance of definite vascular symmetry in several lines of
descent cannot in this case be put down to the action of external conditions
moulding distinct types to one pattern. The one simple and adequate
explanation of the facts is that the various lines of descent started from a
common ancestor with vascular symmetry of this persistent pattern.

Now the whole argument from this class of evidence depends on the
fact that this vascular type is bisymmetrical. Two equivalent and quite
distinct bundles traverse the elongated cotyledon; two traces in the hypo-
cotyl unite to form a tetrarch root. A single type which is symmetrical
about two planes is connected with several symmetrical about one. There
is no difficulty in supposing all these unisymmetrical types to be descended
from the one bisymmetrical type; but it is incredible that the descendants
of distinct unisymmetrical types should all become bisymmetrical struc-
tures of precisely the same kind. The argument may be neglected, but it
cannot be read backwards. It cannot be used to demonstrate the forma-
tion of two cotyledons from one.

I do not mean to assert that the evidence quoted cannot be reconciled
with the hypothesis of a primitive monocotylous angiosperm. The bilat-
eral cotyledon may conceivably represent a terminal member which becomes
modified in one fashion or another into the likeness of a lateral one. But
this interpretation of the facts explains only how an apparently lateral
member may be descended from a terminal one. It is a study in the
derivation of various monocotylous types from a primitive monocotylous
form; it gives no clue to the descent of a race with two cotyledons from
that form.

The same criticism applies to that class of evidence which I have called
general. My reasons for considering monocotyledons as a race specialized
from a dicotylous ancestor by adaptation to the geophilous habit have

i ath UTE I
1g

te

mq

1905] . BRIEFER ARTICLES : 423

already been given in this journal. They need not be repeated here. On
that hypothesis the formation of one cotyledon from two is due to the
necessity of reduction in the aerial organs of seedling geophytes. The
tendency to such reduction is shown in other ways. Among dicotyledons
the few species which have but one cotyledon are geophilous, so—with
one exception—are those which have cotyledons united almost to the top.
In many geophilous forms the cotyledons never appear above ground at
all, and the first leaf is much reduced in size. Adaptation to a geophilous
habit also serves to explain many of the other structural features which are
correlated in monocotyledons with the presence of a single cotyledon,
notably the stem anatomy.

But though the collective weight of evidence in favor of this view
appears to me very great, I have never thought it conclusive. There are
no facts to make it incredible that the primitive angiosperm was monocot-
ylous, and that modern dicotyledons derive their two cotyledons from
division of the original member. That opinion, however, cannot be
deduced from the evidence just given. It is conceivable that the geophilous
habit has served simply to specialize monocotyledons, operating to reduce
the original terminal cotyledon to an apparently lateral one. But that
hypothesis does nothing to explain the rise of dicotyledons from a monocot-
ylous race, and it leaves the very marked similarity in vascular structure
between the primitive liliaceous type and certain geophilous species of
Ranales out of the question. Yet the approach to a true monocotyledonous
structure in some of these forms is most striking, and extends to the mature
as well as the seedling plant.

In conclusion, I wish again to point out that my purpose in this com-
munication has been to correct an inconsistency in the former statement
of my argument, and further to restate a portion of it which has been
misapprehended. I have purposely abstained from criticism of Mr.
Lyon’s alternative hypothesis, and from any attempt to answer his criti-
cisms on mine, except the two which were founded on obscurity in my
previous writings.—ETHEL SARGANT, Quarry Hill, Reigate, England.

CURRENT LETERATURE.
BOOK REVIEWS.
A college botany.

PROFESSOR ATKINSON has revised and elaborated his Elementary Botany
for college use. An elementary college text that includes all of the great divisions
of botany seems to be in demand, at least according to the judgment of publishers.
Perhaps the demand is both true and just, but it is a large one to make of a single
author, who is supposed to be, from the college standpoint, either a morphologist,
a physiologist, or an ecologist. One looks for inequality of grasp under such
circumstances, unless the presentation is so elementary that it hardly belongs to
a modern college course.

In spite of this disadvantage, Professor ATKINSON has covered the whole
general field in a way that indicates an unusually wide familiarity with the various
divisions of the subiect. The parts dealing with physiclogy and morphology are
largely elaborations of the same parts in the elementary text, introducing the new
material suited to more advanced students and bringing certain parts up to date.
The part dealing with ecology, however, has been entirely reorganized, and
represents the first presentation of ecology in an American textbook from the
college standpoint.

In the organization of the text, part one (pp. 135) deals with physiology, part
two (pp. 207) with morphology, part three (pp. 115) with the ecology of plant mem.
bers, part four (pp. 184) with plant associations, and part five (pp. 65) with repre-
sentative families of Angiosperms. The space given to the different parts repre-
sents a balance unusually well-maintained for books of this type. There is
always a temptation to overdo the part in which the author is especially interested.

Professor ATKINSON believes in numerous and good illustrations, with as
many of them original as possible; and hence the volume is full of fresh and
suggestive illustrations, and should be of great service to college classes in ele-
mentary botany.—J. M. C.

Handbook of plant morphology.

The friends of ArTHUR, BARNES, and Coutter’s Handbook of Plant Dis-
section, published in 1886, will be glad to know that this helpful laboratory guide
has been rewritten, thus bringing it again in touch with the best methods of ele-
mentary instruction. The authors have delegated the revision to other hands,
and both for this reason and on account of changes necessitated by recent devel-
opments in botany, it is perhaps fitting that the new edition should appear under

* ATKINSON, GEORGE Francis, A college text-book of botany. pp. xvit737-
figs. 592. New York: Henry Holt & Co. 1905. $2.00.
424 [JUNE

1905] CURRENT LITERATURE 425

a different title.' The most important change noted is the increased number
of types discussed, and the presentation of these in a connected account. In
harmony with this plan many unrelated details, especially concerning the vege-
tative structure of the higher plants, are omitted. Though the number of forms
discussed is increased from eleven to twenty-five, the size of the book is reduced,
having three-fourths the number of pages in Plant Dissection. The types
selected illustrate very well the probable steps in the evolution of plants, and the
discussions are exceedingly clear and suggestive. It seems to the reviewer, how-
ever, that the introduction of a heterogamous confervoid form might have strength-
ened the presentation of the subject of heterogamy among the algae. So long
as teachers have too large classes and too little time for purely inductive study
in the laboratory, some form of written direction seems indispensable. Plant
Morphology meets this demand in a most helpful way. A possible danger may
lie in its excellence, in that weak teachers may depend upon it too fully and neglect
the personal relation which should accompany laboratory study of this character.
—R. B. WY1IE.
NOTES FOR STUDENTS.

Allen? has recently published in full the naar of ns ey of nuclear division

in the pollen mother-cells of Lilim canadense, f 'y announce-
ment which appeared last years This Lee is of especial interest in relation
to the final paper of FARMER and Moore on the reduction divisions in animals

and plants, since Allen reaches ensues different conclusions as to the
events of synapsis and the preparations for the heterotypic mitosis. As FARMER
and Moore also studied a species of Lilium (L. candidum), it seems hardly pos-
sible that both accounts can be correct, so that the line separating the two schools
is sharply drawn in these accounts of a similar form. One school is led by FARMER
and Moore, and finds support in STRASBURGER’s recent paper Ueber reduk-
tionsteilung (1904), and in the recent work of GREGORY on the leptosporangiate
ferns, and of Writi1ams on Dictyota. With ALLEN are associated in the chief
point of dispute (the formation of the bivalent chromosomes of the heterotypic
mitosis) the botanists of the Carnoy Institute, GREGOIRE and BERGHs, and also
OSENBERG.

ALLEN’s account is chiefly remarkable for the detailed study of the events
preceding and following synapsis, which are presented in greater detail than in
any previous eins ie The nucleus of the young pollen mother-cell contains
a network of e irregular masses, derived from the chromosomes of the pre-
ceding eee in gs archesporium, and connected by fibers. As the nucleus
increases in size, the chromatin knots become widely separated, and the nucleoli

«CALDWELL, O. W., Handbook of Plant Morphology. 1r2mo. pp. vili+ 190.
New York: Henry Holt & Co. 1904.

2 ALLEN, C. E., Nuclear division in the pollen mother-cells of Lilium canadense.
Annals of Botany 19:189-258. pls. 6-9. 1905.
3 BOTANICAL GAZETTE 37: 464. 1894.

426 BOTANICAL GAZETTE [JUNE

are clearly distinct from the network. During synapsis the reticulum becomes
transformed into a definite spirem. the fibers thickening apparently by the dis-
tribution of material from the knots, which become less conspicuous. The
fibers now arrange themselves in pairs, and a general contraction of the reticulum
takes place, probably associated with this approximation of the fibers. There
is thus formed a continuous spirem in the nucleus, which is plainly composed
of two slender threads lying side by side and probably with no free ends. The
two threads often run closely parallel, sometimes loosely twisted about one another,
sometimes in contact and apparently fused, ti rather widely separ ated.
It is clear that the double nature of the spirem is not due to a fission, but that
two independent threads are developed out of the reticulum. The two threads
are regarded as of paternal and maternal origin respectively. They gradually
fuse, so that in later stages of synapsis the nucleus appears to contain a relatively
thick single (fusion) spirem, which is shorter and more loosely coiled than in earlier
stages. Each thread of the pair consists of a series of chromatin granules (chro-
momeres) imbedded in a ground substance (linin). As the two threads unite,
the chromomeres generally come together in pairs and fuse to form a single row
of larger chromomeres. The single (fusion) spirem on emerging from synapsis
becomes uniformly distributed throughout the nucleus as a much convoluted
thread. Some of the loops become fastened to the periphery of the nucleus.
There is no regularity in the number of chromatic segments which are formed
later.

While evenly distributed the single (fusion) spirem undergoes a longitudinal
fusion into two threads, which is preceded by a division of each fusion chromomere.
This is the first longitudinal fusion, well known from the descriptions of GREGOIRE,
GuIGNARD, Mortier, and others. The split spirem now segments into the
reduced number of chromatic elements (bivalent chromosomes) characteristic
of the heterotypic mitosis. The first free ends of the segments usually appear
at the periphery. of the nucleus, where it is evident that the split spirem breaks
apart at the loops. The segments of the split spirem now shorten and thicken,
taking on the various forms peculiar to the heterotypic mitosis. They are
obviously pairs of chromosomes, which stand for the full number of sporophytic
elements (24), now associated to form a reduced number (12) of bivalent chromo- .
somes. Shortly after the segmentation of the spirem the chromosomes of each
pair may show evidence of the second longitudinal fission, which is completed
during the metaphase of the heterotypic mitosis. Meanwhile the spindle of the
heterotypic mitosis is being organized, the position of the bivalent chromosomes
is shifted, and they are arranged on the nuclear plate where the elements of each
pair separate as the so-called daughter chromosomes of the heterotypic mitosis.
This first division in the spore mother-cell then simply distributes in two sets.
the twenty-four sporophytic chromosomes which ALLEN believes to have been
derived from a maternal and paternal spirem. The second longitudinal fission
which becomes conspicuous during metaphase of the heterotypic mitosis is clearly
a premature fission of the chromosomes which are to enter the second nuclear

1905] CURRENT LITERATURE 427

division (homotypic). There is no true resting period between the two mitoses,
the chromosomes which leave the heterotypic figure pass morphologically
unchanged into the homotypic.

The chief points of difference between ALLEN’s account and that of FARMER
and Moore are briefly as follows: FARMER and Moore fail to find the organi-
zation of two threads preliminary to the formation of the single spirem which
emerges from synapsis. They recognize only one longitudinal fusion of this
spirem, which is regarded as preliminary to the formation of the chromosomes
of the homotypic : mitosis. The pairs of chromosomes of the heterotypic mitosis.

significance of the reducing division cannot be considered here. The accounts
themselves rest on matters of fact and not of interpretation, and one or the other
is likely to fall, for it is hardly possible that both can be right, especially since they
treat of the same form.

ALLEN presents at the end of his paper a thorough and very interesting dis-
cussion of his conclusions in their relation to problems of heredity and hybridi-
zation.—B. M. Davis.

THE suBJEcT of soil inoculation for legumes has recently acquired a.wide-
spread interest in this country, first, from numerous popular accounts relating
to the subject, and again on account of a widely advertised commercial product.
by which is it claimed the proper bacteria can be introduced into the soil. For
this reason Moore’s account? of his work in this field is of special interest.
The first part of the bulletin is a general historical account dealing with the
various methods by which nitrogen is fixed in the soil, and leading up to the dis-
coveries of HELRIEGEL and WILLFARTH connecting the fixation of nitrogen by
legumes with the tubercles on the roots, and the discovery of bacteria in these
tubercles by Worontn. The chief results of Moore’s investigations may be
briefly stated as follows. The root-tubercle organism exists in three well-defined

h

threads” passing through the tissues of the host. These curious fungus-like
structures have often been observed and figured, and have been the objects of
much discussion. Their explanation is that they are zooglaea-masses composed
of numerous minute bacteria. These minute bacteria soon give rise to larger
rod-shaped forms which may or may not be motile. These finally produce the
branched forms peculiar to the Iegume-nodule. It is only this last form that
is of any benefit to the plants, for in this state the bacteria are broken down and
their contents made available. The name of the organism is changed to Pseudo-
monas radicicola, since the motile rods have flagellae only on one end. The

ooRE, G. T., Soil inoculation for legumes, etc., U. S. Dept. of Agric., Bureau.
of Plant Focus, Bull. 71, pp. 72. pls. 10. 1905.

428 BOTANICAL GAZETTE [JUNE

is only a single species, but several forms or races occur adapted to certain species
of legumes. Their slight racial characteristics may be easily broken down by
cultivation. The results of studies on the nitrogen-fixing power of the bacteria
are extremely interesting, showing that the supposed symbiotic relation between
the organism and the root probably does not exist. It is rather to be regarded
as a parasite, its nitrogen value being merely incidental to the death of the organism.
The nitrogen is fixed by the tubercle-forming bacteria within their bodies. This
was determined by cultures in flasks containing nutrient solutions without nitro-
gen. There was no increase of nitrogen in the solution, but a marked increase
in the organisms themselves. In its biology the organism is therefore considered
a parasite. Later the plant is able to overcome the parasite and profit by the
nitrogen which has been fixed. When grown on nitrogenous media, it was found
that the organism lost both its power of infecting leguminous plants and its power
of fixing nitrogen. In non-nitrogenous media both of these properties were
retained. The failure of NoBBr’s attempts in Germany a few years ago to put
upon the market pure cultures of this organism can probably be attributed to
lack of recognition of this fact. As a result of these studies Moore has devised
a method of putting up for distribution pure cultures of Pseudomonas radicicola,
grown in nitrogen-free media and dried on cotton immersed in the culture. ese
cultures are sent out by the U. S. Department of Agriculture together with pack-
ages of nutrient salts to multiply the organism. The mass-culture thus obtained
is used to inoculate the seed or the soil. Numerous reports from farmers of all
states indicate that this method will prove successful and _practicable.s—H.
HASSELBRING.

WAcHTER® has endeavored to solve some of the problems arising from the
investigations of PuRIEWITscH on the autodepletion of storage organs. The
latter author found inorganic salt solutions to behave as isotonic sugar solutions
in accomplishing an inhibition of autodepletion. For this fact PUuRIEWITSCH
accounted by assuming an incipient plasmolysis of the protoplasm, and in cases
where inhibition is accomplished by dilute salt solutions he resorted to a fluctu-
ating permeability of the plasma for explanation. WAcHTER is apparently skep-
tical of both explanations. For his material the author selected the onion, expect-
ing to avoid such difficulties as are associated with the inversion of insoluble
storage products, being able thus to deal only with diosmosing substance, which
was supposed to be chiefly glucose in the onion. He finds, however, that glucose
is not the chief storage product; that other not directly reducing carbohydrates
are present in fully equal amount, and that they exosmose into water or into salt
solution in far greater proportion than does the glucose. For this fact the author

5 By an oversight, only a portion of this review appeared in the BOTANICAL
GAZETTE for May, p. 371.—Eps.
6 WAcHTER, W., Untersuchungen iiber den Austritt von Zucker aus den Zellen
Speicherorgane von Allium Cepa und Beta vulgaris. Jahrb. Wiss. Bot. 41: 165-220.
g. I. Igo5.

1905] CURRENT LITERATURE 429

remains unable to account. Attempts to correlate with external conditions the

storage organs is possible, while PuRrEWwITsCH believed complete depletion pos-

sible. WACHTER believes from his results that the density of the external medium

is not a factor in depletion, for he found that even with exosmosis inhibited the

density of the cell sap was greater than that of the external fluid. The author

remains uncommitted as to the fluctuating permeability of the plasma, and insists

that further investigation is necessary to answer this question—Raymonp H
OND

THE X-GENERATION and the 2x- eave ag is the title of an interesting
philosophical paper by Lotsy.?. The nuclei of the x-generation contains only
one-half the number of chromosomes as do the ie of the 2x-generation. Only
asexually reproducing organisms have the x-number of chromosom The

which generation should be called the x-generation? If the fern clas is the
x-generation, the prothallium is a 4$x-generation; if the prothallium is the x-gener-
ation, the fern plant is a 2x-generation. The behavior of the chromatin during
fertilization and during the reduction of chromosomes shows that the x-generation
is the primitive generation, and that the 2x-generation is later, and that its double
number of chromosomes is due to fertilization. Thus the ancient question of
the philosophers, which came first, the hen or the egg, can be answered by merely
counting the chromosomes. The 2x-generation cannot exist indefinitely, but
sooner or later must form reproductive cells in which the primitive number o
chromosomes is restored. This reversion to the x-generation consists in the
Separation of the paternal and maternal chromosomes. The process is preceded
by the pairing of these chromosomes, which have remained separated throughout
the vegetative life of the organism. Numerical reduction of chromosomes is the
expression of this pairing. Lorsy does not hesitate to apply the theory to animals,
the body representing the 2x-generation, while the x-generation is reduced to the
sexual cells.—C. J. CHAMBERLAIN.

THomson® has been investigating the megaspore membrane or coat of gym-
nosperms, naturally applying the term to the investing coat of the prothallium
as well as to the coat of the uninucleate spore. This coat is present in all gym-
hosperms except the Taxeae, among which it is entirely eliminated or nearly so.
It consists of two layers, the outer one being suberized, while the inner one is

the megaspore coat resembles in structure and composition the microspore coat.
The only exception to this general character of the megaspore coat is found among

7 Lorsy, J. P., Die x-Generation und die 2x-Generation, eine Arbeitshypothese ”
Biol. Centralbl. 25:97-117. 4 diagrams. 1905.

8 THomson, R. B., The megaspore-membrane of the gymnosperms. Univ. of
Toronto Biol. Series, No. 4, pp- 64. pls. 5. 1905

430 BOTANICAL GAZETTE [JONE

the Araucariae, where the suberized outer layer is wanting. In the forms possess-
ing the usual type of membrane, there is present a more or less well-developed
tapetum, derived from the sporogenous tissue, which is quite distinct from that
derived from nucellar tissue. Using the relative development of the megaspore
coat and of the tapetum as a basis, the author concludes that the Abieteae are the
most ancient group of Coniferales and the Taxeae the most recent; that the
Taxodieae and Podocarpeae are complex, including both ancient and recent
forms; and that = =i se occupy an intermediate position in the phylo-
genetic series.—J. M. C.

Nicotost Roncati? has published an account of the ovule of Anona Cheri-
molia. A row of four megaspores and considerable parietal tissue are formed.
The. innermost megaspore develops a narrow and much elongated embryo sac,
whose antipodals appear to be ephemeral. The first divisions of the endosperm
cells result in a series of walls across the narrow sac, which is thus divided into
a linear series of five or six large chambers. Subsequent divisions fill these
chambers with endosperm tissue. The embryo, as figured, has no definite

nsor, but is at first a globular mass of cells. The ‘‘rumination of the seed,”
characteristic of Anonaceae, is attributed by the author to the invasion of the
perisperm by infoldings, chiefly from the inner integument. If this be true,
it is entirely different from the “rumination” to be observed in Asimina:and in
Torreya. The name “reserve idioblasts” is given to masses of nutritive material
found in abundance in cells along the convolutions of the ‘‘rumination,” an
thought to supply nutrition to the embryo after the endosperm has been digested
and the perisperm contains an insufficient nutritive supply. The paper was
presented to the Academy by Professor CAVARA c

a

Jones"° has published an account of the anatomy of the stem of Lycopodium.
The general conclusions reached are as follows: The development of the vas-
cular system has proceeded along two lines: (1) as the protoxylems increase
in number, a series of alternating bands of xylem and phloem is developed; (2
the phloem is scattered through the mass of xylem in more or less isolated patches,
until a structure is produced that simulates that of Gleichenia. The first form
is developed in creeping stems, the second is characteristic of tropical epiphytes.
The young stem has a triarch or tetrarch structure; and the complex structure
of larger stems is developed from this by the division of one of the —
into two strands, and by the separation of a phloem into two portions; su
quently, another protoxylem divides, and this process, together with the splitting
of the phloem, is repeated until the number of protoxylems and protophloems
reaches the number present in the large stem. The disposition of xylem and
phloem is constantly altering, the length through which a definite portion of
tissue (as a phloem-island) may be traced varying from 0.5 to 8™™.—J. M. C.

-

9 Roncatt, F. Nicolosi, Sviluppo dell’ ovulo e del seme nella Anona C herimolia
Mill. Atti Acad. Gioenia Sci. Nat. Catania IV. 18:Mem. 2. pp. 26. pl. 1. 1904-

1° JONES, CHARLES Epwarpb, The morphology and anatomy of the stem of the
genus Lycopodium. Trans. Linn. Soc. London., II. Bot. '7:15-35. pls. 3-5. 19°5-

a

. r " n a 8s * in
ir rah tee a ie gn Ei es eg ee

1905] CURRENT LITERATURE 431

Rose has published a fourth paper under the title ‘Studies of Mexican and
entral American plants.’’'' The contribution involves twenty-three families,
under which the miscellaneous titles are grouped. Two new genera are described,
one (Ornithocarpa) in Cruciferae, the other (Raimannia) in Onagraceae. The
new species described belong to Ostrya (2), Synthlipsis, Thelypodium, Lepidium,
Echeveria (2), Ribes (7), Neptunia, Cercidium (3), Parosela (12), Lupinus (16),
Indigofera (4), Phaseolus (3), Aeschynomene, Cologania, Crotalaria, Harpalyce,
Willardia, Erythroxylon (3), Cedrela, Polygala, Vitis, Heliocarpus (3), Tilia (2),
Abutilon (3), Kosteletzkya (2), Robinsonella (2), Ceiba (2), Ayenia (2), Melochia,
Taonabo (3), Heterocentron, Conostegia, Monochaetum, Hartmannia ( 3)s
Raimannia (3), Eryngium (6), Prionosciadium (5), Arracacia, Coulterophytum,
Ligusticum, Museniopsis (2), Oaxacana, Roseanthus, and Schizocarpum. The
new genus Raimannia replaces Oenothera as recently cae twelve species
being given as belonging to it; and the name Oenothera replaces Onagra. There
is also a synopsis of the Mexican species of Ribes.—J. M. C.

Oakes Ames’? has begun a sumptuous series of publications dealing with
the taxonomy of Orchidaceae. Its purpose is “‘to illustrate, from type material
when possible, new or recently described orchid species, and species heretofore
inadequately figured; to publish the original descriptions of all species so figured,
with additional characterization, full synonymy, and geographical distribution;
to furnish descriptions and descriptive lists of orchidaceous plants, which may
prove useful in the study of regional floras; and to communicate the results of
critical investigations among special genera.” In the first fascicle nineteen
species are described and illustrated, five of which are new, the plates being drawn
by Blanche Ames and made by the Heliotype Company. There are also the
following contents: A descriptive list of orchidaceous plants Sia in the
Philippine Islands by botanists of the U. S. Government (including sixteen new
species); an Oncidium new to the United States; Contributions toward a mono-
graph of the American mings of Spiranthes; and A synopsis of the genus Spir-
anthes north of Mexico.—J. M. C.

Krenrtz-Gertorr'3 has published a small book dealing with bacteriology.
As indicated by the title, it gives a general condensed view of the modern field of
bacteriology in its relation to the home, the kitchen, the dairy, etc. A general
introduction deals with the evolution of the modern theory of fermentation

™t Rosg, J. N., Studies of Mexican and Central American plants. Contrib. U. S.
Nat. Herb. 8: 281-339. pls. 63-72. figs. 14-19. 1995.

? AMEs, OakEs, Orchidaceae: illustrations and studies of the family Orchidaceae
issuing from the Ames Botanical Laboratory, North Easton, Mass. Fasc. I. pp.
vii+156. pls. 16. Boston and New York: Houghton, Mifflin and Company. 1995.

‘3 KrentTz-GERLorr, FELtx, Bakterien und Hefen insbesondere in ihren Bezie
hungen zur Haus- und Landwirtschaft zu den Gewerben, sowie zur Gesundheitspflege
nach dem gegenwirtigen Stande der Wissenschaft gemeinverstandlich dargestellt.
8vo. pp. roo. Berlin: Otto Salle. 1904

432 BOTANICAL GAZETTE [JUNE

as resulting from PAasTEuR’s work on spontaneous generation. An insight is
given into the relations of molds, fungi, bacteria, and yeasts to each other, with
an outline of their general morphology and classification. Modern methods of
cultivation and the introduction of pure cultures in fermentation industries and
dairies are discussed. A short chapter is devoted to the activity of bacteria in
the soil and their relation to agriculture. Considerable space is finally devoted
to pathogens, and the spread of infectious and contagious diseases. Advice and
methods are given as to their prevention and treatment. A number of illustra-
tions elucidate the text. The whole book represents a good summary of our
present knowledge of micro-organisms in a concise, popular style-—P. HEINE-
MANN.

GoEBEL"+ has investigated the cleistogamous flowers of a variety of plants
from the point of view especially of the adaptation theory, and presents an inter-
esting account of the external condition under which these occur in nature, or
can be made to occur in culture. It appears that a large number of plants have
the ability to produce cleistogamous instead of chasmogamous flowers under the
aig of certain external conditions. Contrary to the belief of Darwin and

, GOEBEL shows that there is no causal connection between the absence of
festa. or the formation of seed, and the appearance of cleistogamous
e latter is due to insufficient food relations, and this may arise from
poor soil, a lack of mineral constituents, or insufficient light. In many plants
the cleistogamous flowers do not set seed. In these cases the appearance of
cleistogamous flowers is not due to the suppression of seed-formation, but GOEBEL
thinks the seed-formation is suppressed by the appearance of cleistogamous
flowers.—W. B. McCativum.

Motiscu's has reported convincing evidence that stems of certain seedlings
respond with positive curvatures when exposed to rays emanating from sealed
tubes containing a mixture of radium and zincsulfid. The seedlings should be
less than 7™ distant from the tubes and preferably about 2°™. Tubes con-
taining radium bromid only (activity 3000) fail to give positive results, as pre-
viously found by Drxon. When radium bromid and zincsulfid are mixed,
cage a steady phosphorescence of considerable intensity develops, due to
the influence of the radium on the zincsulfid. Moriscu therefore regards the
curves pianist as heliotropic, indirectly induced by radium. The interesting
fact is also recorded that while such experiments succeed in the laboratory they
usually fail in the greenhouse. The author believes that the illuminating ges
and other volatile impurities in laboratory air reduce the negative geotropism,
thereby increasing sensitiveness to phosphorescence and heliotropic stimulus.—
Raymonp H. Ponp

4GorBEL, K., Die kleistogamen Bliiten und die Anpassunpstheorie. Biol.
Sc a ene etc. 1904.
1s Moriscu, Hans, Ueber ii ens indirekt hervorgerufen durch Radium.
Ber. Deutsch. Bot. Gesells. 23:2-7. fig. I. 1905.

1905] CURRENT LITERATURE 433

THE FIRST CASE of the regeneration of the whole plant from tendrils is
described by WINKLER.'® V6cuHTING first showed that the tendrils of Vitis
vinifera will produce roots, but on Passiflora coerulea WINKLER finds that the
isolated tendrils will give rise to both roots and shoots. Young tendrils were
cut off and placed in damp sand. They continued to live, but soon became quite
woody, and in the course of some weeks formed a callus at the base. After three

shoots in the same way when similarly treated. The fact is pointed out that
the shoots formed from the primary undivided rors continue to form primary
leaves much longer than those formed on the adult leaves. On isolated pieces
of stem internodes new shoots arise from the callus at the base——W. B. McC
LUM.

PourRIEviTcH’? shows by experiment that the investigations of BONNIER,
and Mancin which have been the basis for the prevailing opinion that the ratio

co, ; : A
©, Joes not vary with temperature include some errors, namely comparisons of
2

incomparable material, and tests of too short duration to permit the test objects
to acquire the temperature intended for the test. From his own investigations,
in which such errors and others are apparently avoided, the author concludes
ma" ; ,
that the racio — changes according to temperature, becoming greater as the
2
latter rises. This influence of temperature is most noticeable in young organs,
and depends upon the kind of nutritive substance present in the tissues. When
this nutritive substance disappears from the tissue, the influence of temperature

SPALDING showed that when roots are wounded near the tip and traumatropic
curvature prevented by placing them in plaster for as long as eight days, they will
when removed from the plaster curve in the same way and with the same reaction
time as they would have done had they been allowed to do so at first. He con-
siders that the latent period has been prolonged all this time. Burns*® has rein-
vestigated this problem and finds that so long as the wounded tissue persists the
stimulus to traumatropic curvature still exists, and unless mechanically prevented
the root will curve. When regeneration of the wounded tissue is completed the
stimulus is removed. There is thus not a prolongation of the latent period, but
a continuation of the stimulus.—W. B. McCaLium.

16 WINKLER, H., Ueber regenerative Sprossbildung an der Ranken, Blattern und
Internodien von Paieliiora coerulea L. Ber. Deutsch. Bot. Gesells. 23:45-48. 1905.
17 Pourtevitcn, M. K., Influence de la température sur la en des plantes.
Ann. Sci. Nat. Bot. IX. 1:1-32. :
2 ey sek ee and its relation to traumatropism. Beih. Bot.
Beata. a: eee figs. 4.. 1904.

434 BOTANICAL GAZETTE [JUNE

PRIANISCHNIKOW"? presents a preliminary report of sand culture experiments
which extend his investigations of 1g00 on the relative value of different phos-
phates in plant nutrition. As in the earlier work he found ammonium sulfate
to be a “‘physiologically acid” salt, so now he = contrary to all expectations,
that ammonium nitrate can also function as a ‘physiologically acid” salt. This
peculiar acidity is much less in the case of ammonium nitrate, however, than in
that of ammonium sulfate. The phrase “physiologically acid” is used in the
sense suggested by ApotpH MEveER as applied to a salt whose basic group is
more rapidly appropriated by the plant than its acid group.—RAymonD H. Ponp.

WorsDELL’”° has discussed the meaning and origin of the phenomenon ordi-
narily called fasciation, including not merely those abnormal appearances to
which the term is ordinarily applied, but also ‘“‘normal fasciation”’ as exemplified
by such stamen clusters as appear in Hypericaceae and Malvaceae. His con-
tention is that in the development of a structure there are two opposing tendencies,
the younger being a tendency to integrity, the latter being a tendency to plurality,
and that the condition of the mature structure is the resultant. The mechanical
cause i. has to do with the distribution and functioning of “growth
comtern.”——1 BT. C.

PincHot** has published a second part of his Primer oj Forestry, dealing
with the practice of forestry, work in the woods, and the relation of the forest to
weather and streams. There is also a short account of forestry at home and

a ume
sacha elementary account of practical forestry. Such literature, prepared
by those who have the most ample information, will do more than anything else
to educate the intelligent public as to the meaning and need of forestry.—J. M. C.
A GENERAL ACCOUNT of the vegetation of the island of Guam has been pub-
lished by W. E. Sarrorp.??_ The account includes some reference to every plant
known to occur on the island, and discusses the principal plants used for food,
ber, oil, starch, sugar, and forage in the Pacific tropical islands recently acquired
by the United States. The method of cultivating and propagating the more
important species is treated in considerable detail. While the paper is somewhat
miscellaneous as to contents, it is full of information for botanists.—J. M. C.

9 PRIANISCHNIKOW, D., Ueber den Einfluss von Ammoniumsalzen auf die
NEO Ss von Phansithousk sive bei héheren Pflanzen. Ber. Deutsch. Bot. Gesells.
23:8-17. 1905.

2° WorSDELL, W. C., “Fasciation:” its meaning and origin. New Phytologist
4:55-74. figs. 17-24. 1905.

2t PINCHOT, GIFFORD, A primer of forestry. Part II. Practical forestry. U. S.
Department of Agriculture, Bureau of Forestry, Bull. 24. part 2. pp. 88. 1905.

22 SAFFORD, WILLIAM Epw1n, The useful plants of the island of Guam, with an
introductory account of the Shyeical features and natural history of the island, of the
chara history of its people, and of their agriculture. Contrib. U. S. Nat.
Herb. 9:1-416. pls. I-69. 1905.

ESE a so eee

1905] CURRENT LITERATURE 435

SMALL MANUAL of poisonous plants has been prepared by A. BERNHARD
SmitTH.?3 It contains plants from all countries, with their active principles and
toxic symptoms, together with the proper treatment in case of poisoning. The
main divisions are four in number: (1) plants acting on the brain; (2) those
acting on the spinal cord; (3) those acting on the heart; and (4) vegetable irri-

ants. Subdivisions on the basis of symptoms lead one easily to the different
groups of plants.—J. M. C.

THE ENT ADDRESS of Professor D. H. Scott as president of the Royal
Microscopical Society dealt with the subject of the so-called Carboniferous ferns,?+
which comprise about half of the total known flora. The recent discoveries of
seed-bearing forms among them were reviewed, and the conclusion reached that
in all probability only a minority of these reputed ferns are really ferns, and that
probably the majority of them are seed-bearing plants. a

ARBER’S has described two new species of the sicieaas Lagenostoma, whose
chief interest is its relation to Lyginodendron. An important point in connection
with these new species is that the seeds, “with or without a cupular investment,”
terminate the ultimate branches of what is probably a modified fertile leaf —
if =e

JoHNsTON?®° has published thirty-five new species preliminary to the publica-
‘tion of a flora of the Islands of Margarita and Coche, Venezuela. Among them
there is a new genus (Anguriopsis) of Cucurbitaceae.—J. M. C
IN CONTINUING his revision of Eucalyptus, Marpen' presents E. amygdalina,
E. linearis, and E. Risdoni, under each species giving the description, synonymy,
range, and afhinity.—J. M. C.
SmitH?® has published an account of three diseases of truck crops in Delaware:
1) a leaf-spot of cucumber, squash, melon, and cantaloupe due to Sphaerella
citrullina; (2) a leaf-spot of the egg-plant due to Ascochyta lycopersici; and (3)

3 SmirH, A. BERNHARD, Poisonous plants of all countries. pp. xv+88. Bristol:
din Wright & Co.
24 Scort, me H., What were the Carboniferous ferns? Jour. Roy.
Micr. Soc. 1905: 137-149. pls. 1-3. we 32-33
25 ARBER, E. A. NEWELL, On some new species of Lagenostoma: a type of
pesdopemon seed from the Saas (abstract). Annals of Botany 19: 326-
325. x
“ Jomsrox - R., New re from the islands of Margarita and Coche, Vene-
zuela. Proc - Acad. 40
27 cues : pa A ee revision of the genus Eucalyptus. Part VI. pls. 4.
ig a by the authority of the Government of the State of New South Wales.
1905. 25. 6d.
8 Surru, C. O., The study of the diseases of some truck crops in Delaware. Bull,
Del. as, . 70. pp. 16. pls. 2. figs. 6. °

436 BOTANICAL GAZETTE [JUNE

a leaf-spot of the bean and cow pea due to Phyllosticta phaseolina. Brief descrip-
tions are given in each case of the fungus itself and its behavior in pure cultures
and inoculation experiments. No experiments in the treatment of these diseases
are reported. The paper should be of interest to mycologists.—E. MEAD WILCOX.

~Rotrs?9 has published his second contribution to the knowledge of a potato
disease, called the rosette by SELBy,3° due to the sterile fungus Rhizoctonia.
The author has conducted extensive experiments both in the treatment of the
disease and in the study of the life history of the fungus. He finds that the cor-
ticium or fruiting stage of the fungus develops freely upon the living stems of
the diseased plants, but considers the sclerotial bodies which are common on both
stems and tubers the most important agents in the distribution of the fungus.
Formalin treatment of the tubers to be planted improved the appearance of the
crop but reduced the yield. The corrosive sublimate solution treatment gave
good returns when the land employed for the crop was new. Liming the soil
and treating the ‘‘seed” tubers with sulfur gave negative results. The following
recommendations are also made: (1) clean tubers should be carefully selected
from healthy tubers for planting, (2) all vines and stems of weeds should be gath-
ered and burned at the end of the season, (3) care must be taken to see that the
fertility of the soil is maintained, that through proper methods of cultivation the
soil is well aérated, and that where irrigation is employed the water is applied
to the land at frequent intervals and in proper amounts, taking care not to allow
the soil immediately about the forming tubers to become too wet.—E. MEAD
WILCox.

29 Rotrs, F. M., Potato failures. A second report. Bull. Col. Exp. Sta. 91.
Pp. 33- pls. 5. 1904. See Bull. 70 Col. Exp. Sta. for the preliminary report upon this
subject.

3° Sze Bulletins 139 and 145 of the Ohio Expériment Station.

NEWS.

F. C. NEwcoMBE has been ne: to a full professorship of ore at the
University of Michigan

F. E. Clements, en of Nebraska, has been appointed associate pro-
fessor of plant physiology

PRoFEssor F. W. OLtver, University College, London, has been elected a
Fellow of the Royal Society.

Jutius WIESNER, priate s of Vienna, has been elected member of the
Royal Danish Academy of Scienc

N. WILLE, University of Chis has been elected foreign member of the
Academy of Sciences at Stockholm.

Dr. F. DELPrNo, ra ae of meer in the University and Director of the
Botanic Garden, died at oe

. CAROTHERS, University of a has been appointed instructor

in apa at Rockford oe. “Bockfor d, Ill.

F. Grace Smirn, University of Chicago, has been appointed instructor in
botany at Smith College, Northampton, Mass.

P. H. Otsson-SEFFER, Stanford University, is to spend three months in the
province of Soconusco, Mexico, in experimental work upon the Mexican rubber
tree

F. D. Heap, University of Nebraska, has been appointed botanist of the
Agricultural Experiment Station and Associate Professor of botany in the Univer-
sity School of Agriculture.

Dr. W. B. McCattum, University of Chicago, has received the first Walker
prize from the Boston Society of Natural History, the title of his paper being
“Physiological analysis of the phenomena of regeneration in plants.”

Tue Brorocicar STATION of the University of Montana, under the director-
ship of M. J. Elrod, has announced its seventh annual session at Flathead Lake,
Bigfork, Montana, extending from July r2 to August 17. The work in botany
will be under the direction of Thomas A. Bonser (University of Chicago) of
Spokane.

THE MINNESOTA SEASIDE STATION, under the directorship of Professor
Conway MacMillan, has announced its fifth session at Port pay I a: British
vageine on the Straits of Juan de Fuca. The session extends fro uly 8

o August 18, and the botanical staff oT of Conway eid Albert :
Sie, E. Tilden, and F. K. Butters.
1905] 437

GENERAL INDEX.

The most ne aged classified entries will be found under Contributors, Per-

Sonals, and Reviews. ew names

and names of new genera, species, and varietics

are printed in pold-face type; synonyms in italics.

Abbé, E., agg of 240
Abutilon
Acacia Greggii, water-conducting system
OL

Acer

‘Achiya, DeBaryana 61; polyandra 6r

Achromatic ap in Pellia, Gregoire and
Ber; ores on

Acrolasia, iydtiery on I

Adiantum, Christ on ste: Hieronymus

Brg genus 366; ae
ria 368; viscidula 367

N., on F

Aletes, hee on 159
Aleurone, Von Spiess on 38
Alga, chemical stimulation, Livingston on

317
Allen, C. E., on nuclear division 425
Allium, periodicity of cell division in root
18; fistulosum, chromosomes in 376
Aloitis, eesae ok on 310
Alternation of , Saya: in animals and
plants 1375 tigate
Prot ‘Society, Com-
Rye See: So; sco ng 320
Pan haa on nm Orchidace € 431
——— respiration 87; Pebaaciogaks

pane sativus, | proteolytic enzymes 409
n

JF

eimia, Dav venport on 376

Aneimites, seeds in, White on 235

Angiosperms, Bonnier on 383; early his-
tory 4

A iopsis 43

Anogra, Rydberg on 159

Anona, Roncati on 430

senuaha ossil

tic, fe floras, Nathorst on 231;
vege etation, Skottsberg on 307
Anthracnose, apple 378; ones: Hume

on 232

438

Apocynum, Greene on 159

Apogamy, in Alchemilla 312; in Hiera-
cium, Ostenfeld on 234

pee scab, Lawrence on 237

Aquilegia, Greene on 159

Arabis, ee on I ee

Arber, E. A. on cas Os 435

Archegonium of "Torreya 168

Arctostaphylos 372

Are

t.

Art

og

schou va on re leaves 71
rracac
Je é. 219, 370, 3855 personal 79,
tee aeodromus 376
Ascteaycetas, Claussen on 375
Aspidium, Hieronymus on a glandu-
losum 367; vena 367
— enium, Chris 15
Ast r, Rydberg o aes
Astragalus, Rydberg on 159; gaviotus
54; Whitneyi pinosus 54
Atkinson, G. F., pcre
lege text book of bota
Autran, =— on Nicolas “Alboft 157
Ayenia 4

“ec A*col-

Bacteria and oxygen pressure 316

ele riology, Kienitz-Gerloff on 431
romus, / ape Peg
so iain growth of, Tk
Barnes, C. R. Hien E53, 223, ny 225, 302;
308, 312, ars, By 318, 381, 382, 383;
personal, 79,
Bateson, W., p te 79
chi Greene on 310

Beal, W. J., on Michigan flora 225
Bedelian, J., on effect of greenhouse cul-
tivation 31
Beih. zum Botanisches orn nina 80
eta st Wieland o
Benson, Margar

ret, clio 240
Beighs, J., on chromosome reduction 316,
376, 377;_and Gregoire on achromatic
figure in Pellia 2

ia 227
Be ong Biological Station for research:

Berry, E. W. 231, 235, 237, 371; 3745 7
tas plants 374

[JUNE

1905]

Berthold, C., ‘Untersuchungen
Phy we der pflanzlichen Oeeea-
csi
Bertrand, N. on orchids and fungi 315
Bicarpellites
Bistorta, ‘ae on 159
Bitter, G., age onal 319, 3
0

wig, H ssytha 239
Bon si ‘hysiological a oe 348
t, G., on angio

sperms and gymno-

‘spent S 303

i hi A, personal 437
Borgese d Jen son, on Danish heath 317
Rotation Gazete length of papers
Botani sien ca of Ameri ea. a.
O, age nas 80; of ‘the Nether-

ands, jpishicavtons of 319
potas of the Central States, meeting

Reticuli, development of sporangium

Boudiera, snare on =
Bowenia, Stopes

Brabenec, F., on bed ike 374
B on on 376

., personal 2
Britton, N. i personal nes ‘a ee

37

Bromelin, effects of toxic de gon 409

Brot — s, V. F., on mo 374

Bro ra personal 2

Herbert. . heaviest pe + ete personal 79

Burgerstein, 'A., “Die Transpiration der
” I

., on heterophylly 77; on
regeneration 433

Burt, E. A. personal 80, 320

Buscalioni ego and, on proteases 323
Bush, B. F n Tradescantia 310

Butkewitach, _on Valen’ gi enzymes 323

Butters, F. K., personal 437

e

Caeoma nitens, seep 2 irc 267

Calamites, Maslen o

Caldwell, J. S. 409

Caldwell, O. W., ‘‘Handbook of plant
morphology”? 424

Calliospora

390; Diphysae 391; Far-
lowii 391;

Soiwest 390

INDEX TO VOLUME XXXIX

439

Campbell, D. H. , personal 3845 on Ophio-
glossaceae and Marsiliac e3

Candolle, C. de, on Pinericvec ea

Cannon, W. A. 39
Carbon, compounds, oxidation of go;
nutriti plants, pena on 72
Cardiff
Cardot, is on on Aho 376
Carduus mariti
Cassie rts E., eee nal 437
_— Shafer on iy wales spindle
tion, Hus
casein iltormis, Boew ig on 239
Castan va am ace, proteolytic
enzym
Castilleia, Greene on 159; Rydberg on
311; ety"
Castle, oe of SEX 304
Ceanothus, Greene. on 15 ydberg on
1595 dentatu 443 “impresus 43; sore-
diatus 4 . oo

Ge Point Lake ethos 319

i ig

Ce

Celt ge On 77; Tg: rnicke 230;
periodicity in divis n, Kellina ton 318

Celtis ig waterconductn Pia

Cera oo opes o

Cercidium 4
Chambers C. J. 685.97; 137; 227, 220,
23 ©, 233, 234, 235) 3 316, 376, 379,
429; person
ae a “Cobh on 156;
endy on 157
Chemistry ny plants, Czapek
Chemotaxis, sperm tutte es oe
sperm Ff Marchanta, —— on 237
Chlorogalum, Gre
Christ, H., on new apse “of xe 159
n Tson

Christma » 207: I 320
hromosomes, Kowalski on oe in
Allium fistulosum, Berghs on 376; in
Convallaria majalis, Berghs on 377;
st , Berghs on 316, Gregoire

316, Strasburger on 31 os
Ehepiier, M. A. 236, 239, 3
— sopsis, Rydberg on Ae californica

Ehveutdnion Greene on 159; Rydberg
on 311; ibosus 50

°Q

on ecology of New Zealand 155;
Waimakariri River basin 157

440

Col, A., on vascular bundles 2 36
Sonnet ape Rydberg on 311
ogani

of fi rs and insects 68

Comandra, Fe aeons n 376

vari tthe a Jeffrey on 306;
src on N 305

Conos

Co ntibutors ran y. C. 219; 370, 385,
Barnes, C. R. 81, 153, 223, 224, 225,
302, a 312, 315, 38975 328, 381, le,

77> £375 227; 229, 230,
pe 234, 235) 236, 316, 376, 379, sim

; Christman, A. H. 267;
M.A eat, sae 382;
68, 73, 75, 76, 77, 78, 158, 225, 226,
301, 302, 306,.31o, 311, 326, a wd
374, 376, 380, 382, 383,

Cowles, H. C Y 72, T5bu) rey, 220,
233, 234, 236, Pare 238, 239; Darbi-
shire, O. V. 356; Davis, B. M. 61, 64,

71, 301, 375, 380, 1355 Dean, A. L.
42 El ie. Dy : Ganong,
W. FL 3145 a FO, 1302:

: sselbrin

413, 214, 31S, 301 ..425- eae i
4313 —— sk. 0295 Jéefirey, EC: 305;
Jensen, G. H. 70, 317; Land, W. JG
74; Coulter and 161; Livingston

22, 1A 237) ig i 316, 297, rag ear
Florenc
W. R.

179, 307; P. ma a

ia 232, 238, "378, 428; Sargant, t, Ethel
20; Shoemaker, D. N. 248; ” Shull,
6. HH. 30 3, 309, Bai Phaser, Rear:
Trow, A. H. Tru and

#

Convallaria majalis, chromosomes 377
ar, ; ae as Schively on Epiphegus

Cooley, Grace E., on Lari
on Philippine ferns 302

ae not 378
n sero ie nd gynodioe

hybrids 3
pa bai pauciflora, development 262
Coulter, J.

420, 439, 431, 434, 435; personal
and Land 161 ieee ie

BOTANICAL GAZETTE

[JUNE

Coulter, S., ie 320
Coulterophytu m 431
se villea tridentata, water-conducting sys-

m 402

ane H. C. 69, 74, 155, 157, 226, 333,

pha 230, 237, 238, 239; personal 79,

orreya 161

ee Makino on 376
Crotalaria 43
Crowell, J. E, singe 240
Cryptant the, Greene
Cryp ce

Cucurbita, maxima
331; \Fepo, lien ko enzym

Cucurbitaceae, embryology, Fave on
ie Bea tube, pres .

Cummings, Clara .. persona

Cyanophceae, hag: n 228; "Phillips on

acharias on 22

Pos a, Hieronymus on 159

pleas ee i White on 374

Cycads, ai Stopes 229

Czapek, F., Sais reaction o

eopistie saneineaks 308; “Biochemie
der Pflanzen”’ 223
D
Dalla Torre, C. G. de, and Harms
“Genera Sh euaatienns 374
aan 0g O. V. 6
n, F., perso
De ln Carota, proteolytic get 329
ses eg a rt, C. re n Aneimia, 376
, B. M. 4, 71, 301, 375) 3

35; sittne! 320; s, oie oy ta

321
‘Deneanien simplex 48
phinium, Greene on 159
Delpino, F, death of 4
7 OLE Chatham Island 157
Dershau, M. von, on nucleolar material

Desert plants, water-conducting systems

s, H., person

DeVnie nal 384
Diatoms fertilization 380; Karsten on

Diels, L., on new genus 310; and Pritzel
6

ecatheo: . on 310
Draba, Rydberg on 159 _

1905]

abble, E., on anatomy of palms 382
Dracaena 372
Durand, E. J.; eae 320
Durand, T., personal 384
Puma P., on Patagonia 158

P
F. S., perso:
Ea ast ina ae on Th robivie 393
Echeveria
Ec ge Ganong on 233; Tansley on

Edaphic a in Flathead- Valley

Elaphoglossum, Hieronymus on 159
A.D.

Elm . 42

Elrod, 'M. in personal 437

Embry , Ly on 311; of Hamamelis
259; of Torreya Ai

Emerson, R. n heredity in bean
hy 304

brids
Enalus acoroides Svedelius on 76
Encelia acton
Encephalartos, Stopes on 229
Endosperm in Hamamelis 258
Energesis 9
eo aa Prantl, “ Pflanzenfamilie in’?
2257 374
prea ni of bromelin 409; pro-

Epilobium: “Rydberg on 159
i

pe egus virginiana, C and Schively

Epiphy: » Ule on 238
a ch Peletrisot on 380
Eriksso o

tnst, A rsonal 240
oi Soe 431
Erysimum, Rydberg on 159

Etysieueske. ke forms, Salmon on

uclisia

Euphorbia, Millspaugh on 159

Eus ene tewart and, on potato
diss

Erythroxylon
ida lay ‘Maiden on 226, 435
E n 310

Evcintien, Teeualé on 301

F
Farr, Edith M., on new iy orga 310
Fasciation, Worsdell Nn 434
awcett, W. and Ronde on Lepanthes
6
—— Margaret C. ‘Life history of

Fermentation 88
Fermi and Svacelbont on proteases 323

INDEX TO VOLUME XXXIX

wine M. L., on Comandra 376
ona

Fertilization, in diatoms 380; in Staphy-
lea 383; in Se Pac een 382; in Sapro-
legniales 61; in Paper a 161

Fischer, ia ate anstcgenall 370

H. n geo 3
eae Valley, i. forests 99, 194,
6

27
Flower-formation, Loew o

Foods, reserve of trees. "Sablon on 315

Forel on flower colors and insects 69
Forestry, oe on 434

Forestry Quart 160

Forests of F. ated Valley, Montana, go,

194, 276; o n, Sheldon on 226;

pene a of a ] pie 32; types in
Michigan 28

Fossil plants, Perkins 0

Fothergilla gardeni, gts iil

Fouquieria splendens, OT ST
ote 403

Fragaria, ger on 378 |

Friedel, Jea ,on
BAe ta

sch, K., on ase Ne canis 380
Fucgia Alboff on 157

G

Galium, ae on 159
Galloway, B. T., 1904 report

eae male,

el R
Geddes, W. N.,

me Ry oti on rs

Georgia flora, Harper

Geotropism, Fitting on em Wendie on 311
Gepp, A. E. S., on Leptosarca 376
Gerassimow, J. J., on growth of Spirogyra

oak Rydberg o

Giltay, hee on ower paciie and insects 69

Gleaso AL ew species 159

Glome rpitcaini, sah on

Goebel, K., = ist ‘praia flowers 432;
mn 314

sonregencrati Stephani on 376

Goroschankin, J., death of 160

Gorup-Besanez, on proteolytic enzymes

442

Gramineae, Studies in 123

Graves, H. S., personal 80

Green, J. on selon ia enzymes 322

Greene, E L ew genera and species
159, 310,

i
Gregoire, V., hromosome reduction
316; and coke on aches figure
in Pella 227
Gregory, R. P., personal 80
Grindelia, Rydberg 0 on 311
Grout, A. J., ‘‘Mosses with a hand lens”

ate
oon rate, Hering on

nogramme, Christ on 1 te Hierony-
15

osperms, Bonnier on 383; mega-
spore ie of 429
m, Correns on 304
Gynoglottis, S Smith on 376

H

Hallier, E., death . 240

Hamamelis, arborea, development ~
virginiana, development i embry
25

9g, endosperm 258, nogeny of
flower 250, pollen tube ee, spermato-
genesis 252
s, H., Dalla Torre and ‘‘Genera
Siphonogamarum” 374
Harpalyce 43%
Harper, R. A 5 fyi 80, 3
Harper, R. M., on flora of Govcs 239;
on new psc 159
Hartman

Hartz, N. sa eae plants 232

Hasselbring, H. -76, 312, 313, 314, 318,
37%) 425

Heald, F. D., personal 4

Heath, Danish, Hikacea cua Jensen on

edgcock, C. G., personal 320
Heinemann, P.
Helianthus, Gleason on 159
Heliocarpus 4 431
Penne ger and ra

eci

Hemale Ww B., on Thiseltonsa 376
Heredity, Castle on 304; Correns on 304
Hering, G., on rate of growth 377

— iy wi aaah edish deal 226

ee on 432

Het

Heterophly Burns on 77

Hicoroides 3

cium , spogamy in 234; partheno-
genesis 23

BOTANICAL GAZETTE

[JUNE

Hieronymus, G., on new species of ferns
15?

Hitchcock, A. $., personal 240,

Hollick, Arthur, personal 80; in cet
ceous flor, 237

Holm The ae

Waris Rydberg on 159
Hol E. W. D., personal 385
Homalobus Rydberg on 159
Hooker, J., Pon 240
Horkelia Rydberg li 50

Howe, M. A., pers
Hume, H. i, anthracnose 8 the pomelo

Has, ‘i. ir a, aries oe on spindle

Hypakiaoon ashen on 304; Lock on
3035 rage studies in 302; Tschermak

By; A Hieronymus on 159

I

ndian Forester 319
nang 431

Insects and: flower colors 6
eeaiemah ae ogue "of Scientific
Literature
soetes, Fie acts fs sperms 381
Istvanff, Gy. de, on root rot of vine 76

be ier Sead ate 76

Jeffrey, E. a 384; on
anatomy = Conifer es 2a

Jennings,

Jensen, G. H.

Jenson, Borgesen ak on Danish heath

Fokus: J. R., on Venezuelan plants 435
Jones, C. E., on anatomy of Lycopodium

430
Jordan, ~ S., on ae 309
Jordan . On
Heaters, Pe Strobili i in 235
K

Karsten, G., on microspores of dia
a and Schenck Weoctationsbilder”

Klerman, W.A., — 384

Kellicott, W. E., on periodicity of cell
division 318

Kew Index 68

gta rloff, on bacteriology 431;

n flower re ‘and insects 69

1905]

Kirkwood, J. E., on embryology of Cucur-
bita
vas saat Newcombe ” ia
niep, H., réle o

378

Kn TEV Deis; on intercalar protoplasm

Koernicke, M, reductio
3813; on lant cain, one

Kohl, F. G., personal 80;
phyceae 228

/

on Cyano-

Komaroy, “Flora Manshuriae’”’
H., and Valeton, ps ae

ava
eat a 431
Kowalski, J., on mitosis 379
L
Lactolase, Staklosa, on 317
Laetiporus, Murrill on 310
Lagenostoma, Arber on
Land, W. J. G. ulter

747 Co and 1
—_ W. H., on Scuaitanis of pain
hte: Green

Laurent, oh on carbon nutrition of green
plants
Lawrence, W. H., on apple canker 378;

- fa ad — on 381

pidiu
piesa Gen on 376

um 4
Lilium canadense, nuclear division 425

INDEX TO VOLUME XXXIX

443

Lycopodium, anatomy of stem, Jones on
430
Lyon, Florence 313; personal
H. L. 365;

Lyon, on Shad donot
embryo 311
MacDouga 1, D. T., personal 80; on dis-

continuous variati lon 309; on mutation

399
Machaeranthera pi nosa 49
MacMillan, C., personal 437

372; 376
Mie double fertiiation 3

Mangrove association, Sc a
Marchantia, chemotaxis of pean
Marine Biological Laboratory, Woods
Holl 31
k ., personal 319
st ean affinities with Ophio~

glossaceae 313
slen, A. J., on Calamites 382
oe Stephani on 376

Massart, J., personal 384
Maury, P., on fossil at 2 me
Maxon 66; perso

W. R. 3 al 79
McCallum, W. B75; 73; ee 434; 423%
a ota 320, 437
Meadow formations in Flathead Valley

194
oe wall of gymnosperms, Thom-
429

Li innaeus, Olsson-Seffer on
gee Mpls deg Oe Pap oe = 238, 297,
16 3; personal 80; on chemical

3
Pee n 317
Lloyd, F. E., personal 79, 384; on pollen
tube 78; and Bigelow ‘‘The teaching
= biology” 153
, R. H., on bamboos 77; on plant
_ breeding 78, 30

wy Ch Aan formation 238
Loma ’ Chris 159
Lotsy, J. 4 on phi ae of generations

a
us, Greene on 159
pene mopsis, “Hieronymu us On 159
Ludwigia ma , Harper on 159
Lupinus 431; se on 159; g
53

M sche bac aolt

Mendelism 3

Mercuric chien toxic action 5
Mertensia, Rydberg on 310

V spinel aig ae speeciay 43
Mesoreanthus, Gre

Metcalf, M. M., “ Organic evolution” 301
Mexican plants, Rose,

Mexico, rusts from

M ,on Glomeropitcanna 376

M ichigan, forest ty = 28; natural his~

ry survey, New me on n 237 rela~

bietaies. Greene on 310

i ntea 246; macrotis 246
, W., personal 319

Misa gh, €. ae , personal 240, 320}.

on species
Min aren ues Station 437
10

Molds, and oxygen pre! 316
Molisch, H., on heliotropism and radium.
432

“444
Monardella, Rydberg on 311; crispa 46;

TO

Monascus, Barkeri 58; purpureus, mor-
phology of 56

Mo anit sacenap 371

Monoc 431

opel trae Fritsch o

Montana, Biological Station 187 forests
of : tathend Valley 99, 194,

Moo ie soil inoculation 371, 427

Neaan T. -» on polarity 75

Mosses, Espen Roth on 22 5

Munroa squarrosa

Mutrbetke: py on > path enogenesis 234

Murrill, W. . n new genera 310

Museniopsis

oe in Alchaanittn 312; MacDougal

Nn 309

N

Nathansohn, A., on permeabilit
—— A. G., on Antarctic fossil a

ace, B., on Dini ca
Nephrodium ronymus on 159
Ne eptunia 431

Neumeister on gf spereccen es lagen
Newcom a

Nitrogen
Nomenclature, Pe Boies princi-

179
shtiea division, Allen o
Nucleolar material, Derchau on 235
Nutrition, Prianischni kow on 434

oO
Oaxaca
Octocnemaceae, Van — on 376
Oenothera, Léveillé on
-» True an I

- 56, 228, Nay Dersonal 320
F. W., personal
Olsson Sefer, a 179 3073 | ceed 437;

373
Sa N., penn 79
. in Torreya 166
ene: affinities with Marsili-

€ 313
Opulaste, ken on 159

in Hamamelis 2
rnithocarpa 430
Oroxylum 372

BOTANICAL GAZETTE

: Parasitism, Marsha

[JUNE

saan C. H., on apogamy 234

n, rom .» pe sine Pe
alana of carbon c unds
Ox ~ygen, effect of eaticten, Friede l on
318

7

Pachy mn oe: on 159

Pachys ma, Farr

cl ‘Drabble 5 oat Stenzel on
231; polyem in 83

ll Ward

Pa oe eS enetieducthing Gees 403

Paros ee

onbennake SSUS 372

ali eet ange in Alchemilla 312; in
‘Van and Hieracium, Murbeck
on 2 Sas in oe ela Winkler on

a9
Peck, C. H., personal 320; on new fungi

37

Peirce, _ les ne 160

Peletrisot, C. N., on Ericaceae 380

Pellia, “achromatic fae. Gregoire and

erghs on 227

ip sl D. P., on anatomy of Coni-
S$ 305; on ‘fossil plants 374

Pinas Diels and Pritzel on 376

Pentstemon, Greene on 159; Rydberg on

31I
Pérez, on flower colors and insects 69
sap A ai on Philippine flora, 302;
€ 302

Perkion. G. “a .. on plants of Tertiary

Pensaret lg of —— membranes,
Nathansohn o

Abbé, 2 rae Arthur, J. C.

; Atkinson, G. F oe Barnes,

Cc. R. 79, 319; Bateso W. 79; Ben-

Personals:
79, 320

son, Margare 240; Bitter, G. 319, 384;
Blackman, F. FP 719; Bonser, T. A. 436;
Brefeld, O. 160; Bristol, C. L. 319;

‘Fe big Soca ings, Clara E. 240;
Darwin, F. 79; Davis, B. M. pon 320;
Delpino, F. 436; DeVries, H. 384;

Durand, E. J. 320; Durand, os ty 2)

Earle, F. S. 320; Elrod, M. =e 436;
Ernst, A. 240, 384; Fernow, B.

160; Garber, J. F. 79; Ga a a

1905]

ete el 160; Graves, H. S.
80; ge tg . P. 80; Haley, E. 240;
Har rper, . 80 0, 320 B.D.
436; nae G.G. nae res
A. S. 240, 384; Hollick, A. 80; Holway,
E. W. D. 385; Hooker, J. 240; Hottes,
C. F. 320; we, A. 240; Hus,
'T. As 1605, Jeffrey, BE. C. 384; Jen-
nings, O. E ; Jordan, E. O ;
Kelle n, . 384; Kohl, F
80; Livingston, B. E. 80; Lloyd, F. E

ta,

a. 240, 320; Newe es 436;
Olive, E. W. 320 liver, oy ; Ww. ei
436; Olsson-Seffer, P. 436; io, N,
79; Overton, J. i Peck, C. H.
320; Peirce, G. J. 160; Pfitzer, E. 384;
Pond, R. 320; Renault, B. 80;
Rusby, H. H. 240; Sargant, Ethel 240;

ar. 152-70 553203

Tra M. 320; Trelease, W.
80, 320; Uhiworm, O. 80; Unde d
E. "M. 80 80; Ward, H.

436; Wile, N. 436; id olfe,
Wi ard, 4 S. 79; Wylie, 19
Petraschewsky, 1 4 OL Bokion: Tespira-

mes
H6

Peziza macrotis 246

Pflanzenfamilien, Engler and Prantl 225
Phaseolus 431; Mu ungo, proteolytic en-
zyMes in 330; vulgaris, proteases 332
Phellodendron 3 oo F
aa hile Rydberg on 159

henol, toxic ac

Philippines, sie of 5
Phillips, O. P
eciosum, sexual repro-

Phra atone deglubens
i resi eographic panes. principles

Pigments 1 Tschirch on 317
‘“Primer of Forestry” 434

ter
S$ 372; Murrayana 283; ponderosa

Pla nt breeding, Lock on 78, 303
Plant World, purchase of 79

INDEX TO VOLUME XXXIX

445

Plateau on flower i and insects 68
el ree aoe

in roo -

myces, Murrill on s

Poiso sar plants, Smith on 435

Poisons, Bi vd Pa i rs caeiaiioes on
to sn

Polari 7 ee n 75
rae Pas ion
Pollen tube, n Cucurbitaceae and Rubi-

aceae, Llo es n 78; in Hamamelis 255
Rolyerihnray3 in che 383; in Sphag-
m 36 “8

hoe.
“re oda, Christ on 159; Hieronymus

ne Hieronymus on 159; acule-
um 366; Lonchitis 366; i. 366
Pond, R. a 73, 232, 238, 378, 4283

personal 32
serene. To on growth and oxygen pres-
su
Pitta See ca Hume on 238; Rolfs on
36; Stewart and ure on 372
“oles

Pourievitch, M. 5 ae Heap On 433

Prianischnikow D, on ans and
lant subi 434

Prionosciadium 4

Pri a el, E., on ee eee 310; Diels and

n new genus 3
Petes erpinaca palustris, Burns on hetero-
phylly 77

Prosopis “om water-conducting sys-

mereary Be an on 321

Proteids, chemistry of 89

Proteolytic enzymes

aoe
Protoplasm, ieicallober, Kny on 236
Prunoides 371

Psammomoya, Diels and ne on 310
Pseudocymopterus, Rydbe 159
Pseudohydnum, Rick on I os

oe
Whit e on 374; Aneimites

235
re 221

Radium and heliotropism, Molisch on 432

consarnas ae on Sisyrolepis 376

Raiman 31

Parente, Brongniartae 3 306; cassiaecola
394; expansa 393; gracilis 393; incon-

spicua 395; i oeieae nar laevis

446

3953 Beer 303; Lysilomae 3092;
Mimosae-sensitivae 394; Pithecolobii
3943 "pule cherrima 395; siliquae 393;
iii 3965 eee 395; Talpa 396;

Rege eran — on 433; Goebel on
314; Winkler on
Rehder, A., on new species 373
ehm, H., on new is es “of fungi 159
Renault, B., geal nal
A, B., F on and, on Lepanthes

O., on bisporangiate _ strobili

Resorcinol, toxic action of ro
Respiration, anaerobic 87; and combus-
on 83; cours

aceae,’ (431)
Atkinson’s “A college sen boo “ f bot-
: “Untersuch-

Pinus” 67; Fischer’s “
zerland”’ Aes Grout’s ‘‘ Mosses with a
hand len 373; Kienitz-Gerloff’s
“Rekterien ey Hefen” 431; Lloyd
and Bigelow’s “The carat . biol-
alf’s volu-

153; Metc

tion” 301; Pinchot’ $s “Primer me for-
estry” 434; Sargent’s “Manual of
trees” 301; “Trees and shrubs” 372;
Schneider’s “‘Tllustriertes Handbuch
der Laubholzkunde” 373; ith’s

Poisonous plants of all countries” 435;
rem “ mbers of commerce and

ntification” 373
‘Shien, Rolfs on 4 3
odophyceae, Sys organs and sporo-
phyte generation
Ribes ast “Heller on ge
— > a new genus of fleshy fungi

Ridille, cay on cg aie 383
Robertson, Agnes, o rreya californi
316; _Sargant a. on poco ney in Zea

Mai
chinacacle
Rolfs, F. M., mo potato fa aie 436
senate F. ‘N. Ano:

oot press

ed, ies types a. —— on 239
Rosa, ppg Shy
Rose, J. N igs eS and Central

American Plants 4 431
Roseanthus

BOTANICAL GAZETTE

[JUNE

Roth, G., on European mosses 225
Rubiaceae, patie tube, Lloyd on 78
Rubioides 371

Rubus, Hiscchand on

Rumination, in pret 430; in Torreya

Rusby, H. H., saga 240

Rusts, from Mexico 385; sexual repro-
duction oc
Rydberg, P. A., on new species 159, 310

S

— L. du, on reserve foods of trees

ene Christ on 159

Saffo - : E., on Ht of Guam 434
Salmon, E. S., on biologic forms 313
Sopindoides 3 371

alan oes Meee salice 61, 300
ga er Ethel 420; personal 240; and
a rtson, on scutellum in Zea Mais

Silene, C. S., on a new Phellodendron
373; “‘ Manual . trees” sot; “Trees
and shrubs”

Schenck, H., ak and, “‘ Vegetations-
bilder” 238
Schively, pes F., Cooke and, on Epi-

Schmidt, J., on the mangrove association
°

Schneider, A., personal 437
Schneider, C. ‘‘K. Illustriertes Handbuch
= ay gee atl 373
Scott, D. H., on Carboniferous ferns 435;
on n strobits of Fe macon salto 76
-J.0

new eae iy
Shaw G. Ra on new p! S 373
cc L., personal o9e
Sheldon, E. P., on peta Fotents 226
hibata, K., on chemotaxis 381

., on flo

Shull, G. H. 303, 309, 312
Sidalc cat Resihincavalg 159

ne o 150
Silver nitrate, ‘ete — 5
Simons, Etoile B., nal
Sisyrolepis, Radler o on 376
Skotts —. C., on South Atlantic and
Antarctic a etea 307
Giclowekin Rydberg on 159
Smith, A. B., on poisonous plants 435
Smith, C. O., on diseases of truck crops
435

1905] INDEX TO VOLUME XXXIX 447
Smith, F. Grace, personal 437 Taonabo 431
Smith John Donnell, personal 160. zo mating parthenogenesis 234
Sm ie io” on Dracaena 373; on Telium 2

enc oglottis 376 Thaxter, R ee personal 320
Smit hsonian ree presentation of ies Scincia 31

herbari Thermometric Oia bate eaety on 232
ayes of Plant Rsged yw _ Physi- Thiselton-Dyer, W. T., personal 68, 24

Soil, inoculation rising on ae —s ‘ge
ti st types 32; relation t

erg on 311
Solidago, Rydberg on 311
Sophia, - ene on ee aoe on 159
Spaulding, P., per.
Spade in Hc enetie 2co; (an
= gin 162

Spe hemotaxis 237, 3
Sphacrales Rydberg on

m, polyembryony 36
ie noe Ahura m, anew cant wgaciatcl on 76
Spinacia maha pro a enzyme 328
Spindle formation, Hus ve
Spiranthes 43t

i cites oan AM on 238
gium of Betaeies °
Spruce formations, in Flathsid Valley 196
is oe G. O., on abso n of electro-
gnetic waves 22
Stach) Hel coed on 14 a on 310
t

Staphyle “ tts, Riddle o
ere? ,, Némec on ree “Hischler
« G., on fossil palm ee 231
Ste i aa F., on new liverw
Stewart, F. rw and Eustace, on ae,

diseases 372
Stictoclypeolum, Rehm on
tigeoclonium, physiology 297; as test
water 348
sa, J., on lactolase 317
Stone, "ident ““The timbers of commerce and

e C., personal 79; on ovules
of jeter 220

Storage organs, beonag = on 428

Strasbu urger, E., n Alc hemilla 312; on

ae ydberg on 159

Svedelius, N., on Enalus sag 76

—_ hm ooh Hesselm 226
wertia, Greene 159

oo, ep

Tansley, A. G., on ecological survey 239;
on ecology 233

20

B., on megaspore wall of

g erm an
Thymol, toxic acti
Tilden, a phine E, “peeuea 437
Tili
Tischi = fs on statolith theory 231
Saal californica, Robertson por 316;

xifolia, Coulter eg Land on

Touterea , Rydberg on 159

oxic c action, effect of Guclabie substances

Toxic agents, effects of action of bromelin
Tracy, S. M., personal te _
Tradescantia, Bush on 310

Transpiration, apparatus for observing

Transpirometer, pig. Me
urce of tel 383

7
es movements, anti-ferment reac-

Trov : 300; on fertilization in
Saprolegniales 61
rue H., and Oglev:

2 levi

Tschermak, E., on mips ek Taces 303;

on hybridizati
Tschirch, A., on prada 317; on roots
Tuberization, Bertrand on 31 5
Tumion taxifolia 161

U"

Uhlworm, O., personal 80
Ule, E., on Amazon “epiphytes 238
Underwood, L. M., perso
Uredinales, erminology of spore-struc-

tures 219
Uredineae, eae pa n on 71; Eriksson
313; sexual reproduction 267
redinium rn
Uredo Aeschyn

menis 392; sensitiva 394

Uromyces, ail ta 387; bauhinii

cola 389; erga sexual reproduction
71; Clitoriae 389; Cologaniae 387,

448

Fabae 388; Hedysari- -paniculati ae
Lupini an Medicagini s-falcatae
mexican ; montanus 386; aio
386; fetes 388; Trifolii 386
Uropyxis Daleae Eysenhardtiae
390 Ni ssoliae 391
Ustilagineae, Clinton on 314
Vv
has ot neygerseist on 311
n Tieghem, Ph., on Octocnemacae hi
Variation, Padinhaee sich MacDougal
tries Ye eet Col on
Ss s Johnston on 435
Vine, as rot, “Eacntt 6

Vines, , on abo enzymes 325
Viola, House on 159
S$ 431

oo Spiess, K., on aleurone 383
Vuillemin, P., on zygospore membrane
318

W
Wachter, W., on storage organs 428

Waller, A. D., on blaze currents 382
Warburg, O., on Philippine plants 30 302
ipo H. M., personal 79; on parasitism

Willesley College, gift to 79

BOTANICAL GAZETTE

[JUNE

bai D., on fossil Lesioty 374; 375; on
A355

90, 194, ot i

Wieland, G. R., on Bennettitales 159
Wiesner, J., personal 437; on leaf ber 381
Wikstroemia, parthenogenesis in
Wilcox, E.\M. 232, 237, 238, ee 378,

43
Willardia 431
Wille, N., personal 43

7
Winkler, H., on parthenogenesis 236; on
regeneration 433
Wolfe, J. J., personal 319; on Nemalion
6

eee E. A., on rock-soil
hee

Woo S., personal 79

Woredl a ree on fiecintion 434

Wri 8

ng yros 7
Wylie, R. B., 78, 424; persona al

319
455 ericana 246; gi-
gantea 246; riders 246

Z
Zacharias, E., on Cyanophyceae 229
Zahlbruc kner, ae on lichens 374
Zamia, Stopes
Zea Mais, Sar; at ad Robertson on 382
Zizyphus lycioides, water-conducting sys-

m 4
Zygospore membrane, Vuillemin on 318

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