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BOTANICAL GAZETTE 


Per) 


THE <= 


BOTANICAL GAZETTE 


EDITORS: 
JOHN M. hE The University of Chicago, Chicago, Il 
CHARLES R. BARNES, The University of Chicago, biicaie: Ill. 
ARTHUR, Purdue University, Lafayette, Ind. 


ASSOCIATE EDITORS: 


CASIMIR mei 
Gen 


j...B. DET 
University of Padua. 


ADOLF ENGL 
University of Berlin. 


LEON GUIGNARD 
L’Ecole ‘de Pharmacie. 


Rosert A. HA 
University of Wisconsin. 


JINzO ge tibial 


Fritz Nou 

University a Bonn. 
VOLNEY M. SPA 

University ab Michigan. 
ROLAND THAXTE 

Harvard University. 
WILLIAM TRELE 

Missouri Tonto Garden. 
H. MARSHALL 

University of Cambridge. 
EUGEN. WARM 

ihivetiey of Copenhagen. 


mperial Dubversity, Tokyo. 
VeIT WITTROCK, Royal Academy of Sciences, Stockholm. 


VOLUME XXVIII 
JULY—DECEMBER, 1899 
cy 


WITH TWENTY-SIX PLATES AND TWENTY-SIX FIGURES IN THE TEXT 


CHICAGO, ILLINOIS 
PUBLISHED BY THE UNIVERSITY OF CHICAGO 


1899 : fom \ 
AP i 
f rm, oH ; 
| dee ilies BE 
Mo. Bot. Garden, Tyo 
1900. a WA 
=. i 


PRINTED AT 


eee The University of Chicago Press ¥ 
i CHICAGO 


TABLE OF CQR IE i: 


PAG 
Studies on reduction in plants (with plates 1-v1), . - Geo. F. Atkinson I 
Flowers and insects. XIX, - - - - - - Charles Robertson 27 
The origin of the leafy sporophyte, - - - - John M. Coulter 46 


The development of the microsporangia and microspores E 
of Hemerocallis fulva (with plates v1, vit), Edward L. Fullmer 81 


The spore-mother cell of Anthoceros (with plates 1x, x), Bradley Moore Davis 89 


The structure and oe eam of mean area tenerum 
(with six figures), . . Le Roy Abrams 110 


The compound oosphere of Albugo bliti (with plates xI-xv), F. Z. Stevens 149, 225 


A bacterial disease of the sugar beet (with plates xvI-xx), 
Clara A. Fuaniechon 177 


Revision of the North American species of Tephrosia, - B. L. Robinson 193 
Note on the development of the holdfasts of certain Floridee 

with plates XXI—xxXIII and five text-figures), —- - Carrie M. Derick 246 
On the toxic effect of deleterious agents on the germina- 

tion and development of certain filamentous fungi, - Jj. F. Clark 289, 378 
The development of the microsporangium and microspores 

in Convallaria and Potamogeton (with plates xxiv, 


XXvV), - - - - - - - - Karl M. Wiegand 328 
Some Rocky Mountain Chrysothamni - - oie Aven Nelson 369 
Studies in Crategus. I, - - - - - - ‘C. D. Beadle 405 


BRIEFER ARTICLES — 
The seedlings of Jatropha ashok: L. and Persea gratissima 
Gartn. (With six figures), Theo. Holm 60 
On the blight of Sorghum, - - Maxime Radais 65 


- Frank Haines Lamb 69 


‘ 


‘ 
' 
' 


Root suckers on Douglas fir, 


Notes on travel. I, II, - - - - David G. Fairchild 122, 203 
Some species of Tetraneuris and its allies - - - Aven Nelson 126 
Pycnanthemum verticillatum, a misinterpreted mint, M. L. Fernald 130 


Three new Choripetalze from North America and Mexico, 
B. L. Robinson 134 
VOLUME Xxvill] v 


~vi CONTENTS [VOLUME XXVIII 4 
The probable causes of the poisonous effects of the darnel : 
(Lolium temulentum L.), - - - P. Guérin 136 — 

Section G (Botany), A. A. A. S., Columbus Meeting, Charles R, Barnes 207 
Botanical Society of America, - - - - Charles R. Barnes 210 
Botanical Club, A. A. A. S., - - - - . - - - | 2i2 

The sexuality of the Tiloptetidaceze - - - Camille Sauvageau 213 
Flower visits of oligotropic bees, - - - Charles Robertson 215 
Quercus ellipsoidalis in tows; - - - - - BT. ee ee 

A newly observed station for Galinsoga hispida, - - B.L. Robinson 216 


A practical reform in the nomenclature of cultivated plants, 
Wilhelm Miller 264 


The botanical garden and institute in Padua, . - J.B. De Toni 268 
Contributions from my herbarium, - - . - W. W. Ashe 270 
Two new Michigan fungi, -— - - * : - B.0O. Longyear 272 
Mexican Fungi. II, “ - = - - EE. W. D. Holway 273 4 


An _ hermaphrodite a 28 in Preissia Seca eg 


one figure), - e B. Townsend 
Some plants recently introduced into Florida, - - WM. L. Fernald 
Some peculiarities in Puccinia teleutospores, —- - H.. H. Hume 


What is Prunus insititia ? « 3 - a a P. A. Rydberg 


Notes on Thorea (with plate xxv), 
George G. Hedgcock and Abel A. Hunter 


Note on corn smut - - - : - - A. S. Hitchcock ’ 
A botanical art gallery, -  § - - - - Conway MacMillan 430 
A new Lilium - - - - - “ - ° C. W. Hyams 431 


‘OPEN LETTERS— 
A new Tilletia on Oryza - - - - . ; F.S. Earle 138 
To bryologists  - - - - - - - John M. Holsinger 275 
CURRENT LITERATURE — 
For titles see index under author’s name and Reviews. Papers noticed in 


“Notes for Students” are indexed under author’s name and subjects. 


NEws — 78, 143, 223, 285, 368, 444. 


VOLUME XXVIII] CONTENTS vii 


DATES OF “PUBLICATION. 


No. 1, July 29; No. 2, August 24; No. 3, September 25; No. 4, November 21; 
No. 5, November 30; No. 6, January 10, 1900. 


ERRATA. 


P. 28, line 1, first column for 17 read 1(?) 

P. 112, line 8, for and in read and as in. 

P. 130, line 6 from above, insert after no. 3142. 

P. 138, insert as heading: A New Tilletia on Oryza. 

P. 180, line 19 from below, for beest read beets. 

P. 180, line 9 from below, for 130,000 read 13,000. 

P. 201, line 13 from below, for Rugel read Shuttleworth. 

P. 251, line 16 from below, for XXI, fg. 50 read XXII, fig. 3- 
P. 260, line 11 from below, for — read 436. 

P. 261, line 7 from above, for plantlet read plantlets. 

P. 261, line 8 from above, after of omit a, and for carpospore read carpospores. 
P. 261, line 9 from below, for holdfast read holdfasts. 

P. 262, line 2 from above, for Scytosiphoul omentarius read Scytosiphon lomentarius.. 
P. 272, line 17 from above, for Bi/tmore read Raleigh. 

P. 274, line 12, for Airtefolit read hirtifolit. 

P. 352, line 8 from above, for 7g. go read fig. 39- 

P. 353, line 2 from above, for fig. ¢7 read fig. go. 

P. 353, line 12 from above, for fg. g2 read fig. 47. 

P. 353, line 5 from below, for fg. 43 read fig. 42. 

P. 354, line 6 from above, for fig. gg read fig 43. 

P. 359, omit line 8 from below. 

P. 359, line 7 from below, for 7g. go read fig. 39. 

P. 359, line 6 from below, fig. 47 read fig. 40. 

P. 359, line 5 from below, for fig. g2 read fig. 47. 

P. 359, line 3 from below, for fg. 43 read fig. 42. 

P. 359, line 2 from below, for fg. ¢¢ read fig. 43. 


Botanical Gazette 


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fol. XXVIII, No. } Issued July 29, 1899 


CONTENTS 


TUDIES ON REDUCTION IN PLANTS (witH PLATES I-VI). Geo. F. Atkinson — - - I 


LOWERS AND INSECTS. XIX. Charles Robertson - is eae - - - 27 
HE ORIGIN OF THE LEAFY SPOROPHYTE. John M. Coulter - - - - 46 


SRIEFER ARTICLES. 


THE SEEDLINGS OF JATROPHA MULTIFIDA L, AND PERSEA GRATISSIMA GARTN. som 
Ftol: 


SIX FIGURES). TZheo. m 60 
ON THE BLIGHT OF SORGHUM. Maxime Rada - - : - - 65 
| Root SucKEeRs ON DoucLas Fir. Frank Hand Lamb - - - - - 69 
a 
SURRENT LITERATURE. 
- BOOK REVIEWS ° a * # 5p > - ~ - 71 
GENERAL PHYSIOLOGY. SoME PoPULAR Books 
MINOR NOTICES 3 Fe ¥ 2. * < M4 - - - 74 
NOTES FOR STUDENTS * " : " * ‘ : : : 74 
NEWS - - - - - - - - - - - - 78 


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ee ee eee 


VOLUME XXVIII NUMBER «1 


MOTANICAL: “€ZAZETTE 


FULY:.1899 


STUDIES ON REDUCTION IN PLANTS.: 
Gro. F. ATKINSON. 


(WITH PLATES’ I-v1). 


I, REDUCING DIVISION IN ARISA-MA TRIPHYLLUM BY RING- 


AND TETRAD- FORMATION DURING SPOROGENESIS. 


Iv connection with some studies carried on under my direc- 
tion upon the embryology of certain of the Aracez, an effort 
was made to investigate carefully the nuclear figures during the 
formation of the pollen from the mother cells of the arche- 


Sporium. In Arsema triphyllum the figures accompanying the 


heterotypic divisions are of such interest that it seems desirable 
to offer the results of the study as a contribution to our knowl- 


-edge of this subject. Especially is this so because the evolu- 


tion of the heterotypic figures is so divergent from that 
described for most of the plants heretofore studied; and in one 


‘phase coincides so plainly with certain of the types found in 
-animals ; while in another phase it departs, so far as I have yet 
-been able to determine, from any type heretofore described. 


Early in July the young leaves and the spadix of Arisema, 
which are to appear the following spring, are formed, though 
quite small. They may be found by breaking the aerial shoot 


‘from the corm and removing several enveloping fleshy bud scales 


which protect the next season’s leafy shoot. The stamens: and 
pistils appear at this time as very minute protuberances on the 


' Paper read before the Botanical es of America, at the Boston meeting, 


‘August 1898. 


—! eng 


2 BOTANICAL GAZETTE [JULY © 


surface of the spadix. During the latter part of July and early | 
in August these become more prominent and begin to take on © 
the character of stamens or pistils, as the case may be.. The ~ 
staminate plants can usually be determined during August and 
in September by the dried remnant of the staminate flower, — 
which in many instances remains attached during the season; or — 
by the cleft in the petiole of the leaf where it emerged. The 
pistillate plants can usually be determined by the presence of 
the forming fruit, though some of the smaller pistillate plants of © 
one season become staminate for the next season, because of 
exhaustion in seed bearing. The archesporium of the staminate _ 
plants develops during July, August, and early September ; or it 
may be mature by September. : 

The material for this study was collected and prepared by — 
myself during September 1897. The staminate plants were : 
brought to the laboratory and the material was fixed on the day 
of collection. The spadix was first removed from the bud so _ 
that the anthers were entirely exposed. One or two anthers — 
from each spadix were removed and crushed on a glass slip in a ! 
drop of water for examination under the microscope to deter-_ ; 
mine the stage of development. In this way all material where 
the pollen was already developed could be discarded, and only _ 
that preserved which was in one or another stage of mitosis. é 
This made it possible to obtain good sections for study on a 
large number of the slides prepared. The nuclear figures can be @ 
quite readily seen in the living condition, but if there was any 
difficulty in determining the stage of mitosis, a drop of a solu- 
tion of chloral hydrate was added, as suggested by Humphrey | 
The spores are formed centrifugally on the spadix, 2. ¢., the sta~ 
mens at the base of the spadix are usually more advanced than 
those toward the apex, so that if the pollen was just formed in 
the lower stamens, the upper ones might show pollen mother 
cells in one or another of the stages of division. There was, 
however, considerable variation in the same stamen in this 
respect, while in a single locule there was little if any variation 

The material selected in this way was placed without further 


eS nO SS ee eee ae Ae ee ee See Te SS a ee 


4 
Z 
: 
: 


yee 


1899] STUDIES ON REDUCTION IN PLANTS 3 


preparation in Flemming’s chrom-osmium-acetic solution for 
twenty-four hours. It was then washed in running water from 
twelve to twenty-four hours, dehydrated with grades of alcohol ; 
decolorized by placing for twenty-four hours in bulk in hydrogen 
peroxid and alcohol, made by using 70 parts of a 95 per cent. 
alcohol and 30 parts of hydrogen peroxid. It was then stored 
in 75 per cent. alcohol after one rinsing. Six months later a 
portion of the material was placed in the hands of a student 
(Miss Susie P. Nichols) for study, and three months later the 
remainder was prepared on the slide for study by Mrs. W. A. 
Murrill under my direction. It was imbedded in paraffin, cut 
with a Minot-Zimmermann microtome 6.6—13.3 m thick, stained, 
some in Flemming’s triple safranin-gentian-violet-orange stain, 
and some with iron hematoxylin. For the present description 
of the material, and for the conclusions presented, the writer 
alone is responsible. 

During the formation of the spirem the linin network gradu- 
ally disappears, and a more or less continuous thread is formed. 
A distinct spirem was not observed, though this phase of mitosis 
has not yet been sufficiently studied. Even at the time of the 
longitudinal cleavage of the thread it was seen to anastomose 
more or less, where, in its windings, portions came in contact. 
At this time it presents the appearance of a very open, irregular 
network. The chromatin granules, at first distributed over the 
linin network, gradually accumulate in small masses scattered 
along the thread. As the spirem thread becomes more distinct 
it shortens, and accompanying this the small masses of chro- 
matin increase in size, giving an appearance of a string of beads, 
or they present an irregular moniliform appearance. Inaddition 
to this beaded appearance of the thread the masses of chromatin 
are more or less irregular or angular. At some of the angles of 
the chromatin masses short delicate threads are attached, prob- 
ably remnants of the linin reticulum which do not participate in 
the formation of the spirem. This, with the angularities of the 
chromatin masses, gives a somewhat ragged contour to the 
spirem. Quite early in the formation of the spirem, and before 


4 BOTANICAL GAZETTE [JULY 


the reticulum has completely disappeared, the thread shows a 
longitudinal splitting which is evident between the chromatin 
masses. In the early stages of the longitudinal splitting of the 
thread I have not seen the small chromatin masses divided. 
Each chromatin mass often. appears to be made up of two 
smaller ones lying side by side, which have fused at the points of 
contact. Even in larger chromatin masses, and among the chro- 
mosomes in Arisaema, there is a strong tendency for those lying 
in contact to fuse more or less, so that the individuals of a single 
mass or of the chromosomes are often more or less obscured. 

While the chromatin masses on the spirem are more or less 
fused, they are small and emarginate at opposite ends, and at a 
point coincident with the axis of the band. By this character 
one can distinguish the double nature, or paired condition of the 
masses. It suggests a division of the original masses, or the 
deposition of the chromatin in paired masses along the thread, 
which fuse more or less at the point of contact. These chro- 
matin masses often stain unevenly, two portions separated by 
the axis of the thread or band taking on a deeper stain. The 
darker points lie in pairs along the thread, each separated by a 
paler chromatin area which runs for a short distance on either 
side along the division of the thread. At the point between the 
deeper staining masses the paler zones fuse. 

As the spirem shortens, these paired chromatin masses are 
brought nearer together. Fusion among them takes place, so 
that the masses increase in length and breadth as the spirem 
broadens. The longitudinal division of the spirem is marked 
here and there, sometimes at quite regular intervals, by rounded, 
elliptical, or oblong openings in the chromatin, while in other 
places, for considerable distances, no line of division can be 
ascertained, though often the difference in the intensity of the 
stain marks the separation of the masses into pairs, while the 
paler surrounding zones are fused. As the band shortens, the 
contour becomes more irregular, showing crenations and irreg- 
ularities, to which are often attached delicate strands, perhaps 
portions of the linin reticulum. 


> See 


renee 


sloyren ae 


ste hs ae 
SAI Sa ae Sia reser oreo pee eed, 


it a a al a el ee ee a ee ee 


Pe ee ee Wee 


1899] STUDIES ON REDUCTION IN PLANTS 5 


The chromosomes are marked off by the band separating into 
sixteen segments. So far as could be determined, this number 
is quite constant. The segments are not formed by an abrupt 
division of the spirem, but it appears as if the chromatin masses 
by shortening unite themselves into elongated areas, which are 
connected for a time by two delicate threads representing the 
original line of longitudinal division. The chromosomes lie 
around the periphery of the nuclear cavity and are quite variable 
in form; this is due to variations in the extent to which longi- 
tudinal fission has proceeded. In a few, longitudinal fission 
is complete, and a pair of long knotted rods is present. In 
others, there is a slight connection at one end or near the middle 
by the fusion of adjacent chromatin. In others, one or both 
ends of the flattened chromosome are indented or forked; in the 
latter case figures approaching the letters Y or X are formed. In 
many of the chromosomes the line of fission is shown by one 
or two small rounded openings or narrow open slits near the 
middle or near the ends; or the opening is long and extends 
from end to end, individuals of the pair being connected only at 
the end, thus resembling long irregular chain links. Some of 
the individuals of the pair lie parallel ; others are separated more 
or less at the middle, when quite well defined rifts are formed. 
Still others resemble chain links twisted half way around, thus 
presenting a figure 8 appearance (jig. 6). 

The length of the chromosomes is now from one third to one 
half or more the diameter of the nuclear cavity, and they show 
the irregularities presented by the spirem band, being knotted or 
angular along the edges or at the end. Delicate linin threads 
are often still attached at the knotted or angular enlargements. 

The chromosomes now shorten, the chromatin becoming 
massed together, until their length is from one sixth to one fourth 
the diameter of the nuclear cavity. A larger proportion of them 
have now opened out in the middle in the form of rings, and, while 
they have shortened in length, their breadth is about the same as 
formerly, or they may bea little broader. This gives a more per- 
fect ring form, but the chromosome is still somewhat longer than its 


6 BOTANICAL GAZETTE [JULY 


breadth, so that the line of longitudinal fission can be determined. 
The line of longitudinal fission is also indicated in other ways. 
Other chromosomes at this time are present in the form of short 
plates, some of which show slight indentations at the end, while in 
others longitudinal fission is complete, and a pair of rods is formed. 

During the process of shortening of the chromosomes, the 
greater part of the chromatin usually becomes massed at four 
definite points in the ring, or plates, or pair of rods. These 
denser masses of chromatin lie near the ends of the rods, or near 
the ends of each lengthened half of the ring or plate, and are 
connected by paler staining areas, the density of which varies in 
different rings. These four masses of chromatin, in the ring or 
in the rods as the case may be, form the tetrad. These rings 


and tetrads in Arisema triphyllum are very distinct, as much so 


as in certain animals. The two lengthened halves of the rings 
are usually more nearly separated along the line of longitudinal 
fission than along the line of transverse division, so that they 
quite readily separate. Each half of the ring is more or less 
crescent shaped, and the ends, instead of being rounded, are 
often quite regularly angular, presenting figures that frequently 
aid one in determining the orientation of the cleavage line. Up 
to this time the chromosomes still occupy their position around 
the periphery of the nuclear cavity, and, up to quite a late 
period, often show fragments of linin attached. 

The nuclear membrane now disappears, and the kinoplasmic 
threads, which are to form the spindle, enter and move the chro- 
mosomes upto the nuclear plate. The threads first radiate 
irregularly in all directions, but converge more and more at two 
poles as they gradually form the spindle. As this takes place, 
there seem to be two centers of force which lie at the poles of 
the spindle, and occasionally this is manifested a short time 
before the spindle is complete by radiations of kinoplasm about 
the poles in the form of rays, which suggests a centrosphere 
figure (jig. 74). This was observed very plainly in one case, 
and perhaps similar figures in other plants have led observers to 
interpret them as centrospheres., 


ee Pe eee ee ese er 


1899] STUDIES ON REDUCTION IN PLANTS 7 


As the chromosomes are drawn to the nuclear plate they lie 
in various positions. At first it was thought that the axis of the 
chromosome, whether a ring, a pair of rods, or plate, lay perpen- 
dicular to the axis of the spindle. I have observed them in this 
position a short time before the spindle is complete. In some 
such cases, the position of the spindle threads suggests that the 
chromosomes are pulled around before division so that the axis 
is parallel with the spindle axis. In other preparations, the 
chromosomes, where they lie close together, tend to fuse to such 
an extent that it isimpossible to determine in what position they 
lie at the nuclear plate, z.e., whether parallel with or perpendicu- 
lar to the axis. In a large number of preparations conditions 
are such as to lead me to believe that the axis of the chromo- 
some is parallel with the axis of the spindle. The chromosomes 
are so numerous and of such size that they cannot all lie at the 
periphery of the plate, but occupy the center of the nuclear 
plate as well, so that an end view of the nuclear plate shows the 
chromosomes quite evenly distributed over this area. In many 
cases a large number of them are fused, and curious figures are 
thus presented. At other times they lie entirely separated, so 
that the individual rings, or pairs of rods, can be seen and 
counted in an end view of the nuclear plate. Where the section 
is cut so as to show an oblique view of the plate, the groups of 
tetrads, or pairs of rods, can be well seen and counted. From 
studies of sections in this direction, it appears that there are 
sixteen groups of tetrads, which would make sixty-four indi- 
vidual chromosomes, thirty-two to be distributed to each daughter 
nucleus of the first division, and perhaps sixteen to each 
daughter nucleus of the second division. A side view of the 
spindles of such preparations, at the nuclear plate stage, shows 
the long axis of the rings or pairs of rods to be parallel with the 
axis of the spindle. Since the chromosomes are distributed 
through the center of the nuclear plate as well, they lie like a 
bundle of chain links. The form of the ring, as suggested 
above, indicates that the axis of the longitudinal cleavage of the 
spirem or of the tetrad lies parallel with the axis of the spindle. 


8 BOTANICAL GAZETTE [JULY 


This is important in determining the character of the first divi- 
sion, as to whether it is to be a longitudinal or a transverse 
division of the chromosomes. 

Not only does the form of the ring aid in determining the 
position of. the chromosomes at the nuclear plate stage of the 
first division; the paired rods also serve to determine this. 
These lie parallel with the axis of the spindle. Even before the 
rings have moved to the nuclear plate there is a great tendency 
for them to separate into longitudinal halves. This occurs also 
at the nuclear plate stage. The tetrads separate more readily 
along the line of longitudinal division’ than they do on the line 
of transverse division. For this’ reason, at the nuclear plate 
stage, the figures with paired rods, rings partly or completely 
separated along the. line of the lengthwise cleavage of the 
spirem, together with the form of the rings, should determine 
the position of the long axis of the chromosomes. The tufts of 
spindle fibers are attached to each individual of the tetrads, and 
transverse or reducing division of the chromosome results as 
these are drawn away from the nuclear plate. In many cases 
the sixty-four tetrads, thirty-two to each pole, move as distinct 
individuals, so that longitudinal, as well as transverse division, 
occurs in the first mitosis of the mother cell nucleus. But while 
transverse division occurs during the first mitosis of the nucleus, 
longitudinal division of the chromosome takes place first. The 
peculiarity of Arisaema, then,.is that, while longitudinal division 
precedes transverse division of the chromosome, both divisions 


occur during the first or heterotypic division, and the real reduc- | 


tion follows soon the pseudo-reduction in the heterotypic 
division, 

The only other account of a reducing division in the first 
mitosis which I have noticed is that by Korschelt in the study 
of the development of the egg of the annelid Orphryotrocha, 
quoted by Wilson in “The cell in development and_inheri- 
tance,” p. 201. During the first mitosis in oogenesis, the spirem 
segments into four chromosomes, which is the normal number 
in the somatic cells. These split, and then-the pairs fuse 


Sa in 


ia epee eer a aes 


eae tee aa ee = 
Pe Bait Ae ee) ceca By Bt oad 


7 
| 
| 
| 


1899 | STUDIES ON REDUCTION IN PLANTS 9 


together again, the four chromosomes arranging themselves in a 
tetrad, two going to the first polar body and two remaining in 
the egg cell. These two split and form another tetrad, two 
chromosomes going to the second polar body and two remaining 
in the egg nucleus. This brings about reduction in the first 
mitosis. 

Beliaieff? for a number of years has maintained that a reduc- 
ing division takes place in plants, and, based on his studies of the 
division of the pollen mother cells of Iris, classifies the modes 
of division of the nucleus in plants into three types: (1) the 
vegetative division; (2) the heterotypic division; and (3) the 
reducing division. The heterotypic division takes place, as is 
well known, in the first mitosis of the pollen mother cell, and 
the reducing division accompanies the second mitosis. 

Calkins? announces reducing division during the second 
mitosis of the pollen mother cell in Pteris and Adiantum. 
Calkins was unable to determine whether the reducing division 
in Pteris and Adiantum occurred during the primary or secondary 
mitosis, since the tetrads were packed so closely in the nuclear 
plate of the first division. During the secondary mitosis the 
diads elongate to forma rod. This elongation he thinks takes 
place in the direction of the axis of the spirem, and would thus 
show that the primary mitosis is an equation division, while 
reduction takes place during the secondary mitosis. Neverthe- 
less he recognizes that it is immaterial whether reduction takes 
place during the first or second division. The formation of tetrads 
in the prophase of the primary mitosis is generally regarded as the 
preparation of the number of the chromosomes which are to be 
eventually distributed to the four daughter nuclei of the second 
mitosis; and where the tetrads are formed by one longitudinal 
and one transverse division of the segment of the spirem, it is 
equivalent to reduction in the prophase of the first division, 

? BELIAIEFF, W.: Ueber die Reductionstheilung des Pflanzenkernes (Vorlaufige 
Mittheilung). Ber. d. deutsch. bot. Gesells. 16 27-34. 189 

3Chromatin reduction and tetrad formation in nesta ees Contrib. Dept. 
Bot. Columbia Univ. 5: 101-115. i. 295, 296. an 


Io BOTANICAL GAZETTE [JULY 


though usually, so far as accounts go, the final separation for the 
reduction does not occur until the second division. In Arisaema 
the mode of tetrad formation by longitudinal and transverse 
division is such as to effect reduction in the prophase of the first 
division, while during metaphase and anaphase the reduction is 
completed by the separation of the tetrad of the transversely 
divided diad. 


II. REDUCING DIVISION OF THE CHROMOSOMES IN 7R/LLIUM 
GRANDIFLORUM DURING SPOROGENESIS. _ 


In studying the development of the pollen of 7yil/ium grandi- 
florum, the chromosomes were found to be of such large size as 
to offer, possibly, a good opportunity for an investigation which 
might throw some light on certain of the problems of mitosis 
during sporogenesis. The preliminary studies were made upon 
material collected during the autumn and winter of 1896-7. 
With the experience of one season’s work it was possible to plan 
for the collection of a quantity of material in different stages of 
development. The young flowers of Trillium begin their develop- 
ment in Juneand July ofthe previous year, and by autumn they are 
well formed, though they remain protected in the sheathing bud 
scales during the winter. So far as I have found, the arche- 
sporium is completed and the period of growth of the pollen 
mother cells is closed with the oncoming of winter, if not before. 
The period during which the division of the pollen mother cells 
takes place extends over seven or eight months. This is due 
not only to some variation in the time of maturing of different 
plants in different localities, but also to temperature. In the open 
woods, on rather high ground, where the plants are protected 
from cold north and west winds and are exposed to the warmth 
of the sun, the pollen is often mature in the latter part of 
September. In the cool ravines and open places exposed to 
north winds, the pollen is formed in warm days during February, 
March, or April, according to the season, and to some extent 
also according to the state of maturation in different individuals. 

Very cold weather checks the progress of mitosis, but when 


1899 ] STUDIES ON REDUCTION IN PLANTS “EE 


the mother cells have reached the proper state of maturation a 
few warm days in the winter or early spring months are sufficient 
for the two successive divisions. It appears that during tne 
winter season, when the nights are cool and the days warm, the 
mitotic figures may be prolonged for several days. For this. 
reason it is not difficult to prepare a large quantity of material 
in different stages of division. For the present study the 
material was collected during the month of February 1898. 
Some of this was growing in a garden where it was transplanted 
the year before, while the bulk of the material was collected 
from wooded ravines, from one to two miles distant from the 
laboratory, the plants being easily found by raking off the leaf 
covering on the ground. 

All the material was examined before fixing in order to 
know just the stage of division in individual plants, and even 
assorted into lots showing the spirem stage, first and second 
division, for convenience in cutting material of any desired stage. 
During division the chromosomes are so large and the cytoplasm 
so clear, one can easily determine the different stages with a 
one sixth objective, so that desired material could be assorted 
into that showing prophase, nuclear plate, anaphase, etc. In 
each individual flower, the tip of one of the stamens was removed, 
crushed in a drop of water on the slide, and examined first with 
a two thirds objective, then with a one sixth. In this way unde- 
sirable material could be rejected. 

Before placing in the Flemming solution, sometimes the tips 
of all the anthers were cut off, while other material was 
placed in the fixing solution with the anthers closed. After the 
removal of the sepals, petals, and the apex of the pistil, so that 
the fixing solution could enter the locules of the young carpels, 
the entire andreecium still attached to the receptacle was thrown 
into the solution. This made it possible to section several 
anthers of an individual flower together. 

The fixing solution used was Flemming’s chrom-osmium-acetic 
solution. The material remained in this from fifteen to twenty- 
four hours, was washed from twelve to twenty hours in cold 


12 ». BOTANICAL GAZETTE | JULY 


running water, dehydrated in grades of alcohol to 95 per cent., 
then decolorized in mass in alcohol of 70 per cent. which was 
diluted with hydrogen peroxid. It was then passed back to 
alcohol of commercial strength, then through absolute alcohol, 
and cedar oil ; finally it was infiltrated with paraffin and imbedded. 
It was then stored in the paraffin blocks for a, few weeks or 
months as time became available for cutting and staining. The 


material was cut on a Minot-Zimmermann microtome and stained, 
some with the triple stain (safranin-gentian-violet-orange) and: 


some with iron-hematoxylin. 

The early stages of the spirem have not yet been studied, 
but the spirem forms a broad and probably continuous band. 
Before it segments, it often shows a differentiation, where not 


overstained, into a ground substance giving a pale purple or pale 


violet reaction with the gentian violet, and more deeply stained 


bodies which are in pairs and appear at quite regular intervals in 
the band. The line of separation which runs along between the 


pairs of denser chromatin masses marks the line of longitudinal 


division of the spirem, though the ground substance often shows 
no division line. As the spirem matures, here and there are 


seen short openings along the middle line. This is especially 
well marked as the spirem segments into the chromosomes. 
The individual chromosomes often show traces of longitudinal 


division, by short openings near one end, or at the middle, oftener 
at the ends, where there is a slight indentation; or, the division 


proceeding deeper, the end is more or less forked, resulting in 
Y- or U-shaped figures. 


The spirem segments into about six chromosomes. They 


are at first rather long and become somewhat shorter and 


broader as they move to the nuclear plate of the spindle. 

At this time the usual changes in the nucleus take place. 
The membrane disappears and the threads of kinoplasm move 
in, showing first a radiating arrangement and gradually moving 
to converge into the two poles as the spindle is formed; the 
chromosomes are drawn toward the center and are finally oriented 
in the nuclear plate. The chromosomes are broad, flattened, 


1899] STUDIES ON REDUCTION IN PLANTS 13 


and irregularly oblong. There. is considerable variation in size. 

They are very characteristic in form and structure. A few 
show narrow slit-like openings in the middle. A few of this 
form become very short and the opening then is somewhat 
rounded, so that a ring form is the result. This is comparatively 
rare. Others are divided at the ends, some showing a slight 
emargination, while some are more or less deeply forked, show- 
ing a tendency to form XandY figures. Combinations of these 
two types are also found, so that there will be an opening in the 
middle, while the ends are somewhat forked, or one end is deeply 
divided. More characteristic, however, is the tendency to a 
differentiation in the density of the chromatin, which is especially 
marked when the chromatin is not stained too deeply. This dif- 
ferentiation of the chromatin is of the same kind as that mani- 


fested in the spirem and shows the paired condition of the 
chromosomes. The ground substance of the chromosome in 


these cases shows uniformly a paler tint and is translucent, while 
the denser masses of chromatin stain very dark and are more or 
less opaque. These chromatin masses are paired just as they are 
in the spirem, and probably result from the division of single 
chromatin masses in the early fission of the spirem. When these 
chromatin masses are well marked in the chromosomes, they 
seem to be uniformly of the same number so far as observed. 
There are often found well marked pairs of chromatin masses, 
each mass lying in the edge of the bar, four at the ends (two at 
each end) and four near the middle (two on each side near the 
middle), making eight in all. 
Very frequently the edge of the bar is undulate, the prom- 
inences on the edges occurring at the location of the chromatin 
masses. The apparent uniformity in the number of these 
chromatin masses in the chromosomes is, perhaps, of consider- 
able interest, and one is led to inquire whether they represent the 
units of the chromosomes or whether each one is a member of a 
tetrad group. If the latter is the true interpretation, then there 
would be in each chromosome of Trillium grandifiorum two 
united tetrads, or the chromosomes in the prophase of the 


14 BOTANICAL GAZETTE [JULY 


heterotypic division would be quadrivalent, instead of bivalent, 
as in the case of the normal tetrad. 

As the chromosomes are drawn toward the nuclear plate, they 
become bent, so that each one represents a short arc of a circle 


t 


or a broad open U form, as if drawn more forcibly by threads 


attached to the middle portion. As they are oriented on the 


nuclear plate transverse to the axis of the spindle they at first’ = 


stand in various positions. Some lie tangentially, with the convex 
side toward the axis of the spindle and with both ends at the 


periphery, while others lie so that one end is directed toward © 


the axis, with the other end at the periphery. In this way 
they are sometimes ‘‘convolute” or more or less “ imbricate”’ 
in the nuclear plate, and may remain so for some time during 
the metaphase, or while separation at the nuclear plate is begin- 
ning to take place. Before separation has proceeded far, how- 


ever, the chromosomes are usually more bent upon themselves, — 


with the free ends of the U nearer together and directed out- 
ward in the usual way. 
While the monaster is forming, and many of the chromo- 


ER ERT es 


sainanaoraerpene sete Sp mere 


somes stand with one end directed toward the axis of the i 
spindle and the other radiating therefrom, certain ones which are . 
strongly lobed at the ends present a figure which leads one to 


think that the ends might be separated first, and that the 


chromosomes might then, in the anaphase, move to the poles in 
the form of aU, but with the concave or open side directed © 
toward the poles, or that the inner ends might be separated first, — 


somes has begun in such figures, and it is suggested that they 


are not yet drawn into proper position in the nuclear plate. 


As separation of the paired chromosomes begins, the larget - 


number of spindle threads appear to be attached in one tuft at 


the middle. The flattened U-shaped chromosome now broadens 
at the middle, in response to the tension of the tuft of threads" 
attached at this point on either half, and the chromatin substance 


is soon drawn out into a short process on either side which 


x 


a eR Ee ee ge ney 


1899 | STUDIES ON REDUCTION IN PLANTS 15 


becomes the point of the V-shaped daughter chromosome, or 
forms a slight projection on the convex side of the U-shaped 
ones if this form is retained. No pull is exerted on the end of 
the chromosome, the ends of the pairs remain united, or in 
position at the periphery of the nuclear plate, while the middle 
portions are drawn toward the opposite poles, and in opening 
out thus, the dividing chromosome forms the diamond-shaped 
figure when seen from in front (fig. 78). 

During the anaphase the chromosomes are V-shaped or 
U-shaped, and show considerable irregularity in contour, being 
more or less nodulose, often showing still the four dense 
chromatin masses. Some close up behind and form rings, 
some divide in front and form two rods. As the chromosomes 
approach the poles the polar ends converge, so that the chromo- 
somes lie close together around the periphery of the ends of the 
spindle. Lying in this position the daughter nucleus is formed. 
The form ofthe daughter nucleus is somewhat like the half ofa 
biconvex lens, the convexity being outward, while the truncated 
end lies toward the cell plate now formed by the connecting 
spindle threads. The form and position of the chromosomes 
give to the daughter nucleus its shape, for as the closed ends 
of the V- or U-shaped chromosomes converge at the apex of 
the cone, the spreading arms of the open end cause the nucleus 
to broaden out on the side facing the cell plate. The nuclear 
cavity now appears and the chromosomes are lying on its 
periphery against the nuclear membrane. They become usually 
more irregular in form, with angular points on the edge to which 
appear to be attached delicate threads connecting with the 
nuclear membrane, or reaching to an adjacent chromosome. 

Where the nucleus is small by the close crowding of the 
large chromosomes, it is quite impossible to determine whether 
the chromosomes unite in such a way as to form a spirem. The 
nucleus does not, however, pass into a resting stage with the 
linin reticulum upon which the chromatin is distributed, but the 
chromatin bands remain intact. In the large nuclei, where the 
chromosomes do not lie so close together, they appear in most 


16 ; BOTANICAL GAZETTE [JULY r 


cases to remain distinct, and in the form of V- or U-shaped 
bands, or often they are horseshoe-shaped, and sometimes form 
rings. In some cases it appears as if the arms of adjacent 
chromosomes coming in contact had fused at the end, but in no 
case could I see that they were thus united around the nucleus 
to form a continuous spirem, as described by Mottier for 


Podophyllum. Sometimes it appeared as if two had united by . 


their ends to form a very large ring. In other cases the 
chromosomes may be permanently separated into two groups 
‘during this period. 


As the nuclear cavity is formed and the first spindle is dis- 


appearing, the free ends of the chromosomes sometimes bend 
inward partly over the truncate side of the cavity, and at other 
times the chromosomes do not occupy the regular position which 
they usually show when they have reached the poles. In these 
cases ends of the chromosomes may be fused here and there, but 
in all stages several free ends are to be seen, and the figures pre- 
sented by the nucleus are such as to lead one to believe that the 
chromosomes remain distinct through this phase. If the 
daughter nucleus is not elongated when first formed, it very soon 
begins to elongate in a plane parallel with the cell plates, so that 


it becomes nearly or quite twice as long as its diameter, and it is 
more or less inequilateral, the convex side being toward the — 
periphery of the primary mother cell, while the plane side faces — 
the cell plate. In Opening out in this way the chromosomes 
become more and more distinct. The elongation of the daughter a 
nucleus often takes place while the chromosomes are moving t0 


the poles (fig. aiy., Ei such cases the chromosomes are more 


easily followed, and the evidence is quite convincing in support 4 
of the view that the individuality of the chromosomes is pre — 


served from the anaphase of the first division through to the 


prophase of the second mitosis. From the V- and U-shaped forms — 
possessed by the chromosomes as they go to the poles, many 
of them change to horseshoe form, or some to complete rings — 
by the free ends converging while the arms part slightly at the — 


middle portion. In a number of cases the chromosomes divide 


Peas” aera ? 


1899] STUDIES ON REDUCTION IN PLANTS 17 


transversely at the convex end and the two rods lie side by side 
or near each other. The nuclear membrane now disappears and 
the kinoplasmic threads enter to form the spindle for the second 
division. 

The elongation of the nucleus while the chromosomes are 
moving to the poles shows that the forces are in play which 
form the spindle for the second division. It is evident also 
that the chromosomes remain distinct, and that they are soon to 
be again separated transversely. 

The elongated form of the nucleus marks the position of 
the spindle for the second division, and the poles of the spindle 
lie near the poles of the elongated nucleus. The spindle is 
therefore inequilateral, the poles being curved toward the cell 
plate of the first spindle figure. At the same time the chromo- 
somes begin to move inward to the cell plate, and in doing so 
show the same general form which they possessed during the 
anaphase of the heterotypic division, which it seems they possess 
through the short period which intervenes before the formation 
of the second spindle figure. 

During this time there is no longitudinal cleavage of the 
chromosomes. Even should the chromosomes form a continu- 
ous band in the daughter nucleus of the first division, there is no 
longitudinal splitting of the same at this time. Sometimes there 
is an appearance of a longitudinal fission of the chromosomes 
where the edges seem to be more deeply stained than a middle 
zone along the axis. This was found to be due, however, to the 
deeper staining of an outer layer of the large chromosome, and in 
properly stained preparations can be seen at any stage of mitosis. 

If longitudinal fission of the chromosomes takes place during 
the second division, it would then be sought for in the nuclear 
plate stage. 

Since the chromosomes during the prophase of the second 
mitosis are of the same general form as those of the anaphase of 
the first division, and very likely preserve their identity through 
the short intervening period, they should be chiefly V- or U- 
Shaped. This is the case, as the examination of a large number 


18 BOTANICAL GAZETTE [JULY 


of preparations proves, while a few are ring form (fg. 30), and 
still others, having divided transversely at the apex of the V or 
the closed end of the U, exist as a pair of rods. 

As the chromosomes approach the nuclear plate the arms of 
the V- or U-shaped ones close together, and, usually at the same 
time, transverse division takes place at the closed end. In this 
way all the chromosomes become of nearly uniform shape, con- 
sisting of two parallel or nearly parallel rods, which become 
more or less fused along the line of contact. The result of this 
is to form a double chromosome, very broad and irregularly 
oblong, which resembles in a striking manner the paired chro- 
mosomes at the nuclear plate of the first or heterotypic division, 
though they are formed as a result of folding and transverse 


division instead of by longitudinal cleavage. As the paired — 


chromosomes are oriented on the nuclear plate, they usually 


become bent in such a way that the convex side lies toward the 


‘axis of the spindle, while the ends are directed outward, and the 
axis of the double chromosome is perpendicular to the axis of 
the spindle. The figure therefore presented by the metaphase 
of the second division is very much like that of the heterotypic 
‘division, and the only way in which one can determine that 


‘these belong to the second division is by the fact that there are 


two such spindles within the wall of the mother cell. Variations 
in their position in the monaster occur. One end of the chromo- 


some may be directed inward toward the axis of the spindle, so : ' 
that a few of them may be somewhat convolute; or the bent end : 
may lie tangentially at the periphery of the spindle; or the 
chromosome may be nearly straight and standing on one end, 
‘while the other end radiates outward. As the position of the | 
chromosome varies at the nuclear plate, so the figures presented ' 


during the anaphase vary, but the result is the same in each case. 


The broad chromosomes lying thus at the nuclear plate, their ! 


edges face the poles of the spindle; the threads of the spindle 


which pull on the chromosomes are attached to that portion of a 
the chromosome on either side which lies near the periphery of — 
the spindle. On those which are bent so that both arms of the — 


Se a 


1899] STUDIES ON REDUCTION IN PLANTS 19 


U radiate equally, the threads are attached at the middle; on 
those which stand so that the arms radiate unequally the threads 
are attached somewhere between the end and the middle; while 
those which stand on the end have the threads attached at the 
end of each edge. The pull of the threads attached at the dif- 
ferent points on the differently oriented pairs of chromosomes 
separate the individuals of the pair, the separation beginning at 
the point of attachment of the thread, and, as the portions are 
drawn toward the poles, the liberation proceeds until the mem- 
bers of the pair are no longer in contact. The different figures 
presented may be grouped all in three types, the V or U, the 
hook or the V with unequal arms, and the straight rods. These 
as can be readily seen are dependent on the point of attachment 
of the spindle threads. 

The result of this separation of the individuals of the paired 
chromosomes in the second mitosis is a reducing division of the 
chromosomes, or a qualitative reduction of the chromatin sub- 
stance; for, as we have seen, the paired chromosomes in the 
second mitosis are formed by the looping of longer chromo- 
somes, which often open out at the bent end before reaching the 
nuclear plate. The second division results simply in the separa- 
tion of the arms of this loop, and the distribution of each toa 
different daughter nucleus. It may be admitted here, then, with 
a feeling of reasonable confidence, that in. Trillium grandiflorum 
a reducing division of the chromosomes, or in other words, a 
qualitative reduction of the chromatin takes place during the 
second division of the nucleus in the development of the pollen 
Or microspores. One may well be cautioned against a hasty 
judgment in the interpretation of the figures presented during 
Sporogenesis, because of what seems to be contradictory evi- 
dence given by different investigators upon the question as to 
whether or not a reducing division of the chromosomes takes 
place in plants, and it is only after careful study of an excellent 
series of preparations that I am led to present this as my convic- 
tion of the nature of the process as it occurs in Trillium. The 
large size and small number of the chromosomes, as well as 


20 BOTANICAL GAZETTE [JULY 


their form, have contributed in no small degree to the results 
obtained. 

As is well known, the investigations of Mottier+ upon division 
of the pollen mother cells in Podophyllum, led Strasburger® to 
believe that a reducing division of the chromosomes occurred 
during the second mitosis of the pollen mother cell.. But more 
recent researches by Mottier, especially upon the condition of 
the chromosomes in the daughter nucleus and the origin of the 
chromosomes for the second division, led Strasburger® to recede 
from his former position and to reaffirm his conviction that the 
second division is the result of a longitudinal cleavage. Accord- 
ing to Mottier, the chromosomes unite in the daughter nucleus 
of the first division in such a way as to form a continuous single 
band, z.¢., a band made of single chromosomes united end to 
end, the arms of the V-shaped chromosomes and of the paired 
ones, where transverse division has taken place, open out to 
join with the arms of neighboring ones. This band forms the 
spirem, which splits longitudinally before it segments into the 
chromosomes for the second division. Asa result of this proc- 
ess of longitudinal fission of the spirem in Podophyllum, the 
segments or chromosomes are paired, but the pairs in the case 
of Podophyllum would then be the result of longitudinal fission, 


The separation of the members of the pair at the nuclear plate : 
during the second mitosis would not then be here a reducing — 


division. 


These conclusions of Strasburger and Mottier led me to ; 
study very carefully the same stage in Trillium. As I have © 


indicated above, the evidence seems to me taeShow that the 


chromosomes retain their individuality through the daughter — 
nuclei from the anaphase of the first division to the prophase of 


4MotTtier, D.M.—Beitrige zur Kenntniss der Kerntheilung in den Pollen-mut- 


si einiger Dikotylen und Monokotylen. Jahrb. f. wiss. Bot. 30: 169-204. A433 
97 


SSTRASBURGER, E.—Ueber Cytoplasmastructuren, Kern- und Zelltheilung. Jahrb. 


f. wiss. Bot. 30:375-405. 1897. 


°STRASBURGER, E. and Mortier, D, M.—Ueber den zweiten Theilungsschritt im 
Pollenmutterzellen. Ber. d. deutsch. bot. Gesells, 15 : 327-332. pl. 75. 1897. 


Se IE ee eee NY 


ss a 


sa 


1899 | STUDIES ON REDUCTION IN PLANTS 21 


the second. It might, however, be admitted that they unite to 
form a continuous spirem, without invalidating the conclusions 
reached in this study, for I have not found the slightest evidence 
of a longitudinal fission during the daughter nucleus stage. On 
the other hand the chromosomes, as the nucleus opens out, 
appear in the same form and number as shown at the close of 
the first division, that is, in the form of a letter V or U, or in 
rings, or even in the form of paired rods, which form more or 
less the same figure, and agree in this respect with the same 
types described by Beliaieff for the same stages. 

The origin and form of the chromosomes in the first division 
of the pollen mother cell of Trillium, and the figures presented at 
the nuclear plate, show that the first division is heterotypic, 
though the transverse division of the chromosomes at this time, 
which indicates the tetrad character, is rarely present. The 
figures presented by the second division are exceedingly interest- 
ing, since they suggest the heterotypic division also, as is indi- 
cated by rings, in some cases, which are formed by the closing 
of the open ends of a V or U figure. There is thus a semblance 
of a heterotypic figure, with reducing or transverse division 
during the second mitosis in the sporogenesis of Trillium, and it 
would be interesting to know if the heterotypic figure described 
by Farmer’ during the second mitosis in the sporogenesis of 
some liverworts, results in qualitative reduction. The figures pre- 
sented by the chromosomes at the nuclear plate of. the second 
division are strikingly similar to those which exist during the 
true heterotypic mitosis, and the separation of the chromosomes 
gives, during the anaphase, figures exactly like those shown 
during the first division. 

A larger number of chromosomes in the second division, 
perhaps, are of the hooked form, and some are of the rod form, 
though both the hook and rod form are found rarely in the first 
division. It would appear that in Trillium, as well as in Arisema, 
as shown by my studies on reducing division in that genus, the 


7 FARMER, J. B.—On spore formation and nuclear division in the Hepatica. 
Ann. Bot. 9: 469-523. pl. 76-78. 18 95. 


22 BOTANICAL GAZETTE fe 


form of the chromosomes and the mode of separation at the 
nuclear plate would not permit of classification into the types 
suggested by Beliaieff.* It is quite reasonable to believe that in 
mitosis, where the size and proportional dimensions of the chro- 
mosomes vary, as they are so well known to do, a great variety 
in the form of the chromosomes may exist, and that, correspond- 
ingly, there may be great variation in different species mani- 
fested in chromatin figures and evolutions, even where the same 
result is finally obtained. 

In the peculiarities of the second mitosis of the pollen 
mother cellin Z7illium grandiflorum, there is presented a distinct 
type in the evolutions of the chromosomes during the reducing 
division. Considerable interest attaches also to the peculiarity 
of the large chromosomes during the anaphase and metaphase 
of the first mitosis, where there is an appearance of eight denser 
portions of chromatin in the chromosomes, arranged in four 
pairs in such a way as to suggest two tetrad groups in a single 
segment of the spirem. In connection with this, it is interesting 
to note, that, while in Zriliium gvandifiorum during the first | 
mitosis in the pollen mother cell there are only six chromo- 
somes, in a number of the Liliacee there are.twelve. This 
indicates, possibly, that the segments of the spirem in Trillium ' 
here represent four chromosomes, fused end to end (quadri- 
valent), instead of two (bivalent), which is usually the case as 4 
result of the pseudo-reduction. Avisema triphyllum, according tQ_ 
my studies on this species, also presents a distinct type, in that — 
the reducing division, though following the longitudinal] division, © 
occurs during the first mitosis. a 

That there should be variations in the evolutions performed 
by the chromosomes in different plants, such as to represent | 
different types, is what one might expect, not only in view of the — 
great variations in the size and form of the chromosomes 12 — 
different plants, representing different types of chromosomes, but — 
also in view of the tendency to variation so manifest in many of | 


re: *BELIAIEFF, W.—Ueber die Reductionstheilung des Pflanzenkernes (Vorlaufige : : 
Mittheilung). Ber. d. deutsch. bot. Gesells. 16 27-34. 1898 t 


1899] STUDIES ON REDUCTION IN PLANTS 23 


the phenomena of plant life, extending even to nuclear phenom- 
ena, illustrated by the different types in fertilization, as shown 
by the investigations of Ikeno? in Cycas, Shaw’ in Onoclea, and 
Blackman* in Pinus sylvestris, as well as by Miss Ferguson™ in 
Pinus Strobus. 

Is it not well to inquire if some of the divergent and con- 
tradictory results regarding the behavior of the chromosomes 
obtained by different investigators, when dealing with different 
plants, are not due to the fact that we are dealing with different 
types in some cases? Are there not among plants different 
types of chromatin reduction? So that in one type there is 
represented a mass reduction, or quantitative reduction of the 
chromatin; in another type a pseudo-reduction, or numerical 
reduction only of the chromosomes; and in another type a 
qualitative reduction of the chromatin or reducing division of 
the chromosomes? Touching the hereditary or constitutional 
influences of fertilization, we recognize different types in plants, 
as shown by close- and cross-fertilization; and different types 
also in the mechanism for bringing about pollination. 

Some of the bewilderment which now surrounds certain 
phases of the study of the morphology of the nucleus will, I 
believe, disappear, if we recognize that there is such a thing as 
a reducing division or qualitative reduction in plants as repre- 
sented by such types as Trillium, Arisema, Adiantum, Pteris, 
Iris, etc.; that there are plants in which only a quantitative or 
mimerical reduction occurs, represented by such a type as Podo- 
phyllum ; and possibly that there is still another type, where 

° IKENO, S.—Untersuchungen iiber die Entwickelung der Geschlechtsorgane und 

—602 


den Vorgang der Befruchtung bei Cycas revoluta. Jahrb. f. wiss. Bot. 32 2557 
bl. 8-10. 1808. 
é © SHaw, W. R.—The Fertilization of Onoclea. Ann. Bot. 12 : 261-285. pi. 79. 
1898, ~ 

* BLACKMAN, V. H.—On the cytological features of fertilization and related 
pines in Pinus sylvestris L. Phil. Trans. Roy. Soc. B. 190: 395-426. pl. 72-74. 


** Miss Ferguson’s studies were carried on under my direction in the Bot. Lab. 
Cornell University, and a a paper, yet unpublished, was read before the Bot. Soc. Am. 
Aug. 1898, entitled “A preliminary note on fertilization in the white pine.” 


24 BOTANICAL GAZETTE [JULY 


in the same plant qualitative reduction may take place in some 
cells, while quantitative or numerical reduction only takes place 
in others. This seems to me, as a working hypothesis, more 
reasonable than to insist, because one type has been found 
in one or several plants, that all plants must conform to it. 


CORNELL UNIVERSITY. 


EXPLANATION OF PLATES I-VI. 


(All the figures are drawn to the same scale, using a Zeiss microscope, compen- 
sation ocular 12, and the 2™" homogeneous immersion objective, the image being 
down even with the base of the microscope. The figures are reproduced from 
these drawings without any reduction.) 
PLATES I AND Il, Ariz: triphyll: 


_ 


Fic. 1. Spirem stage of nucleus showing the dividing thread with chromatin 
masses on it. The masses of chromatin are partly divided, and appear as paired 
masses along the thread. 

Fic. 2. Portions of partly formed spirem, showing indications of longitudinal 
division. 

I Early stage of chromosomes, just after segmentation of spirem band, 
een different stages of longitudinal division, forming oblong plates with ends 
maaan ae bik or those approaching X and U forms, where the division is more 

ents of linin are still attached. 


gre 4, © 6 7. Different stages, showing the gradual shortening of the chromo- 


somes in some cases, in others different forms of chromosomes, in same stage 45 
those in fg. 3. In figs. g and 5, some are opening out to form rings; in fg. 6 one of 


the oblong rings is twisted to form a figure 8, and in fig. 7 one shows two openings 
along the line of longitudinal division. All of the chromosomes in these figures are 


os more or less irregular, sph ah and show ee of the linin attached. 


s. 8, 9, 10. Chromosomes much shortened, an large number of them = 


nda 
in te ring form. In fg. “9 two have suo divided to form rods, two form 


oblong plates with forked ends, and one forms a ring, while in fg. zo nearly all are in Z 
the ring form. Some of them are still angular, and show the peculiar form of somany 


of the rings of Ariseema. The tetrads are being formed by the accumulation of the 


chromatin in denser portions near the ends of the rods, and near the ends of each — 
half of the rings, the paler zone across the middle showing the line of transvers¢ 


division. Linin threads are still attached to some of the angles. The nuclear 
membrane is still intact, and the ch 


from the 


Fie: II, 12, 13. The nuclear sie trae has disappeared, and threads of king - 


m are entering to form the spindle; chromosomes in the form o 


pl . 
rods, or angular plates, which are solid or with a central perforation indicating the — 


ring form 


Se ene 


romosomes are arranged around the periphery % 
the ea eat in the positions occupied by them when they separated as segments — 


eo ee ee 


i 


i 
4 
F 
4 
: 
‘ 
: 
A 


1899 ] STUDIES ON REDUCTION IN PLANTS 25 


Fic. 14. Spindle with chromosomes approaching the nuclear plate. The poles of 

the pes show Sasa! lines of protoplasm indicating centers of force 
mosomes in the nuclear plate stage, but they are in this wigs 

tion so va ie fase it is impossible to see the way in which division takes plac : 

Fics. 16, 17. Chromosomes beginning to separate at the nuclear hae rings, 
paired rods, and tetrads can be seen. The rings and paired rods are lying with the 
long axis parallel with the axis of the spindle, pena aes division has taken place 
in some and is taking place in others, so that the halves of rings form crescent- 
shaped rods, the tetrads are indicated by oe staining portions of the rods or 
ings. 


Fics. 18, 19. A little later condition of the same (metaphase) stage; longitudinal 
division has taken place so that nearly all the rods or halves of the rings have sep- 
arated, and by the pulling of the spindle threads the tetrads are being drawn apart 
pert to the axis of the chromosome, to bring about the reducing division. 

. Polar view of the metaphase, showing that the chromosomes are 
snciines: al ‘aiciah the nuclear plate. Sixteen groups can be counted, bearing in 
mind that a pair of rods, or a tetrad, makes a group. Some of the sivonsiciaes are 
turned somewhat igi especially in fg. 20, which was an ee view of the 
plate, and in some cases the paired rods are seen only from the 

Fic, 22. The bees have nearly separated by transverse div is 

Fics. 23, 24. Tetrads, about 32 on each side, sEyrenenrer 7 ole during the 
anaphase of the first mitosi 


PLATES III-VI, Trillium grandifiorum, 
—_ 


Fic. 1. Portion of the spirem with partial longitudinal division, and a few chro- 
mosomes. 

2, 3, 4, 5. i. stage in the formation of the chromosomes just after the 
ee of the spirem, showing, in some, openings along the middle line, in 
others the ends a. saan the line of cngiteaiant division. Pairs of deeply 
Staining chromatin masses are evident in some of the chromosomes, frequently four 
ee or Meee masses in a single chromosom 

6, 7, 8,9, 10. The nuclear Sean has disappeared, and the chromo- 
Somes are being drawn up to the nuclear plate. 

Fics. 11-17. Various figures showing the metaphase stage, the orientation of the 
chromosomes in the nuclear plate ; in some, as in fg. 77 and 77, the four paired 
masses of denser chromatin are well seen; in fg. 77 the pull of the spindle ge is 
drawing out the middle portion on either side of the broad chromosome, the firs 
Step in the separation of the longit itiaatly divided seginent. 

Fics. 18, 19. Two different stages in the separation of the chromosomes in the V 
form as they are pulled toward the poles 

Fic. 20. Cell plate formed alter first mitosis, and U or horseshoe-shaped chromo- 
Somes lying in position as they approached the poles. 

Fics. 21, 22. Oblique view of two poles, to show the position of the chromo- 
Somes as they come to the poles; in fig. 27 t e group has already elongated some- 
what in the direction of the axis of the spindle for the second mitosis. 


26 BOTANICAL GAZETTE [JULY 


S. 23, 24, 25, 26. Position of the chromosomes in the daughter nucleus after 
first mitosis; nuclear membrane formed. 

Fic. 27. Nuclear membrane has disappeared at the prophase of the second 
mitosis, and the chromosomes, still showing the form and characters present when 
reaching the poles after the first mitosis, are satan, to move to the nuclear plate 
of the second mitosis. 

Gs. 28, 29, 30. Later stages as the chromosomes are approaching the nuclear 
plate. The U form is closing up by the folding of the arms, and in some, transverse 
division has taken place so that they form a pair of rods; in a few the ends of the 
arms met and form a ring, which simulates the form of chromosomes in the 
heterotypic division. 

Fic. 31. End view of nuclear plate stage in second mitosis. 

IG. 32. Side view with spindle established, the chromosomes presenting much 
the same form as in the first mitosis, from the closing up of the arms of the U and 
the cross division at the junction of the arms. 

S. 33, 34, 35. Different stages in the och of the transversely divided 
a sieet at the nuclear plate in second m 

Fic. 36. Three different forms of the renee Sethe as they are ae to the 
poles in the second mitosis; the V form; hooked, or J form: ; and the rod form, 
depending on the point of attachment of the spindle threads; all in he same 
pee 

8, 39. End view of spindle showing chromosomes approaching the poles 
at the on of the second mitosis. 


BOTANICAL GAZETTE, XXVIII. PLATE ‘T. 


ATKINSON on REDUCTION of CHROMOSOMES 


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BOTANICAL GAZETTE, XXVIII. PLATE 


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ATKINSON on REDUCTION of CHROMOSOMES 


BOTANICAL GAZETTE, XXVIII. 


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ATKINSON on REDUCTION of CHROMOSOMES 


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BOTANICAL GAZETTE, XXVIII. PLATE IV 


GF Atkinson dél. ad nat 


Lifh-AnstvE.A Funke Leipzig 


ATKINSON on REDUCTION of CHROMOSOMES 


Pe ae sere 


BOTANICAL GAZETTE, XXVIII. ‘PLATE V. 


29 28 


Fins aa os eee ae, RE 
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ATKINSON on REDUCTION of CHROMOSOMES 


BOTANICAL GAZETTE, XXVIII. 


PLATE V1. 


fh AnstVEA FUNKE, G02 


Lith AnstVitur 


ATKINSON on REDUCTION of CHROMOSOMES 


FLOWERS AND INSECTS. XIX. 
CHARLES ROBERTSON. 


1. Comparison of the genera of bees observed in Low Germany 
and in Illinois, with the number of species of each and their flower 
visits—The results credited to Miiller are taken from the Fertil- 
zation of Flowers. They are based on observations made by 
Herr Borgstette at Teklenburg, in the north of Westphalia, and 
by Miiller at Lippstadt and in Sauerland, in the central and 
southern parts of the same region, as well as observations made 
by him in Thiiringia. My results are based on observations made 
within ten miles of Carlinville. Each species of bee is credited 
with a visit for each of the species of plants on whose flowers it 
has been taken. : 


II. On the flower visits of oligotropic bees. — Those bees which 
visit a wide circle of flowers Loew* calls polytropic. On the 
other hand, the bees which restrict their visits to a few flowers 
he calls oligotropic. Cases are given by quite a number of 
authors, but, as far as I can learn, they are cited as mere curi- 
osities; and, as if to keep them more interesting by surrounding 
them with mystery, the facts which give them significance are 
omitted. The fact that a species of bee is found on the flowers 
of one or a few species of plants may only indicate that the bee 
is rare, or that the entomologist does not know where to look 
for it. Inthe economy of the host-bees (those not inquiline) 
the most important flowers are those from which the female gets 
the pollen upon which her brood is fed, and we need not trouble 
ourselves with any cases, or give special names to them, unless 
it is particularly specified that the female collects the pollen. 
The more often the female visits a flower without collecting any 
pollen, the stronger becomes the presumption that there is 

*Blumenbesuch von Insekten an Freilandpflanzen. Jahr. Bot. Gartens Berlin 
St Isea 
1899] 27 


————_.. 


28 ‘BOTANICAL GAZETTE 


Westphalia and Thiiringia Macoupin County, Illinois" 
: No. species Visits No, species "Visits 

Sphecodes . - - 17 28 12 
Prosopis - - - 15 88 7 
Colletes - - - - 4 16 14 
Halictus - - - 32 440 30 
Augochlora”- - . : _— —_ 5 
Agapostemon - - 4 
ena - - - 51 219 42 
Parandrena - - - I 
ia - - - - — I 
Panurginus - - - — —_— 9 
Perdita - - - - a= —- I 
valliopsis . —, — 3 
Rhophites - - 2 8 at 
Rhophitoides - - I 2 — 
Halictoide - I I 
Panurgus” - - 2 16 — 
Dasypoda - - - I 7 — 
ilissa - - 3 16 —- 
Macrop) . - é I 4 I 
Ceratina - - - - I 3 2 
Xylocop - - - — — I 
jucera - 2 2 I 15 —_ 
mpho a ea eae ky 
Melissodes - - — — 18 
Synhalo oe — — 4 
Xenoglossa - - - — 2 
Entechni - - — eae I 
nthophora - . 7 5 Eo ames * 5 
Saropoda - - “ I 9 os 
Melecta - = m 2 3 I 
Bombomelecta ~— - - —_ ee I 
Crocisa - - - I I — 
Epeolus - = - = 7 2 12 
Nomada- - - - 21 85 17 
Heriade - - - I 13 3 
Chelost - - = 3 25 peo 
Andronicus” - - “ ae I I 
Alcidam - - —— sess 2 
in 13 100 10 
Megachile “ - - 9 77 15 
Chalcodoma - - e I I — 
Diph is - - - I I 5 puns 
Anthidium - - - 3 16 I 
S li - - - - 3 12 2 
Coelioxys ce ea 6 28 7 
Neopasites - - « re ne 2 
Bombus - - - - 13 457 8 

Psithyrus- - - 4 52 3 $7 

Ape As - I 189 I 78 

Totale a +s - 205 1,981 251 : 
Not accounted for - 210 : 
2,191 


1899 ] FLOWERS AND INSECTS 29 


another flower which she does visit for that purpose, and which, 
therefore, holds a more important relation to her species. Accord- 
ingly, I propose to consider those examples in which the female 
collects pollen of one species, or several species of the same 
genus or natural family, the relationship of the plants being such 
as to give significance to the cases. On the other hand, if a bee 
uses the pollen of only two plants of different families, I assume 
that it is essentially polytropic, and that the few visits are merely 
connected with the fact that it is rare or has a short flight. Of 
course there still remains a strong contrast between the visits of 
a bee which flies only a month or two and one which flies through- 
out the season. As a rule, if a bee has a long flight it must be 
regarded as polytropic, unless the flowers on which it depends 
have a long blooming time. Of the thirty-nine species of Halic- 
tus and the allied Augochlora and Agapostemon, I regard only 
one as oligotropic, Halictus nelumbonis. {t has a comparatively 
short flight, while the blooming seasons of the Nymphzacez are 
long. When a genus of plants has more than one closely allied 
species, the difference between a monotropic and an oligotropic 
bee may depend merely upon the accident that only one species 
occurs in the neighborhood. My observations show that an 
oligotropic American bee will gather the pollen of a closely 
related introduced European plant of the same genus. 

The relations of the host-bees to the flowers from which they 
get pollen are quite analogous to the relations of parasites to their 
hosts, of phytophagous insects to their food plants, or of pre- 
daceous insects to the insects upon which they feed or with which 
they provision their nests. How the bees maintain these rela- 
tions is much easier to understand, since the flowers are modified 
in such a way as to facilitate their visits. 

Any ecological position is of advantage only to a limited 
number of individuals. As soon as this optimum number is 
passed, anything which will enable a set of individuals to get 
along without coming into competition with the dominant form 
will be of advantage to them, and their preservation will depend 
"pon their adopting this course. A characteristic which would 


30 BOTANICAL GAZETTE [JULY 


be a disadvantage before the optimum is reached, may be an 
advantage after the optimum is passed. Whatever may be the 
characteristic which enables this set of individuals to hold its 
own in a new ecological position, I think the principal circum- 
stance which accounts for the adoption of a new mode of life is 
the pressure of competition. The dominant form retains the 
original position, the other becomes modified (specialized) in 
adaptation to the newly acquired position. 

In my neighborhood there are thirty-five species of Andrena, 
which complete their flight from March 17 to July 14. These 
succeed one another, so that not more than twenty-one would 
be in competition at the same time, if their habits were the 
same. Ten begin their flight in March, seventeen in April, 
seven in May, and one in June. 

Of thirty-three species whose habits are pretty well knows, 
nineteen are polytropic and fourteen oligotropic, in the sens 
in which I use those terms. Four of the oligotropic species get 
pollen from plants of the same genus, but each of the other tel 
has its own flower, so there are eleven sets which are absolutely 
without competition among themselves. I think it is clear that 
so many species could hardly flourish in the same locality and 
complete their flight in so short a time, if all were in competition 
for the pollen of the same flowers. 

The average maximum flight of the females is forty-eight days 
Now suppose that, on account of the pressure of competition 
one of these shifts to a different phenological position. Of OF 
flowers whose pollen is so situated that the bee can readily col 
lect it, only those are available whose pollen is produced m 
abundance between the time the female is impregnated and the 
end of the time of flight. To use human terms, the bee mn 
choose between a limited number of flowers, and is in no wise iret 
to regulate its habits according to mere whim. oe 

From the above considerations I do not accept the views ® 
Kerner, although they are the ones adopted by Knuth? © 

* Natural History of Plants 2: 206. 1894. ae 

3Handbuch der Bliitenbiologie 1 : 106, 114. 1898. 


y ty 


BN a a a a a Oh ke a te sil i " 


1899 | _ FLOWERS AND INSECTS 31 


explain such cases. Kerner says: ‘ The flowers of the common 
bryony, Bryonia dioica, are not less remarkable. They occur on 
two kinds of plants, z. ¢., on one plant are developed only stami- 
nate and on the other only pistillate flowers, and since the pollen 
is not powdery, and therefore not scattered by wind, it must be 
carried by insects from plant to plant if the ovules are to mature. 
But the flowers, especially the pistillate ones, are very insignifi- 
cant, green in color, with faint smell, and they are half hidden 
under the foliage. Many insects fly past them without noticing 
them. They are almost exclusively visited by one of the Hyme- 
noptera, viz., Andrena florea, and it can find them in the most out- 
of-the-way places. This can hardly be accounted for except by 
supposing that the scent of bryony flowers 1s perceived by these 
particular bees and not by other insects.’’ He admits these con- 


clusions must be accepted with discretion. Andrena florea gets 


its pollen exclusively from staminate plants of bryony. How 
much better do we understand the case if we admit that the 
scent of the flowers is perceived by the bee? Tne mud-dauber 
makes its nests of mud and fills them with flower-spiders, which 
are so near like the color of the flowers which they frequent that 
they are enabled to capture their prey by lying in wait. Do we 
explain the case if we say that Pelopceus perceives the scent of 
mud and Thomisidz ? 

As for out-of-the-way places, my observations indicate that, 
as a rule, oligotropic bees nest in the neighborhood where their 
food plants occur, and that, when the brood emerges next year, 
it finds the flowers in bloom, and that near by. 

Asa typical case of an oligotropic bee, Emphor bombiformis 
may be mentioned. Both sexes occur in abundance on flowers 
of Hibiscus lasiocarpus, the female collecting the pollen, the males 
often spending the night in the flowers. The bees do not occur 
“xcept when the Hibiscus is in bloom. Within several yards of 
the Hibiscus I have seen the female making nests in a dry bank, 
oe ing water to soften the earth she was excavating. The bees 
coming out next year find the Hibiscus in bloom near by. The 
Only visits to other flowers I have seen the bees make were to 


i ULY & 
32 BOTANICAL GAZETTE [J | 


those in the neighborhood of the Hibiscus. Thus a | 
single female sucking the nectar of Cephalanthus a 1S, : _ 
another that of Vernonia fasciculata, as wellas a single male suck _ 


ee aa 
ing nectar of /pomewa pandurata. The outside visits in no y 


| ibiscus. For 
modify the essential relation of the bee to the Hibiscus 


s ok li- 
myself, I do not believe in the absolutely exclusive visits of oli 


hy 
gotropic bees to their pollen flowers, and I see ng <a "d | 
they should be expected. If the plants from which a bee g 


ay ion of 
pollen are common and widely distributed, the a 
flowers to which it occasionally resorts is much ae alll 
case like Hibiscus, Indeed, it strikes me that it is an adva 


me owels — 
for the males and unimpregnated females to visit other a 
and not interfere with the females which are collecting p 


: llen 
Some bees which stick their pollen with honey get a io 
from nectarless flowers, and so are compelled to visit other 


; 's, with pollen 
ers for nectar. I have seen Macropis steironematis, with p 


balls on her legs, sucking nectar of Melilotus alba. 


. : ts e 
_In case of this Macropis and Steironema, Kerner might sa 


the bee perceived or liked yellow flowers, but all of the ae 
visits I have seen this bee make were to white flowers, Ceanot 

Melilotus alba, Apocynum. 
bloom in the neighborhood. 


. alle 
I shall now give the cases of oligotropic bees mentioned ? 


: é rity of 
Lubbock,* on authority of Miller ; by Loew, on authority P 
Schmiedeknecht ; and by Knuth® from various sources. 
Andrena florea _ Visits exclusively Bryonia dioica. i 
hattorfiana = « " Scabiosa ( Knautia ) arvens® : 
Halictoides “ “ Campanula spp. 
Cilissa melanura “ as Lythrum Salicaria. : 
Macropis labiata " a Lysimachia vulgaris. . 
Osmia adunca . “ Echium. 7 
‘British Wild Flowers in Relation to Insects 21. 1875. 


’Blumenbestch von Insekten an Freilandpflanzen. Jahrb. Bot. Gartens 


3:274 (72). 1884 


° Handbuch der Bliitenbiologie I:114. 1898, 


: in 
In these cases Steironema was 


ee eee A 


a 
’ 


TT Cee en ee Se ee 


| 
| 
: 


1899] FLOWERS AND INSECTS 3 
Andrena nasuta visits Anchusa officinalis. 
cineraria = Salix. 
lapponica ss Vaccinium. 
cettii ee _ Scabiosa ( Knautia). 
hattorfiana 4 Scabiosa 
orea 2: Bryonia 
alpina ' Campanula. 
curvungula ° Campanula. 
austriaca ip Umbellifere. 
lucens " Umbelliferz. 
Andrena florea visits exclusively Bryonia dioica. 
hattorfiana “ “ Scabiosa (Knautia) arvensis. 
cettil . " Scabiosa (Knautia) arvensis. 
" " Anchusa officinalis. 
Bombus gerstaeckeri “ . Aconitum lycoctonum. 
Cilissa melanura almost Lythrum Salicaria. 
Macropis labiata f . Lysimachia vulgaris. 
Osmia adunca As Echium. 
cementaria -. ‘ “ Echium. 


.Andrena florea, mentioned in the three lists, collects pollen 
of bryony and has been found on no other flowers. 

Andrena hattorfiana: both sexes visit Scabiosa arvensis, the 
female collecting pollen. Miller found a female on Dianthus 
carthustanorum and a male on Jastone montana. 

Halictoides dentiventris: Miller captured both sexes on Cam- 
panula rotundifolia and trachelium, but not collecting pollen. He 
Says that at St. Petersburg Morawitz found it only on Campanula. 
In the Alps, Miller observed this species collecting pollen of 
P otentilla grandiflora and Hypocheris uniflora and visiting seven 
other flowers. i 

Cilissa melanura collects pollen of Lythrum Salicaria, the males 
sucking. Miiller saw the female sucking on flowers of Leontodon 
hirtus. : 

Macropis labiata: males and females visit Lysimachia vulgaris, 
the females collecting pollen. Males suck on Gnanthe fistulosa, 
Rhamnus Jrangula, Rubus fruticosus. . 

Osmia adunca: Miller saw both sexes on Echium vulgare 
and says it feeds its young exclusively on honey and pollen of 


34 BOTANICAL GAZETTE | JULY 


Echium, but under Vicia Cracca this species is indicated as collect- 
ing pollen. In the Berlin garden Loew found it collecting pollen 
of Nepeta Mussint. In the two latter cases there is some error, of | 
the bee is not oligotropic. So of the cases mentioned by Lub- — 
bock, only one is exclusive, and two are not even oligotropic. 

Andrena cettu: females collect pollen of Scabiosa arvensis, 

Bombus gerstaeckeri: why Knuth says this species visits exclu- 
sively Aconitum lycoctonum 1 do not know, for on page 191 itis 
distinctly stated that the males and workers visit A. Napellus, but 
nothing is said about pollen-collecting. 

Osmia cementaria: males and females suck and collect pollen 
on Echium vulgare; males suck on Trifolium arvense. | 

Of the cases mentioned by Knuth, excepting Andrena nasuity 
only two are exclusive. Osmia aduncaand Bombus gerstaeckeri att 
not good cases. In the other cases the females collect the pollen : 
exclusively from the plants set opposite them, and the cases aft 
not essentially modified by occasional visits for nectar to othet 
flowers. I hold that Macropis labiata is as good a case as Andrent — 
florea. 

Of the cases mentioned by Loew, three have been passed 
upon. I know of nothing against any of them except Andreni 
cmneraria. The female collects pollen of Salix, but also of Za 
axacum officinale, so 1 should exclude it. 

In the observation of the insect visits of flowers correc 
determinations are very important, for otherwise the records are 
wrong. One has to be sure that the bee is actually collecting — 
pollen, for often a female bee will suck nectar from a flowet 
when her scopz are full of pollen from another species. On i 7 
other hand, there is danger of assuming that a bee is oligotropi ) 
from too few observations. 

A neighborhood where the flora and insect fauna are in thei 
normal condition is more favorable for correct observatiom | 
Yes eh ee 7 for, when the flowers upon which a eo | 
forebd (a Sait dowets hich oeioiy ict 
most cases the f a _— originally it did not visit. “4 

ormer is more likely to happen. 


Pe es ee gee 


1899] FLOWERS AND INSECTS 35 


A bee may be regarded as oligotropic: (1) When the female 
collects the pollen of the plants in question and is not known to 
collect pollen of any other plants. (2) When the bee does not 
occur except during the blooming season of the flowers. If the 
female is shown to occur after the flowers have quit blooming, 
the case is very doubtful. (3) When the bee is frequent upon 
the flowers, and more or less rare upon other flowers, at any rate 
except inthe neighborhood of the food flowers. The case is also 
doubtful if it is shown that the distribution of the bee extends 
greatly beyond the plants upon which it is supposed to depend. 

~ In the more satisfactory cases, if any one should say that he 
had observed the bee collecting pollen from a quite unrelated 
flower, I would not accept the determination, or, if that were 
beyond question, the opinion that the pollen came from the 
flower on which the bee was taken. 

Below I give a list of bees which I regard as oligotropic in 
the above sense. When I have observed the female collecting 
pollen from more than one species of a genus, I give the genus; 
when from more than one genus, I give the family. The details 
will be given elsewhere. 

In Prosopis the females are destitute of pollinigerous appa- 
ratus, their nests being provisioned with a paste of honey and 
pollen. I know of no way to distinguish the flowers which the 
females visit for this purpose from those which they visit in only 
an incidental way, so. I assume that a species of this genus is 
oligotropic only so long as it is found exclusively on flowers of 
One species or group. This may be assumed for either sex so 
long as the condition holds, as in case of P. zllinoensis, of which 
I do not know the females. P. nelumbonis has always seemed to 
me to be the best case of an oligotropic Prosopis. 

I have never believed that our species of. Epeolus were 
cuckoos of Colletes, because there are more common species of 
the former than of the latter genus, and their phenological posi- 
tions do not show the same correlations which exist between 
Andrena and Nomada, Megachile and Coelioxys. Besides, the 
Maximum of Epeolus does not approximate that of any other 


36 BOTANICAL GAZETTE [yuLy 


é 1g 
3) ra] a | 2 = 
eee) Gsle |e 
oe Plants ag aang for ‘s 2s sf 2 | 
3 | $2) 5 | Seige 
& Bee | Bm | 25 | go 
3 o> C's | Se 1s 
z, i & (e) A 
Colletes aestivalis - - | Heuchera hispida - q eee . ‘ 
latitarsis —- Physalis— - - 5 = — 3 
willistonii § - | Physalis lanceolata - | ; 
americanus - Composite- —- 8 |—| 2 3 as 
atus - Ss . “6s 2 4 peal I a ‘ 
compactus. - Y - - > ne ss 1 ae 
eulophi - " - 3 |—- | 3) 
Andrenaarabis - —- Arabis levigat A pee ae 
erigenie - - | Claytonia Virgini nic q ae 2 a 
geranii - Hydr slp a ‘appendic' um} I para ee i 
g. mac = 4+ Geran I pe BS ie 
polemonii Pac atien : focar I a 3 
Spirzeana - - | Spirzea Arun ‘ 1 namie oS 3 te 
viole - - Viola cucullata - “ I 2 Be, 3}. rie 
+ ash coca > RIDE: « - : . 4 I he ; 9. 
illino - - . - 4 I oe: a 
site ‘a - - 4 mae ae oa 
salicis - * és 2 : 4 - 4 te i : 
asonii_- - | Umbelliferze - - 3 mga eas 
ziziae - . Pic : i 5 — | >) 
rudbeckize - | Rudbeckia hirta - - I | — I a. 
alicie - Composite — - - See 
asteris = - - = ‘ . Fe eer in 
helianthi Seal ‘ 2 i. 2 | oe 
nubecula - te ‘ 8 ‘ 4 1 | 74 
pulchella “s : i alae tions 2) 
solidaginis —- “ ie. 6 |— | 1) ae 
+ elo andrenoide Salix - - - - 3 I oe, 9 
cropis steironematis - Steironema-_—«- 3 | —- | hee 
Hali ictus nelumbonis .- Ny phzeaceze ~ 3 — ae x 
.Megachile exilis - Campanula Americana Oe cae Gow: 
_ pugnata = - Compositze - - 4|/— I CA 
Panurginus famcai ae tiloba : ee ea * ie 
‘ ‘albitarsis - Com 7m Pos “ 2 — 4 en 
8 Gus - - - : 4 on 
positarum * “ k : 5 — 3 . 
labrosiformis  - «“ eo | ed eee 
tudbeckiz - “ 4};—|—-|= 
rugosus - s “ ‘ ‘i 4 pls 20 
Paterna. - “ 2 |—-.| 458 
Ssa pruinosa  - | Cucurbit — | 3 | ee 
Emphor bombiformis - Hibiscus Pepo (elt) - ‘ — | = ae 
Anthophora walshii —_- ssia rista y [—4 eee 
Perdita Octomaculata - Co omposita: : s 3 of) ae 
Halictoides marginatus Helianthus - - 3 1(— tre 
otal SS Se ce gee, ee 


1899 | FLOWERS AND INSECTS 37 
m os] 

ee ee 

bs] Ug | ee] = fo 

5 ooh me oo oe _ 

ES 2. Ce eee 
Plants visited by females for = 5 oo | on | Se 
Bes pollen on). Se lee les 

Me os n o ° 
y Re pects eg 
a 2 $ oH mek 
g ie on | S22 | ge 

4 | & te o a 
Mellisodes desponsa-~ - - 2 I 2 
illinoensis  - eo pinnate - 1 j|— | — I I 
agilis - - Compo sitae - 6 |— | 12} 10 | ‘22 
americana - - - - 9 2 I 3 
coloradensis” - ts : : 7 | — 6 I 7 
pennsylvanica af ee 6 bs 9 44 12 
simillima = - sh : : Gat 12 41238 


genus of bees on which it might be supposed to be inquiline. 
Then they are more abundant than would be expected of inqui- 
line bees. Mr. Ashmead’s observations confirmed my views, 
and I have never doubted their correctness since I first read an 
account of them. In Psyche, for March 1894, p. 41, he states 
that he found Z. donatus making nests in the ground and provis- 
ioning them with a honey-paste. Epeolus thus comes under the 
Same category as Prosopis and is treated the same way in the 
table. 

The cuckoo bees of the genus Nomada hold no particular 
relations to flowers except through their hosts. However, they 
Show considerable differences. NV. vincta, which is common on 
Helianthus and was taken once on Coreopsis, is, I think, an inqui- 
line of Andrena helianthi, both bees occurring at the same time, 
in the same neighborhood, and on the same flowers. 


Bee 


At least females visit exclusively 


Prosopis nelumbonis 
ha = - = 
ilinoensis - 

Epeolus helianthi 


ec 
pusillus 
Nomada vincta 


Be EOE, 


Nympheza 

Thaspium a aureum m trifoliatum - 
Umbel 

Heltaathas’ grosse- -serratus - 


QO 
° 
3 
a) 
fo} 
zg 
& 


Composit * - . 


Other fis, visited 
No. spp. by male 

4 

I ets 

5 — 

I I 

4 ae 
13 3 

2 sce 

4 — 

3 oes 


38 BOTANICAL GAZETTE [JULY 


III. Competition of flowers for the visits of bees.— It is a ques 
tion to what extent groups of plants adapted to certain kinds of — 
bees should be regarded as in competition and to what extent — 
they should be regarded as mutually helpful. We will suppose — 
a case in which a plant whose flowers may be visited by beesis 
introduced into a region where all visitors must be acquired, 
If the region contains no flowers, there will be no bees to acquire, — 
On the other hand, it seems to me that the more nearly the 
flora retains its original characteristics the more bees there will — 
be and the more chances there will be of the new flower acquit 
ing bees as visitors. My view is that a patch of plants adapted 
to bees of certain kinds will be more abundantly visited, if it is : 
surrounded by plants depending on bees of the same kinds, than 
if the neighboring grounds are unoccupied. There will be more ~ 
of these bees in the neighborhood. In the table there are fifty- 
two species which get pollen from particular plants. As far as 
the data are correct, we take it for granted that the presence 
and abundance of these bees in a given locality depend on the | 
presence and abundance of the flowers from which they get theif _ 
pollen. One object in making the table is to show that the 
plants growing in the neighborhood of plants visited by oligo- 
tropic bees gain a certain number of bee visits. The table — 
shows that these plants gain 204 visits in this way. Itis expected, 
however, that some of the visits enumerated in the second an¢ 
third columns will have to be transferred to the first. Exclud | 
ing these columns, the neighboring unrelated plants gain 110 
visits from the es. 


for insects t 
pollen, altho 


1899] FLOWERS AND INSECTS 39 


and dry that it is apt to be blown away as soon as it is liberated 
from the anthers. The first step in the development of entomo- 
philous flowers was the secretion of nectar somewhere about the 
stamens and pistils, correlated with the modification of the flower 
so as to afford convenient resting places for insects, and the 
pollen becoming more adhesive, so that it would remain on the 
anthers after dehiscense and become attached finally to the 
bodies of the guests. The object of insect visits being the 
nectar, modifications favoring cross-pollination resulted in the 
various forms of diclinism and dichogamy. The perfection of 
nectar-bearing flowers naturally reached a high grade in the less 
specialized groups of plants, as, for example, the orchids, and 
was most frequently associated with the less specialized antho- 
philous insects. 

Along with the development of convenient landing places 
and sticky pollen, there has no doubt been an increasing number 
of insects which resorted to flowers for pollen. Finally, the 
most highly specialized of anthophilous insects, the Hymenop- 
tera, gave rise to a still more highly specialized group of insects 
which adopted the habit of provisioning their nests with nectar 
and pollen. Along with the acquisition of this habit the bees 
developed a coat of feathery hairs to which the pollen might 
cling, these hairs on certain parts of their bodies, as the hind 
legs and ventral surface of the abdomen, being greatly modified 
to form special pollen-carrying apparatus called scope. Thus 
the pollen became absolutely essential in the economy of the 
most highly specialized anthophilous insects. To the flowers, 
on the other hand, the bees became the most important visitors, 
because they had to resort to flowers more frequently than other 
Msects, and because they were provided with a coat specially 
fitted to retain the pollen, and at the same time exerted them- 
selves to get the coat as full of pollen as possible. 

That the development of entomophilous flowers with sticky 
pollen preceded the development of the bees is indicated by the 
fact that the less specialized bees only collect adhesive pollen. 
The most highly specialized bees, however, have acquired the 


flowers. They fairly monopolized the staminate flowers, W 


40 BOTANICAL GAZETTE [JULY 


habit of sticking the pollen with honey, and so can use that of 
anemophilous plants. 

Those flowers, however, which, through their nectar and cor 
related modifications, were the best fitted to use the services of 
ordinary insects for cross-pollination, were the least fitted to | 
utilize the insects which were the highest product of anthophilow’ 
development. Strange as it may seem, the characters which hit- 
dered them from availing themselves of these services were the 
very characters which are considered the highest adaptations for 
cross-pollination, viz., diclinism, dichogamy, and large size. Ob | 
the other hand, the forms which have enabled flowers most readily 
to avail themselves of the services of bees are the very charac 
ters which have been interpreted as adaptations for self-polline 
tion and geitonogamy, viz., small size, homogamy, and the — 
aggregation of dichogamous and other flowers in close clustets. | 

If an insect in search of nectar visits a dicecious or othe 
diclinous plant, it is not hard to understand how it is likely t® 


the pistillate flowers were visited by an entirely different set 
insects. In the table there are six species of bees which 
their pollen exclusively from dicecious species, Salix and Spi 
Aruncus. Of the plants furnishing pollen to oligotropic — 
these are the least able to utilize these bees on account of thet 
dicecism. 

Dichogamous flowers are at somewhat of a disadvantage 
utilizing pollen-collecting bees from the fact that the bees 4 
more apt to pay attention to the flowers which are discharging. 
pollen and neglect those in the other stage. In Jmpatiens ful : 
and J. pallida | have observed that Megachile brevis collects * 
pollen from flowers in the first stage and avoids those * 


1899] FLOWERS AND INSECTS 41 


receptive stigmas, because she instantly perceives that the anthers 
are gone. Apis mellificaand Bombus virginicus do the same when 
collecting the pollen of Z. falva. In Campanula Americana, which 
is also proterandrous, the oligotropic Megachile exilis cleans the 
pollen from the style-brushes before the stigma opens, and avoids 
the old flowers. In Lobelia syphilitica | have seen little bees col- 
lecting the pollen which was pushed out of the anther tube before 
the stigma appeared. Inthe proterandrous Monarda Bradburiana 
I have seen small bees collecting pollen directly from the 
anthers, avoiding the old flowers. The strongly dichogamous 


flowers mentioned in the table are not so well adapted to utilize 


their'special visitors as are the homogamous ones, such as Viola, 


. Psoralea, Hibiscus, Cassia, because in the latter the bees cannot 


collect the pollen without touching the stigmas. 

Some dichogamous flowers may make effective use of the 
pollen-collecting bees, as in the case of Nymphea reniformis, 
which, in my opinion, is proterogynous and without nectar. By 
a sudden bending of the filaments, bees alighting on the anthers 
are let down into the stigmatic basin before they discover that the 
pollen is not being discharged. Of course, in other dichogamous 
flowers the bees may visit the flowers in the pistillate stage before 
they discover that the pollen is gone, or for nectar, but my 
observations have convinced me that this is not the rule, for if 
they do not know exactly what they are doing and how to do it, 
they act just like it. On their pollen-collecting expeditions they 
do not make many mistakes or waste much time. 

Even some homogamous flowers are so-large that the smaller 
bees may collect their pollen without touching the stigmas. 
This may not matter so much if the flowers are visited by large 
bees, which are more effective. But the smaller flower may, in 
many cases, utilize the large bees as well and the smaller ones 
better, So I think the influence of the pollen-collecting bees is 
in favor of the smaller homogamous flowers. 

Under the influence of the nectar-sucking, less specialized, 
anthophilous insects the highest development is found in 
diclinous, dichogamous, and hercogamous flowers with highly 


42 BOTANICAL GAZETT [JULY 


specialized nectaries and precise localization of pollen contact. 
'In the less specialized plants, this kind of adaptation early 
reached the highest degree of perfection in the case of the 
orchids. But, as far as I know, no orchid holds an important 
relation in the economy of any bee. 

Under the influence of the female bees, the most highly spe- 
cialized of anthophilous insects, the highest development is 
found in homogamous flowers without nectar, such as Desmodium 
and Cassia. 

Since bees have entered the field, many flowers seem to have 
been at a disadvantage in gaining their services, because the sta- 
mens were so few that they could not offer pollen in paying 
quantities. And in many cases the stamens were covered by 
gale and carina, so that, to collect the pollen, the bee would 
have to spend much time going to the bottom of every flower. 
This difficulty was obviated by lengthening the stamens, reduc- 
ing the size of the flowers, and crowding the flowers so that the 
bees could run over or around the inflorescences and sweep Up 
immense quantities of pollen. Inflorescences of this kind are 
found in Cornus, Hydrangea and Viburnum. 

Here we find an explanation of the fact that certain Legt- 


minose and Labiate have abandoned their galez and caring, 


exposing their stamens, and contracting their infloresences into 


head-like or flat-topped clusters, as in Amorpha, Petalostemom, 


Lophanthus, Mentha, Blephilia, and Pycnanthemum. Contraty 


to Miiller, I think Delpino is right in regarding Mentha as one — 


of the most highly specialized of the Labiate, and I incline to 


the same opinion regarding the above genera of Leguminos® — 
These cases are obscured by the fact that the arrangements per “ 
mit the visits of a lot of less specialized insects. Nevertheles* 


I think the bees have determined the result. 


In the case of Lobelia I have mentioned that small bees col 
lect the pollen pushed out of the tube before the stigma appears. 
In the Composite we find plants perhaps best adapted to attract 
and utilize the pollen-collecting bees, and the table shows that 
they have among their visitors more oligotropic. bees than any 


ei ee Ss le AS a 


1899] FLOWERS AND INSECTS 43 


other group of native plants, and that, too, in spite of their 
dichogamy. If the flowers were greatly scattered, they no doubt 
would not attract so many bees, and the bees could carry off the 
pollen and not render any service by visiting the flowers after 
the stigmas appeared. But, as a result of the reduction of the 
flowers in size and the crowding of them in heads, we find a 
circle of flowers, each one of which ejects the, contents of five 
anthers in a convenient mass. Just without is a circle of flowers | 
with protruding stigmas. Bees sweep over the disk, filling their 
pollen-scope with the greatest facility, at the same time effec- 
tually pollinating the neighboring stigmas. 

As the homogamous flowers have largely been given over as 
adaptations to autogamy, so the crowded inflorescences have 
been given over as adaptations to geitonogamy. As a category 
Ido not accept Kerner’s geitonogamy. Kerner regards most of 
the crowded inflorescences as adaptations for geitonogamy, and 
founds a special category for their reception. This is accepted 
by Knuth and is incorporated in his recent Handbuch.” I do 
not believe in any adaptations for geitonogamy. I do not deny 
that it occurs, and under pseudo-ecological conditions may be 
advantageous, but it is only the name of an accident and does 
ahi account for any floral adaptations. Kerner does not makea 
distinction between a structure, or habit, which has a certain 
effect, and one which may be conceived to be developed for a 
certain purpose, or selected on a certain condition. He even 
Speaks of a “contrivance for securing hybridization.” Under 
“contrivances whereby the pollen is protected against wet” he 
Says: “In Podophyllum peltatum the pollen is sheltered by the 
bell-shaped flower, but in addition to this the peltate foliage- 
tes a also Spread out over the flowers and act as umbrellas.” 
ee ~ Category of protection by isolation in “_— he 
Avene a number of ordinary water plants and says: ‘Flies 
se pete which come through the air for honey and pollen 
valida cad Wetors, promoting, as they do, a crossing of the 

> Snails, centipedes, etc., are, on the other hand, kept 

7 Handbuch der Bliitenbiologie 1:51. 


44 BOTANICAL GAZETTE [JULY 


back by the water.” He gives no evidence that this protec: 
tion has anything to do with the fact that the plants have 
acquired an aquatic location. He uses trivial and accidental 
effects as a basis for interpretation of all kinds of ecological 
phenomena. 

While it is true that adaptations for cross-pollination are 
more apparent in the less specialized plants depending on the 
less specialized anthophilous insects, it does not follow that the 
adaptations of the highest plants in relation to the highest 
insects, though more obscure, are to be interpreted as arrange 
ments for autogamy and geitonogamy. 


V. On the supposed pollen-carrying apparatus of flies and birds 
—In regard to the plumose ariste of such genera of Syrphide 
as Volucella and Sericomyia, Loew® observes that the structure 
appears of no use to the flies, but is of importance in the trans 
fer of pollen. And he regards them, as well as the hairy coat 
on the lower part of the face, as an adaptation for carrying pol 
len. In the same connection he mentions the hairy eyes of ce 
tain species, though he does not go so far as to consider this as 
an adaptation for the same purpose. 

In the Entomological News 4:323. 1895, under the title 
Insects as pollenizers, Mr. J. B. Smith mentions that some Dipte® — 
have compound hairs, similar to those found in the Apid@. 
The author does not say exactly what he does mean, but I have 
always regarded the note as implying the view that these hairs 
were so modified for carrying pollen. ‘| 

In the American Naturalist 28: 680-681. 1874, Mr. J. L. Ham | 
cock speaks of certain ‘‘repositories”” on the head of the ruby : 
throated humming bird, and throughout his paper seems 1 | 
imply that the feathers, etc., are specially modified for carrying | 
pollen. As Mr. Darwin says, proof of the existence of suc F 
adaptations would be fatal to the theory of natural selection. "| 
have always regarded these statements as mere teleological os 
osities, but in his Handbuch Knuth has adopted Loew’s views | 

*Jahr, Bot. Gartens Berlin 6:114. 1886. I 


—_— = 


| 
4 
4 
k 
3 
; 


1899] FLOWERS AND INSECTS 45 


which has the effect of giving them some standing among the 
fundamental principles of flower-and-insect ecology. 

The existence of branched hairs in the bees may properly be 
interpreted as an adaptation for carrying pollen, because the 
bees use them for that purpose, and the importance of the hairs 
is evident, in view of the economy of the insects. They cannot 
in any way be interpreted as existing for the benefit of the 
flowers. It could be of no advantage to flies and birds to carry 
pollen, since they make no use of it. However, it might be 
claimed that these guests derived an indirect benefit from the 
pollination of their favorite plants. But their relations to flowers 
are not close enough to make their existence depend upon the 
pollination and preservation of any particular species. 

An examination of the inquiline bees will lead to the conclu- 
sion that the several genera are not related to one another but 
have arisen independently from different groups of host bees. 
It will also lead to the conclusion that they have all lost their 
hairy coats, or tend to do so, as in Psithyrus. To my mind the 
fact that these bees began to lose their coats as they abandoned 
their pollen-collecting habits, involves a clear refutation of the 
claims that any structures on flies and birds were developed for 
the purpose of carrying pollen. 


CARLINVILLE, TEL. 


THE ORIGIN OF THE LEAFY SPOROPHYTE. 
JoHN M. CouLtTeER. 


ATTENTION has been called afresh to this exceedingly inter- 
esting and obscure problem by the discussion of alternation 
of generations by Professor Bower in his recent presidential 
address," and in papers of Dr. Klebs,? and Dr. Lang.3 The 
remarks of Professor Bower are largely in defense of his theory 
of the antithetic origin of the sporophyte, which had been 
attacked by Dr. Scott in his presidential address of two years 
before in restating Pringsheim’s theory of homologous alterna- 
tion. In defending his position, Professor Bower discusses argu- 
ments derived from the behavior of algz and certain fungi, from 
bryophytes, and from apogamy and apospory. He claims that 
those alge and phycomycetes which show subdivision of the 
zygote into spores appear to offer the “ key to the enigma” of 
the origin of the sporophyte, but he makes no further claim for 
these “fruit bodies” than that they suggest the way in which 
the sporophyte may have arisen, his view not at all involving 
the idea that these “fruit bodies” and the sporophyte are homo 
genetic. He calls attention to the fact that knowledge of cyto 
logical phenomena among alge and fungi is far too meageh 
especially in connection with the divisions of the zygote referred 
to. If reduction is found to occur in connection with the zygote 
divisions, in such forms as CEdogonium and Coleochete, there 
would be a reasonable foundation for the belief that the “ fruit i 
bodies” are the correlatives of a sporophyte, the beginning of 
neutral generation. ; 

In reference to the bryophytes, Professor Bower sees if 
them a good illustration of the origin of the sporophyte by = 

* Nature, Nov. 17, Noy. 24, Dec. 1. 1898. . 

? Annals of Botany r2: 570-583. 1898. 

3 Annals of Botany r2: 583-592. 1898. 

46 [July 


1899] ORIGIN OF THE LEAFY SPOROPHYTE 47 


progressive antithetic alternation.” He calls attention to the 
remarkable constancy of alternation in this group, apogamy and 
apospory being singularly absent. This undeviating alternation 
he suggests may be accounted for by the dependence of the 
sporophyte, which is in an ‘‘equable physiological condition.” 
As a contrast to this, the independence of the pteridophyte 
sporophyte, and its exposure to varied conditions, may have 
caused more freely unusual developments. The primitive pteri- 
dophyte, however, was probably in a dependent condition, as the 
embryos of modern pteridophytes are. 

Apogamy and apospory Professor Bower would regard as 
“abnormalities,” calling attention to the fact that these phenom- 
ena have their ‘“‘headquarters” in the leptosporangiate ferns, a 
peculiarly specialized phylum with many other abnormalities. 
Even when apogamy occurs the archegonia are first produced, 
indicating the “first intention” of the plant; and in both apog- 
amy and apospory the growths may be very anomalous. 

In this connection, Professor Bower makes a very interesting 
Suggestion, based upon the experiments of Dr. Lang and others. 
He observes that apogamy is induced by prevention of contact 
with fluid water (‘rendering fertilization impossible”), exposure 
to direct sunlight, and possibly to certain temperature conditions. 
All this leads to a ‘‘plethoric”’ state, which he thinks may be a 
necessary condition preceding apogamy, as opposed to deficient 
nutrition, which precedes apospory, the latter being ‘‘a physio- 
logical tefuge for the destitute plant.’’ He suggests that nuclear 
changes may accompany these conditions, plethora doubling the 
chromosomes, and hence inducing the development of a sporo- 
phyte ; and deficient nutrition reducing the chromosomes, thus 
making a gametophyte possible. Of course it remains to be 
Proved that nuclear instability, coming to be well recognized, is 
Connected with disturbed nutrition, and also whether a smaller 
or larger number of chromosomes necessarily determine a gamet- 
ophyte or a sporophyte. 

- cn whole, therefore, Professor Bower still maintains that 
phyte is the result of the gradual elaboration of the 


48 BOTANICAL GAZETTE [JULY 


zygote, ‘‘a fresh phase having thus been gradually interpolated,” 
in other words, that its origin is antithetic. It would seem that 
in his opinion the sporophyte has probably appeared in just one 
way. This does not mean that all sporophyte plants are homo- 
genetic, but that all have had an origin similar to that of the sporo- 
gonium of bryophytes. Professor Bower acknowledges that 
the present tendency is toward a comprehensive polyphyletic — 
view as regards alternation, stating that “when difficulties arise 
refuge is taken in the plausible suggestion of distinct lines of 
descent.” 

Dr. Lang’s paper is rather a presentation of current views 
than an expression of opinion in reference to any of them. He 
recognizes the fact that the regularity of the zygote product 
in such forms as CEdogonium and Coleochzte represents a life 
history decidedly different from the homologous alternation of 
sexual and asexual plants in most thallophytes. From one point 
of view this zygote product is merely a reduced asexual indi- 
vidual; from another point of view it is not a reduced asexual 
individual, but a special adaptation to multiply the product of 
fertilization. The former is the theory of homologous origin: 
the latter the theory of antithetic origin. Certainly the facts 
of morphology do not decide which theory is correct. Dr 
Lang calls attention to the fact that in considering alterna 
tion the possible polyphyletic origin of the archegoniates must 
be kept in mind, as the pteridophytes may represent an entirely 
distinct line from the bryophytes, as suggested by Goebel. 
In spite of Professor Bower’s disposition of apogamy as 4 
argument, Dr. Lang thinks that experiments with this phenome — 
non indicate so clearly that the gametophyte may assume char- 
acters of the sporophyte under suitable conditions, almost # 
complete series of transitions between gametophyte and spor” 
phyte having been observed, that such a general property of the 
fern gametophyte cannot be disregarded in the discussion, evel 
though the phenomenon may be called teratological. He thinks 
that apogamy Suggests the homology of the gametophyte and 
sporophyte, and may suggest how pteridophytes could have 


* 


1899] ORIGIN OF THE LEAFY SPOROPHYTE 49 


been derived from alge forms, and how alternation in ferns 
might have arisen if it did not come antithetically. 

The paper of Dr. Klebs deals with the subject of alternation 
of generations in thallophytes, and therefore concerns this present 
discussion but indirectly. His experiments among the lower 
forms, as is well known, have proved that there is no such rigidity 
in life histories as was once supposed. As a consequence, he 
does not consider that there is any such thing even as a regular 
homologous alternation of sexual and asexual phases. He thinks 
that experiments may prove that the so-called ‘fruit bodies” 
of such forms as Edogonium and Coleochete may turn out to be 
the result of certain conditions, rather than an inevitable part of 
the life history. He seems to consider that the origin of pteri- 
dophytes probably has nothing to do with that of bryophytes, 
and that there is at present no clue whatsoever as to the origin 
of the former. Such a peculiar structure in common as the 
archegonium he suggests may be a purely parallel development, 
without necessarily indicating any phylogenetic connection. 

It will be seen from the above papers that, while the origin 
of the sporogonium of bryophytes seems to be suggested, the 
origin of the leafy sporophyte is too obscure to justify any 
definite claim, According to Bower it is most probable that it is 
developed from such a sporogonium structure as is displayed by 
the bryophytes today; according to Lang and Klebs there is a 
Possibility that it may have had an entirely independent origin, 
and may never have been in the sporogonium condition. 

It 1S recognized that there are peculiar difficulties in the dis- 
ain of such a subject. Although the morphology of the 
“xisting representatives of the various groups is fairly well known, 
plas two enormous gaps in our knowledge which. make a 
biases Sr es impossible. One of these gaps is the ancient 
ape Riad the bryophyte and pteridophyte lines. For instance, 

in that the pteridophytes were well represented in the 


‘pal i ae ; : 
Paleozoic, probably even in its earliest periods. This represents 


such 


ee a tremendous stretch of time that almost any change, how- 
r 


e i . . . i 
xtensive, may have been possible'in any given form. It is 


Mo. Bot. Garden, 
1906. 


5° BOTANICAL GAZETTE [JULY 


not through lack of time, therefore, that one would suggest that 
it is unlikely for a leafy sporophyte to have been developed from 
a sporogonium. From the fact that our earliest evidences of the 
pteridophytes show them to have been about as highly differ- 
entiated as they are now, it is evident that the evolution of the 
line reaches very far back. It is probably hopeless to expect 
that this gap in our knowledge will be filled. 

The other gap is in reference to cytological details. The 
whole subject of alternation of generations seems to be s0 
bound up with nuclear changes that a knowledge of these it 
the thallophytes becomes a very great desideratum. This gap 
in our knowledge is likely to be filled up rapidly. It may be 
that we have been too rigid in our use of the number relations 
of chromosomes as distinguishing gametophytes from sporo- 
phytes. Be this as it may, there is enough in the testimony 
associating the doubling and the reduction of chromosomes 
with the sporophyte and gametophyte stages to justify such use. 
It would seem that an investigation into the nuclear changes 
which occur in the “fruit bodies” of such forms as CEdogonium 
and Coleochete would go far toward settling the antithetic origin 
of such a structure at least as the sporogonium of bryophytes 

It is not my purpose in this paper to traverse ground which 
has been gone over so recently and so ably, but merely to dis- 
cuss certain facts and possibilities in connection with the leafy 
sporophyte that may be suggestive. In discussing the origin of 
such a structure as the leafy sporophyte where there is 1° 


possible direct evidence, and where every view must be hypothet- 
ical 


coordinate the facts and to suggest lines of research. 


No structure among plants seems to have left so little trace 


of its origin as the leafy sporophyte of pteridophytes and 
Spermatophytes. The evolution of the leafless sporophyte : 
bryophytes seems traceable from an oospore which directly 
organizes a group of sporogenous cells. Sterilization of the 


peripheral cells would result in a simple spore-case like that of 


» It seems necessary to consider all possible alternatives: 
The chief service which these various alternatives render is 0 


a See ee ee ee a a Se 


1899 ] ORIGIN OF THE LEAFY SPOROPHYTE 51 
y 


Riccia, while further encroachment upon the sporogenous tissue, 
with more or less differentiation of the sterile tissue, would 
account for the series of sporogonia displayed by bryophytes. 
Whether the origin of this structure is to be regarded as homol- 
ogous or antithetic is not pertinent to the present discussion, 
but it seems reasonable to see in it an entirely new structure 
developed by the oospore, and in no way homogenetic or even 
homologous with the gametophyte. It has been noted that the 
argument drawn from apogamy in favor of homologous origin 
finds little or no application among bryophytes, for the origin of 
the sporogonium seems to be as fixed as the origin of any plant 
structure can be. 

It has been common to regard the distinct sporophyte as 
having been established once for all by the bryophytes, and the 
sporophytes of the higher groups to have been derived from 
those of the bryophytes. In searching for the origin of the 
leafy sporophyte, therefore, attention has been focused upon 
the sporogonia of bryophytes, and the Anthoceros forms have 
been selected as most nearly representing the ancestral condition. 

The doctrine that any plant structure, however important, 
can have but one phylogeny, is hardly tenable at present. That 
heterospory has appeared independently in several lines has 
become evident ; and that it has resulted more than once in seed 
formation is hardly less evident. The conditions which 
determined these modifications must have been common enough 
to have established similar results more than once. Why the 
Sporophyte may not fall in the same category is not clear. 
Professor Bower’s statement that the polyphyletic origin of a 
ihe a is an easy escape from difficulties suggests caution, 
Ase - not close the door to the fact that nature may have 

€ same easy way out of difficulties. 
id the sporophytes of bryophytes and pterido- 
ae co eg abide to have nothing in common except that - 
be he from ie oospore and represent an asexua 
oe mae hese facts are important, but so are the numerous 
in which they differ sharply. There are also asexual 


52 BOTANICAL GAZETTE [JuLy 


generations derived from oospores among thallophytes, but 
regular alternation of sexual and asexual generations is not 
definitely established. When alternation becomes definite the 
sporophyte is a recognizable structure, but that this structure 
must have been established just once or in just one way is far 
from necessary. 

It may be well to contrast the leafless and leafy sporophytes. 
In the former case the structure is never independent of the 
gametophyte, develops no lateral members, has nothing com 
parable to sporangia, and its whole tendency is to render complex 
the spore-producing region. In the latter case the sporophyte 
is dependent upon the gametophyte only in its embryonic 
stage, develops prominent lateral members, has distinct simple 
sporangia, and its whole tendency is to render complex the 
sterile or nutritive tissues. As one traces the evolution of the 
bryophyte sporogonia they give evidence of increasing com 
plexity and hence rigidity, and little promise of originating such 
a diverse tendency as that shown by the sporophyte of pterido- 
phytes. The mosses are conceded to be a highly specialized, 
and hence non-productive line, the legitimate outcome of the 
whole bryophyte tendency. Why the liverwort lines may ae 
also be regarded as highly specialized and hence non-productive 
does not seem clear. It is true that the Anthoceros forms show? 
sporophyte tendency unlike the others, and that if such a spol 
phyte should become independent and put out leaves, and if the 
continuously developing spore region should be restricted and 


broken up into simple sporangia which should associate them 


selves with the leaves, we might have something like the existing 


leafy sporophytes. But there is no evidence that these thing? 


ever happened. On the contrary, the sporophyte of Anthocero® 


would seem to be as hopelessly specialized as that of other lines. 


It is true that allthe things referred to above may have happene® 
and Anthoceros may be the nearest living suggestion of 


archetypal pteridophyte, but the case is not so clear that cae 


eyes should be shut to other possibilities. : 
If the bryophyte sporogonium is responsible for the leafy 


| 


ee a ee ee eee Ta 


; 
) 
: 
: 
{ 
; 
: 
, 
| 


1899] ORIGIN OF THE LEAFY SPOROPHYTE 53 


sporophyte, then it is evident,as Bower has shown, that the 
leaves of the latter are the result of progressive sterilization, 
and are secondary structures of the sporophyte. But if some 
other origin of the leafy sporophyte is possible, the leaves may 
not have arisen as secondary structures. 

It may be well to trace briefly the origin of gametophyte 
leaves, as exhibited by the mosses, since the sequence of events 
seems fairly clear,and may prove suggestive. Among the 
Riccia forms the thallose body produces sex organs and does 
chlorophyll work with no special differentiation of regions. 
From this condition there is evident a tendency to segregate the 
sex organs into definite regions, so that eventually the region of 
the body devoted to sex organs becomes quite distinct. The 
differentiation of a sex organ region is still further emphasized 
by its separation from the rest of the body by being carried up 
upon a vertical branch, an extreme case being displayed by 
Marchantia. Asa result, the chief chlorophyll work and the 
production of sex organs are distinctly set apart by the organi- 
zation of a gametophore arising from the thallus. 

The gametophore, primarily a sex organ branch, proves to 
be more favorable for the display of chlorophyll tissue than the 
thallus, and the simple leaves of mosses appear, supplementing 
the chlorophyll work of the thallus. In sphagnums the thallose 
body continues associated with the leafy gametophore. In the 
true mosses, however, the chlorophyll work of the gametophyte 
'S more or less given over to the gametophore leaves, and the 
thallus region is reduced to the so-called ‘‘protonema.” Ina 
very true sense, therefore, the gametophyte is always a thallus, 
special vertical or radial branches being developed in liverworts 
as gametophores, and in mosses as leafy gametophores. The 
loose habit of homologizing the leafy ‘‘moss plant” with a liver- 
wort thallus on the one hand, anda fern prothallium on the other, 
's not merely bad morphology, but is apt to be very misleading. 

The suggestion to be obtained from this history is that leaves 
may develop in response to more favorable conditions for their 
work, and such development may result in the great reduction 


54 BOTANICAL GAZETTE [yuty 


of chlorophyll work done by the less favored region, and its con 
sequent simplification. It is evident that with the exchange of — 
an aquatic for a terrestrial habit the thallose body would not be 
a favorable type for chlorophyll work, and that the development 
of chlorophyll tissue upon erect structures of various kinds might 
follow. Among bryophytes the erect structure laid hold of is 
the gametophore, and not the sporogonium. I grant that this 
same reasoning would make the sporogonium of the Anthoceros 
forms a specially well adapted erect structure for the develop- 
ment of leaf tissue and hence leaves. The objection, however, 
is that the sporogonia of bryophytes are most persistently spore- 
bearing structures and nothing else, every tendency towards ) 
more complex organization having spore production and spore : 
dispersal in view; and that such specialized structures are not ~ 
apt to be productive of new lines of development. 

In considering, therefore, whether it is possible to disregard 
the bryophytes in our search for the origin of the leafy spore 
phyte, we are largely influenced by the fact that the bryophy te 
sporophyte, throughout its whole history, is dominated by 4 
tendency which does not appear in the pteridophyte sporophyt®. — 
Before the establishment of alternate generations the plant body — 
may be said to have had three functions, namely, chlorophyll 
work, and the production of gametes and spores. The appear 
ance of the bryophyte Sporogonium was dominated by the separa 
tion of spore formation from the other functions, chlorophyll 
work being retained by the gametophyte, along with gamete 
production. Attention has been focused so long upon me 
gametes and spores as the two dominant factors in differentia 
tion that it is hard to conceive of the possibility of the domina 
tion of another factor. It is entirely conceivable, however, that — 
another form of differentiation may have occurred, dominated _ 
by the needs of the chlorophyll work, and not by spore produc: 
tion. Certainly a great need for change, when aquatic conditio™ 
were exchanged for terrestrial, was in connection with the displ4y _ 
of chlorophyll tissue. It would seem as if the bryophytes had | 
laid emphasis upon spore production, and therefore never became | 


ee ee ee ee a ee eT ee, 


ia a a a a a lI 


BE Ee ee ae Ee Se EL ee a CEST ARS a ae eee ee ae ep Late ee en ee ee ee 


| 
4 
‘ 
7 
3 
' 


1899] ORIGIN OF THE Le:AFY SPOROPHYTE 55 


organized for the fullest use of terrestrial conditions ; while the 
pteridophytes laid emphasis upon chlorophyll work, and became 
highly organized for terrestrial life. It would seem possible, 
therefore, with the three factors to take into account, that two 
distinct asexual lines may have been organized, distinct in the 
factor selected to dominate. 

Such a conception may be simple enough, but it is hardly 
worthy of consideration without more practical statement. If 
more favorable structures can be developed in response to the 
needs of spores or gametes, there seems to be no good reason 
why more favorable structures may not be developed in response 
to the needs of chlorophyll work. If such a response in struc- 
ture is possible, it would naturally express itself first in develop- 
ing the largest display of chlorophyll tissue in the most favor- 
able region of the body, which would gradually become 
differentiated more and more distinctly from the rest of the 
body. It does not seem clear why the appearance of an erect 
leafy axis, bearing neither gametes nor spores, is not quite as 
Supposable as the appearance of a sporophore with neither 
gametes nor leaves, ora gametophore with neither spores nor 
leaves. 

Of course such a leafy axis would be an integral part of the 
thallus body from which it was developed, and in no sense a 
distinct « generation,” any more than the leafy gametophore and 
the protonema of mosses are distinct generations. Upon such a 
leafy axis spores would find a more favorable position than upon 
the ordinary thallus body, and eventually they would be segre- 
gated upon the leafy axis, developing in connection with chloro- 
phyll tissue just as they had in the thallus body. In such condi- 
sons comparatively simple sporangia would be developed, being 
entirely subordinated to the nutritive tissues. A parallel case 
is found in the gametophore of mosses, which also prove favor- 
able for leaf development; or even in the sporogonia of certain 
ophytes, which also prove favorable for chlorophyll tissue, 

this 's rigidly subordinated to the work of spore production. 

With the development of a leafy axis bearing spores, there is 


56 BOTANICAL GAZETTE [JULY 


no reason why it should not become independent of the thallus 
body which produced it, as the leafy gametophore of mosses 
becomes independent of the protonema. The great difference 
in the final result in the two, cases arises from the fact that in 
mosses the protonema is without gametes or spores ; while in the 
case we are supposing the thallus body produces gametes, and 
the leafy axis spores. That a thallus body can directly produce 
just such a leafy axis bearing spores is testified to by the numer 
ous cases of apogamy observed among pteridophytes. In fact, 
the theoretical life history we have been tracing is concretely 
represented by the life history of a fern in which apogamy has 
occurred. 

If the phenomenon of apogamy represents the primitive 
status of the leafy sporoyhyte, it remains to imagine how this 
spore-bearing leafy axis could have become the usual product of 
the oospore. We find no trouble in believing that the usual 00s 
pore product frequently appears apogamously, for this has been 
demonstrated; but to imagine a general primitive apogamous habit 
of origin gradually passing into a predominant oospore habit of 
origin is difficult. In the condition supposed, namely, a thallus 
body producing gametes, and a special leafy axis bearing spores; 
7ygotes and spores would have the same power, the germinatiot 
of each resulting first in the thallus body and afterwards the 
leafy axis. If real alternation can be brought about by such a 
condition, the thallus portion of the zygote product and the 
_ leafy axis portion of the Spore product must be gradually elimi- 
nated. In other words, the tendency would be to eliminate that 
particular region which is.concerned in producing the reproduc 
tive body. Perhaps such a tendency is no more difficult © — 
understand than the fact that a spore produces a gametophyt€ 
rather than a sporophyte, and a zygote produces a sporophyte — 
rather than a gametophyte. A common explanation has been 
that a zygote, for some reason, stops reproducing the plant body 
Which organizes it, and beg; irely new 
structure, which certainly seems to have been the case in the 
formation of the Sporogonia of bryophytes. It would seem 1° 


ee ae ee ee eee 


EEE Le OE NS eT ee ee ee ee 


1899] ORIGIN OF THE LEAFY SPOROPHYTE 57 


more difficult for a zygote to stop producing one distinct portion 
of the plant body, and to continue producing the other. 

Why in both cases it tends to produce the structure less imme- 
diately related to it, rather than the one which has originated it, 
is a question which cannot be answered at present. Cytology 
may offer certain suggestions, but they are vague as yet. The 
fact that the chromosomes are doubled in number by the process 
of fertilization, and are reduced again in the sporogenous tissue 
may have some bearing on the question. It seems clear that in 
all life histories where the sexual act occurs there must bea cor- 
responding reduction division somewhere. In distinct alterna- 
tion of generations, the ‘“ doubling”’ and the “reduction” are 
associated with the two generations. But before distinct alter- 
nation was established « doubling” and “reduction”? must have 
occurred, and there is no present reason to doubt that in such 
case reduction often, if not generally, occurred in connection 
with the development of spores. When, therefore, the zygote 
was restricted to one region of the body, and the spore to a very 
distinct region, the alternation of « doubling” and “reduction” _ 
might well develop into an alternation of generations. 

The very interesting results obtained by Strasburger and 
Farmer in their Study of Fucus, which show that the reduction 
division in that plant occurs in connection with the development 
of the sex organs, may be correlated with the absence of spores. 
Such an observation emphasizes the fact that reduction must 
occur somewhere, and if sporogenous tissue is not developed, it 
etn seem more likely to occur in gametogenous tissue, repre- 
oe a new cell sequence, than in ordinary nutritive tissue. 
ao an origin of the leafy sporophyte, it would follow 
thus dak ee are not secondary, but primary structures, and 
ledece Las ylls have arisen from the differentiation of foliage 

ing sporangia, a state of things certainly suggested 


ioe ust Primitive pteridophytes known. It would further 
Cw that the Evolution of the strobilus has followed the 
development 


of foliage leaves, a view in accordance with the 


old Be 
re morphology, Such a view would make intelligible the 


58 BOTANICAL GAZETTE [yuLy 


great “gap” recognized as existing between bryophytes and 
pteridophytes, as the two groups would not be phylogenetically 
connected, and would have developed along very divergent lines 
from the first. It would mean that at least two independent 
sporophyte lines have appeared, the bryophyte line probably 
with an antithetic origin, and the pteridophyte line possibly with 
an homologous origin. The great prominence of the latter ling 
with its spermatophyte sequence, is correlated with the develop 
ment of a vascular system, and it would seem as though the 
evolution of an elaborate vascular system must have depended 
upon the domination of chlorophyll work. | 

Perhaps one of the strongest arguments against the poly 
phyletic origin of archegoniate plants is the constant characte 
of the archegonium. It would seem to some inconceivable that 
an organ so definite and so characteristic, and so unlike anything 
in thallophytes, could have appeared in two independent lines. 
However, the possibility of two independent appearances of such 
an organ would depend upon its origin, a subject of great obscurity: 
That it has been derived in some way from the oogonium of 
thallophytes seems hardly to be questioned, and that it is one.tl 
the results of the exchange of aquatic for terrestrial habits sees 
hardly less doubtful. That the archegonium represents a gO! 
of oogonia protected by a layer of sterile tissue seems to be# 
reasonable suggestion, and that the differentiation of this steril 
protective layer into neck and venter would follow naturally 
from the exclusive functioning of the innermost oogonium sect 
probable enough. The conditions which induced this protectio® 
of aggregated oogonia, however, could hardly be claimed ® 


leat 


algal evidence that can be presented, as in the case of the 
less sporophyte. It must be remembered, however, althou 
may be regarded as a convenient refuge for all theories 
phylogeny, that we are dealing with a structure whose origin 
very ancient. Why the alge continue to give suggestions a 


Sa Sr tt Sl aie alae il a ee ko aa 


1899] ORIGIN OF THE LEAFY SPOROPHYTE 59 


the origin of the bryophyte sporogonium, and, so far as known, 
give no intimation of the independent origin of the leafy sporo- 
phyte, isa pertinent question. It seems to be also true, how- 
ever, that the bryophytes give no clear suggestions as to the 
origin of the leafy sporophyte, and we are left to imagine the 
method of its origin from either group. 

In thinking of this possible disconnection of the bryophyte 
and pteridophyte lines, it may be well to recall the similar 
experience of the gymnosperm and angiosperm lines. Certainly 
the gymnosperms and angiosperms seem to have more characters 
in common than do the bryophytes and pteridophytes, and seem 
to be more insistent in their demand for a common phylogeny ; 
yet that the gymnosperms represent at least one independent 
phylum can hardly be longer doubted. 

All such discussion is, of course, very vague and general, 
and may not commend itself to many as profitable. But it 
Serves its purpose in stating the problem, and in presenting the 
possible alternative solutions. We have been in danger of 
restricting the operations of evolution too rigidly, making the 
lines of advance too few, and forgetting the possibilities of 
change during the enormous stretches of time. The polyphyletic 
origin of similar structures and of similar groups makes the 
Problems of phylogeny immensely more complex, but is probably 
much more consistent with the facts. 


THE UNIVERSITY oF CHICAGO. 


Perorern ARTICLE S$. 


THE SEEDLINGS OF JATROPHA MULTIFIDA UL. AND 
PERSEA GRATISSIMA GARTN. 


(WITH SIX FIGURES) 


Two mopes of germinating are characteristic of the dicotyledons, 
with the cotyledons above or under ground. ‘The first is undoubtedly 
the commonest. In this case the cotyledons, as a rule, are freed from 
the seed-coat and develop as two, seldom only one or several, leaf-like 
organs. In the other case, however, the cotyledons either remail 
enclosed in the seed or become free, but stay under ground. hes 
are the principal forms of germination which Klebs? has ascribed 1 
the dicotyledonous orders, and it is interesting to see that both forms 
may occur not only within allied genera, but even among species 0! 
the same genus. Furthermore, the germination itself exhibits a nul 
ber of biological variations in regard to the relative development of 
the primary root, the hypocotyl, and the cotyledons. But it would 
seem very difficult to find any deviation from the rules given above, # 
least from the first, in which the cotyledons are above ground and 
free, while the second comprises two possibilities, enclosed of iret . 
cotyledons. When the seed-leaves are carried up above ground, this 


*KLEBs, GEorG: Beitrage zur Morphologie und Biologie der Keimung- Unt 
such. aus d. Botan. Inst. 


Tiibingen 1: 536-635. 1885 ‘ 
P * MULLER, FR: Keimung der Bicuiba. Berichte d. deutsch. botan. Ges: a 
7. 


as l yu he 


I 


1899] BRIEFER ARTICLES 61 


hypocotyl and*.two deeply lobed cotyledons, which, although above 
ground, do not leave the seed. To this instance may, however, be 
added a second, but of an order very remote from the Myristicacee. 
A few weeks ago Mr. G. W. 
Oliver, of the botanical gar- 
den at Washington, D. C., 
called my attention to some 
very odd-looking seedlings of 
Jatropha multifida, which were 
kindly submitted to the writer 
for closer study. . The germi- 
nation of this plant had, so 
far, only been very briefly 
mentioned by Sir John Lub- 
bock,? who merely recorded it 
aS an exception from that of 
the other Euphorbiacee. The 
seedling develops as follows. 
Next to the primary root 
the hypocotyl increases very 
considerably in length and 
penetrates the soil by making 
a strong curvature until the 
seed becomes raised above 
ground, while simultaneously 
the petioles of the cotyledons 
ave reached their final de- 


in ae 
kisiguns and the first leaves, Fic. 1.—Seedling of Jatropha multifida L; 
Mg Opposite and of approx- natural size. 


62 BOTANICAL GAZETTE [JULY 


the cotyledons drop off, with their leaves still enclosed. The accom: 
panying drawing (fg. 7) illustrates a seedling with the cotyledons, 
Cot, attached, and the first leaves, Z*, almost expanded. The primary 
root, &, is persistent and branches very soon, while a few secondary 
ones proceed from the base of the hypocotyl. The very long and thick 
hypocotyl, H, is green and glabrous; there is an early development of 
cork-layers ; the bark-parenchyma is broad and traversed by a number 
of laticiferous ducts: The mestome bundles are of normal structure 
and between these are numerous strata of interfascicular cambium. A 
pith occupies the center of the hypocotyl; its cells are like those of 
the bark, thin-walled and filled with starch, but there are no 
laticiferous ducts. As stated above, the cotyledons have long 
petioles and their blades (fg. 2) are mostly oblong witha 
short point, but without any lobation as was observed in 
bicuiba. They are somewhat fleshy and pale in color; never 
ie oc. theless, stomata are present on the upper surface and not on 
Blade of a the lower. Along the ribs on the upper face, two kinds of 
cotyledon ; hairs are visible. These are either long, multicellular, and 
natural size. nointed, or unicellular and almost globular in shape. 
latter (fig. 7) are quite abundant in contrast to the first, and in some 
places they covered the surface just above the ribs. They represent 
glandular hairs, but their function GR 
could not be ascertained. The 
lower surface of the blades, which 
lie close up to the endosperm, is 
wholly glabrous. A typical pali- 
sade tissue was observed, covering 
a rather dense pneumatic tissue, 
and both contained abundant 
deposits of starch; _laticiferous Fic. 3.—Epidermis with glandular ies 
ducts were also very frequent. from the cotyledons of Jatropha multifile 
This seedling has no mate * 24° : 
among the other species of Jatropha, which have been studied hereto: 
fore. Sir John Lubbock figures and describes 7. Curcas L. and J: 
podagrica Hook., both of which possess stout and long hypocotyls 
furthermore, their cotyledons are free and provided with distin 
petioles, although not equaling in length those of /. multifida. 
Another and very peculiar manner of germinating was observed . 
Persea gratissima Gartn., of which several seedlings were cultivated * 


oO 


1899] BRIEFER ARTICLES 63 


the same time in the botanical garden at Washington, D.C. It is 
strange that the Lauraceze have been almost entirely passed by in works 


dealing with seedlings, none having been 
recorded either by Klebs (/. ¢.) or Sir John 
Lubbock (/. ¢.). Schacht‘ appears to be 
the only author who has given us some 
information about them; he states that the 
seed of Persea gratissima germinates while 
the fruit is still attached to the tree. This 
author observed, also, that in this species 
of Persea the plumule attains a very early 
development with a number of leaves, 
similar to Juglans and Tropzolum. Hay- 


* é 
ing detected a few other peculiarities con eee | areas 


nected with the germination of Persea, and 
having been unable to find any figure of 
this, I take this opportunity to publish 
and illustrate my observations, together 
with the still more remarkable case of 
germination just described. 

In Persea gratissima there is no endo- 
‘perm, and the large cotyledons remain 
enclosed by the seed-coat. No hypocotyl 
develops during the germination, but the 
plumule §tOws out very soon as a shoot 
with several leaves, while the primary root 
at the same time has attained a considerable 


strong lateral roots arranged in whorls of 
tata to five or more. In the accom- 
? ying drawing ( Jig. 4) the plumule has 


dey. : 

Pe aS a single shoot, and it is very 
8€ to notice that t 

leaves, 7 he very first four 


imitating 
On the ot 


Fic. 4.—Seedling of Persea 
gratissima Gartn., natural size. 


64 BOTANICAL GAZETTE [yuLy 


leaves, while in Juglans and Carya, for instance, all the first leaves are 
scale or bristle-like. : 

en, however, the plumule does not develop as a single shoot 
but as a complex of ramifications, the first leaves become almos 
suppressed and appear only as small and rather broad 
scales (fig. 5). In this case, the shoots have pushed 
out very freely from the axils of the lowest leaves, and 


these lateral branches, there is a bud observable it 
the axil of each of the two cotyledons (jg. 6), which 
is evidently ready to develop if the 


become injured. The first developed Z wees € ; 
leaves were perfectly glabrous, in oe 


Fic. 5.—Plumule contrast to the succeeding ones. Fic. 6.—Abit 
of seedling of Per- Th 


sea gratissima; nat- of Persea 
eerece lower surface, but none on_ the ©°ty!edon 
upper. The mesophyll formed a 
homogeneous tissue filled with starch, and. there was 
no 


Z3, the first, seco’ 
and third leaf; mag 
indication either of collenchyma or stereome nified. 

above or underneath the mestome bundles. 

In Lindera and Sassafras, at least in their North American rep 
sentatives, the germination takes place underground, but the plum 
develops as a single shoot. The first leaves are bristle-shaped, 
ceeded by a few whose shape is approximately the same as that of the 
typical leaf. The three-lobed leaf of S. officinale Nees, howevet * 
only seldom observed in the first year of the seedling, those develop” 
ing at this Stage being ovate and entire—Tueo. Hom, Brooklanh 
0 8 aE 


1899 ] BRIEFER ARTICLES 65 


ON THE BLIGHT OF SORGHUM. 


THE tissues of the different organs of sorghum, under certain con- 
ditions which are not as yet perfectly known, may-become the seat of 
an intense production of red pigment which impregnates them. ‘The 
cells die and disintegrate, the disease being known as sorghum blight 
(sorgho bralt, Hirsebrand). | 
- The disease was first described in Italy by Palmeri and Comes,’ 
who have attributed it to the development of Saccharomycetes and 
bacteria. Later, in America, Burrill (1887) studied anew the blight, 
and, after having isolated sporogenous bacteria from the infected 
tissues, he attempted the inoculation of healthy plants. The results 
were variable; however, the author inferred the parasitism of the 
Bacillus Sorghi, nov. spec., and prescribed some measures for the pres- 
ervation of plantations from the invasion of the disease. Analogous 
experiments by Kellerman and Swingle’*, more convincing than the 
preceding, because of the number of infections obtained, led to the 
same conclusions. More recently, Bruyning* has examined some 
plants of sorghum attacked by the blight, and formulated some con- 
clusions different from those of the preceding authors. After having 
isolated several bacteria from the red tissues Bruyning retained two 
species which he considered the only factors of the fermentative disease, 
but these species are chromogenous outside the sorghum. These 
Microbes acted symbiotically, superposing their respective pigments, 
the one yellow, the other red, the mixture giving the coloration observed 
in the tissues. These views were not supported by any experiment of 
Moculation, ) 
ie. a studied some specimens of blighted sorghum sa 
Oe thea §etia. aay able to convince myself that in this case 
i see, ee of the blight were caused by the parasitic ss aa 
€ tissues of the plant. 


I  . Fie r 
oo oe preliminari Sopra alcuni fenomeni di fermentazione del Sorgo saccarino 
ese oa d fis, e mat. di Napoli, fase. 12, 7 : Sete 
: port of Bot. Depart. of the Kansas Stat. Agri. Coll. 1888. 
briilure du Sorgho, etc., et les bactéries qui la provoquent (Arch. Néerland. 
* PP: 297-330. 1898) he 
Tmacy ems were sent to the botanical laboratory of the School of 
With th Aes Professor Trabut. Professor Guignard had the kindness to entrust me 
~€ examination of them, See ek eRe 


, fai 


66 BOTANICAL GAZETTE [yuLy 


Already, by microscopic examination, it had been ascertained that 
in many regions of the central parenchyma of the stem the cells and 
the intercellular spaces enclosed masses of a small ovoid yeast, measur: 
ing from 1.5 to 2.54 on an average. It was, moreover, the only 
microorganism which could be directly observed, even with great mag: 
nification and with the aid of staining reagents. 

It was easy to isolate this yeast in a pure state by taking up the 
germs with a sterilized needle from the center of the stem, the section 
being made with a flamed scalpel, or by removing small cylinders of 
the red tissue from the pith by means of a sterilized trocar. The first 
culture liquid was a 5 per cent. glucose bouillon. Some subsequent 
isolations by means of Petri dishes with bouillon sugar gelatin have 
given, to the exclusion of all other organisms, white colonies formed 
by a yeast morphologically identical with that which had been observed 
in the diseased sorghum. 

Cultivated on the surface of carrot, potato, and on different gelatin 


in the tissues of the plant. Sown in unfermented grape juice, 0 in 
various artificially sweetened liquids suitable for the culture of yeast 
this organism shows a feeble alcoholic fermentation. ‘I'he fermenting 
power is slightly increased by a series of re-sowings in the samé 
medium. My attempts to obtain ascospores have been fruitless, and I 
am not able at present to classify this yeast with the true Saccharo- 
iyces, 

The mere fact of isolating a yeast from the tissues of the blighted 
sorghum i is not sufficient to enable one to conclude that this yeast isa 
parasite, and that it brings about the symptoms of the disease. It is 
known, in fact, that a large number of these ferments may be encoll 
tered on the surface of stems and leaves; it would not be astonishing. 
therefore, if some dead tissues, still containing in their cells a part 
their reserve sugar, were invaded by the saprophytic development ‘ oft 
superficial yeast. The following experiments, however, show that had 
tissues of the plant, and in them brings about the phenomena of the 
blight. . 

Sorghum plants raised from seed and cultivated in a hothouse que 
ing the months of November and December 1898 and January 1899 
were inoculated with pure cultures. All aseptic precautions were 


1899] BRIEFER ARTICLES 67 


taken to avoid the introduction of foreign microbes. I used Dr. 
Roux’s sterilizable syringe with a fine injection needle, in order to 
reduce to the minimum the wound necessary for the introduction of 
the culture. Only the stems were inoculated. The epidermis was 
exposed by cutting a small flap in the leaf sheath, carefully disinfecting 
by means of a red-hot iron, and, after the inoculation, the opening sealed 
by means of warm wax or sterilized paper. The plants experimented 
upon were successively examined at intervals of time varying from five 
days to two months. In every case the yeast had developed and mul- 
tiplied in the intercellular spaces, and in the interior of the living 
cells to a distance of 10 to 1 5™" above and below the point of inocu- 
lation. 

The microscopical appearance was that which may be observed in the’ 
tissues of sorghum spontaneously infected ; the lesion was also rend- 
ered visible in the same way by the red coloration of the parenchyma 
and fibrovascular bundles. These latter transfuse the coloring matter 
the whole length of the internode, and quite beyond the infected 
region, so that the appearance of the pigment at a certain point of the 
tissue is not a sure sign that the parasite is present at that point. On 
the surface of the stem may be observed long red lines corresponding 
to the outer bundles and their contiguous parenchyma seen through 
the transparent stem. 

In these experiments, the isolation of the parasite from the affected 
pith has shown, by comparison with the control, that the parasitic 
Yeast was the same as that in the cultures used as a starting point. 

t is probable that the reserve sugars of the cells of the sorghum 
“onstitute the principal food of the parasite. Unfortunately the volume 
of infected tissue, limited and especially difficult to separate out, has 
not permitted the changes carried on by the parasite in the chemical 
“omposition of the plant to be estimated in this particular. 
= Jaa capable of producing in aie sorghum similar phe- 
Eade a ? The following experiments answer’ th the 
healthy = oo have been made ‘aseptically in the stems of 
chatnpagne't ta with pure cultures of wine yeast (round yeast of 

‘5 pee: Owzy]). The parasitism has been established under the 
sacs ky oy a before, the yeast developing in the intercellalar 

€ pith cells of the stem with the accompanying produc- 


tion =e : 
be Of the characteristic red pigment, and with transfusion by the 
Ndles of the internode. 


68 BOTANICAL GAZETTE [yuLy 


_ On the other hand, what is the origin of the pigment? Itisa 
general observation that wounds inflicted on the tissues of the sorghum 
developed a red coloration around the injured part. It is important 
to understand precisely in the preceding experiments the role of the 
local lesion induced by the inoculating needle. Aseptic punctures in 
the pith of healthy sorghum stems were made under conditions identi- 
cal with those of the preceding experiments, the inoculating fluid alone 
being omitted. Under these conditions the pigment appeared in the 
wounded cells, but it was not abundant and was rigorously confined to 
the wound. The quantity of coloring matter thus produced is not 
capable of being carried by the bundles and of spreading beyond the 
actual point of the lesion. The experiment shows, however, that the 
chromogenous property belongs to the wounded cells of the sorghum. 

From the preceding facts it may be concluded : 

1. That yeasts may develop in the living cells of the sorghum. 

2. That the parasitism of these yeasts may bring about an intensé 
red coloration of the plant tissues, this coloration being the same 4s 
that which may be observed in the disease of sorghum called the blight. 
The production of the pigment appertains to the affected cells, and the 
parasite takes part only through the lesion which it produces. 

_ These results confirm the old hypothesis of Palmeri and Comés, — 
who, observing the fermentative phenomena of the red juice of the 
pith of the blighted sorghum, had inferred from it the parasitic action 
of the Saccharomycetes without giving it experimental proof. € 
same facts, moreover, are not contradictory of the experiments of Bur: 
rill, Kellerman, and Swingle. In fact, it may be concluded that, the 
ted coloration being the result of a chromogenous function character 
istic of the wounded cells of the plant, different parasites, yeasts 
bacteria, may, by developing in the tissues, induce by continued lesio® 
the production of a considerable quantity of the pigment. 

_ On the contrary, it is necessary to make complete reservations * _ 
to the conclusions of Bruyning, who, attributing to the bacteria them 
selves the chromogenous function, denies to all micro-organisms 1ac¥ 
ing this function while outside the host plant the power of inducing the | 


aes of sorghum blight.—Maxime Ranals, School of Pharma), ~ 
aris. 


1899 | BRIEFER ARTICLES 69 


ROOT SUCKERS ON DOUGLAS FIR. 


THE occurrence of stool shoots among deciduous species of trees is 
very common, and their production by the forester is resorted to for 
the reproduction of many species. Among conifers the formation of 
stool shoots is limited to a few species. The California redwood 
(Sequoia sempervirens) is very commonly reproduced in this way. Much 
less common than stool shoots are root shoots, often known as root 
suckers or suckers. Among conifers only the redwood, the California 
nutmeg (Zumion Californica), and the short-leaf or yellow pine (Pinus 
echinata Miller) have so far been reported as producing root suckers. 

For several years the woodsmen of western Washington have 
recognized in the fir forests a curious growth which they have called 
“sap suckers.” As we see them in the forest they appear as a broken 
stub ranging in height from 0.6 to 3.5". Without leaves or branches, 
they appear entirely lifeless until cut into with an axe. An examina- 
tion shows that they are covered with living bark and beneath that a 
living woody tissue very hard and with a grain of very fine and intri- 
cate burl. 

The sap suckers vary greatly in size and external appearance. The 
diameters range from 30 to 60™ and the height from 0.6 to 3.5”. 
Investigation has shown that they are connected with one of the main 
Toots of the parent fir, their distance from the parent trunk varying 
from 0.6 to 4.5", an average distance being about 1.8". The living 
bark bears little resemblance to the bark of the fir except in color. 
The clefts are much finer and the plates much smaller. The living 
—, in every instance, forms only an enveloping sheath about a 
pen ae Bee It varies in thickness from 12 to 50" and there is 
a eins in the disposition of this living sheath. at begins at or 
he . ace of the ground and grows upward, entirely encircling 
wes vie ies nat In vaeecap cases where the stub was very short the 
ptottiberancs * :d =a it over, forming sometimes a hy globular 
bad ED os oat 0.6" high and sometimes a column 3.5" high; but 
veda. a. Cea oad of cases the burl covering has extended to only 

€ height of the stub, so that the rotting dead end still 
Protrudes above. 
we ad es is only a secondary growth on an ordinary root 
Ormation of these on the roots of the Douglas fir, Pseu- 


dotsy a . ‘ 
82 taxifolia, was proven by examination. In the dense woods of 


7oO BOTANICAL GAZETTE [uty 


this region it is hard to distinguish the suckers from ordinary seed- 
lings. Like most seedlings, those of the fir are very weak, and when they 
reach a maximum diameter of about 25™ they die and begin to decay. 
At this time the formation of the secondary burl covering begins. The 
nourishment before given the sucker is utilized in covering over the 
dead and decaying stub with this new live growth. 

In one instance a dead sucker had been covered with a primary burl 
covering to a height of 1.5". This had died, the bark had fallen of, 
and a second burl covering, still alive, had covered this over to 4 
height of 0.9". 

These suckers are found only in the moistest and densest forests. 
Even under the proper conditions they are very rare. They were 
observed in the summer of 1898, ten to fifteen miles from tide water, 
Chehalis county, Washington.— Frank Haines Lamp, Biltmore, N.C. 


MURRENT LITERATURE. 
BOOK REVIEWS. 


General physiology. 


THE RECEPTION by both European and American biologists of Ver- 
worn’s Allgemeine Physiologie upon its publication late in 1895 was very 
cordial. The book awakened so widespread an interest that within three 
years a second edition was issued. This second edition has now been trans- 
lated and edited by Dr., Frederic S. Lee, of Columbia University.’ In its 
English dress the book will doubtless commend itself still further to English 
readers. Certainly it will put within reach of general readers a work that 
will give a better idea of the scope of physiology, and one that will present 
to special students readable and stimulating discussions of physiological 
problems, 

The work may be called a treatise upon “general” physiology only in 
the peculiar sense that it discusses the general as opposed to the special 
functions of the cell. It does not deal with the functions of organs at all. 
iC the whole the phrase general cell physiology would seem to describe it 

tter, 


__ Itis a pleasure to find an animal physiologist who is yet to a reasonable 
degree familiar with plant physiology. But it is not for the views of plant 
function hor even for the facts adduced that the book may be commended to 
botanists. In these fields, indeed, one feels that Professor Verworn is travers- 
ing rather unfamiliar ground, in which he sometimes loses his way, to the 
leading astray of the unwary. Nevertheless the ability and suggestiveness of 
the book make it one which botanists inclined at all to physiology will do 
te to read. Professor Verworn has a luminous way of saying things, and 
vane What he says sometimes suggests what not to say, more often his apt 
applications indicate others which he has not expressed. 

The subjects, treated, after an interesting introductory chapter on the 
nd methods of physiological research, are as follows: living substance ; 
Mposition and the differences between living and lifeless substances; 
entary vital phenomena, namely the phenomena of metabolism, of the 
8¢s of form, and of the transformation of energy; the general conditions 

Tt 
lated Mi, 2 Max.—General physiology, an outline of the science of life. Trans- 
xvi tee re ome German edition and edited by FREDERIC S. LEE. 8vo. pp- 
1899] ‘205. London and New York: The Macmillan Company. $4.00. 


its co 


chan 


72 BOTANICAL GAZETTE JULY 


of life, including a discussion of the origin of life and the history of death; 
stimuli and their actions; and the mechanism of life. 

No detailed discussion of the book need be entered upon, since itis 
already fairly well known in the original form. . The translation has bee 
admirably done by Professor Lee. The smooth and readable German of Ver 
worn has been converted into easy and idiomatic English.—C. R. B. 


Some popular books. 


ALICE LOUNSBERRY is the author of A guide to the wild flowers? recenlly 
published. Mrs, Ellis Bowan has had charge of the illustrations, which 
consist of sixty-four colored and one hundred black and white plates, and 
fifty-four diagrams. Dr, N. L. Britton has written a brief introduction. The 
numerous attempts to provide easy and interesting ways of recognizing plants 
indicate'a real demand and one that is very hopeful.. Nature study is finding 
a prominent place in the schools, and any book which stimulates it properly 
is to be commended. The plants selected for this book are well illustrated 
and fairly wéll described, and should be recognized easily by the intelligent 
observer. Although strictly taxonomic, the plants are presented in ecologic 
groups; as for example, “plants growing in water,” “ plants growing in mols 
soil,” “ plants growing in dry soil,” etc., etc.—J. M. C 


what things are and what they are for, so far as current knowledge §%% 
The titles of the chapters are suggestive of the topical character of the bod 
A few of them are as follows: crocuses, dandelions, the flowering of the 
forest trees, green leaves at work, grasses, climbing plants, a handf : 
weeds, in winter woods, etc. The photographs are especially excellent, ™ 
_ Some of our common plants are made to stand out with remarkable distint 
ness. The book can be commended to all those who wish to come into “ 
tact with nature in an untechnical way, and also to teachers in charge" 
nature study.—J, M. C, a 
DWARD KNOBEL has attempted to make the identification of the ee 
sedges, and rushes of the northern United States an easy matter.’ The plate 
7A guide to the. wild flowers. 8vo. pp. xvii+ 347. New York: Frederick & 
Stokes Company. 1899. ‘ tee 
* Field, forest, and wayside flowers. 8vo, pp.xvi-+ 411. New York: The = 
& Taylor Company. 1899. $1.50. 7 ‘ 
“The grasses, sedges, and’ rushes of the northern U. S. 8vo. pp. 78» #O 
Boston: Bradlee Whidden, 1899. $1.00. . a 


1899] CURRENT. LITERATURE 73 


are numerous, and oné of the features of the book is the series of marginal 
illustrations of essential structures opposite each description. Just how easy 
this will make the determination of species in these perplexing groups can 
be known only after experience. It is a question whether anything in clear- 
ness has been gained by substituting ear and earlet for spike and spikelet— 


J.M.C 


A NEW NATURE READER has been provided by Kate Louise Brown, 
under the title Zhe plant baby and its friends3 While the title suggests that 
the author has concerned herself with plant seedlings, a reading shows that 
the adult plant has received most consideration. Facts concerning many 
common plants are presented in the form of stories intended to be used as 
supplementary matter with younger children ; and for such purpose the book 
should find a place. _ Some objectionable features of many books intended to 
assist children in nature study are happily absent from this book. Among 
these may be mentioned the treatment of fertilization and nomenclature. The 
stories concerning pollination are told in such an interesting way that even 
the youngest pupils should be interested. In some cases attempts at color- 
ing as well as attempts at drawing lessons in morals from the plants studied 
have done violence to the facts, and the unfounded story of Egyptian mummy 
wheat is told once more. But aside from a few such errors it should be said 
that the plan and spirit of the book will make it very helpful with younger 
pupils —Ort1s W. CaLpwELL. : 


FREDERICK LERoy SARGENT has produced an interesting little book 
for young people entitled Ze corn plants, their uses and ways of life® Of 
Course the phrase “corn plants”’ means cereals. The first pages are devoted 
to the mythology of the cereals, embracing the stories of classical antiquity to 
which allusions are so frequent in more modern literature. 

___ For the botanist the chief interest is found in those pages which discuss 
the ecological adaptations of these important plants: A great deal of atten- 
Hon 'S 8lven to such"questions as protection against wind, weight, and excesses 
of moisture and dryness, several new drawings illustrating the structures which 
ra the plants in these respects. Few plants seem to be more interesting 
ze the ecological standpoint, when one considers that they are grown in 

Posed places, absolutely unassisted by close relation with other plants 

€ to modify the effect of the wind or the sunlight. | ; 
Peis section on the advantages of cereals as food-plants the author 
resting descriptions of the methods of using céreals and the extent 
Cie plant baby and its friends. 8vo. pp. 155. Boston: Silver, Burdett & 
Y- 1899. 48 cents, 

6 - 

wo plants, their uses and ways of lifé. 8vo. pp. v-+106. figs. 32 
Oughton, Mifflin & Company; 1899. 75 cents. aes 


74 BOTANICAL GAZETTE {your 


and geographical distribution of the various sorts. The book is full of good 
information put in an attractive style and should find abundant welcome— _ 
O. W. CALDWELL. 


MINOR NOTICES. 


THE THIRD FASCICLE of “Illustrations de la Flore du Congo,” by Wik 
deman and Durand, has just appeared, containing twelve plates, with des — 
criptive text. The plates are exceedingly handsome, and their number ha 
now reached thirty-six.—J. M. C. 


A NEw classification of the Leucobryacezx is proposed by M. Jules — 
Cardot in Revue Bryologique 2631-8. pl. 7. 1899. It is based chiefly upo 
the anatomical characters of the leaves as shown by cross-sections, such 
the presence or absence of sclereides, and the form and arrangement of tit 
chlorophyllose cells.—C. R. B 

SOME RESEARCHES of Loeb upon the influence of alkalies and acils 
upon embryonal development and growth’ led to results which may hae 


NOTES FOR STUDENTS. 


. pears lea 
GEORGE J. PEIRCE has been studying the nature of thea Pe 
and fungus in lichens? 


€ author also affirms that the central body of the gonidil 


as Ramalina, Usnea, and Spherophorus, is a nucleus, not a pyrene! . 


HERMANN VON SCHRENK® has been investigating a disease of 7% ee 
known as peckiness, and also a similar disease ot Libocedrus ss y 
both cases the wood is destroyed in localized areas, which are surround —e 


7 Archiv f, Entw.-mechanik der Organismen 7:631-641. pi. 7. 1898. 
* Proc. Calif. Acad. Sci. II. x: 207-240, pls. 41. 1899. 


9Eleventh Ann. Rep. Mo. Bot. Gard. I-55. pis. 6. 1899. 


a a a a 


RT IRAE, I PRS ee ee a ae ee ee eT Re MPS Mee Fe ew eR ee 


1899] CURRENT LITERATURE 75 


wood apparently sound. In both, also, a fungus mycelium was found, which 
flourishes within the diseased centers, but no characters appeared by which 
it could be determined. It is interesting to note the fact that these two treés, 
similarly affected, are the representatives of a race largely extinct, and grow 
in different parts of the country. 

The same author (of. czt. 6-70) has also discovered a peculiar sclerotioid 
disease of beech roots. Small tubercles were found attached to the fibrous 
roots, and upon examination these were found to be a twisted and contorted 
mass of rootlets invested by a sheath. It was impossible to determine the 
fungus, but it seemed probable to the author that it is distinct from those 
forming the mycorhiza.— J. M. C 


MEssrs. FARMER and WILLIAMS have continued their contributions to 
the life-history and cytology of the Fucacez by publishing a second paper” 
which deals with the development of the oogonia and oospheres, the phenom- 
ena of fertilization, and the early segmentation stages of the oospore, espe- 
cially in the genera Fucus, Ascophyllum, and Pelvetia. The more interesting 
subjects discussed are the reduction in the number of chromosomes, the 
appearance and behavior of centrospheres and centrosomes, the formation of 
the walls of the oogonia, oospheres, and germinating oospore, and the part 
played by mucilage in bringing about the extrusion of the oospheres. 

The reduction of chromosomes occurs in the papilla which is cut off to 
form the rudiment of an oogonium. This was also shown by Strasburger to be 
the case in Fucus. In the three successive mitoses by which the eight 
magnon nuclei are formed, the reduced number of chromosomes is retained. 
Prominent centrospheres in the form of converging cytoplasmic radiations 
“re present in every mitosis, whether of the oogonium or segmenting oospore. 
ee Could not be traced back to preexisting centrospheres, but appeared to 
‘rise independently by the activity of the protoplasm, and to disappear again 
nhl Though the centrospheres are cytoplasmic, the spindle 
between them is intranuclear in origin. Centrosome-like structures were 
— seen within the centrospheres, but their number was not constant. 
aps was no indication that the centrospheres of the first oosporic division 

erived from the spermatozoid. 

Se Peas polyspermy were observed that it is certain but one a 

newly ones eee on oosphere. The existence of a wall aroun : ¢ 

ttaheServest C €8§s was conclusively demonstrated ; such eggs burst : - 

treated merely a. water to fresh, whereas unfertilized eggs similarly 

anes nterestin ? sen os ee ilizati observed in 

Halidrys, Ges & aterce with regard to fertilization were : 
PS of gyrating spermatozoids were seen distributed over the 


10 Phj 
hil. Trans. Roy. Soc., London, B. 190 : 623-645. 1898. 


76 BOTANICAL GAZETTE [yun 

surfaces of the oospheres, each attached by one cilium. Suddenly the sper 
matozoids leave the egg “like a crowd of startled birds,” the egg at the same 
time becoming covered with conical projections. The authors consider thi 
phenomenon as marking the moment of fertilization, and as a proof of a defi 
nite repulsion of the supernumerary spermatozoids.— WILSON R. SMITH 


revives the term “epithelium,” previously used by Warming and Goebel, 1 
assigning to the layer a digestive function, as did Guignard. ; 

The antipodal cells, variable, as usual, both in number and size, are sai | 
to be digestive in some plants, and in others, conductive in function. Thel 
densely staining protoplasm and their tendency to burrow back into tht 


“pseudo-chalazal’’ region are cited in support of this view. Moreover, ti | 


instead of being mutually hostile, are in perfect harmony. Would the _ : 
say that the “epithelial layer” is not epidermis, merely because it has here? 


the same process in a macrospore; and it would also have become eV" 
that he was not confounding the two terms “ oosphere”’ and “ macrosp . 
ne vested MLLE. MATHILDE : Sur la structure et les fonctions de l’assise ee 
liale et des antipodes chez les Composées. Journ. de Botanique 12: 374-384 © 
13+ 9-17, 49-59, 87-96. 1899. 


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1899] CURRENT LITERATURE 77 


We cannot help wishing that the author had given complete drawings of 
some of the more striking ovules, such, for example, as Dah/ia gracilis, with 
the cell contents accurately shown. When a point is proven “by the appear- 
ance of the protoplasm, and the distribution of starch and other foods in the 
ovule,” one quite naturally expects in the drawings definite delineation. In 
this, however, the six plates are sadly lacking. 

he author has touched an important and suggestive field. Results from 
a comparative study of other groups ought to be easily accumulated from the 
many permanent preparations of ovules and embryo sacs in the possession of 
various botanists, which eet repay reexamination from this new point of 
view.— W. D, MERR 


NEWS. 


Dr. M. TREvs, director of the botanical gardens at Buitenzorg, has beta 
elected a member of the Royal Society of London. 


Mr. H. G, TIMBERLAKE, instructor in botany in the University of Mut | 
gan, has accepted a corresponding position in the University of Wisconsin. 


A BIOGRAPHICAL SKETCH, with portrait, of the late Edward Lewis Stu 
tevant is published by C. S, Plumb in the tenth report of the Missouri Bota _ 
ical Garden. 


Mr. Ernst A, Bessey has been appointed assistant in the a 
Vegetable Physiology and Pathology of the United States Department 
Agriculture. 


M. Ep. PRILLIEUX, the eminent French phytopathologist, has zs 
elected a member of the French Academy of Sciences, in the room t 
late Charles Naudin. 


PROFESSOR W. A, SETCHELL and other botanists of the va t 
California are about to leave on an expedition to study the flora of the t 
tian islands. — Science, June 23. : 


y at 

PROFESSOR P. H. Rots has been appointed professor of botany 4 

Clemson College and botanist to the Agricultural Experiment Station ®— 
South Carolina, to succeed Dr. A. P. Anderson. a. 


. ) . 

PROFESSOR Henry G. Jesup, for twenty-two years professor of botany” 
Dartmouth College, Hanover, N. H., has resigned, and Mr. G. T. Moore 
Harvard University, has been appointed instructor in botany. 


THE PROPRIETORS of Nature announce that they are about to reise 
Sowerby’s English Botany, third edition with supplement, containing ~— 
tions and life size figures of every British plant, hand colored. YS 
varies from 16 to 19 guineas, according to binding. : - 
THE INSTRUCTORS and fellows of the University of Chicago ee : 
received botanical appointments for the coming year are as follows: io | 
W. Caldwell, professor of botany, State Normal School, Charleston, Ill; Fe 
G. Coulter, assistant in charge of botany, Syracuse University ; Florent 
Lyon, assistant in botany, Smith College; Dr. W. D. Merrell, an 


78 


ee Oe ae Re ey ee Fe oe ee ee ey Se 


1899 NEWS 79 


charge of botany, University of Rochester; Herbert F. Roberts, assistant in 
Shaw School of Botany; Dr. Wilson R. Smith, instructor in charge of botany, 
McMaster University, Toronto, Canada. 


THE BoTANICAL SEMINAR of the University of Nebraska is pushing work 
on the survey of the state and hopes to send additional parts of the “Flora” to 
the printer before the year is over. The Regents of the University made a 
small grant for a printing fund for the survey at their last meeting. 


BY THE WILL of the late Professor O. C. Marsh, Yale University is given 
his fine residence with the extensive grounds and greenhouses for a botanic 
garden. The house may be used as the residence of the director, or as a 
botanical laboratory. It is to be hoped that this gift will stimulate Yale to 
develop its botanical work more worthily. 


Dr. Epwarp A. Burt, through the courtesy of Dr. W. G. Farlow, has 
been studying the Thelephoracee of the Curtis herbarium and published 
exsiccati in the cryptogamic herbarium at Harvard University. He expects 
to examine types and authentic specimens at Kew during July, spending the 
remainder of the summer in Sweden collecting and studying fleshy fungi. 


AT CoRNELL University: Mr. W. A. Murrill has been appointed for 
the year as assistant cryptogamic botanist to the Experiment Station, to 
take charge of the work of Dr. B, M. Duggar during his absence in Europe. 
Messrs. Heinrich Hasselbring, Judson F. Clark, and George J. Hastings 
have been appointed assistants in botany. Dr. K. M. Wiegand has been 
Promoted to an instructorship in botany. ? 


AT THE UNIVERSITY OF MICHIGAN: Professor V. M. Spalding, who has 
been in California during the year, has returned to the East and will resume 
we work next autumn. Professor F. C. Newcombe will spend the summer in 
Paris to study the scientific institutes of that city. Dr. J. B. Pollock will 
have charge of the botanical work in the University summer school. Dr, Julia 


nad and Mr. Pond will do work for the U. S. Fish Commission at Put- 
in-Bay, 


THE PLANS of the botanical staff of the University of Iowa for the summer 
ra oeutiies Professor B. Shimek will be engaged in special studies of 
tis mere Problems in Iowa, under direction of the U. S. Department of Agricul- 

ae Savage will probably complete his studies of the Mosses and 
Mr 


there for report to the United States Department of 
» and incidentally picking up the fungi of the region. 


80 BOTANICAL GAZETTE [JULY 


Tue RuHoDE ISLAND College of Agriculture and Mechanic Arts, with the 
cooperation of Hon.»Thomas .B.. Stockwell, state commissioner of pub ; 
schools, and Dr. Horace S. Tarbell, superintendent of schools in Provident, 
proposes to open a summer school for nature study at Kingston, R. I, from 
July 5 to 19, 1899. A general summer school is not contemplated, and te 
work offered by the various departments constitutes a single course dealing 
solely with local phenomena in their adaptability to the teaching of nature 
study. The distinctive feature will be the study of living nature rather than 
post-mortem biology. Plants will be studied in the field, supplemental 
laboratory work when necessary to a clear understanding of the subjet 
treated, No attempt will be made to determine species, but it is prop 
to treat the following topics: relation of plants to each other, relation 

lants to animals, oe for cross pollination, and adaptations for 
distribution of seeds and fruit ere will be no entrance examination 
no special requirements for engi of the course. 


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Vol. XXVIII AUGUST 1899 No. 2 


THE 


BOTANICAL GAZETTE 


EDITORS 
JOHN M. COULTER, 7ike University of Chicago, Chicago, Il. 
CHARLES R. BARNES, Zie University of Chicago, Chicago, 11. 
J. C. ARTHUR, Purdue University, Lafayette, Ind. 


ASSOCIATE EDITORS 


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& University of Bonn 
J. B. DeETONI VOLNEY M. oa 
5 Univer a of Padua University of Michigan 
—- SDOLF ENGLE : ROLAND THAXTER 
aa Duar of Berlin Harvard University 
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__3N26 MatsumuRA EUGEN. WARMIN 
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CONTENTS 


THE DEVELOPMENT OF THE MICROSPORANGIA AND MICROSPORES OF HEME- 
al Ri 


Pr OCALLIS FULVA (WITH PLATES Vil-vill), Edward L. Fullm - 81 
a HE SPORE-MOTHER-CELL OF ANTHOCEROS, oe FROM THE HULL 
_ _ Borantcar Lazoratory. XV. (WITH PLATES IX AND x.) Bra -y Moré Davis es 
STRUCTURE AND DEVELOPMENT OF CRYPTOMITRIUM “TENERUM (WITH 
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RIEFER ARTICLES. 
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VOLUME XXVIII ; NUMBER 2 


MeTANICAL (GAZETTE 
‘ AUGUST #899 


d THE DEVELOPMENT OF THE MICROSPORANGIA 
» AND MICROSPORES OF HEMEROCALLIS FULVA:* 


EDWARD L. FULLMER. 


(WITH PLATES viI—vi1). 


ALtHoucH Hemerocallis fulva has been a frequent subject of 
Study, the results reported in reference to the phenomena of 
_karyokinesis and spore formation do not seem to agree. In the 
: Cytologische Studien,? this plant is described with multipolar 
Spindles in the reproductive cells during the early division stages, 
no centrosomes being found in such cells. The division of the 


feproductive cells of the higher plants studied, according to the 
Tesults of the sever 


-mother-cells of Hemerocallis are not very favor- 
ogical study, the chromosomes being small and the 
as cult to differentiate on account of a band of dark 
= © Material found close dround each nucleus; nor are the 
e piadle threads so thick as those usually found. The dark band 
only an artifact, caused by the chromic acid of the kill- 
ae a The material ‘for this study was killed in chrom: 

# acid, = after being imbedded in paraffin was ine we 
and 144 thick. The stains used were iron-alum-hamatoxylin ; 

* Contribution from the bo 
* Jahrbiicher fiir wiss. Bot. 


spindles diffi 


tanical laboratory of Ohio State University. 
30: 1-268. A/. 78. 1897. 
Se 


82 BOTANICAL GAZETTE [avcust 


Flemming’s triple stain; a combination of anilin-safranin with 
iron-alum-hematoxylin; and anilin-safranin followed by picte- 
nigrosin. The last named gave the best results in staining the 
incipient spindles. 


DEVELOPMENT OF THE MICROSPORANGIA. 


A cross section of a very young stamen, at the point where 


the microsporangia are to be formed, shows merely a rectangular 
area which consists of epidermal and general tissue cells. Three 
or four hypodermal cells of each sporangium become differenti: 
ated as archesporial cells. These divide by periclinal division, 
giving rise to the primary sporogenous cells and the primal) 
tapetal layer. The cells of the primary sporogenous tissue mul 
tiply rapidly, forming only sporogenous cells. The division 0! 
these cells is practically complete when the primary tapetl 
layer begins to divide. While the sporogenous cells are enlarg: 
ing the division of the primary tapetal layer takes place, ye 
ing a wall layer and an inner layer. The inner layer divides 
once, forming an intermediate or middle layer, and the layet 
which develops into the peripheral part of the tapetum a 
axial part being derived from the adjacent general tissue. 


DIVISION OF THE MICROSPORE MOTHER CELL. 

A typical spore mother cell when division begins (fg) 
shows a large nucleus in the center, with radiations in the cyt? 
plasm extending from the nucleus outward toward the cell wal 
There is usually a layer of deeper staining cytoplasm next * 
nucleus, and one or more large nucleoli just within the nucle 
membrane. The young spindle is bipolar from its first apper 
ance, each pole being dome-shaped for a time ( figs. 10). 
spindle encloses the nucleus through which the chromosom™ 
are scattered, and the layer of deep staining cytoplasm _ 
moved outward from the nucleus spindle lies wholly within ™ 
The radiations above described appear to be of the same ea 
and to have the same function as those described in animal c” 
such as fish eggs, If they are consumed in spindle formation 


: 
; 
| 


1899] MICROSPORANGIA OF HEMEROCALLIS FULVA 83 


they must be drawn entirely within the dark layer before the 
nuclear membrane has disappeared. 

A careful study was made of preparations containing cells in 
various stages of division, many of which showed distinctly 
dome-shaped bipolar spindles just after the nuclear membrane 
had disappeared. Occasionally a nucleus was observed which 
seemed to have radiations extending outward, but these were 
never sufficiently distinct to be considered as forming a multi- 
polar spindle. The spindle was not differentiated by the stains 
in any case until after the nuclear membrane had almost 
disappeared, as the dark band obscured the poles. At first 
these alone would be visible, since the remainder of the spindle 
could scarcely be distinguished from the nuclear membrane 
because of their close proximity; the central part of the spindle 
perhaps being in contact with the nuclear membrane. The 
spindle fibers at this stage are delicate, the spindle being in 
Process of formation. The fact that the nuclear membrane 
disappears while the spindle is forming lends support to the 
theory that the material of the nuclear membrane is consumed 
In spindle formation. 

As the poles separate, the spindle gradually becomes pointed 
and seems to grow somewhat narrower in the middle portion, so 
that it is not as wide as was the original nucleus ( figs. I-7, 
10-12). ‘The chromosomes are drawn into the equatorial plane 
(figs. 8,74) soon after the spindle becomes elongated to definite 
ia Small deep-staining bodies which have the appearance 
ye gaa are often found at the poles in the various stages 

. — yaaa divisions (Jigs. 9: 9 i Ea) ‘ 
idler ce 3: i vision. of the spore-mother-cell the bi 
Re as oe successively (fig. 22), but almost always 
Ge ies Y (figs. 34, 76), the spindles usually being parallel 
16), and oa (Jig. 15), but occasionally obliquely placed (fig. 

€n persisting even after division is complete (fg: 


17). 
ah e - Phenomena of the first and second divisions are the 
band j Xcept that in the latter the nuclei are smaller and the dark 


1s i 
not so conspicuous, 


84 BOTANICAL GAZETTE [AvGUST 


Four microspores (figs. 18, zg) are usually produced from 
each spore-mother-cell, but occasionally five, six, and even eight 
are formed (figs. 20-27). Miss Lyon? also found this pecul- 
iarity in Euphorbia corollata, which in the case of Hemerocallis 
has been known for some time. The extra nuclei were fist 
described as being produced by one or more of the four tetrad 
nuclei dividing by karyokinesis. Recently, however, they have 
been described by Juel+ as being formed from chromosome 
which became isolated in one of the divisions of the spore 
mother-cell. Many tetrads having supernumerary nuclei wert 
examined, only a few of which seemed to show definitely how the 
extra number of nuclei might have arisen. In all cases whet | 
the origin was indicated by spindles or otherwise they seemed 
to have been produced by a subsequent karyokinesis. The extra 
nuclei in figs. 20 and 27 may be a result either of the indirect 
division of one of the cells of the tetrad, or of the threefold 
division of one of the nuclei at the second division. The 
division of one nucleus into three or more is quite common 
animal cells in case of pathological tissue. Wilson,’ in speaking. 
of cases of pathological mitosis, says: 

The abnormal forms of mitoses are arranged by Hanseman in two aft 
eral groups, as follows: (1) asymmetrical mitoses, in which the chromosom® 


is formed. .:. ... Lustig and Galeotti (’93) showed that the unequal 
ution of chromatin is correlated with and probably caused by aoe 
inequality in the centrosomes, which causes an asymmetrical developme™ 
the amphiaster. a 
The same author refers to the discovery of Galeott! 
asymmetrical mitoses may be artificially produced in the 
thelial cells of salamanders by treatment with various @ 
Guignard?® finds very irregular and also multipolar spindles sale 
3 A contribution to the life history of Euphorbia corollata. BOT. Gaz. 25:41? 
8. | ; 
‘Jahrbiicher fiir wiss. Bot. 30 : 205-226. 1897. 
> The cell in development and inheritance, 67-68. 1896. 
° Ann. Sci. Nat. Bot. VIII. 6:177-220, 7. g-r7. 1898. 


1899] MICROSPORANGIA OF HEMEROCALLIS FULVA 85 


spore-mother-cells of Mymphea alba and Limodorum abortivum, 
and also in the tapetal cells of Magnolta Yulan. A number 
of spindles from cells of the above-named plants and also from 
the cells of Nuphar luteum, having well defined centrospheres and 
radiations around the poles, are shown. 

If the fifth nucleus in figs. 20 and 27 was formed by the 
division of one of the tetrad nuclei, the division was probably 
normal. In this case the presence of spindles suggesting tri- 
polar mitosis would be explained by the persistence of spindle 
structures as in fig. 77. Fig. 22 is similar to figs. 20 and 22, 
except that one of the cells resulting from the first division of 
the spore-mother-cell did not divide. /%g. 23 may have arisen 
by any one of the processes described under figs. 20 and 27, 
but since there is no spindle connecting either of the two smaller 
nuclei with one of the larger, they may have arisen from a chro- 
mosome which was isolated in the first division of the spore- 
mother-cell, forming a nucleus and afterward dividing. In this 
case but one of the cells resulting from the first division could 
have divided, as there are only three other nuclei present. How- 
ever, the three are nearly uniform in size, and all have the general 
appearance of tetrad nuclei, and not the appearance of a nucleus 
formed in the first division, as the elongated nucleus in fig. 22, 
which has the usual shape of such nuclei. 

Hig: 24 shows four nuclei, two of which are dividing by 
karyokinesis, The appearance and position of the cell walls, as 
well a the unequal size of the nuclei, suggests that the two nuclei 
tae from the first division of the spore-mother-cell were of 
oo By the subsequent division of the nuclei thus 
again dec pairs of nuclei resulted; the larger pair of a are 

y Su ee by karyokinesis. This figure may also be explained 
an ead the two smaller nuclei to have been produced from 
could =e op paamigag In this case the spore-mother-cell 
One ce “i ee but once, and is now dividing a second time: 
niche ve nuclei in fig. 25 is dividing. The source of this 

not be determined with certainty ; but it may have been 


forme . . 
¢ cither by the indirect division or by the fragmentation 


86 BOTANICAL GAZETTE | aucust 


of one of the tetrad nuclei, more probably the smallest one 
The fact that several, perhaps the normal number of chromosomes 
are present opposes the view that it arose from an isolated 
chromosome. Six nuclei are found in fig. 26, the origin of 
four being shown by spindles. The two extra nuclei are quite 
small, containing but little chromatin, and may have been pto- 
duced by isolated chromosomes as above described. However. 
they were produced in this manner the smallest of the fou 
nuclei must have arisen by pathological mitosis by an unequal 
distribution of chromatin, for it is very small and contains but 
little chromatin. Figs. 27 and 28 show tetrads with supernumer 
ary cells. The cells of such tetrads are seldom of equal sid 
one or more of the cells usually containing a small nucleus. 


THE DEVELOPMENT OF THE MALE GAMETOPHYTE. 


The microspores soon after their separation and rounding of 
acquire the general shape and characteristic markings of the 
mature pollen grain, gradually becoming very much enlarged. 
(figs. 28, 30). A single elongated nucleus, which seems to have 
a small amount of chromatin, is found in the center of the spot 
until growth is far advanced - after which the division into genera 
tive and tube nuclei occurs (fig. 30). Until recently it ee 
thought that the tube nucleus never divides, but Chamberlait 
found many cases of such division in Lilium tigrinum and L. i : 
tum. Smith® found the same thing to be of frequent occurrent | 
in Eichhornia crassipes. emerocallis also shows this peculiatt) 
(figs. 31-35), many pollen grains having two, and a few havilg 
three, four, or even six tube nuclei. These were formed ?) : 
direct division in all the cases that I observed, as were ae 
reported by Chamberlain. Fig. 33 shows a very irregulat : 
nucleus, 

If the tube nucleus is the homologue of the cover cell of 
antheridium of. Marsilia, and represents the wall of an anther 
ium, it might sometimes divide through reversion to its se : 

7 Bor. Gaz. 23 > 423-430. 1897. 

* Bot. Gaz. 25: 324-337. 1898. 


: 
: 
: 
jj 
4 


22S ee ae 


Bots : 
Bee ee 


ois 


4 


1899 | MICROSPORANGIA OF HEMEROCALLIS FULVA 87 


primitive condition. Since extra tube nuclei are produced only 
by fragmentation, so far as known, such division may representa 
pathological condition. It is possible that direct division in the 
higher plants never represents anything more. 


SUMMARY. 

1. Three or four hypodermal cells of each sporangium 
become differentiated as the archesporial cells. The wall of a 
sporangium consists of three layers exclusive of the epidermis. 
The tapetum is a physiological rather than a morphological 
structure, the peripheral part being organized from the wall 
layers and the axial part from the general tissue. 

2. The spindle appears bipolar from its first appearance, 
being dome-shaped in the early stages. No trace of multipolar 
spindles was observed. The spindles often persist for a consid- 
erable time after division is complete. Bodies having the 
appearance of centrosomes are frequently seen at the poles. 

3. The Origin of the supernumerary microspores was not 
absolutely ‘determined. In many cases where their origin was 
indicated by spindles or otherwise they appeared to arise by the 
indirect division of one of the tetrad nuclei. 

* The tube nucleus frequently divides by direct division, 
forming sometimes as many as Six or eight nuclei. 

: In conclusion I wish to express my thanks to Dr. W. A. 
Kellerman and Mr. J. H. Schaffner for valuable assistance and 
criticism, 


Co_umsus, O. 


EXPLANATION OF PLATES VII AND VIII. 

ae psi seh drawn with a camera lucida and are reduced to three 
objective an i mena size. Fig.32 was drawn with a Bausch and Lomb ?2 
22, 23 with a ea He. 2 ocular (X 1200); figs. 3) 4s 91 10, TF. 15s 24% 
with a Ba €itz |, objective and a Leitz no. 4 ocular (< 1600); all others 
usch and Lomb ,, objective and a Zeiss no. 6 ocular (X 1600). 
FIG. 1. Micros . ee ae Ps d 

pore mother-cell in very early stage of first division, show 


ing dark b. : 
to the cel] oo nucleus and cytoplasmic radiations extending outward 


88 BOTANICAL GAZETTE [aucust 


Fic. 2. Loose mother skein stage, showing a dome-shaped bipolar spindle 
which is situated entirely within the dark band. 

Fics. 3-5. Successive division stages, showing centrosome-like bodies a 
poles of dome-shaped spindles. 

Fic. 6. Same as above, but having no bodies visible at the poles. 

Fic. 7. Mother star stage; spindle pointed, having centrosome-like 
bodies, and with radiations at lower pole. 

etakinesis stage; microspores separated. 

Fic. 9. Near close of metakinesis; centrosome-like bodies and radi 
tions present. a 

Fig. to. Early stage of second division, showing dome-shaped spindle 
with centrosome-like bodies; dark band not so prominent as in first division. 

11. Showing one cell in a later stage of division than is the other. 

Fics, 12-14. Successive division stages. : 

Fic. 15. Loose daughter skein stage, showing an isolated chromosome; 
spindles parallel. 

Fic. 16. Same as above, showing spindles lying obliquely to one ane 

Fig. 17. Tetrad having nuclei variable in size and showing persisting. 
spindles. : 

Figs. 18-19. Later tetrad stages. 


e another. 


PLATE VIII, 
GS. 20-21. Mother cells with five nuclei, three of which are connectél 
by spindles, showing their common origin. 
1G. 22. Same as above, except that one nucleus did not divi 
division of spore-mother-cell. 
FG. 23. Five nuclei in mother-cell, two pairs of whi 
spindles. 
Fic. 24. Four nuclei in tetrad, two of which are dividing b : 
Fic. 25. Mother-cell with five nuclei; one is dividing by karyokines® 
Fic. 26. Mother-cell with six nuclei; the origin of two not shown. 
Fics. 27-28. Five- and six-celled mother cells. 
FG. 29. Microspore soon after separation. 
Frc. 30. Normal mature pollen grain having one generat 
nucleus. 
Fic. 31. Tube nucleus of pollen grain dividing by direct division. 
Figs. 32, 34. Pollen grains with one generative and two tube nucle! 
Fig. 33. Pollen grain with a very irregular tube nucleus. 
Fic. 35. Pollen grain with three tube nuclei; generative 1 
shown. 


de after fist 


ch are connected bf 


y karyokines® 
ive and one W 


cleus 


Fig. 36. Six tube nuclei in pollen grain. 


BOTANICAL GAZETTE, XXVIII. PLATE Vil. 


NN  omacaaae 


Vi/1. 


PLATE 


BOTANICAL GAZETTE, XXV/I1. 


FULLMER on HEMEROCALLIS 


THE SPORE-MOTHER-CELL OF ANTHOCEROS. 


CONTRIBUTIONS FROM THE HULL BOTANICAL 
LABORATORY. XV. 


BRADLEY MooORE DaAvIs. 
(WITH PLATES IX AND X) 


Anyone who has ever examined sections of the sporogonium 
of Anthoceros must have been impressed with the extreme beauty 
of the spore-mother-cells as they are exhibited in all stages of 
development throughout the length of this structure. The excep- 
tional chloroplast and the various conditions illustrating the 
division of the cell-contents to form the spores are perhaps the 
most striking features presented. These led the writer to attempt 
a detailed examination in the hope that light might be thrown 
on certain problems that interest the student of plant cytology 

The work of Farmer (’94 and ’g5) is the only contribution 
of a detailed nature upon the cytology of the Hepatice. He 
did not study Anthoceros but confined himself chiefly to certain 
tSalloid Jungermanniacee and reported some very interesting 
and remarkable conditions. The present investigation does not 
“gree with his accounts of the processes of nuclear division pre- 
Be hy such forms as Pallavacinia, Aneura, Fossombronia, 
- However, at the outset the writer wishes plies hose 
forms aa at important differences between such divergent 
The eke . and members ot the. Junge 
taught us to of the past few oo cytology have ale y 
trated by eigh of generalizing upon the conditions illus- 

°Y particular cases. 
aa oo deals with the single species Anthoceros 
where ice oe being collected at Woods Hole, Mass., 
ments on — xhe preliminary studies and exper 
1899] ods of fixation. The best results were obtained 
89 


90 BOTANICAL GAZETTE [avons 


from specimens that had been fixed for twenty-four hours ina 
weak Flemming solution of the following formula: 1 per cet 
chromic acid 25°, 1 per cent. glacial acetic acid 10%, 1 per ceil 
osmic acid 10°, water 55°°. Much material, however, was killed 


killing agents. The results of these experiments are brougit 
together at the end of the paper and present some interestilf 
data. 
Sections were cut 5 and 7.5 ~ thick from material embeddel 


ing to the principles of Haidenhain. Acid fuchsin followed bj 
methyl green or gentian violet presented interesting examples# 
differential staining, but the results were not as good as tho 
given by Flemming’s triple stain. 

At the outset it will be well to state that the number® 
chromosomes for A. /aevis appears to be four and eight in ie 
_ 8ametophyte and sporophyte respectively. The count in me 
gametophyte is based solely upon observations cf many ie. 
figures in the cells of a developing antheridium, but it mee 
with the number presented in the nuclear figures i _ 
spore-mother-cell. The number eight for the sporophytic g® | 
eration was presented quite frequently and in different regio® : 
of that structure. Thus the archesporium just before the diffet 
entiation of the spore-mother-cells, the tissue below the at 


$3 
it 


time when the spore-mother-cell is differentiated until the spore 
are fully ripe. It is possible from this fact to relate te 
other with great exactness the events that take place 8 © 
Spore-mother-cell matures. 


| 
| 
| 


1899 | SPORE-MOTHER-CELL OF ANTHOCEROS gI 


As is well known, the sporogonium of Anthoceros grows from 
a region of embryonic or meristematic tissue in the slightly con- 
stricted portion of the structure just above the foot. From this 
region there rises a central columella around which are arranged 
in the form of a hollow cylinder the cells destined to develop 
into spores and sterile cells (elaters). Recently published 
accounts of the structure and development of the sporogonium 
by Campbell (95, pp. 130-131) describe this hollow cylinder as 
two layers of cells thick and composed of groups of spore-mother- 
cells separated by clusters of sterile cells that later become the 
elaters. The writer’s preparations indicate far less regularity of 
structure than this description would imply. In many cases the 
Spore-mother-cells formed a single layer around the columella 
and the amount of sterile tissue was very insignificant. Only 
eccasionally was the layer of cells distinctly double and then 
Na irregularly so. There appears, therefore, to be considerable 
Variation in the complexity of the sporogenous tissue in Antho- 
feros, and the simplest conditions approach very closely the 
“rangement of cells reported by Campbell (98) for Dendro- 
ceros, 

The cubical cells of the archesporium frequently contain 
nuclear figures, but the spindles are very small and unsatisfac- 
tory for study. Such a spindle is shown in fig. 7, with the 
oo separating into two groups and about to Pe to 
we oe cytoplasm is granular and contains minute 

“Ss. The writer was not able to find a trace of the chloro- 
plast _ these cells, but looks forward to a further study of this 
Point in living material and with special methods of staining. 

8 ae cells become very quickly differentiated 
Mee in size. Their nuclei are not at first to be 
hee ¢d from those in the cells of the archesporium, for 
at are small and contain the one prominent nucleolus and an ill- 
. en ete linin threads. The most asta Seth change 
Plast. This 2 e appearance in fixed material of the chloro- 
bhincs ia Tucture is at first a slightly denser mass of pig: 

€ other contents of the cell, but so small that it 


g2 BOTANICAL GAZETTE [avert 


can scarcely be distinguished from differentiated granular region © 
of the cytoplasm. Indeed, the first clear proof of its preset” 
in my preparations has been the sharp staining of the starch 
grains (purple with gentian violet) that are found almost imme: | 
diately in the interior. The reader may see in fig. 3 such at | 
early stage in the development of the chloroplast, and shouli | 
notice that the three small starch grains lie in a homogene 
matrix of much the same consistency as the surrounding cyte | 
plasm. The chloroplast in cells younger than that shownil 
fig. 3 could not be recognized with certainty by the methods 
staining employed in the present study. - 

The chloroplast once differentiated soon becomes the mo | 
conspicuous object in the spore-mother-cell by virtue of itso | 
and brilliant differentiation with stains (figs. 4, 5) pect 
arities become more apparent as it enlarges. The outline» | 
sharply marked but it has been impossible to establish a distin® 
bounding membrane. The interior contains beautifully define! 
starch grains that become very numerous as the chloropl# 
increases in size and finally fill by far the greater part of th 
enclosed region. Each starch grain at first occupies @ 1 
cavity surrounded by delicate films of protoplasm that app™ 
as strands in the sections figured (figs. 5-9). The chlorop 


has therefore a honeycomb-like structure, each cavity Dem 


chloroplast where they merge into the densely staining fil 
the exterior. But the bounding film grades off insensibly ™ 
the cytoplasm and it was not possible to distinguish by 4%) 
reaction the strands inside the chloroplast from the protel 
outside. 

The first division of the chloroplast takes place very ee 
after the structure reaches a certain size. Preparation ae 
process is usually indicated by the chloroplast elongatit 
bending around the nucleus in the form of a thick cr 


1899 ] SPORE-MOTHER-CELL OF ANTHOCEROS 93 


The phenomenon of division is one of simple constriction in the 
middle region and final separation of the halves as the furrow 
deepens around the chloroplast. It is very plain that the line of 
fission is determined by the furrow and not from the interior of 
the chloroplast. Therefore if any particular region of the 
protoplasm is actively concerned in this process one would 
naturally first seek the agent in the peculiar film bounding the 
chloroplast. However there is little in the structure of this 
region to indicate such important activities. 

After the first division of the chloroplast the two portions 
usually separate and pass to opposite ends of the elongated spore- 
mother-cell ( fig. 6). Each has a structure quite similar to the 
parent chloroplast except that the starch grains become even 
more numerous and prominent and the fine strands and films of 
protoplasm separating the starch grains are less clearly defined. 
oon after this first division of the chloroplast there takes place 
that peculiar condition of the nucleus termed synapsis, but this 
phenomenon had best be considered in a later portion of the 
Paper that deals with nuclear activities. There is a well defined 
period when the spore-mother-cell has only two chloroplasts, and 
no longitudinal section of the sporogonium will fail to exhibit a 
fegion in which all of the cells are in this condition. As a spore- 
mother-cell grows older, and increases still further in size, indica- 
tions of the approaching second division of the chloroplast 
become manifest. 

The second division of the chloroplast repeats exactly the 
Phenomenon of the first division. As a rule both structures 
“oncerned are active at the same time, but one may be in a more 
Pe ae of fission than the other. The proportions of the 
OE eae at this time are more nearly equal (As: 7) 
portion tis it aia condition, z. ¢., the cell is| broader in as 
tributed ou Sati As a result the four chioroprast are dis- 
ive the centrally placed nucleus in such a pane 
divisicnaot a the greater part of one of the four tetrahedra 
Fation: of ty 1€ spore-mother-cell that are present after she sepa- 

€ cell contents to form the spores. Sometimes the 


94 BOTANICAL GAZETTE [avers 


second division of the chloroplast follows so closely after th 
first that the final products are arranged for a short time inthe 
form of a bow around the nucleus (fig. 9). Such conditions 
are however only transitory and result from the elongated form 
of such cells. 

The structure of the four daughter chloroplasts exhibits this 
difference from that of the mother structure shown in figs. 44 
The protoplasmic material that formerly filled the spaces betwee 
the starch grains as a spongy network of strands and films nor 
appears to be entirely absent or present only in a much reduce! 
form. The chloroplast has the structure of a vesicle filled with 
starch grains. Itis interesting to contrast this condition with the 
conditions exhibited in the youngest spore-mother-cells. Ther 
the bulk of the chloroplast was made of protoplasm and the fer 
starch grains were insignificant. But in the oldest spore-mothet 
cells the chloroplast has become one great storage vesicle 0 
starch, conspicuous by the absence of that protoplasmic differet 
tiation which is so characteristic of the chloroplast whereve! 
found, 

The problem of the division of the chloroplast and the dis 
tribution of the resultant daughter plastids through the cell 8 
full of interest. Does the development of the furrow and the 
final fission of the chloroplast result from the activity o the 
protoplasm in or around the structure? It is conceivable 
the division has its cause merely in the advantage that mgr 
come from a more even distribution of the chlorophyll throug? 
the cytoplasm. Thus it is obvious that the arrangement show 
in fig. 5 is very unsymmetrical. Perhaps a strain may be exerted 
upon a chloroplast bent around the nucleus in such a mane 
and finally result in its fission, and then lead to a redistributio® 
of the cell contents to bring about a certain balance in thee 
The difficulty of this view as applied to Anthoceros lies 10 * 
- fact that the number of chloroplasts is fixed at four, corresp : 
ing exactly to the number of spores. How could a nume 
coincidence so important in phylogeny be left to the play o 
forces merely acting for symmetry and the advantage : 


1899] SPORE-MOTHER-CELL OF ANTHOCEROS 95 


distribution of chlorophyll throughout the cell? But if we 
assign the cause of the division to the protoplasm in or around 
the chloroplast there arise complications very difficult to explain. 
The four chloroplasts are in the cell long before the nucleus 
divides. Can it be supposed that cytoplasm would be intrusted 
with so important a task as the preparation of a chloroplast for 
each of the four nuclei that are later to preside over the spores 
before there is any indication that such nuclear division is to 
take place? The process reverses what would appear to be the 
natural course of events; 7. ¢., one would suppose that the divi- 
sion of the nucleus would determine the position of the four 
Spores and that the cell contents would arrange themselves later 
with reference to the nuclei. Perhaps this really is the fact, and it 
may be merely chance that each of the four nuclei finds a single 
chloroplast to accompany it in the spore, but such coincidence 
of numbers would be very curious. A somewhat similar prob- 
lem is presented by the oospore of Coleochaete, where, according 
‘o Oltmanns, eight chloroplasts are formed before the nucleus 
divides, and then as cell division progresses the nuclei are dis- 
tributed symmetrically until each becomes associated with one 
chloroplast. 

The structure and behavior of the chloroplast suggests some 
extremely interesting lines of research. Is there such a sub- 
mae as plastidplasm, a particular form of protoplasm with 
morphological characters that may distinguish it from tropho- 
Hoon de centrospheres, and other differentiated struc- 
siti cg = If there is a plastidplasm, what form does it 
Re cea e ee of ontogeny, when chlorophyll and other 
eh a e i sent? Is the plastid a permanent organ of the 
region of ce aps generally supposed As ae plastid is a 
aah atecaieg where the pigment is gathered, and 

y ‘ne seat of metabolic activity, it is possible that 


these : : ' 
even eis might produce its form and structure. It is 

nceivable that the di ioe : i 
tid may not lie 
iD peculj ifferentiation of a plas y 


effects , aig of protoplasm itself, but represent the outward 
ov the metabolic phenomena concerned with its pigment. 


i 


96 BOTANICAL GAZETTE [AUGUST 


It appears to the writer that plant cytology has a very important 
field open for investigation dealing with the structure of the 
chromatophore and its place in ontogeny. 

We pass now to the second part of the paper, which is to 
consider the behavior of the nucleus in sporogenesis. The 
structure of the spore-mother-cell and process of nuclear division 
in Anthoceros was first described by Strasburger (’8o, p. 162). 
The following account entirely supports the essentials of those 
observations, but the present investigation attempts a molt 
detailed examination of nuclear activities, involving the prob 
lems of spindle formation, synapsis, the succession of mucleat 
divisions, and formation of the walls between the spores. 

The fact has already been stated that eight chromosomes at : 
present in the sporophyte, and consequently enter the nucleus 
of the spore-mother-cell. This nucleus in a resting condition § 
very similar to the nuclei of the archesporium and_ shows vel) 
little structural differentiation (fig. 3). It is small and its linin 
network is very inconspicuous, but the nucleolus is prominent 

Coincident with the appearance and increase in size of the | 
chloroplast the nucleus enlarges (figs. 4, 5, 6) and the threads ol 
linin become very prominent. They are so exceedingly small, 
however, that it was not possible to determine accurately the | 
structure, even with a Zeiss apochromatic immersion lens under 
the magnification of 2250 diameters. Dark-staining minute 
bodies along the linin thread are presumably granules of chro- 
matin, but their distribution could not be ascertained with or 
tainty, nor was it possible to follow the convolutions of - 
Spirem. 4 


1899] SPORE-MOTHER-CELL OF ANTHOCEROS 97 


in fig. 7. Synapsis in Anthoceros seems to occur after the first 
division of the chloroplast, and it is very apt to persist through 
to the second division. The contracted threads of linin then 
gradually extend outward through the nucleus (fig. 8), and 
finally a spirem condition is attained similar in all outward 
appearances to that present in the nucleus before synapsis. 

The nucleolus after synapsis is often found somewhat frag- 
mented, as is shown in fig. zo. Whether or not the nucleolus 
partly dissolves and contributes material to the linin thread, as 
has been suggested for certain forms (e. g., Lilium), could not be 
determined for Anthoceros. 

Synapsis in Anthoceros does not appear to the writer to be 
an artifact, and in this opinion he agrees with the views 
*xpressed by a number of investigators who have studied this 
phenomenon in other types. Particularly favorable opportunities 
we presented for the solution of this problem in Anthoceros, 
Bese one may subject at one time all conditions of the spore- 
nother-cells to the same fixing fluid. Under these conditions 
*ynapsis always appears in that particular region of the sporo- 
“a s cells in the condition just before and during 
basis Sesion ot the chloroplast. No nuclei were ever 
inte _ Synapsis in other parts of the sporogonium, and 
disturbed “ala and older cells immediately adjoining the 
thread. oe presented the typical fully expanded spirem 
ing uid shit - - these cells had been bathed in the same fix- 
of the slides identical treatment in the preparation 

€ nucle; a f course - may be claimed that the contents of 
synapsis pecs he subject to shrinkage at the time when 
of fixing iis. es : ene be understood that ae a variety 
chrom-acetic ae lea a Pico acetic, corrosive sublimate, 
nately the niall si erkel’s, all gave identical results. hae es 
4 detailed stud ey a ie nuclear elements in Anthoceros se 
and led to 7 eae this sateresting proGess extremely difficult, 
We have ee. conclusions as to the meaning of synapsis. 
*Pore-mother-cel] hat events take place during the growth of the 
in the following order: first, an increase in 


98 BOTANICAL GAZETTE [ AUGUST 
size of the nucleus, the assumption of the spirem condition, and 
the first division of the chloroplast; second, sudden synapsis at 
the time when there are two chloroplasts in the cell, and often 
continuing until after the second division of these structures; 
third, gradual emergence of the nucleus from synapsis. These 
changes present the spore-mother-cell ready for nuclear division 
with the following arrangement of the cell contents. The fou 
chloroplasts are distributed symmetrically, with the nucleus it 
the center of the cell. In this condition the cell remains fora 
period somewhat longer than that of the synapsis, and then pre 
pares for the first division of the nucleus. 

It is difficult to recognize the earliest indications that the 
nucleus is approaching prophase of division. A somewhat later 
stage is very conspicuous, when the outline of the nucleus 
becomes angular and a mat of delicate threads surrounds the 
structure (fig. 72). There is a period, however, previous to this 
condition, when the nuclear membrane appears much less firm 
and somewhat irregular in outline. This structure follows the 
Spirem stage such as is shown in fig. To, and_ precedes the 
unmistakable prophase conditions illustrated by figs. 12, 13: - 
appearance is given in fig. zz, and the following peculiamti® 
should be noted, viz., an irregularly outlined nuclear membfaté 
a faint linin network, and fragmentary nucleolus. But perhaps 
the most important characteristics appear outside of the nucleat 
membrane in the cytoplasm as a delicate web of fibrils closely 
applied to the nucleus. These fibers are so delicate as almost” 
defy the reproduction that has been attempted in fig. If. They 
presage the development of the spindle. ; 


In the prophases the nucleus may exhibit some extreme 
varied appearances. The form is very irregular, takin | | 
curious wavy angles. Outside of the nucleus there is 21" | 


of protoplasm manifestly very different in structure from 
neighboring cytoplasm» In overstained preparations this regime 
Ss very conspicuous, forming a sort of zone around the nucle 
but the details of its structure can only be made 0 


| lla 
favorable preparations. Then it is seen to possess 4 fibeilif 


ut in vel - 


Ms 
‘gid é 


1899] SPORE-MOTHER-CELL OF ANTHOCEROS 99 


structure. The delicate threads lie all around the nucleus, but 
they accumulate conspicuously at the pointed projections (jigs. 
12, 13). 

At this time one is apt to find the linin network contracted 
away from the nuclear membrane and gathered in the form of a 
confused tangle in the central region somewhere near the 
nucleolus (fig. 73). The chromatic material becomes more 
prominent and from the ill-defined mass in the interior there 
emerge four deeply staining chromosomes (figs. 12,74). When 
the chromosomes are fully differentiated one may expect to find 
one or two of the angles of the nucleus more prominent than 
the others, and the fibrils around them taking on the appearance 
of spindle fibers (figs. 73, 74). The process of spindle differ- 
entiation is a gradual one, and only in the later stages is it pos- 
sible to feel sure of the direction that the axis will assume. I, 
is seldom that the two poles appear from the beginning so nearly 
°pposite one another as to have a common axis. It is more 
usual for the spindle to be somewhat bent at first, as is shown in 


fg. 14. However, ultimately the two poles arrange themselves 


‘o form a symmetrical spindle of the form illustrated by figs. 75, 
16, 17, 78. 

Me fully developed spindle has an interesting structure, with 
several clearly marked features well shown by fig. 8. The poles 
. oe periiaps slightly convex ; they are never pointed. 
Brest es are conspicuous around the nuclear plate ( figs. 16- 

), but what relation these bear to the chromosomes could not 
be determined. 

. — : the condition reported by Farmer (94 and ’05) for 
Poskiake the Hepaticae, notably Pellia, Pallavicinia, and 
Whether oo it became a very important problem to determine 
the — centrospheres were ever present at the poles of 
care for te The writer searched his preparations with great 
do not ae structures, but came to the conclusion that es 
the Spindle ¢ ey Anthoceros, Sometimes the cytoplasm aroun 
shown in fi Ontains large deeply staining granules, such as are 
© 47, and these may occupy positions near the ends 


100 BOTANICAL GAZETTE | AUGUST 


of the spindles, but extensive observations would convince 


anyone that they have nothing whatever to do with centrosomes 7 


The protoplasm in the vicinity of the pole is often very dense 
in structure, containing no granules and few vacuoles. It fre 
quently presents the appearance shown in jig. 16, but it never 
exhibited indications of that differentiation expected of 3 
centrosphere. 

It will be apparent to the reader from this account that the 
spindle of Anthoceros during the first division follows a history 


closely parallel to that described for homologous cells in Lilium — 
by Mottier ('97), Hemerocallis by Juel (’97), Equisetum by 


Osterhout (’97), and Cobea by Lawson (’98). That is to s# 
the spindle is organized by numerous delicate fibrils of proto 
plasm that develop conspicuously during prophase in the cyto 
plasm around the nuclear membrane. The fibrils are at first 
somewhat irregularly distributed, but finally become arranged It 
the form characteristic of the respective spindles. 


Strasburger in the third edition of the Lehrbuch der Botany 


1898, p. 67, has introduced the term filarplasm to be applie 


to protoplasm having the form and activities above described 


and so clearly established by the researches of his students 
during the past three years. The writer understands that filar- 


plasm is supposed to be made up of the substance designated by 


the older term kinoplasm. However, filarplasm has morphologt | 
cal characters, as indicated by its name, and these are the three 
or fibril-like structures. The term therefore expresses admirably 


the facts of morphology without implying or assigning phys” 


logical activities to the substance. 
It will be very gratifying if future investigation should 


appears to extend the range of mitoses associated with filarp! 


into a group of plants much lower than the lowest previ : 


that 


reported (pteridophytes by Osterhout). It is also significa 


it should be a class containing one large order, the Junge 


; : eked 
niacez, where according to Farmer centrospheres are very ses 


estab 
lish filarplasm as an element in the cells of higher plants diffe 


entiated from other forms of protoplasm. This investigatio? 


& 


Pe ea Se 


1899] SPORE-MOTHER-CELL OF ANTHOCEROS 101 


features of the mitotic figures. There should be types of plants 
that will allow us to form some idea of the relationship between 
filarplasm and the asters and centrospheres present in certain 
thallophytes, in other words solve the problems of their places 
in phylogeny. Perhaps the future study of Hepatice will throw 
some light on this question. 

The two sets of daughter chromosomes that result from the 
division at metaphase pass very quickly from the nuclear plate to 
the poles of the spindle. Here they may frequently be found 
clustered together ina region of dense protoplasm that surrounds 
the group, as is shown in fig. ro. The development of a nuclear 
membrane around the chromosomes finally organizes the two 
daughter nuclei which are connected for a short time by the 
spindle fibers. These latter structures gradually fade away, 
beginning at the poles (fig. 20), and finally entirely disappear. 
The daughter nuclei then lie in undifferentiated cytoplasm usually — 
occupying positions between the chloroplasts somewhat as is indi- 
cated in fig. ar. 

. iy daughter nuclei following the first division pass into a 
eras developed resting condition. The chromosomes break up 
ss 3 number of chromatin granules that become distributed over 
rh network. A prominent nucleolus appears. In sections 
— one may always find spore-mother-cells con- 
ne elas nuclei, although this condition is quickly replaced by 
ceding second division. 
en ih mitosis in the spore-mother-cell involves both 
a but the two nuclear figures are developed 
by Ges ci y of one another. The two spindles are never united 
re ers. Sometimes the two spindles will lie almost side 
Y side, but usually they are placed igh les to each other. 
Frequently oe. y are placed at right ang - es ; 
1h OF bac... €ction of a spore-mother-cell will give a po ar . 
view of the a and the neighboring section a longitudinal 
‘js tg . se An instance of this character has been chosen 
in fig. > € principal features of their structure and is shown 
# By comparing fig. 24 with illustrations of the first 
- 16-18) it will be seen that the spindles of the 


102 BOTANICAL GAZETTE [AUGUST 


second mitosis are much smaller. However they exhibit a 
essentially similar structure, having flattened poles without cen- 
trospheres. Several prominent fibrils make up the center of the 
spindle which is bordered by an ill-defined set of mantle fibers, 
The polar view of a spindle shown at the left of fig. 24 demon 
strates that four chromosomes are present at the nuclear plate, 
the same number that appears during the first mitosis. Prophast 
and anaphase conditions of the second mitosis are found otly 
with great difficulty and are very unsatisfactory for study becast 
of the small size of the elements involved. 

The four nuclei that result from the second mitosis associate 
themselves each with one chloroplast, and with these become dis- 
tributed symmetrically through the cell, so that the protoplast 
naturally segregates into four regions representing what are later 
to become the tetrahedral division of the spore-mother-cell (figs 
26, 27). The spindle fibers disappear completely. 

The problem of the splitting of the chromosomes engage! 
the writer’s attention, but it must be plain that Anthoceros * 


not a favorable subject for the study of this process. The neatly. 


spherical form of the chromosomes offers immense difficultits ! 
orientation. Farmer (’95) reported some peculiar conditions " 
the forms studied by him, which he considers as illustrations 
the ‘‘heterotype”’ division described by Flemming. They res 


from the habit that the chromosomes have of doubling on the™ | 
chromo | 


selves and then being pulled apart as V-shaped daughter 
somes. The figures are very complex, but Farmer ass 
that the division is really longitudinal and not transverse, so that 


it cannot be interpreted as qualitative. In Anthoceros all eviden® 


ures & 


the daughter chro ;, eight in number, arranged in four pa : 
Such a Stage is shown in jig. 23, and the grouping certainly 1” 


cates that each of the original four chromosomes has divi 


tudinally into halves. It is not at all unusual to find HE) | 


sets of daughter chromosomes placed as in fig. 22, which ™ : 


ne 


j 


} 


: 
a 
a 
: 
4 
py 
a 
a 


1899] SPORE-MOTHER-CELL OF ANTHOCEROS 103 


suggest transverse fission, but the writer is not willing to accept 
this evidence, as such an arrangement would naturally appear 
when the daughter chromosomes pass away from the nuclear 
plate to the poles of the spindle. 

We have now finished our account of nuclear activities in 
the spore-mother-cell, but there remains for consideration the 
description of the manner in which the cell contents are divided 
to form the spores. After the two mitoses each of the four 
nuclei lies in a region of dense protoplasm at the side of a chlo- 
roplast towards the interior of the cell (fig. 25). The bulk of 
the protoplasm is therefore collected into four masses somewhat 
apart from one another but connected by very numerous delicate 
filaments. The cross filaments are very conspicuous but irregu- 
lar in their arrangement, frequently anastomosing. As shown in 
hig. 25 they are not confined to the vicinity of the nuclei but con- 
nect all portions of the separated regions of protoplasm. They 
do not resemble spindle fibers, being much thicker, but have 
instead the appearance of strands of cytoplasm. Following 
the condition shown in jig. 25 one may find stages similar to 
fg. 26. It is plain that the protoplasmic strands have spread 
sideways and fused with one another so that there is now present 
a film of protoplasm between tetrahedral regions of the spore- 
Mother-cell. This film marks exactly the position that is 
finally to be occupied by cell walls when the spores are fully 
organized. 


spe Petiliarity of the process just described lies chiefly in 
is the ms to be its entire independence of spindle fibers. It 
iiecatie accepted view that the walls crossing spore- 
that GH ee pollen-mother-cells are derived from cell plates 
The writ ee the fusion of spindle fibers after anaphase. 
tion oie. thinks that no investigator has described a condi- 
the Be. to Anthoceros. The present studies indicate a 

€s of the two successive mitoses completely disap- 


pear, ets 

the w This 1S certainly a very difficult point to determine, but 
r ; t 

c iter feels confident that the anastomosing strands which 

Onnec ‘ 


t the four Masses of protoplasm in the spore-mother-cell, 


104 BOTANICAL GAZETTE [avawst 


as has been shown in figs. 25, 26, are formed entirely independ: 
ently of spindie fibers. How then are the walls formed? 

It is certain that the film of protoplasm indicated in jig. 2 
thickens and finally gives place toa straight wall, at first delicate 
( fig. 27) but gradually becoming firmer until its cellulose nature 
is unmistakable. 

The phenomenon is typical of one of the processes recently 
discussed by Strasburger (’98), in which a cell wall is formedin 
the interior of the protoplasm. It appears as if the film of proto 
plasm exhibits the activities present in the ‘ Hautschicht” when 
it lays down or increases the thickness of a cell wall in the mat- 
ner known as apposition, involving, at least in part, the change 
of its own substance into cellulose. ; 

With the separation of its contents the spore-mother-cell as 
a unit ceases to exist,and a new set of activities begins that maj 
very properly be reserved for discussion in another paper. Some — 
interesting events take place in the spore as it ripens, but technical 
difficulties interfere greatly with their elucidation. : 

The peculiar fact that the sporogonium of Anthoceros 
presents spore-mother-cells in all stages of development makes 
it possible to contrast the times occupied by the various changes 
One cannot establish the actual duration of any process, bit . 
within certain limits it is possible to determine the relative periods 
of each event. It must be assumed that the rate of gre 


the spore-mother-cell to the period of the first division of 
chloroplast. The division of the chloroplast occupies 6- 
The nucleus is in synapsis 12-20 units. For 30-50 units thee 
has two chloroplasts. It takes about 2 5 units after synapsis © 
Produce conditions favorable for mitosis, The first miter 
occupies 1-3 units, and the two daughter nuclei rest for : 


1899] SPORE-MOTHER-CELL OF ANTHOCEROS 105 


units before the second mitosis which is more rapid than the first. 
There is finally a period of 3-6 units before the spores are 
definitely organized by the partition walls across the spore- 
mother-cell. 

The results presented by this investigation indicate that the 
Hepatice are likely to furnish some very interesting material 
for future researches in cytology. From superficial examination 
it appears probable that the Ricciacee and Marchantiacee in 
general present mitoses similar to Anthoceros, but we may hope 
that some forms exist that will harmonize the peculiarities 
described for the Jungermanniacez with the conditions found in 
Anthoceros. 

EXPERIMENTAL TECHNIQUE. 


As stated in the beginning of the paper the writer experi- 
mented with a number of fixing agents to determine as precisely 
%s possible their merits or faults. Of these Flemming’s formula 
“esignated “weak ” gave decidedly better results than any other. 
Anthoceros, and presumably for other types, the chief test of 
a fixing fluid is its effect upon the achromatic parts of a nuclear 
figure. Chromosomes are the least difficult of all the nuclear 
— to preserve. Preparations will not infrequently present 
a — of nuclear plates when the spindles are manifestly 
wee dition. The following is a brief statement of the 
cell. Several fluids upon nuclear figures in the spore-mother- 
oo acid fixes filarplasm but the safranin stains dif- 
Spindle a soa gentian violet does not hold well in the 
days in ie i sections fastened to the slide be left several 
Sentian violet lemming the staining qualities with safranin and 

are much improved although, it is doubtful if 


they ¢ 
a made as good as those presented by Flemming fixed 


Merkel’s fluid 
Platinum chlorid 
Petiods (36 hou 
badly fixed 


(1 per cent. chromic acid 12°, I per cent. 
2°, water 72°) even when used for long 
ts) is thoroughly unsatisfactory. The spindles 


106 BOTANICAL GAZETTE [AuGUST 


Boveri's picro-acetic acid gives beautifully bleached tissue, 
but achromatic regions are not clearly differentiated although 
chromatic elements stain well. 

Sublimate-acetic (5 per cent. glacial acetic acid in saturated 
solution of corrosive sublimate) is not good. Nuclear membranes 
and filarplasm-are very poorly preserved. 

Hermann’s fluid is very much like Flemming’s in its effects 
and is thoroughly satisfactory. 

The osmic acid of the Flemming’s and Hermann’s mixtures 
appears to give them certain advantages over all other fluids. 
Although they may not kill and preserve tissue better than some 
other agents, as for example chrom-acetic acid, certain stains, 
safranin and gentian violet, differentiate all structures of the cell 
very much better when they have been used. 


SUMMARY. 


_ The number of chromsomes is eight for the sporophyte and 
four in the gametophyte. 

The chloroplast appears rather suddenly in the spore-mom 
cell as a differentiated region of the protoplasm, containing 
several starch grains. When fully developed it has a honeycom . 
structure, each cavity being occupied by a grain of starch. . 

The division of the chloroplast is one of simple fissi0% 
apparently through forces acting outside of the structure 
and perhaps concerned with the film of protoplasm that tee 
rounds it. 

Synapsis occurs in the nucleus soon after the first division 
the chloroplast. It is not an artifact. — 

- The second division of the chloroplast presents the spore 
mother-cell ready for the division of the nucleus. | The fo : 
chloroplasts, hardly more than vesicles filled with starch gas 
are arranged symmetrically in the cytoplasm. with the nuclee 
in the center of the cell. ; 

The resting nucleus has a nucleolus and a spirem threa 
however, is so small that details of structure could not 
mined. 


thet: 


d, which 
be dete 


1899] SPORE-MOTHER-CELL OF ANTHOCEROS 107 


Prophase conditions of the nucleus show the presence of a 
mesh of delicate fibrils(filarplasm) around the structure. The 
nuclear membrane is at first irregularly angular, but finally two 
poles of a spindle are differentiated. 

The metaphase presents a spindle with flattened poles, 
entirely lacking bodies that might be interpreted as centrospheres 
or centrosomes. 

- There is a period of rest after the first mitosis when each 
daughter nucleus has a nucleolus and spirem thread. The two 
mitoses are, therefore, successive. 

The second mitosis presents structural features quite identi- 
cal with the first. 

- Thechromosomes are four in number in each mitosis. They 
appear to split longitudinally. 

All traces of the spindles become lost soon after each mitosis. 
~ The fully mature spore-mother-cell presents four chloroplasts, 
each with a single nucleus on the interior side. | The protoplasm 
segregates in these four regions of the cell, leaving spaces that 
are crossed by many anastomosing strands of cytoplasm. These 
strands cannot be traced from spindle fibers and appear to have 
He connection with filarplasm. 

The walls separating the cell contents into four spores are 
a. from films of protoplasm that appear between the chlo- 
2 asts with their respective nuclei. The films are formed by 
Cie of strands of cytoplasm that cross the spaces 

€ four regions of cell contents. 

The UNIVERSITY oF CHICAGo. 


LITERATURE CITED. 


95) Mosses and ferns. 
~~~ (98) On th 
€ structure and development of Dendroceros. Jour. Linn. 


Campgery ( 


33: 467 
FARMER and Re 
Phylla N 
FARMER (; 


EVES(’94) On the occurrence of centrospheres in Pe//ia epi- 

fes. Ann. Bot. 8: 210. 

9) Studies in Hepatice : On Pallavicinia decipiens Mitten. Ann. 

— (os) 5; | 
Pore formation and karyokinesis in Hepaticae. Ann. Bot. 9 : 363.- 


108 BOTANICAL GAZETTE [avcust 


(’95) On spore formation and nuclear division in Hepatice, Ann. 
Bot. 9 :46 

JuEL(’97) Die Kerntheilung in den Pollenmutterzellen von Hemerocallis fulta 
und die bei denselben auftretenden Unregelmassigkeiten. Jahrb, f. wiss. 
Hot, 90+ Fi: 

LAWSON (’98) Some observations on the development of the karyokinetic 
spindle in the pollen-mother-cells of Cob@a scandens Cav. Proc. Calif. 
Acad. Sci. Botany 1: 169. 

Mortikr (’97) Beitrage zur Kenntniss der Kerntheilung in den Pollenmutter 
zellen einiger Dikotylen und Monokotylen. Jahrb. f. wiss. Bot. 30: 

OsTERHOUT (’97) Ueber Entstehung der karyokinetischen spindle bei Equ- 
setum. Jahrb. f. wiss. Bot. 30: 5. 

STRASBURGER (’80) Zellbildung und Zelltheilung. 

(’988) Lehrbuch der Botanik, dritte Auflage. 

(’98b) Die pflanzlichen Zellhaute. Jahrb. f. wiss. Bot. 31: — 


EXPLANATION OF PLATES IX AND X. 


All preparations were studied under a Zeiss oil immersion 2” ape 1,30 


with compensation-oculars. Figures were sketched with an Abbé camera | 


at the following magnifications, Rigs. 3-7, 9, 10, 12, 10-21, 24-2]) magnified 
1000 diameters; 7, 2,8, 77, 73, 74, 15. 22, 1500 diameters and fig. 23; 2250 
diameters. 

All preparations except that shown in fig. 2 from material fixed! 
Flemming, sectioned Sy thick and stained on the slide; figs. 4) 45) 
and 23 from slides stained with iron-alum-haematoxylin after the mete” 
Haidenhain ; all other figures from preparations stained with Flemmings 
triple stain, safranin, gentian violet, and orange G. 


PEATE LX. 


Fic. 1. Nuclear figures from archesporium, just after the spli 


: : each 
chromosomes whose number is large, apparently about sixteen, eight for nat 


daughter nucleus. 
Fic. 2. Nuclear figure from antheridium, four chromosomes. 
FIG. 3. Spore-mother-cell, early condition of chloroplast with 
four grains of starch, ; 
1G. 4. Spore-mother-cell in more advanced condition than Ig J 
network more prominent, chloroplast elongating preparatory to division + °" 
grains prominent. 
G. 5. First division of chloroplast. 
Fic. 6. Two chloroplasts, linin network prominent. 
FIG. 7. Synapsis, two chloroplasts. 
Fic. 8. Nucleus emerging from synapsis. 


wt 


n weak 


tting of the 


three OT 


BOTANICAL GAZETTE, XXVIII. 


pee . 


Laie 


ak tain 
EAP ue 


es 


cy 


s 


fh Ane 


“ 
pees 


ROS. 


. 
. 


. DAVIS on ANTHOCE 


ae tee Pen monn Ne ge ee ag OL ea ee er a a 
oe 


pied ee ka AM es 5 ok al aa 


BOTANICAL GAZETTE, XX VII. 


DAVIS on ANTHOCEROS. 


PLATE &. 


re a a a ee pe ee a 


1899] SPORE-MOTHER-CELL OF ANTHOCEROS 109 


Fic. 9. Second division of the chloroplasts. 

Fic. 10. Spore-mother-cell at maturity, four chloroplasts. 

Fig. 11. First indication of approaching mitosis, accumulations of filar- 
plasm around the nucleus whose membrane appears less clearly defined. 

Fig. 12. Prophase, filarplasm well differentiated. 

Fig. 13. Prophase, one pole of spindle developed. 

Fic. 14. Late prophase, the two poles of the spindle differentiated, four 
chromosomes assembled in the equatorial region. 


PLATE X. 

Fic. 15. Early metaphase, four chromosomes at nuclear plate. 

FiG. 16. Metaphase, strongly developed mantle fibers. 

Fic. 17. Metaphase, granular cytoplasm around the nuclear figure. 

Fig. 18. Late metaphase, after splitting of the chromosomes, very clearly 
defined spindle. 

Fig. tg. Anaphase, spindle still clearly defined. 

FiG. 20. Telophase, trace of spindle in equatorial region. 
Fig. 21. Spore-mother-cell, daughter nuclei after first mitosis in rest- 
‘ng condition, four chloroplasts. 

FiG. 22. Nuclear plate at first mitosis, side view. 
Shes os Nuclear plate of second mitosis viewed from pole of spindle, 

cating longitudinal splitting of the chromosomes. 
ae spies sections of same spore-mother-cell during second 
the 7: mis on plate of one spindle with four chromosomes viewed from 

: ; Side view of the other spindle. 
ee Center. of spore-mother-cell showing three nuclei, cytoplasm 
th sated around the nuclei and chloroplasts, delicate strands connecting 

€ regions, 
in riick indication of the position of the walls that are to be formed 
‘o form a delic er-cell ; the coalescence of connecting strands of cytoplasm 

ate film. 


1G. 27. Final separati 
mother-cel], paration of the protoplasmic contents of the spore 


THE STRUCTURE AND DEVELOPMENT OF CRYPIO 
MITRIUM TENERUM. 

LE Roy ABRAMS. 

(WITH SIX FIGURES) 


THE genus Cryptomitrium, represented by the single specie 
C. tenerum (Hook.) Austin, has not yet been thoroughly studied, 
and as the plant has been collected in several localities m# 
Stanford University,.at the suggestion of Dr. Campbell a stud} 
of its structure and development was undertaken, in order, 
possible, to determine its relationship to the other Marchantiaceé | 

The first collections were made late in the spring of 1898 
at which time the plants were mature, and the spores alos 
ripe. Some of these plants were placed in alcohol, whi 
others were allowed to become dry, and the earth upon ™ 
they were growing was kept until the following Sept agi: 
when the work upon the plant was taken up. A consid 
number of the mature sporogonial receptacles were also put ¥ 
dry, inorder to study the germination of the spores. 

Some of the pieces of earth with the dried specimens 
them were thoroughly soaked, and then kept well m i 
Within a day or two the tips of the apparently dried-Up 
became green and fresh, and in about two weeks the am 
began to form. All of the material for study was © 
in this way, until after the rains came. Then comsit 
material was collected from out of doors, where it d 
much faster and was healthier than that grown in the 
atory. 


0 


THE THALLUS. . 

Cryptomitrium, like most other Marchantiace®, * 
dichotomously branched thallus, which in this spect 
thin and delicate. The smooth glossy appearance, P. 
one can easily distinguish the sterile plants from * 
IIo 


1899] DEVELOPMENT OF CRYPTOMITRIUM III 


Californica Hampe, a species almost always associated with it, 
is due for the most part to the minute stomata. 

These stomata are surrounded by eight, occasionally seven, 
very symmetrically arranged guard cells (a, fig. z), and not by 
five or six, as stated by Stephani. Each opens into a well- 
developed air chamber (4, fig. 7), 
the boundaries or walls of which 
can be seen easily with a hand 
lens, forming a fine network 
under the epidermis. 

These air chambers are much 
the same as in Fimbdriaria Calt- 
fornica. They are distributed 
irregularly throughout the green 
tissue. Only a single layer of 
cells Separates them, and often one 

> arty is connected with another. 3 

_ Their development begins a little 

; ao back from the apical ht 

: than in the above mentioned 

ccaieb described by Campbell.? Fic. 1.— Stomata of the thallus. @, as 

med general appearance and _ seen from the surface. X600. 4, trans- 

external characters of the ventral. Vem section. X 600. 

a ab been quite thoroughly and accurately described 
ephani (Joc. cit.), and as their development does not differ 

of other allied genera it need not be repeated here. 

ht be expected, both kinds of root hairs, those with 


nn iii 


‘ 
oS 


ap Feceptacle. The development and composition 
oil bodies found in the Hepatic have been thoroughly 


ae ae a 58-60. 1892. : 
‘ ELL, D. H.: Mosses and Ferns, p. 48. 1895. 


od m : Die Oelkérper der Lebermoose. Flora 4a: 1874. 


112 BOTANICAL GAZETTE [aveust 


THE SEXUAL ORGANS. 


Cryptomitrium tenerum is monoecious. The antheridia forma 
single row just back of the sporogonial receptacle. They 
are sunk deep in the thallus, and each one is marked on the 
surface by a small conical ostiolum. These ostiola are vey 
inconspicuous, however, and their presence can scarcely be | 
detected with a hand lens. : 

The antheridia are developed before the female receptacle, 
and in Fimbriaria and other allied genera, they are formed on 
dorsal side just back of the apical cell (a, fig. 2). The prim 


a 


Fic. 2.— Antheridia. a, longitudinal section of apex of thallus with 
antheridia ; x, apical cell. X 600. 4, c, d, successive stages of youns 
% 600. ¢, full-grown antheridium. < 480. 


antheridial cell, when it can first be distinguished from so 
cells around it, is a little larger and stains. more deeply 
these cells. The first division (a, fig. 2) 1s transverse 


1899] DEVELOPMENT OF CRYPTOMITRIUM LES 


divides the primary cell into the stalk or pedicel cell and the 
antheridial’ cell proper. The antheridial cell thus formed is 
then divided into three cells (a, fig. 2) by two transverse divi- 
sions. The next two divisions are longitudinal medial and are 
at right angles to each other, so that each of the three original 
cells is divided into four. In many cases, however, only the 
two lower cells are divided in this way (4, fig. 2), the top cell 
remaining undivided for a longer time. The four cells formed 
from the central cell by these longitudinal divisions are each 
again divided longitudinally, thus separating the sperm cells 
_ from those that go to make up a portion of the antheridial wall 
 (¢,fig. 2). The upper and lower cells take no part in forming 
the sperm cells, but form, respectively, the upper and lower por- 
_ tions of the antheridial wall. The remainder of the development 
, does not differ materially from that of Fimbriaria Californica as 
| described by Campbell.t| The top of the antheridial wall is not 
prolonged, however, as in that species, but is only a single row 
, of cells, as in Marchantia. In fact, the full-grown antheridium 
; (¢, fig. 2) resembles very closely that of Marchantia. 
| oe Sporogonial receptacle, or carpocephalum, is of Leit- 
— «&eb's “Composite’”’ types The apical cell of the thallus forms 
the growing point of the receptacle, but instead of remaining a 
single cell it divides into two cells. Each of these again 
ne : like manner. Finally one of the four cells thus 
me ivides into two, so that there are five growing points. 
cases this last division does not take place, so that 
— ae Only four growing points. Five is the usual num- 
fue for I found only two specimens out of the great 
os ig sl aig examined that had only four growing points. 
: eptacles, 1S Somewhat emphasized in the half-grown 
ee Sine or, at this time, the lobes between the growing 
ol the sah ld developed than the rest, so that the undersice 
Ave Ptacle has five quite prominent projections or folds. 
“ceptacle develops, however, these disappear. 
“CAMPBELL, D, i: 
> Leitcer: Ubtersuc 


Mosses and Ferns 50, 51. 1895. 


hungen iiber die Lebermoose 6: 32. 


114 BOTANIUAL GAZETTE [avcust 


The dorsal growth of the receptacle is excessive, while the 
ventral growth is limited to a few layers of cells. Consequently 
the apical cells (a, fig. 3) lie very close to the stalk. The 
lacune or airchambers are for the most part confined toa single 
layer. They are extremely large, however, and are separated 


FiG. 3.—a, longitudinal section of sporogonial receptacle; % apical ‘a 
stoma. X 80. 4, c, longitudinal section of stoma. X 600. 4, transverse 5° 
receptacle; /, furrow of peduncle. X 80 


from one another by a single row of cells. -Each ait oe 
iS connected with the exterior by means of well-dev" 
stomata. These peculiar breathing pores are present 19 


1899] DEVELOPMENT OF CRYPTOMITRIUM 115 


of the Marchantiacee. They are almost cylindrical, and are 
composed of several rows of cells, which are formed from the 
original guard cells. These, instead of remaining single, divide 
by means of inclined walls into several cells. Generally four 
cells are cut off on the upper side (4, c, fig. 3), and two on the 
lower side of each original guard cell. 

Stephani, Joc. cit, reports that there are two furrows on the 
ventral side of the stalk or peduncle of. the receptacle, but I was 
unable to find such to be the case. In all the specimens exam- 
ined there was only one. Mr. Howe,’ also, reports only one 
furrow in the specimens examined by him. Seen in cross-section 
(d, fig. 3) it resembles very closely that of Fimbriaria and 
Duvalia as figured by Leitgeb.?. The root hairs, all of which are 
tuberculate, are found in this furrow. At the base of the recep- 
tacle these branch, one branch going to each lobe between the 
growing points. 

The archegonia, although they are on the underside of the 
receptacle, are in reality on the dorsal side, for they are formed 
actopetally just back of each apical cell. Hence there are five 
‘OWS or groups of archegonia (d, fig. 3). In each of these rows 
there are usually three or four archegonia. 

/ oh * development of the archegonium corresponds very closely 
aa ns of other Marchantiacee. The primary cell becomes 
arger than the neighboring cells, and. the cell contents 

ie meh denser, so that it stains very deeply. The first 
Ss a 's transverse. The outer cell forms the archegonium 
dis ies, the Stalk. Strasburger® states that in Marchantia 
‘t. Ms hi 1S again divided by a wall parallel to the first, and 
Ze Sse cell of the two thus formed forms the foot of the 
1. Janczewski? describes the same thing in Prezssia 

This second division does not take place in Crypto- 


Mowe, M.A.: Erythea 5: 87, 


Oy, 88. 1897. 

. MEITCER: 

ae Untersuchungen iiber die Lebermoose 6: PL. 4s AES. Js 20+ 
ASBURGER : Jahrb. f. wiss, Bot. 7: 416. 


SJax 
CZEWSKI: Bot. Zeit.— : 418, 1872. 


116 BOTANICAL GAZETTE | avGust 


mitrium (a, fig. 4). Campbell’? also states that it does not 
occur in Targionia nor in /2mbriaria Californica. The remainder 
of the development does not differ materially from that of: 
typical archegonium of any of the Marchantiacea, and it need 
not be repeated here. 


: 
; 
: 
| 


Fic. 4.—Archegonia. a, 4, c, d, ¢, f, longitudinal sections of nce ae 
*, apical cell; /, /, 7, cover cell; v, ventral canal cell; g, transverse section ° 
an archegonium about the age of e. a, b,c, X 600; d,e, X 480; 4% x e 


The cover cell, which in other forms studied” divides ? 
four cells immediately after the neck has been separated 
the venter, remains undivided for a considerably longer tims 
this species (4, c, d, /, fig. 4). Unfortunately I was " : 
determine just when this division took place. Several arch $ 
were obtained at the age of the one represented in d, fig. # 

CAMPBELL : of. cit. 52. 1895. 

“CAMPBELL : of. cit. 30. 1895. 


1899} DEVELOPMENT OF CRYPTOMITRIUM ti7 


in every case the cover cell still remained undivided. The 
stages in which the division had taken place (e, fig. ¢) were too 
old to determine with any degree of accuracy when this division 
occurred, and it is to be regretted that no intermediate stages 
were obtained. 

Gayet,” in a recent article, which has been reviewed by 
Campbell, states that the archegonia of the Hepatice have a 
distinct apical growth, the same as in the Musci. His con- 
clusions, which are contrary to those of Janczewski,’* Campbell, 
and others, are not confirmed by my own observations. While 
I did not make a very careful study of this point, I could find 
nothing that indicated an apical cell. The cover cell, which 
Gayet claims to be the apical cell, does not have the appearance 
of one. It is much smaller than the upper cells of the neck, and 
no way do these cells look as though they had been cut off 
from it. The fact that the cover cell remains undivided for a 
considerable time in this species might favor the idea of apical 
growth, were it not for the fact that the cover cell in this species 
's even smaller than in other species that have been studied 
(4, fig. 4). It looks as if it were lying dormant and not as if it 
Were an active apical cell. 

The usual nu 
Marchantiacese. 
are present, 


mber of neck canal cells is eight, as in the other 
In some cases, however (f, fig. 4), only seven 


THE SPOROPHYTE. 


ae Owing to the fact that the embryo in nearly every case lies 

eS aaa with the stalk, its development has been com- 

Marked re out. One is at once struck with the 

F805 ; id shows to the embryo of Targionia.’* 

its otiginal - ertilization ie egg cell enlarges to nearly twice 

enlarged ii es The first division is transverse and divides the 

: Nay. nto two almost equal cells (a, fig. 5). The next 
aes Sci. Nat. Bot. VIII. —:——. 1897. 

Se 8: 428-431. 1897. 

I: loc. cit, 


— SAMPBELL - %P. cit. 60, 61. 1895, 


118 BOTANICAL GAZETTE [aveust 


division is longitudinal (4, fig. 5). This is followed by another 
longitudinal division which is at right angles to it. Each of the 
eight cells thus formed is then divided into two slightly unequal 
cells by a longitudinal division (e, fig. 5). The first longitudiml 
wall is often inclined so that the top cell. (c, fig. 5) resembles 
very much a two-sided apical cell. The remaining divisions at 


Meeks Ay 12) 


Soe 
if ke pean vt 


4 
‘ 
| 


al section. X 600. 7 


Fic. 5.—Embryo. a, 4, very young stages in longitudin 300. ah 


somewhat older. X 480. d, longitudinal section of still older stage x 
transverse sections. X 480. | 


very irregular and are difficult to follow. The young go 
takes on a more and more elongated form. Finally, the 
portion almost ceases to grow, so that the embryo ae 
dumb-bell-shaped (a, fig. 6). The upper portion is t form 
archesporium, and at about this time, or even earlier (d, J 


a definite row of cells, which becomes the capsule we 
formed around the outside. Usually, the first transverse df 
marks the separation of the capsule and the foot, but 
cases (d, fig. 5) this cell remained undivided after the ie 
longitudinal divisions, so that at the base of the embry¢ 
four large cells. : 

Soon after the capsule wall is formed the archespor P 
can easily be distinguished, for their protoplasm becomes: 


1899] DEVELOPMENT OF CRYPTOMITRKIUM 119 


and both it and the cell walls, which become very gelatinous, 
stain deeply (a, 6, fig. 6). These gelatinous walls soon dissolve, 
so that the archesporial cells. are set free. Two sorts of cells 
can easily be made out at this time. The one, the spore 


fo 
@ o@ © 
Op - e, 
e 290 


L_\, a 
Caratece 
ee a 


S 
ta 


heb Ew, 
tion of me Embryo. eed longitudinal section of half-grown embryo. X 300. 4, por- 
dit owe Section showing capsule wall and archesporial cells. X 480. % 
al section of 


a nearly mature sporogonium, showing spore-mother-cells and 
*@ operculum. X 150. d, young elater. X 600. @ spore-mother-cells. 
. 


x Young elaters 


: cells, are almost spherical. Their nuclei are large and — 
(¢, fg. 6), and are surrounded by closely reticulated 

| The other, the young elater cells (d, fig. 6), are 
Their nuclei, though quite distinct, are much smaller 

of the Spore-mother-cells. 


. 


120 BOTANICAL GAZETTE [avcust 


/ The foot of the sporogonium (c, fig. 6) is not so wel 
developed as in Targionia, being not more than one thirds 
large as the capsule, which is large and globular. At matutity 
the capsule is regularly dehiscent at its apex by an operculum 
This operculum, as stated by Howe (doc. cit.) , is composed o! | 
two rows of cells (0, fig. 6), while the remainder of the capsill 
wall is, for the most part, only one cell thick. Near its base, 
however, an apparently continuous ring, composed of only & 
two rows of very small cells occurred in all the specimen 
examined, 

The ripe spores germinate very slowly. During the months 
of October, November, and December, several cultures wet 
made of spores which had ripened in the previous April, and it 
no case did they germinate until eighteen or twenty days alter 
they were sown. Their germination and manner of growth cot 
respond very closely to that of Targionia. 


SUMMARY. 

In comparing Cryptomitrium tenerum with the other Marcha 
tiacee, I found, as Stephani claimed, that it was undoubted} 
very closely related to Duvalia. I had no specimens 
Duvalia, however, and was dependent upon Leitgeb’s”™ descr 
tion. Both genera are moncecious. Both have the same minute 
stomata surrounded by seven or eight very symmetti 
arranged guard cells. Stephani states that Cryptomitrium’ 
two furrows in the peduncle, while Duvalia has only one fe 
only one furrow was present in the specimens I exalt 
that neither this difference between the genera, nor the dite 
ence in number of guard cells that Stephani described, existe 

The receptacle of Duvalia is nearly spherical, w r 
Cryptomitrium is disk shape. In other respects the recep A, 
resemble each other very much externally ; but in the ei 
ment of the receptacles there is a great difference. ee 
according to Leitgeb’s account, belongs to the type of Be é 
chantiacez, which has the growing point of the receptacle ® 


© LEITGER : of. cit. 87-90. 


a 
si 


1899] DEVELOPMENT OF CRYPTOMITRIUM 121 


forward margin. In Cryptomitrium the receptacle is a branch 
system, such as Leitgeb attributes to Marchantia and one or 
two other genera. This fact alone would be enough to out- 
weigh all the minor external characters referred to, were it not 
for the fact that, although Leitgeb puts Fimbriaria in the same 
type as Duvalia, Campbell (0. cé¢.) found that the receptacle 
of Fimbriaria Californica belongs to the ‘‘ Composite,” or branch- 
ing type. 

While one hesitates to criticise the classical work of Leitgeb, 
the quality and accuracy of which is for the most part remark- 
able, it does not seem reasonable that plants resembling each 
other as closely as Cryptomitrium and Duvalia should differ 
so much in respect to the growth of their receptacles, to say 
nothing of the fact that species of the same genus, Fimbri- 
aria, should also have this difference. It would seem more 
probable that Leitgeb was mistaken. Probably if one should 
carefully examine Duvalia and also the species of Fimbriaria 
which were studied by Leitgeb, it would be found that these too 
have as many growing points as there are groups of archegonia. 
Should this not be the case the apparent close relation between 
sa Cryptomitrium would be Only. apparent, and the 
oc oe perhaps, have to be considered more nearly 

cary farchantia, and Fimbriaria Californica could then no 

b€ considered as a Fimbriaria, for the difference in the 

‘wo kinds of receptacles is too great to occur within the same 
genus, | 


L ~ 
ELAND STANFORD JR. UNIVERSITY. 


BRIEFER ARTICLES 


NOTES OF TRAVEL. I. 


VENEZUELA. 


Tue aim of the expedition with which the writer is connected, 8 
planned by Mr. Barbour Lathrop, of Chicago, and carried out at his 
own expense, has forbidden any exhaustive research into the botanical | 
resources of South American countries. It has permitted rapid com 
parisons, however, and it is these comparative sketches which it ® 
believed will interest American botanists. 

The first approach to a great continent, if it has thousands 0 


square miles of unexplored territory in it, as South America has, 5 
ela, satis | 


The steep mountains behind the town shut it in like a green wall, 
the low hanging clouds and dark rainy valleys, into one of . 
famous railroad to Caracas disappears, are characteristically tropic . 
By characteristically tropical the writer may give a wrong impress 
since what could be characterized as tropical in one region might 
be true of another. The xerophytes are as abundant in the tropics ® : 
in temperate regions, although in the popular mind the 
acteristic of the tropics. Venezuela landscapes show 4 
portion of xerophytes than I had expected to see, and a ten 
tram ride to the small bathing place of Moquendo gave mea good oppo 
tunity of seeing the characteristic cactus vegetation of the ° 
Almost barren patches of reddish-brown soil and frequent signs of pram 
fires on the hillside surprise one, while the tufted grasses, agave” 
cacti give the whole a decidedly arid look. The climate of La a 
is a dangerous one for foreigners, as the malarial fevers there pen 
severe. We were informed, however, by intelligent English 
living in Caracas that the latter are no more severe than rhe 
acas itself. From my friend’s most uncomfortable experien 
evident that the capital hasa serious form of malarial fever, 2 ° 
care must be exercised to avoid exposure after sunset. favor | 
122 ae 


1899} BRIEFER ARTICLES 123 


The La Guayra and Caracas railway has some of the most pictur- 
esque scenery in the world. - Twenty-three miles of track are neces- 
sary to cover seven as the crow flies, and the curves and ziyzags along 
the coast give glimpses of great grandeur. The disappointing part of 
the landscape lies in barren soil and unmistakable signs of aridity. 
Curious cereuses and acacias or Prosopis, and a gigantic species of Ascle- 
pias with flowers three times the size of our A. Cornufti attract one’s 
attention, while the fine-leaved forest trees in the valleys give the land- 
scapeéa much more northern aspect than would be expected. ‘Iwo 
views strike the traveler most favorably ; one from a curve in the road 
which overlooks the coast, where, spread out below, are plantations 
of sugar cane and banana, fringed with the most graceful of cocoanut 
palms that stand out like dark green plumes against the white surf ; 
and the second, some distance nearer Caracas, where the road crosses 
aravine which drops into a narrow valley, 1500 feet deep and com- 
pletely clothed with forest. 

The vegetation effect, while most impressive, is not truly luxuriant, 
amd unmistakable signs of aridity are everywhere present. Curious 
arid ridges and isolated peaks along the sides of scantily covered val- 
leys give the impression of poor soil and rapid erosion. There isa 
Tumor that these arid patches were once wooded, but that injudicious 
Po constant forest fires, and later prairie fires have denuded them. 
bai atts the ? Venezuelan botanical authority in Caracas, does not 
The this. He declares they have been barren from prehistoric 


Caracas lies 2632 feet above the sea, surrounded by barren hillsides 


wh : 

Ptheciadg are covered with dense forest. 
; “te Is very little of botanical interest in Caracas itself. A few 
'nterestin 


through a gardens lie across the small stream which flows 
cultivated he = One In particular contained a number of curious 
seen, Ma 2 “4 In it was the most remarkable fountain I have ever 
4 small nbain ace iit aia imitation of a boa constrictor spouted 
Wes of storks (Of water. Around, on little artificial islands, were stat- 
were large oe the act of swallowing. Growing on these islands 
Tus, while i tase Oranges in fruit, and tufts of Egyptian papy- 
Plants and Chin ihe the basin a row of blossoming strawberry 
he coffee =e hibiscus bushes had been planted. ~ 

few Within th ‘a et Sefior R. Dalla Costa Mosquera, one of the 

© city limits, is well worth a visit and has in it a magnifi- 


124 BOTANICAL GAZETTE [ AUGUST 


cent avenue of St. Domingo mahogany trees. It illustrates the cofiee 
culture of Venezuela very well, which is in marked contrast to that ol 
Brazil by its employment of shade trees. ‘These cast a relatively deep 
shade over the whole plantation, and gave the impression of a thickly 
planted grove. In Brazil no shade trees are employed, and in Ceylon 
and Java they are planted very sparsely among the coffee trees. I 
seriously questioned the advisability of such heavy shading and was — 
informed simply that it was considered advantageous. Some of the 
best coffee in the world is grown in Venezuela, but very little of it 
reaches the American market because the latter demands principally — 
the cheaper Brazilian sorts. On the best Venezuelan estates the 
method of pulping the coffee berry before drying is in use, while the 
majority of Brazilian coffee growers still cling to the old method of 
drying the berry first and removing the dried pulp afterward. There 
are large Brazilian estates where this method: has been given up and the 
best machinery is in use. 

Nothing was seen in Caracas of the cocoa industry, although some 
of the finest cocoa in the world is grown along the Venezuelan st 
coast and in the interior about Maracaibo. 

In every city one of the most interesting places for a bota 
the market, and in Caracas it is characterized by an extraordinar 
of flowers. Tuberoses, white double violets, delicate purple Bi: 
Easter lilies, and curious bouquets made up of double columbines | 
marigolds and lilies surrounded with tissue-paper lace were most I 
favor at carnival time. Curious pear-shaped, thick-skinned shaddocks 
which are used for preserve making ; long, chocolate-brown, melon: 
like squashes, with orange-yellow flesh ; immense green water oo 
with squash-like meat, showing how careless the growers are about ! - 
interbreeding of their squashes and melons; bright yellow © of : 
mas” with orange-colored, mealy flesh, related to the curcumas . 
Peru, which are used in the manufacture of a favorite ice ; peaches: i. 
apples of inferior quality grown in the mountains of the inte 
together with the usual number of vegetables of quite inferior I 
ties compose the piles of produce on the well regulated stalls. is : 
part of the market is much more appetizing than the other ee 
where the most disgusting looking strings of salted meat are hung | 
sale. 

There is in Caracas a National Society of Agriculture 
to diffuse intelligence regarding the culture of agricultural pr 


nist is : 
y show 


tg 
4 


which ai 


1899] BRIEFER ARTICLES 125 


to teach the use of fertilizers and soiling crops, etc. But it is govern- 
mental, and when that is said of almost any concern in South America, 
it means that it is subject to rapid political changes. 

The courtesy of Mr. Alamo, Assistant Secretary of Agriculture and 
Mr. Romero, secretary of the society, could scarcely have been 
greater. 

To any foreign scientist, Dr. A. Ernst, the professor of botany in 
the University, is an invaluable acquaintance. His long experience 
with Venezuelan conditions and his fund of information on the botan- 
ical resources of the country are the result of a wide acquaintance and 
humerous expeditions into the interior. His advanced years make it 
impossible for him to continue his work as a collector, but his vigor- 
ous mind and excellent memory make his suggestions most valuable 
to a traveling scientist. 

As a young German botanist he came to Venezuela on a collecting 
trip and was invited ‘by the government to remain. Revolution has 
followed revolution, but, through his refusal to meddle in the politics 
. the country in the first place, and his undeniable ability as a scien- 
ust in the second, he has kept the position he now holds as one of the 
— profound scientists and highly esteemed citizens of the republic. 
hae ee in an attempt to educate Venezuelan youth to an 
en ne otany have not, it must be regretted, left him hope- 
a. outcome, and dizal what the writer could learn there is 
key courage foreign scientists to engage In governmental 

enezuela. 
oo? aon from Caracas into the interior, one English, 
erman ; neither, however, takes the traveler anywhere near 


the : A 
oF ay Interesting region of the Sierra Nevada, which lies, by mule 


'n order to explore these interior regions the traveler must 
atrange an fair knowledge of the Spanish language and should 
dimension oo. € in square leather boxes not over two feet 1n largest 
ble wit] be of - Ae donkey or mule transport. The food obtaae: 

€ very poorest quality ; and, as the peomes live In huts 


126 BOTANICAL GAZETTE [avcust 


of most unsanitary character, great care will be necessary to avoid con 
tracting the numerous diseases associated with such conditions. 

Caracas is not a favorable place from which to explore the resourees 
of Venezuela. It lies too far from the most interesting portions of the 
country. The Orinoco can be ascended better from Trinidad, and the 
Sierra Nevada requires an expedition on muleback to reach it. From 
descriptions given by travelers on the Orinoco and its branches, tle 
dangers from fever in the forest regions of Venezuela are very gretl 
and anyone undertaking their exploration risks his life. Mr & 
André, whose travels into the interior have been as extensive as any 0 
recent years, said he would not think of taking with him any perso 
who had not lived at least two years in the tropics and become atdl: 
mated as far as possible to conditions similar to those in Venezuela— 
Davip G. Faircuitp, U.S. Department of Agriculture. 


SOME SPECIES OF TETRANEURIS AND ITS ALLIES. 


We sometimes hear the statement that the difficulties for tit 
systematic botanist are being multiplied by the breaking up of somal) 
of the old genera and the creation of new species from former aggre 
gates, but practical experience shows, it seems to me, that sé 
gation, when based on describable characters, certainly simplifies. oe 
replacing of the untenable Ac#ine//a by TZetraneuris, Rydber gia, a | 
Picradenia (Pitt..3: 265), is a case in point. 

The reduction of several good species to one (an aggregate) mas 
necessary a description so general that the amateur in the field has. : 
difficulty in placing the most aberrant form until he collects 4s 
of specimens clearly unlike. In the past, reduction of speci ® 
often occurred because certain ones were rare and hence not i 
represented in the herbaria, but it seems unfair to eliminate ae 
simply because it exists in a locality not easily accessible 01M 
visited. Le 

is 


Being located in the center of distribution of Tetranenris ao 
allies, I became interested in the group. The following notes 
descriptions are offered as supplementary to Dr. Greene's valu . 
paper cited above. oo 


on 


Bis 


_ TETRANEURIS ACAULIS (Pursh) Greene, Pitt. 3: 265. 1898 


Galardia acaulis Pursh, F). 2:743. 1814. Actinella acaulis Nutt 


7.&GF | 
23381. 1842, etc. oe 


1899] BRIEFER ARTICLES 127 


As limited by the earlier writers this is a variable but a recognizable 
species. When many of the following were incorporated, the diffi- 
culties of the field botanist were multiplied several fold in respect to 
this species. 


TEYTRANEURIS ACAULIS cespitosa, n. var.—Strongly matted, depres- © 
sed-spreading, the numerous branches of the caudex much thickened 
by the imbricated leaf-bases: leaves very numerous and crowded, 
densely silky-lanate as are also the scapes and involucre: heads nearly 
sessile or on scapes 3-6™ long. 

That specimens of this variety exist in some herbaria as 7. acau/is is’ 
possible, though in the large series in the Herb. Mo. Bot. Garden none were 
found. Its matted habit, silky-lanate leaves and very short scapes easily 
Separate it. It occurs sparingly on sandy ridges in the foothills. Laramie 
hills nos. 1890 and 4314 represent it. 


Tetraneuris simplex, n. sp.—Tap root vertical, short, compara- 
tively small with few or many secondary roots: caudex short, consist- 
‘ng of one or more thick crowns which are densely covered with brown 
dead leaf-bases: leaves appressed-pubescent (not silky), nearly naked 
arsely long-hairy, glabrate in age when the fine puncta- 
i vident, crowded on the crowns, ascending or erect, 
near spatulate, tapering only slightly to the margined base, sub- 
meat” long : Scapes simple, single from the crowns, 15-25 high, 
‘lender, erect, lightly pubescent below, becoming silky or lanate above 
wy on the involucre : head large, 2.5—4™ across; rays with a broad 
ligule (5-8"") : akene pubescent. 


ding the two in the field. 7. acaulis is always cespitose, often in 
are shorter, the heads smaller, and the rays 
mplex. The leaves of the latter are comparatively 


€ first, strongly in contrast to the silky or even lanate pubes- 
er, 


Slabrate from th 
“ence of the oth 


sy i aks nave stthie been inclined to call this 7: acaulis is shown 
l. scatosa 5 = of - occur in the herbaria just as often ticketed 

‘hether Z; in 2 ep linearis, though to these it is not so closely related. 
© determine, 5, *S or ZT. simplex is the original Galardia acaulis is difficult 
> ve term « pilosa” in the original description, and the 


128 BOTANICAL CAZETTE [avcust 


agreement in Nutt. Gen. 173 and T. & G. FI. that the leaves are“ sericeous!y 
(silky) villous,” and that the plants are aggregated in dense tufts, suggests 
the separation that is now proposed. 

Besides a large series of plants from near Laramie, specimens af 1 
simplex have been examined as follows: T. A. Williams, Pine Ridge, Neb; | 
J. Schenck, Neb., 1893; H. J. Webber, Pine Ridge, Neb., 1889; Henny 
Engelmann, North Fork of the Platte, 1858; A. S. Hitchcock, Kan, 1845, 
no. 289; C. H. Thompson, Kan., 1893, no. 169; Capt. Bryan's Expedition, 
Lower Pole Creek, Wyo., 1858; R. S. Williams, Great Falls, Mont., 189) 
no. 82; G. E. Osterhout, Livermore, Colo., 1898; M. E. Jones, Cheyenit 
Cafion, Colo., 1878 ; Hall & Harbour, no. 275. 


Tetraneuris incana, n.sp.—Root rather slender, simple or branched: : 


~ 


the crowns, silvery-white with an appressed pubescence, linear-oblat: 
ceolate, 2-4™ long: scapes naked, single from the crowns, sles 
curved-ascending, 1-2 high, the fine silvery pubescence sii 
spreading: involucre silvery-silky, bracts few, shorter than the 1 hip 
disk, the outer oblong, obtuse, the inner spatulate, scarious marg! 
rays few, the ligule as long as the disk: disk corollas sprinkled ® : 
resinous globules and toward the summit strongly thickened by a den 4 
penicillate, glandular beard: pappus scales oblong, aristat: - 
slender, nearly as long as the corolla, pubescent. 

This rare species is strongly marked in its close, silvery pub 
nearly simple caudex, its silvery involucre, and its dense coat of gla 
on the corollas. The only collections of it at hand are no. 393) © Nels 
near Fairbanks, July 11, 1894 ; no. 5006 (type number), by Mr. Elias aie 
Wallace creek, July 30, 1898; and a specimen by Mrs. Muth, Lewis & i 
co., Mont. Its habitat is white clay ridges among the barren hills. 


escenses 


TETRANEURIS TORREVANA (Nutt.) Greene, l. c. 

Actinella Torreyana Nutt., Trans. Am. Phil. Soc. 7 +379: ae et | 

A strong species of the central-eastern Rocky mountains: wee 
eral forms, but always tufted, strongly punctate and neatly = i | 
except on the caudex ; somewhat variable as to the width ~ ee 
of the leaves. My no. 48ro, from the Platte hills neat wee 
June 18, 1898, are nearly typical ; nos. 4571 and 4747) Jun 
the Tertiary clays of south-central Wyoming have broader, 


1899 ] BRIEFER ARTICLES 129 


than the original description permits. A form represented by no. 
4327 and some earlier collections, from the limestone ledges of the 
Laramie hills, is strongly matted and has the branches of the caudex 

enormously thickened and protected by the densely lanate leaf-bases. 

' Add to this its large root, broad green leaves, and the copious secre- 
tion of its punctate glands, and it might well stand as var. glandulosa. 
It is in this species that the salient character of the genus (4-nerved 
ligules) often fails ; 5-8 nerves are not infrequent. 


etraneuris Mancosensis, n. sp.— Tufted, with woody root and mul- 
ticipital caudex, the short.thickened crowns clothed with the expanded, 
membranous, lanate leaf-bases: leaves glabrous, 4-8" long, crowded 
on the crowns, linear or linear-oblanceolate, acute or cuspidate, rather 
minutely punctate: stems few to several, bearing a few (usually 2) dis- 
‘ant leaves, 2" in length (including the long monocephalous peduncle) : 
heads large, disk about 1™ high; involucre silky-lanate, the bracts in two 
a three tows, the inner oblong or somewhat expanded upwards by the 
— margins: palee of the pappus oblong-elliptic with an acu- 
mination as long as the body proper, equaling the disk corollas: 
ligules of the rays Is—rgm™ long, 6-8™ broad 
Paani by Professor C:S. Crandall, Mancos, Colo., June 29, 1898, and 
Bearer to ge aloe Scaposa Hinearis Nutt. It is in fact, however, much 
ously pula gata from which its slenderer, longer, and less conspicu- 
long-ped © ‘faves, its nearly glabrous two- or three-leaved stems, its 
eta heads, and long pappus palez at once separate it. 


T " 
FTRANEURIS LaNnaTA (Nutt.) Greene, 1. c. 
: lla lanata Nutt 


: mls Ge ‘ 
are plant of the arid interior, on dry ridges on the high plains. 
ne following collections of it have been secured by the writer: 
from Ft, . from Green river, in 1897 and 1898 respectively ; 4607 
8er, June 9, 1898 are young specimens, but probably the 
‘though these of less of the wool is permanent even on the leaves, 
oe ®w some punctation on the glabrate areas. 
panna GRANDIFLORa (T. & G.) Greene, l. c. 
This aaa T. & G., Journ, Bost. Soc. Nat. Hist. 5:110. 1847. 
ges, *S In abundance in the alpine regions of all our mountain 


man 


Picrap 
‘athentic — Ricuarpsonn Hook., Fl. 1: 317. p/. 208. 1833.— That 
[7S oens Of this occur in this range is possible, but it seems 


130 BOTANICAL GAZETTE | avonst 


quite certain that the wide range attributed to this species in Gray: 
Synoptical Flora is due to other species being included. With t 
erection of P. floribunda (Gray) Greene and P. canescens (Eaton) Greett 
into species the area covered has also been segregated. But even allt 
the establishment of the two following species some Rocky moutta 
, are still left to represent the origi 


forms, such as my no. 
species. 

PICRADENIA LIGULAFLORA Aven Nelson, Bull. Torr. Bot. Claba | 
378. 1898. 

This species is proving to be far more common than was at fis | 
suspected. As to habitat see notes on the following species. 

Picradenia macrantha, n. sp.— Caudex branched, each branch s 
mounted by a few to several crowns; crowns clothed with the lana 
leaf-bases: stems single from each crown, erect, tascicled, sparse 
pubescent, somewhat striate, about 15°" high : leaves glabrate, not coe 
spicuously punctate, rather numerous on both crowns and siti 
slender petioled, the blade variously parted into linear divisions 
of them pedately trifid, some of the stem leaves pinnately P 4 
twice trifid, the uppermost sometimes simple: heads large, peduncle 
one to five on each stem (generally two or three): involucre ge 
than the disk flowers, outer bracts lanceolate, nearly glabrous, © 
for half their length, 6™ long; inner oblong, acute, scarious me é 
rays 6-10, chrome yellow, the ligule 15-18" long and one thi 
broad : pappus scales 5-7, lanceolate, shorter than the corolla. 


habitat of this is open, stony slopes in the mountains or hills, 
flora occurs on dry, clayey, alkaline ridges or flats on the ope? sist | 
number 4830, Fort Steele, June 18, 1898. What seems to be the er 
is no. 1688, Centennial hills, Aug., 1895 ; also South Park, Cole» 
Marcus E. Jones.—AveNn NELSON, University of Wyoming. 


PYCNANTHEMUM VERTICILLATUM, A MISINTERPR®™ 
MINT. 
to 4 


Mr. W. W. Ecc.LeEston has called my attention 
themum, abundant about Rutland, Vermont, which has ss 
by recent botanists with both ?. muticum Pers. and P. gues 
but appears different from either of those specie - 


1899 ] BRIEFER ARTICLES 131 


the plant matches the original plate published by Michaux‘ with his 
description of Brachystemum verticillatum. Fortunately a portion of 
Michaux’s type specimen is preserved in the Gray Herbarium, and a 
careful comparison with that leaves no doubt that Mr. Eggleston’s 
plant isthe same. Another Vermont specimen collected by Robbins, 
anda number of specimens from more southern states possess the 
same characters as the Michaux plant, and with it represent a species 
fairly distinct from either P. muticum or P. T. orreyt. 

Brachystemum verticillatum was described by Michaux from the 
mountains of Pennsylvania and Carolina. It was soon transferred to 
Pyenanthemum by Persoon,? who merely accepted Michaux’s descrip: 
ton without comment. Pursh in his Flora, took up the name, modi- 


M In 1891 Otto Kuntze® took up the obsolete’ generic name Koed/za 
eee Setierred to it Michaux’s Brachystemum verticillatum, 
muticum, and % statement it is apparent that he meant Brachystemum 
Derticillaty Was merely following Dr. Gray in regarding that and &. 
itehnes. sy the same. In 1894, in their list of the plants of 
different 5 oe Irginia, Dr. Small and Miss Vail treated the plants as 
cum Michs “a Britton! there transferring Brachystemum muti- 
Benth.) ess Oellia. A. Zorreyi Kuntze (Pycnanthemum Torreyi 

so then recognized as a distinct species. In the Botan- 


*MIcux,, F], 9. 
»Flia: 6. pi, 37, 4 BENTH. Z. ¢, 330. 


5 DC. Prodr. 12: 190. 
’ The 60. K. Rev. Gen. 2: 520. 
Te is ]j 
ace '§ little reason to suppose that the name Aeellia will be generally 
ough 


* Pes, Syn. 2: 128, 
* BENTH. Lab, 328. 


sretsight Was P 
to Engle 

8 
Morncy Meth. 4o7, 


*Brrrp * 
©N in Smal! and Vail, Mem. Torr. Bot. Cl. qi 145. 


132 BOTANICAL GAZETTE {avcusr 


ical Club Check List, however, P. Torreyi Benth. is reduced tof 
verticillata. Thus within three years this plant, which was quite. 
obscure to the great monographer of the Labiate, under the inspi- 
ing influence of a resuscitated generic name was combined with # 
least two other species: Otto Kuntze, following Dr. Gray, considett 
the plant a form of Pycnanthemum muticum ; then it was treated 1 
species distinct from both P. muficum and P. Torreyi; then dui 
the same year it was united with P. Zorreyz. ; 
As stated above, a study of the type of Brachystemum verticillates: 
Michx. shows that plant to be fairly distinct from either his 3. 
cum or the later Pycnanthemum Torreyi Benth. The follows 
description and notes may help to distinguish this plant, and show tt 
points of similarity and of difference between it and those with whict 
it has been associated. 
y and rathet | 


PYCNANTHEMUM VERTICILLATUM Pers.—Stem_ shortl 
corymbos: 


closely pubescent especially above, the branches loosely sub : 
* leaves ovate-lanceolate or lanceolate, entire or slightly toothed, 
sessile, mostly glabrous; the reduced upper ones subtending . 
rather. dense terminal or verticillastrate glomerules, densely shat 
pubescent : bracts of the glomerules ovate-lanceolate with subulate" 
more or less pubescent on the backs, and with ciliate margins: oe 
5.5"" long, with the 5 hispid lance-subulate teeth about equal : 
7 or 8"" long, the upper lip short-oblong or obovate, the lower 
the oblong middle lobe twice exceeding the lateral ones: * 
included, scarcely 1™™ long. —Syn. 2: 128 ; Benth. in 
190 ; Gray, Am. Jour. Sc. 42: 47. P. danceolatum Benth. Lab. 
part, not Pursh. /P. muticum Gray, Syn. Fl. 2: 355) in part, 
Brachystemum verticillatum Michx. Fl. 2:6. pe 31: 
cillata Kuntze, Rev. Gen. 2:520 (as to synonym only); i di 
Vail, Mem. Torr. Bot. Cl. 4: 146 (at least as to Farmer Mie 
but not as to note, which apparently refers to Pycnanthemum | 
toides Gray); Britton, Mem. Torr. Bot. Cl. 5: 28° (exclnt 
Pycnanthemum Torreyi Benth.).—Specimens examined : a 
(Michaux); Colchester, Vermont (Rodédins) ; Rutland, Roy! 
Clarendon Springs, Vermont (W. W. Eggleston); Be | 
setts (C. W. Jenks and C. W. Swan); Dedham, Mase 
Alice Browne) ; Providence, Rhode Island (George There 


* Mem. Torr. Bot. Cl. 5: 280. 


1899] BRIEFER ARTICLES 133 


York (/. Carey); Easton, Pennsylvania (Z. C. Porter); Farmer Mt., 
Carroll co., Virginia ( John K. Smait). 

Habitally Pycnanthemum verticillatum should rarely be confused 
with either P. muticum or P. Torreyt. P. muticum has broader ovate- 
serrate leaves, the upper distinctly whitened ; the bracts of the glom- 
erules are linear or lance-attenuate ; andthe long stamensare generally 
much exserted. Though in foliage P. verticillatum is approached by 
P. muticum, var. pilosum Gray, that form is readily distinguished by its 
characteristic pubescence and bracts. 

tom the somewhat similar 7. lanceolatum, to which Bentham once 
referred it, P. verticillatum is distinguished by its broader leaves, the 
Upper ones pubescent ; by its more open inflorescence ; by the longer 
more acuminate and less pubescent bracts of the glomerule ; and by 
the sharper calyx-teeth and smaller corolla. 

Some forms of Pycnanthemum Torreyi approach P. vertictllatum, 
but usually that Species may be readily told by its thinner narrowly 
lanceolate leaves, distinctly tapering at the base, the upper mostly 
slabrous ; and y the narrower bracts of the glomerule. The stamens 
of the wo plants present a striking difference. In P. Zorreyd they are 


long-exserted, equaling the upper lip of the corolla; while in 2. vertt- 
cllatum they are 


the corolla, 


Verticillatym 
Ti orreyi 


, 


More re. ‘ 
Senus — to treat it asa species. The specific characters in the 
<a are not so fixed as would be convenient for 


co » DUt as species go in the group, P. /anceolatum 
and P, “inifolinm i P 8 Adal! 3 : 
m S€ems Ww 


™m, Camby idge, Mass. 


134 BOTANICAL GAZETTE [avctst 


THREE NEW CHORIPETALAE FROM NORTH AMERIG 
D MEXICO. 


” Silene rectiramea. — Cespitose perennial, 2 to 3°” high: stems st 
eral from a multicipital caudex, covered at the base by the pale scarious 
persistent scale-like ciliated bases of the earliest leaves, terete or elliptt 
in section, slightly striate in a dried state, pubescent and more or Its 
viscid especially above, sometimes simple to the inflorescence, som 
times branched from every node; lower internodes relatively shot 
often curved, the middle and upper elongated, 6 to 8™ long, ™ | 
exceeding the leaves, remarkably straight ; branches solitary or opp 
site at the nodes, diverging from the stem at a uniform angle of about 
45°; their internodes‘also elongated and very straight: leaves of the 
stem about five pairs; the lower ones, like the radical, oblanceolalé, 
2.4 to 4™ long, 4 to 7 broad, the middle and upper lance-oblong ® 
linear, all acute, 1-nerved, obscurely pinnate-veined, minutely papillos 
and pulverulent-puberulent under a strong lens: bracts lance-linea,_ 
often purplish, 7 to 9™ long; bractlets similar, 2" long : flows 
terminal on the divergent branches of and open flat-topped cymes 
in weaker stems reduced to a terminal and one or two short-pee® 
lateral ones; calyx cylindric in anthesis, white and papery but vet 
with light purple, 9™" long, 1o-nerved ; the nerves opposite the a 
branching freely, intermediate ones subsimple; teeth orbicular 
incurved margins: gynophore in anthesis 1.7, in fruit 2.5 long 
1 to 1.1 long, glabrous except externally at the very base ; claws 
ulate, subauriculate at the summit, 3-veined ; blade short, 2 aS 
long, bifid a fourth of its length: stamens 10, equal : carpels 3: 4 
sule ovoid, 7"" long, at maturity 1-celled to the very base : seeds 
dish-brown, tubercles in few concentric series, those of the dorsal 162 
enlarged and forming a more or less definite crest. 


z 


Collected by Professor D. T. MacDougal about the Grand Cafion of . 
Colorado in Arizona, altitude 2150™, 28 June, 1898, no. 181. 
ray. 

This species stands near S. verecunda Watson, ag 
gated very straight branches and delicate papery calyx, which, altho pete 
rowly cylindric to obovoid, shows no indication of the tightening * : i 
tion about the carpophore which is to be noticed in 5S. verecunda. pee | 
S. rectiramea were distributed in Mr. MacDougal’s interesting ee il | 
but were determined only to the genus. Mr. A. A. Helle 0d 
charge of the identification of the sets, has courteously waivel 
author his right to characterize.this species. 


1899 | BRIEFER ARTICLES 135 


‘Arabis Crandallii.— Cespitose perennial, 3°" high, pale green and 
hoary puberulent throughout, with minute stellate interplexed hairs: 
stems numerous (20 or more), slender, terete, from a loosely mul- 
ticipital caudex: root single, vertical: radical leaves oblanceolate- 
spatulate, 1.5 to 1.8 long, 3 to 4™ broad, entire, acutish, cuneate- 
attenuate at the base, concolorous, minutely stellate-tomentulose on 
both surfaces, I-nerved ; the cauline (about 8 on each stem) similar but 
shorter and more oblong, sessile by a subamplexicaul base: pedicels 
ascending or appressed, 5 to 6™™ long, slightly enlarged at the summit : 
sepals oblong, obtuse, stellate-puberulent, often purplish-tinged, 3™ 
long: petals obovate, cuneate, white, roseate-tinged, twice as long as 
the calyx : pods erect, slender, subtorulose in dried specimens, 2.5 to 
4" long, 1™" broad, flattened ; seeds (immature) uniseriate in each cell 
and nearly or quite as broad as the septum. 

Collected by Professor C. S. Crandall at Cimmarron, Montrose co., Col- 
“Ee attitude 2100", 18 May, 1898, no. 6. Type in herb. Gray. 

Was nid Species most early approaches the Canadian A. Macounn 
“s which, however, it is clearly distinguished by its fine stellate 
pubescence, shorter erect pods, and larger leaves. 


$-foliolate j oe 
fololate in the manner of AL. sensitiva (a diminutive fourth leaflet occa- 


Pre aa leaflets glabrous, glaucous, coriaceous, oblong, entire, 
very He nes, acute to rounded at the apex, subcordate and 
Pe a the base, oblong, 3 to 10™ long, nearly half as broad ; 
35 ‘ag i slender, wiry, 3 to 8™ long ; secondary rhachises 2 to 
along the ae uncles slender, ascending, fascicled by 3’s, 4’s, and 5’s 
fated, ora Pen of the branch, and forming a loose, elon- 
» teem y iad inflorescence, leafy towards the base: heads 
Perfect aie a diameter (incl. the long stamens), roseate ; flowers 
date. ces calyx campanulate, less than 1" long, cuspi- 
corolla 2.7™™ long, glabrous, 3-4-nerved and 3-4- 
ate-deltoid, a third as long as the tube: pods 2.5 to 
‘and wholly unarmed both as to valves and replum, 
ity ; too acuminate at the tip, 3-4-jointed, Indian-brown 

> Valves falling away in segments. 


136 BOTANICAL GAZETTE [ aveust 


Common on hills near Acapulco, Mexico, where collected by Dr. Edwarl 
Palmer between October 1894 and March 1895, no. 29 

I am indebted to Dr. Rose for calling my attention to this species, Dr 
Palmer's specimen having been undistributed in the Gray Herbarium at the 
time of my recent revision of the genus. Types in herb. Gray and herb. U.S 
National Museum. — B. L. RoBrnson. 


THE PROBABLE CAUSES OF THE POISONOUS EFFECI 
OF THE DARNEL (LOLIUM TEMULENTUM L). 


THE presence of a poisonous principle in the darnel has been wel 
known since the earliest investigations of the subject, and  receil 
experiments confirm this fact. According to Hofmeister,’ the darne 
contains two active principles: semu/in, obtained by this author ® 
chloroplatinate, which acts upon the nervous system ; and the othe 
determined by the oily substances and fatty acids which are contaie! 
in the seed in large proportion, which attacks the alimentary canal 

In the course of our researches upon the seed integuments and tit 
pericarp of grasses, we have had occasion to note the practically es 
stant presence in the seeds of the darnel of a fungus to which it se 
reasonable to us to assign the poisonous effects. This fungus, ~ 
is always present in the form of mycelial filaments, appears at an © 
stage in the interior of the ovary. In the first stages of its - : 
ment it invades the entire nucellus. At the time that the ke 
integument of the ovule disappears, the nucellus itself is we 

cde 


which, obliterated in the maturing of the grain, constitute the bar 


the outermost endosperm. It is in this zone that we have ” f the 
in the mature seed. After the removal of the diverse coatings® 
fruit, the hyphe which constitute this fungus zone appear exw 
filaments, generally very long, more or less branched and int 1 
with one another. We have found this disposition of the fang ne 
material from Bolivia, Brazil, Chili, Abyssinia, Persia, ae fort 
Portugal, Sweden, Germany, and many localities in France. from be 
seeds of most diverse origin the mycelial zone was lacking ee 4 


* Archiv. f. exp. Path. u. Pharm. g0:—. 1892. 


1899] BRIEFER ARTICLES 137 


three. This observation has been confirmed in other species of Lolium, 
to wit, Z perenne L., L. arvense With. (var. of LZ. temulentum), L. lini- 
cola Sond. It is only exceptionally that the first of these contains the 
parasite. The rest are infected to the same degree as L. ¢emulentum. 
When one observes that the species reported poisonous are the very 
ones in which we have found the parasite, it seems reasonable to ask 
whether the temulin of Hofmeister is not a result of the action of the 
fungus upon the nitrogenous materials in the peripheral region of the 
seed. 


This fungus, of whose nature we are not yet satisfied, may not in 
any case be identified with the Exdoconidium temulentum of Prillieux 
and Delacroix. The latter attacks the seeds of rye which it clearly 
deforms, the infected grains becoming smaller and lighter than the 
hormal ones. The grains of the darnel show no such deformation. 
Further, in the rye grains thus attacked, and called “ setg/e enivrant,” 
the protecting layer has generally disappeared, and all the external 
part of the endosperm has been invaded by the parasite. In the dar- 
ve the endosperm suffers no alteration from the action of the fungus, 
— layer itself remaining perfectly intact. Since our obser- 
ee were made the same mycelial layer has been noted by Hanausek 
nd Nestler? and before them by Vogl.3 
— Tesults are practically alike. However, the other authors have 
. bed the fungus for Z. semulentum alone.—P. GuEkrIn, Préparateur 

Ecole Supé 


rieure de Pharmacie de Paris. 
2 1 iy : 
- Berichte der deutschen botanische Gesellschaft @:-. pl. De ash 0 ai 


> Die wichti 
Wichtigsten vegetablischen Nahrungs- und Genussmittel. 1898. 


PPEN LETTERS: 


IN YOUR June number and ina Aulletin of the South Carolina Agrictl 
tural Experiment Station, Dr. A. P. Anderson quotes me as identifying the 
Tilletia found on rice in South Carolina with Z72/etia corona Serib. The 
resemblance is certainly striking, but in writing Dr. Anderson I did not intend 
to express a final opinion in the matter. I had not at that time seen@ 
description of the Japanese 7i//etia horrida Tak. It now seems to me - 
the differences in the manner of affecting the host plant, the spore mass beiig 
included by the glumes in 7. Aorrida and conspicuously exserted in 7. carats 
should be considered of sufficient weight to separate the species a 
until such time as their life histories can be carefully studied. Theref ee 
should prefer to call the South Carolina specimens 77//etia horrida Ta 5 
my opinion much more confusion is occasioned by the hasty Gain 
many forms under one common name than by tentatively recognizing. 
many forms as independent species. 

the name 7. Nea eaeg seems to be antedated by Arthur's ye 
rotundata (Prel. List lowa Uredine, Nov. 1884), described _ Towa spe 
mens on Panicum irrigatum. This species has recently bee rll & 
543 Of Economic Fungi, under the name Tié/etia rotundata "(aa 
Ev.—F.S. EARLE, Auburn, Ala. 


SeRRENT LITERATURE. 
BOOK REVIEWS. 
The Sandusky flora. 


A LocaL flora of much interest has been published by the Ohio Academy 
of Science as the first of a series of “special papers,” the second and third 
tumbers of which are devoted to insects. This initial effort reflects credit 
upon the Academy, both for the subject-matter and for the manner of its pres- 
entation. The volume of 167 pages is thoroughly satisfactory in its typo- 
sraphical and press work, and records a piece of careful local exploration 
worthy of the excellent setting. , 

The flora of Sandusky and vicinity as here presented is the result of the 
labors of Mr. E. L. Moseley ' during the last seven years, and is based upon 
the herbarium of the Sandusky high school, while specimens of most of the 
ron forms are also deposited in the Gray herbarium of Harvard University 
and in the herbarium of the Ohio University. 

The catalogue proper, which is well indexed, is preceded by thirty-four 
tial text, in which a number of interesting matters pertaining to the flora 
eae —; It adds not a little interest to learn that the flora of Snagueiey 
biieha a in the humber of its species, having more than is given in those 
a. by David F. Day for the Buffalo region, by Wm. R. Dudley 
ter ro hia and by the local Academy of Science for the Roches- 
erally lar, a mati these regions border on the great lakes and are sev- 
es not ne an that included for Sandusky. Even the whole of England 
about San, co % hundred species of phanerogams more than are found 
explored and a ‘The islands included in the area, a half dozen having been 
of the flora — with special care, do not add materially to the richness 
Surprising . a have only a few plants not found on the mainland. The 
itions in part a of ss sacri is ascribed to favorable physiographical con- 
5 dusky ‘hin wes a especially to the climate. The summer is longer at 
4 month longer — points along the shore of Lake Erie, being much over 
Average temperat an at Buffalo, and the spring and summer have a higher 
Statistical evide ure. The reasons for this state of things are discussed and 

nce presented, 


* Mosete 
Y : 
Sowing ies. a Sandusky flora, a catalogue of the flowering plants and ferns 
. county, tivation in Erie county, Ohio, and the peninsula and islands o 


wa : 
Bro, PP. 167, 1 Paper No.1, Ohio Academy of Science ; Wooster, 1899. 
1899} . 


139 


140 BOTANICAL GAZETTE [AUGUST 


The nomenclature of the catalogue is that of the Kew /ndex, with sme 
concession to the last edition of Gray’s Manual. The enumeration begins 
with Botrychium ternatum Swartz and ends with Xanthium Canadense Wi) 
Altogether it is an excellent catalogue, and reflects credit upon the industry 
and ability of the author, and upon the enterprise of the Academy of Science. 


Anatomy of the dicotyledons. 


A SENTENCE of Radlkofer, which the author thinks prophetic—“l 
next hundred years belong to the anatomical method” — inspired Solereder 
to bring together the immense mass of material regarding the anatomy of the 
stem and leaves of the dicotyledons into a handbook for botanical labow 
tories? To the previously accumulated knowledge of anatomy the @ 
himself has been a notable contributor. 

e labor of compilation alone must have been immense. Hert #* 
brought together in systematic form the data regarding each of the familie 
of dicotyledons. The author first presents a synoposis of the chief anatom 
cal characters of the family as a whole; then gives an extended account of 
the leaf structure, followed by a similar presentation regarding the ste 
Each section closes with a thorough bibliography. ae 

In an introductory chapter Dr. Solereder explains what is meant by 
anatomical method, and discusses the more important anatomical en: 
and their value in taxonomy. An extensive closing chapter (75. pas . 
devoted to a synopsis of the various anatomical features, with reference 
their occurrence in certain families, genera, and species—@ sort of compal® 
tive anatomy. : ined © 

Though one may doubt whether the anatomical method Is re ered 
play the important réle in taxonomy which Radlkofer and his pup : 5 wel 
believe, and may easily find matter for criticism in the introduction, the 
as flaws in the details regarding structure, the enormous _ ye 
author has performed in the production of the work disarms ge rele 
evokes only praise. The book is unquestionably a most useful one 19" 
ence, not only to the systematist, but to the histologist and to be ee 
as well. It will prove indispensable in every botanical departmt’ 
active work is in progress and will doubtless demonstrate its value 
to day. 1} inde® 

As a reference book it has one serious defect, the want of af sve tie 
The index only includes the families, and while ordinarily this will 8! 7 
desired clue, the value of the book would be greatly enhanced by 40°" 

* SOLEREDER, HANS: Systematische Anatomie der Dicotyledonen: pee 
buch fiir Laboratorien der wissenschaftlichen und angewandten we 
xii + 984. figs. 189. Stuttgart: Ferdinand Enke. 1898-1899. Me 


—— 


1899] . ; CURRENT LITERATURE I41 


each genus and species and of each author mentioned. The Bavarian 
Academy of Sciences, which has assisted in its publication, might well have 
increased its subvention if necessary to provide such an index.—C., R. B. 


Speculative biology. 


In 1875 Pfliiger propounded a hypothesis regarding the constitution of 
organized bodies which may be described as the hypothesis of chemical con- 
tinuity. Impressed with the extensive polymerization among carbon com- 
pounds, especially the proteids, he ventured the suggestion that in an organism 
polymerization may progress indefnitely, so that the whole protoplasm is not 
an aggregate of similar molecules having definite molecular weight but may 
form a single giant chemical molecule. This theory has found few adherents. 
It is accepted in toto, however, by Dr. Georg Hérmann, who proceeds in a 
recent book? to show its adequacy to explain certain biological problems, 
and, therefore, its inherent probability. 

He applies it to the transmission of the impulse in nerve and the phe- 
nomena of nerve section; to the contraction of muscle and the discharge 
from the electric organs of fishes; and discusses the structure of the cell and 
- Gitepra of the protoplasm “ from the standpoint of the principle of chem- 
‘cal continuity,” (Hypothesis — principle: are they synonymous?) 
ro aa the book is pure speculation, and must not be taken as anything 
hick oe fs fear me author does not always remember the sandy founda- 
diss oe as sie building. The various ingenious diagrams, sepreeeptDe 
whieh os Ivers interesting forms and positions lend an air of verisimilitude 

ch might deceive the very elect. 
sled vag as necessary ; speculation is indispensable in the 
srrioksis AN naa hypotheses by the investigator ; but it may . 
while. Until ae whether the publication of a speculation is ever wort 
speculation of . me more intimate knowledge of the chemistry of proteids, 
*aoity init vi, e kind here set forth must be regarded as little more than 

: ation of spirit—_C. R. B é 


ae NOTES FOR STUDENTS. 
St ote his observations on the agencies by which insects are 
“alviq a paabile Professor J. Plateau now gives a large number made on 
ment that the oi on Hydrangea opuloides,* confirming his previous state- 
aS Y are chiefly attracted by the sense of sight. Neither the col- 
ORMANN, Grorc 


der lebenden s : Die Kontinuitat der Atomverkettung ein Strukturp’ sone 
Aa ubstanz. 8yo. pp. iv-+118. figs. 32. Jena: Gustav Fischer. 1899. 7 
*Mém, 

Soc. Zool, de France 11 : 339-375. fig. 4. 1808. 


142 BOTANICAL GAZETTE [averst | 


ored bracts in the former nor the conspicuous sterile flowers in the latte — 
plant can be regarded as “vexillary.” In both cases the pollinating insets 
make their way at once to the flowers which contain the honey without bem 
visibly guided by the showy organs in either case; while if these are remove! 
it does not appear to make any material difference in the number of insets 
which visit the inflorescence.— Jour. Roy. Mic. Soc. 1899. 298. 


IN A NOTABLE paper on cellulose enzymes, Professor F. C. Newcomit 
clearly demonstrates the existence of cytohydrolvtic enzymes distinct free 
diastase, especially in the seedlings of white lupine and date palm- 4 
enzymes, which in some plants are doubtless mixed with diastase, “act®™ 
starch so feebly and on reserve cellulose so energetically that they ate toe | 
regarded as cytase as distinguished from diastase.’ In all cases the oo 
walls first become hyaline, then more and more transparent, finally seeming | 
to melt away in solution. Besides the clear proof of the existence of the log 
suspected cytase, the paper adds much to our knowledge of the distribut® 
of cellulose enzymes.— C. R. B , 
states that his obs 
ic membrats 
at least a 


Dr. A. M. Bousier in a brief paper on the pyrenoid® 
vations “ prove the existence in pyrenoids of an external plasm 
perfectly differentiated and independent of the chromatophore, 
mature stage of development. This membrane encloses a Jeucoplast, 
accumulates starch, with a crystalloid at the center.—— C. R. 5. 


aquatic plants.’— C. R. B. 


flight 

THE CHIEF VALUE of Kolkwitz’s recent paper on the influence ° se 

the respiration of fungi® is due to the refinement of technique emplo 
the degree of accuracy attained. In these respects It SUF 


is the first extensive accurate study of the effect of light upon ™ 
tory activities of plant protoplasm and of animal protoplasm a5" i 
mals are prone to move and then by their varying activities 
any conclusions as to the effect of light alone. Severed ap - vet 
unsuitable, as diffusion at the cut end is abnormal and quant 

Fungi having false parenchyma are unsuitable since the intercel 


> Annals of Botany 13: 49-81. 1899. 

° Bulletin de l’Herbier Bossier 7451-458. 1896. 

7 Bulletin de I’'Herbier Boissier 7:—. 1899. 

*Kotkwitz, R: Ueber den Einfluss des Lichtes auf di 


e Athmung = be 
Pilze: Jahr. f. wiss. bot. 33 : 129-165. : 


1899] CURRENT LITERATURE 143 


may suffer change from light. Therefore the author selects such fungi as 
produce a loosely woven mycelium that spreads itself out openly to the light 
(Aspergillus, Penicillium, bacteria), and measures such activities as are solely 
dependent on light. As sources of error he recognized chiefly the evolution 
of CO, through decomposition of oxalic acid or of dead parts, and errors 
introduced by variation in temperature. The classical method of Petten- 
kofer (1862) is adopted and the amount of evolved CO, is determined by 
titration with oxalic acid. To secure greater accuracy the gas was forced, 
not drawn, through the apparatus at the rate of three, four, or five liters per 
hour as desired. The process and apparatus are described in great detail. 
Suffice it to say here that exceeding care was given to every feature. The 
culture vessel, of special design, presented a great surface to the light while 
of but small capacity. In order to reduce the absorption by glass the walls 
of the vessel were very thin. The feature wherein this study chiefly surpasses 
previous work is in the regulation of the temperature of the culture during 
€xperimentation. This was accomplished by immersing the culture vessel in 
‘tank containing six liters of water and keeping it at a constant temperature 
electricity, automatically regulated by a very ingeniouscontrivance. Lest 
the thin layer of water covering the culture flask should vary in temperature 
<. water was continually agitated by a paddle operated by a SurRine. 
"i iy air was warmed to the temperature of the water. In this way 
icht 1on was from one tenth to one thirtieth of a degree C. The electric 
ae Was Constant in quantity and quality, thus avoiding the variations inevi- 
in the use of sunlight. Estimations of CO, were made every two min- 
el ey aan announces as a result of his labors that light, under the 
as ployed, increases respiration about 10 per cent. The effect is 
Nid cs ey oF old cultures, richly or poorly nourished fungi, and in 
secondary eae The influence of light during long periods when 
Weiven =f 1 a arise was not investigated. An excellent bibliography 
- L. STEVENS. 
the Wea fon cAxoxomr interest are as follows: In continuing his flora of 
the A ‘alice (Symbole Antillane), URBAN, in the second part, resents 
»Tepr 
24! describes about 
Rico 
erma(Rhamnacez). Linpav presents the Polygonacee, 
Containing —. and sixty-six species, the great genus Coccoloba 
“Presented by tw ty of them. ScHLECHTER presents the Asclepiadacee, 
being Mean... “one genera and eighty-eight species, the largest genus 
Rew genera oa sis Mess thirty-four species, eighteen of which are new. es 
Mag. Tokyo a; ablished, Tainionema and Decastelma.— K. MIYAKE (20 : 
‘T~4. £2. 3. 1899) has described a new genus of Hepatice, 


144 BOTANICAL GAZETTE [avers 


very closely resembling Pellia. It is said' to have a spermatozoid me 
larger than that of Pellia, which has heretofore been credited with the lage 
spermatozoids among the Hepatice. The new genus is known as Matinee, 
in honor of Makino the discover: Specimens without sporogonia had alreat 
been described by Stephani as Peé/ia crispata, so that the name stands 
M. crispata (St.) Miyaki.—In Proc. Amer Acad. (34: 507-534. 1899) Be 
INSON and GREENMAN publish revisions of Montanoa, Perymenium, and Ze 
uzania, long a puzzling series of Mexican and tropical American composite 
Montanoa is recognized as containing thirty-two species, nine of which are ne 
Perymenium has twenty-six species, ten of which are new, and Zalusans 
has twelve species, two of which arenew. The same authors (did. 534-50 
have published a synopsis of the genus Verdesina, which has not been treat 
as a whole since 1836 (DC. Prodr.), at which time thirty-three species wer 
recognized, all but two being American. Now the genus is conceded ue 
exclusively American, and contains 10g species, more than 7° per cent ® 
which are local. The greatest display of species 1s in the uplands of cent 
and southern Mexico, where 40 per cent. of the species are endemic. se 
synopsis twenty-five new species are described.— GREENMAN (ibid. 566-57 ; 
has published some new and critical Mexican species, the new are 
bering twenty.— ELtas NELSon has published a revision of the ph ah 
western North America. He recognizes thirty-eight species, nineteen of . 
are new. The paper is a master’s thesis in the University of Wie 
is published in the ninth report of the Wyoming Agricultural Colles 
mie, Wyoming.— E. P. BICKNELL, in continuing his studies of Sis 
(Bull. Torr. Bot. Club 26:297-300. 1899), has described four new Spi : 
from Michigan.— A. A. HELLER (bed. 312-315) has described of 
new species from western North America.—- AVEN NELSON (Erythea? * 
1899) has discussed the western species of Aragal/us (Oxytropis) Or 
eight new species ; and has also described (did. 65-70) five pa 
Oreocarya, two of Cryptanthe, and one of Allocarya.—JAREP ‘Ss a 
18. Div. Agrost. U. S. Depart. Agric.) has published a synopsis of 


Sitanion, recognizing twenty-three species, twenty of which are dei 


in in 
GUIGNARD has recently studied the reduction of chromatin wo 
sucha study 


major®, This plant has proved exceptionally favorable for suc nega” 
the number of chromosomes, twelve in the sporophyte and six in! 
phyte, is the smallest yet reported for any flowering plant. al 

First division.—In the prophase the spirem splits longi a 
segments into six primary chromosomes each of which consists © shor? 
During the succeeding contraction and growth, each 


of these pieces° : 
9 fas major Aree 

: Le développement du pollen et la réduction dans le Nee 
microscopique 2 : 455-509. 1899. 


1899] CURRENT LITERATURE 145 


double row of chromatin granules, a preparation for a second splitting, so 
that the primary chromosomes are to be regarded as quadruple. As the 
primary chromosome separates into its two parts (secondary chromosomes), 
the splitting already inaugurated by the fission of the granules begins to take 
place, but is not entirely completed, since the two chromosomes remain united 
at their extremities, thus forming a V with its apex attached to the contractile 
threads of the spindle. Each daughter nucleus receives six double (second- 
ary) chromosomes. 

Second division.—In the second division, six V-shaped chromosomes 
appear. At the point of the V there is an interruption in the linin support and 
everything favors the conclusion that these are the secondary chromosomes 
of the first division which have not lost their individuality. No longitudinal 
division takes place at this time, there being merely a distribution of the two 
parts of the V-shaped double chromosome. Thus the two divisions merely 
distribute the four parts of the primary quadruple chromosome (tetrad), which 
were already defined in the prophase of the first division. It is evident that 
there can be no qualitative reduction. 

: Miss Sargant ® both figured and described a second fission of the chroma- 
z A in Lilium Martagon, and called attention to the quadruple nature 
<ce Preaek chromosome. Guignard believes that his results agree with 
neh Se of Pallavicinia, Brauer’s of Ascaris, Meves’ of Salaman- 
a different mp with Belajeff's description of Iris, although that writer has given 
cell, An €xplanation of the origin of the chromosomes in the pollen mother 
diagrams ame review of the chromosome problem, illustrated by very clear 

The ee no means the least important part of the work. he 
of Lilium hay ied results of Nawaschin and Guignard on the fertilization 
tilization the : a confirmed by Miss Sargant," who finds that during fer- 
male ile € nucleus is applied to the female nucleus, while the second 
Polar nuclei w, applied fo both the polar nuclei. In one case, in which igs 
united them ae _ yet in contact, the much elongated “ antherozoid 
Pollen tube ie : brid . In several preparations it was noted that - 
Since both pes, Pe eiization had taken place, contained two small ree 

are ies. nuclei are already accounted for, it is suggested that 
CHAMBERLATN, y due to the division of the tube nucleus.—Cuas. J. 


toa 

Ann, Bot. 17; 187-224. 1897, 
"On th ‘ 
Pa nh * Presence of two vermiform nuclei in the fertilized embryo-sac of Lilium 
roc. Royal Soc, 65 : 163-165. 189 


NEWS. 


A BIOGRAPHY and bibliography of the late Professor William Nylande 
appears in the June number of the Revue général de Botanique (rx: 218-297) 


IN AN APPENDIX to the Bud/etin of miscellaneous information issued 
the botanical department of Trinidad, the superintendent, Mr. J, Ha 
is giving a description of the West Indian and Trinidad ferns. 


Messrs. CARLTON R. BALL, of Iowa Agriculture College, Elmer D. Me | 
rell, of the University of Maine, and P. Beveridge Kennedy, of Cornell Unive 
sity, have been appointed assistants in the Division of Agrostology, U* 
Department of Agriculture. 

has be 
Bouldet,® 
unt of 


Dr. FRANCIS RAMALEY, of the University of Minnesota, 
appointed professor of biology in the University of Colorado, at 
succession to Professor John Gardiner, who has retired on acco 
tinued ill-health, having held the chair since 1889. 

THE Académie Internationale de Géographie Botanique has exe 
its international scientific medal upon Professor John M. ene 
medal is awarded by the Academy “in recognition of services “ g 
science by those who cultivate and advance its various branches. 


tive study of species and of organic variation, is announce® © ith 

& Sons, New York City. It is entitled «« Statistical methods 

reference to biological variation.” os 
PRoFEssor Dr. EpuARD JANCZEWSKI, Cracow, Austria, 's e 


study of the geographical distribution and the forms of Ribes TH" 


desires to examine the forms growing in the United States, “_ 
the var. subglandulosum. Anyone who can furnish a range OF © 
favor by © 


specimens and viable seeds of these forms will confer 4 
cating with Professor Janczewski. 
ee Fitzpatrich 


A“ MANUAL of the flowering plants of Iowa,” by *+ rivate © 
Lamoni, Ia., is announced. The publication of this work 1s 4 P 
prise, and intending purchasers should address the author. -afly de 
ing the Polypetalz, is not a mere list, but each species ce ic 
together with notes on habitat and distribution, with suitable ap . 
1899 ] 146 


1899] NEWS 147 


A SERIOUS DISEASE of the peach appeared in Michigan and other states 
several years ago and seems to be. spreading. It is known to growers as 
“little peach” for the reason that the fruit of the affected trees seldom 
attains more than one fourth normal size. The Division of Vegetable Physi- 
ology and Pathology of Washingtoh is making a study of the disease this 
year, the work being placed in charge of Mr. Merton B. Waite, who is now 
in the field. 


Mr. HERMANN von SCHRENK has been made a special agent in the 
Division of Vegetable Physiology and Pathology of Washington and author- 
ized by the Secretary of Agriculture to make such studies of the diseases of 
forest trees and timber as may be directed by the chief of the division to 
which he has been appointed. This important line of work has been much 
neglected in this country, and it is hoped that with the start made it will soon 
have the attention it deserves. 


Mr. W. A. OrToN, who has for several years been associated with Pro- 
fessor L. R. Jones of the University of Vermont, has been made an assistant 
= the Division of Vegetable Physiology and Pathology, U. S. Department of 
Agriculture, Mr. Orton will spend the greater part of the summer in South 
Carolina and other southern states, investigating the diseases of cotton. He 
= Row at James island, S. C., making some preliminary studies of a disease 
of sea island cotton which has prevailed in that section for several years. 


tees ee Bacon Evis, wife of Mr. J. B. Ellis, of Newfield, N. J., 
Whil vee ome on July 18, after a long illness, at the age of sixty-eight. 
© Mts. Ellis was not known as a botanist, it was her cooperation which 
SS eat issue by her husband of the two great sets of fungi 
ios in a orth American Fungi and Fungi Columbiani, The 3000 port- 
arranged ies 99 € sets were mounted were made by her own hands ; and she 
Mens sent out . ded the pockets for about three fourths of the 200,000 spect- 
ognition, in those books. Her labor of love certainly deserves public 
Garden, eENATIONAL conference on hybridizatién was held in Chiswick 
tural Society oe July 11 and 12, under the auspices of the Royal Horticul- 
Vinery in th “ n exhibition of hybrids was given on the first day in the large 
., .vatdens. The conference was attended by many distinguished 
ists. Mr. H. J. Webber was the delegate from this 
'€ aid of lantern illustrations he described the hybridization 
Timent of A . Mr. Swingle are prosecuting under the charge of the 
ble to be = Professor L. H. Bailey sent a paper, but was 
. ht in person, 


Tue Y 
botany uae CORPORATION, dt a recent meeting, voted that the chair of 
¥the late Professor Daniel C. Eaton until his death should 
* 


148 BOTANICAL GAZETTE [av 


hereafter be known as the Eaton Professorship of Botany. It may 
recalled that this professorship was founded and endowed bya relative and 
by friends of Professor Eaton in the year 1864, although it has never bet 
distinguished by a name. . The extensive and valuable botanical library and 
herbarium which Eaton accumulated have been donated to the university by 
his family, and Mrs. Eaton has placed in the botanical laboratories a bromt 
tablet to her husband’s memory, and, in addition, has founded. a gradualt 
scholarship in botany.—£rythea, for August. 


PROFESSOR BESSEY has been called upon to serve as the Acting Chance 


will make it necessary for him to be relieved of mu 
in the laboratories and lecture room, the regents have elected Dr. August 
Rimbach (Ph.D., Jena 1887) ad interim Instructor in vegetable physiol) 
and pathology. Dr. Rimbach is known as the author of many board 
papers in the Berichte der deutschen botanischen Gesellschaft and other Ger 
man botanical journals. He was a pupil of Stahl, Detmer, 
and Sachs, and was Professor of Botany in the University of Cuenca, 
dor, from 1889 to 1894. He has traveled extensively in South Amen® 
engaged in the study of flora of tropical and alpine regions. 


Mr. J. N. Rose has just returned from a three months’ trip t 
Mexico, bringing about goo species of dried plants, many living i“ 
cially agaves, and plant photographs. His collection is not so 1atg® 
rich in new species as the one of 1897, but it will doubtless help ue 
many puzzles which have long worried botanists dealing with Mexical 
Besides rediscovering Echinocactus Parryt, he collected several , 
lost or hitherto unknown to American herbaria. About 200 


Calientes, Jalapa, City of Mexico, Oaxaca, Cuernavaca, Pachut 
Monte, Guadalajara, Tequila, San Luis Potosi, an . 
from which many types have beentaken. Mr. Rose also made : 


Mr. Rose was accompanied by Dr. Walter 
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Vol. XXVIII SEPTEMBER 1899 No. 3 


THE 


BOTANICAL GAZETTE 


EDITORS 3 

JOHN M. COULTER, 7ic University of Chicago, Chicago, Til. 
CHARLES R. BARNES, The University of Chicago, Chicago, Il. 

J. C. ARTHUR, Purdue University, Lafayette, Ind. 


ASSOCIATE EDITORS 
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Geneva 


BR University of Bonn 
DeTONI VOLNEY M. SPALDING 
ADOLF ENGLER ROLAND THAXTER 
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| age de Pharmacie, Paris Missouri Botanical Garden 
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CONTENTS 


COMPOUND OOSPHERE OF ALBUGO BLITI. ContTRIBUTION FROM THE HULL 
Botanica LABORATORY. XVI. (WITH PLATES xI-xv.) F. L. Stevens - 149 


L fs. “cepa OF THE SUGAR BEET. Daten PLATES cae on Clara A. 


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MON OF THE NORTH AMERICAN SPECIES OF TEPHROSIA. B.L. Robinson- 193 
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VOLUME XXVIII NUMBER 3 


SOTANICAL GAZETTE 


SEPTEMBER 1899 


THE COMPOUND OOSPHERE OF ALBUGO BLITI. 


CONTRIBUTION FROM THE HULL BOTANICAL LABORA- 
: ZORY. XVI 

F. L. STEVENS. 
(WITH PLATES XI-Xv) 
7 Tie Tesults of this investi 
in their essential features 
Processes connected with 


gation introduce factors entirely 
and at variance with the cytologi- 
fertilization previously described 
» enocytic or other forms of plants or animals. The title 
: ntradiction of terms. The writer 
ession compound oosphere, but it 
al at least for the present, to modify the term 


Rew term indicating the conditions and the process 

2 ation Presented in this paper. | 

: ™pound Sosphere is one containing several or many fune- 

Sexual nuclei. This idea violates the present conception of 

cell as it exists throughout the plant king- 

€, So far as the writer is able to judge, the 
149 


“Structure of that 
aod furthermor 


150 BOTANICAL GAZETTE [SEPTEMBER 


ova of animals present no such peculiarity. If there is any 
character that defines the oosphere and ovum it appears tobe the 
presence in the cytoplasm of a single female nucleus that is nor 
mally fertilized by a single male nucleus. In Albugo Bliti Biv, 
however, the mature oosphere contains many female nuclei, aid 
fertilization is effected by the discharge of many male nuclei from 
the antheridial tube and their subsequent fusion with the female 
nuclei in pairs. An oospore results from this multiple sexta 
act with about one hundred fusion nuclei, which remain in the 
resting condition until germination. The existence of such cot 
ditions must be supported by strong evidence, and great cau 
should be exercised in interpreting the data upon which the 
conclusions are based. 
One is partially prepared, however, for the acceptance of such 
conditions as these by the thought that the form under conside 
ation isa coenocyte, and that comparatively little is known 
the behavior of the nuclei and cytoplasm in such structures , 
Excellent summaries of current knowledge are given by Hus 
phrey (’92), Zimmerman (96) ,and Wager (’96). It is os 
Sary, in view of the existence of these accounts, to enter eo 
details here. Suffice it to say that in several (Monoblepha 
dinacee, Entomophthoracez, and Chytridinez) of the gees 
coenocytic groups the behavior of the nuclei in fertilization” 
practically unknown. In those groups of which there is mF 
knowledge (Saprolegniacez, Peronosporee, Zygomycetess For | 
Siphonez) concordant results have not yet been attained ae 
example, in the Saprolegniacee the question is still in ; 
whether or not fertilization occurs (Hartog ’95, Trow 95): 
the Siphonez the two most comprehensive papers (Belint 
Oltmanns ’95) upon the one genus (Vaucheria) that 
investigated disagree essentially as to the events leading = 
development of the oosphere. The process of fertilizatio” 
cribed for the Zygomycetes by Léger (’95) involves the 
phenomenon of the fusion of nuclear complexes. 
The pioneer work on the histology of the Peronospor 
by Wager on Peronospora parasitica in 1890, was follow” 


1899] THE COMPOUND OOSPHERE OF ALBUGO BLITI 151 


article by the same author in 1896 on Albugo candidus. Berlese 
in 1898 published an article on the entire group. It appears that 
most of the study of the histology of coenocytic fungi has been 
concerned with the Peronosporez, but even here the data are yet 
too scanty to admit of any wide generalizations. The improve- 
ments in technique during recent years should be held in mind, 
as much of the discrepancy between the earlier and later results 
may be thus explained. 

Wager’s work (796) should be consulted for a comprehensive 
summary of the knowledge up to the time when he published the 
results of his investigations. For A/bugo candidus his own research 
shows a condition where the antheridial tube liberates one sperm 
nucleus which fuses with the solitary female nucleus in the 
Soplasm. He employed corrosive sublimate in saturated aqueous 
solution as a fixing agent, and stained the sections with Hartog’s 
mgrosin-carmine. In his earlier work on Peronospora Wager 
describes a multinucleate oogonium and antheridium. The nuclei 
. “ maturing oogonium pass to the periphery, where they 
divide mitotically. Two or three then return to the center and 
Probably fuse, as only one nucleus is found there at a later stage. 
The antheridial nuclei divide simultaneously with those of the 
®ogonium. The antheridi 


th al tube contains one or more nuclei, 
© antheridium finall 


stage. Thi aed having many less than it had at an earlier 
“eae “ soe was fixed by either absolute alcohol or 
mic acid, and stained with Kleinenberg’s haematoxylin. 
ogy Sives four figures to illustrate the development and 
=: cathe - Por tulace, but his description of the nuclear 
Albugo se ‘on 1s not illustrated. All statements concerning 
tj m to have been based on either this species or herba- 
of ai of other species. He studied also four species 
alcoho]. re Fora killing agent he used either 95 per cent. 
’ alcoholic corrosive sublimate, Flemming’s solution, or 
: ined with F lemming’s triple stain or Hartog’s 


armine, Further reference will be made to these arti- 
In the ape 
a ' p r. 


Csults Presented in this paper were obtained mainly 


152 BOTANICAL GAZETTE [ SEPTEMBER 


from material fixed in chrom-acetic acid, cut in serial sections i 
paraffine, and stained on the slide by Flemming’s triple stain 
For full details regarding methods the reader is referred to the 
end of the paper. This investigation was begun in 1897, om 
year being spent in the botanical laboratory of the Ohio State 
University through the kindness of Dr. W. A. Kellerman. Lat 
indebted to Mr. J. H. Schaffner for many courtesies during my 
work at the same institution. Since the summer of 1898 tht 
study has been continued under the direction of Dr. Bradley 
Moore Davis in the Hull Botanical laboratory of the University 
of Chicago, where I have also received helpful advice and a) 
gestions from Dr. J. M. Coulter and the members of the bola 
cal staff. I wish to express my thanks to my wife for much kind 
assistance, in particular for the preparation of Plate XV. 


4 


The character of the mycelium of Albugo varies with e 
nature of the host tissue. The hyphae are slender where tt 
cells of the host are thick-walled and placed close togethe | 
while in loose tissue they may swell to a considerable diamete 
fig. 42 shows the general structure of the hyphae, the roundit 
nuclei, each with a prominent nucleolus and nena. 
distributed irregularly through the vacuolate cytoplasm. * 
single nucleus is represented in fig. rz. It is worthy of atten , 
principally because of its very faint linin network. It has® 
actual diameter of from 2~2. 5M. 

The oogonium may be terminal or intercalary, its 
simply the greatly expanded mycelial wall, as is ev! 
the frequent persistence of haustoria on its surface. ee 
early stages of developing oogonia may be distinguish i 
enlarged mycelia by certain peculiarities of the protoplas® fe 
43.) The nuclei are elongated, the vacuoles are angular al | 
torted, and the cytoplasm is drawn into stringy bands; 
which gives evidence of a disturbance not present in the 0 
mycelium. These peculiarities are frequently evident © 
tative hyphae a distance of 200 from the developing ° 


DEVELOPMENT OF THE OOGONIUM AND ANTHERIDIUM. 


denced 


& 


1899] THE COMPOUND OOSPHERE OF ALBUGO BLITI 153 


Similar appearances were noted by Istvanffi (’95) and Wager 
(96), and their explanation is undoubtedly the true one, namely, 
that the protoplasm was rapidly flowing from the mycelium to 
fll the enlarging oogonium. When sufficient nucleated proto- 
plasm has entered the developing oogonium, this structure is cut 
off from the hyphae by a septum at the point of enlargement 
(fig. 44). The oogonium is now fully differentiated from the 
vegetative hyphae, the nuclei recover their original form and lie 
ina coarsely vacuolate and somewhat granular cytoplasm. The 
general appearance of the oogonium and its contents may be 
seen in fig. 45. 
. The antheridium develops simultaneously with the oogo- 
uum, but gives no evidence of the flowing of the protoplasm 
— the growing structure. Probably owing to the small size of 
this organ there is but little disturbance as it fills with proto- 
plasm. It becomes cut off from the parent hypha and the con- 
tents are similar in appearance to those of the oogonium as is 
shown in Fig. 45. 
The most conspicuous feature in this early development of 
eae is the increase in the size of the nuclei. This 
Sama Somewhat rapidly just before the oogonium has 
ies full size. As the nuclei grow larger the linin net- 
©omes much more prominent, until finally it assumes a 


Ver sae 
es characteristic structure in the form of large meshes, the 
teads bein 


the 


eee of early mitosis. While the nuclei are pass- 

the oogonium wall thickens slightly. 

: that the number of nuclei may be deter- 

the ‘gags oe accurately. The count ranged from 226 

ce :.. number being quite exceptional. Making 
€ fact that several nuclei may readily have been 


Min 
1 333, 
allowan 


154 BOTANICAL GAZETTE | SEPTEMBER 


counted twice in adjacent sections, it would perhaps be fairto 
place the average at about 250. Wager found 115 in A. candidu, 
and Berlese 200 in A. Portulace. It is very difficult to makean _ 
accurate count in the antheridium, because this structure isso — 
small and of such irregular shape that it is impossible to recog: 
nize its limits in adjacent sections. However, an average based 
on several counts indicates that the number is likely to be about 
35. This number is considerably greater than that suggested 
by either Wager or Berlese, both of whom report about 12 mud 
in the antheridium. : 
When the oogonial nuclei are passing into the spirem cont! 

tion the Hautschicht seems to be closely appressed to the wall 
the vicinity of the antheridium. This fact is demonstrated mo 
clearly in preparations where there has been slight collapse 0! 
the contents of the oogonium ( figs. 47, 48) and the protoplasm has ; 
shrunken away from the wall everywhere excepting at the pois | 
opposite the antheridium. This adherence of the Hautschicht 
correlated with a very marked granulation of the cytoplasm # 
this region, a phenomenon also noted by Wager, and one whit 
seems to be significant. It suggests that a cellulose enn 
is secreted to dissolve the wall of the oogonium. As | i 
cated in figs. 48, 49, 50, this wall frequently shows the marks © : 
corrosion over a considerable area, always at a point oppos | 
the antheridium. This interesting process results in 4 neat ‘ ; 
foration, through which the cytoplasm of the oogonium flows a 
as to form a very conspicuous swollen papilla within the ” 4 
idium. Various stages in this process are shown in jigs — 
It is difficult to explain this phenomenon. The initiatory — 

the perforation of the oogonium wall seems without ag 
taken by the protoplasm of the oogonium itself. But oy 
the significance of the pushing of the cytoplasm of the ee 7 
into the antheridium to form the peculiar bubble-like papi 2 
The structure, both wall and contents, stains deeply, thus a 
ing very conspicuous, while its extremely frequent -« ue 
as well as its presence in other species, seems to in eer 

is not abnormal. The papilla wall is so extremely thin : 


1899] THE COMPOUND OOSPHERE OF ALBUGO BLITI 155 


may only conjecture whether or not it is derived from the anther- 
idium wall, The history of the papilla beyond the stage shown 
in fig. 54 is not clear. In this condition the structure is very 
thin-walled and its contents vacuolate, resembling a large irreg- 
ular compound bubble. Whether the delicate wall now bursts or 
the contents are gradually withdrawn into the oogonium is uncer- 
tain. At all events, the papilla in later stages leaves no trace of 
its former existence. There follows at a later period conditions 
(figs. 56, 57) which show that there is certainly a movement of 
the cytoplasm in the opposite direction, the antheridium extend- 
ing a process with a cell wall through the opening into the oogo- 
tum. The first surge of the cytoplasm from the oogonium into 
the antheridium may be due simply to the unequal conditions of 
turgor in the two structures, but it is possible that there is also a 
phylogenetic significance in the phenomenon. The occurrence 
of similar structures in A. candidus and A. Portulace shows it to 
te of some import. Such a papilla in a much less highly devel- 
“a form is figured by Wager and referred to as the receptive 

pila. 

The antheridial tube presses into the oogonium in the form, 
aa thin-walled process (figs. 56,57). It is filled with 
ie aaa that greedily absorbs and retains stain, and is 
rina‘ per y @ sheath of dense oogonial cytoplasm. The nuclei 
ae = ag antheridium, none entering the tube at this time, 

ie indistinguishable in size and structure from those in the 

‘wpa They are also in the spirem condition, similar to 

Scribed for the oogonial nuclei, The description of the 
further develo 6 ae Z foll 

Oh toa pent of the antheridial tube is deferred, to follow 

of the differentiation of the compound oosphere. 


D 
'FFERENTIATION OF THE COMPOUND OOSPHERE. 


The : : 
Previous description carries the history of the sex organs 


up to ‘ 
ody time when the antheridial tube has penetrated the 
Cor, l m one fourth or one fifth the diameter of that structure. 


related wit 


°Ccurs the jog the further development of the tube there 


erentiation of the periplasm and ooplasm, and the 


156 BOTANICAL GAZETTE (SEPTEMBER 


extrusion of the nuclei from the central region of the oogonium 
The process consists essentially in a centripetal movement ol 
the cytoplasm, and results in a massing of this cytoplasm inthe 
center of the oogonium in such a manner that the vacuoles and 
nuclei are carried to the periphery of the denser central portion 
thus developed. Behrens (’g90, 315) describes a somewhat similar 
condition in Vaucheria as follows: ‘‘ Der ganze Vorgang bestelt 
also in der Abldésung des gréssten Theils der Protoplasten von det 
Wand durch Vacuolisation der wandstandigen Plasmaschichtea.” 
This curious phenomenon was noted by Wager in A. candidit 
and subsequently by Berlese in A. Portulace. The process ® 
heretofore described is simple. In A. Bliti it is complex, bi : 
unique and full of interest; and as a complete knowledge * 
essential to an understanding of the further development of the | 
oosphere a detailed description must be given. 2 
The first hint of the centripetal aggregation is found al 
tendency of the cytoplasm to depart from the even distributio® 
shown by a young oogonium; and to collect in masses through 
out the interior (fig. 58). These denser portions run togethes 
forming fewer but larger masses (fig. 59). Thus several eee 
nent aggregations of cytoplasm may be formed, separated ins 
one another and from the wall by vacuoles of varying sizes VS 
60). These denser regions are homogeneous in structure, 
taining minute vacuoles of uniform size evenly distributed * 
matrix of cytoplasm free from granules. The dense egies 
tain no nuclei, because these are forced from the dense cyt 
to a position on its periphery. The dense centers — ove 
forcing out the vacuoles. This may result immediately 
condition shown in fig.67, but frequently the coalescence . : 
more slowly and irregularly, and often a reniform mass 1 ge 
the indentation on one side marking the juncture yet % ee 
The last gap narrows until only a few vacuoles pera ee 
its track (fig. 67), and these soon float outward sige is 
of cytoplasm, the rudimentary oosphere (jigs. 62, “ _ 
the vacuoles pass outward they often leave captive a ce 
wake (figs. 61, 62, 64), but these soon follow. A typ! 


1899] THE COMPOUND OOSPHERE OF ALBUGO BLITI 157 


of the resulting condition is presented in fig. 64. The outer region 
ofthe oogonium, the rudimentary periplasm, is coarsely. vacuo- 
late, presenting a conspicuous contrast to the dense central mass. 
Along the boundary between these two regions are gathered most 
oi the nuclei (fig. 6g). 
The next stage in the differentiation of the oosphere is. con- 
spicuous and clearly characterized. It ends in producing a dis- 
tinct differentiation between the oosphere and the periplasm 
“fg. 65). This condition is brought about primarily by the mar- 
shaling of the nuclei into an oval or an irregular hollow sphere; 
a section of which is shown in jig. 65, while a somewhat earlier 
Stage is to be seen in Jig. 64. Both figures illustrate the one 
important fact that all or nearly al] of the nuclei are at the 
boundary of the central dense mass. The latter figure in addition 
shows that there is a sharp line of demarcation between the 
3 lasm and periplasm. There are usually a few scattered nuclei 
1 the periplasm, and occasionally one finds a nucleus in the 
Sosphere that has not passed out as rapidly as the others. 
Important changes occur in the cytoplasm while the nuclei are 
‘vanging themselves into a hollow sphere. At the beginning of 
this process the region that is to become periplasm is coarsely 
“chap in marked contrast to the dense cytoplasm of the 
meet ut the two regions blend gradually together where they 
(Jigs. 62, 64). Later when the hollow sphere of nuclei 
ome ty regular in outline dense granular cytoplasm ig 
ie : gap and between the nuclei (figs. 65, 68, 69). 
differentiated : er of the rudimentary periplasm also becomes 
region of th peo a film more densely granulated than any other 
sphere oC and finally determines the limit of the 
Critical ae fre is not yet an organized wall, and the most 
: y Teveals nothing more than a dense film of proto- 
7oNation, * 1S convenient to call this condition the stage of 
The Position 
Vents lead 


ading to 
"RQ this ¢ 


of the stage of zonation in the sequence of 
_to the differentiation of the oosphere is clear. 
endition comes the characteristic and sharp limitation 


158 BOTANICAL GAZETTE [SEPTEMBER 


between ooplasm and periplasm which is maintained until 
maturity, while before zonation such a differentiation did not 
exist. The process of differentiation is gradual, and a seriesol 
developmental stages has been obtained which seems complete. 
This period is the only one where the ooplasm possesses very 
few nuclei or none, and it is impossible to regard it as beinga 
period later than stages which present zonation and also contail 
50-100 nuclei (figs. 68, 69, 70). The development of the 
antheridial tube is such as to lend strongest support to tht 
sequence above indicated, since the tube is shorter in stages p= 
ceding zonation and longer in stages following it (see plates), 
thus affording strong corroborative evidence. While the differ | 
entiating line is characteristic of zonation the paucity of nucle 
in the ooplasm is equally so. The sharper the differentiatio® 
the fewer the nuclei, and when zonation was very definite ese 
could be found, and it is probable that when this stage is at its 
highest development there are no nuclei in the ooplasm. There 
is some evidence, however, that makes it seem possible that o7¢ 
and even two spindles sometimes remain in the ooplasm, but ths 
is uncertain. a 

No mention has been made, as yet, of the division of the nucle 
of the oogonium. This mitosis closely accompanies the proces 
of zonationas is indicated in most of the figures. These two even 
apparently take place nearly simultaneously. The oe 


60, 6r. At the time of complete zonation t 
metaphase and lie close to the line that separat 
from the periplasm (figs. 65, 66). Spindles are ae 
that actually cross this line at right angles, so that one bs, 
in the ooplasm and the other in the periplasm (78: A a 
The mitoses that take place at this period mark an see ? 
characteristic phase in the history of the 00g 0h an 
dividing nuclei that lie tangential to or wholly a : 
boundary line between the ooplasm and periplae | 
oe : es 
daughter nuclei in the periplasm. Each of t his the oosph® : 


ntly! 


cross the line (fig. 66) gives one daughter nucleus 


1899] THE COMPOUND OOSPHERE OF ALBUGO BLITI 159 


and the other to the periplasm, and the line of differentiation is 
sharply defined and unmistakable. Nuclei may be observed in 
every phase of this mitosis, and the daughter nuclei may be 
found in all stages of reorganization, one of each pair in the 
ooplasm the other in the periplasm. The writer was unable to 
detect any difference between the mitoses that occur strictly in 
the periplasm and those that contribute daughter nuclei to the 
oosphere. Asa result of the division a large number of nuclei 
pass into the ooplasm, thus producing a multinucleate cell con- 
taining by actual count an average of 45 to 55 nuclei (not 
- than 40 or more than 60). The oosphere is thus a coenocyte; 
instead of the uninucleate cell which one would expect there is 
found a multinucleate structure, to designate which the writer 
has used the term compound oosphere. 

Because of the importance of the anomalous compound 
Sosphere and the peculiarities of its development, it seems best 
to discuss at length difficulties that might be suggested. It may 
be claimed that amistake has been made in the sequence of 
Sree, and that the multinucleate condition of the central 
“Bion does not follow, but precedes zonation. This objection 
aoa for four reasons: (1) the sequence is complete up to 
saa there is no place for a multinucleate central region 
white va ? the studies in cytology make it certain ee sa 
Phase of the Sealine ie i is ponies eae 
thus renderin as an in igs of the age o 8 ‘ 
ble: £3). § any misunderstanding of the sequence imposs 
standing of ig ctation Picea is necessary to the “— 
(4) the _. that are positively and unmistakably older; 

opment of the antheridial tube and central body 

with the views presented. It would seem impossible 
Stake in seriation has been made. It may be claimed 
™ sity, os nuclei seen in the mature oosphere are not 
idence to . Salt = by the knife. There is abundant 
- «Often die in ophaibds this claim, for the nuclei in west 
Must have be : is no that, if carried in by the knife, they 
©n carried in two opposite directions by the same 


ta mi 
“lat the 


160 BOTANICAL GAZETTE [SEPTEMBER 


stroke. It may be well to state that specimens of every structure 
or stage represented or described in this article are preserved, 
not in one, but in several mounts, and for most of the important 
stages in many preparations. No isolated or single fact is any- 
where used either to support or destroy any theory. 

With the entrance of the nuclei differentiation of the oosphere 
is complete. The ooplasm is of very fine, even texture, made 
up of such small meshes that the vacuoles are never more that 
half the size of the nuclei, and there are no prominent granules 
or oil drops shown by the Flemming stains at this time (figs.6e 
69). The periplasm is loosely vacuolate, the strands are often 
granular, and the nuclei are frequently at their intersection, — 
often in bunches. : 

The nuclei of the antheridium usually divide simultaneously 
with those of the oogonium, this being so constantly true see 
from a glimpse of one organ the condition of the other could 
be predicted. No difference between mitosis in the oogontum 
and in the antheridium could be observed. At the time of 
differentiation of the oosphere the contents of the anther! 
tube stain deeply, but it has not been possible to demonstrate 
nuclei in its interior at this age. When the nuclei line up P* 
paratory to zonation the antheridial tube has usually peneue® 
the periplasm almost to the outer boundary of the oosphe ; 
and as it later pushes into the oosphere, during the ae 
of the mitosis, it somewhat indents the boundary film. © 
oogonium was seen that had two antheridial tubes pe ge 
the periplasm from opposite sides, but neither had yet Pl 
the boundary of the oosphere. Another stage Wa " 
where an unemptied tube lay tangential to the 005 
apparently the unfavored of two competitors. Still 
presented a stage of fertilization in which two tubes wee 
taneously opening into one oosphere. 

Minute globules of brownish color are to 
cytoplasm of the oogonium from its earliest devel 
zonation. In all early stages they are indefinite ! 
irregular in size, but seldom larger than one © 


! 


be seen it 
lopment up 
n num . 
f the ® 


1899] THE COMPOUND OOSPHERE OF ALBUGO BLITI 161 


When the ooplasm is first differentiated it possesses a small num- 
ber of these globules which are irregularly distributed. At this 
time and in earlier conditions there is nothing to indicate that 
these structures have any peculiar significance. In their color 
and form they suggest minute drops of some oil-like substance 
that has remained undissolved under all the treatment experi- 
enced by the preparation. However, at the time of zonation, 
and in certain later stages, there is only one such dropor globule, 
where there were previously several, and that one is always in 
the geometrical center of the oosphere, and is surrounded by a 
differentiated region of cytoplasm (figs. 69-71, 74). Whether 
®r not this one central globule, which is a constant feature of the 
oosphere from zonation until just before fertilization, is developed 
“of a fusion of the drops previously present could not be deter- 
mined with certainty. This is strongly suggested, however, by 
appearances like those noted in jig. 75, which was drawn from a 
young oosphere, and seems to indicate that several minute drops 


Were fusing to form the central globule. 


Asther AOE ae : ‘ 4 


QGAtuUuIcys 


ot Surrounding it becomes more dense, although still 
a away gradually toward the outside (figs. 47, 69, 71, 74)- 
ble “eaty observed this globule was surrounded by the remarka- 
in . a dense cytoplasm which differs from ordinary ooplasm 
nuclej Naa more darkly and contains fewer vacuoles and 
develo nen ne whole central structure is at its maximum 
(figs . It is a very conspicuous object within the oosphere 
of —- a There occurs also further slight differentiation 
the — oplasm in the immediate vicinity of the globule where 
inner re taken more faintly or has a yellowish tint. This 
(fig. 69) ve also shows a radiate structure under the low power 
in drawin ut the highest magnification, such as that employed 
nite “ig fg. 77 (3300 diameters), failed to demonstrate defi- 
Oosphere - ‘8 77 shows in detail this peculiar region of the 
Tegion ga is the vacuolate ooplasm, then comes the 
“nse cytoplasm with few vacuoles, and finally in the 


162 BOTANICAL GAZETTE [SEPTEMBER 


center is the opaque globule immediately surrounded by « 
lightly stained zone. After reaching the condition of maximum 
development the structure rapidly loses character, disap 
pearing entirely just before the antheridial tube discharges its 
contents. 
A summary of the history of the central structure may be 
given as follows: it first appears in zonation, and reaches is 
maximum development when the daughter nuclei of the fis! 
mitosis pass into the oosphere; after that it rapidly degeneratts 
although traces of its presence sometimes persist nearly to the 
time of fertilization. A body of apparently similar nature ¥® ] 
mentioned by Wager as occurring in the oosphere of A. candids 
and my own as yet incomplete observations on that species indi | 
cate that the body seen in A. candidus and the central globule o | 
A. Bliti are homologous structures, although they differ much 0 
certain details. I find also a structure very like the one above 
described in the oosphere of A. 7; ragopogonts and A. Portulat 
and believe that we have here an organ of the oosphere, perhaps 4 
regularly present in the whole genus Albugo, if not ™ the | 
oospheres of other related genera. It appears with such co 
stancy at certain important stages in the life history of the spe 
cies, and passes through such a definite course of developmet : 
that its presence seems to be of importance. May it not “ 
It should be called to mind in this connection that Dango : 
noted in each of the numerous oospheres, in certain eet : 
the Saprolegniaceez and Peronsporee, a cent 
appeared just before fertilization. Various interpre 
been given to such structures by different writers, early fs 
ers mistaking it for a nucleus. Dangeard supposed - “ 
but Wager thought that Dangeard was probably mistaken 
that the structure is truly a central body such a8 95 
found in A. candidus. The descriptions and time of at 
make it seem quite possible that the body noted in the o 
g of the Society 


*MR. SWINGLE expressed such views at the meetin 
Morphology and Physiology at Ithaca, December 1897- 


1899] THE COMPOUND OOSPHERE OF ALBUGO BLITI 163 


legniacee may be homologized with the central globule of the 
oosphere of Albugo. 
It is premature to discuss the function of this body until its 
relations in other species have been closely studied. As described 
by Wager it appears to be intimately connected with the behav- 
ior of the sexual nuclei; but there appears to be no such rela- 
tion in A. Buitt, where its function is perhaps that of an organizer 
in the oosphere. The body first appears when the oosphere is 
well defined and is most highly differentiated during the entrance 
of the daughter nuclei. At this time also occurs the formation of 
a thin film of denser protoplasm which definitely bounds the 
oosphere. A great dynamic change occurs in the oogonium 
When the ooplasm and periplasm are differentiated and the zone 
of cytoplasm separating the two regions is formed, and in addi- 
tion there is that remarkable division of the nuclei in such a 
manner that approximately fifty daughter nuclei are always cast 
into the ooplasm. Simultaneous with these activities, existing 
when they are at their maximum and disappearing when they 
®ease, there is formed this peculiar structure, which is so definite 
ne character and so constantly present that it seems to have some 
functional importance, 

In view of 
logical 
the wri 


the morphological character and possible physio- 
Value of this central structure to the coenocytic oosphere 
is bate . tures. to abel deus for it the name ce@nocentrum. It 
the iad ere to emphasize the difference in structure between 
Nucleus ea and a nucleus. The globule is distinctly not a 
Stains diffe it is much smaller than any nuclei that were seen, 
i » and is structureless and unchanging. The 
Since these " eer be a nucleus with the globule as a nucleolus, 
OF its [. fail to have the internal structure of a nucleus 

‘"8 membrane. There is no definite demarcation 


from ¢ : 
71,74, rounding ooplasm, as is plainly shown in figs. 47, 


In os 
— 75 other Structures are shown, the nature of which is 
uted in ik They are small granules which lie profusely distrib- 
“ cytoplasmic Strands in all ages of the oogonium, but 


he Sur 
75: 


. 


164 BOTANICAL GAZETTE [SEPTEMBER 


are particularly noticeable in the fine meshed area of the 
oosphere. They do not appear when stained in the ordinary 
way by Flemming’s triple stain, but seem always present and com 
spicuous if stained by Heidenhain’s hematoxylin, when they 
are black ( fig. 75). A hematoxylin preparation when bleacheé 
and restained for a long time in safranin shows numerous rou 
red bodies in the same position. In appearance they are a trifle 
longer than broad and often double, as though two were lying 
end to end, reminding one of large bacilli. It does not seem 
probable that these coalesce to form the central dot, as might 
be suggested by fig. 75, since they have a different reaction t0 


stain. 


SIMULTANEOUS MITOSES IN THE OOGONIUM. 


During the differentiation of the oosphere the nuclei in the 
oogonium divide once ( figs. 58-62, 65-67), the mitosis occurilg | 
about simultaneously for all of the nuclei, cases of independes 
division of single nuclei never being found. Almost invariably # 
oogonium in the condition of zonation presents the nucle! # 
metaphase, or just passing into anaphase ( fig. 65). In ee 
stages, just before zonation, when the cytoplasm is massed we 
one or several centers, the nuclei are usually in prophase Ve 
59-62), but metaphase may be present even as early * i. 
beginning of this process (fig. 58). It is apparent that anaes 
is never reached until after zonation, and that the mitoses eA 
when the cytoplasm commences to collect in mae 
nuclear division always occurs when the antheridial “ te 
the position shown in fig. 59. It would seem, ede sat 
differentiation of the oosphere would take place rapidly 1 . 
consists merely in a floating out of the vacuoles ant Sa 
the interior region of the oogonium, but mitosis 15 Pe 4 


less rapid. If changed conditions should hasten 0! ss 8 
process of zonation, one would expect a variation oe sah 
exist in the time correlations between zonation and the 


The spirem condition of the nucleus has been ma 


the other stages of nuclear division have not been 


1899] THE COMPOUND OOSPHERE OF ALBUGO BLITI 165 


detail. Preparatory to the formation of the spindle the nucleus 
elongates, its membrane being pulled out in two directions, while 
the chromatin collects in minute globules in the linin thread. 
These are at first irregular in size, but gradually become less 
ftumerous and more uniform, presumably fusing with one another 
until a small number of nearly spherical bodies, the chromosomes, 
are present in the nucleus. /%g. g shows a condition with the 
chromatin granules and the linin network still evident, while in 
fig. § the linin strands have almost disappeared. As the chromo- 
somes perfect their organization they approach the equator of 
the now elongated nucleus (fig. 6), and there appear at the 
poles two round bodies which lie within the nuclear membrane, 
These bodies stain red with Flemming’s triple stain and are 
Constantly present at this period of the prophase. Although 
not observed earlier they persist and become more prominent 
m the later stages. The spindle fibers first appear a. little 
later at the poles of the elongated nucleus, from whence they 
seem to grow toward the equator. They are entirely intra- 
— and there is a distinct space between them and the 
larly one (fig. 9). The chromosomes are at first irreg- 
achromat; sh throughout the equatorial region, but when the 
themselves een becomes more distinctly developed they group 
somes are i nuclear plate and divide. As fe chromo; 
determine on Spherical and very small it was impossible to 
Fests in a cl : oral ot their division. The mature spindle 
with the coma distinctly inside of the nuclear membrane, 
Must be # a odies very definitely outlined. That these bodies 
at certain & : pee eats is evident from their constancy 
anaphase P eg m the mitosis, e. g., from late prophase to late 
Auclear is intranuclear, it is not surprising that exttar 
tions Present ae pve be absent, and in fact the only radia- 
0 have been © the spindle fibers. These structures seem not 
Previously described for this group of fungi. 
a. show a condition very commonly seen. ee 
at the cente eS prominent, the chromosomes are masse 
",and the spindle fibers are very slightly or not at all 


igs, 
“clear 


166 BOTANICAL GAZETTE [ SEPTEMBER 


differentiated. The dark strand shown in the lower half of fp — 
8 is probably like the less conspicuous one in a similar position 
in fig. 5. Both may be considered as the remains of a spitem 
thread, such as is shown in fig. g. ig. 7 is open to a similat 
explanation. The difference in the shape of these two figuresis 
noteworthy, since it is probably due to their position in the 
oogonium. The spindle shown in fig. 8 lay in a strand of pet | 
plasm which supported an oosphere, similar to the strands shows 
in fig. 62. It is probable that the length of the spindle is due 
to the tension to which it was subjected. Fig. 7 was from’ 
crowded bunch of nuclei, and could not elongate. The apps 
ance shown in this figure might tend to support the idea that the 
spindle fibers are formed from the linin thread, a view enles 
tained by Wager, but disputed by Berlese. The question p© 
sents so many difficulties that the writer does not feel warrante! 
in expressing an opinion. 

The nucleolus at the time of late propha eee 
small, but often quite as large as when the nucleus 1s | 
spirem condition. It may be found throughout all stages of 
mitoses. ig. 12 shows the splitting of the chromo ; 
jig. 13 may be recognized as a condition immediately ee | 
membrane is still intact and encloses the nucleolus which — 
outside of the spindle, and the centrosomes are at their maxinil® | 
definiteness. It is interesting to note in passing a 
nuclei lying very near to the antheridial tube are usually on 
a full phase in advance of other oogonial nuclei in ess 

ar * 
fact strikingly apparent when the majority of the nuclel c 
metaphase. 

The chromosomes, after the division of th 
move poleward with unequal rapidity, the poles 
character, and the nuclear membrane is no 10 
boundary of the nucleus being marked by the Sf pul 
(fig. 14). With the loss of the membrane the wed 5 
structure assumes, and retains through later stage pe 5 | 
of staining more darkly, a character particularly aah 
the regions where the chromosomes lie ( figs. aS ee . 


se is sometime 


spindle a 


1899] THE COMPOUND OOSPHERE OF ALBUGO BLITI 167 


The nucleolus may travel poleward with one group of chro- 
mosomes, or break into two, either at this time or earlier, thus 
allowing a small nucleolus to go to either pole. It is easily dis- 
tinguished from the chromosomes by stain reaction and usually 
also by its size. At this stage there is at each apex a round 
body of the size and shape of the centrosomes, but scarcely dis 
tinguishable from the chromosomes except through position, 
Even in very late anaphase faint fibers may be seen connecting 
the daughter nuclei (jigs. 15, 16). When the chromosomes 
reach the end of the spindle they become indistinguishably min- 
gled and massed (fig. 76), but the nucleolus often stands out 
very distinctly by virtue of its color and size. 
fter the two groups of chromosomes are sufficiently sepa- 
rated the spindle fibers collapse in the middle ( fig. 78), and the 
daughter nuclei become distinctly organized. Each rounds off 
and contains a dark somewhat crescent-shaped mass of chroma- 
tin on the side that is turned away from its sister nucleus. This 
condition is often very noticeable in the differentiated oosphere 
when several daughter nuclei may be observed, each with its 
dark half centerward. The explanation of this condition is not 
far to seek, The sister nuclei lie in the periplasm with their 
red outward, plainly showing that the former 
€ lay across the line that separated the ooplasm 
of the oo: a These conditions present strong evidence 
ria of the nuclei in the oosphere. 
Sei Or not the centrosome of Albugo persists 8 per 
§an of the cell is a question that as yet is impossible 
ba th i The structure SO prominent at metaphase is not Been 
might ei . nucleus ; the conditions, however, are such that it 
Wnoticed . hidden among the chromatin granules and pass 
Negative a. So small and its stain reaction so uncertain that 
‘i shispe os valueless. eos 
the condit; ; € spindle figure may be greatly mo oe 
or example, if the nuclei happen to be in pro- 
hase when the centripetal rush of cytoplasm 
on due to the movement of the protoplasm seems 


Mitotic figur 
from the per 


to answer, 


re) 
Ons; 
or metap 


°CCUrs the tensj 


168 BOTANICAL GAZETTE [SEPTEMBER 


to act conjointly with the normal elongating forces, thus pro- 
ducing extraordinarily long spindles (fig. 62). On the com 
trary, if the nuclei reach the border of the central mass ina 
earlier stage of mitosis no such forces obtain. Spindles caught 
in the first massing of the cytoplasm are often distorted ant 
bent like the letter 4 owing undoubtedly to torsion caused by 
the vacuoles as they move outward. ie 


MATURATION OF THE COMPOUND OOSPHERE AND OF THE 
ANTHERIDIUM. 


The multinucleate or compound oosphere when completely 
differentiated contains by actual count an average of 455 
nuclei. These are found in various conditions of reorganizatioa 
following the mitosis at zonation, and they rapidly assume th 
typical condition of a resting nucleus, each showing a promt 
nucleolus and very faint linin network. A nuclear membratteS 
sharply differentiated. Presently the linin network becom 
more prominent and a spirem condition is reached, ver Bi 
that first observed in the oogonium. A mitosis now one ae 
the oosphere affecting all of its nuclei, and is similar ba : 
important details to that just described for the oogonit 
as illustrated in jigs. 22-30. The nuclear figure stains 
more faintly than that of the previous division, the ; 
appearing lighter and skeleton-like in comparison with 
the first mitoses. The only other important differences 


“ests 


1899] THE COMPOUND OOSPHERE OF ALBUGO BLITI 169 


esting to note that the two nuclear divisions in the antheridium 
and in the oogonium are similar in character and proceed simul- 
taneously. The antheridial tube at the time of the differentia- 
tion of the oosphere lies in the periplasm, with its apex close to 
the bounding film of the oosphere. It now pushes into the 
oosphere, increasing in diameter as it advances. Ititakes safranin 
stain greedily from this time until it discharges its contents, but. 
if the stain be thoroughly extracted in acid alcohol and the 
preparation treated with gentian violet the contents become 
clear. The tube when fully developed is seen to contain numer- 
ous nuclei. A glance at figs. 73, 76 will give a clear notion of 
this condition. It will be seen that many nuclei are massed 
near the tip of the tube and that others are apparently entering 
at the base. It is impossible to determine their number by 
actual count, Owing to the crowded condition (jigs. 77, 85, etc.). 
However, as there are about 35 nuclei originally present in an 
antheridium, and these divide twice, there must be altogether 
ent 140. Of these, 20 or 30 perhaps remain in the anther- 
idium Proper, leaving a little more than 100 to pass into the tube. 
The antheridial tube pushes toward the center of the oosphere 
during the Second mitosis (fig. 70), and arriving nearly at the 
“enter its tip swells, becoming nearly globular. In this condi- 
tion the end of the tube is covered by a very thin wall which is 

tely visible, and yet holds within a dense mass of sperm nuclei 
S77). 


When the male n 
the ch 


anterior ; » and later pointed at both ends, and the 
énd is seen to contain the nucleolus around which is 

 * Massed a densely staining substance, probably chromatin. igs. 
rtd aly nuclei from both the base and tip of the same tube, 
Which is illustrated in Jig. 73. In the narrow entrance and 

*! Portion of the tube the nuclei are necessarily arranged in 
a ile, but as its diameter enlarges they become massed in 
ei and the tip is so closely packed with nuclei that 
| one forcibly of the appearance of a raspberry with its 


Sin 


170 BOTANICAL GAZETTE [ SEPTEMBER 


drupelets (fig. 77). Two sections of the same antheridial tube 
are shown in figs. 77, 78, one at the tip showing numerous nucle 
surrounded by a very delicate membrane, the other near the 
base giving a view of the narrow nearly empty cavity and the 
thick wall. As the tube enlarges the protoplasm in the anther 
idium proper becomes more and more vacuolate, but its conten’ 
never entirely leave the structure (figs. Soa, 86). The film 
separating ooplasm from periplasm is but slightly if at all 

changed by the entrance of the antheridial tube and during the 
maturation of the oosphere. The periplasm likewise shows 10 — 
important changes. Some of its nuclei divide mitotically, # 
the number does not seem to increase materially. Most of them 
remain in a resting condition. One case was observed wher 
every nucleus in the periplasm was undergoing mitotic division 
simultaneously with those of the oosphere, but this must be 
regarded as a very excéptional instance. 


FERTILIZATION. 


100 in number, are in resting condition. The antheri 
filled by an approximately equal number of male nuclei, 4 . 
tip has swollen so that the contents are separated fot : 
ooplasm by only the thinnest of walls. The wall finally 
ishes and the contents of the tube are free to mingle ee 
cytoplasm of the oosphere (figs. 80, 82). The pore 
move through the ooplasm toward the female nuclei, oe 
being often marked by a streak of denser cytoplasm. ak 
no visible cause of this movement, but as the male and ! i 
pronuclei differ in form a chemotropic influence may perhaps 
safely inferred. Longitudinal sections of antheridial tubes 7 
80, 82) sometimes show the nuclei pouring out, and pee =. 
or oblique sections (figs. 84, 85, 86, 77, 79) a thee 


proof of a discharge of many nuclei. There is in ange 
sections unmistakable evidence of a multinucleate disch of 
Sections 


the antheridium into a multinucleate oosphere. 


1899] THE COMPOUND OOSPHERE OF ALBUGO BLITI 171 


antheridial tube similar to those figured, both transverse and 
longitudinal, are not uncommon in the writer’s preparations, and 
many have been carefully studied. No antheridial tube was 
found which gave any evidence of the possibility of the dis- 
charge of but one functional sperm nucleus. 

When the sperm nuclei emerge from the tube their nucleoli 
are in the anterior ends, and later there appears prominently in 
the same region a substance that stains like chromatin. As the 
sperm nucleus approaches the female nucleus a faint linin net- 
work becomes visible fies. 73, 34; 35). When the sex nuclei 
first come in contact the male is the smaller, but later they 
become approximately equal in size. It seems probable that the 
female nucleus actually decreases slightly in size during this 
equalization. The nuclei do not immediately fuse, though both 
“¢ apparently in resting condition. All stages of fusion can be 
easily observed. The sexual nuclei are pressed together, the 
‘perm nucleus first assuming a spherical form, the bounding 
membrane disappears at the point of contact, and there results one 
dumb-bell-shaped nucleus (figs. 37, 38). As the coalescence 
becomes more complete the fusion nucleus takes ona spherical 
form, and presents the structure of a resting nucleus. No 
etal regarding the fate of the linin network were obtained, 
mica to the extreme minuteness of these structures and the 
ae difficulty in Staining them in a manner adequate to 
‘iakeama (Aes. 35-40). Fusion must be a process of dasa 
the — as fr Vela the advance made in other structures of 

pene : ing es = yecmimation, 
Pairing of a ew of an entire section of an oospore during the 
lumber of e Sexual nuclei is shown in fig. 88. A count of the 
a0 average ‘glee all of the sections of such an oosper gives 
Sperm nucle; avout 100. There seems to be a slight excess of 
ing the ac. ccasional small unpaired nuclei are found dur- 
antheridium ee: there are also several nuclei left in the 
Pore in a er and in the base of the tube. Sections of the 
antheridial ee the nuclei are fusing present no trace of the 

€ inside of its wall, although it is easily traced 


172 BOTANICAL GAZETTE [SEPTEMBER 


through the periplasm (fg. 97). Judging from this condition 
the terminal portion of the tube must vanish immediately after 
giving up its contents. The portion imbedded in the periplasm 
becomes thickened, resembling the primitive wall; but it seems 
never to attain the character of the mature epispore, asis the 
case in so many other species of Albugo, Indeed, no traces of 
the antheridial tube were ever seen in ripe spores. : 
The character of the ooplasm changes when the antheridial 
tube opens. As may be seen by comparing /ig. 70 with figs. 80, 
82, 84, 85 the vacuoles increase considerably in size and become — 
irregular in form. The most striking feature of this later condi- 
tion, however, is the tendency of the contents of the oosphere 
to break away from the periplasm (fig. 80), a phenomenon never 
met in younger stages. This indicates that changes have 
occurred at the boundary between the ooplasm and_periplast 
Indeed, it is at this time that a true wall may be first observed : 
around the oosphere. It will be remembered that previously ade 
periplasm and ooplasm were separated only by the delicate film | 
that appeared during zonation; but now for the first time adit 
tinct wall is present around the ooplasm, and its advent ee 
to be correlated with the opening of the antheridial tube. 
The wall occupies precisely the position of the film botnet 
the ooplasm and periplasm, and is probably formed by ah | 
development of that structure. Its intermediate position betwes! 
the ooplasm and periplasm and the apparent organic connie’ 
with both leads to the belief that it is the product of the ie 2 
action of both regions, rather than of either ooplasm a 
plasm alone. Since this wall remains perfectly distinct ch 
walls that are formed later, it will hereafter be called the 
tive wall. This term is used simply for convenienc’ . 
paper. Further study of related forms may esta att | 
homologies and lead to further classification. The 
wall is very clear and homogeneous in structure, €? hicknes 
striations, and shows great regularity of curve and - ee 
From this time on the condition of the developing walls 
as an index to the age of the oospore. 7 


1899] THE COMPOUND OOSPHERE OF ALBUGO BLITI 173 


A stage of somewhat frequent occurrence is shown in fig. 87. 
Judging from the presence of the primitive wall, the character of 
the ooplasm, and the absence of the antheridial tube, it must 
follow the opening of the latter, and since the nuclei are not yet 
paired must precede the condition shown in fig. 88. There are 
two possible explanations for this condition, consisting as it does 
of an oospore containing several groups of nuclei, each cluster 
imbedded in a mass of denser cytoplasm. Perhaps these nuclei 
are gathering the cytoplasm about themselves, a phenomenon of 
rather frequent occurrence with sexual nuclei; or it may indicate 
the breaking up of the mass of nuclei and cytoplasm that was 
released from the antheridial tube. The latter explanation 
seems more probable. If it be true, a stage similar to that 
shown in fig. 88 would result through a further fragmentation of 
these nucleated masses of denser cytoplasm. 

The previous pages have dealt entirely with descriptions of 
the antheridial tube, the discharge of its multinucleate contents, 
and: the Subsequent fusion of sexual nuclei in pairs. For the 
a of completeness, and in view of the peculiarity of the con- 
ditions and the general, if not universal, belief in a simple pro- 
hs % fertilization, involving only two sexual nuclei, it seems 
desirable to discuss the possibility of such an occurrence taking 
~ in the oosphere, together with the events already described. 
eli fertilization predicates the existence of one jeutete 
ews ay . oosphere i one male nucleus in the antheridium ; 
ture and e alone, or if with others at least different a struc- 
ation the o Action from them. Subsequent to its final differenti- 
Stage “5a os Sidi contains less than 40 nuclei. Ata later 
but Neither - aera: and oosphere contain about 100 peter 
the others Seal single nucleus differing in appearance i 

le cosphere iso uninucleated oosphere exists it must be bel ore 
shows is fully differentiated. A glance over the drawings 
there ever — uninucleate stage is represented, nor was 
MOSt Dersiste slightest hint of such a condition found cn a 

nt search, involving hundreds of oogonia. That . 
(have existed and escaped observation seems very 


174 BOTANICAL GAZETTE [SEPTEMBER 


improbable. The impossibility of the central body either being 
or containing a nucleus has been sufficiently discussed on 2 
previous page. The search for the single nuclei proved in vail. 
The refuge left for an adherent to the idea of a simple fertili 
zation, involving only two sexual nuclei, lies in the assumption 
that the nuclei of the compound oosphere ( figs. 68, 69) have 
descended from a fusion nucleus, which, owing to its rapidity of 
development may have escaped observation in earlier conditions. — 
That is to say, fertilization might have occurred at a stage simi- 
lar to that presented in fig. 6g or earlier. If this were true we 
would expect to find the ooplasm presenting 2, 4, 8, 16, 32, 4 
etc., nuclei, in stages following the division of such a fusioa 
nucleus. As a matter of fact no such conditions were ever 
observed, or is there the slightest evidence that they could be 
present. The oosphere when first differentiated contains 40-32 
nuclei, derived from the mitotic figures that line up in the mat 
ner shown in figs. 64,65. This number is increased to about 100 
by the mitoses in the compound oosphere (jig. 70), and thet 
comes the observed act of fertilization (fig. 85), the discharge a 
from the antheridial tube of a large number of sperm nuclei and 
the subsequent fusion of these in pairs (fig. 88) with the female i 
nuclei. Previous to the act of fertilization the antheridial 7 1 
gradually fills with nuclei as it presses deeper into the ooplas®: | 
There is of course a time when the tube contains a single nuclets a 
but this is when it is about one third the size finally reached, am a 
long before it shows any indication of opening. : os 
It is true that ly i enesis the dense cy top 
very early in oog — 

the interior of the oogonium may contain a small and very" 
able number of nuclei, as is shown in figs. 62, 62s 64. Oe : 
can be no doubt that these conditions represent part of ger a 
ess of zonation, and they have been discussed in that sect an 
this paper headed “Differentiation of the compound oosph a 
It is very probable that stages similar to these might ein the 
where there is only one nucleus left behind in the oospher ge 
process of zonation, but the condition of the antheridial 


ae oe 
all the further history of the oosphere show that this is »° i 


1899] THE COMPOUND OOSPHERE OF ALBUGO BLITI 175 


time when an act of fertilization could possibly take place. 
Moreover, when only one or two nuclei are present they are 
always peripheral, which would not be expected if they resulted 
by the division of a fusion nucleus. Again, these scattered 
nuclei are always in the same condition as those near the 
outside of the developing oosphere, and this is almost invariably 
ametaphase of mitosis. This coincidence is inexplicable on the 
basis of their being the result of the division of a fusion nucleus, 
but it follows as a necessity from the explanation offered in this 
paper. If they are the product of one nucleus, which has under- 
gone three or four divisions, we have to assume not only the 
existence of the sex nuclei, their fusion, and the fusion nucleus, 
but also the resting, prophase, telophase, and anaphase conditions 
in the formation of 2, 4, 8, 16, and 32-celled stages. An anaphase 
nucleus is never seen in an oogonium except when the nuclei 
divide simultaneously either during the first or the second mitosis. 
The first anaphase always occurs when the general appearance 
S that shown in jig. 67,7. e., when the nuclei are completely 
lined upand the ooplasm is well differentiated. The second 
*pPeats in the oosphere after its complete differentiation ( figs. 70, 
4). It is impossible that the 50 nuclei of the oosphere can 
have been derived from a single hypothetical fusion nucleus. 

If attention is turned to the antheridial tube it might be sug- 
:. fertilization could take place at an early period, when 
ee eg are like those shown in figs. 62, 64. But it is only 
always ed ¥ plain the fact that the antheridial tube is 
tion cn: short at this time, and invariably occues the posi- 
Tarely if a. ae 64. The wall of the tube is thick, the oe 
Cation that m contains nuclei, and there is not the slightest indi- 
that it ae y at all ready toopen. If it be assumed, however, 
Nucleus that release, in some manner difficult to detect, a single 
female eke, ad the male nucleus, and which fuses with we 

e develo is Ow can the continued growth of the tube an 
70,73 i of such peculiar conditions as are shown in ~~ 
tube a” 50, 82, 83, etc., be explained? Why does the 
grow after functioning only to meet the difficulty 


176 BOTANICAL GAZETTE [SEPTEMBER 


of disposing of its comparatively massive body and numerous 
nuclei in the ooplasm? Why do its nuclei later assume 2 
specialized form, resembling sperms (jig. 85)? 

In considering the positive side of the argument, in favor of 
a multinucleate fusion, no step is left to be filled by assumption. 
All of the stages were seen repeatedly and the correlations are 
perfect. The antheridial tube opens at the culmination of a 
period of gradual development which has been completely traced. 
After it has emptied its contents it immediately disappeats. 
The oosphere has likewise passed through a series of remarkable 
but perfectly graded conditions with all the steps of develop 
ment clearly shown. Coincident with the opening of the anthet- 
idial tube certain marked changes appear in the cytoplasm; the 
oospore wall is formed, the ooplasm immediately becomes much 
vacuolate where it was previously dense and uniformly constatl 
in character. The discharge from the antheridial tube introduc’ 
into the oospherea large number of nuclei clearly different inform | 
from those previously there. These sperm nuclei are seen in ay 
positions of exit from the tube, and finally become se distributed . 
as to indicate with certainty that they approach the female : 
nuclei. | 

At a stage positively older (judging by the development 
the primitive wall), the oosphere is found full of fusing paits . 
nuclei. That these are not nuclei dividing amitotically 15 PP" ;: 
by the number of nuclei in the oospore decreasin~ rather iv, : 
increasing, and also by the evidence presented through deta™ . 
study. 


THE UNIVERSITY OF CHICAGO. 


[ Zo be concluded. | 


Ye 
'\ 


STEVENS on ALBUGO. 


N 
N 
: 
be 
SN 
S 
N 
Pa 
S 
= 
L 
: 
> 
a) 
4 


PLATE Xil 


PLATE Xill 


N 
N 
S 
as 


’ 


BOTANICAL GAZETTE. 


PLATE XIV 


Ny 
N 
. 
> 
af 
N 
N 
. 
& 
= 
& 
: 
& 
1) 
el 


ZETTE, XXVIII 


PLATE XV 


SE A ee mM eR TS SHS Sa SMe SEES TM er mee air Pmen ven hi y net Ree eee mt aeN Re eee eae 


oa ae is 


TP Ped Saeel Oets RPE, koe 


4 


A BACTERIAL DISEASE OF THE SUGAR BEET: 
CLARA A. CUNNINGHAM. 
(WITH PLATES XVI-Xx) 


Ix the autumn of 1890 Professor H. A. Huston, chemist of 
the Indiana Experiment Station, noticed that the analyses of 
some sugar beets showed a much lower per cent. of sugar than 
others, and the difference seemed to be associated with a slight 
change from the usual appearance of the tissues of the root. This 
observation led to a microscopical examination of the affected 
beets by Dr. J. C. Arthur, who discovered the presence of bac- 
= in the tissues, to which, after further study, was attributed 
their abnormal condition, During the year 1891-2 the character- 
oi is the disease were studied by Dr. Arthur and Miss 
Katherine E. Golden, and the results published in the form of a 
bulletin in 1892.2 

: This preliminary series of investigations determined that the 
Rg Was associated with a specific germ, which could be 
‘ Polated from the diseased tissue. 
oe ee of the beet had been reported from any 
ork. rat 4 in America at the time of the publication of this 
of the bine most Kramer, in 1891, reported a bacterial disease 
simultaneou ag attacking the fodder beets of Russia, and almost 
of the ace 4 i Paul Sorauer, of Germany, reported a disease 

Uer gives “eg that country. In the Export of wee 

d By kis Ueda mnt the disease of the fodder ste 
SUgar beet es bacteriosis gummosis,” and that of = 
Read arly named by himself ‘‘bacteriose gummosIs, 
Fitton ag a for the Promotion of Agricultural Science at the Boston 
Fein no, — Of the sugar-beet root. Purdue University Agric. Exper. Station, Bul- 


7E 
1899) *port, 1894, no, 30. 


178 BOTANICAL GAZETTE [sermon 


are identical, and, perhaps, very closely related to the bacteral 
disease of sugar cane known as ‘‘sereh.” 
The diseased beets, as observed in Russia, are described : 
having dried leaves with withered heart leaves. The rootsof 
badly diseased beets were so tough they could scarcely be 
broken, the broken surfaces soon turning black. These bees 
produced a pathogenic effect on cattle to which they were fel 
Many of the diseased beets, when first sectioned, appeared pe 
fectly sound, but after a few minutes the fibrovascular bundles i 
turned dark and a syrup-like gum exuded from the cells. hh 
other beets the tissue was sometimes completely broken dow. : 
Dr. Sorauer says: ‘The similarity between the beet and th j 
sugar-cane disease ‘sereh’ consists in the destruction of the cane 
sugar and the increase of the invert sugar as well as in the color 
ing of the vascular bundles and the entrance of bacteria. He 
also believes that the disease discovered in America by Arthar | 
and Golden may be the same as that determined by Kramer 
himself in Europe. ; 
Mr. Walter Busse, in 1895, took up anew the stu | 
bacteriosis gummosis of the sugar beet, the material for study 
being sent him by Dr. Sorauer. In describing the diseased! 
he speaks of the gum-like fluid as follows: “500m ate. 
drops appear on the surface of the sectioned beet they 
covered by a thin black membrane, which consists of 
black, round bodies of different sizes.” : Bey 
90 The get OF Mir: Busse was, first, to determine the ee 
bacterium common to all the diseased beets by the sepa’ 
the germ from the diseased tissue; and, second, to cen a 
that this germ was the specific cause of the disease by 
healthy beets with the germ. In the first series of e ve 
three germs were isolated. Two of these were eT ‘o 
the third form was kept for further observation. Leet 
appeared as short rods 1.72-2m long and 0.8 progr 


dy of the 


solutions, producing an abundance of gas. a 
but from othér diseased beets a second gas-producing ~— 


1899] BACTERIAL DISEASE OF THE SUGAR BEET 179 


isolated, resembling the first in form and arrangement, but 
smaller, being 1.5 w—1.75 w long and 0.7—0.8 # broad. This form 
produced an acid reaction on different media, did not liquefy 
gelatin, and grew better at 12°-14° than at 22°. Mr. Busse is 
inclined to believe that the second form is a variety of the first, 
which he designates as Bacillus Bete. He has demonstrated that 
this second form produces the disease known as dbacteriosis gum- 
mosis, and believes that this germ is a saprophyte which becomes 
a parasite in the tissues of the beet. 

Erwin F, Smith,‘ in speaking of the bacterial diseases of the 
sugar beet as reported from Europe and America, is of the 
pinion that the diseased condition of the beets studied by ° 
Arthur and Golden is due to some other cause than a bacterial 
one. He states that it is highly improbable that the root could 
be attacked by an organism which invades its tissues and yet 
does not break them down. Mention is made of the fact of the 
Presence of small bodies in the tissue of healthy beet roots 
which have the appearence of bacteria, but which are probably 
“ystalloid bodies. A paper by Dr. Smith was presented at the 
‘pe of the Society for Plant Morphology and Physiology 
“ i eae 1897, calling attention to the “existence, in parts 
nia nited States, of a disease of the sugar beet resembling if , 
a €ntical with that described by Kramer and Sorauer in 

91-2, and more recently by Busse.” 

ne ae fall of 1896 I had the opportunity to continue the 
I a 10n of the bacterial disease of the sugar beet observed 
study ag eae ts Much of the value of my experimental 
Sheen disease is due to the suggestions of Dr. Arthur, to 
l also desir indebted for kindly help and criticism of my work. 
fessor a ” Press my gratitude to Professor Burrage, Pro- 
°n, Miss Golden, and Mr. H. L. Bryan, also of 
ty, for important suggestions. My investiga- 
€ been continued from 1896 to the present time, 
NO positive evidence that the sugar-beet disease 


‘ © Same as that described by Sorauer and Busse 
Sviry EF. he 


‘Urdue Univers; 
tions, Which hay 
ie Fesulted in 
0 Indiana is th 


Nat. 30:716-729. Sept. 1896. 


180 BOTANICAL GAZETTE [SEPTEMBER 


of Europe. The points of similarity will be noted in the follow | 
ing description of the disease and of the germ by whichitispm — 
duced. | 


GENERAL DESCRIPTION OF THE DISEASE. 


About the middle of September 1896 several diseased bees 
were found in the field of cultivated beets on the grounds of the 
Purdue Experiment Station. The disease attacks the whole 
beet plant, causing a peculiar appearance of the leaves, so that 
with a little practice the diseased beets can be distinguished — 
readily from the healthy ones as they grow in the field. The 
outer, older leaves soon die away, and the intermediate and 
heart leaves are left wrinkled, curled, rather flabby than turges 
cent, and of a yellowish-green color. This wrinkled appearaice 
is caused by blister-like patches being formed between the | 
veins of the leaf, and the whole has been described as resembling 
a Savoy cabbage leaf. See photographs of leaves, plate XVI, 
also photographs of diseased beest, p/ates XVII, and XVII # 

The appearance of the exterior of a beet root when disease 
is not materially different from that of the healthy beet. Kk | 
perhaps not quite as brittle. A decisive test for the wee 
found in the appearance that the root shows when ae 
The fibrovascular bundles appear as dark rings in the oe fai : 
They grow almost black after being exposed to the air forST 
minutes. These rings are quite distinct from the cream col 
fibrovascular bundles of healthy beets (plate XIX). t og 

In 1896 in a field of beets covering an area of mee 
acre and containing approximately 130,000 beets, oi ss 
diseased and several slightly affected ones were foul 
was a smaller number than had been found on the ge te 
in previous” years, and can perhaps be accounted < 
climatic conditions being so favorable to plant BF : The 
preceding summer, there being an abundance of me 
number of diseased beets increased, however, beige! 
time. geased the 


Frost seems to be much more injurious to the di 


ies 
* 


Of melted 


1899] BACTERIAL DISEASE OF THE SUGAR BEET 181 


to the healthy beets. The heart leaves of the diseased beets 
were more easily injured by the frost. It is characteristic of the 
disease that the leaves of badly diseased roots die away until no 
green leaves remain, leaving an apparently dead root in the soil, 
though its tissues will be found to be firm and not in the least 
broken down. The early frosts hasten the destruction of the 
leaves. Both diseased and healthy roots show an acid reaction, 
the diseased seeming slightly more acid than the healthy. 


ETIOLOGICAT HISTOLOGY. 


Comparative study was made of sections taken from both 
diseased beet roots and healthy beet roots, also sections from 
leaf and leaf-stalk of both diseased and healthy beets. In all 
these sections small round bodies were seen in the cell sub- 
stance. These bodies were found to be protein by their reac- 
tion to iodine. They measured from 2-4m in diameter and 
turned yellow when treated with iodine. In the tissues of 
diseased beets other bodies were found which were smaller and 
of a different refractive power and arrangement. These bodies 
‘tained in gentian violet like bacteria, and looked almost like 
“rococci when imbedded in the cell substance, but when free 
nthe water were easily distinguished as small motile bacilli. 


SEPARATION OF THE SPECIFIC GERM. 


he first Steps im the Separation of the germ were as follows. 


T 

“seased beet was selected, a thin knife sterilized in the flame 
<a “move all parts of the beet exposed to the air. A 
epee of beet was then removed from the heart of the root 
ey “ansferred by means of a sterilized platinum wire to tubes 
8elatin or agar. The first series of cultures was made 
ting Selatin tubes with pieces of diseased tissue, at the 
a inoculating a number of tubes in the same manner with 
a beet ‘as a control. The following tables ite 
Tesults i of a Series of such inoculations. The first shows ed 
Tsults of Cultures made from diseased tissue; the second 

° cultures made from healthy are 


182 BOTANICAL GAZETTE [SEPTEMBER 
TABLE I. 


CULTURES ON ARTIFICIAL MEDIA FROM DISEASED AND 
HEALTHY BEETS. 


Number 
Dates and media of Results 
cultures : 
| eee : 
FROM DISEASED BEETS. : 


ee 
Sept. 20 10 Oct, Oct. 10 
3 Gelatin - - Chivaccuteae growth Contaminated 
t. 2 3 eta ee 
Gelatin - - Characteristic growth Still pure 
Oct. 2 4 Oct. 2 
Glycerin gelatin Characteristic growth 
Oct. 24 4 Oct.:2 
Glucose gelatin Characteristic growth 
| init 
FROM HEALTHY BEETS. E 
Sept. 20 Oct. Oct. 10 a 
O Gelatin - - 4 - growth : . No grow? : 
chs, 2 Oct. 45 e ; 
Gelatin - - 3 No growth . No go : 
Oct. 24 Oct. 28 j 
Glycerin gelatin 4 No growth ] 
Oct." a 
Glucose gelatin ) 4 
| 4 


All the above cultures were made in standard gelatin not it ) 


ted or in standard agar to which had been added 5 pee 
glycerin or 5 per cent. glucose. The growth in successitl 7 
ulations was the same in all cases. Some creamy white ee 
grew out on either side of the diseased tissue, and in the 
of a few days were surrounded by a lens-shape¢ °F | 
in the gelatin caused by the gas that was given off int cbt 
When the bubbles reached the surface the growth w 
in white rings around the tube (p/ade XX, By eel 

October 15 several Aries were made in sands 
In some of these a growth resulted but no successtt ea 
were made. One of these cultures was used to in . 
beet in the field. 


5 Photograph of tubes at this stage shown in plate XX. 


1899] BACTERIAL DISEASE OF THE SUGAR BEET 183 


November 3 stab cultures were made from a tube of glycerin 
agar inoculated with diseased tissue. These were all contami- 
nated with the exception of one which formed a perfectly color- 
less layer, gelatinous in consistence, on agar and sterilized beet. . 
This form will be spoken of later. 
No growth resulted from a series of cultures made in 10 per 
cent. cane sugar gelatin. 
In December a diseased beet, which was frozen the previous 
aight, was brought in from the field. Pieces of this tissue were 
transferred with the usual precautions to tubes of melted gelatin 
‘o which had been added 5 per cent. of cane sugar. The growth 
in these cultures was rapid and gas was given off in large quan- 
ties, Stab cultures were made from these and appeared exactly 
uniform, and just the same in appearance as stab agar cultures 
made directly from unfrozen diseased beet, with one exception 
meach case. From one of these exceptional tubes the perfectly 
colorless gelatinous form spoken of above was found. This form 
"as also obtained in one of the stabs taken directly from the 
diseased beet, 
lag above inoculations the growth was much more rapid 

Previous cultures, probably because the tissues were 
broken down by freezing so that the germ could escape more 
“asily into the ; : 

Surrounding medium. 
: Searal Series of cultures was made at the same time om 
¢t in which the disease was produced by inoculation. 


fers . wth was similar to that of the preceding series. Trans- 
above *n from these were uniform and similar to those described 


ae inoculations a plug was removed from the healthy 

e plug aa @ sterilized knife, the inoculating material inserted, 

Page nile aced and covered with cotton. The table on the next 
The Sey results of inoculations of three beets in the field. 

ing the a oculated with diseased tissue was slower in show- 

ON gelatin “Se than the one inoculated with the germ growing 
ake TY because of the time required for the germ 
way through the cell walls of the tissue. 


154 BOTANICAL GAZETTE | SEPTEMBER 


TABLE II. 


RESULTS OF INOCULATING BEETS IN THE FIELD. 


October 22 November 19 November 28 


Beet 
_ Date of 
inoculation 
Source 
of germ 


1 tOct Isolated Yellowish color of | Disease quite | Removed from the . 
germ leaves noticed evident field | 


—_e-or oo eer 
2| Oct. | Diseased | Slight indication | Disease quite | Removed from ® 
tissue i field 


17 of the disease 
TE 
a1: Oct Healthy No change No change No change 
17 tissue 


—_—$—<————e 7 
The heart leaves of the inoculated beets showed the effet 
of the disease in the slight blister-like areas on their surfaces 
The beet inoculated with the gelatin containing the germ, aa! 
the one inoculated with healthy tissue, were brought pes: 
laboratory November 28, as the progress of the disease coul 
not be followed out of doors because of injury by frost La 
beets were placed in culture jars of water in the gree ee | 
where the healthy beet after some time decayed, and theo 


crinkled and faded, but gradually assumed a vine 
darker green appearance, but the plant was sti 
growth. ; 
INOCULATIONS IN THE GREENHOUSE. = i 

The method of inoculation of beets in the greenhous® 2 
similar to that of inoculation of beets in the field. As ; Fhe 7 
of these trials several beets seemed to show the effec 
disease to a slight degree. 2 , 


DESCRIPTION OF THE GERM. — tbadt 

The germ as isolated from the diseased tissu¢ ! 
lus measuring from 0.9-1.0—1.3 » in length, and 0.5-°: nase 
When taken from the culture media the gerne at oe 
singly or in pairs, and possess individual motion. a 


1899] BACTERIAL DISEASE OF THE SUGAR BEET 185 


seems to revolve more or less irregularly on its axis. The germ 

stained well with all the common bacterial stains. No process 

of staining showed the presence of spores or flagella ( plate 
EFFECT OF LIGHT ON GROWTH. 

The germ grew better in the dark than in the light. Germs 
taken from old, dried out cultures were smaller than when grown 
on a moist substratum. Desiccation also injures the capability 
of the germ for motion. Germs taken from an old culture and 
examined in a drop of water were less motile than those taken 
from a fresh bouillon culture. 

The germ grows better at a temperature of 12°-14° than at 
21°. Stab cultures in agar grew slowly at body temperature. 
The germ grown in bouillon and exposed to a temperature of 
100° for five minutes was killed. 


GROWTH OF THE GERM ON DIFFERENT CULTURE MEDIA. 


Stab cultures of gelatin showed a thin grayish-white layer 
baad surface, and extending down the line of inoculation. As 
the cultures Stew older the color darkened to a deep cream. 
The gelatin was liquefied in the course of several weeks. When 
Sige gelatin was inoculated and then allowed to solidify there 
nai growth throughout the gelatin in streaks and films. 

‘SS on gelatin plates were not distinctly outlined and were 
seonnmes accompanied by a disagreeable odor. The germ 
Soa Stow better on agar than gelatin. Agar to which had 

eed 3 per cent. of cane sugar or glucose seemed to 
Paps lavor its growth. The growth on slant agar was drab- 
growth og Smooth margins, and a slow and not luxuriant 
ta 6 © growth was not viscous. 
layers Sar plates the colonies have their origin in the deeper 
of the agar where they are generally elliptical. When 
ot f : ; i i d grayish- 
White Coloni urface they spread out in their round gr y 
..» With compact creamy-white centers. In bouillon 
fluid “eg observed after two or three days. No turbidity of the 
observed, but a sediment was deposited in the bottom 


186 BOTANICAL GAZETTE [ SEPTEMBER 


of the tube. Masses of zoogloea were sometimes found in 
old bouillon cultures. The germ grew well on sterilized sugar 
beet, and deposited a sediment in sterilized beet juice. The 
_ germ grew on sterilized apple, potato, and turnip. A rawpotato 
was broken open and inoculated with the germ, There wasa 
slight growth developed. A raw sugar beet was inoculated 
The germ grew, causing a black coloration of the fibrovasculat 
bundles, and in a microscopical examination was seen to have 
entered the tissue. | 

NITRATE SOLUTION, 


This solution was prepared using 1000 distilled water, | 
gram peptone, and 1 gram potassium nitrate. Tubes of this 
solution in which the germ had grown for three days when tested 
showed that the nitrate had been fairly well reduced. 


ACID AND ALKALINE MEDIA. 


It has been stated that the beet root is acid to the extent of 


little over 1 per cent. Because of this fact experiments were made | 
with acid and alkaline media in order to determine which e 
two would be more favorable to the growth of the gam : 
bouillon, to which had been added 1 per cent. malic acid, thesoluti® 

was made neutral; 5 per cent. acid solutions were not rend a. 
neutral. The solution was not made turbid by the growth wee : 
germ. In 1 percent. alkaline solution of bouillon the gem a 
more motile than in 1 per cent. acid solution. In iv : 
alkaline solutions the germ was more motile and larger, than | 
grown in acid media, measuring from I.1I-1.9 p long; and 09 . 
broad. In 5 per cent. alkaline media the sediment ae : 
was quite viscous. Zoogloea masses were found more” or 
abundantly in all the alkaline cultures. These solulio® 
not made acid in reaction by the growth of the germ. 


STARCH SOLUTIONS. 
f starch fits 


In a solution composed of one part each oO 
and bouillon, the germ grew but the starch was 
These cultures were tested for starch and gave 


not re 


a gecide® 


1899] BACTERIAL DISEASE OF THE SUGAR BEET 187 


reaction; they were also tested with Fehling’s solution for 
glucose, but gave no reaction. The germ grew well in wort 
gelatin.” When this gelatin was melted, and after being inocu- 
lated was allowed to solidify, the germ grew under the surface, 
producing bubbles of gas all through the gelatin. 

Tests made of bouillon containing cane sugar in which the 
germ had grown gave a reaction for glucose. In order to deter- 
mine if the enzyme existed outside the cell, a solution in which 
the germ had grown was filtered through a porous cup. This 
filtrate added to a 5 per cent. cane sugar solution, and tested 
with Fehling’s solution for glucose gave no reaction. Further 
*xperiments are necessary before deciding definitely in regard 
to the enzyme properties of the germ. 


CELLULOSE SOLUTIONS. 
As the germ penetrates the cell wall of the plant in some 
"ay, €xperiments were made to determine its effect on cel- 


— For this a special nutrient solution was prepared as 
Ollows :7 


ee a ee 
a : : - - 2.5 gram 
Magnesium sulfate, ‘ : : os 
Calcium phosphate, - : S é . 2 ac f 
Ammonium sulfate, a er 
Sodium chlorid, ‘ s : ‘ ‘ s 1.25“ 
Beef extract, . ; % : : és aes 


Swedish filter. 


* ® this solution the growth was very slow, and a very small 
ting of gas was produced, and the cellulose slightly broken 


FERMENTATION.® 


of the abundance of gas produced by the germ in 
Special fermentation solutions were prepared. The 
“Wort 8elatin was 
iy, ™ la fermentati 


: Because 
"S growth, 


made by adding 10 per cent. of gelatin to wort. 
on de la cellulose. Centralblatt fiir Parasitenkunde 2: 358. 
* Paste } 
0 fermen re H.and EMMa: A ‘report concerning gases produced by bacteria 
Centralb. f, Parasitenkunde 2:707. Dec. 1896. 


188 BOTANICAL GAZETTE | SEPTEMBER 


gas produced by the growth of the germ in these solutions was 
analyzed by Mr. H.S. Bryan, of Purdue University, the results o 
which are given in Table III. The tests were made for CO,,0,), 
NH,, and CO, the difference being considered as hydrogea 
In the first analysis a much larger amount of nitrogen wa 
obtained than in any of the other cultures. The cultures for 
this analysis were several weeks old, and were perhaps not 
trustworthy. 

There were some irregularities in the amount of gas prv- 
duced, which cannot be accounted for. At one time 2 per cent. 
cane sugar bouillon containing no pepton when inoculated 
gave a large amount of gas, 20° being collected in each fer 
mentation tube; 2 per cent. cane sugar bouillon containing % 
pepton, inoculated at another time under exactly similar condi 
tions, gave only 2° of gas in each fermentation tube. 

The germ grown in bouillon to which had been added 2p 
cent. glucose at one time gave a very large amount of gas: # 
another time there was not enough gas produced to be analyze 7 
The gas produced by the germ as determined by analyses 
composed of a very small amount of oxygen, less ion Js . 
cent., carbon dioxide approximately 44 per cent., nitroget " ; 
per cent., and hydrogen approximately 30 per cent. bee 
tion was produced in sterilized beet juice, Pasteur’s solution, 
maple sap. No fermentation was produced by the 
the germ in bouillon to which no sugar had been added. 


SUMMARY. 
mination 0 
bacteria * 
ot broke! 
pe te 


It has been determined that a microscopical exa 
the tissues of diseased beets reveals the presence of 
the cells of the plant. The tissues of the plant are ® 
down, and the bacteria in all parts of the plant appeat “ ot 
same. Transfers of diseased tissue to the healthy rt ad ] 
resulted in changed appearances of the plant which 1 
almost certainly that the disease was transmitted. the plast 1 

The manner in which the germ finds entrance to to th 
has not been determined. The conditions most favore ‘ 


189 


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Igo BOTANICAL GAZETTE | SEPTEMBER 


attack are those resulting from drought with succeeding low 
temperature. 

The fact that the germ breaks down cellulose slowly explains 
the manner of its progress from one cell to another, 

Experiments have shown that the germ in a medium of low 
per cent. acid grows nearly or quite as well as in one of alkaline 
nature, so that the acid element of the beet root does not offer 
material resistance to the germ. 

The germ converts cane sugar to glucose in the process of 
producing gas. The amount of gas produced is not constant, 
but the reasons for this irregularity have not been determined. 

The germ grows well with any form of sugar and especially 
well in media containing cane sugar. This fact makes it seem 
probable that the germ is especially at home on those media 
which contain sugar in some form, although it will keep alive om 
media without sugar, and after cultivation for a time on such 
media will adapt itself to the conditions presented. 


ANOTHER ORGANISM SEPARATED FROM THE SUGAR BEET. 


The colorless gelatinous form separated from the beet root 
in connection with the disease germ was at first thought to be 
an undescribed germ or rather the product of a germ, for only @ 
few bacterial bodies could be detected under the microscop® 
even when comparatively large masses of the substance Wer 3 
placed in the field. The organism appeared as small bacilli of : 
micrococci, ek 

The mass resembles the form of Leuconostoc so common! 
the vicinity of sugar refineries. Under the microscope, however, 
no streptococci were found, which characterizes Lene 
under the microscope. The gelatinous substance is pegs 
water and alcohol; in the latter it turns to a milk-white ee 
stance before it dissolves. The substance increased Tap a 
bulk when grown on sterilized beet. The mass did not dry ° 
for months after the substratum had become dry and hard. 

The substance grew well on 10 per cent. cane sugar pa 
The growth was slow at first, but after a week or tw? 1g 


: 
F 


Kray 


1899] BACTERIAL DISEASE OF THE SUGAR BEET IgI 


measuring a quarter of an inch in thickness and three fourths of 
asinch in circumference, collected on the surface of the medium 
instab and slant cultures. In case of stab cultures the agar was 
broken vertically along the line of inoculation. The colorless 
growth followed this break in the agar, and as the substratum 
vecame hard the mass collected as a colorless semi-fluid in the 
bottom of the test tube. 

On slant agar there was a thin colorless layer, imparting a 
fuorescent hue to the medium. In agar plate cultures the 
oganism formed small round colonies about the size of a pin 
tead, resembling a small drop of water. These colonies were 
‘ometimes found with the disease germ, in plate cultures taken 
directly from the beet. It also grew on sterilized potato, and 
‘0 some extent on gelatin. Immediately after separation from 
the beet root the organism produced fermentation, but the power 
Was lost after a time. Staining revealed only a structureless 
Mass containing a few bacteria-like bodies, 
| oui Httle effect on the substance. Sections of 
linc, Ic . e organism was growing have been kept in the 
story until they are quite dried out, and the gelatinous mass 
8 stil] apparent, 
vA =e a form of Leuconostoc, it is interesting to 

ased beet roots, 


PurpuE University, 
Lafayette, Ind. 


BIBLIOGRAPHY. 


rat, ae and Gotpew, KATHERINE E, Diseases of the sugar beet 
shia UL. 39 Ind. Agr. Exp. Sta. 54-58. 1892. 
ag tb. der deutschen landwirtschaft. Gesellschaft 1892. Abs. 
—“ Bp nift f. Pflanzenkrankh, 2:280. 1892. 
In i « is : : 
= x, der “Sereh” des Zuckerrohrs verwandte Krankheiterschein- 
ce Uckerriiben, Export no. 30. 1892. 
8: 40-4, eoereciches landwirdschaftl. Centralblatt 2: 30-36. 1891; 
- 1891, 
Couey, K. © ba. : 
1890 : 99, 1891 ases of the sugar beet root. Proc. Ind. Acad. Sciences 


. 


192 BOTANICAL GAZETTE [SEPTEMBER 


Busse, WALTER. Bact. Studien iib. d. Gummosis d. Zuckerriibe. Zeitschrift _ 
f. Pflanzenkrankh. 7: 65-77, 149-155. 1897. Abs. in Centralbl. £ 
Bakteriologie und Parasitenkunde 3: 680-682. 1897. og 

SmiTH, Erwin F. The bacterial diseases of plants. II. Amer, Nat, 3o; : 
716-723. 1896. : 

The bacterial disease of sugar beets in the United States, Pre 

sented before the Society for Plant Morphology and Physiology in 1897. _ 

Ref. in Science 7: 118. 1898. z 


EXPLANATION OF PLATES XVI-XX. ; 


' PLATE XVI. e 

Leaves of healthy and diseased beets, as they appeared when brought in 

from the field. The three diseased leaves can readily be distinguished from 
the two healthy ones because of their blistered and crinkled surface. 


PLATE XVII, 

A diseased beet brought in from the field. The root is quite firm, none 
of the tissue being broken down. The leaves hanging down are quite dead 
and dry. The erect heart leaves are alive and show the characteristic 
crinkled surfaces. A 

PLATE XVIII. 

A, a diseased beet with the dead leaves removed. 8B, a healthy beet of 

same size and stage of growth, also with the dead leaves removed. 


PLATE XIX, 

A, cross sections through the crown of healthy and diseased bet 

4, healthy root. 4, diseased root, characterized by the black rings of vasculat 

tissue. : and 
&, cross sections through the central portion of the same diseased 

healthy beet roots. a, healthy root. 4, diseased root. 

C, cross sections near the tip of the same roots. 4, healthy r00t : 

diseased root. 

PLATE XX. figs 

A, longitudinal sections of the same healthy and diseased beet roots . 

ured in the preceding plate. a, healthy beet. 4, diseased root. «mila 


upward by the production of gas. 
B, the disease germ stained with carbol fuchsin ; 
graphed. : 


“imperfectly pho 


CUNNINGHAM on SUGAR BEET 


BOTANICAL GAZETTE’ XXV/// PLATE XVII 


CUNNINGHAM on SUGAR BEET 


WIANITAL GAZETTE, XXVIII PLATE XVIII 


CUNNINGHAM on SUGAR BEET 


WAL GAZETTE, XXVIII PLATE XIX 


ee eae a! 


PLATE XX 


WOTANICAL GAZETTE, XXVIII 


CUNNINGHAM on SUGAR BEET 


REVISION OF THE NORTH AMERICAN SPECIES OF 
TEPHROSIA. 


B. L. ROBINSON. 


Waite the greater part of the difficulties of the genus 
Tephrosia lie, happily, beyond the geographical limits of the 
North American flora, yet the dozen species which inhabit 
sandy regions in our southern states are sufficiently variable in 
their foliage and similar in their floral and carpological structure 
tolead to diverse views on their specific limits and proper arrange- 
went. Miss Vail’s recent monograph? of these species, although 
sounding in long and detailed descriptions, fails to bring out 
early the primary divisions of the genus. The introductory 
*nopsis— which, although arranged in perfect accord with the 
meal sequence of the species in the paper, is styled an“ arti- 
‘: key” —is based chiefly upon the length and density of 
a features which are too variable and confluent 

aay diagnostic characters of the first rank. It seems to 

stem a that the sympodial and monopodial structure of the 
| Ciently nits difference much more evident and, indeed, suffi- 

“Shige to divide our species at once into two well- 
ts, eg 7 After this first division the size of the flow- 

emplo cg the inflorescence, and nature of the foliage may 

In mi hae greater diagnostic effect. 
Craceg er nation of species it is believed that Miss Vail’s 

is only separated artificially from her C. Small. 
ky adduced are chiefly the “truly prostrate ” habit of 
the eee and its narrower, more numerous leaflets. 
: to eam of the leaflets, this is (according to: Miss 
; Paely no ver, in C. Smallii, and 5 to 12™™ in C. floridana, 
Rid 'y Striking difference. In number the leaflets are 


na 
The distinctions 


AS to 
x“ V. ail) 


: "A tevis, nC. Smallii from 3 to 11, while in Mr. Nash’s 
>to on f ; 

Ps 25336, the North American species of the genus Cracca. Bull. Torr. Bot. 
ees 


193 


194 BOTANICAL GAZETTE [SEPTEMBER 


authenticated specimens of C. floridana they vary from 7 to 13, 
and had these specimens been younger, it is safe to surmise, 
leaves with fewer leaflets would have been found, an inference 
amply supported by the development in related species. 
Regarding the supposed assurgent habit of C. Smallii (a species 
not only distinguished in the field, but called to scientific notice 
by Mr. A. H. Curtiss), it may be said that the main stems bear 
evidence of being prostrate except near the tip. This is shows 
not only by the curvature and position of the leaves and pedun- 
cles, but by the almost constant presence of clinging particles 
of sand which adhere, even in the type specimens, to the lower 
surface of the stem, and are lacking upon the uppet surface, 
This position of the stem is fully confirmed by field notes 
kindly furnished by Mr. Curtiss. The assurgent character |s, 
therefore, confined to the leaves (which are said to be somewhat 
ascending but less erect than in 7. ambigua), the peduncles, and 
the growing tips of the stems. On the other hand, specimens 
of C. floridana, although said to have prostrate stems and leaves, 
give by no means the impression of a closely prostrate plant, 
an idea which is at once conveyed by specimens of T. cluyw 
phylla. (nthe absence of more telling morphological features 
the two species C. floridana and C. Smallii are here united. 


: nee kes s rth’s 

Miss Vail is certainly in error in interpreting yee 

T. angustissima, her characterization evidently being peer 
co 


Mr. Curtiss’ plants so named. Shuttleworth’s typ® ii 
on the Miami river by Rugel, is in the Gray herbarium, es : 
clearly a linear-leaved form of 7. purpurea Pers. with whic os 
shares the characteristic small flowers, of which one OF hi 
axillary, and the rest borne in a very slend ~ 
the other hand Curtiss’ nos. 584 and 5708, 
appear to be nothing but smoothish narrow-leaved : speci 
not differing by a single morphological character ee sat 
value. In her key (p. 26), Miss Vail distinguishes “ad 
(her Cracca angustissima) from T. ambigua on the Bre (p. 32) 
it is erect, and has linear leaflets, but in her description P 

it is characterized as prostrate. 


just me 


} 
4 
: 
E 
; 


; Sonia Necker, Elem. 3: 36 


: “aa 


1899] NORTH AMERICAN SPECIES OF TEPHROSIA 195 


I. Rugela Shuttleworth is an interesting and hitherto unde- 
sribed species possessing the stem-structure of § Brissonia 
but the habit and foliage of 7. chrysophylla, which is of § REIN- 
PRIA. 

l’ tenella Gray seems to have been founded upon a juvenile 
state of T. purpurea, to which (under the name of T. leptostachya 
DC.) it was reduced by Bentham in Mart. Fl. Bras. 151: 48. T. 
purpurea, however, is highly variable, at least in foliage, and 
certainly merits further study with more copious material than is 
now available. 

The writer is grateful to Drs. Britton and Small for the loan 
of the larger part of the North American Tephrosias from the 
terbarium of the New York Botanical Garden. These have 
keen of material assistance in the present revision. 

As the genus is here interpreted in its generally accepted 
“nificance, it is useless to take space for a generic description. 
Ka ning the species here described, the tropical 7. cenerea 

‘fas been found in the southern states, but only on ballast 
9 (Alabama, Dr. C. Mohr). It is of the § RernertA, and 
= es numerous narrowly oblanceolate-linear leaflets, which 

aty-pubescent upon both surfaces. 


ak Brissonia DC, Stem monopodial : racemes terminal or 
large ¥ Rever opposite the leaves: flowers (in our species) 
Petals 1 to 1.7 long.—Prodr. 2: 249, in part ; 

~ im Engl. & Prantl, Pflanzent. 3:. Ab. 3. 269. . Bris 


*Pp 
ods glabrous at maturity : racemes loose: southwestern. 


om Gray. Undershrub, cinereous with fine appressed 
lite, a. Several, 4 to 6% high, suberect : leaves petio- 
Hien lolate; leaflets narrowly oblong, 1.6 to 3.2% in 
long, about °s both terminal and axillary: pods 5 to 5.7™ 
Seeded. pj. Wright. 2:36; Walp. Ann. 4: 489. 
22; 28 Rita Kuntze, Rey, Gen. 1:175; Vail, Bull. Torr. Club 
type locality is erroneously stated to be New 


Sonoita valley, close to the southern boundary of 


196 BOTANICAL GAZETTE [SEPTEMBER 


Arizona, Wright, no. 965, Rothrock, no. 685. (Northern Mexico.) 

* * Pods permanently pubescent or puberulent: inflorescence short and 
dense. 

+- Calyx-lobes ovate-lanceolate to lanceolate: pubescence gray: petioles 
rarely 6™™ in length. 

T. VIRGINIANA Pers. (Goat’s RUE.) Stems several, 3 to 4.5 
high, erect from a stout knotted lignescent root: pubescence 
fine, soft, somewhat variable in quantity, often copious toward 
the summit: leaflets 11 to 23, oblong to elliptical, green and 
scarcely pubescent above, somewhat paler and _ soft villous 
beneath, 2.5 to. 3.2™ long; petioles very short; stipules cadu- 
cous: flowers borne partly in pairs or singly in the upper axils 
but chiefly in a short dense raceme little raised above the sur- 
rounding leaves: calyx hairy, the teeth caudate-acuminate: 
petals white or pale yellow with purplish tinge: hirsute pods 
soon spreading or divaricate.—Syn. 2: 329; Pursh, Fl. 2:489; Ell. 
Sk. 2:245; Torr. & Gray, Fl. 1: 295; Wats. & Coulter in Gray, 
Man. ed. 6, 133; Meehan, Nat. Fl. 1:81. pl. 27. T. virginica Bigel. 
Fl. Bost. ed. 3, 296. Galega virginica L. Spec. ed. 2, 2: 1062; 
Hill. Veg. Syst. 21. p/. 55, fz. Cracca virginiana L. Spec. 2+75?) 
Vail, l. c. 27,—Dry open woods especially in sandy soil, com- 
mon; New England to the north shore of Lake Erie, then 
to Texas and Florida; fl. May, June; fr. July to September. be 

Var. HOLOSERICEA Torr. & Gray. Leaflets inclining to 
narrow and acute; pubescence more copious, long and silky 
even woolly on the pods.— Fl. 1: 296. 7. holosericea se e 
Acad. Philad. 7:105. Cyracca virginiana var. holosericea ‘at 
l.c.—Arkansas, Nuttall, Marcy Exp., and Texas, Hall. Sim 
but less marked forms in Wisconsin and W. New York. 

+ + Calyx-lobes very narrow, almost filiform: pubescence tawny: 

++ Petioles 1.7 to 4.2™ long: leaflets oblong: southwestern. 

T. LEUCAN from a. lignesee 

. THA HBK. Erect, branched fro 

; : : th surfaces 
base: leaflets 15 to 20, oblong, appressed-villous on bo dae. 
nearly concolorous, I.9 to 3.2°™ long: racemes capitate, P ae 
late, chiefly terminal; linear filiform bracts considerably 2 
ing the buds: petals white with or without a purple ange: 


1899] NORTH AMERICAN SPECIES OF TEPHROSIA 197 


tattow, 5 to 6.2°" long, soon divaricate.— Nov. Gen. & Spec. 
§:460. fl. 577; Gray, Pl. Wright. 2: 36; Torr. Bot. Mex. Bound. 
s1. Cracca leucantha Kuntze, 1. c.; Vail, 1. c.—Mountainous 
regions in S. Arizona, Rothrock, Lemmon, Pringle. (Mex. where 
frst collected by Humboldt & Bonpland. ) 


++ ++ Petioles 4 to 17™™ long: leaflets obovate: Florida. 


‘I, Rugelii SHurrtewortH in herb. Stems several, decum- 
tent or suberect from a lignescent stock, finely appressed-pubes- 
cent with bronze-colored hairs: leaves 3-11-foliolate ; stipules 
persistent, 4 to 6™™ long; leaflets obovate, retuse, mucronulate, 
fnely appressed-pubescent and yellowish-green above, decidedly 
jalet, cinereous and very veiny beneath, 8 to 17™™ long, half 
* broad : flowers borne chiefly in pairs in the upper axils or 
‘oming a subcapitate raceme at the summit of the stem: calyx 
avny-villous, 5™™ long, its narrow teeth subequal: petals prob- 
” Purple: pods somewhat falcate, 3.8™ long, 5" broad, 
‘omentulose. —In pine woods on the Manatee river, S.W. Florida, 
, Pi, no. 156, June 1845. Type in herb. Gray. A character- 
fet nt teeording to our present knowledge, thoroughly dis- 
i ony with the habit of 7. chrysophylla, from which it differs 
ce °nopodial stem and axillary flowers, as well as in the 
ere pubescence upon the upper surface of the leaves. 


. rh REIneRrA BS G. 251, in part. Stem sympodial, the 
3 rae at one or more nodes terminating in a raceme which 
Sera development of an axillary bud at its base becomes 
PPosite oe Some of the racemes thus appear to arise 
: Ped in - hag (The sympodial structure is tardily devel- 
a: " Purpurea, which during its first season sometimes 
“Meret any a terminal raceme. This species, however, may 
-y alllly distinoui 
its er guished from those of the preceding section by 
ch M ow ers, which are only 6 to 8.5™™ long. )—Reimerta 
meth. Suppl. 44. 
* 


+ Leaflets ( Flowers large: petals 1 to 1.7°™ long. 


with rare €xceptions) exceeding the short petioles. 


198 BOTANICAL GAZETTE (SEPTEMBER 


+ + Flowers not numerous, borne singly or in pairs at the nodes of the 
racemes, 
== Stem covered at least below with a short dense bronze-colored tomen- 
tum: leaflets thickish, of firm or subcoriaceous texture, glabrous and 
finely reticulated above. 
a. Leaves prostrate, essentially sessile; leaflets seldom more than 7. 
T. cHRysOPHYLLA Pursh. Perennial herb with spreading 
prostrate freely branched somewhat flexuous or geniculate stems, 
subsessile, 2—7-foliate leaves, and obovate leaflets (1.3 to * 
long): few-flowered racemes opposite the leaves; peduncles 
scarcely ancipital, 5 to 8.8 long: petals white, changing to red: 
pubescence on the lower surface of the leaflets dense, sericeou’, 
somewhat canescent but with a slight golden sheen: pods 3.4 
4.2™ long, 8—10-seeded.— Fl. 2: 489; Ell. Sk. 2: 246; Torr. & 
Gray, Fl. 1: 297; Chapm. Fl. 95. TZ. prostrata Nutt. Gen. 24120. 
Cracca chrysophylla Kuntze, |. c. 174; Vail, 1. c. 34.—Dry pie 
woods, Georgia, Boykin, Forbes, to Florida, where apparently 
common, “and westward” acc. to Chapman, but probably in refer- 
ence to 7. Smallit. 
—’ Var. Chapmanni. Plant smaller, leaflets 6 to 13™ long, half 
as broad: pods only 1.9™ long, 5—8-seeded.— Cracca chorysophyl 
var. Chapmanni Vail, 1. c.—St. Josephs, Florida, Dr. Chapman. 
mostly 7 to 1! 


6. Leaves, at least in some cases, ascending, petiolate ; leaflets 
pecies in 


~ T. Smallii, n. comb.—Similar to the preceding $ 
many ways, but stouter, with more numerous and longer (oblong 
or elliptic rather than obovate) leaflets: peduncles cee y 
ancipital above, becoming 5 to 20™ in length.— Crate 
Small, Bull. Torr. Club. 21: 303. C. Smallii and ©. J" 
Vail, 1. c. 33, 35.— Pine barrens in sand, Georgia, Boh 
Florida, Curtiss, Nash, and Louisiana, Dr. Ingalls. 
use of zntermedia in Tephrosia necessitates the adoption 
second specific name. 


of the 


her small, diy 


preadig 


= = Stems very slender, sparingly pubescent : leaflets rat 
tical, thin, 
T. misprpua Pers. 1. c. Stems several, branched, : 
and ascending from a thickish somewhat fusiform 1 


The previous . 


t, finely 


1899] NORTH AMERICAN SPECIES OF TEPHROSIA 199 


pubescent or glabrate ; hairs sometimes spreading but usually 
appressed: leaflets 11 to 17, elliptic-oblong to linear-lanceolate, 
thinnish, usually deflexed, appressed-pubescent or glabrous 
above, slightly paler appressed-villous beneath, usually rounded 
md mucronulate at the apex: petals at. maturity purple, 
sometimes 1.6™ in length: pods 8-10 seeded, covered with short 
appressed or more often spreading hairs.—Pursh, Fl. 2: 489; 
Ell. Sk. 2:245; Torr. & Gray, Fl. 1: 296, excl. vars.; Chapm. 
fl 93. T. gracilis Nutt. Gen. 2: I1g. 1818. TZ. elegans Nutt. 
Jour. Acad. Philad. 7: 105. Galega hispidula Michx. Fl. 2:68. 
Cracca hispidula Kuntze, l. c. 175 ; Vail, l. c. 33—-Sandy barrens, 
Virginia and N. Carolina, Curtis, to Florida and Louisiana, Hale. 
=== Stems rather stout, covered with a copious coarse tawny spreading 
pubescence; leaflets sparingly villous along the midnerve above or 
*ppressed-pubescent over the entire upper surface. 


T. vitLosa Pers. l.c.% Stem 3 to 9 long, sprawling, tawny- 
site: leaflets 3 to 17, elliptic or obovate, rounded and apicu- 
at the end, villous beneath, more or less appressed-pubes- 
» sotapla upper surface, about 2.5°™ long, a third to half 
ancpital - one tawny-pubescent: peduncles long, somewhat 
talyx Likes e | ted few-flowered raceme surpassing the leaves: 
(at least j with long filiform tips: petals pale or more often 
age) purple red.— 7. spicata Torr. & Gray, Fl. 1: 296; 
bm. Fl. 95; Wats. & Coulter in Gray, Man. ed. 6, 133. 7. 
inn, 
—” originally described by Michaux (1803) as Galega villosa, with 
is plant = Floridam.” There is absolutely no evidence that Mic aux 
2S hed neat of the Asiatic G. villosa L. The name 7% ern 
American eg Persoon in his Synopsis (1807) and is there used exclusively 


Poted 


4 C nam he plant of the Old World, although possessing an 
: Sod that ae ‘a not brought under Tephrosia until later and, it is believed, 
ve receive another Specific designation. 


200 BOTANICAL GAZETTE [SEPTEMBER 


paucifoha Nutt. Gen. 2: 119. ? ZL. hispida DC. Prodr. 2:250, 
T. mollissima Bertol. Bot. Miscel. 9: 10. pl. 3. f. 2, and Bot. Zeit 
9'902, acc. to Gray. Galega spicata Walt. Car. 188. G. villose 
Michx. Fl. 2:67. G. paucifolia M. A. Curtis, Jour. Bost. Nat. 
Hist. Soc. 1:122. ? Crafordia bractiata Raf. Specch. 1: 156 
Cracca spicata Kuntze, |. c.; Vail, 1. c. 30.— Dry sandy ground, 
common, Delaware to Florida, west to Arkansas (ace. to Les 
quereux) and Louisiana, Hale ; fl. May to July. 
— Var. flexuosa. Leaflets linear to lance-linear, acute, the 
terminal one much elongated.— 7. flexuosa Chapm. acc. to Tor. 
& Gray, l.c. 297, insynon. T. hispidula var. y Torr. & Gray, & 
Cracca spicata var. flexuosa A. M. Vail, 1. c.— Florida, Chapman ; 
poorly known and perhaps distinct. A similar but nearly 
glabrous form has been found in Alabama by Gates. 
++ ++ Flowers numerous, the middle ones borne in threes and fours: leaflets 
also numerous, g to 27, linear-oblong, 2.5 to 3.8™ in length. ; 
T. onoprycuorpes Nutt. Rather stout for the genus, erect 
or nearly so: stem terete, geniculate, producing from near the 
summit a long-peduncled erect many-flowered raceme: leaflets 
oblong, silky beneath, obtuse or retuse at the apex, cuneate at 
the base, 2.5 to 3.8°™ long, a fourth as wide: flowers pale chang: 
ing to red or at length purple: pods secund, finely appressed: 
pubescent.—Jour. Acad. Philad. 7: 104; Torr. & Gray; ie 
1 296; Engelm. & Gray, Pl. Lindh. 1: 6, 335 Gray in 
Pl. Tex. 7; Chapm. Fl. ed. 2, 615. 7. angustifolia and © 
multifora Featherman, Bot. Rep. Louisiana Univ. 1870: | 
to Gray, Am. Jour. Sci. III., 2: 375. Cvracca onobrychoides Kun 
l.c.; Vail, 1. c. 29.—Dry plains, Arkansas, Nuttall, Harv’), 
Louisiana, Hale, and Texas. 
in 7, Lint 
+ + Petioles longer than the leaflets (rarely equaled by them 
heimeri). 
++ Pods 6 to 8.5™™ broad : leaflets suborbicular. 
herb with 


to 


T. LinpuEimeri Gray. Soft-pubescent perennial ee : 
long reclining branched stems and 5—13-foliolate leaves + fe 

broadly obovate, rounded at the end, subcuneate at the 
sericeous upon the lower surface ; linear striate attenuat 


’ 


e stipules 


este 


19] NORTH AMERICAN SPECIES OF TEPHROSIA 201 


very long: flowers rather numerous in erect pedunculate racemes, 
purple: pods velvety-tomentose.— PI. Lindh. 2: 172; Torr. 
Bot. Mex. Bound. 51. Cracca Lindheimeri Kuntze, l.c.; Vail, 
Le, 28. Sandy prairies, southwestern Texas, Lindheimer, Wright, 
Palmer, Havard, Fuchs. (Adj. Mex. where first collected by 


++ ++ Pods about 4™™ broad. 


T. AMBIGUA Chapm. Stems prostrate or ascending, from a 
: éeep woody root, copiously pubescent to nearly glabrous: leaves 
“et $-I3-foliolate, petioles 3.8 to 6.3°™ long; leaflets obovate 
to oblong, obtuse or obtusish, 2.5 to 3.8°™ long, 6 to 17™™ broad, 
sparingly appressed-pubescent or glabrate above, veins red and 
sme paler, appressed-pubescent beneath; stipules 4 to 8.5™™ 
length: peduncles long, ancipital, remotely 3—6-flowered : calyx 
,) Small: petals purple : pods narrow, appressed-pubescent, 
4 *any-seeded— F l. 96; Wood, Bot. & Flor. g5. TI. hispidula 
ae & Gray, Fl. 1: 296. Galega ambigua M. A. Cuttis, 
: i Nat. Hist. Soc. 1: 121. Cracca ambigua Kuntze, |. c. 
My ail, 1. c. 32, N, Carolina, Curtis, to Florida and Missis- 
“PPh Pollard , common, 
oe . Very slender: leaflets lance-linear, acute or 
ae os a ibd Jae in breadth.— Cracca angustissima Vail, 
Clie mA  angustissima Rugel.— Dry pine barrens near Eau 
{ huly. EP yet, Florida, 4. A. Curtiss, nos. 584, 5708; fl. 


‘ : * * Flowers smaller; petals 6 to 8.5™™ long. 
— Pers. Slender, flexuous, branching from near 
’ OF ascen Neg at first filiform at length lignescent: stem 
“$-foliolate .. finely appressed-pubescent: earliest leaves 
, & oblong, rety ater Ones 7—19-foliolate: leaflets very varia 
€W.) linea “S, or in var. angustissima (7. angustissima 
“A cae acute; stipules filiform; flowers small, 
st pair usually axillary, the others forming a 
pods spreading or nodding, 4™" 
pubescent under a lens, 3-5-seeded.— Syn. 2: 


202 BOTANICAL GAZETTE [SEPTEMBER 


329; Hook f. & Jacks. Ind. Kew. 2: 1045. T. deptostachya DC. 
Prodr. 2: 251. 1825; Chapm. Fl. ed. 2, 616. TZ. adscendens 
Macfad. Fl. Jam. 257. 7. tenella Gray, Pl. Wright. 2: 36. T. 
angustissima Shuttlew. in Chapm. 1. c. 96. Cracca purpurea L. 
Spec. 2: 752; Vail, 1.c. 31. Galega piscatoria Ait. Kew. 3: 71. 
—Sandy ridges, Florida, Rugel, Curtiss, no. 584*, Garber, Simpson, 
also from W. Texas, Havard, to Arizona. (Tropics of both 
hemispheres ). 


GRAY HERBARIUM. 


BeieErER ARTICLES: 


BQLES OF TRAVEL. II. 


PAYTA AND THE DESERT REGION OF PERU. 


Wuen Mr. Barbour Lathrop, with whom the writer is traveling as 
botanical assistant, first decided to go via Panama to the west coast of 
South America, he remarked that he would show him Payta, the driest, 
ost forsaken spot in the world. He would defy even a botanist to 
fnd so much as a single living wild plant. The donkeys of Payta are 
repted, like the locusts during early days in Kansas, to eat any green 
paint in sight, 
Fayta lies less than five degrees south of the equator in the dry 
eg of Peru, on a Coast, steadily rising from the sea in some parts, which 
tas risen as much as forty feet within historic times. So infrequent 
2h the rains On this coast that, when they do come, the whole native 
Population, with crucifixes and musical instruments, goes out to wel- 
“me the river as it slowly forces its way along the bed which for 
| Pe - been as dry as the surrounding desert. This coming 
; h generat * leates heavy rains on the west slope of the Andes and 
¥ followed by showers in the region about Payta. 

mS left Panama it was rumored there had been rain at Payta. 
: we aes Lathrop’s disappointment, these rumors were verified 
Pte sal ee off the coast, and with the glasses discovered in one 
; alleys a green algaroba shrub (Prosopis). By looking 


; ing Noon e 
7 kiled 1 Ss rain had fallen for eight years. The seventh year had 


a form, after . aud a strip of land on each side which is overflowed 
io ty the hy Subsidence of the stream, which runs only a month or 
vated land of the country back of Payta. The long 

203 


204 BOTANICAL GAZETTE [SEPTEMBER 


rooted Peruvian cotton is able to maintain itself for seven years in this 
dried-out river bed and yields profitable crops of the colored, short 
staple cotton, which is used as an adulterant for wool, occupying a 
place in the wool rather than the cotton market. A stroll to the toy 
of the nearest hill at Payta showed plainly that the rain had been a 
heavy one, for, scattered over the nearly level table land were the hard 
baked remains of unmistakable mud puddles. In these, strange as it 


Fic. 1.— Photograph of garden at Mollendo, produced by irrigaio® 


th 
seemed to us, no plants were found, although scattered owe 
sand and gravel all about were young seedlings and even acts 
grasses. ust: 
The flora of Payta would not be a difficult one to write up “al 
ively, provided one were on the spot at the right time. _ sti 
pamphlet we were able to press all the phanerogamous P plan 
were found, without any difficulty. These plants com 
whose seeds must have remained dormant since the las 


rl th ei 
: F encasing 
years before, and perennials which have kept alive by 


1899 | BRIEFER ARTICLES 205 


sues in thick layers of impermeable cork. There is something 
remarkable in the ability which these desert shrubs have of reducing 
their transpiration surface to such a degree that they can withstand the 
mtense insolation of this tropical region, and the even more trying 
influence of an extremely dry atmosphere. It is highly probable that 
they are able, during the winter season, to absorb moisture from the 
logs which are blown in from the ocean. Owing to the cool currents 


j 


ae i 
barren desert, otograph taken just outside of garden fence, showing the completely 


Of air wh; 
“Slag and winter across these deserts, they are not 
The collection [aa as would be expected in this latitude. 
two Perennials _. provisional flora of Payta—consists 
“Attals, three ce : Be and an undetermined shrub, and seven 
a beautifal © has: a lupine, a caryophyll, a seedling amaranth, 
t¥0 day. yellow flowered oxalis. Our visit to Payta was twenty- 
The i h €rain, and the grasses and oxalis were in full bloom. 
Be ita of delicate yellow blossoms scattered over the 
es, and slender blades of grass so far apart that 


after th 
Ae ht of 
Peet yb 


206 BOTANICAL GAZETTE [SEPTEMBER 


they looked like a very “bad catch” at lawn seeding, is one which few 
who have not been in the deserts in spring can imagine. Unlike the 
desert regions of our own country, with their sage brush, yuccas, and 
host of small tufts of grass and sedge, these deserts are for miles abso- 
lutely without a living plant. For days we steamed down the coast, 
but aside from an occasional garden made by irrigation in the neigh 
borhood of the towns, we saw no green plant of any description. 
From about Payta in Peru, to Carrizal in Chili, representing fourteen 
days of steamer travel, the coast presents one unbroken line of desert. 
_At Arica it is broken by a small fertile valley, and at Carrizal the 
desert ends in a scanty vegetation of cacti and low growing cushion- 
like perennials. 

At Mollendo we had an opportunity to see what this des 
have been had there been an abundance of rain. The two phot 
graphs are taken within a hundred meters of each other. One shows 
a private garden in Mollendo with apples, peaches, grapes, passion 
fruits, figs; in short a good collection of fruit and shade trees. The 
other is a representative view of the surrounding country 4 barren 4% 
a fresh lava bed. 

There are below this desert at Arica, and doubtless at other points, 
underground sources of water, for large pepper trees which a eh 
planted in the town square are growing as finely as they do in Gibral- 
tar, or southern California, and overshadowing the little clubhouse * 


ert might 


such a cool agreeable climate as this “zona sicca,” °F 
western South America. The contrast between the west and 
coasts of the continent at the same latitude is very T@ the same 
S. off Brazil, duck suits are necessary for comfort, while at : 
latitude off Peru thin flannels are quite comfortable. descrif 
For a systematic botanist, as may be judged from the abov® jologi- 
tion, there is not much of interest in this region, but from a phys! 
cal point of view it will yield some very interesting facts. ntion % 
Dr. Schimper, in his Pfanzengeographie p- 679, calls se studied 
the fact that this desert region of Peru has been very pun num 
from an ecological standpoint. It is probable that 4 ee s of squat 
ber of species will be found along this coast, and hundreés jp 
miles are absolutely without a living plant for year 
localities, however, favored by the fogs, are COV 


ered in the 


1899] BRIEFER ARTICLES 207 


season (our summer) by grasses in sufficient quantities to support 
nemerous small herds of cattle. 

The most favorable point from which to begin an ecological study 
of this desert region would be Payta and the inland town Piura behind 
i, which can be reached by railway. Having made arrangements at 
Miura for mules and a guide, the towns of Pacasmayo and Salaverry 
vould not be too far apart to serve as centers for operation down the 
coast. The discomforts of travel through this desert, I understand, are 
tot such as should deter any enterprising botanist from exploring it. 
the expenses, including steamboat travel for which the charges are 
‘wenty pounds sterling from Panama to Callao, would be easily within. 
five dollars a day.—Davip G. FAIRCHILD. 


SECTION G (BOTANY), A. A. A. S., COLUMBUS MEETING. 


THE meeting of section G began on Monday, August 21, in Town- 
“end Hall of the Ohio State University, by a brief session for organi- 
~_— In the afternoon at four o’clock, in Botanical Hall, the vice 
President, Dr. CuariEes REID Barnes, delivered an address on Zhe 
Progress and problems of plant physiology. At the close of the address, 

i ot the section were voted to the speaker. 

eading of papers began on Tuesday, when the following were 

Presented in full, or in abstract, or by title: 
“alias The fertilization of Albugo blitt. 
Le “oped : The embryo sac of Leucocrinum montanum. 

; K: Notes on subterranean organs. 


Wt Bey, 
ith their cea Some monstrosities in spikelets of Eragostis and Setaria, 
in 


Cuar : 
AD ee E. Bessey : Studies of vegetation of the high Nebraska plains. 
7 — The tamarack swamp in Ohio. 

Byron eset The breeding of apples for the northwest plains. 

fetes = TED: Field experiments with “nitragin” and other germ 
Henry |, 
g - BoLiey - 
“Witonments, ae 
W 


2 a 
Sy ywas d 


The duration of bacterial existence under trial 


y ignated Sudiivant Day, and was used to com- 
2 1883), two Ry S. Sullivant (died 1873) and Leo Lesquereux (died 
2 Through th a aa bryologists who were long residents of Columbus. 
* Initative and energy of Mrs. E. G. Britton and the 


208 BOTANICAL GAZETTE [SEPTEMBER 


assistance of a number of botanists, an exhibition of many interesting 
bryological books and pamphlets, type specimens and original drawings 
of mosses, photographs, portraits, and autographs of bryologists, maps 
of distribution, etc., was given during the day in the large lecture room 
in Botanical Hall, where Wednesday’s sessions were held. Portraits 
of Sullivant and Lesquereux, loaned by their daughters, formed the 
center of interest. ‘This exhibition attracted much attention and was 
warmly commended. 

The exercises in honor of Sullivant and Lesquereux were as follows: 

Professor W. A. Kellerman read a portion of Gray’s tribute to Sul: 
livant. 

Professor C. R. Barnes read a biography of Lesquereux. 

Mrs. E. G. Britton gave a brief account of the species of mosses 
named for Sullivant. 

A letter was read from Professor Arthur Hollick regarding the 
paleobotanical work of Lesquereux. 

Professor IL. M. Underwood gave a brief outline of the progress ® 
the study of the Hepaticze of North America, and Mrs. Britton for the 
Musci. Both addresses were illustrated by the books, pamphlets 
photographs and maps of the exhibition. 

Professor F. E. Lloyd exhibited the plates and type specimens : 
the twelve new species of liverworts described by Dr. M. A. Howe @ 
his recent monograph. 

Professor W. A. Kellerman presented to each member ™ we 
tion a set of six species associated with the names of psa 
Riddell, an early Ohio botanist. The specimens were from type! a8 
ties in most cases. They were Sudlivantia Sullivantit, Lo abe 
livantii, Arabis patens, Solidago Ohionis, S. Riddellii, and 7 
nivale, 

Mrs. E. G. Britton also distributed specimens of 
Ohioense and Bryoxiphium Norvegicum. 

The following papers were also read : 


f the sec 


Orthotrichu® 


CHARLES Mone: Notes on the moss flora of Alabam 

A.J. Grout: Suggestions for a more satisfactory 
pleurocarpous mosses. 

Bruce Fink: Notes concerning the study of lic 
Mississippi valley. 

W. C. STEVENS: Botanical teaching in the seco 

IDA CLENDENIN: Botanical teaching in the secondary sch 


a. 

classification. of the 
fe 

hen distribution in 


ndary schools. 
ools. 


1899] BRIEFER ARTICLES 209 


The two papers on botanical teaching had been prepared by 
request and evoked an interesting discussion. 

On Thursday the following papers were read : 

H. A. HARDING: On the occurrence of the black rot of cabbage in 


CHARLES E, Bessey: One thousand miles for a fern. 

Watter T. SwINGLE: A summary of our knowledge of the fig. 

Wa. TRELEASE : The classification of botanical publications. 

As a result of the discussion of this paper, transmitted by the 

Botanical Society of America, the following resolution was adopted : 
Resolved, That Section G recommends as a basis for the classification of 

* botanical library the decimal system now in common use in the United 

States. The section requests that the suggestions embodied in Dr. Trelease’s 

theme for classification be adapted, so far as possible, to that system, and 

pag paper be then published in Science for the purpose of eliciting dis- 


Epwin B. CopELAND: The geotropism of the hypocotyl of cucurbits. 
A.F. Woops: The destruction of chlorophyll by oxidizing enzymes. 


Apropos of this paper the following action was taken : 
Resolved, That Section G ex 
2 eminent physiolo 
Physio} 


press its gratification at the appointment of 
gical chemist to the staff of the Division of Vegetable 
sy and Pathology of the Department of Agriculture. 

The secretary was ins 


tructed to communicate this resolution to the 
tary of Agriculture 


C.0. 
ee gaa The effect of hydrocyanic acid gas upon the germina- 


oe: Some physiological effects of hydrocyanic acid gas 
oo a was read from the Cambridge Botanical Supply 
American b been publishing the card edition of the bibliography 
" Bibliogra a Bader the direction of the Section’s Committee 

ect Te that the publishers must terminate the present 
Of sy st . lose of 1899 on account of the fact that the num: 
Alter discussion a vee insufficient to meet the cost of publication. 

TM of Dp M © committee was increased to five by the appoint- 
. * MacDougal, of New York Botanical Garden, and J. F. 
Its a of the Buffalo Botanical Garden. The section 
to “ogg of the card index and empowered the commit- 

€ for its continuance in any feasible way. 


director 


210 BOTANICAL GAZETTE | SEPTEMBER 


On Friday the following papers were read : 


J. C..ARTHUR: The cultures of Uredinez in 1899. 

FRANCIS E, LLoyp: The embryology of Vat/lantia hispida. 

A. D. SELBY: The flora of Franklin county, Ohio. 

Erwin F. SmitH: The fungous infestations of agricultural soils in the 
United States. 

. E..Bessey: Are the trees advancing or retreating upon the Nebraska 

plains? : 

Wm. SAUNDERS: Useful trees and shrubs for the northwest plains of 
Canada. 

H. L. Botyey and L. R. WALDRON: The occurrence of calcium oxalate 
and lignin during the differentiation of the buds of Prunus Americana. 

HERMANN VON SCHRENK: Two diseases of Juniperus. 

Ws. B. Stewart: Etiolative reactions of Sarracenia and Oxalis. 

JuLia B. CLirForD: The mycorhiza of Tipularia. 

Henry KRAEMER: The crystals in Datura Stramonium. 


At 3 p.m. on Friday the section adjourned, sine die—C. R. B. 


BOTANICAL SOCIETY OF AMERICA. 


THE fifth annual meeting was held in Columbus, Ohio, August 18, 


1g, under the presidency of Dr. LuciEN Marcus UNDERWOOD. ig 
sessions were held in buildings of the Ohio State University. * 
address of the retiring president, DR. NATHANIEL LORD BRITTON, ase 
The development of the New York Botanical Garden Was ge ay 
the chapel of University Hall on Friday evening. It vet OF ; 
with its nat 

ddress 


beauties, the progress of the planting, buildings, etc. he close 
be published in full in a later number-of the GAZETTE. jee 
of the address the thanks of the audience were voted ga 
for the interesting and able presentation of facts regarding ™ a 
institution of which he is director. The following pape eer 
in full or in abstract or by title before the society. Abstracts : 
of them will be found in later pages: 
CHARLES E. Bessey: Afetaly and dieciousness. 
BRADLEY M. Davis: The sfore-mother-cells of Anthocerds. 
DANIEL T. MACDouGAL: Symbiosis and saprophytism. 
Davip M. Morrier: The effect of centrifugal force upon 
NATHANIEL L. BRITTON: The American species of Arise” 


1899] BRIEFER ARTICLES 211 


JosepH C. ARTHUR: Zhe Uredinee occurring upon Phragmites, Spartina, 
and Arundinaria in America. 

Byron D. HALSTED: Distribution of American Erysiphee. 

BraDLey M. Davis: Gametes and gametangia of the Phycomycetes. 

WILLIAM TRELEASE: CJassification of botanical publications. The society 
requested that this paper be read also before section G, A. A. A 

DanteL T. MacDouGaL: Eviolative reactions. 

Lucien M. UNDERWOOD: The foundations of genera in ferns. 

The officers elected for 1900 are: BENJAMIN LINCOLN RoBINSON, 
pretdent; Byron Davip HALSTED, vice president; ARTHUR HOLLICK, 
treasurer; GEORGE FRANCIS ATKINSON, secretary; B. ‘T. GALLOWAY 
and D. P. PENHALLOW, councillors. 

The new members elected were : J. M. Macoun, Geological Survey, 
Ottawa, Can.; W. J. Beal, Agricultural College, Mich.; C. F. Mills- 
paugh, Field Columbian Museum, Chicago; Marshall A. Howe, 
Columbia University, New York. 

Amendments to the constitution were adopted, creating as addi- 
tonal classes of members, life members, associates, and patrons. The 
new sections are as follows : 

Opa . Members-—Any member of the Society may become a 
eine Lis y the payment to the treasurer of one hundred dollars at any 
ais € membership fees shall be added to the permanent fund of 

ety, 


g a #. Associates.— Associates of the Society may be elected in the 
| a er prescribed for members, Before the first of January follow- 
anual Non, each Associate shall pay into the treasury of the Society 
Divi dues to the amount of five dollars. Associates shall have all-the 
] . aay except that of voting, and of holding office. Subse- 
the List of ti tion of this provision members shall be chosen only from 
ae Seiten lates, : 
: te payment to the treasurer of a and ioe less 
is consti ed and fifty dollars at any one time, or a bequest of such sum, 
Mall be agg donor a Patron of the Society. The names of Patrons 
: 1 sy , h the annual lists of officers and members, and the 

Society, Patro heoag to receive copies of all the publications of the 
ae ns fees shall be added to the permanent fund of the Society. 
ten the treasurer shows funds on hand amounting to 
iy was . ‘aa $1260 is on deposit in savings banks. be com- 
May be sat to invest funds where a higher rate of interest 
Society are €d than from savings banks. The total assets of the 
ee how $1517.59.—C. R 8 


212 BOTANICAL GAZETTE [SEPTEMBER 
BOTANICAL CLUB, A.A. A. S. 


Unper the presidency of Dr. Byron B. Halsted, with Professor A. 
D. Selby acting as secretary, the Botanical Club listened to the follow. 
ing brief communications: 


C. E. Bessey: A greasewood compass plant. 

C. E. Bessey: A visit to the original station of the Rydberg cottonwood. 

N. L. Britton: Report on Mr. Heller's botanical exploration of Porto Rico. 

F.S. EARLE: Tomato fruit rot. 

W. J. Beat: The botanical club of the Michigan Agricultural Be: 

Wma. SAUNDERS: The arboretum and botanic garden of the C 
Experimental Farms, Ottawa, Canada. 

LiLoyD : On two hitherto confused species of Lycopodium. 

L. M. UNDERWOOD: What shall we regard as generic types? 

L. C. Consett: A device for registering plant growth. 

A.D. SeLBy: The introduced species of Lactuca in Ohi 

O. F. Cook: Notes on some of the work of the Division of Botany of the 
U.S. Department of Agriculture. 
' Tuomas A. WILLIAMS: Some features of the investigations on grass 
and forage plants, in charge of the Division of Agrostology, U. S. Depart 
ment of Agriculture. 

J. W. T. Duvet: A brief embryological study of Lactuca Scariola L. 

N. L. BrRITTon: Notes on the northern species of “ 

N. L. Brirron: Remarks on some species of Querc 

W. J. BEAL: The introduction of Cabomba pat in Michigan. 

C. E. Bessey: The wilting of Cleome integrifolia. 


W. A. KELLERMAN: Labels for living plants. acd 
H. L. BoLttey: The position of the fungi in the plant system 4S - 
b 
y the work on the organisms of nitrification. distribution df 


L. M. UNDERWOOD: Summary of our knowledge of the 
fungi in America. 

C. E. Bessey: The powdery mildew of Polygonum aviculare. 

A. D. SELBY: On Plasmopara Cubensis. 

A. S. Hitcucock : Distribution of some Kansas plants. + plants 

W. R. Lazensy: Unusual development of leaves and growth ! P 
from cuttings. 

A. D. Hopkins : Some botanical notes by an entomologist. 

A. S. Hircucock: Some wheat crosses. 


On Thursday morning the Club adjourned sine die. 


1899} BRIEFER ARTICLES 213 


THE SEXUALITY OF THE TILOPTERIDACEZ:" 


Ectocarpus pusillus Griffiths or Acinetospora pusilla Bornet possesses 
two forms of reproductive organs, the unilocular and plurilocular spo- 
rangia, whose elements present conditions intermediate between zoo- 
spores and aplanospores. I have recently encountered a third form of 
reproductive organ, the monosporangium, identical with that of Hap/o- 
jera Vidovichit Bornet or Heterospora Vidovichii Kuckuck, and this 
last species ought henceforth to be placed in the genus Acinetospora. 

These monospores are uninucleate, like the so-called oospheres of 
Saphospora speciosa (said to be the sexual form of Haplospora globosa), 
sad are covered by a membrane derived from the interior of the 
shosporangium, as are likewise the 4-nucleate monospores of Hapdo- 
pera globosa. Since they possess a membrane before their dehiscence 
they cannot be fertilized. On account of the frothy structure of their 
protoplasm and the great variation in their dimensions, they resemble 
neither oospheres nor spores. They germinate readily in cultures and 
teelop litle plants that bear the same kind of reproductive organs. 

ia them as gemmae or propaguda, and the organ that contains 
ee P ‘eudosporangium. From what we know of the Cutleriacee 
bsporg sel It is possible that the plurilocular sporangia of Acine- 
Psi metale organs whose oospheres germinate partheno- 
The wee € am will undoubtedly be found some day. 
lor Tlopteris fonclusion as to the nature of the monospores is justified 
and Haplospora. We are no longer able to admit that 
form ora qeures and oospheres similarly situated, of like 
and identical ee ventions, with the same protoplasmic structure 
lly while : ods of germination, but that the toi germinate 
{ Satisfactor € oospheres develop parthenogenetically. It seems 
~— Propagula ang . to say that one or the other of the elements are 
"© germinate at the 4-nucleate propagula are those that have begun 
upon the mother plant. 

We shall say then th Pp 
tige the ealy 4 : n that according to the present state of our knowl- 
and a. representatives of the Tilopteridacee, Tilopteris, 
‘ttheridia of Tilo Pree themselves solely vegetatively. The 
, th rudimentar Pteris give rise to true antherozoids that really seem 
a y structures, but it is by no means certain that they 
Bora et Ota is a résumé by the author of the paper entitled “‘ Les Acineto- 
WD, Bradley M. Dae’ Tilopteridacées,” Jour. de Bot. 13: —. 1899. Translated 


214 BOTANICAL GAZETTE [SEPTEMBER 


are the same as those organs of Haplospora, however similar the type. 
We do not know of female organs, but we may foresee that they 
would have the form of plurilocular sporangia with a median cavity 
and a separate dehiscence for each little cell. Similarly one may 
foresee that the antheridia of Acinetospora will be provided witha 
median cavity and that the dehiscence will be common and terminal. 
The Tilopteridacee have no affinity with the Fucacez with which 
they have often been placed in the classical works. On the contrary 
they approach very closely the Ectocarpacee and Cutleriacee. They 
may be divided into two tribes, the ACINETOSPORE# (genus Acineto- 
spora), more closely related to the first ; and the HAPLOSPORE# (genera 
Tilopteris and Haplospora), more closely related to the second. 
When these plants shall show us organs that are as yet unknown We 
_ shall without doubt have to raise the preceding tribes to the rank of 
families, the Acinetosporacee and Haplosporacee. I have shown in 
the table below a way in which the affinities of these plants may be 
represented.—CaMILLE SAUVAGEAU, Dijon, France. 


Ectocarpus 


fronds monosiphonic 


_—— 


Choristocarpus Acinetospora 
ous 
propagula exogenou propagula endogen 
pet from an stn cell. growth tric chothallic. 


Antheridia with Tilopteris 
a central cavity 20, 
ogee from Haplospora 


ase 


Cutleriacee 


Sphacelariacee —— 


BRIEFER ARTICLES 215 


FLOWER VISITS OF OLIGOTROPIC BEES. 


Since the proof of my last paper in this journal was read I have 
had occasion to doubt the correctness of the statements regarding 
Epeolus, on pages 35-37, and have asked Mr. Ashmead if he still held 
ihe views credited to him. In a letter of August 2, he writes that 
the nests were evidently made by Zxtechnia taurea and that the 
Epeolus was merely entering them, not making them, as he supposed 
athe time. Epeolus, therefore, comes under the same category as 
Nomada. After all, the phenological position of the genus corre- 
sponds pretty well with that of Melissodes upon which most of the 
species are probably inquiline. 

In the table of oligotropic bees Xenoglossa cucurbitarum should be 
ncluded. Lately I have found it collecting pollen of Cucurbita pepo. 
It also visits Citrullus vulgaris, Asclepias Cornuti, Ipomea nil and J. 
fandurata, It has been taken at Ames, Iowa, by Miss Alice M. Beach, 
on flowers of “summer squash ;” at Mesilla, New Mexico, by Mr. 
Cockerell, on flowers of Cucurbita perennis; at Metropolis, Ill., by 
Mr. Hart, ou Martynia proboscidea, and is mentioned in the GAZETTE 
17: 65) under the MS. name X. brevicornis. 1 suspect that all of our 
‘Peciés of Xenoglossa get their pollen exclusively from Cucurbitacez. 
~CHARLEs RoBERTSon, Carlinville, Ils. 


QUERCUS ELLIPSOIDALIS IN IOWA. 


Mr. WILLIAM D, Barnes, of Morgan Park, Illinois, has placed in my 


ay €ns of an undetermined oak, collected by him in 1895 at 
Scott county, Iowa, which proves to be Q. ellipsoidalis Hill. 

Of Scott robe nigh collaborators are preparing a catalogue of the plants 
re uscatine counties for the Davenport Academy of Sciences, 

2ote accom = unable to determine the name of this oak. The field 
ith the sa os = Specimen reads : “Tree with smooth bark, and 
leaves quite ae aspect of Q. palustris.” It isa fruiting branch, the 
Chicago, Nee — rather narrower than those commonly found near 
St0 size m ray ee f frequently be seen on individual branches, or 
"y characterize nearly an entire tree. The acorn is one of 

Closely resembling the one figured in Plate J/, ¢, 


a zal kind, 
: YTANIC. 
ALG ETTE, March 1899.— E. J. HILL, Chicago. 


AZ 


216 BOTANICAL GAZETTE [ SEPTEMBER 


A NEWLY OBSERVED STATION FOR GAZINSOGA 
HISPIDA 


SOME time ago the writer’ called attention to the fact that besides 
Galinsoga parviflora Cav. and its var. hispida DC., another quite dis 
tinct species, G. Azspida Benth., had been found in the Atlantic states, 
occurring at Camden, New Jersey, where it was collected on waste lands 
by Mr. C. F. Parker. This species was again found last October by 
Mr. Howard Schriver at several points in Cumberland, Maryland. Mr. 
Schriver, noticing its differences from G. parviflora Cav., sent specimens 
to the U. S. National Museum, whence they were kindly forwarded t 
the writer by Dr. J. N. Rose. 

Mr. Schriver reports the plant as not only forming a large mass of 
vegetation (5 or 6 meters long) near a “ bonded warehouse” or sort 
of cattle depot, but also extending down the banks of an adjacent 
stream where individuals were scattered on steep cliffy slopes of Helder- 
berg limestone, even reaching the water’s edge. A sporadic specimen 
was also found on a country road not far away. While the presence in 
the neighborhood of the cattle yard, a railway, and a distillery would 
readily account for the introduction of plants from a distance, the — 
ent species in the luxuriance of its growth and tendency to spread = 
the indigenous vegetation would suggest that it has found congenial 
conditions and is likely to persist. The species is readily distinguished 
by its “pink” or “red” rays (drying purple) and by it short i 
which is about half the length of the achene. In foliage and ml 
copious spreading pubescence it resembles G. parvifiora, bho hispide 
DC., but both G. parviflora and its variety have white rays 
longer pappus (nearly or quite the length of the achene). 

As G. hispida Benth. is likely to be found at other static 
middle Atlantic states it may be worth while to cite its more impo 
synomymy, which is as follows: 

Vargasia caracasana T)C. Prodr. 5:676. 1836. 

Galinsoga hispida Benth. Bot. Sulph. 119. 1844- 

G. brachystephana Regel, Ind. Sem. Hort. Turic. 2. 184° pot. Fe 

G. caracasana Sch. Bip. Linnaea 34: 529; also Bull. Soc: : 
18:86. 1865.—B. L. Rosinson, Gray Herbarium. 


and much 


tations in the 


* Proc. Am. Acad. 29: 326. 


CURRENT LITERATURE. 
BOOK REVIEWS. 
Engler and Prantl’s Pflanzenfamilien. 


___ Titi great work has been noticed from time to time in the BOTANICAL 
: GAZETTE as the various parts have appeared. But now that Volumes II-IV 
_ ‘Secomplete, which contain the siphonogams, the time seems appropriate 
4 for a more extended notice. The first part appeared in 1887, and twelve 
_ eas later the three volumes of siphonogams were finished. The publication 
tthe three volumes of Bentham and Hooker’s Genera Plantarum, covering 
4 the same ground, but with no such breadth of treatment, extended from 
1862-1883, a period of twenty-one years. There is no definite statement as 

‘the completion of Volume I, devoted to cryptogams, but several sections 
4 a been published, and other parts are appearing with reasonable 


-— Sofaras stat 
‘SS the indexes and the cryptogams, 


are 6997 in number, the original illustrations 3026 (woodcuts 3023, 
* ete and the individual figures 19,366. The total price is AZ 436, 
ow even half morocco volumes 47 474.50. ; 

ng Considers such details he is impressed by the magnitude of the 


and still more by the organizing power which has kept the large plans 
m 


Gr, 
oe Previous general works in its spirit, it is alone in the 
id beauty of ts illustrations. Every family is thoroughly and 


217 | SEPTEMBER 


218 BOTANICAL GAZETTE | serrennet 


admirably illustrated, and we venture the prediction that many of these figures 
will become classic in future texts. No such collection of figures represent. 
ing the plant kingdom exists, and they give a conception of plants in general 
that can be obtained from no other publication. The figures and text include 
not merely those structures which may be said to have taxonomic importance, 
but anatomical peculiarities of each family are set forth. All through the 
work the ecological standpoint is prominent, and the sections on geographical 
distribution are among the most valuable. 

It is to be expected,that the treatment is unequal, and the different pars 
of very different degrees of merit, but with fifty-seven collaborators this could — 
not be avoided. It seems to most botanists far more important to complete 
a work within a reasonable time, and so establish a usable datum-line, than 
to drag it out indefinitely and allow one part to be out of date before another — 
is published. In general the treatment will be regarded as conservative, 
there being apparent no desire for change if existing lines can be used at all 
In so delicate a matter as nomenclature, as is well known, the “Berlin rules,” 
which are in fact the Engler rules, are drawn up in the spirit of compromise, 
not going to either exireme, and probably satisfying neither set of extremists 
No set of rules proposed, however, has had as yet such a tremendous advat- 
tage of general usage as this great work will compel for the Berlin rules 

It is impossible to mention in detail the views advanced as tothe evolutien 
ot plant groups. There will be much difference of opinion as to minor points, 
for many smaller groups, through lack of adequate investigation, had to be 
“lumped,” but in the judgment of the reviewer the main lines of evolution 
suggested will stand, which are in brief as follows: spiral arrangement and ‘ 
indefinite numbers to cyclic arrangement and definite numbers ; naked flowers : 
to differentiation of calyx and corolla; apocarpy to syncarpy; pale ; 
sympetaly ; hypogyny to epigyny; actinomorphy to zygomorphy. - 

h can be no doubt, but 


” 


cases of ‘reduced flowers’? occur there : 
sae imitive in 
great majority of so-called cases of reduction are really primitive 


acter seems hardly less doubtful.— J. M. C. oe 


Ferments and fermentation. ee 

THE attention which the various problems connected with poe 2 
have received during the past decade and the interest, both then doobly 
practical, which attaches to the investigation of these problems pare. 
welcome a book on the soluble ferments from the hand of Prot ane 
Reynolds Green.t In it he has sought to bring together, 5° wie esses of 
the results already reached, and to indicate the view of the DF 


‘GREEN, J. REYNOLDS: The soluble ferments and fermentati Th Macmilla® 
480. Cambridge: The University Press. 1899. 125. [New York: 
Company. | 


) CURRENT LITERATURE 219 
fermentation to which these results point. For it can hardly be said that the 
sults now attained furnish any adequate explanation of fermentation ; though 
__ theyremove it more completely from the realm of so-called ‘‘vital”’ action, 
they refer it to the category of equally inexplicable catalytic phenomena. 
After discussing the nature of fermentation and its relation to enzymes, 
_ the author gives a detailed account of diastase (60 pp.), inulase, cytase, sugar- 
_ @litting enzymes, glucoside-splitting enzymes, proteolytic enzymes (57 pp.), 
_ fatsplitting enzymes, clotting enzymes (48 pp.), ammoniacal fermentation, 
_ midases, alcoholic fermentation, and the fermentative power of protoplasm, 
4 The work closes with chapters on the secretion, constitution, and mode of 
a ation of enzymes. Of course the treatment of these topics includes a dis- 
_ Gission of the discovery, occurrence, preparation, and behavior of enzymes in 
_ ‘eth animal and plant bodies. 
Awork like this is not open to adverse criticism. ° It is rather to be com- 
mended without stint. Indeed, every physiologist will be thankful to know 
and to have at hand this compact but full summary of researches, accompa- 
: = itis, by an extensive bibliography, leading to further details in the 
‘ginal papers. One notices with pleasure that even the results contradictory 
to the general trend of investigation are clearly stated. This engenders con- 
the fairness of the work ; a confidence which the closest scrutiny 


Py The bibliography, which the author modestly says is not exhaustive, is 
2 | be po ended. Its extent will be appreciated when it is stated 
| wh ae about 800 titles! The citations would have been improved 

taken from orm system been followed. Here is an assortment of five styles 
: L(BBo) 2313 (4) (1) Zeit. f, Biologie Bd. x. 92; (2) Zeitsch. f. klin. Med. 
- (08y4), or ' © Cent. f. Bact. 1891. Bd. 10. 401; (4) Cent. f. Bakt. 15 

them, Gesell s er. d. deut. chem. Gesell. 23 (1890). 3689; (6) Ber. d. deut. 
ey 95), 1433. This will be looked upon by many as of small 
the defect ig Pag so it is, in comparison with the value of the work. But 
pity that it OP eaeadl and so easily avoided that it seems the greater 
to mar so good a bibliography. Moreover, calling attention 


it here ma 
eed to lay i . to emphasize a point which many scientific writers sorely 
, 4 7 
U 


Users of this wes. ._- 

Wal Sez, 5 work will be thankful that the editor of the Cambridge Nat- 
the aut [. of which this is oue, had an index prepared, in spite 

that the peeing that one was unnecessary. They will only be sorry 

‘thors, for ls not fuller, and that it does not include the names of 


: whi ae 
hich the bibliography must be consulted.—C. R B 


220 BOTANICAL GAZETTE [SEPTEMBER 


NOTES FOR STUDENTS. 


BEFOKE THE Botanical Society of America, at the Columbus meeting 
Professor D. T. MacDougal read a paper on Symbiosis and saprophytism, oh 
which the following is a synopsis : 

At the last meeting of the society I read a paper in description of my work 
upon a large number of herbaceous mycorhizal plants,? and a short note was 
presented before the meeting of the Society for Plant Physiology and Mor 
phology, in December 1898, in which a delimitation of the terms saprophyte 
and symbiosis was attempted. Attention was also called to the fact that 
only two seed plants, Wullschlgelia and Cephalanthera, may be truly des 
ignated as saprophytes, all other species of so-called holosaprophytes being 
symbiotic with mycorhizal fungi. a 

. During the last year my efforts have been directed first to ascertain the 
adaptations undergone by these true saprophytes, and compare such changes 
with those undergone by mycorhizal forms. Secondly, evidence which might 
have a bearing upon the physiological relations of a seed plant and its 
mycorhizal fungus has been carefully sought for. 

Cephalanthera, the saprophyte examined, showed alterations in structure 
generally similar to those of mycorhizal forms; but, in exception to the major 
ity of chlorophylless species, it retains the stomata of the leaves, and has 
developed no underground transpiratory organs. ky 

The roots, unlike those of most mycorhizal forms, are deeply buried in the 
soil, on account of which the number of good herbarium specimens to be 
found is extremely small. Two types of these organs are present : a fibeenyy 
with a reduction and fusion of the stelar components, and with radially elo® 

gated cortex. This variation has hitherto been regarded as a result dm 
presence of a fungus as in mycorhiza, while as a matter of fact It's goed 

tation to humus foods. The second type of root is devoted to storage an silk ; 
a normal multiplication of the cortical elements. Both types per 
with the two kinds of roots formed by Wullschlegelia, which still 


e chemical 
ferentially 


have taken place, entailing also unusual osmotic conditions. 
two seed saprephytes live like fungi, although capable 0 

The coralloid formations on the offsets of Calypso, which are 
Norway to Washington, have been examined, and the result of ne eh 
ment of these adventitious mycorhizas is quite similar a coupled 
described in Aplectrum. The occurrence of the coralloid mycorhiz@ BY” © 


-® Published in the Annals of Botany 13:—. March 1899- 
\ 


1899] * CURRENT LITERATURE 22% 


witha diminution of the leaves, the roots, and the storage organs, indicating 
ani acquisition of highiy organized food-material ; a variation which 
wil have the same ultimate influence on the species as in Aplectrum, and is 
toubtless responsible for many of the so-called aberrant forms of leaves and 
fowers reported. 

This species and Corallorhiza Arizonica were examined with especial 


tas lost the stomata. The subterranean rhizome and its coralloid branches 
we each furnished with separate types of motile stomata, however, which 
fanction during the entire year, but clearly serve transpiratory and respiratory 
needs alone, 


The arrangement of the fungus is one which has been found in nearly all 
. ete endotropic mycorhizas hitherto examined, and its study has yielded 
me important results, 
The hyphae of the fungus extend themselves in comparatively straight 
threads longitudinally toward the tip of the organ in the sub-epidermal region 
“ttecortex, and this portion remains alive even in old members constituting 
what may be designated as the “vegetative mycelium.” The entrance of the 
gtd vegetative mycelium into the young cortical cells causes almost 
nee mm the character of the latter, and no important interchange 
: the two plants ensues in this region. 

” Srowth of the root and the contained vegetative mycelium goes on, 
ea “Sagi me mycelium sends out hyphal branches which extend 
: hgccy: - epidermis and may or may not traverse the hairs into the 
: wother set of very clearly absorbing organs. At the same time 

®t up and eae ches penetrate to the median region of the cortex and both 
tracted to the ae profound disturbances. The tips of the hyphe are 

A i Geied 0 of the nuclei. Dense coils and sometimes large 
| which serve as organs of interchange between the fungus 
: these seins Pa The starch contained in the cortical cells inhabited by 
"600, the organs of a the fungus; then, with the maturity of this general 
mterchange and the large amount of contained proteid 
a? te higher plant. 
~The seg Slant ; aracter of the partnership of the two plants : 

: fingus, and yields Eruiics a habitat for the vegetative mycelium of the 
feagus takes up sa pertain carbohydrate foods, principally starch. The 

“AS, conducts Say Products from the soil by means of its external 
mat and man €m to the inner branches in the cortex of the higher 

mactures.proteids, of hich ion i d in its own met- 

ee of course, but th es ty) — a portion Is use 
4 lfollowsas a nec € greater part is yielded to the seed plant. : 
that mycorhis “4 ary Corollary from the above conclusions, that Frank's 
a’ adaptations are fungus traps and that the seed plant 


: ” St free and beco 


222 BOTANICAL GAZETTE [SEPTEMBER 


derives the entire advantage from the association is no longer tenable, Like 
wise the theory of Janse, that endotropic fungi which form mycorhizas are 
negatively chemotropic to oxygen and bear the same relation to the seed plant 
as the organism in the leguminous tubercle, is not capable of universal appli- 
cation. Such relation has been proven by Nobbe and Hiltner between Podo- 
carpus and the peronosporous fungus which forms endotropic mycorhiza with 
it, but in no other instance. 

At this stage of our investigations, thea, two distinct physiological types at 
endotropic mycorhizas are recognized: One adapted for nitrogen fixation, 
and a second for the absorption and modification, perhaps oxidation, of humes 
products by the fungus and their liberation in the tissues of the higher plant 
The greater number of examples are included under the last type. 

Ec.otropic or sheathing mycorhizas such as Monotropa perhaps appro 
imate more nearly to the latter type.3 


HE PAPER read before the Botanical Society of America, at Columbus, 
entitled “ The effect of centrifugal force upon the cell,” by Professor D. M. 
Mottier, discussed in detail the effect of centrifugal force varying from 1800 
to 1900 times that of gravity, acting for definite short periods of time, ap 
cells of certain alge, mosses, and phanerogams. In all cells operated with, : 
the movable plasmic contents, together with the inclusions, were made to 
fall into a compact mass at the end of the cell. In cells which were not 
killed outright or too badly injured, so that death resulted soon pages 
the displaced cell-contents gradually redistributed themselves in due cou 
of time. ae 

The most strikingly interesting phenomena are presented by dividing, 
cells and in the behavior of the nucleolus. In dividing cells of ae 
as Cladophora and Spirogyra, the cell-wall in process of formation “ak 
time of centrifugal action was never completed. The chloroplast pS 
and other displaced contents, on becoming redistributed, pass ieee we 
the opening in the partly formed cell wall provided this opens a 
ll : 


too small. edistributed, 
meen 


amount of chlorophyll, and a larger one partly colors 

chlorophyll. In certain cells of the plerome of Zea Mays nbrane inte 

ogams, the nucleolus was thrown out through the nuclear mem 

cytoplasm. These nucleoli never re-entered the nucleus. the dividi"g 
Other important observations were presented touching it ae 

cell and nucleus.‘ ical Ch 
3The full paper will be published in the Budletin of the Torrey ae a 
*This paper is published in the Annals of Botany for Septembet 


NEWS. 


Dr. E. B. CopELAND of the State Normal School, Chico, California, has 
been appointed instructor in botany in the University of West Virginia. 


Dk. Henry C. Cow.es, of the University of Chicago, has spent several 
Weeks with a party of advanced students at Marquette, Mich., prosecuting 
ecological studies on the adjacent flora. 


Proressor P. H. Rous, formerly of Lake City, Florida, has accepted 
2 position at Clemson College and Experiment Station. His post office is 
| Gemson College, S. C. 


| TW Sutrusontan Instrrution is soon to appoint an assistant in crypto- 
gamic botany, with a salary of $75 a month. The civil service examination 

_ "ilbe held November 5 and 6. 

WE LEARN FROM Science that Aven Nelson, botanist of the Wyoming 

periment Station, is making this summer an extended survey of Yellow- 

“one National Park and adjacent forest reserves. 


THE OFFICERS-ELECT of the Botanical Club of the A. A. A. S. are Pro- 
W FS, Earle, of Auburn, Ala., president, Professor A. D. Selby, of 
"ster, O., vice president; and Professor F. E. Lloyd, of New York, 
‘retary, ‘ 


MR. Ge 


ORGE T. Moore, who has Been an assistant in the cryptogamic 
ry at Harvard University, has accepted am instructorship in Dart- 
College, in charge of botany. This appointment recognizes botany 
a8 a sister department in zoology. 


: i . . . 
yay SUMMER has witnessed the establishment of another inland biologi- 
~Eigenmann pine ib at Winona lake, under the direction of Professor 


_ With Professor D, M. Mottier in charge of botany. It is closely 
eile With Indiana Waiver 


sity. 

T : 
bas . DePARrMeNT OF Botany of the Marine Biological Laboratory 
| Yery successful session. Dr. Davis and Mr. Moore, of the staff, 


ne "ie Atkinson, MacFarlane, Mottier, Penhallow, Kraemer, and 

MacDougal, Shaw, Townsend, and Smith made shorter visits to 

Never before have so many botanists been present during 
\ 


223 [SEPTEMBER 


224 BOTANICAL GAZETTE 


“ MouLpbs, MILDEWS, AND MusHrooms,” is the alliterative title of a 
guide to the systematic study of the fungi and mycetozoa and their liters — 
ture, by Professor Lucien Marcus Underwood of Columbia, author of Ox 
Native Ferns and their Allies. \t is to be issued shortly by Messrs. Henry — 
Holt & Company. : 


Dr. WALTER S. SWINGLE has returned to this country after several 
months of travel abroad. He has brought with him many interesting and 
valuable economic plants that will be given an opportunity to make this 
country their home. The date palm and the truffle are, perhaps, the best- 
known forms whose introduction into this country would be greatly welcomed. 


THE SECRETARY of Agriculture has planned “a publication which shall 
contain a résumé of the achievements of the United States in every branch 
science as related to agriculture during the nineteenth century, for distribu- 
tion at the Paris Exposition.”” Each of the bureaus and divisions charged 
with scientific work has been directed to contribute one or ype 


land owners to apply the principles of forestry. The division solicits cores 
pondence regarding every such effort in order that it may make a pr 
showing of the extent of this sort of work. 


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OCTOBER 1899 No. | 


THE 


TANICAL GAZETTE — 


EDITORS 
M. OULTER, The University of Chicago, Chicago, Ill. Ces 
RLES R. BARNES, Zhe University of Chicago, Chicago, Hl. ee 


J go ARTHUR, Purdue Oniversity, Lalgperte Ind. 


ASSOCIATE EDITORS 


DeCANDOLLE FRITZ NOL 
A nies = Soon 
- VOLNEY M. SPAL 
ry 9 of Padua Oni a ¥ / Michiga ite 
at ROLAND THAXTER | 
he of Berlin Ha sets University 
Zz WILLIAM TRELEASE 
de Pharmacie, Paris Missouri “Baa Garden 
ARPER H. MARSHALL WA oa 
¥ oy Wisconsin Conte ve Cambridge : 
oo. EUGEN. WARM Set ia 
niversity, Tokyo Un pe tas Cees ae 


s EIT WITTROCK 
yal Academy of Sciences, Stockholm 


e) CHICAGO, ILLINOIS 2 eee : oe 
ublishey by the Gnibversity of Chicago ee 


(be Aniversitp of Chicags eres 


vaso 1899 BY THE UNIVERSITY OF CHICAGO 


7 
a 


i) 


: ; 4 
Botanica 
 Fournal Embracing all Departments of Botanical Science 
Single Numbers, 40 Cents 


ription price must be paid in advance. No numbers are sent nr the expiration 
of the time paid for. No reduction is made to dealers or agent 


FOREIGN AGENTS: 
WESLEY & Son, 28 Essex Germany — GEBRUDER BoORNTRAEGER, Berlin 
18 Shillings, SW. 46, Schiuchoabeden 17a. 18 Marks 


Issued November 23, 1899 


& CONTENTS 


as a ace BLITI. CoNnTRIBUTION FROM THE. HULL 
ABORATORY, XV L. Stevens 225 

DEVELOPMENT ¢ OF THE HOLDFASTS OF rigrtve ages FLORUEA 3 
$ XXI-XXIII AND FIVE TEXT- -FIGURES.) Carrie M. Deri 246 
ES. 


CAL Rerorn IN THE NOMENCLATURE OF CULTIVATED PLANtTs. Wilhelm 


264 

AL GARDEN AND INSTITUTE IN Papua. /. B. DeToni - : +e 
* FROM MY HERBARIUM. W. W. Ashe - : : * ree 
HIGAN FuNGI, 2. O. on oi - - - * : 272 

- W.D. Holw 273 

S. John M. oo. - - 275 

: 276 


ial AND SmpvnE An Eco.ocicaL TEXT-BooK 
TH’S pe eeon: 


€ (not is than 50 co 1 ecial order in advance 
pies) will be printed on y upon sp hers 
five aoe oon covers) will be furnished grafts to authors of — articles ; 


50 roo 150 sees 
oe : : $2.25 “17 $a00 
tke Cs i haben neat $1.50 eo? 
: GazerrE cover)... $1.00 $1.50 $2.25 Aaite 


tributors are 


mes with particular 
Natale age to write scientific and proper na 


t 
nin the pages of the GazeTTE. Manuscripts should be sent to 
of 


hould be sent to the : 
. wil be replaced fre vee only when am G is gute oe thirty days after receipt of ~ 


ity Press Divito all business correspondence should be addressed to The — 
and ion, Chicago, Ill. 
a atom be made payable to The University of 


Orrice at Curca AGO, ILL,, AS SECOND-CLASS POSTAL MATTER.| 


Every Botanist 


Should be familiar with 
the prominent works of 


GEBRUDER BORNTRAEGER 
Publishers. 


JAHRBUCHER FUR WISSENSCHAFTLICHE BOTANIK, begriindet we) 
Pringsheim, herausgegeben von W. Pfeffer, Leipzig, und E. Strasberg 
Bonn. : 
JUST’S BOTANISCHER JAHRESBERICHT, herausgegeben von E. Koka 
Referirt alle botanischen Arbeiten. 


BERICHTE DER DEUTSCHEN BOTANISCHEN GESELLSCHAFT. 
BERICHTE DER DEUTSCHEN PHARMACEUTISCHEN GESELLSCHAFT. 
VERHANDLUNGEN DES BOTANISCHEN VEREINS DER PROVES 
BRANDENBURG. 

DIE GARTENKUNST, Zeitschrift fiir Gartenkunst und Gartentechnik. 
oe 

Symbolae Antillanae seu Fundamenta Florae Indtae occidentalis, edidit Ing. © q 

ol. I, fasc. i. Mk. 10.80. Will be continued. 

Syllabus der Vorlesungen iiber specielle und medicinisch 
tanik von Ad. Engler, Berlin. Zweite Auflage. Cart. 


-pharmaceutise™ of 
Mk. 3.80 
Lehrbuch der ikologischen Pflanzengeographie von Eug. Warming Kopel" 
Deutsche Ausgabe von Dr. Knoblauch. Cloth, Mk. 8.00. — 
flanzen und Mise 


Kleinasiens Naturschatze, seine wichtigsten Thiere, Kulturp c 1 
nd 2 Plane 


schatze, von K. Kannenberg. Mit 31 Illustrationen u 


Mk. 14.00. 


Write for free Catalogue; postpaid. Address: 1 
ishers 
Gebriider Borntraeger, Publis 4 
BERLIN, > W 7 


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There is also the L[ntroductory Year for young pupils, which gives 
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Morgan Hall jor girls, West Hall and East Hall for boys, 
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The laboratories are amply furnished with apparatus for 
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The library contains four thousand well-selected books for 
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a" unequaled opportunity (1) jor students who desire to 

"“b deficiencies in college preparation or prepare fi al 
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For ven : 
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The Dean of the Academy 


MORGAN PARK, ILLINOIS 


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NUMBER 4” 


BNICAL CsAZETTE 


OCTOBER 1899 


OMPOUND OOSPHERE OF ALBUGO BLITI. 
BUTION FROM THE HULL BOTANICAL LABORA- 
ORY AVE; 
F. L. STEVENS. 
[Concluded from p. 176.] 
RIPENING OF THE OOSPORE. 


Process by which the oospore attains its mature condition 
y divided into three stages, which are marked by changes 


~~ nand the exospore nearly completed, and (3) a period 
lich the Secondary endospore is formed and the oospore 

'Y Tipe and ready for its winter rest. 

ei in the oosphere withdraw from the immediate 


“ve previously enlarged and the cytoplasm is more 
a0 at any earlier stage. The primitive wall becomes 
“075 thick 


. 


225 


226 BOTANICAL GAZETTE [ocToper 


At this time a peculiar substance appears in the interstices of 
the ooplasm and fills the large vacuoles. It is not seen in younger 
stages, but is constant at this period and takes a rusty color when 
Flemming’s triple stain is used. When the substance first 
appears the tint is barely perceptible, but later it assumes a brick- 
red color. Slightly stained sections show a film in the vacuoles, 
often so curled up that an edge view is obtained, thus showing 
that the vacuoles actually contain a stainable substance. This 
substance is present in stages such as those represented in igs. 
ee YA 959 94- 

As the primitive wall reaches maturity it becomes covered 
by a layer of semi-transparent substance which is penetrated by 
straight pores. The condition is illustrated in fig. 92, where 4 
portion of this layer lies in such a position that both a surface 
and an edge view appear at one time. As the pores are close 
together and straight the structure reminds one of the hymenial 
surface of the Polyporei. Since the main substance of the layer 
is nearly invisible, the pores appear dark and look like papilla 
or cilia growing out of the primitive wall when seen in edge 
(fig. 92). A condition was noted by Wager in A. candids, and 
referred to as the “columnar condition,” which from the dese’ 
tions and drawings seems similar to this. There appc? eae : 
what later in this semi-transparent layer peculiar saucer-shaped 
masses of a dark substance of unknown nature (jig. 91): — 
are the first indication of the ridges so characteristic of 


mature epispore. There is but little periplasm at this 
y still undim" 


htly and show 


only a membrane and nucleolus. They frequently mass an 
in bunches, but there is no evidence that they 
furnish material for the exospore as is described for A. é 
To complete the endospore the saucer-shaped Bd 
ously mentioned (jig. gr), extend laterally and run es shows : 
forming the ridges. This gives the epispore the wee 
in fig. 93. Nuclei persist much longer outside of the “llowed 
but they seem to be functionless, and their history was not 


previr 


_ e] THE COMPOUND OOSPHERE OF ALBUGO BLITI 227 


q further. The mature endospore consists of two layers that may 
_ tecalled the primary and secondary endospores according to | 
their development. The primary endospore appears as an even 
ayer lining the primitive wall, at first agreeing precisely in stain 
_ faction with the gelatinous material of the vacuoles, which is 
Still present (fig. 93). The primary endospore, however, soon 
_ Wtluses this stain and gives the reaction of cellulose with chlor- 
4 iodide of zinc. A variation in the normal sequence was found in 
_ M€oospore in that the epispore was completed before any trace 
_ Of endospore was to be seen. 
j The conspicuous feature of the third period is the laying down 
_ ofthe secondary endospore inside of the first. It is of cellulose, 
and is about equal to the primary endospore in thickness. The 
q Wo layers of the fully developed endospore are clearly evident 
; when stained, also when they cleave apart in a perfectly regular 
mtd as they frequently do ( fig. 97). There seems to be a 
__ Pase in the laying down of cellulose when the primary endospore 
3 “completed, as is shown by the fact that perceptible changes 
| Sur in the development of the epispore after the first and 
E belore the second endospore is formed. Fig. 94 shows the 
4 — trace of the secondary endospore that was seen. When 
_ “second layer of cellulose is completed the gelatinous material 
: completely disappeared from the vacuoles. The fact that 


this 
Substance appears just before the laying down of the primary 


oo young, and the disappearance of all of the gelatinous 
“multaneously with the completion of the secondary 
a significant facts, They indicate that this substance 
Of the , Y connected with the formation of the cellulose walls 
ees to the a ppears to be transferred directly from the 
Re cellulose. exterior of the protoplasm, there to change to 


ol iia food substances accumulate in the center of 
Wotie 5 taking the form of one very large irregularly shaped 
Mtaining ie Around this mass lies a zone of cylopaee 
: © fusion-nuclei in resting condition. Their number 


228 BOTANICAL GAZETTE [ocToRER 


is about one hundred, so it is not probable that there has bees 
any multiplication since fertilization. Mitotic figures have never 
been seen in the oospore after that act, and it is probable that 
the oospore persists in this condition until the following spring. 


OILS AND OTHER FOODS. 


A thorough discussion of this subject would involve a much 
more elaborate microchemical study than was undertaken. It 
seems desirable, however, to record a few observations since 
these extra-protoplasmic substances are very conspicuous at cer 
tain periods of development and their presence often introduces 
serious difficulties of technique. No writer has given the sub- 
ject the attention that it deserves, and detailed study would 
doubtless yield valuable results. 

There are three substances which, when present, always 
appear as nearly spherical globules with the general appearance 
of oil, and they have undoubtedly been described as such by 
many previous observers. Further study may show that they 
are not true oils, and this seems quite probable, as they do ” : 
answer to all of the microchemical tests; but in the lack of 4 7 
precise knowledge of their chemical nature they may be ae 
sidered here under that name. All of them are found pe 
serial sections cut from paraffin, that is, in material which 4 
passed in bulk through the weak alcohols and rested in 7§ P* : 
cent. alcohol for weeks or months; which has passed i 
paraffin through chloroform and been cleared of out 
immersion for a few moments in xylol; and finally whic 
undergone the baths accessory to the stain employed. at 

The first oil is very abundant, existing in far greater deve 
than either of the others, and during more stages ae ee the 
opment of the fungus. It is found in the yous ee 
oosphere, and the periplasm. Diagrams showing “6 oa 
tion in the important periods of development are ae che age 
plates. fig. 46 presents it for an oogonium of ysis sonatio® 
shown in fig. 45. Fig. 63 illustrates its distribution @ we 
and it: should be noticed that the drops are ean = 


: ie) THE COMPOUND OOSPHERE OF ALBUGO BLITI . 229 


meshes of the ooplasm are fine, but large in the periplasm 
where the meshes are more coarse. . Fig. 72 shows the condition 
ofan oogonium at a stage similar to fig. 70, a period some- 
what later than zonation. The distribution is much the same 
because the protoplasm has changed but little. fig. &7, a stage 
ter fertilization, indicates that as the meshes of the ooplasm 
become coarser the globules become larger; apparently some of 
the globules fuse to form the larger drops. A later stage 
similar to fig. 93 shows that the oil of the periplasm is nearly 
sthausted during the building up of the epispore. 

This most common oil is found in unstained sections as 
— Olackish or brown drops when the material has been killed by 
Flemming’s fixing fluid. It is usually absent from material fixed 
by chrom-acetic acid, or if present is in very small quantity. 
tthe material is allowed to remain several (six) days in the 
chrom-acetic fixing agent the oil appears in considerable quan- 
ity, very much as it does in Flemming’s material, but of lighter 
= olor instead of brown. When material fails to show the 
ae seems to be really absent, because several days’ immersion 
a cent. osmic acid solution fails to show its presence. 
eretore, it seems that the osmic acid of the Flemming’s fixing 
og the long immersion in chrom-acetic acid, so acts on 
st oll as to render it less easily soluble in the fluids that it 
: Sag fhe killing and the time when the sectians are 
: tiful a 7 oil when present takes a characteristic ane peau: 
he — from the safranin in the Flemming’s triple 
_Sombination. If it be ver dark from the action of the 
SMic acid, it is fre ‘ d er- 
“Mid to all quently necessary to employ hydrogen p 
May be le blackness, in order that nas best results 
ee i. from the stains. It is partially on account : 

at material killed by Flemming’s fluid is not so 


Sora) 
 €ttic i cytological study as that fixed by the chrom- 


- The Second oijl-lj 
'Y, possibly ¢; 
y ei 

* evident in oe 


ke substance is present only in very small 
ht or ten small globules in an oogonium. 
h Flemming and chrom-acetic material, is 


niet 
Fr 


230 BOTANICAL GAZETTE [ocroner 


stained black by Heidenhain’s haematoxylin, but may be distin 
guished from the first oil by the fact that it does not take the saf- 
ranin when applied in the usual way. It is figured in the plates 
as black dots (figs. 51, 56, 59, 61, 64, 68, 69) ; in the antheridium 
in fig. 55; in the periplasm in fig. 80. No difference in stain 
reaction was observed between this oil-like substance and the — 
central globule of the coenocentrum, and they may be of similar 
nature, 

The third oil is only found in. the maturing oospore, first 
appearing while the secondary endospore wall is developing as 
globules of a clear honey-yellow color in the meshes of the proto- 
plasm ( fig. 95). It soon accumulates at the wall, and fig. 94 shows 
large drops in close contact with the forming secondary endo- 
spore, which has broken away from the primary endospore, prob- 
ably owing to the impact of the knife in cutting. The drops 
finally become larger and more numerous, as is shown in fig. 96. 
From a study of subsequent stages it seems probable that pi 
oil drops later break away from the wall irregularly, and unite nd 
form the several large drops that are often found in colder 
oospores. This results finally in the condition shown in fig. 9h 
where the entire central region is occupied by a curious irregular 
globular structure which stains much as the oils have stained in 
previous stages, but which is certainly not of fluid consistency: — 
It probably represents reserve food material. If this spre 
represents a genuine oil drop in the living spore, we may 
to do here with the shrunken and collapsed vesicle oF membrane 
that encased it in life, and whose contents have been “ 2 
the processes of technique. Its appearance would accor ee ” 
with this view. From its size and the time of devee a e 
the oospore it is surely the structure described as oil bx pT 3 
investigators, and is characteristic of the winter oospore: 


GENERAL CONSIDERATIONS. . 
eneral view of the : 
to previous ? 
= possible 


: It seems well before closing to take a g 
facts that have been presented, and of their relation 
knowledge; also to point out more clearly annie 


1899] THE COMPOUND OOSPHERE OF ALBUGO BLITI 237 


in earlier pages the broader significance of the phenomena 
described. 

A mitotic division of the nuclei of fungi seems to have been 
first noticed by Sadeback (’83) in Exoascus, and has since been 
observed in various forms and carefully studied and described in 

_ afew papers. 
The occurrence of a mitotic division in the oogonium of 
_ Albugo was discovered by Wager (96) in A. candidus, and con- 
firmed by Berlese (98) in A. Portulace. Wager notes that the 
muclei enlarge and divide, leaving one daughter nucleus imbedded 
inthe central body while the other nuclei pass to the periplasm. 
The one division increases the nuclei from about 115 to about 
double that number. Berlese says the nuclei divide several 
times in A. Portulace, increasing the number from 30 or 40 to 
about 200. The account given in the present paper describes two 
*pproximately simultaneous divisions affecting all of the nuclei 
® both antheridium and oogonium, and these mitoses result in 
ie formation of the sexual elements, numerous male and female 
nuclei, The second division is strikingly different from the first 
Be erence of the nuclear elements, particularly the 
tna of = This condition suggested the possibility of a sgeieee 
evidence a. but careful study revealed na: Conve se 
a ee - The mitoses are characterized by the intranuclear 
| ation of the spindle, the intranuclear centrosomes, the per- 
5 ~Maence of the nucleolus, and the entire absence of extra- 
 ‘Welear radiation. 
get described the disappearance of the nucleolus in early 
followed by the formation of chromosomes and then 
- the spin B  vclopment. He inclines toward the view that 
WW to bia ‘ €rived from the linin. The membrane persists 
ie ‘iad 2 and Wager did not follow the division seem 
a dissppea, Observations similar to those of Wager, pein 
indle a of the nucleolus in prophase, argues that ; : 
in 4. Blin tf der ived from it. The behavick of the nuc eo 
"entioneg ; 's different from that described by the writers 
: » in that the structure remains apparent inside of the 


a aia i ial 


232 BOTANICAL GAZETTE [octosex 


nucleus through all stages of mitosis. However, since Wager 
studied this question only incidentally, and as Berlese gives no 
figures and his account is very brief, a detailed comparison 
of the species is impossible. Berlese reports from twelve to 
sixteen chromosomes, and seems to have been able to count 
them during the fusion of the sexual nuclei. Neither Berlese 
nor Wager give details of the mitosis later than prophase. 

Wager reports that as near as he can estimate there are from 
twelve to sixteen chromosomes shown in the mitotic figures. In 
A. Bit six are found with certainty in some anaphase nuclei, 
and twelve appear in some metaphase nuclei with equal certainty. 
It may be that when twelve are counted the chromosomes have 
already divided, and that they really belong to two rather thas 
to one nucleus. However, this is not certain, and there is some 
evidence that makes it appear that there is a reduction in the 
number of chromosomes during the first mitosis, but this canmot 
be considered as proved. 

Centrosomes were not observed in either A. candidus ot 4. 
Portulacé, or in any other of the Phycomycetes, so far as the 
writer is aware; but they have been described in earlier a8 
for Ascomycetes (Gjurasin ’93, Harper ’95), and for Basidomy- 
cetes (Wager ’92, Juel ’98). a P 

With such fundamental differences as have been indicated, 
is useless to attempt to establish a type of mitosis for pois" : 
or to attempt to determine the relationship of the Aaa gene 
cytology. The type here described for A. Bit, while ae 
ing a few deviations from the mode of mitosis in vogue a 
higher plants, is in no way a departure from the forms ¥ 
known among the lower types of plants and animals. i 

It is impossible to generalize on the facts of mori : 
sented in this paper, because simple processes of mae : 
have been described in all forms where homology ee of 
sought. Until the behavior of the nuclei of on Lee 2 
Albugo is known, it is impossible to say which is the eee 
form of the genus, A. candidus or A. Bliti. From mee in most 
study of A. candidus it can be affirmed that A. Bua oe 


1899] THE COMPOUND OOSPHERE OF ALBUGO BLITI 233 


ways far more favorable for study ; its nuclei are larger, its peri- 
plasm more abundant, its developmental stages more ‘strongly 
marked, and its antheridial tube larger. Albugo candidus, how- 
ever, has a remarkable coenocentrum, which will be much easier 
ostudy than that of A. Bliti, owing to its much greater size and 
more pronounced stain reaction. A problem of great impor- 
tance lies in a comparative study of the ccenocentra of the 
genus, 


The characteristic massing of the cytoplasm to form a rudi- 
mentary oosphere in 4. Bhiti, differing thus from the vacuolate 
oosphere of A. candidus, is not a wider divergence than might be 
expected in different Species; nor is the variation in the cceno- 
‘eatrum more than what might be regarded as a specific differ- 
ence. If such variations are found to be more marked in other 
Species the Way may be clear to trace the relationship between 
plants with one oosphere and those with several oospheres in 

tach ogonium ; between forms which differentiate their peri- 
F plasm alter the manner of Vaucheria, and others that follow the 
tof the Saprolegniacez in a parietal rather than a central 
massing. It must be left to future research to make clear the 
ae that must exist between the multiple fertilization 

| ed by Albugo Bliti and simple acts of fusion between 
; “xual elements, 


Ttmay be that cytological investigation will show remarkable 
_ Vafiations in 


dei: many respects in this genus, and establish a chain 
wwe forms. The Saprolegniacez are said to range from 
omplete sexual fertilization. Should Albugo 
ly rich in habits the present knowledge of 
much increased. 


METHODs, MATERIALS, AND STAINING REACTIONS. 
© materj 
- al 
Sisteq a 


Pon which this investigation was based con- 


ems, and flower clusters of Amaranthus retro- 
, ba. oA. hybridus L. bearing the fungus. It was collected 
SES the my = Y,, Columbus, Ohio, and Chicago, III. In all 
Peles seemed to be unquestionably Adbugo Bliti Biv. 


234 BOTANICAL GAZETTE [octoses 


(C. Amaranti Schw. C. amarantacearum Zal\,). The form described 
by Zalewski (’83) as a different species was not met. Oospores 
are very abundant on both leaves and stems, producing on the 
former characteristic blister-like patches that assume a blackish 
hue if the oospores ripen in sufficient quantity. In the stems 
their presence may be predicted from peculiar swellings, usually 
accompanied by a reddish coloration, the entire plant often 
being thus affected. In partially diseased plants the oospores 
are likely to be found in the inflorescence, which reacts much as 
does the stem, becoming swollen and red. The parts most favor- 
able for study are the stems, but leaves and flowers often set 
tion more easily. In killing the material undesirable parts were 
cut away, and the portions apparently favorable were cut into 
small bits; leaves were scored, and stems and peduncles were 
cut in pieces about 2 to 4°™ long, deep incisions being made 
every 2™™ to give ready access to the killing agent. The killing 
with suitable solutions was apparently perfect, and was as good 4 
midway between incisions as where the solution immediately 
reached the tissue. : 
The killing agent giving the best results was chrom-aceti¢ 
acid of the following formula: chromic acid 0.8 per cent., acell 
acid 0.5 per cent. in water. The material was usually eft in this 
solution from twelve to eighteen hours, then washed in five of . 
six changes of water, allowing about two hours between change 
It was then successively transferred to 12, 25, 5% 
alcohol, remaining about two hours in each grade, ‘ 
in the last grade until it was practicable to imbed in pat . 
variation in the above method, by which the material page 
six days in chrom-acetic acid gave interesting results. 
dered theoil in the protoplasm much less soluble, 
the loss of many of the details of the mitoses. 
Flemming’s chrom-osmic-acetic acid was em 
same manner as the chrom-acetic acid, but was 
since the sections were very much darkened by t 
thus necessitating elaborate methods of bleac 
desirable stain could be obtained. 


ployed i” 
not so ust g 
he osmic a — 
hing before * 


19] THE COMPOUND OOSPHERE OF ALBUGO BLITI 235 

Other killing agents used were corrosive sublimate in satu- 
rated aqueous solution; hot corrosive-acetic-sublimate in alco- 
hol; Carnoy’s fluid; absolute alcohol; Hermann’s fluid; and 
Merkel’s fluid. Most of these gave far inferior results to that 
obtained by the chrom-acetic acid, and none surpassed it in 
effect 


Inorder to imbed in paraffin the material was transferred 
through 85, 95, and 100 per cent. alcohol to a mixture of absolute 
alcohol and chloroform, first of one third then of two thirds 
strength of chloroform. The specimens were left about two- 
hours ineach fluid, and were finally placed in pure chloroform. 
After the material had been left in chloroform for an hour a 
quantity of paraffin was added, and two hours later the material 
m= wamedon the bath. After two hours of gentle heat it was 
“moved to a warmer position, and later the most of the chloro- 
frm was poured off, melted paraffin substituted, and the whole 
j kept in the bath at a temperature of about 55°. It was found 

*antageous during the whole process of imbedding to use 
ey lion dishes, as by this means most or all of the chloro- 
aay off gradually by evaporation. Material was left 
twice ER 2 mont paraffin three or four hours, the paraffin being 
— Chlorof snged in the meantime to insure the removal of all 

~ a. It was then cast in a cake, the final paraffin hav- 
Bt melting Pvint of about 62°. Sections 3 to 5 win thickness 
Sig gil Jung sliding microtome and fastened to the slide 


t Mayer’s albumen fixative. 
aL Flemming: 


results. This stain demands the greatest atten- 


Peal use or failure is inevitable, as is well known by all 
8 a es In general, the best results were attained by a 
bs prolon a sa in safranin, followed by a rinsing more or 
ker of bien mM acid alcohol. The time here is entirely a mat- 
“20st usua a varying with the result desired ; 30-90 sec. was 
: the Slide After running down through the alcohols 


iia Placed in saturated solution of gentian-violet for 
5 min. Jt was then rinsed in water and placed in orange 


fon. 


236 BOTANICAL GAZETTE locroser 


G from 5-25 sec.; a longer time may do no harm but probably 
10-15 sec. is always sufficient. Wipe away excess of liquid and 
flood the slide twice with absolute alcohol, allowing the second 
lot of alcohol to remain on the slide until sufficient of the gentian- 
violet has been removed. The time required will depend on the 
material, the length of time it was in the gentian-violet, and the 
result desired. Drain rapidly with filter paper and flood with — 
clove oil for one minute; drain, follow by cedar oil and cover — 
in xylol balsam. If properly stained the host cell wall should 
be a light violet, the chromatin of the spirem and the chrome 
somes blue, nucleolus and centrosomes red, and cytoplasm 
slightly yellowish. 

Hematoxylin stain was used, applied after the method of 
Heidenhain (iron-alum 2 hrs., hematoxylin 12-18 hrs., followed 
by the slow extraction of iron alum till the proper degree 1s 
reached). This treatment gave some beautiful results in contrast 
with the Flemming stain, and was particularly valuable in demon 
strating the achromatic portion of the nuclear figure. Hartog’s 
nigrosin-carmine stain, as used by Wager and Berlese, ¥# 
tried repeatedly on corrosive sublimate material, but the results 
were far inferior to those afforded by Flemming’s tpl = 
Heidenhain’s hematoxylin stains. However, it is possible © ” 
demonstrate, even with this combination, the presence of a - 
nuclei in the oosphere and in the antheridial tube, and to rec 
soni ee ; : ‘c figures. thet 
ognize the principal features of the mitotic Mg! - Ehrlich, 
stains employed were Delafeld’s haematoxylin, Biondi’ — 

: = inferior in thet 
and cyanin-erythrosin, but they were distinctly 1n 
results. 

The following stain reactions were presente‘ - 
arations, and were attained by the Flem 
unless otherwise stated: chromatin blue or v!0 


Heidenhain’s hematoxylin ; nucleoli red, black wis pe datk 

hematoxylin; centrosomes as nucleoli; ie n page 16} 
: : . e Oo if ey 

blue ; cytoplasm yellowish ; granules, mention ee EE 


are only seen in preparations stained with Heide 
toxylin, and then black. 


1899] THE COMPOUND OOSPHERE OF ALBUGO BLITI 237 


SUMMARY. 


1. The oogonium when cut off from the parent hypha con- 
lains about 300 nuclei, which enlarge and divide mitotically 
while the oosphere is being differentiated. 

2, The oosphere is differentiated through a massing of the 
cytoplasm of the oogonium. By this process the nuclei, usually in 
stages of mitosis, together with the vacuoles, are expelled from 
the central region, and there results a dense and coarsely vacuo- 
te periplaam. This condition occurs when the antheridial 
tube is very short. 

3. There is a stage called zonation in which the nuclei, usually 
metaphase, are lined up around the ooplasm, some of the 
spindles lying across the definite boundary that separates ooplasm 
tom periplasm. In the telophase of this mitosis about fifty 
daughter nuclei are found in the ooplasm. 

4 The antheridium contains at first about thirty-five nuclei 
omy divide twice mitotically, and simultaneously with the 
“sion in the oogonium and oosphere. 

oo. i” the entrance of the antheridial tube a papilla 
Projecting from the oogonium into the antheridium. 
6. The antheridial tube penetrates slowly, reaching the 
| *oplasm at the time of zonation, later entering the oosphere and 
—Ppearing as a conspicuously multinucleate structure. When it 
ao there are discharged about one hundred male nuclei which 
With the female nuclei in pairs. 
: a oc nuclei differ in form; the sperm being elon- 
a 3 € egg spherical. 
ae central body, the ccenocentrum, bpabbee a as 
: ‘ 5 Matures and disappears before fertilization. Its 
iy unknown. There is some evidence of its being a 
— SYNamic Femter of the comp d h 
9. The ioc: 7 re oosp ae oe 
oth a. ‘ are alike in the oogonium and antheridiu é 
ba os intra-nuclear and there are no extra-nuclear 
Md are —.. centrosomes are very prominent at metaphase, 
Ss uclear. They could not be distinguished in the 


238 BOTANICAL GAZETTE [octoper 


resting nucleus. -The nuclear membrane persists until after 
metaphase and the nucleolus is present throughout the division, 
10. The primitive wall of the oospore first appears when the 
antheridial tube opens. . Later the epispore is laid down uponit 
by the periplasm. 
11. Two endospores are formed by the ooplasm after the 
development in the vacuoles of a peculiar substance which dis- 
appears as the endospores reach maturity. 
12. After the complete encasement of the oospore it becomes 
rapidly filled with food-stuffs. A large central oil-like drop is 
present during the winter condition, 
13. The fusion nuclei pass the winter in the resting comer 
without further perceptible change. 


THE UNIVERSITY OF CHICAGO. 


BIBLIOGRAPHY. 
BEHRENS, go. Einige Beobachtungen iiber die Entwickelung des Oogons 
und der Oosphire von Vaucheria. Ber. d. deut. bot. Gesells. 8: 1 
BERLESE, 98. Ueber die Befruchtung und Entwickelung der Oosphiire bei : 
den Peronosporen. Jahrb. f. wiss. Bot. 31: 159. ’ 
DANGEARD, go. Recherches histologiques sur les champignons. Le Ba 4 
1890: 125. ‘ 
oe 93. Ueber die Kernteilung in den Schlauchen von Pesisa ves : 
a Bull. Ber. d. deut. bot. Gesells. 11: 113. é : 
ee 95. On the cytology of the vegetative and reproductive “— - 
the Saprolegniacee. Trans. Roy. Irish Acad. 30: 645: « 
HUMPHREY, 92. The Saprolegniacee of the United States with notes 
other species. Trans. Am. Phil. Soc. 17: 63-149. 
HARPER, 95. Beitrag zur Kenntniss der Kernteilung und Spo 
Ascus. Ber. d. deut. bot. Gesells. 13: 67. 
ISTVANFFI, 95. Ueber die Rolle der Zellkerns bei der Entwickelung OH 
Pilze. Ber. d. deut. bot. Gesells. 13: 456. 
JUEL, 98. Die Kernteilung in den Basidien und die 
myceten. Jahr. f. wiss. bot. 32: 361. 
LEGER, 95. Structure et developement de la zygospore du 
dis, Rev. gén. Bot..7: 481. 


renbildung a 


Phylogenie der Basidir - 


Sporidinia sr 


1899] THE COMPOUND OOSPHERE OF ALBUGO BLITI 239 


OurMaNNS, 95. Ueber die Entwickelung der sexualorgane bei Vaucheria. 

Flora 80 ; 

SADEBECK, 83. Untersuchungen iiber die Pilzgattung Exoascus. Jahrb. 

der wiss. Anstalten zu Hamburg —: Io1. 

Trow, 95. The karyology of Saprolegniaceze. Ann. Bot. 9: 609. 

—— 95. Trans. Roy. Irish Acad. 30: 640. 

WaGer, 90. Observations on the structure of the nuclei in Peronospora 

parasitica, and on their behavior during the formation of the oospore, 

Ann, Bot. 4: 127. 

—— 9. On the nuclei in Hymenomycetes. Ann. Bot. 6: 146. 

_—— 96. On the structure and reproduction of Cystopus candidus Lév. 
Ann. Bot 10; 295. : 

IALEWSKI, 83. Zur Kenntniss der Gattung Cystopus. Bot. Cent. 15: 215. 

TMMERMANN, 96. Physiologie des pflanzlichen Zellkernes 129, 146. 


EXPLANATION OF PLATES XI-XV. 

All figures are from material killed in chrom-acetic acid and stained with 
Fleming's triple stain, unless otherwise indicated. The figures were sketched 
Abbé camera, using the following combinations of lenses: Zeiss 2" 
3 . 1.30, with compensating ocular 18 and 12; also Bausch & Lomb 4, 
: cleaned no. 4. These combinations give respectively the following 
_ '"“ignifications when projected to the table level: 3300, 2200, and 1500 
there “me Plate XI i not reduced in reproduction. All other plates are 
: ourths of the original scale. 


PLATE XI. 
Al drawings repres 


 alied «: ent a magnification of 3300 diameters, and were 
: with Zeiss lenses, 


ee. 1 : : 
oF - Resting nucleus in mycelium, linin faint, nucleolus prominent. 
6.2. Nucleus in flowin 


aes g cytoplasm near entrance of oogonium. 
the Nucleus 'n antheridium, spirem stage, membrane faint. 
: hatin "eed small drops accumulated on the linin network, the 
Fig. Som net, nucleus becoming spindle-shaped, no centrosome — 
Pete, chromant somewhat later, linin threads have almost entirely disap- 
TeMains of ge Scattered through nucleus, nucleolus to the right, 
Fis. 6. 4 ‘nin strand to the left 
a Sram more spindle-shaped, globules arranged in the equato- 
RE, no Spindle bea round bodies about equal to the chromatin dots in 
*etbrane. €rs, the definite line bounding the whole of the nuclear 


zz 


240 BOTANICAL GAZETTE [octToser 


Fig. 7. Similar to 7g. 5. The longitudinal lines probably chromatin which 
has not reached the equator, nucleolus at the left. This nucleus was crowded 
by a mass of others, hence its short form. 

Fie 8. Similar to fg. 7. Fibers visible toward the poles. This spindle 
was formed in one of the strands supporting the forming oogonium, and was 
consequently much elongated by tension. It is from the oogonium shown 
in fig. 62. 

FIG. 9. Chromosomes at the equator, spindle fibers very apparent at the 
poles, but not visible at the equator, nucleolus to left, membrane intact and 
inclosing the spindle. 

Fic. 10. From oogonium shown in fig. 60. Cross section of a spit 
dle, twelve chromosomes apparent (stained by haematoxylin from Flemming’s 
material), 

Fic. 11. Spindle mature, chromosomes closely grouped at the equator, 
centrosomes prominent, spindle brilliant and clear, nuclear membrane at 
ent but poorly stained (fig 73). 

Fic. 12. Chromosomes splitting, membrane _ visible with nucleolus 
enclosed, 

Fic. 13. Nucleolus large, nuclear membrane very definite, daughter 
chromosomes ready to separate. Only those in the highest focus aii gee 
several others being found at a deeper focus. The stain was particularly to 
show the membrane, and was not suitable for centrosomes. 

Fig. 14. Anaphase: chromosomes separating, nucleus lying about mid: 
way, centrosomes still visible, whole nucleus staining dark and membrane 
indistinguishable from spindle fibers. moe 

Fig. 15. Chromosomes nearing the poles, centrosomes not citing 
from them, nucleolus thidway, slight traces of spindle fibers stretching a 
the middle space, cytoplasm in the ends of the nucleus stains darker 
that of the central area. 

Fig. 16. Whole spindle elongated, chromosomes massing t08 


poles. ie 
Compare witt 


Fig. 17. Similar to fig, 76, but in a crowded position. 
7 


cheat 
- 


; the age 
Figs. 18 and 1g are from the periplasm of an oogonium of abou ; 


shown in fig. 67, slightly younger than shown in fig. 68. 
Fic. 18. Spindle fibers collapsing in the middle leadi 
ration of the daughter nuclei, the fibers constituting the or) 
brane of the daughter nucleus. ai 
§ ee chromo 


ng to the sepa" . 
gin of the men oa 


Fig. 19. A young daughter nucleus, nucleolus, membr 
somes. 


ime] THE COMPOUND OOSPHER« OF ALBUGO BLITI 241 


Fig. 20. Same as fig. 79 in resting stage, nucleolus prominent, linin 

faint. 

Figs. 21 and 22 are from the same oosphere, and in the condition shown 

in figs. 68-70. 

Fig. 21. Same as fig. 20 but passing into the spirem stage. 

Fic. 22, Nucleus elongating preparatory to division. Compare fig. g. 

Fic, 23. Breaking of skein into chromosomes, centrosomes apparent. 

Fig. 24. Spindle forming inside of nuclear membrane, nucleolus lying 

outside of the spindle. 

Fig. 25. From an oospore of the condition shown in fg. 70. Spindle 

'ying completely inside of the nuclear membrane, chromosomes grouping at 

the equator, and centrosomes well defined. 

Fi. 26. Metaphase: chromosomes splitting, membrane, centrosomes, and 

fibers clear, Fibers apparently of about the same number as the chromosomes. 

Fig. 27. Daughter chromosomes ready to leave the equator. 

Fig, 28. Anaphase: chromosomes well separated, centrosomes visible, 

Spindle fibers crossing the middle space, chromosomes six in number. From 

Sime oospore as figs 24 and 27. Compare with fg. 76. 

Fig. 29. Later anaphase: chromosomes near the poles, area in which 

they rest darker stained than central portion. 

Fig. 30. Similar tofig.z8. Spindle fibers collapsed and daughter nuclei 

tady to separate. From same oospore as fig. 29. 

oo male nucleus from the entrance of antheridial tube. This 

€ same as the one marked X in fig. 73. 

Fi, 32. A nucleus (sperm) in the tip of the same tube that contained that 
m fig. 37, also marked X, Wall of the antheridial tube, dense 

Foplasm su 


son, Tounding nuclei, sperm with nucleolus and mass of chromatin 
_ “anterior end, 


Fig, : 
rey se Tube open, elongated and pointed sperms escaping, showing a 
MBit network, one female nucleus shown. This is part of the 
pes F gs own in fig. 8&5. 
Ig, ; 
than ; 34. A sperm approaching a female nucleus, linin more prominent 
Nn fig. 3 2, 
Fig, : 
Mound, cay Sperm in contact with a female nucleus, becoming more nearly 
; > ts linin still more prominent. 
: GS, ‘ 
Hk t0 40. Various stages of fusion, from oospores of the general 
“Rote Na represented in fig. go. During fusion the whole nucleus becomes 
cae Y Stained with gentian-violet. 


er aee 
- Otdinary ®nocentrum with globule at its center, female nucleus near by, 


vacu 
oles of the Oosphere near the margin. 


242 BOTANICAL GAZETIE [octonss 


PLATE XII, 

Magnification in all figures 1500 diameters. 

Fie. 42. A portion of mycelium, showing nuclei with prominent nucleoli, 

Fic. 43. Nuclei and cytoplasm flowing into a developing oogonium, nucle 
and vacuoles elongated and angular, nuclei too darkly stained to show 
structure. 

Fig. 44. The septum below the oogonium, a portion of mycelium and 
posterior end of odgonium, oogonial nuclei assuming the spirem stage 
mycelial nuclei still in resting condition with prominent nucleoli bet 
enlarged. 

Fic. 45. Oogonium and antheridium, nuclei of both in the spirem stage. 

Fig. 46. Shows location of oil drops in an oogonium of the condition pre 
sented in fig. g5. They were all drawn at one focus; a slight change of 
focus would have brought vastly more into view. 

Fic. 47. Showing adhesion of the oogonial Hautschicht to its wall in the 
neighborhood of the antheridium, nuclei overstained. 

Fic. 48. Adhesion as in fig. ¢7. Oogonial wall partly and irregularly 
corroded away on the side toward the oogonium. 

Figs. 49-55. Stages in the perforation of the wall preparatory t the 
entrance of the antheridial tube. : 

Fig. 49. Optical section of papilla, wall partly corroded away and bulging 
toward the antheridium. The dense protoplasm represented in black was 
stained a deep red by the safranin. 

FiG. 50. Sectional view of a condition slightly older than jig: #9: ox 
shaded portion of the separating wall took the stain differently from the 
and was apparently in the last stages of dissolution. 

Fic. 51. A papilla of different shape in a stage similar to the last. 

Fic. 52. Optical section, stain as in fig. 49. coe : 

F1G. 53. A papilla becoming bubble-like, walls very thin, slight'Y°™" 
as though due to imperfect killing, watery vacuoles apparent, their © 
staining homogeneously with the sia ee airections, 

Fig. 54. Bubble-like papilla expanding irregularly in all di 
tents highly vacuolate, wall extremely thin. igi ng in eithet 

Fig. 55. Wall almost perforated, but showing no marked bulg! 
direction, 0, oogonial, a, antheridial side. punded 

Figs. 56, 57. Young antheridial tubes with a cellulose wall and s 
by a dense sheath of the protoplasm of the ng ae voachiol 

Fic. 58. Protoplasm beginning to collect in masses, ee fixed 
metaphase, some shown’ in polar view (preparation from mat ae 
Flemming’s agent and stained by haematoxylin). 


1899] THE COMPOUND OOSPHERE OF ALBUGO BLITI 243 


Fig. $9. An intercalary oogonium, slightly older condition than fg. 58, 
cytoplasm distinctly in masses, nuclei in early prophase, antheridial tube just 
entering bearing no nuclei, a haustorium at extreme right. 


PLATE XII. 

Magnification in figs. 60, 67, 62, 63, 64, 65, 68, 69, 70, 72, 1500 diameters ; 
nfigs. 66, 67,77, 3300 diameters; in fig. 77, 2200 diameters. 
FiG, 60. Nuclei in metaphase, protoplasm massed in a few centers, 
spindles very clear and brilliant (stain hematoxylin from material fixed in 
Flemming’s agent), 
Fig. 61. Nuclei in late prophase, the protoplasmic masses coalesced to 
form one, vacuoles mark the juncture last made, a few nuclei not yet floated out. 
Fig. 62. Nuclei approaching metaphase, spindles much elongated, all not 
yet extruded from the central region, mitosis in the antheridium, antheridial 
tube shown at its typical position for this stage. 

Fic. 63. Diagram of disposition of oil drops in an oogonium of the con- 
ftion shown in fig. 65 (zonation). 
. Fic. 64. Nuclei nearly at metaphase, zonation almost complete, oil-like 
drop to the tight in ooplasm, antheridial tube present showing no nuclei. 
Fic. 65, Zonation : nuclei near metaphase, ooplasm sharply differentiated, 


plasm (stained by hematoxylin from Flemming material). See also fig. 64. 
‘ 66. Shows spindle in metaphase lying across the film between 
and periplasm, 
*i6. 67. Nucleus in anaphase directly across the boundary film of the 
: » also half of a late anaphase cut diagonally. 
; — » 69. Consecutive sections of an oogonium just after the division 
by * *) showing the position and shape of the antheridial tube. 7g. 
we, the cenocentrum which had just passed its maximum development 
fig. 72). 
a> Tey. = a: ; 
a eal tube and differentiating oosphere, oosphere nuclet 
- itd shows *§.74 was taken from an adjacent section of the same oogonium 
. the ccenocentrum, 
IG, 
— Panded aC letely developed ccenocentrum, the central globule sur- 
Wi thes ree regions of differentiated protoplasm. Froma stage of zona- 
Fe about “nuclei were in anaphase (as in fig. 67) and the daughter nuclei 
to pass into the ooplasm. 


Fig, —w 
Be 72. Diagram of distribution of oil at the stage shown in fig. 79- 
malt 4 Antheridium and tube of the age shown in fig. 79 showing 


: é ; 
* Contents; Z, film; 9, one of the female nuclei; f, periplasm; 


’ Fepresented also in Plate XT, figs. 31, 32. 


244 BOTANICAL GAZETTE * [octos 


PLATE XIV, 

Magnification in figs. 74, 80, 82,87, 88, 1500 diameters; fig. 76, 2200 
diameters ; figs. 75, 77, 78, 4, 85, 86, 3300 diameters. 

Fig. 74. Coenocentrum and differentiated oosphere with dividing nuclei. 
The broken empty portion is where the protoplasm shrunk away from the 
autheridial tube. Section adjacent to that shown in fig. 70. 

Fig. 75. Ccenocentrum, in an oogonium, at anaphase period of zonation, 
consisting of three small globules evidently fusing, surrounded by a region of 
denser i siaaea granules resembling small oil drops scattered through the 
ooplasm 

1G. 76. Similar to fg. 77 but somewhat older, antheridial tube showing 
many nuclei, slightly torn near tip, antheridium becoming vacuolate ; p, peri- 
plasm ; 4, film; 9, female nucleus. 

Fic. 76 a. Portion of the tip of the same tube, found in an adjacent 
section. 

G. 77. View of the end of an unopened antheridial tube, wall not 
se sperms very numerous and crowded, each showing a dark nucleolus, 
ooplasm slightly shrunken away from the tube. 

Fic. 78. Section near the base of the same antheridial tube, showing 
very thick wall in contrast with the extremely thin film covering the gies 
Fig.77 


Fig. 79. Diagram to show position of preceding sections. 
low the or 


from a section just above the line aa, fig. 78 from one just be 
One intermediate section was not drawn. 

Figs. 80, 80a. Adjacent and approximately longitudi 
opening antheridial tube of the same oogonium, showing t 
and the increased vacuolation of the ooplasm which frequently 
from the primitive wall. 

Fig. 81. Diagram of the distribution of oi] in an oogonium of 
shown in fig. 8o. 

Fig, 82. Antheridial tube discharging sperms, 
primitive wall very young, ooplasm becoming vacuolate, 
slightly oblique so that its base is in an adjacent section. 

Fic. 83. Diagram to show relation of sections pres¢ 
86: fig. 84 was cut from above the line aa, and tangentia 
opening tube ; fig. 85 from between the lines; /7g- 86 from jus 
0, 


nal sections of the 
he primit tive wall 
shrinks away 


the age 


antheridium separset : 
antheridial tube - 


nted in figs: 5¢ e 
al tothe OP 
t below he it 2 


] tube and 
Fic. 84 (see fig. 837). Sperm nuclei leaving the antheridia 


approaching the female nuclei. The tube mF 
Fig. 85 (see fig. 87). Mass of sperms escaping from t ee ee 
be traced to the left as a mass of darkly stained structure 


i) THE COMPOUND OOSPHERE OF ALBUGO BLITI 245 


The sperms ina mass appear dark but individually are hyaline except at 
anterior end which bears the nucleolus. See also fig. 37, Plate XJ. 

Fic. 86 (see fig 83). The base of the antheridial tubé filled with dark 
staining cytoplasm and few nuclei. 


Fic. 87. Just after discharge from antheridial tube, periplasm as in fg. 

#2, several small masses of nuclei apparently both male and female sur- 

founded by denser cytoplasm (these masses seem to arise by the breaking 

_ Martofthe contents of the antheridial tube and soon disappear), primitive 

_ wall well developed, antheridium above, ooplasm coarsely and irregularly 
vacuolate, 


Fig. 88. Nuclei fusing in pairs, primitive wall distinctly thickened. See 
_ Plate XI, figs. 36-40. 
PLATE XV. 

All figures magnified 1500 diameters. 

Fig. 89, Protoplasm collecting in dense network toward the center of 
the oospore, leaving light peripheral strands where the endospore is soon to 

I. 

& Fig. 90, Shows the remains of the antheridial tube in the periplasm, with 
_ *Mtrace of its former presence in the ooplasm. The oospore is of the age 


town in fig. 88 


Fi, gt, Primitive wall mature, exospore forming, now consisting of a 
Porous semi-transparent mass with imbedded disks which are to form the 

"ges, vacuoles in ©ospore filled with gelatinous substance, nuclei of proto- 
plasm overstained, 
Fig. 92, Somewhat 
: Suspore more clearly, 
‘ad giving a Surface yi 
Mie. 93. £ 
- Sloontaining 


younger than the last figure, showing structure of 
presents edge view and also a fragment bent back 
€w, gelatinous substance in vacuoles. 

“ospore nearly formed, primary endospore complete, vacuoles 
F 8elatinous substance. 

newle, > Padimentary secondary endospore, gelatinous substance in 


FIG. 95. Oi forming in oospore. 


1G, 
With ng io Spore Walls complete, including double endospore, vacuoles 
ls, Aisin aaee substance, oil accumulating in large drops on the endospore 
Fig, 


97. Winter Conditions of exospore, the ridges higher than in previous 


Rages, | ; j 7 
We central oil-like mass, nuclei in sporoplasm. Section not directly 
™ le so that the endospore appears thicker than it really is. 
*9. Exterior of ripe oospore. 


a 


NOTES ON THE DEVELOPMENT OF THE HOLDFASTS 
OF CERTAIN FLORIDE#:, 
Git wie M. DERICK, 
(WITH PLATES -XXI—XXIII AND FIVE TEXT-FIGURES) 


Wir the exception of passing references in various works, 
two articles, the one by Borge (1), the other by Strémfelt (8), 
include, I believe, all that has been written upon the holdfasts of 
the alg. The former deals with a few members of the Chlore- 
phycee ; the latter is very comprehensive, but it is without illus- 
trations and gives no specific details. Therefore, the study of 
the development of the holdfasts of some nearly related species 
of the Rhodophycez has seemed advisable. 

The observations described in this paper were made at the 
Marine Biological Laboratory, Woods Hole, Mass., during the 
summers of 1896 and 1897, and the work was finished in the 
Botanical Laboratory of McGill College. | 

Cultures of the spores of several species were made ae | 
various conditions. Ordinary glass object-slides were git 3 : 
flat porcelain dishes, either white or painted black. The vesse j 
were filled with filtered sea-water and in them were laid 2 7 
bearing ripe spores. The spores usually sowed i 
twenty-four hours, in which case the plants were removed. . 
of the dishes were fed by a gentle stream of running es 
others were disturbed only three times a day, when ppd 
was drawn off by a siphon and replaced by filtered se . 
The latter method proved much the better of the two. ier 

The color of the background had no effect upon the Cultures 
ment of the spores, nor were the plantlets heliotropic- se 
kept in a shaded place flourished best, even 4 short ao 0 
direct sunlight killing plantlets. It is to be pases je 
record of variations in the temperature and the dens! fs 
water was kept. Oltmanns (6) has conclusively show? 


of the 
t such 
[ocros 
246 


ees alee 


BOTANICAL GAZETTE, XXVIII ae : PLATE XXI 


PLATE XXII 


NY 
XN 
x 
rs 
>< 
= 
'& 
N 
S 
= 
< 
: 
N 
S 
& 


PLATE XXT 


NY 
N 
N 
> 
md 
N 
N 
N 
S 
: 
S 
= 
SS 
S) 
% 


1899] DEVELOPMENT OF HOLDFASTS OF FLORIDEZ 247 


variations are most important factors in the life and distribution 
of marine alge, but a. comparison of many cultures of the same 
species indicates that differences in the conditions which pre- 
vailed in the laboratory affected neither the form nor the order 
of development of the plantlet. It was difficult to keep the 
young plants healthy for more than three or four weeks; and 
during a few days of intense heat, in August 1896, all ceased to 
grow, were attacked by bacteria, and finally died. 

Both carpospores and tetraspores were germinated and most 
closely resembled one another in their development. In some 
tases, however, tetraspores attached themselves to the substratum 
less readily than carpospores. This is regarded by Brannon 
(2) as an adaptation to the immediate distribution of the species. 

Every day, slides upon which spores were growing were 
examined and drawings of living plantlets were made. Perma- 
wnt mounts for comparative study were prepared at regular 
imervals and mature holdfasts were preserved in alcohol or in 
} per cent. formalin in sea water. 

The species selected for investigation belonged to the Rho- 
dymeniales, with the exception of one member of the Rhodo- 


: Ey tase Rhabdonia tenera J. Ag. The carpospores of this 
| Pat attach themselves firmly to the slides in nine hours; a very 
— Mlicate outer layer, probably of mucilage, may be observed, but 


oa 


eh 


‘at hardly be distinguished from the cell-wall (Pl. XXV, fig. r). 


ees an irregular layer of a coarsely granular substance 
beret the plantlets, and in one, a thick mucilaginous disk 
“yohagd at the base ; these appearances were most aon a 
(Pr. is were doubtless due to slightly abnormal conditions 
ho 7, 8). After attaching themselves, the spores 

: Wenty-t oy *Bter upon a segmentation stage, and within 
the : a hours two divisions are made. The first separates 

PI Xu by means of a vertical wall into two equal cells 

* Figs. 2,3). Three and four-celled stages result from 
tight division of the two primary cells in vertical planes 
by obli ngles to the first wall; other cells are cut off from these 
We walls, and thus an irregular spherical mass 1s formed. 


248 BOTANICAL GAZETTE [ocToses 


In about thirteen days differentiation begins. Four basal cells 
elongate to form the primary root-cells, the position of which 
bears no definite relation to the direction of the supply of 
light (71. XXT, figs. 7, 8). Occasionally only one or two pri- 
mary rhizoids are thus formed (Pl. XXI, figs. 9, 10, 1). 
Unfortunately, the plantlets died at this stage, and only by 
analogy could the subsequent history of the holdfasts be 
determined. However, the close resemblance between the pri- 
mary root-cells and, mature holdfasts of Rhabdonia and of the 
two species to be considered next justifies the assumption that 
the intermediate stages closely agree. It is interesting to note 
that, according to Osterhout (7), the earliest divisions in the 
tetraspores which give rise to proliferations do not succeed one 
another in the definite order just described, but the spores divide 
“by means of oblique walls which do not occur in regular suc 
cession.” 

Very similar to that of Rhabdonia is the early history of 
Lomentaria uncinata Menegh. and of Champia parvula Harty. The 
carpospores of Lomentaria, after secreting a wall, attach them 
selves to the substratum by a very thin mucilaginous secretion 
(Pl. XXT, fig. 12), which sometimes persists after the first 
thizoids have been formed (Pl. XXV, fig. 15). The — 
tion proceeds rapidly, but primary root-cells do not arise pes 
a later date than in Rhabdonia (Pl. XAT, figs. 13: 14). 
four basal root-cells, dividing by horizontal walls, ; 
short, closely-appressed filaments, which soon branch pseu af 
dichotomously (Pl. XX7, figs. 15, 76) and forma ae 
Strémfelt (8) regards the branching of the rhizoidal per é 
dichotomous and says that the increase in circumference : c 
mature holdfast is due almost entirely to marginal growth. pie? 
tative reproduction may take place not only by po 
separation of the various upright branches, which sprin 
tiously from the surface of the holdfast (P/. XX, fig: 2? 
oy means of stolons, which are sent out in 
and form secondary holdfasts (PZ. XXV, fig: 21) 548 parc 

Davis’s account (4) of the plantlets of Champa 


become — 


discoid holdfast. 


various directions 


1899] DEVELOPMENT OF HOLDFASTS OF FLORIDEZ 249 


_ developing under natural conditions, accurately describes those 
grown in the laboratory. A somewhat spherical mass composed 
_ ofsixteen cells results from the segmentation of the spore, then 
the first indication of a permanent holdfast appears as a slight 
projection of each of the four basal cells. The primary root- 
cells, dividing at right angles to their longest axis, form two- 
celled filaments, which branch monopodially and give rise to a 
j broad spreading. holdfast. In section the mature holdfast 
_ appears parenchymatous (P/. XX7, figs. 77, 18), but it is often 
_ possible to distinguish the component filaments (P2. XXY, fig. 20). 
' Exceptional plantlets produce one instead ot four primary root- 
j cells, but the later stages conform to the type (Pl. XX, fig. 79). 
__ Inall cases the cells of the holdfast are paler than those of the 
; frond, and the chromatophores of young specimens are in close 
_ eontact with the walls. . 
: From the foregoing it will be seen that Rhabdonia, Lomen- 
‘ “ra, and Champia agree (az) in passing through a segmenta- 
ee stage, resulting in a somewhat spherical mass of cells, (4) 
im the elongation of four basal cells, and (c) in the subsequent 
_ tevelopment of four primary rhizoids, which branch repeatedly 
and finally form a large discoid holdfast, composed of pseudo- 
‘ parenchymatous tissue. 
: "marked contrast are several members of the Rhodome 
' - The spores of Chondria tenuissima (Good. et Wood.) C. 
FE ae ose Chondria dasyphila Ag. germinate very readily. The 
« : em of the spores and the history of the development 
i las '20ids are alike in the two species. The spores, which 
eo and have coarsely granular contents, quickly and 
Sie attach themselves to the substratum, doubtless by means 
‘i hin and uniform layer of mucilage, though no secretion 
fin the cell-wall is perce tible (PLXXT, fig. 23). They 
a Son divide into t : lightly con- 

ee wo unequal cells separated by a slightly 

2d the parallel to the first follow (Pl. XXV/, FE. ue 
Primary oo cell of the resulting filament elongates into the 

oid (PL. XX7, figs. 25, 27). Sometimes the basal cell 

® branch dichotomously ; but, as the branches do not 


250 BOTANICAL GAZETTE [ocrones 


arise simultaneously nor are they separated by a wall parallel to 
the longer axis of the basal cell, it is evident that the branching 
of the filaments is strictly monopodial (P/. XXT, figs. 26, 32, 35). 
The upper cells of the plantlet soon cut off pericentral cells, 
and at the same period the first rhizoid divides repeatedly so as 
to form a multicellular, monosiphonous filament (Pl. XXI, fig. 
28, as): Secondary rhizoids are next developed, either as out- 
growths from the primary root-cell or from a pericentral cell imme- 
diately above the basal cell (P/. XXTJ, fig. 31). Long slender 
rhizoids may be formed before a clasping-disk arises, but, both ia 
plantlets developing in the laboratory (2. XXT, figs. 29, 33,36) 
and in those growing in a state of nature (Pl. XXY, figs. 30,34 
_ 39, 40), the primary rhizoid generally remains very short and by 
means of oblique walls produces discoid holdfasts at the tip 
The size and efficiency of this holdfast are increased by meals 
of free secondary rhizoids (P/. XXT, fig. 33) or by intracuticula 
filaments (P?. XXT, fig. 34), both of which have their ongin ® 
pericentral cells near the base of the plantlet. The secondary 
rhizoids develop in the same manner as the primary and vote 
bine with them to form a large, irregular, discoid holdfast, ® 
which it is possible to distinguish few of the component a 
ments (P/. XX/, fig. 37). The mature holdfast, though adhering 
closely to its host, does not penetrate it, but a cutin 
the cortical layers often results from the contact. As in Lomen 
taria, etc., the large holdfast gives rise to several 
branches; and great powers of vegetative f 
implied by the stores of floridean starch with 
are often charged. The chromatophores are closely a 


er radial 


in the peripheral protoplasm, especially next to the ine eres 
and the lateral walls, and often indicate the plane ee wee 
scold, 


wall will soon be formed. They are large and 5 
resembling those of the cortical cells of the frond f 
the anastomosing filamentous chromatophores of the a ia 
Very different from the short, multicellular, Phe Pol 
oids of the Chondriez are the primary holdfasts of oup is 
siphoniee, The form taken as a type of the satel 


ather that 


tral cells 
ir : 


a 


ization of 


adventitious — 
eproduction use 
which the cells 
geregates 


‘i 
y 
= 
i 


: 
? 


7] 


1899] DEVELOPMENT OF HOLDFASTS OF FLORIDEA 251 


Polysiphonia violacea (Roth) Grev. In a few hours after they 
ae sown, both carpospores and tetraspores attach themselves 
to the substratum by a coarsely granular, mucilaginous secretion, 
which completely covers the spore (P/. XAT, fig. gr). In optical 
section this envelope appears densely granular at the margin, 
while a clear amorphous area imtervenes between the outer layer 
and the developing plantlet (Pi. XXT, fig. 44). The spore divides 
isto two unequal cells, of which the smaller soon becomes 
slightly pointed and finally grows into the primary root-cell (2. 
: XX, fig. 42). Divisions parallel to the first ensue (Pl. XX, 
igs. 43-45), and the basal-cell elongates so as to form the first 
—— thizoid piercing the mucilaginous sheath, which finally disinte- 
_ Sfatesand disappears. A six-celled stage is often reached before 
divisions in planes at an angle to the first occur. Generally, 
segmentation continues for some time and the siphons are clearly 
marked off from the:central axis before a second rhizoid arises. 
| As in Chondria, this has its origin in an unsegmented cell adja- 
«Cnt to the primary root-cell. The former differs from the 
later only in its brighter color and denser cell-contents; and, 
_ “maining undivided, it forms a component part of the holdfast 
| wae 50). The protoplasmic connections between these 
oe. ate obvious, but they are difficult to trace between the 
= cells of the plantlet (Pl. XXI, figs. 45,47). When three or 
B weeks old the young plant develops several rhizoids spring- 
." the primary or secondary root-cell (Pi. XXT, 
elds Ccasionally, however, the segmentation stage of the 
: a. and the growth of the frond begins before multipli- 
q . rhizoids takes place (Pl. XXII, jig. 10). At the 
’ as .. trond other rhizoids are sent out by corticating cells 

Parated from these by a wall, a protoplasmic connection 
. | = aaa No intracuticular filaments are developed 
’ Noldfast &pendence of the rhizoidal constituents of the ipaietie 
oth Practically preserved (Pi. XXIL, fig. 6). In addition 


¢ thizo; : 
. Zoids near the base of the plant, any corticating cell ofa 
mbent branch 


Bigg; may produce a secondary holdfast and thus 
a m the extensi 


on of the colony. As Strémfelt (8) noted, 


252 BOTANICAL GAZETTE [octoper 


all the rhizoids are unicellular and unbranched; and, although 
an apparent tendency to branch may be observed occasionally, 
the lobes are not separated from the main portion by a wall, even 
the protoplasmic contents being undivided (Pl. XXV/, fig. 9), 
All the rhizoids eventually develop terminal clasping-disks. 
The first indication of such a structure may appear in plantlets 
four days old, but, as a rule, the primary rhizoids do not undergo 
modification until several days later. The disks begin asa sim- 
ple enlargement of the tip of the rhizoid (Pl. XX, figs. 48-50; 
AX, fig. 1), become deeply lobed, and assume a very irregular 
outline (Pl. XXT, figs. 52-56). The cell-contents of the rhizoid 
extend into the lobes, but no division takes place (Pl. XAIL 
fig. 8). Great variations in the length attained by the primary 
rhizoids occur both in plants grown under natural conditions and 
in laboratory cultures (P/. XX/, figs. 46, 48; XXII, fig. 1). The 
cause of such variations has not been determined, but it is prob- 
able that contact irritation may be the most important factor 10 
the formation of the disks. This view is supported by the fact 
that disks are sometimes produced on the sides of rhizoids when 
these come in contact with a firm substance (77. XXII, figs. 4s 


5). The length of the secondary rhizoids depends upo? the. 


distance of the parent cells from the substratum, and as rad 
contact is established broad clasping-disks are formed, whi 
mechanically cohering with one another and with the eee 
disk produce a very strong holdfast. The rhizoids are paler 
color than the rest of the plantlet, having less dense conten 
and fewer chromatophores. 

The rhizoids of Polysiphonia violacea never pe 


e latter 
host-plant; but at the point of contact the susface of ee de 
is often dark brown and cutinized, while the outer comic 

| tism occuss 


are destit _ But incipient parasi 
itute of chromatophores pices om nodosim™ 


in Polysiphonia fastigiata Grev., growing on Ascop here species: 


Stack. Gibson (5), in writing of the histology of this 


noticed that “the attachment of the epiphyte to 0 
very intimate. Root-filaments given off from the : 
frond penetrate deeply into the tissue of the host, a 


netrate the — 


1899] DEVELOPMENT OF HOLDFASTS OF FLORIDEA 253 


among the cortical cells and medullary hyphe. The root-fila- 
ments have very thick cell-walls and central cells only, these 
being much elongated.” The ends of the rhizoids are swollen 
and in close contact with the cells of the host (seat figs. 4, 5, p. 
256), but no haustoria penetrate the walls of the latter. In one 
imstance a unique variation occurred.’ A few intracuticular fila- 
ments, descending from the corticating cells of the Polysiphonia, 
fan parallel to the main axis of the rhizoid throughout its length. 
The host suffers no serious injury, only a depression and cutini- 
ation of the surface with a very slight disorganization of the 
cortical cells at the point of penetration. Though the associa- 
tion of the two plants does not justify the assumption of com- 
plete parasitism, the symbiotic relation existing between them is 
Much more intimate than that observed between Ascophyllum 
and any of the truly epiphytic alga. According to Brebner (3),a 
‘milar relation exists between Dumontia filiformis and its host, 

serratus. 
Dasya elegans (Martens) C. Ag. was the third species of the 
odomelaceze examined. The spores attach themselves by 
1 * mucilaginous secretion much less definite in form and less 
_ *Petsistent than in Polysiphonia violacea (Pl. XXII, fig. 30). The 
_ ‘Pore elongates before the division takes place, and in many 
F "aa the very young plantlet assumes an hour-glass shape, 
q — with a delicate mucilaginous secretion at either end. 
' a. "o are parallel to one another, forming a long fila- 
: of the Aeg y, of which one terminal cell becomes the apical cell 
= Nd, the other the primary root-cell (Pl. XXL, figs. 31, 

—. Twelve or more parallel divisions may — 
— cell elongates and forms a rhizoid terminating 
 Petiod (PL st such a modification may appear at - earlier 
i. AM, fig. 34). The multicellular disks, which de 
td of : og are like those of Chondria. The root-cell ort e 
4 old broadens and becomes slightly lobed; oblique 
| als eg the lower corners of the cell; division is continued 
ellular disk with a mucilaginous margin results (PI. 


U, 
P3941; XXUL, figs. 2, 3 7,8,9). The primary Toot 


254 BOTANICAL GAZETTE [octopEs 


cell often branches, each portion giving rise to a disk (Pl. XXI/, 
jig. 42; XXII, fig. 4). Although the primary rhizoid usually 
remains short, in some instances it attains considerable length’ — 
before undergoing division or forming a disk (Pl. XXU/, figs. 37, 
38). While these changes are taking place, the cell adjacent to 
the basal cell sends out rhizoids similar to those arising from the _ 
primary root-cell (Pl. XX///, figs. 1, 5,6). These various root- 
filaments combine to form the primary holdfast, which is after- 
wards strengthened by multicellular branching rhizoids, springing 
from the basal corticating cells of the frond. The course of the 
filaments may be traced for some distance in the holdfast, but it 
is difficult to distinguish between those cells which have their 
origin in the primary disk and those which are derived from the 
corticating filaments. The difficulty in determining the relation- 
ship of the parts is increased by secondary lateral connections, 
which are developed between the corticating cells (PI. XXII, fg. 
43). The marginal cells of the mature holdfast are larger and 
broader in proportion to their length than the corticating cells 
of the frond, and have denser cell contents, but the chromato 
phores of both are separate disks, while those of the — 
siphon are the anastomosing filaments characteristic of hose : 
the Rhodomelacee. A creeping tendency may be exhibited : 
an early age, very young plantlets sometimes developing ™° 
distinct holdfasts of almost equal importance (Fl. XXIII ft 
As in the other species described, many branches arise pea 
massive rounded holdfast, probably springing adventitiony 
from the surface of the latter. 

A comparison of the three eee 
described will show that they agree in forming 4 primary disk: 
cell, which elongates into a rhizoid terminating ina eae a 
and in developing secondary rhizoids, which are sent ee 
root-cell, the cell adjacent to it, and the cortical ce = a 
of the frond. But, while the rhizoids of Polysiphoni r ia 
cellular, unbranched, and free, those of Dasya an pee 
multicellular, branched, and aggregated into a com : 
which in section resembles parenchymatous tissue. 


species of the Rhodomelace® 


pact Cc 


1899] DEVELOPMENT OF HOLDFASTS OF FLORIDE 255 


___ The species remaining for consideration belong to the Cera- 
_ -miacee. Cultures of Spermothamnion Turneri Aresch. were unsuc- 
cessful, only one healthy plantlet having been obtained. It 
developed two small disks, the one a terminal primary holdfast, 
the other a secondary structure arising from one of the middle 
wllsofthe filament (P/. XX//L, fig. gz). As Strémfelt has pointed 
out, the mature rhizoids are short, unicellular, unbranched 
_ ogans, terminating in a lobed disk in which delicate threads of 
_ protoplasm can be traced (Pl. XXIII, figs. 42, 43). The rhizoids 
ue especially abundant near the base of the plants, but, as is 
well known, any cell of a procumbent branch may give rise to 
_ these simple bodies. Now and then, an ordinary branch of the 
_ ftond produces at its apex a much-lobed twisted disk, which 
- tlosely embraces a neighboring branch as a tendril would a sup- 
| port (PL. XXII, jig. 40). ; 

Very few spores of Griffithsia Bornetiana Farlow germinated, 
and the resulting plantlets were short-lived. It is, therefore, 
possible to describe each step in the development of the hold- 
Gow Two unequal cells arise from the first division (teat fig. 
pe * Monosiphonous filament composed of rather globose 
4 cells is formed, the basal cell of which elongates and becomes 
: the root-cell (text fig. 2). The structure of mature holdfasts 
Mould lead one to Suppose that the primary root-cell divides 
p eedly, forming a broad spreading mass of large-celled 
t oY Ep tous tissue. The holdfasts of a young plant 
F thes ® 4 small piece of bone (teat fig. 3) and of older 
/} g owing on Zostera marina L. differed only in size, that of 
2 a! being very large with an abundance of adventitious 
| Hac. gs from its margin. Strémfelt (8) places the 
ae thsia with Ceramium and Callithamnion in a group 


: guish 


‘ Mina” Eatations were seen. The cells of the holdfasts are 
ay :.. and have very dense granular contents, prob- 

oR tlie . bs Supply of reserve food-material. 
“y allied genus, Callithamnion, differs greatly from 


256 BOTANICAL GAZETTE [octorer 


Griffithsia in the form and history of the holdfast. Though the 
spores of Callithamnion Borreri Ag. developed in the laboratory 
with difficulty, cultures sufficient to illustrate the order of devel- 
opment succeeded. After attaching themselves to the slide by 
an almost imperceptible secretion, the spores elongate and 
become pointed at both ends. The first division is parallel to 
the shorter axis (P/. XX//J, fig. 29), and by subsequent partitiona 


C. SALD. ae) / 


Fic. 1. Griffithsia Bornetiana Farlow, germinating tetraspore 
ing. X 333. 

Fic. 2. G. Bornetiana, plantlet 9 days old. X 213. 

Fic. 3. G. Bornetiana, holdfast of small plant grow 

Fics. 4, 5. Polysiphonia fastigiata Greville, growl 
Stack.; in fg. 5 the cells of P. fastigiata are shaded in, t 
merely outlined. & 213. 


g days after 9% 


ng on Ascophyllum gre 
hose of Ascophy as 
of the two f— 

Ju 


monosiphonous filament is formed. The smaller 


minal cells is the primary root-cell (77. XXII, fi: 32): chs 
from one set of cultures, the first cell of the hoi 
a very thick mucilaginous wall, is cut off from 


) DEVELOPMENT OF HOLDFASTS OF FLORIDE 257 


extremity of the root-cell and separated from it by a wall (PZ. 
ANU, figs. 37-33, 37,38). A multicellular holdfast results from 
the division and branching of the primary root-cell and of the 
oldest cell of the disk (Pt. XXT//, figs. 34, 36), and is further 
_ srengthened by a few corticating rhizoidal filaments. In mature 
_ Callithamnion Baileyi Harv. the intracuticular rhizoids are more 
_fmerous. They arise from the lower angles of the central cells 
from the basal cells of the branches. Thence they descend 
through the walls of the monosiphonous frond to the holdfast, 
_ Where they branch freely and spread out in various directions, 
_ lorming, with the filaments arising from the cells of the primary 
tisk, a flat circular holdfast (Pl. XXII, figs. 35, 39). 

The carpospores of Spyridia filamentosa (Wulf.) Harv. germi- 
mie very readily. The mucilaginous secretion, by which the 
‘Peres are first fastened to the substratum, can hardly be dis- 
‘nguished from the wall of the spore (Pi. XXII, fig. 11). The 
eo frst forms two unequal cells, the smaller of which becomes 
the pamary root-cell, the larger divides by parallel walls so as to 
form a short filament (PZ. AXT/, figs. r2-14). Asa rule, no long 
Aan thizoid is formed, but occasional exceptions are found (PZ. 
‘ AM, fig. 78). Though variations may occur (2. XXV//, fig. 15), 
1 first Corticating cells are usually cut off from the upper 
hg the central cells before the formation of the discoidal 
_ oSast begins (Pl. XXTI, jig. 16). Sooner or later, however, the 
aa oaelaa divides in several planes parallel to the longer 
q tds of . and thus produces a flat multicellular disk, the 
21) 7 oa separated by walls (P?. XXTI, figs. 15,17, 19 
4 i. ~ tst the primary disk is but slightly lobed, but the 
; Ito. 8!Ve rise to short filamentous outgrowths, which branch 
j oy (Pl. XXII, figs. 21, 22). Rarely a lateral 
" Weaflam: a the primary root-cell, which is then prolonged 
| by Retin. (72. XT, fig. 23). The cells of the primary hold: 
a to divide, and with them are combined _rhizoi- 
» Which have their origin in the corticating cells of 
Odes of the frond and grow through the cell-walls 
tratum ot XX jigs. 24, 25). Free filamentous 


258 BOTANICAL GAZETTE [ocTopsr 


outgrowths of the larger cortical cells develop disks and enter 
into close mechanical union with the main holdfast, adding greatly 
to its efficiency. The secondary rhizoids thus formed are destitute 
of cortications (P/. XXT/, fig. 29) and are easily distinguished 
from those ordinary branches which, coming in contact with the 
substratum, develop holdfasts (P. XXVI, figs. 26, 28a). Spyridia 
creeps not only by means of such branches but by the aid of 
masses of uncorticated hair-like rhizoids, which may be formed at 
any point (Pi. XXII, fig. 286). One curious instance was noted 
of short rhizoidal outgrowths from each of the central cells ofa 
trailing branch, the cortical cells being unmodified (71. XXII, 
Jig. 27). The differences in the chromatophores are similar to 
those noted in the Rhodomelacez. ; 

Ceramium differs in many respects from the other genera 
described. Both Ceramium rubrum (Huds.) C. Ag. and ¢. alee 
tum Harv. were carefully studied and found to agree closely i 
so far as the early development of the plantlet and the forma- 
tion of the rhizoid are concerned. Shortly after it is sow”, the 
Spore produces a cell-wall (P/. XX///, fig. 24) and a well-defined 
temporary holdfast. The latter is a granular disk of — 
thickness, and mucilaginous in character, attached to the base 0 
the spore (Pl. XXTI/, fig. ro). The granules, embedded inne 
matrix, are often arranged in lines ‘radiating from the axis to 2 
rim of the disk (Pl. XXTV/, fig. 25). The distances pace 
granules being least in vertical planes, the disk appears den a 
when viewed from the side. This peculiar body does not a 
to the ordinary tests for cellulose, and is not dissolved after P 
longed treatment with dilute potassium hydrate. It is 


; - 
neither by Hanstein’s aniline blue nor safranin; but the gO ss 


lar portions stain deeply with haematoxylin and with Congo 


paged fore, 
and the whole with Bismarck-brown. The disk a me: | 
distinct from the cellulose wall of the spore and heer 


ally from ordinary vegetable mucilages, though 
closely allied to the latter. This temporaty t. but has 
peculiar to plants existing in an unnatural environmen" ve 
been found in very young plants growing on Chordati2- 


holdfast is 


fi 


| DEVELOPMENT OF HOLDFASTS OF FLORIDEA 259 


spore develops, the first rhizoid pierces the disk, which then 
becomes disintegrated and finally disappears (Pl. XXV/I/, fig. 
Fi) 


After attaching itself in the manner described, the spore 
elongates and divides several times in parallel planes at right 
_ mgles to its longer axis (P/. XXVII, figs. 11-13). The basal cell, 
_ fowing rapidly, produces a multicellular rhizoid at an early 
_ e; but large plantlets, which have already cut off corticating 
tills, occasionally show little or no tendency to form rhizoids 
i (Pl. XXIII, figs. 14,215,218). The primary root-cell branches into 
_ *veral rhizoids, which are increased in number by outgrowths 
from the cell adjoining the first root-cell (P/. XXT//I, figs. 17, 19, 
_ %,27). Still later, the cortications near the base of the plantlet 
develop multicellular branching rhizoids of great length. All 
femain free throughout the life of the plant, and both primary 
ad secondary rhizoids branch monopodially near the tip, and 
give rise to large multicellular disks of irregular outline 
(7. XXIII, figs. 16, 22, 23, 28). These indented clasping-disks 
ue closely crowded together, cohering so as to form a large 
munded holdfast, in which the various elements may be clearly 
‘inguished. As both of the species are upright in habit, no 
_ “condary holdfasts are developed at any point of the mature 
mAs in several other genera, the chromotaphores of the 
) ale of the holdfasts resemble those of the corticating 
' aie; 1 that those of the central axis, the former being 
ai 27 © latter irregular branching bands (2. XXV//, jigs. 17 


nd evident, therefore, that the species of the Ceramiacee 
ined differ greatly both in the manner of development and 
ot te holdfast, agreeing only in the production of one 
Points . Spermothamnion Turneri forms at various 
oes nos ae unicellular rhizoids with terminal disks, branching 
 banetiong Mt, and cortications are not developed. Griffithsia 
ena puitices a large spreading holdfast composed entirely 
q tell, oY matous tissue arising from the primary root- 
: amnion, Spyridia, and Ceramium have primary 


260 BOTANICAL GAZETTE [ocrosss 


root-cells, from which spring rhizoids terminating in multi- 
cellular disks. Others originate in the cell adjacent to the basal 
cell and in the cortications. In addition the first two possess a 
strengthening mass of intracuticular root-fibers, but Ceramiumis 
quite destitute of them. 

Thus, while of some value in showing relationships, it will 
be seen that the chief interest ina comparative study of the devel- 
oping spores and holdfasts of the Florideze would be in varia- 
tions dependent upon differences in light, temperature, or the 
density of the surrounding medium, and in adaptations to vege 
tative reproduction. 

In closing, I would acknowledge my indebtedness to Dr. 
Setchell, who, in 1895, suggested the holdfasts of the Rhodo- 
phycez as a subject that would repay investigation; to the late 
Dr. Humphrey, under whose helpful and suggestive direction the 
work described in this paper was practically begun; and to 
Professor Penhallow for kind advice. 


McGILL UNIVERSITY, MONTREAL. 


LITERATURE REFERRED TO. 
1. BorGe, O.—Uber die Rhizoidenbildung bei einigen fadenforms® 
Chlorophyceen. Upsala, 1894. 
2. BRANNON, M. A.—The structure and development 
americana Harv. Annals of Botany 11:1. 1897. 


of Grinnellia 


BREBNER, G.—On the origin of the filamentous thallus of Dumontis a 


filiformis. Jour. Linn. Soc. 30: —. 1895. Har. 
4. Davis, B. Mi— Development of the frond of Champia parvult 
from the carpospore. Annals of Botany 6 : 339. 1892. “is 


5. Gipson, R. J. H.— Notes on the histology of Polysiphonta 
(Roth.) aie. Jour. of Bot. 29: 129. 1891. 

6. OLTMANNS, Fr.— Uber die Cultur- und Lebensbedingunges 
Meeresalgen. Jahrb. fiir wiss. Bot. 23 : 349. 1892. 


ca tener J. AE 
7. OsTERHOUT, W. J. V.—On the life-history of Rhabdonta 


Annals of Botan : + 1866. 
any 10: 403. 189 torgane det : 


8. STROMFELT, H. F. G.— Untersuchungen iiber die Haf 
Algen. Bot. Centralbl. 33 : 381. 1888. 


Sai TRS OT is dee a in Rs. 0 ASR ales 


DEVELOPMENT OF HOLDFASTS OF FLORIDEA 261 


EXPLANATION OF PLATES XXI-XXIII. 


PLATE AX, 


Fig. 1. Rhabdonia tenera J. Ag., carpospore 9 hrs, after sowing. X 334. 
Fic. 2. R. tenera, 2 days old. X 334. 

Figs. 3-11. 2. tenera, from 4 to 17 days old. X 334. 

Fig. 12. Lomentaria uncinata Menegh., carpospore. X 334. 

Figs, 13-15, Z. uncinata, 6 days old. X 334. 

Fig. 16. L. uncinata, an unusually large specimen, 6 days old. X 213. 
Figs, 17-19. Champia parvula Harv., plantlet resulting from the ger- 
_ ‘Gination of a carpospore, 8 days old. xX 334. 


Fic. 20. C. parvula, holdfast of a plantlet growing on Polysiphonia. 
X 213. ' 


Fics, 21, 22, Lomentaria uncinata, bases of mature plants. 
Fig. 23, Chondria tenuissima (Good. et Wood) C. Ag., tetraspore shortly 
twas sown. X 233 


Figs. 24-28, ¢. tenutssima, plantlets developed from carpospores, from 
104 days old. x 233. 
Fig. 29, ¢. tenuissima, plantlet resulting from the germination of a tetra- 
Sore, 10 days old. x 233. 

—— Fie30, ¢ tenuissima found growing on mature Chondria. X 233. 

Figs, UE a tenuissima, 3 days old. X 233. 
Fig. 33. ¢ tenuissima, 


Fig, 34. C. tenuissima, 14 days old. X 213. 
Figs, 35, 


' 3. C, lenutssima, 5 days old. X 213. 
: “i 2 ‘enuissima, portion of mature holdfast, stained with methyl 
13. 


12 days old. X 213. 


Oa, C. tenuissima, holdfast of plants found growing on mature 
ined with methyl blue. x 333. 

4156. Polysiphonia violacea (Roth) Greville. 

ae Carpospore and plantlets from 1 to 5 days old. X 400. 
ntlet 7 days old. x 633. 


ee Figs, 5 : 
X too, 47, 48. Plantlets found growing on Scytosiphon lomentartus Ag. 


Ke 
ne aia’ lantlet, result of germinating carpospore. X 213. 


96. Ends of primary rhizoids of plantlets 12 days old. X 400. 


262 _ BOTANICAL GAZETTE [octossr 


PLATE XXII, 

Figs. 1-10, Polysiphonia violacea (Roth) Grev. 

Fic. 1. Plantlet growing on Scytosiphoul omentarius Ag. X 400. 

Fig. 2. Plantlet 3 days old. X 400. 

Fig. 3. Plantlet 9 days old. X 400. 

Fics. 4, 5. Plantlets 12 days old. x 233. 

Fic. 6. Holdfast showing rhizoids springing from corticating cells. X 54. 

Fic. 7. Plantlet 12 days old. x 213. 

Fic. 8. Clasping disk of rhizoid of plant growing on Zostera marina. 
X 400. 

Figs. 9, 10. Plantlets 9 and 11 days old. X 400. 

FIGs. 11-29. Spyridia filamentosa (Wulf.) Harv. 

Fig. 11. Carpospore. X 400. 

Figs, 12-18. Plantlets from 2 to 12 days old. x 400. 

FIGs. 19-23. Bases of plantlets, showing primary cells and clasping disks. 
X 400. 

FiG, 24. Base of rhizoidal branch (c. of fig. 28). X 213. 

Fig. 25. Edge of mature holdfast, in optical section. X 157: ; 

Fig. 26. Transverse section of frond, showing that the origin of rhizoidal 
branches is in the corticating cells; somewhat magnified. 

Fig. 27. Procumbent branch, near 78a, with rhizoidal outgrowths ssi 
central cell. x 333. 

Fig. 28. Creeping branch, slightly magnified. 

F1G, 29. Uncorticated secondary rhizoid with disk. X 333- 

Figs. 30-43. Dasya elegans (Martens) C. Ag. 

Figs. 30, 31. Germinating tetraspore. X 400. 

FIG. 32. Segmenting carpospore 2 days old. X 400. 

Figs. 33-35. Segmenting tetraspore. X 400. 

Figs, 36-38. Plantlets 12 days old, resulting from the 
tetraspores. x 400. 

Figs. 39-42. Plantlets 17 days old, resulting fr 
Carpospores ; figs. 79, 42 show only the root-cell and disk. X 49% 

FIG. 43. Portion of mature holdfasi, in optical section. X 233 


Fane ee A Tg EET Le Oa 


germinatio® of 


ia OS Aang eae a 


ae 
om the germ 


PLATE XXIII. ing 108 she . 

ulti ie 

Figs. 1-9. Dasya elegans (Martens) C. Ag. Plantlets se root-cell and 4 

germination of carpospores; in some cases only the prim 
disk are shown; from 12 to 20 days old. X400. 


1899] DEVELOPMENT OF HOLDFASTS OF FLORIDEZ 263 


Figs. 10-23. Ceramium rubrum (Huds.) C. Ag. 

Fic. 10. Germinating carpospore 2 days after sowing. X 633. 

Figs, 11-17. Plantlets from 3 to 8 days old ; in figs. 75, 76 only the basal 
cells and primary rhizoids are shown. X 400. 

Fig. 18. Plantlet found growing on Polysiphonia. X 400. 

Figs. 19, 20. Plantlets 9 days old. X 400. 

Fic, 21. Plantlet growing on Polysiphonia. X 400. 

Fig, 22, Plantlet 5 days old, showing early branching of the primary 
thizoid. X 400. 

Fic. 23. Holdfast of a rather young plant. X 213. 

Figs. 24-28. Ceramium strictum (Harv). 

Fic. 24. Carpospore 36 hours after it was sown. X 400. 

Fig.25. Plantlet 3 daysold. X 233. 

Figs. 26,27. Plantlets about 7 days old. X 400. 

Fig, 28, Primary root-cell and disk of mature plant. X 400. 

: ag 29-32. Callithamnion Borreri Ag., plantlets from 2 to 6 days old. 


FIG. 33. C. Borreri, plantlet showing basal cell and rudimentary disk, 6 
days old, x 213. 
Fi. 34. C. Borreri, primary disk of plantlet g days old. X 333- 
Pie. 35. C. Baileyi Hary., mature holdfast, in optical section. X 213. 
) Fics, 36, 37. C. Borreri, base of plantlets 6 days old, each showing pri- 
May disk. X goo, 
F * 
ay 38. C. Borreri, plantlet 6 days old. X 213. 
mg © Boiley:, a portion of a mature holdfast. X 100. 
Ne 4. Spermothamnion Turneri Aresch., branch with clasping disk. 


F 
ie Gal. S. Ti urnert, plantlet 6 days old; nine cells of the filament are 
Ee Tepresented, x 213 


Figs, 
42, 43. S. Turneri, mature holdfast. X 213. 


BRIEFER ARTICERS 


A PRACTICAL REFORM IN THE NOMENCLATURE OF 
CULTIVATED PLANTS. 


SoME years ago the Society of American Florists adopted Nichol- 
son’s Dictionary of Gardening as its authority for the names of cult 
vated plants until Jzdex Kewensis should be completed. Jndex Kewen 
sis has been finished for several years, but no florist, nurserymal, 0 
seedsman has standardized the names in his catalogue until in the cas’ 
about to be described. Moreover, no tradesman, so far as I know, has 
ever tried to be absolutely consistent in his names or to follow any ont 
botanical authority. Nevertheless, the seedsmen, nurserymen, and 
florists are bringing up the perplexing problems of nomenclature, 
making resolutions, formulating rules, appointing committees, and 
adopting standards. An intelligent minority is always pressing , 
reform. Standards are adopted and no one follows them. Will they 
ever be followed? Some say no, and affirm that there are essential 
elements in trade that will always make horticulture and botany ©02 
flict more or less. I have long thought otherwise, and now me 
record an experiment that seems to show an entirely practical way 
standardizing the nomenclature of trade catalogues. 

It seemed to me very important that some particular ee 
should be compared with Jndex Kewensis, and every name ce ould 
conform with it. Every name not found in /vdex Kemer if hat- 
then be compared with Nicholson’s Dictionary of Gar dening a0 
monized with that, if possible. This process has actually 
in the catalogue of F. H. Horsford, of Charlotte, Vt. : 

hould be pt 
proved to be an interesting one, and its main features . a 
on record, for some of our best horticultural firms, I i : 
ing to standardize their catalogues, if only they are shont claturs 
doit. We should bear in mind that the principles of eo 
so familiar to every botanist, are entirely unfamiliar to the bus) “ 
culturist with a living to make. . 

At the outset one might readily imagine that ay ; 
seedsman, or florist who has access to /udex Kewensts an 

264 


The casé 


nursery™a : 
Nichols 
[ocrom 


1899] BRIEFER ARTICLES 265 


can do this kind of work himself for his own catalogue. This idea 
will have to be modified. Some tradesmen can do it themselves, but 
most cannot. However, all the important ones can hire it done, and 
cheaply. It is a job that would be exceedingly dry and uncongenial 
to many excellent business men. Competent students, however, can 
be found at the universities who are making their way, and would be 
glad of such work. A thousand names can be standardized for five 
dollars, at the rate of twenty cents an hour for twenty-five hours. This 
does not include the task of rearranging names in alphabetical order, 
othe reading of proof (as some: cataloguers may prefer to do this 
themselves), but only the work of supplying the information necessary 
to the cataloguer. 

There are about seven hundred species in the Horsford catalogue, 
and only twenty-five of those names are not to be found in Jndex 
Kewensis or Nicholson. This is less than 4 per cent., which is surpris- 
ngly low when one reflects on the great number of novelties since 
1893; but Mr. Horsford sells largely of native plants, and these have 
Teceived comparatively few trade names. Moreover, a goodly propor- 
tion of these twenty-five missing names are those of hardy native ferns. 
Index Rewensis has no ferns. 

One barely begins to compare the names of a catalogue with /ndex 
‘ae when he is confronted with an important problem of which 
this is Picture : 
Alyssum saxatile Cran tz = 
Alyssum Saxatile Linnzeus. 
Now, how does the « 
“aployer has in 
'Us? Probabl 
¢ have bot 


A. gemonense. 


standardizing clerk” know whether his 
his nursery the Alyssum saxatile of Crantz or of Lin- 
y he could give a shrewd guess. Possibly he may 
: hy of the original descriptions at hand, and the plants also, bat 
: this ig ue ate iad against it. But, putting such considerations aside, 
of” a matter of identification, not of nomenclature, and the distine- 
3 thing neg a two kinds of work must be grasped at the outset, or 
' , = done. The duty of the nomenclature clerk is clear. 
3 ints in ordi at the plant in the nursery is the one that /ydex Kewensts 
: tnsidered by 7, type. The names in ordinary type, he knows, are 
 Mdlics are S : ndex Kewensis to be the tenable ones, while all those by 
al € can C en It is to Mr. Horsford’s business interest to lo 
Mperly os age Whether the AZyssum saxatile in his nursery 1S 

ot is really Alyssum gemonense. Most tradesmen, 


266 BOTANICAL GAZETTE [octoper 


however, do not have the time, the training, or the books to determine 
all their plants. Identification is the work of the botanist, and the day 
will come, I hope, when all the plants in the nurseries may be identi- 
fied: by specialists. Such work, however, is many times more costly 
than merely following a uniform system of nomenclature. The dis 
tinction between identification and nomenclature cannot be urged too 
strongly. 

Another case is also interesting and occurs frequently: 

Aconitum autumnale Lindley = A. Fischeri. 

Aconitum autumnale Reichenbach = A. Napellus. 

In this case Judex Kewensis does not give any Aconitum autumnale 
in ordinary type, and therefore it recognizes no good species of that 
name. Here, again, the duty of the nomenclature clerk is clear, and 
he writes: 

Aconitum autumnale (Lindl. or Reich.?). 

It is not for him to decide whether the plant in the nursery is really 
A. Fischeri or A. Napellus. He has called the attention of the nursery- 
man to the question, and leaves it open. The nurseryman, perhaps, 
cannot settle the question while his catalogue is going to press, and he 
follows the suggestion of the nomenclature clerk literally. Perhaps he 
may not be able to settle the point for several years, but trade reason: 
are constantly urging him to get the point settled. Meanwhile hi 
consistent and honest to indicate a doubt. Two entirely i 
things have been cultivated under the name of Aconitum ee 
but no one will suspect it if the fact were concealed. Honest @0 
inspires confidence. ae 

The next point will have great weight with the horticnl NT 
Horsford catalogues Anemone montana and Anemone se : 
different things, but Zzdex Kewensis says that the first 18 4 Rage 3 
the second. The nomenclature clerk allows Mr. Horsford od the 
two distinct things under the same names as before, but one 
entries now reads: but hott 
Anemone montana (A. sylvestris according to /ndex Kewenst, : 
culturally distinct with me). be shows 

This is perfectly clear, but too long, 
presently. The important things to note are two. 
man is as free as before to differ in opinion from t™ he opinion ¢. 
ity, but now he is consistent throughout, and supplies the 


1899] BRIEFER ARTICLES 267 


the recognized authority as well as his own. Secondly, every name 
that appeared in the old catalogue appears in the new, but many of 
the old names now appear as synonyms or cross-references. No trade 
name that means money need be omitted. A shorter method of 
expressing a difference of opinion from official standards is to put an 
explanatory note at the beginning of the catalogue to this general 
ect: “Names in brackets show a difference of opinion.” Thus we 
catalogue : 

Anemone montana [.4. sy/ves¢ris]. 


This means that Zxdex Kewensis considers A. sylvestris to be the 
proper name of the species and 4. montana the same thing, or perhaps 
only a botanical variety, while we consider that the two things are dis- 
tinct for horticultural purposes. Instead of suppressing the opinions 
ofothers that conflict with our own, we tolerate them both, and place 
them side by side. 
The commonest situation that needs change is shown by the fol- 
lowing example: Mr. Horsford advertises for sale Achillea Eupatorium. 
* Kewensis says this equals A. filipendulina. The best way for him 
'0 do is to advertise 
Achillea filipendulina. (A. Eupatorium.) 
This makes a great many changes in the alphabetical arrangement 
Species, and sometimes of genera. 
ag way to do is to advertise under the old name, with the 
bite parenthesis, and perhaps in different type, and an expla- 
theo some prominent place of the device used consistently 
cal a the catalogue. This is a far less satisfactory mice: The 
* ing that can be said for it is that it supplies the information. 
2 spied would be cheaper in some cases than revolutionizing a 
vell gue, but if a thing is worth doing at all it is worth doing 
hada points may be briefly mentioned. The name of ee 
: isolutely oii would better not be given in trade catalogues ene 
and aaa rat makes a catalogue look too dry and techn 
E €. There is no officially accepted authority fof popu 
3 Inder iy bid for Names of varieties, whether botanical or horticultural. 
: —e 1S Not supposed to take account of anything below the 
aig gives some varieties and also popular names. : 
appens that Jndex Kewensis gives a species twice } 


es, 
It 


268 BOTANICAL GAZETTE [octonsr 


ordinary type as if both were tenable. For instance, there is a Cam- 

panula strigosa of Vahl and a Campanula strigosa of Solander, both in 

ordinary type. In such a case the nomenclature clerk may write: 
Campanula strigosa (Solander or Vahl ?),. 


It is well to explain in the beginning of a catalogue which names 
are the proper ones and which are the synonyms. The latter are com- 
monly in italics. It makes little difference how a catalogue is arranged, 
provided that there is a full index somewhere. Few indexes are 
enough. The Horsford catalogue has no index, and there are seven 
departments, the arrangement being alphabetical under each depart 
ment. The fact that there are seven departments should therefore be 
prominently stated, and the seven departments listed in the space of aa 
inch or two in such a way that the mind can take in the whole scheme. 
— WILHELM MILLER, Cornell University. 


THE BOTANICAL GARDEN AND INSTITUTE IN PADUA. 


Tue readers of the BoTANICAL GAZETTE may be interested to heat 
something of the ancient Botanical Garden of the University of _ 
instituted by the Venetian Senate in a decree of the pesto 
of June, 1545, through the wise forethought of Francis Bonaf 
in 1543. : 

Pe director, Professor P. A. Saccardo, who has recently improved 
the Institute and the Garden, published some interesting notices ate 
the 350th anniversary of its foundation from which I take the gre# 

part of this note.’ 

Professor Saccardo’s activity turned, in the first place, to eat 
the library, initiated in 1770 by one of his predecessors, John ae 
and enriched afterwards by Professor Bonato and Professor ae 
so that it contains already more than 10,000 volumes. e acs 
books, besides about forty periodical reviews and many aus 5, Vhs 
I must mention the oldest botanical book with instructive figures, 
Herbarium Apuleji Platonict, printed in Rome in 1479- of works 

The director has filled up during recent years the we 
on the floras, especially on the foreign ones, to make apes 


tl saa 
CCL dalla 
anno C Jiotype 


*SAccaRDO, P. A.: L’Orto botanico di Padova nei 1895 ( i 


: ight bh 
fondazione). Padua. 1895. Quarto, with one topographical and elg 
plates, 


1899] BRIEFER ARTICLES 269 


of setting in order the herbaria, which he was then disposed to begin 
and which is now well advanced. 

A hall, built in 1842 as a’greenhouse, was arranged in 1892 to 
contain the general herbarium, consisting of 396 packets disposed 
horizontally in appropriate compartments of two great cases, with about 
44,900 species represented by 60,000 specimens ; the Dalmatian herba- 
rium, composed of 37 packets with 2500 species and 10,000 specimens ; 
and the cryptogamic herbarium, composed of commercial collections 
and those presented to the University. 

The phanerogamic herbarium, especially from the Venetian prov- 
incts (65 packets, 3500 species, 10,000 specimens) is Saccardo’s own 
and is placed in a great hall which was adapted in 1880 as a laboratory 
and contains also collections and materials necessary for scientific 
mstruction. The students do their laboratory work there, under the 
attendant’s guidance. 

The mycologic herbarium, which is also the property of the director, 
deserves Particular mention. It is in the director’s room, in 66 cases 
bX 36X 23%), and represents more than 30,000 specimens, many 
of which come from mycologists, some very rare. Saccardo’s herba- 
“um and mycologic library (300 volumes and 2300 pamphlets) are the 
portant scientific material with which that clever mycologist wrote 
the classic Sylloge Fungorum. In the director’s room near the library 
hg archives of the garden where there are the interesting autographs 
oe Prospero Alpino, Cesalpino, and Pontedera. 
ba *ssor Saccardo has also increased the collection of portraits of 
De Vast ollection initiated by his predecessor, Professor Robert 
Slani. It is really well furnished, especially through gifts made 
4 agg oh of Palermo, the son of the late Bee eee 
Seal of ains about 600 portraits, among whic 


Baron To 


Amer: merican botanists and of botanists who have studied the 
eS flora. 
Qt : 
i. he lecture hall, built in 1842, which can contain two hundred 
: , there ar ; 


Portraits of seventeen professors of botany, in oil ee 
Three weekly lessons on general botany are given in 
a Week ‘at professor ; two free professors give free leseons 
) esp ecially to naturalists and chemists ; Dr. Adrian Fiori, 

‘to the chair, delivers a course on cryptogams and plant pathol- 


ian i 
Ds on Writer delivers a course on plant physiology, with ap aa 
to @gTiculture, 


~ © 8nd-white 


7 {two 


ty 


270 BOTANICAL GAZETTE [octoper 


So much for the Institute; but a great deal might be added as to 
the Garden and its greenhouses, which are rich in interesting plants* 
Classic plants are a Chamerops humilis L. var. arborescens, 9.5" high, 
planted about 1585, and visited September 27, 1796, by Goethe, 
wherefore it is known as ‘‘Goethe’s palm tree;”’ a Zecoma grandijiora 
Del., admired by Goethe for its beautiful flowering ; a very old Viler 
Agnus-castus L. (about 345 years old); an Araucaria excelsa R. Br. 20° 
high, kept in a special greenhouse; many very beautiful trees (Gymmo- 

cladus Canadensis Lam., Gingko biloba L., Diospyrus Lotus \.., Carya 
oliveformis Nutt., etc.). The greenhouses also are furnished with beat 
tiful plants, among them an Astrocaryon Chonta Matt., a Cycas circinalis 
L., a Cycas revoluta Thunb., a Pandanus utilis Bory, a Livistona 
australts R. Br., many Cactaceee and Orchidee. 

More than 5700 plants are cultivated in pots, to which we must add 
110 old trees in the open air, 412 younger trees and shrubs, and 26 old 
greenhouse trees. — J. B. DeTon1, Padua, /taly. 


} 

CONTRIBUTIONS FROM MY HERBARIUM. 
Crataegus Sauratonae, n. sp.— A small tree 3-4” in height, with 
oval crown and ascending or spreading branches, the branches gen 
erally very crooked, as well as the slender twigs; twigs ash-grJ : 
color, and armed, though sparingly, with stout gray oF reddish 5 50 
the twig of the season glabrous and red-brown : leaves glabrous, = 
long, obovate or elliptic, or rhombic-ovate, acute and see oe 
above the middle, mostly entire towards the narrow base, with © 
or four pairs of prominent veins ; the slender petiole 0.5~* and 
stipules, bud scales, and floral bracts not conspicuously en : 
early deciduous: flowers in rather small glabrous phe 
entire, lanceolate, glabrous ; pedicels 1.5-3™ long, ete ; 
fruit about 12™" in diameter, or more ; styles four oT yi 


North Carolina; on the tributaries of the Neuse river, : 

and along streams in Caswell county, N. C.; growing with 

L., the white oak, and shag-bark hickory. del gia” 
e 


Crataegus wirits 


: i : storiche ¢@ °° 
For the accounts of these see R. de Visiani: Di alcune meee oli me 
dino di Padova. Padua, 1856.— G. B. DeToni: Alberi € fruticl 
giardini di Padova, Padua, 1887. 


i899] BRIEFER ARTICLES 271 


CkaTABGUS COLLINA Chapman, which has been reported by Mr.C. D. 
feadle from the mountains of North Carolina, is not uncommon in 
that state as far eastward as Durham county, generally growing along 
the edges of fields or in coppice woods. 

CratakGus VaILIAE Britton seems to be quite distinct from the 
dosely related C. uniflora Moench., often having long, erect, virgate 
tranches, and becoming a tall shrub; while C. uniflora is generally 
lower, seldom more than 1™ in height, with spreading or horizontal 
ranches. It is found in North Carolina as far eastward as Durham 
and Raleigh, 

Crataegus Chapmani (Beadle), n. comb. Crataegus tomentosa Chap- 
mani Beadle, Bor. Gaz. 25°: 36. 1898. This tree is clearly worthy of 
pecific rank. The much broader leaves and more prominent veins, 
sualler fruit in larger corymbs, and more slender spines separate it at 
mee from all forms of C Yomentosa. In leaf characters and especially 
im the numerous pairs of prominent veins there is much resemblance 
0 punctata Jacq. I find Crataegus Chapmani to be not uncommon 
a Ashe county, N. C., and Grayson county, southwestern Virginia. 

__ Fraxtyus prorunpa Bush.— 
Oly from the Jo 

‘ Ihave observed 
 Thorth as Gre 
- “stieed on the 
mas 


This tree has hitherto been reported 
wer part of the Mississippi valley and the Gulf region. 
it, however, in a few places along the Atlantic coast as 
at Pungo swamp, Washington county, N. C. So far as 
Atlantic coast the tree is confined to the largest river 
and the deeper flat swamps with stiff soils, growing with oaks, 
and occasionally loblolly pine. 
E Phang Texana Buckl.—I reported this tree as occurring east of 
E au... mountains, in this journal two years ago (24; 376. 
3 ledmont region of the Carolinas and Virginia. I have 
Mey detected it on the Atlantic coastal plain, in Onslow 
: » Within twenty miles of the Atlantic coast, where I saw a 


pep of trees, some of the specimens being 35” high, and 
Tin diameter. 


| ificent 
Sore than 


RIA PAROLINAE-SEPTeNTRIONALIS Ashe.— This tree proves to 
‘ Ath Atlant Nn in certain portions of the Piedmont regions of the 
‘ Ie stat It prefers dry, rocky soils, steep declivities, and 
dy ridges, though it occasionally enters lowlands. 
ange of low slaty and rocky hills which extends with 
m Maryland to middle South Carolina the tree is fre- 


cet 
ates 


272 BOTANICAL GAZETTE [ocTopER 


quent ; while in central and northwestern Georgia it is more common, 
and it occurs, though locally, in middle Tennessee. 

FOTHERGILLA MONTICOLA Ashe.— The recent discovery of this local 
shrub at Chapel Hill, N. C., makes another station for it about 150 
miles east of any previously reported locality. It grows there ona rocky 
hillside with Rhododendron Catawbiense Mx., and the chestnut oak. This 
is also the most eastward station for Rhododendron Catawbiense Mx. 
Dr. J. K. Small reported Crowder’s mountain as being the most east 
ward station, but Chapel Hill is 140 miles further east, and has an 
elevation rooo feet less than that of Crowder’s mountain, being only 
500 feet above sea level. The Systematic Flora (2: 42) gives the plant 
as occurring only at high elevations. This rhododendron is also 
found abundantly along the Oconneechee hills, twelve miles north: 
west of Chapel Hill, and at a slightly higher elevation. With it at this 
place is Aconitum reclinatum Gray, one of the most local species of 
the genus, and hitherto supposed to be confined to higher elevations, 
5000-6500 feet, in the southern Alleghanies.—W. W. ASHE, Bilt- 
more, NV. C. 


TWO NEW MICHIGAN FUNGI. 
—2.5 broad, thin, conver 


Tubaria luteoalba, n. sp.—Pileus 1 
) sometimes 


becoming plane, finally centrally depressed, the margin e 
becoming partly or wholly upturned, hygrophanous, white, la 
yellowish, silky-squamulose near the margin from the res ae 
veil, margin striate when moist: stem 1.5-2™ long, 0.3-0:5 ic 
hollow, slightly enlarged at base, whitish, silky, downy a ond "es : 
curved: lamell adnato-decurrent, 0.2-0.4™ broad, subdistant, : 
nearly white but soon ochraceous from the spores: spores ellip a 
4-5 X 6-8p.—On decaying stems and leaves of weeds and Sees he 
low wet ground near Michigan Agricultural College, April weg is 
This fungus resembles 7. furfuracea in form se ee an it 
smaller, lighter in color, and in every way more delicate oe 
species. The spores are also smaller and lighter in eee me i. 
autochthona it differs in its larger size, form of stem, and ha a 
veil sometime forms a fibrous zone on the stem. It has not 
lected in any other locality. cools per 
Galera crispa, n. sp.—Pileus 1.5-3-5™ broad, seipaessee rivalos 
sistently conico-campanulate, subacute, uneven and somew 


1899] BRIEFER ARTICLES 273 


ochraceous-brown on disk, lighter toward the margin which becomes 
cenulate and upturned in older specimens, slightly pruinose at first, 
mugulose and a little paler when dry: lamelle slightly adnexed, not 
cowded, rather narrow, interspaced with anastomosing veins, much 
crisped, at first nearly white, then becoming ferruginous from the 
spores: stem 7-10 long, o.1-0.3™ thick, tapering from the somewhat 
bulbous base, yellowish-white, pruinose at base, hollow, fragile: spores 
Sion broad, 12-16 long.— In grass in dooryards and pastures, June 
and July, 

___ The specific name is based on the peculiar character of the gills 
Which are always crisped as soon as the pileus isexpanded. Professor 
Charles H. Peck, to whom specimens of the above fungi were referred, 
and who very kindly reported on them, suggested that this might be a 
tariety of G. dateritia, unless the peculiar character of the lamellz 
_ Moved to be constant. The fact that specimens possessing this char- 
_ ‘ter have been collected in the same localities during the past three 
“asons seems sufficient proof of the constancy of this character, and 
_ ““asequently, would indicate that this form is worthy of specific rank. 
~B.0. LoNcyrar, Michigan Agricultural College. 


MEXICAN FUNGI. II. 


, Tar following species of Ustilaginez were collected by me in 1896 
re v8, and all, except the three marked with an asterisk, have been 
ay by P. Hennings, of Berlin. 

¢ snaRO RaBennorstiana P. Henn. On Panicum filiforme. 
if “Jara, Oct. 12, 1896. 

a PAMPARUM Speg. On Sefaria. City of Mexico, Oct. 2, 


Usr 


2 — Uter P. Henn. On Chloris submutica. City of Mexico, 
9 


b 
Ustina 


| WMerico, 0 Dieretiana P. Henn. On Tripsacum dactyloides. City 


Oct iT 896. 


Usty 
| Me ‘AGO Hivarraé P. Henn. On Ailaria cenchroides. City of 
= Ico, Oct. 2, 18 6 


y - Ustizagg cop 
City of Me 


1p 
wee For previous pape 


OGINIS P. Henn. On A gopogon cenchroides P. 
xico, Oct. 3, 1896. 


Tr see Bor, Gaz. 24:23. 1897. 


274 BOTANICAL GAZETTE [ocToner 


UstiLaco Brunk! Ell. and Gallw. In culms of Andropogon per- 
foratus. Cardenas, Mexico, Oct. 22, 1898. 
UstTILAGo NuDA (Jens.) Kell. & Swingle. On Hordeum vulgare. 
Toluca, Sept. 19, 1898. 
Ustitaco Crencuri Lagerh. On Cenchrus (tribuloides?). Carde- 
nas, Oct. 22, 1898. 
Ustilago Holwayana P. Henn., n. sp. Soris in spicis, es 
destruentibus, longe cylindraceis, primo membrana pallide cinerea 
velatis, atris; sporis subglobosis vel ellipsoideis, subacutangulis, att 
violaceis, 13-16 X 12-15; episporo verrucoso, atro.—In spicis Pat 
palit velutint. Patzcuaro, Michoacan, Oct. 19, 1898. 
Ustilago Andropoginis-hirtefolii P. Henn., n. sp. Soris in panicalis, 
ovaria destruentibus, longe cylindraceis, membrana cinnamonea vestitis, 
dein pulverulentis, atro-olivaceis vel fusco-olivaceis, 9-14/; episporo 
granuloso verrucoso.— In paniculis Andropogonis hirtifolii pubifiore. 
Patzcuaro, Michoacan, Oct. 20, 1898. Apparently quite distinct from 
all the species known on Andropogon. : 
Ustilago Panici-proliferi P. Henn., n. sp. Soris in paniculis, € 
omnino destruentibus, membrano cinerea tectis, dein pulverulents 
atro-olivaceis; sporis subglobosis, acutangulis, pallide olivaceo-fusts 
intus granulatis, 7-9; episporo levi.—In paniculis i prolifer 
acuminati. City of Mexico, Oct. 10, 1898. This species differs we 
U. Panici-miliacei (Pers.) Wint. in its much smaller spores, and in as 
ing the sori covered with an ashy-gray membrane; and ror U. eh 
Josa P. Henn. and U. pustulata Tr. & Earle in its smooth lighter 
ored spores. The sori and general appearance are also different. 


exit 
*Ustitaco Partatoret F. de Wald. On Aumex Me | 
Meissn. Toluca, Mexico, Sept. 19, 1898. = ieee OM re 
*Ustitaco Tritici (Pers.) Jens. On Z7iecum vuigare. : 


Mexico, Oct. 11, 1898. ae ; 
*UstiLaco Zem (Beckm.) Ung. On Zea Mays. City 
Oct 10, 1808. 
CINTRACTIA AXICOLA Cornu. On Fimbristylts. | 
ico, Sept. 26, 1898. port 
Beracens Saison (Berk.) P. Henn. On Rhynchon” 
Jalapa, Mexico, Oct. 3, 1898.—E. W. D. HoLway, 


Cuernavaca, aed : 


Decorah, low 


OPEN LETTERS. 


TO BRYOLOGISTS. 


Tue undersigned desires to make a full canvass of all working bryologists 
and collectors of mosses, both American and foreign, believing that a better 
general acquaintance will further both the friendly and scientific spirit. To 
this end the following data are cordially solicited from all who have not already 
sent them ; 


1. Name in full; age; vocation. 

2. Average time your vocation annually permits you to devote to mosses. 

3. A list of your bryological publications ; alsoa list of exsiccati you may 
tave distributed, 

‘ Have you worked on any foreign moss flora? State in which geo- 
Sphical region you are most interested. 

§- Which genera, or larger groups, are you making your specialty ? 

- Are you desirous to have referred to you for critical examination 
*pecies that fall in the line of your special interest ? 

7. Do you exchange mosses? If so, what material do you offer? 

A circular embodying these questions was sent out in June of the preaee 
ae ‘Was answered so approvingly and enthusiastically that I feel pagel 
j me m making an additional effort to make the data as complete as possible, 
{ oe collected into permanent form, and to place them finally 
q hands of each one interested.—Joun M. HouzinceR, Winona, Minn. 


CURRENT LITERATURE. 


BOOK REVIEWS. 


Botanical teaching. 


A ToPIc which is or should be of deep interest to professional botanists 
in our higher institutions is the nature and quality of the botanical instruction 
in the secondary schools. Many of them have shown their interest by per 
sonal endeavor to help teachers to improve the scope of their courses and 
secure proper equipment of laboratories for instruction, This endeavor's 
bearing fruit, and the rapid improvement in botanical teaching augurs well 
for future development. Besides the innumerable fugitive addresses 
teachers’ associations and institutes, several modern text-books and laboratory 
guides are helping along the good work. : 

Professor Ganong now contributes a book which will do much to asst q 
teachers to strengthen their botanical work. This book, The Teaching P 


Botanist, has a pedagogical purpose, as distinct from the texts and labor 
The first is made up 


they will become more widely efficient than heretofore. 
proclaimed the new gospel will be delighted to have this boo 
may refer teachers seeking such help. It will save reams 0 
hours of talk. 

If we may choose among the good things in this 
“What botany is of most worth,” “On things essentia 
teaching,” and “On some common errors prejudicial to good 
ing” are probably the most useful. But the suggestions On “ee 
description, on laboratories and their equipment, on collections, an 
are excellent and sure to be helpful. 

The second part consists of ‘‘an outline for a synt 
in the science of botany,” conforming to the sabre 

ivisi rst t 
part. This course consists of two divisions, the 5 of the larger 


k to which they 
£ letters and 


part, the essays OF 


hetic element’ argh 
s elucidated yexeal 


of plants. The course is divided into various topics with mer pupil 
tions and directions which might be put directly i : 
These are followed by notes regarding the materia 
on the pedagogical import of the various points called fon, focros® 


1899] CURRENT LITERATURE 277 


Doubtless few teachers will want to follow exactly this course, nor does 
the author expect them to do so; but many will certainly derive great help 
by selecting from it the topics appropriate to their own conditions and having 
dearly before them the didactic value of the laboratory work. 

It may be worth while to point out that the author’s principles, which one 
tannot escape, do not compel the conclusion that it is best to begin the ele- 
mentary course with a study of seeds. The teacher who now begins by 
mtroducing the student to the simple algz need not feel that he must aban- 
ton this method. The excellent principles presented in the second essay 
may be as well developed by another method. Aanad it is only fair to say that 
Professor Ganong advises each teacher to make out his own course. 

University men will do well to read Dr. Ganong’s essays and recommend 

book to every teacher of botany.—C. R. B. 


Buds and stipules. 


Sir Joun Luspock has published a book with the above title in the well- 
te International Scientific Series.t There is little or no attempt to give 
“ything new, but rather to place before the world in a somewhat popular 
syle the most interesting results of his previous study.” 

The author was led to study stipules by the observation of Vaucher that 
sme tock-roses have stipules and others not; the question arose: why? 
The Study of stipules led on to a study of buds, especially their protective 
Structures, 

The order of the chapters does not seem particularly logical, and there 


‘pail more repetition than is needed, even in a popular work. The 
; the dey 


Setting; 008 Chapter on the structure of buds, many species being 
| _ ee Six treats of the forms of stipules, and it is shown oi 
Uses of ig 's. In the seventh chapter the author discusses the su : 
Poteet the buds a Their general use he conceives to be to cover an 
» "Lustoc - They are often important also as organs of photosynthes!s ; 
4 Sg On buds and stipules. Crown 8vo. pp- xix + 239. pl 4 
"0 stipule et Megan Paul, Trench, Triibner & Co. L’t’d. 1899. 55 
S.C I-IV. Jour. Linn, Soc. Bot. 28, 30, 33- 1890, 1894, 1897. 


| 


278 BOTANICAL GAZETTE [octomm 


they may become tendrils, or spines, or glandular organs; or they maybe 
rudiments, looking back to organs of use in another form. 

There is a chapter also on fhe nature of stipules. There are three views 
as to what stipules are: (1) they are appendages of the leaves (Van Tieghem, 
Baillon, Gray); (2) they are autonomous organs, analogous to leaves (Liné 
ley) ; (3) they are an integral part of the leaf. Lubbock holds the third view, 
The first view he regards as untenable because stipules originate indepent- 4 
ently of leaves and often before them; the second because the stipule 
bundles are derived from the foliar bundles. ; 

The book is full of illustrations and very suggestive, though it seems that 
there is too great a certainty as to just what everything is for—HenriG 
CowLeEs. : 


An ecological text-book. 


AMONG THE recent text-books for secondary schools none is so dominateé 
by the new ecological standpoint as the book just issued by Dr. ace uM. 
Coulter. This is one of the series of “Twentieth Century Text-books, ® 
course of publication by Messrs. D. Appleton & Co.’ 

It is the first of a pair of books, each representing work for half 1 
but independent. The second, with the title Plant Structures, is to be issued 
shortly. It is to be dominated by morphology as the first is pips 
ecology. Inthe judgment of Dr. Coulter the order in which he pee 
books is the proper one for presentation in an elementary howe: d 
sequence is likely to meet with the criticism that the student, ignorance 
plant structure and without wide acquaintance with plant groups, |S 
to appreciate ecological phenomena and _ principles. The author De ae 
the advantages which counterbalance the disadvantages are (1) the agile ] 
of a true conception of plants in nature, (2) acquaintance with the puna : 
lems of plant -physiognomy, and (3) the avoidance of the use of the iy 
microscope at the outset. : | 

Though the physiognomy of vegetation is an interesting ys ae 
most important phase of botany, it is doubtful whether at the Pe a 
the subject is well enough organized to justify its dominating grit yeas 
course. It is still more doubtful whether it will be possible fo 
to find teachers capable of presenting it. Granting the ecologica 
be the ideal botanical course, the question is whether We. ce 
away from the floristic or pseudo-taxonomic teaching t0 psig an to he 
reach so remote an ideal. The writer has already committ ghould be first 
view that the simpler morphology and physiological topics: ss pedagne® 
presented in an elementary course and therefore only states WF pie 
’ 3Coutrer, J. M: Plant Relations, a first book of botany. 120 eee 

figs. 206. New York: D. Appleton & Company, 1899. 


1849] CURRENT LITERATURE 279 


pmblem. To its solution Dr. W. F. Ganong contributes interesting argu- 
ments in a book elsewhere reviewed. 
Readers of Plant Relations will be impressed by the terse and lucid style. 
Though the utmost condensation has been necessary, the author has pre- 
served a simplicity of language and has attained a degree of accuracy which 
leaves nothing to be desired. The book is also striking in the number and 
beauty of its many illustrations, of which a large part are original. Amon 
the finest ones are those derived from Schimper’s recent treatise, Pfanzen- 
Seographie, 
A very useful pamphlet of twenty pages, embodying suggestions to 
teachers for the use of the book, is designed to accompany it. It contains 
helpful remarks regarding the laboratory and field work which the author 
~ of course, shall be prosecuted as the foundation for the study of 
text, 


Weare sure that live school-teachers will welcome this book because it 
Pesents a new view of the plant world, valuable for instructional purposes 
and hitherto too much overlooked. University teachers will receive it gladly 
because it emphasizes one of the vital aspects of botany, and makes more 
. the crusade against the cut and dried formalism of “analysis.” — 


Cytological technique. 


INVESTIGATIONS upon the structure of protoplasm demand not only 
seme skill in mechanical manipulation but also a knowledge of the prin- 
~ underlying fixing, staining, and other details of microtechnique. A 
meent book hy Dr. Alfred Fischer puts the whole subject of microtechnique 
‘Pon a firmer and more philosophical basis and gives an up-to-date discus- 
So of modern theories of protoplasmic structure.* 
and descr; ai fixing agents, considers in detail the solutions in common use 
bumose their action upon the various cell contents, as peptone, proal- 
» Nucleic acid, nuclein, etc., etc. The numerous experiments with 
' what Ces of known chemical composition should be of value in determining 
© to be regarded as artifacts and what as structural elements of the 
hose engaged in cytological work. : 
ei Pages) is devoted to staining. Both theory and practice 
Ott of the ¢ din detail. Some of the topics are as follows: T he washing 
‘ XINg agent, and its significance in theories of staining ; staining 
taining solutions without differentiation; double staining with 
: Solutions ; Simultaneous double staining with mixed stains ; impreg- 
Themen AUTRED.: Fixirung, Firbung und Bau des Protoplasmas. preg 
Phe ae a sae Technik und Theorie in der neueren Zellforschung. 5V0 P 
‘8S. 21. Leipzig: Gustav Fischer. 1899. 4/11. 


280 BOTANICAL GAZETTE [OCTOBER 


nation ; objections to the physical theory of staining; chromatin and the 
fundamental doctrines of staining. Here again experiments upon substances 
of known chemical composition,occupy a large part of the space. 

Part III deals with the structure of protoplasm. Spindles, centrosomes, 
and radiations are thoroughly discussed and artificial figures are compared 
with those occurring normally. Chromatin is treated in the paragraphs on 
granules. The various theories of the structure of protoplasm, as the granula 
theory, the network theory, the filar theory, and the foam-structure theory, art 
critically reviewed.—CuaAs. J. CHAMBERLAIN. 


Knuth’s Handbook. 


STUDENTS of the interrelations between plants and their pollinators, con- 
stituting a branch of what the Germans call “ biology,” and what Americans 
are coming to call “ecology,” have learned their first lessons in large part 
from Christian Konrad Sprengel, once rector of the Lutheran Stadtschule at 
Spandau, Charles Darwin, and Hermann Miiller, late Professor in the Real- 
schule at Lippstadt. Some years since, Sprengel’s book, « Das entdeckte 

-Geheimnis der Natur im Bau und in der Befruchtung der Blumen,” ws 
reissued by Professor Paul Knuth, of the Ober-Realschule at Kiel; and - 
English translation of Miiller’s ‘“Befruchtung der Blumen” has brought his 
work within reach of many persons not familiar with the German language. 
It appears that the original edition is no longer procurable, 
Knuth set himself the task of revising and reissuing it. 
branch of science has been so great in the last quarter century, bowers? 'g 
he has found it better to write an independent work,° based on Miller 
writings, but brought up to date. os 

For reasons not perfectly clear to the uninitiated, this has been divided 
into three parts: an introduction and bibliography, pollination i 
in Europe and the arctic region, and extra-European Sst escab 
of flowers. The first two volumes are now issued, in three parts, ae : 
comprehensive index. The third volume is announced as in course © igs 
ration, and will be received with no little satisfaction on its completion ‘ a 
lines of botanical work are so fascinating or so accessible to the beginn me 
pollination studies, and with this book before him he should be able 
to sift the known from the new in his observations, so that the a much 
added to the former in suitable published form. It is doubtless pt st 
hope for an English translation, but the absence of one Is gre patt 
indication of the urgent need of a working knowledge ot Ce ee 

_ of every student ambitious to distinguish himself in modern scien aon 
—WILLIAM TRELEASE. Her 
yon Her 

SKNuTH, PAUL: Handbuch der Blutenbiologie unter Zugrandeleg nas 
manu Miiller’s Werk, “ Die Befruchtung der Blumen durch Insekten. a | 

Leipzig. 1898-9. 


1899] CURRENT LITERATURE 


MINOR NOTICES. 


A monoGraPH of the hypogaeous fungi of California has been published 
by Mr. H. W. Harkness.° One hundred and eight species are described, of 
which fifty-eight are new, with five new genera. There is unfortunately no 
mtificial key or synopsis, so that the work will be of little value except to the 
specialist, and we fear he will have occasion to complain of the brevity of the 
diagnoses of new genera and species. These rarely exceed two or three lines, 
with few or no explanatory remarks.—C. R. B 


__ F.Y. Covitte and G. N. Rose have published a list of plants collected 
by Mr. and Mrs. Leonhard Stejneger on the Commander islands during 1895 

| amd 1897, These islands bear no trees, due, the authors think, to the violent 
wet winds that Sweep over them during the winter. The geographic relation- 
_ Ship of the flora is primarily Kamchatkan, with strong Aleutian and arctic 
elements, and there is almost no insular specialization. The list is reprinted 
The fur seals and fur-seal islands of the North Pacific Ocean, part 4, pp. 
3§2-361. 1899.— J. M. C. 
“BoTanizinG” is the title of the new book by the author of Zhe 
Botanical Collector’s Handbook, Professor W. W. Bailey, of Brown Univer- 
‘ i It embraces the material of the former volume but is completely 
_ ‘“wmtten and greatly improved. The amateur will here find the completest 
‘ and best directions for all kinds of botanical collecting. The author has been 
3 eased by various specialists, some making only general suggestions, others 
3 Meparing the sections relating to particular families of phanerogams and the 
: larger groups of cryptogams. Fifteen illustrations show useful apparatus for 
_ Jn of specimens. An index should have been added. The author 


: ’ 
a to delete the words “the steppes of Asia or” on page 4, 
—_— Ps : B. 


NOTES FOR STUDENTS 


at : oy aaa PAPER read before the Botanical Society of eee 
- Betaly ar: . Charles E. Bessey discussed the probable significance . 
ies with diclinism and diceciousness. He suggeste 
Ser flowers _~ plants, as shrubs and trees, as well as those epee 
Comp bi ay dispense with petals. Petaly is apparently correlate wi 
%» and apetaly with anemophily. Apetaly and diclinism appear 
alk togressively increasing aphanisis. Lists of apetalous and 
ants Were given and discussed. 
eas - Calif, Acad. Sci 
oui ale Botanizing : a guide to field-collecting and herbarium work. 
42. figs. 15. Providence: Preston & Rounds. 1899. 


- ILL. Bot. 1:241-286, db/. col. pl. 42-45. 1899. 


282 BOTANICAL GAZETTE [octonsr 


AT THE COLUMBUS MEETING of the A. A. A.S., H. L. Bolley presented 
Section G a paper by Lawrence Waldron on “ The occurrence of calcium 
oxalate and lignin during the differentiation of the buds of Prunus Ama 
cana.” It was found that the crystals of calcium oxalate occur in quite ser 
prising abundance in the meristematic tissues of the bud, and in the vey 
youngest stages of the scales of the bud; and that the oxalate become 
lessened in proportionate quantity as the tissues develop. Lignification 
the hairs and scales of the bud commences at a very early period of ther 
development. While it is usually assumed that calcium oxalate is a wast 
product of metabolism, its occurrence in such large quantities in the mete 
stematic cells of the bud and scales would seem to indicate a question a 
whether it has not a definite value at this point at this particular time in the 
life history of the plant. 


PROTHALLIA OF LycopopIuM are so rare that the present pape 
although based upon only half a dozen specimens, is a valuable contribution! 
These prothallia resemble those of Botrychium Virgintanum as described by 
Jeffreys. A vertical section shows a limiting layer of colorless cells, above 
which are several cells invested by an endophytic fungus. The cells of the 
upper half of the prothallium are entirely free from the fungus. The anthe- 
ridium is developed from a single superficial cell and at maturity does eo 
project above the surface. The archegonium, also developed from 4 ~ 
superficial cell, projects considerably. There are six or eight canal ¢ 
the lowest presumably a ventral canal cell. ae 

The value of the prothallium as a taxonomic character IS disc 


: ‘um whi 
some length, and the writer concludes that species of Lycopodium 
possess similar prothallia cannot, on that ground alone, be — 

um in form, 


ace 
r ; 
ytic habit— Cuas. | ; 


CHAMBERLAIN. . 
AN EXCEEDINGLY interesting account of the fertilization of _ 
spermum Bohneri by W. Schmidle® has recently appeared. ye agreting 
worked on material gathered in Germany and his reer : : 
with my own studies in respect to the presence and behavior ® : 
Ann. Bot. 19:77 


3 


®Lanc, W. H.: The prothallus of Lycopodium clavatum L. 
317. 1899. rinsertion ™ 

9W.ScHMIDLE : Einiges iiber die Befruchtung, Keimung, und ost 
Batrachospermum. Bot. Zeit. 57':125. 1899. 


ss 


thea] CURRENT LITERATURE 283 
phores in the trichogyne and antherozoids, are very different as regards the 
_ wtivities of the nuclei and processes of fertilization. 

The antherozoids of B. Bohner? differ from any that I have ever seen in hav- 
ing almost invariably two nuclei. When the antherozoid fuses with the tricho- 
gyne the nucleus nearest to the point of application passes into that structure 

amd is usually followed by the second, which however may remain behind in 
the antherozoid. 

__ Thetrichogyne and carpogonium of 2. Bohneri have only one nucleus which 
lies in the carpogonium. One of the nuclei from an antherozoid passes the 
_ agthof the trichogyne into th pogonium through the constriction between 
_ thetwo structures. Fertilization takes places in the carpogonium, where the 
_ male nucleus fuses with the female. 

___ The passage of the male nucleus causes the protoplasm in the trichogyne 
_ ‘Wgather into a peculiar dense mass which indicates a certain stage in the 
_ process of fertilization, 


Alter fertilization the ca i i 
2 rpogonium becomes cut off from the trichogyne, 
and the late : sal 


€r may then contain one or more nuclei introduced from one or 
iter antherozoids that fuse with it. Extensive fragmentation of these 
r. 


may occur late 
___ The phenomena describ 
. tage af the processes of fe 
 Oltmanns and Ww 
Thave 


ed are in all respects compatible with our knowl- 
ttilization. They accord fully with the accounts of 
ille for other members of the Rhodophycee. 
realized for a long time that my account of Batrachospermum * left 
pomtered in a very unsatisfactory state. The conditions that 
Were exceptional and at variance with those so generally present 
However, at the time, I was convinced of the 
d since that publication I have several times 
ways finding the same structures that I figured 


ss of my position, an 
~ 4 MY material, al 


to the nucleus that I have supposed 
Such a stage would be exceedingly 


"© Rot wish 1b ese: : : 
“iy 8 mutise the work of Schmidle further than to suggest 


» together with an absolutely complete 
ss of fertilization, 


fertilization of Batrachos 


: Ustrating the proce 
“Davis: The 


permum. Ann, of Bot. 10:49. 1896. 


284 . BOTANICAL GAZETTE [ocronss 


These can only be obtained from material very carefully killed and fixed, 
perhaps after special methods. I must believe that we shall not feel sure of 
the processes in Batrachospermum until the technique of the investigation is 
developed té6 a point much superior to that of either Schmidle or myself.— 
BRADLEY MOORE Davis. 


BARTHOLD HANSTEEN publishes the extended results of his researcheson 
the synthesis of proteids in green phanerogams in Pringsheim’s Jahrbiicher* 
He used Lemna minor, Vicia Faba, and Ricinus communis as experimental 
plants. His summary we translate: 

1. In general, at least, light plays no direct réle in the synthesis of proteids 
in the bodies of green phanerogamous plants. In these the formation of 
proteids occurs in active cells without the influence of light and independent 
of the time of year, if only suitable conditions for growth be present. 

. Glutamin, asparagin, urea, ammonium chlorid or ammonium sulfate 
combine with available grape sugar or—at least the four last named nitrogea 
compounds—with the direct reducing sugar formed in the digestion of 
starch. 

d. Urea or glycocoll generally unite either with available cane sugat ® 
probably indirectly reducing sugar. 

2. The chemical nature of the immediately available carbohydrate is not 
unimportant for proteid synthesis; on it primarily depends whether the 
formation of proteid is effected or not. : 

3. The various amides (amido-acids) or nitrogen compounds are ogee 
ally not physiologically equivalent for proteid formation. The best suited 
for this purpose is urea, whose transformation into proteid occurs a5 Oe 
getically with cane sugar as with grape sugar. On the contrary, api 
alanin, and creatin cannot be looked upon as materials so suitable for af 
making; for even under the most favorable conditions, and equally whet 
direct or indirect reducing sugar in suitable amounts is simultan . 
accumulated in the cells, proteid formation from these compounds tele 


™ Jahrb. f. wiss. Bot. 33 : 417-486. 1899. 


NEWS. 


Mk. ABeL A, HunTeR has been appointed botanical collector for the 
University of Nebraska. 


Tar University of Geneva has conferred the degree of Ph.D. honoris 
fausa upon M, Casimir de Candolle. 


Dr. Cart E. CorrENs has been advanced to the assistant professorship 
of botany in the University of Tiibingen. 

Dr. WILHELM FIGDOR has been appointed docent for anatomy and 
physiology of plants in the University of Vienna. 


HouGHTON, MIFFLIN & Co.’s fall announcements include a book entitled 
Animal and plant lore, by Mrs. Fanny D. Bergen. 


Dr. J. M. JANSE, of the botanical garden at Buitenzorg, Java, has been 
*ppointed professor of botany in the University of Leiden. 


; nta 
Ds. K. GiesennaGEn, of Munich, has received from the governme 
sibvention of 4/6000 for an investigating tour to Malacca. 


Dr. L. HILTNER has been appointed director of the hectetoe 
Rory of the biological division of the Imperial Bureau of Hygiene in bertin. 


PROPESSOR GEORGE W. Marrtrn, teacher of biology in the sep Hb 
*, ool, has been appointed professor of biology in Vanderbilt Univer- 
‘ wy, Nashville, Tennessee, . 

Pit PeNGLe has returned from his fiteentli year in the collection 
4 ' Mexican plants. His health has not been good, but, as usual, : deieesi 
mith * collection of choice plants. 


q Dr, Pav Kuru, of Kiel, has returned to his home from his pene 
—. the world, He has brought with him abundant ecological ma 

a Java, Japan, and California. arg 

a RECENT FIRE which destroyed the entire stock of Jacob spe es 

»M Lincoln. Neb., all the unsold copies of Pound and Clements 


abhy of Nebraska were burned. 
: Iniver- 
hy ress N. BERLESE, heretofore professor of botany in the U nile 
= * Camerino, has been appointed professor of natural sciences 
‘ * Lyceum, D t 


r. J. B. DeToni has been appointed to the position 
285 


9) 


286 BOTANICAL GAZETTE [octoper 


vacated by Professor Berlese. Dr. DeToni’s address, however, will still be 


On May I the north wing of the great temperate house at Kew Gardens 
was opened to the public. The center of this house was built in 1862, and 
the south wing in1894. The building is 628 feet long, with a maximum width 
of 164 feet and a height of 60 feet. It covers one and two thirds acres, and 
has cost £43,000. 


THE INTERNATIONAL scientific medal of the Academie Internationale & 
Geographie Botanigue has been conferred upon Dr. N. M. Glatfelter, of St 
Louis, for his work upon Salix, and upon Dr. Roscoe Pound, of Lincoln, Neb. 
for his phytogeographical researches. Fifteen investigators in Europe have 
been similarly honored. 


THE LINNEAN MEDAL has been awarded to Mr. J. G. Baker, late keeper 
of the herbarium and library of the Kew Gardens, “ for his services to Dotan 
during a long series of years, especially his writings on ferns and petaloid 
monocotyledons, serviceable alike to botanists and cultivators.” The pres 
tation occurred at the anniversary meeting, May 24. 


THE MACMILLAN CoMPANY announce for publication this autumn 4 
Handbook of North American Myxomycetes by Professor Thos. H. Macbride, 
of the University of Iowa. The work includes descriptions of all yi 
hitherto described from North America, with brief synonymy, accompat 
by diagnostic notes. The work is to be illustrated by nineteen full-pag® 
plates. 


A ein, the 
THE cooperation of a local mountain club, Dr. R. von Wettstein, © 


Y 
director of the Vienna botanical garden, has been enabled to establish a ii 
logical experiment station in the Tyrolese central Al 
Hiitte”’ in the Gschnitzthal, at an elevation of 2300". . 
has been fitted up for a laboratory. Research will b 
production of species by direct adaptation. 


Mr. C. L. Potiarp of the National Museum, aided by 
Greene, has undertaken the distribution of authentic sets of N 
Violacez. Forty sets are in preparation and the muscu” 
return for an equivalent amount of selected duplicates, either 10 
or other groups. Decades will be issued at irregular intervals, 
material can be secured and labels printed. 


ffers oo 
the Violace® 
as rapidly . 
oe Js oft 
THE POSSIBILITY of obtaining separates of articles from eae jibe 
renders possible the carrying on of investigations apart tog “rectates : 
ries. We welcome, therefore, every establishment which ae . 
diffusion of such literature. Mr. A. I. Eriksson, Tufts mee separates 
into the business of dealing in natural history books and aut : 


, i899] NEWS 287 


ie will act as agent for authors who wish to dispose of any of their publica- 
tons, A catalogue will be issued shortly. 


AT THE MEETING of the St. Louis Academy of Science, held on the even- 
ing of October 16, Dr. H. von Schrenk presented some notes on Arceuthobium 
fusillum, which was found in Maine during the past summer, growing on the 
white spruce along the seacoast. The trees which are attacked form large 
witches’ brooms, the branches of which are much longer than the normal 
tranches. The manner in which the seeds are distributed was briefly 
described, and seeds were exhibited adhering to branches of the white spruce. 


Mr. 0. F. Coox, of the Division of Botany, Department of Agriculture, 
_ tas been detailed to make a preliminary examination of the plant products 
_ of Puerto Rico with reference to the introduction of new and useful tropical 
_ plants into that island. Mr. Cook is accompanied by Mr. G. N. Collins of 
_ the Department of Agriculture as photographer, and by Mr. George P. Gall, 
Who is sent by the Smithsonian Institution to collect material for the National 
Herbarium. The expedition left New York on October 28 by the United 
States transport MacPherson.— Science, Nov. 3, 1899. 


THE DISTINGUISHED French horticulturist Henri Lévéque de Vilmorin 
/ Msstricken by apoplexy and died at Verriéres on August 24, in his fifty- 
“venth year. He visited this country in 1893, when many of our botanists 
the pleasure of meeting him at Madison, Wisconsin. Henry Vilmorin 
_ “Snot only the head of the historic house of Vilmorin & Co., the largest 
: sed firm in France, but personally directed in large part the numerous 
’ “periments for the improvement of cultivated plants which the firm were 
; “ontinually conducting at Antibes, Verriéres, and Ferme de St. Fiacre. bee 
; — was widely honored by scientific societies and has made valuable con- 
8s to horticultural literature. 
ON Marcu 3, 


1899, a bill was passed by Congress providing “that on or 
anuary I 


_1, 1903, the fence around the Botanical Garden shall be 
E Tinie. Provided that at the first session of the LVI Congress the Joint 


OM War new garden from becoming of the greatest eS oe 
"tumeg . T. SWINGLE, of the U.S. Department of pomemeraer’ 
®t, Where 2) dia months’ travel in Europe, North Africa, and 

© has been Studying the agriculture and horticulture, with a 


288 BOTANICAL GAZETTE [ocroser 


view of introducing new agricultural industries into America. The journey 
was under the direction of the Section of Plant and Seed Introduction of the 


own southwestern states. Incidentally he noted many points of great biolog- 
ical interest. The caprification of the fig is still practiced as described 
Aristotle more than two thousand years ago, and a careful study of the com 
mensalism and symbiosis of the fig plant and Blastophaga is by no means 
superfluous, but on the contrary very much needed, since all previous students 
have studied it at the same time of the year, and many doubtful points remain 
to be cleared up. Mr. Swingle is going to California very shortly to study 
the fig industry of our own Pacific coast. | 


Proressor W. A. SETCHELL, Dr. W. L. Jepson, Mr. L. E. Hunt, and 
Mr. A. A. Lawson, of the University of California, have returned to Berkeley 
from a botanical expedition to Unalaska. Dr. Jepson studied the flowering 
plants, Professor Setchell and Mr. Lawson the flowerless : 
Hunt, who is in the Civil Engineering Department, determined altitu 
took the photographs of plant communities, etc. The party remained gt 
laska for eight weeks and carried out its work as planned, collecting 
oughly in the neighborhood of Unalaska bay, making extensive field 
and securing a fairly full collection of photographs. Professor Setchell 
Unalaska for about three weeks, on a trip to St. Michael and Cape 
collecting plants of all kinds and making notes as to points of 
and ecology. Returning, the party went from Unalaska to Sitka! 
coast, collecting at Unga, Karluk, Kodiak, Orca, Juneau, and 
were thus able to trace many plants of the shores along a conside 
of the Alaska coast, and to note the changes in habit and the @ 
altitudinal distribution. There is a very considerable amoun 
accumulated and it will not be known until it is carefully ba 
much of it is new, or just to what extent it will throw light 0 ‘ 
tribution. The collections of marine algz, taken in connection 
collections made in Alaska, Washington, California, and ‘Mexi 
last four or five years, it is hoped, will indicate the limits of the 
floras of the Pacific coast of North America when they ar 
mined and tabulated, and will afford the basis for some exact I 
causes of demarcation.—Sczence, Oct. 13, 1899- 


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Il NOVEMBER 1899 ~——=SNo. 5 


THE are 


EDITORS 
l. COULTER, 7ve University of Chicago, Chicago, il. es 
SHARLES R. BARNES, Zhe University of Chicago, Chicago, Ml. 25 


: ; J.C. ARTHUR, Purdue University, Lafayette, Ind. 


ASSOCIATE EDITORS 


{IR eo Le FRITZ NOL 
Geneva Crier Z Bonn 


| os VOLNEY M. {SPALD 
= go Padua Pceen - Michigan 
wi ROLAND. THAXTER 
: si “dl Berlin Harvard University 
WIGNARD WILLIAM TRELEASE a 
ole de Pharmacie, Paris Mi. aes — Garde 
po. ORPER H. MARSHALL W 
ersity of Wisconsin rr = Camir idge Re 
MURA EUGEN. wl ) 
og i Driers, sag ae of Copenhagen 


T WITTROCK 
~ Royal yaa of Sciences, Stockholm oe %, = e 


CHICAGO, ILLINOIS 
Mblishey by the University of Chicago 


be Aniversitp of Chicags Press 


PYRIGHT 1899 BY THE UNIVERSITY OF CHICAGO 


y 


Nem | : 4 . | 
' Botanical Gazette 
Monthly Fournal Lmbracing all 2 of Botanical Science 
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1 | No, 5 Issued November 30, 1899 
ele, 


CONTENTS 


XIC EFFECT OF DELETERIOUS AGENTS ON THE GERMINATION 
‘VELOPMENT OF CERTAIN FILAMENTOUS FUNGI. /. —. Clark - 


LOPMENT OF THE MICROSPORANGIUM AND MICROSPORES IN CON- 
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ANTS RECENTLY INTRODUCED INTO FLorIpA. MM. LZ. Fernald - : ie de 
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OTICES . : = . . i 5 u - 364 
p R STUDENTS ; : ON ey 
ig ‘ ; : : Y 6 < . 368 


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ot 


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CONTENTS: J 
Bibliography History Biographies and Portraits f 
f Herbals Early Botanical Science ue 
Linnaeus Handbooks Dictionaries ‘ 
HISTOR 1 Microscopy Morphology aad Physiology 4 
ie Y and s Botanical classification Nomenclature - 
7 SCENTIFIC Descriptive Botany Encyclopaedic works J 
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. Fossil Plants § Transactions of ocientitic a 
OTANy J Periodicals Applied Botany Botanical 1 Gardens 
| Omarig of 3000 f Agriculture and Horticulture to the end of the 18th century J 
1 ates under oo Gardening Ornamental Plants Food Plan af 
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e XXVIII NUMBER 5 


THE GERMINATION AND DEVELOPMENT 
CERTAIN FILAMENTOUS FUNGI: 
ter. CLARE ; 


INTRODUCTORY. 


E object of the writer in undertaking the investigation, 
sults of which are presented in this paper, was to deter- 


ent years the study of plant pathology has come to be 
the most important in the whole range of botanical 
It Seems desirable, therefore, that all possible light 
thrown upon the toxic properties of the various . 
in combating fungus pests. It is also very desir- 
1 a Scientific point of view, to throw all light possible 
toblem as to the element or group of elements to the 
‘Which the toxic properties of the compound are to 


to the Progress of modern physical chemistry, plant 
4re now enabled to make up solutions of all chemi- 


P sented to the Faculty of Cornell University for the degree Master 


289 


290 BOTANICAL GAZETTE [Novenser | 


cal agents having the same number of molecules present in 
equal volumes of the various solutions. This is a decided step 
in advance, inasmuch as it enables investigators to compare the — 
properties of a molecule of any substance with those of any 
other molecule, a comparison obviously impossible under the 
old method of making up solutions of a certain per cent. by 
weight. The vital error in making a comparison between per 
centage solutions is readily seen when we recall that a I per 
cent. solution of formaldehyde contains over eight times a& 
many molecules per cc. as a I per cent. solution of mercuric 
chloride. — 
These equi-molecular solutions are made by dissolving @ 
many grams of the compound as there are units in its molecular — 
weight in 1000 grams of the solvent. Such a solution is termed 5 
a ‘‘normal solution” of that compound, and it is represented col . 


veniently by the formula ~. Diluted to half this strength we 
get , or one half normal solution, and so on. Solutions 


stronger than normal may be made by using double or quadruple 
the number of grams specified to the 1000% of the solvent, 


giving le and S: respectively, of the substance. 
I I : 
as show! 


Still more recent chemical and physical research h a 
m 


that in the case of very many substances in solution, 
the molecules of the dissolved substance are no longer f 
as such, but have become divided into two or more parts. : 
part-molecules have been termed ions. To illustrate, gas 
pose that 36.37 grams of pure HCI have been added to 
cc. of water; we know that the HCI is no longer all pi 
whole molecules. It fact, we have every reason to 


Th ey 


about 80 per cent. of it has become éonized into H at 

+ a 
the H ions being charged with positive electricity, 
having a corresponding negative charge. The 109°" cf 


stances capable of ionization are similarly ce ; 


1899] TOXIC EFFECT OF DELETERIOUS AGENTS 291 


towards the cathode when a current of electricity is passed 
through the solution, while those charged with negative elec- 
tricity migrate in the opposite direction towards the anode. As 
we have already said, a solution of HCl is about 80 per cent. 
ionized. This percentage of ionization increases with each 
; a n 
ee of dilution, becoming practically complete at eo 
‘the case of HCl. Limited space forbids further discussion of 
this interesting and important phenomenon. For further infor- 
‘mation the reader is referred to the excellent papers by Drs. 
Kahlenberg and True (’96) and Mr. F. D. Heald (’96), or to 
the more recent text-books on physical chemistry. Suffice to 
Sayin this place that the ionization of the molecule has enabled 
chemists and physiologists to determine in many cases the toxic 
, dlement or group in poisonous compounds. We shall have occa- 
og to refer frequently to this matter in discussing the experi- 
‘Mental data presented in this paper. 

_ Ithas been the writer’s aim to supplement the work of Drs. 
/Mahlenberg and True, and Heald on phanerogams, and Drs. 
Kronig and Paul (’97) on bacteria, by applying the theory of 
— of the molecule to the study of the physiology of 
cides, 


METHODS. 

Selection of forms—In the selection of forms the following 
Were given particular consideration: (@) regularity Of 
ation, (6) ability to grow normally in liquid media, (c) 
fo fruit normally in a saturated atmosphere. After 
silentation with a large number of forms the following 
chosen as being well suited for the work: Aspergillus flavus 
nk, Sterigmatocystis nigra v. Tieghem, Edocephalum albr- 
accardo, and Penicillium glaucum Link. otrytis vulgaris 
S afterwards chosen as a fifth form, it being entirely satis- 
in regard to germination and mycelial development, and 
y desirable because of its semi-parasitic habit; it, how- 
iled to fruit in cell cultures. Pure cultures of these 


292 BOTANICAL GAZETTE [NOVEMBER 


molds were obtained and renewed from week to week. The 
spores used in inoculating the cultures in the experimental work 
were taken from fresh tubes in which the fungus had been grow- 
ing seven to fourteen days. A solid medium made by adding 
128" of agar to a liter of sugar beet infusion was found tobe 
very satisfactory for stock cultures. 

Selection of medium.—The object being to test the effect of 
the deleterious agents on the fungi under as nearly normal con- 
ditions of development as possible, the selection of a suitable 
medium was of primary importance. Many preliminary cultures 
were made with various media, including distilled water, infusions 
of potato, celery, sugar beet, prune, and bean (stems, pods, and 
mature seeds), besides various others compounded from inors 
ganic salts, sugars, asparagin, etc. The results in germination 
and development were very varied. In distilled water Sterig- 
matocystis and Penicillium failed to germinate in 24 hours at 
28° C., and of Botrytis, which did the best of the five forms, but 
40 per cent. germinated in that time. Mycelial development ® 
all was meager, and fruiting generally mil. Very minute Linge 
tities of the deleterious agents were found to inhibit germin® 
tion, but the death-point of the spore was found in the ae 
dichloracetic acid, potassium hydroxid, and cobaltous oe 4 
to be the same as in the medium finally selected. The ge a 
compounded from salts, sugars, etc., were more satisfactory » 
vegetable infusions, however, were superior to all others. ib 
infusion of sugar beet was ultimately chosen as The fe 
whole the medium best suited for the forms used. 4d above 
that, in the case of the three typical poison tione 
the concentration causing the death of the J age pre wa 4 
same, whether the agent was dissolved in distilled " 
beet infusion, is very important in that it shows that pied a 
which was used throughout the study, does not <e os 
change the toxic properties of these agents oe ye hs 
spores. The infusion of sugar beet was prepared Ke we 
450 grams of the root in a liter of water for 3 house ee of t¥O 
was then strained, cooled, and stirred “up with the vias : : 


hog] TOXIC EFFECT OF DELETERIOUS AGENTS 293 


| eggs, after which it was boiled, strained, filtered, and poured 
into flasks. After thorough sterilization the infusion was ready 
loruse. In order to get the greatest possible uniformity it was 
found desirable to make up four or five liters at atime. In this 
medium the spores of all the forms used germinated quite uni- 
_ formly in from 3 to 8 hours (according to the species), grew 
_fpidly, and fruited normally (except Botrytis) in from 18 to 
- hours at a temperature of 28°C. 
Method of culture —The van Tieghem hanging-drop culture 
: was found to be entirely satisfactory. The cylinder part, which 
_ Woof glass, had an internal diameter of 17.5™™, and a height of 
— to, This cell, with a capacity of over 2.5 cubic centimeters, 
mrovided an abundance of oxygen for the normal development of 
the fungi. The cylinders were cemented to the slip by means 
of beeswax, and the cell was completed by sealing the cover to 
the top of the cylinder by means of a ring of vaseline, applied 
ty inverting it on a glass slip covered by a thin layer of melted 
| Wseline. A small nick was made in this ring so that when the 
_ twas applied a minute opening might be left through which 
_ He expanding air could pass when the cultures were placed in 
og thermostat. An hour later they were carefully examined, 
such as had not already become hermetically sealed were 
made so by tapping with a pencil over the tiny opening 
“erred to above. This precaution prevents much trouble and 
When culturing with volatile substances. For convenience 
dling and €xamining under the microscope two cells were 
_ on each slip. The cells were permanently labeled by 
: ing lettered and numbered labels to the ends of the slips. 
Precaution against accidents duplicate cultures were not 
os on the same slip. , 


hg ae Cover. Four or five drops of the same solution were 
SM in the bottom of the cell. The spores of the fungus to 


294 BOTANICAL GAZETTE [NOVEMBER 


be tested were transferred from a pure culture to the hanging- . 
drop by means of a sterile platinum needle, the utmost care 
being taken to prevent the adherence of the spores in bunchesin 
making the inoculation. The cover bearing the culture was 
then inverted on the cell and gently pressed until completely 
closed, except for the minute opening already fully described. 
When a set of cultures was complete all were placed ina ther — 
mostat which was kept at a constant temperature of 28°C 

Care of cells, covers, pipettes, etc-—After completion of a 
series of cultures, all bottles, rods, etc., were thoroughly washed x 
and placed in running water for several hours, then dried and 
placed in a dry oven at 160°C. When the cultures in the cells 4 
matured, the covers were removed and the cells were thor 4 
oughly washed under the water tap, wiped, air-dried, and finally 4 
placed in the dry oven at 110°-120°C. for an hour. - 
insured thorough sterilization, and at the same time drove off the 
last trace of any volatile substance that might have escaped the 
washing. The covers were first boiled in strong KOH, then in 
several changes of water; this was followed by boiling in suas 
H,SO,+K,Cr,O,. They were again thoroughly rinsed : 
again boiled in four changes of clear water, rinsed in 95 set 
alcohol, wiped, and sterilized at 160°C. The pipettes : 
cleaned by forcing water through them for an hou 
to the water tap. They were then sterilized in a steat be 

An occasional culture was found to be contaminat 
bacteria, due no doubt to dust particles bearing spe 
in contact with the cultures in the making. Such co 


however, by bacteria or fungi amounted to less bp I ie 
f preliminary r 


Vapor pressures in the cell. 
cal chemists that every liquid has a certain vapor i. 3 vapor 
water at standard atmospheric pressure and 28 C. stance 
pressure of 28™" of mercury. Any addition fee 
substances to this water will lower its vapor Pree T se vapor 
substance added be hygroscopic the lowering Te 


=) TOXIC EFFECT OF DELETERIOUS AGENTS 295 


} pressure may be very great, as is the case with KOH and 
H,SO,. On the other hand, should the substance be quite vola- 
tile the increased vapor pressure of the substance added may 

_ more than counterbalance the lowering of the vapor pressure of 

the water, as is the case when ammonia or alcohol is mixed with 

water. To recapitulate in brief, the addition of any substance 

_ orsubstances to water gives a mixture with a vapor pressure at 

_ ‘ariance with that of pure water. This vapor pressure may be 

_ eater or less than that of pure water, depending on the physi- 

_ tal properties of the substance or substances added. Thus, a 

: hanging-drop containing KOH in a cell in which water has been 

; placed below (as is the usual method), absorbs moisture by rea- 

_ Son of its low vapor pressure. Indeed, there will be a constant 

_ distillation of water vapor from the water below to the culture 

. drop, until all the water has passed up or the drop, becoming too 

‘ large to “hang,” falls to the bottom of the cell. With alcohol 

the reverse takes place. No sooner is such a culture made up 

than the alcohol begins to distill from the hanging-drop, and it 
fas not, even for an hour, the concentration supposed to be 

Present. Such a culture as a test of toxicity is valueless. 

Were it possible to have all hanging-drops of exactly the 

~ ‘i, and exactly the same quantity of water below, we 

should expect uniformity in results. Such uniformity, however, 

Would be useless, perhaps worse than useless, as it might, pre- 

attention being’ called to the fundamental error of the 


s n 
found that of four cultures of Macrosporium in a Srohee 
wlan 

pe. Metiatic chlorid, three grew and one failed; of 


Cultures 7? 
409600 


five grew and three failed; of ten cul- 


| 204800 four grew and six failed; of four cultures in 


296 BOTANICAL GAZETTE [NOVEMBER 


n “ : ; ; 
——— one grew and three failed! Witha strictly non-volatile 
102400 


substance we should expect less variation, because with less dif- 
ference between the vapor pressures of the hanging-drop and 
that of the water below, the changes in the concentration of the 
deleterious agent in the hanging-drop would necessarily take 
place more slowly. Of the volatile properties of HgCl, we 
shall speak later. 

In looking for a method that would meet every requirement 
of the case, many preliminary tests. were made with all five 
molds and with different deleterious agents. Two such tests 
with potassium cyanid are here presented in detail. It is of 
interest to note that while KCN is not in itself a volatile com- 
- pound, in aqueous solution more or less hydrolysis takes place, 
resulting in the formation of a corresponding amount of HCN 
(Shields ’93), which is quite volatile. Hence, aqueous solu 
tions of KCN are in their behavior quite typical of volatile com- 
pounds, 

Column 1 gives the culture label by means of which the ie 
tures were identified; column 2 the concentrations of KC} 
in the hanging-drops of the various cultures; column ; me 
solution—if any—used in the bottom of the cell. “Dry 
implies that no water or other solution was placed in the ee? 
‘of the cell. Under the head “germination” the percentage 
spores germinated is given for three observations, at 12, ee 
36 hours, respectively. Under “development”’ the lenge 
germ tubes of spores showing an average development Is 8 
in micromillimeters. in the 

These data not only show that cultures having ae 
bottoms of the cells are unreliable and vary acco ae 
amount of water present, but that “dry” cells oe cult 2 

of the , 
s but one fourth eo 
” former, how” 
of the KCN present in cultures labeled ‘‘A9, the he hanging” 
ever, had several drops of the solution from which ' : d 
drop was made placed in the bottom of the cell wens 
customary drop of water. The striking difference 1? 


results : 


ate 


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NOM -+ NOISQUNI 1am NI SQU71IONaasV 


[NOVEMBER 


BOTANICAL GAZETTE 


298 


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NOM + NOISOANI LOA NI WAITTIOINGd 


1899] TOXIC EFFECT OF DELETERIOUS AGENTS 299 


needs no further comment. The results with. Penicillium and 
other forms abundantly confirmed the test, and lead to the con- 
clusion that when water is used at the bottom of cells containing 


vapor pressures are being tested, erratic germinations are to be expected, 
and in the case of a test of any substance —particularly those of high 
w low vapor pressure —the toxic properties shown will be less than a 
correct method would indicate. 
In the regular work reported in this paper, a few drops of the 
same solution as that used in the hanging-drop of any particular 
cell was in every case placed in the bottom of that cell, thus 
establishing complete equilibrium of vapor pressures in the cells, 
and thereby preventing changes in the concentration of the solu- 
tion: under test as must take place when this precaution is not 
observed, By this method it was found quite feasible to make 
hanging-drop cultures of any composition: whatever, ranging 
from ‘— (70 per cent.) solution of alcohol to a a (21 per 
‘nt. solution of potassium hydroxid. 
Stock. solutions of chemicals.—These were prepared in all cases 
; by a responsible chemist from the purest chemicals obtainable. 
It was found desirable to have stock solutions in highly con- 
. *entrated form so that all necessary diluting could be made by 
_-Wding beet infusion, The stock solution of HCl, for example, 
Sntained 25 grams pure HCl per too®. HCI contains 3.58 
oe cent. HC] by weight. Such a solution was gotten by taking 
: ¥ of the stock solution and diluting, by adding beet infusion, to 
a. 2 Was gotten by diluting 10° to 20°, and so on. 
Chemical a 


nal and y egular cultures—Inasmuch as the work was in most 


hanging-drop cultures and solutions of substances having very high | 


300 BOTANICAL GAZETTE [NOVEMBER 


various chemicals to this group of plants; for it was of course 
early learned that the data worked out for phanerogams by 
Kahlenberg and True (’96) and Heald (’96) were a contrast rather 
than a comparison when placed beside the facts learned by 
experiment from the fungi. Heald (’96) called attention to this 
fact on finding a fungus growing vigorously in a solution of HC 
that had killed the root of Pisum. The data obtained from the 
trial cultures proved invaluable in making up the regular cultures, 
as it was then possible in most cases to make up a seriesof 
twelve dilutions in no. 1 of which the spores would certainly be 
killed, while in no. 12 the fungus would be practically unharmed. 

In the regular work all cultures were made up in duplicate, 
including duplicate checks in pure beet infusion. Important 
points were quite frequently checked over in duplicate and some- 
times in quadruplicate. This was also done in the case of unés- 
pected developments; for instance, the writer was surprised 10 
find the spores of Penicillium showing so great a specific resist- 
ance to acetic acid. Repeated checking, however, proved the 
correctness of the first observation. 

One of the most marked features of the entire work was the 
regularity in results. It is true that the cultures nearest the 
inhibiting point in a few cases varied to the extent that gee 
nation of a number of spores took place in one culture wil 
the duplicate failed. Such were not considered evraitt. 8 
however, both cultures of a certain strength grew and one ee 
weaker concentration failed, the latter would be ern 
erratic. Such erratic cultures, however, did not exceed ae 
in a work requiring upwards of forty-five hundred pene 
tures aside from preliminary work. Such as did ee st 
doubtless due to oversight in cleaning the cells or other a¢ cay 

Examination of cultures, noting results, cc inter . 
made up in the early part of the day, and were examine ‘oui | | 
vals of from three to six hours for the fourteen hours “ spores be 
and at longer intervals until the fungus had matured, or cc 
in the cultures which had failed to germinate were fra tage f 
pure beet infusion to test their vitality. The perce = 


1899] TOXIC EFFECT OF DELETERIOUS AGENTS 301 


spores germinated in the various cultures was noted on two 
occasions, first when from 30 per cent. to 70 per cent. of those 
in the checks had germinated, and again a few hours later. A 
similar method was adopted to indicate the early mycelial devel- 
opment. At some time after germination the germ tubes show- 
ing an average development were measured and noted; this was 
repeated a few hours later, and sometimes a third measurement 
was made. The first appearance of conidia was also noted, but 
a this very frequently occurred at night (between 15 and 22 
hours after inoculation) the point was not so well noted. 
In the earlier work, the cultures in which the spores failed to \ 
seminate were opened at 72 hours and a number of the spores | 
were transferred by means of a sterile platinum needle to a | 
hanging-drop of pure beet infusion in a clean cell, in order to 
test whether they were killed by the agent or merely inhibited. | 
It per cent. or more survived, the culture was classed as 
Wed; if none at all or less than 1 per cent. survived, they’ 
Nae classed as filed. In most of the work, however, the trans- 
fers were made at 48 hours, it being found that all spores that 
aay germinate did so in less than 36 hours and usually much 


The cultures which germinated in the presence of the dele- 
a terious agent were likewise divided into two classes: (1) those 
; Which although they may have been retarded or stimulated in 
4 mycelial development by the agent, finally matured a fair crop 
|) dia in about the norma! time; (2) those, which, although 
, 7 8etminated and continued to grow, presented a markedly 
. ay °F retarded mycelial development, and generally failed 
“ww, Detween these two classes came—as might be sup- 
7 a tumber of cultures which were very difficult to classify. 
a, Cases there would be an apparently normal mycelial 
7 Cince a. but almost total supression of fruiting. In other 
| on "regular, meager, and even yeast-like mycelium would 
Surprise by finally developing a number of apparently 


Cultures ; hours. 
i aa M alcohol, formaldeh H,O KCN were transferred at 72 hours, 
ye : others at 48 hours, e yde, 22a) and Ww 


302 BOTANICAL GAZETTE [NOVEMBER 


normal fruits. In general, a fungus was said to be “injured” 
by that concentration of the deleterious agent which prevented 
its classification in class I as above. 

In discriminating between class III (inhibited spores) and 
class IV (killed spores) attention is again called to the impor 
tance of avoiding dunches of spores in making the inoculation. 
In many solutions such bunches— doubtless containing air— 
float on the surface of the drop and fail to receive the full 
influence of the agent. When spores from such a culture are 
transferred to pure beet infusion to test their vitality the bunch 
may be broken up in the process and the spores germinate — 
readily while all others are dead. Much can be done with care 
in making the inoculation, but at best it is a serious source of 
error, and it has been on this account, and on account of the 
impossibility of transferring the spores without taking with 
them small quantities of the agent, that the inhibiting concentra- 
tion rather than that causing death has been adopted as the chief 
critical point in discussing the experimental data. us 

To illustrate what has been said regarding t 
notes on cultures, a typical left-hand page of the cu 
book is here reproduced. This will also aid the reade 
stand the classification of the experimental data 
The opposite page was always reserved for more exten a 
on points observed from time to time in the progress Of © 
experiment. 

Sources of error—Before proceeding t 
experimental data it might be well to mention 
of error observed during the preliminary stu 
_ against in the progress of the work. 

1. Xylonite cells were found to hav 
on some fungi when used for hanging-drop cu 

2. Bacterial contamination. oS 

3. Lack of equilibrium in vapor pressures in the cells. : ef 

: details of itl 


he taking of 
Iture note- 


r to under- 


oa discussion of the 
briefly the source 
dy and | 


e an injurious jabnent 
Itures at 28 C. : 


4. Deterioration of stock solutions (see 
ment with KCN). 
5. Use of impure vaseline for sealing cells. 


TOXIC EFFECT OF DELETERIOUS AGENTS 303 


1899) 


z 
” ee ” ae ZIV 105% 
” + 9» mi ” = 11V 66bz ~ 
= 
[\pateq|) say] gh ye} possay|suesy, | 8 oly Lovz 
[‘azour| Auvur (say bz |S unra8 |yS ‘say |z1 ‘say| gh ye] paszroy|suvry ] 2 6V S6bz = 
A[uo : 
sum mat | ze | gr | zz z get 4 zz ne Vv €6bz ot 
‘say ZZ ye .. 
PeiN7z [TPM gr OLI zz oor ad I et ~ LV 16bz g : 
QzI a 
gf on zz 09 zI oor A os 8 os OV 6ghz cy 
gz 
lz 06 ee oLI ZI Sz 9 ool | g cL 9 = SV Lgbz 
zis 
Lz or Ez OgI a of 9 0g 9 = by Sgbz 
zz oo zz 00z ZI of 9 Sg 9 — tv Egbz = 
aoeds oars ri 
0} pa}jiuo gboz 
gayediidap ya oo a4 O1z ZI of 9 0g 9 _ ZV 1ghz 
JO pl00ax] zz co Zz 00z zI of 9 $8 9 FQsoD— IV 6Lbz 
| a | *say a ‘say | a "say % | "sry | w “say 
mon | th es BR | FG f= 
dojaaap [ero uonvuruds 
"OD _S% *eanjvssduray, “deat ur 


6681 ‘12 ‘w 6 yw paanyny 


Lauda NI sa TTIDNaasV 


304 BOTANICAL GAZETTE [Novewmn 


6. Use of spores of uncertain age and vitality. 
7. Use of culture media unsuited for the normal development 
of the fungi tested. 
8. Imperfect sealing of the cells, due generally to the raising 
of the cover by expansion of contained air when the cultures 
were placed in the thermostat. 
9. Transference of bunches of spores in making inoculations. 


EXPERIMENTAL DATA AND DISCUSSION. 


Details regarding the critical points (¢. ¢., concentrations 
causing injury, inhibition, and death) of the various fungi in the 
more important of the thirty-seven compounds tested may be 
found by referring to the diagrams prepared to accompany this 
paper. This device, primarily intended to conserve space by 
giving.in a condensed form the various specific resistances of 
the different molds will also be useful, it is hoped, in conveying 
to the reader, by means of the eye, a general impression as to 
the relative toxic properties of the agents tested. It should be 
distinctly borne in mind in consulting these diagrams that each 
vertical line represents a doubling of the number of molecules 
present in the solutions in passing toward the right, the vere 
space between two vertical lines representing one concentratio?: 
The relative average toxic properties of the more poisonous 
agents are further graphically depicted by means of he 
charts in the final installment of this paper. Ee 

ACIDS. oe 

In the case of acids, diagrams have been prepared: sO 
the resistances of the individual molds in the eight acice 
diagrams will be found to accompany the diagrams a 
various acids on pages 307-308. In these diagrams a ee 
been made to emphasize the fact that the aie ead of 
concentration in passing to the right, by placing at oe ete 
the columns the proportions of molecules present in Se ore 
tion in terms of x; x being in every case the number an 

“ _ solution. Thus a normal S00" 
262144 = 


cules present in a 


)| 


TOXIC EFFECT OF DELETERIOUS AGENTS 305 


c 2H . 
always contains 262144% molecules, ng contains 5242882 mole- 


cules, and so on. 


Hydrochloric acid, HCl; 70, 230, 614. This acid, on account | 
of its very high ionization at the critical points, and because of | 


iSvery wide use in physiological and chemical investigations 


by other workers, has been taken as a standard by which we shall | 


compare the others. In making these comparisons, HCl, being 
‘ the most highly ionized of all acids, is assumed to represent the 
_Yalue of ionic H. This is a purely arbitrary assumption, and 
the reader will bear in mind that when we speak of ‘the toxic 
Value of ionic H” we mean the nearest approach we could make 
| to determining its toxic value for the molds, viz., that of 91% 
| tinic H +-9% HCL. 
[The following “ coefficients’? have been worked out for 
: HC with these five molds: coefficient of injury, 70; coefficient 
. of inhibition, 230; coefficient of death-point, 614. 


These coefficients mean that on an average for the five forms 


ied 2. 


2048 of a normal solution caused distinct injury to the 


| taltures, 23° 


614 
2048 


tion of the Spores, and 


pew of 2—has been used throughout this paper in determining 
“ents. As a matter of convenience the numerator only 
expressed in discussing the three critical points of the 
'S agents, giving as it does at a glance the correct relative 
: J of the agents. The absolute value of any coefficient 
(in cither mold or agent) be determined by simply supply- 
‘Omitted denominator, 2048, the resulting fraction being 
“TY Case that proportion of a normal solution. In dis- 
Ng the 
oe . 


2048 Of a normal solution inhibited the germina- 


of a normal solution killed the 
‘Pores, The denominator of the fractions, 2048—the eleventh | 


a 


306 BOTANICAL GAZETTE [ NOVEMBER 


From time to time attention will be called to the wonderful 
resistance of the fungi to many deleterious agents as compared 
with that of the higher plants. The comparison (or contrast ’) 
in resistance to ionic H is as 800: 1, the average death-point for 
the five species of molds being re HCI, while that of three spe- 
cies of phanerogams is ar HCl. (Heald, ’96, p. 152.) 

On the whole, HCI was at the same time the most completely 
ionized and the least toxic of the acids tested. Sterigmatocystis. 
proved most resistant, for although germination and early myce- 

n n seal | 
— was required 


lial development were distinctly retarded by ery 


to inhibit all the spores, and - for forty-eight hours to kill 


them. Cultures in =, although much retarded in early develop- 
the checks, and 
unt of mycelium 
first noticed in 


ment, had at forty-eight hours quite overtaken 
at seventy-two hours far surpassed them in amo 
produced. Distinct retardation of fruiting was 
o mature its fruit 


hd a : ‘ 
can) Ge required nearly double the time t 


EXPLANATION OF DIAGRAMS. oe 

The initials A., S., C., B., P. stand for the respective i ie 
names of the fungi used. fa 

The fraction of a normal solution placed at the head 9 " 

refers to the space between the vertical lines over which pe 
this space is depicted the result of culturing the various 1 

concentration of the different agents by the symbols de oe 

ot Two lines indicate normal or almost normal develop 


—— Three lines indicate distinct injury- 


Four lines indicate very great injury. 
mination. — 


HM Alternate blocks indicate total inhibition of 8& ee 

ed for vitality 
mum Solid black indicates death of spore when tested S33 
48 hours. 


1899] TOXIC EFFECT OF DELETERIOUS AGENTS 


(Edocephalum ___Sterigmatocystis Aspergillus 
> 2 e q fh: oer es 
em ey Wy an a = tre it Pz = 
feos 2 Te okehies mes op He FSH 
yes 2H OQ site atta vines le ea os 08 0 URe em 
“50 on 4-5 ZOOL Ae Sa “8o8 
| | | re 
| | | a 
a 16% bf 
a | 32x 
! 64x 
Tn Lt | 
25627 o14 
Hanae Lt | 
LT iL | 5122 ve 
| | 1024.t 
a 2048.7 L 
Bene 
ie 4 a nents 
ae a 8192.7 : 
ae a 8 16384-r 
=a 32768 * 
HHH % 655367 
Py [cs es - = & = 
= w1072-r 
SHH ian 
saa 2b62r44r 
Sa — x. — | 
= & 524288.r 
os -— 
= a 
1048576.r 
‘ 20971524 


DIAGRAM I 


BOTANICAL GAZETTE 
Penicillium = Botrytis 
Xx x 
rz et iP 2 mrt 
oO ™~ ew Sb oO TPR e SE Py 
. eo gem C) @ 9) Wee Mee 
is a — ee QO > ae « i ©) 
& 5 Se mee Seo 
Wx 
1x 
or 
qr 
8x 
16x 
}— 4-4 +4 
32% 
mT 
64x 
128r 
++ 
2562 
5r2t f 
10247 : 
2048.5 
Sinise be 
4096" 
dl 81922 
163845 ¥ 
Seanez 3a70ar 4 
65536 
1310724 
| _-—_-—— 
2621447 
Bee 524288." 
| o @ @ 8 3 ee 
10485764 
20971522 


DIAGRAM II 


[ NOVEMEER 


4 H,S0, 


tae) a " 


ale Sia Le 


els #la ols 


“|e 


DIAGRAM III 


310 BOTANICAL GAZETTE [NOVEMBES 


8 eee eee 
n 
2048 
BS Bie ieee eee 
1024 

Te 
* 
512 
ee oT TT 
as6 
Bini Reels ereeeeie. 
128 
ee eee Slee eee ” 


it 


Sis 


| 2 2 ee — —€ i Md 
16 
Lh 
| 2 @ 8 2 3 —e a ae bd 
|i gee eesti 
: 4 
= 
2... ee ee ee se ud 
me 
2 
a 


Ty UPD pie 


DIAGRAM IV : 


K,CrO, VT2V 7 
seo Bur> wre Bude wsoturd vofn> woud 
tT | 1] 
| MAE TTT TTT TET 
H ++ ima ae | 
aH HiT rT 
rtd [ti | magiy 
| | | 
HT | | 
| 
} | 
a 
i 
HA 
a P35 
LL = SS eee 
Lia aa + 
oo i OS Re 


DIAGRAM V 


ale S18 


olR 


S|2 le 


[NOVEMBER 


312 BOTANICAL GAZETTE 
Cu(NO 3). CuSO, FeSO, CoSO, NeSO, 
webu teun> we Bunr> we > 


- 


SSS 


DIAGRAM VI 


Bl* Gla 


ela sla ola Gia Sia Lia 


ele 


313 


DIAGRAM VII 


1899] TOXIC EFFECT OF DELETERIOUS AGENTS 
Strych. 8,0, Sod. sal. ZnSO, KI 
sxSur wae Sa wok n> yo fu > a: > 
n 
524288 
: n 
ll an 
n 
| | 131072 
n 
| | 65536 
n 
lll 32768 
n 
16384 
n 
8192 
TTT n 
4096 
2 Ee ied 3 
2048 
ain n 
1024 
| Bias see = re 
: 512 
Sia Ht 
256 
Simin ase HT] 
BP 3s 128 
———_— in — 7 n 
ef ala 64 
Sine | le a Lh eae 
8 sf 32 
tt ct GG BE Eee rT 2 
ls S| 16 
a ——J co ee ee et n 
| *p d ele 8 
a = ee i ee n 
4 
2 BN ie Bee's 
e@ il ae bil 
eS 2 
2 fe rs a ee in 
aR 
= th a 
nd a a Se 
e & i 
es | a aae. i 
& = ‘sain 
~ 3 
bo tn I 
RE a5 BH is 
| a aT 


314 


BOTANICAL GAZETTE 


_Trichlor. Dichlor. 


y 


> sit km > ~~ 


“* 


i | 
i 
. 
_— 


4 


DIAGRAM VIII 


[ NOVEMBER 


899] TOXIC EFFECT OF DELETERIOUS AGENTS 315 


- compared with the checks ; while < the limiting culture, failed 


to fruit in six days. 

With HCl a sixth form, Rhizopus nigricans, was tested. Like 
its near relatives the mucors, it makes very unsatisfactory growth 
ina liquid medium, and was therefore discarded. Its resistance 
to HCl proved to be practically the same as that of Gidocepha- 
um, The limiting culture of Aspergillus, = , developed a 
 tumber of abnormal conidiophores, each bearing 1-4 sterig- 
mata, on the sixth day. 

In the case of HCl, as in the case of all other acids tested, 
 mattempt was made to determine the relative toxic value of the 
anion. The potassium salt was used. This salt becomes highly ' 


| lonized into K and Cl ions in moderately concentrated solutions. 
Ina : KCI solution 84 per cent. of the salt is thus ionized. In 
. 20a concentration of HCl 89 per cent. is ionized ~ H 
—{dClions. (Kohlrausch, ’85.) In the former we have K and 


(lions and some KCl; in the latter, H and Cl ions and some 
| HCL It is evident therefore that the essential difference 


+ 
; tetween these solutions is the replacing of H in the latter by K, 
am the HCl by KCl in the un-ionized portion. The significant 
Pomt is that the concentration of Cl ions is practically the same 


i : both. The : HCI solution is fatal to the spores of Aspergil- 


: *S while © KCI solution has practically no injurious influence. 
: ay this fungus germinates, grows, and fruits normally in a! 
a ei of double this concentration of KCl. Hence we ee 
i as Cl ion is relatively harmless to this mold. Similar tests | 
' 2a its low toxic value to all the molds used. 
: ES lute toxic value of ionic Cl, and other weakly see 
New > however, very difficult to determine. ae this ee 
a es ould be remembered that the potassium sips a 
4 inn Sie are, although highly, far from being a “/ 
; at the concentrations permitting germination of mo 


316 BOTANICAL GAZETTE [NOVEMBER 


spores. Then, too, we are, perhaps, not justified in assuming 


that ionic K is entirely non-toxic; for although potassium is a 
necessary food element for all plants, may it not be that great 


concentrations of even so good a thing as ionic K may be bad 
for fungi? Later, this will be shown to be the case with iron, 
an element which, while absolutely necessary in small quantities 
for all plants (Molisch, ’94), is quite toxic in excess for both 
fungi and higher plants. 

Experiments with Aspergillus and CEdocephalum in KG 
solutions, however, enable us to say quite positively that for 
these forms the anion Cl has at most /ess than one thirty-second 


the toxic value of ionic H, and may therefore be disregarded in 
a discussion of the toxic properties of HCl. 

Nitric acid, HNO,; 48, r4z, 384. Ionized in almost the 
_ same proportion as HCl (Kohlrausch, ’85), HNO, proved much 


more toxic. Inasmuch as the concentration of H ions is pla 

' . xT . * “| n- 
tically the same as in HCl, and the NO, ion 1s practically vt 
toxic, having a toxic value of less than one thirty-second tha 


oo oe ‘ ie of 
ionic H, we must look for an explanation in the toxic valu 


the un-ionized molecule, HNO,. This was found to be approx 


ae 
mately 7.7 times that of ionic H?. In other word gree 
of HNO, loses nearly seven eighths of its toxic properties 
: iy 1? ) found that 
molds on becoming ionized. Krénig and Paul (’97 
‘ hours 
anthrax spores immersed in a solution of HNO, for two 
were entirely destroyed. A similar immersion A ee 
preparation of spores in the same concentration of H Ce 
the survival of 385 colonies. Preparations of spores - 


in te solutions of these acids, however, showed far less 2 
nae 1 distinctly mo 
in toxic properties, although the HNO, was still di + the Jat 
toxic. This was evidently due to the fact ne 7 | ioniteds 
concentration the acids were both much more highly eae 


?See table I, p. 325. 


7) TOXIC EFFECT OF DELETERIOUS AGENTS 317 


hence both more nearly approximated the toxic value of ionic 


H. This toxic power of the un-ionized molecule these workers 
termed its “specific poisonous effect.” 

According to this reasoning, then, were HCl and HNO, both 
completely ionized we should expect to find them equally toxic. 
_ This matter has been fully tested by Kahlenberg and True (’96). 


They found that toward Lupinus which is killed by a —*_ con- 
3200 


centration of ionic H, they and all other completely ionized 
acids with non-toxic anions had exactly the same toxic value. 
The influence of HNO, on germination was much the same 


nH ' ° 
a that of HCl, ead causing distinct retardation of both germi- 


_ tation and early growth. Later, however, a marked difference 
the appearance of the cultures manifested itself. Cultures 
‘ontaining the HNO, in = to Pe concentration produced in _ 
_ ‘Marly every case a much heavier mycelium than in the corre- 
ponding cultures in HCI. Fruiting was retarded, but not so 
 ‘Seatly retarded as was usually the case where mycelial develop- 
“Ment Was so Strongly stimulated. 

oo The stimulation of mycelial development was possibly due to 
the non-toxic nitrogenous NO, ion. Cultures of Aspergillus and 
Eocephalum in = solutions of KNO,, however, did not estab- 
ih this View, as they did not greatly differ from those in similar 
ahaa of KCl and K,SO,, although the concentration 
__$ 0s would in this case be some forty times as great as in 
“Cultures. of HNO, showing the most stimulation. A more 
_) Pfoposition is that it was due to the same factor as the 
‘the : @ toxicity, viz., the un-ionized molecules. The fact that 
«tal appearance of the cultures resembled that of cultures 
«ted by the oxidizing poisons considered later would suggest 
7 Ff the nitric acid molecule as being the active influ- 
th The fact that fruiting was not greatly retarded, consider 
— mycelial development, is in harmony with this 
n 


318 BOTANICAL GAZETTE [NOVEMBER 

Sulfuric acid, % H,SO,; 61, 205,589. As will be observed, 
of this agent is based on the half molecule in order to make it 
strictly comparable with the other acids, the others being all 
monobasic. 

aE <n 
H,SO, becomes ionized first into H and HSO,, ions, but as 
fe - 

dilution increases the HSO, ion further breaks up into H and 
~SO,. The ionization of H,SO, at the average inhibiting point is 
about 62 per cent. only (Kohlrausch ’85). Each 100 molecules, 
then at this concentration breaks up into approximately 124 H, 
76 HSO, and 24 SO, ions. This solution having a greater toxic 


value than a similar concentration of ionic Hi and the anion being 
practically non-toxic, the excess of toxic properties must be due 
to the partially ionized group HSO,. By referring to table I it 
will be seen that the toxic value if this anion is approximately 


1.3 in terms of ionic H. 
In this as in the other mineral acids, Sterigmatocystis proved 


the most resistant, - being necessary to kill. Botrytis was the 
most easily killed, ~ being fatal. CEdocephalum, although requit- 


ing concentration to kill the spores, was considerably injured 


By so greatly injured by S , and produced a very light yeast- 
like mycelium in Sn which on the third day practically ceased 


growing. On the whole, H 250, retarded germination less ah 


HCl and HNO,. ac 
Acetic acid, ‘CH ,COOH; 25.6, 83, 314. This acid at 


inhibiting point, - — is but 2 per cent. ionized (Koh '85): 


t as 

The toxic value of oF anion was found to be about qs 1, H, ne < 
so small a proportion of the acid is ionized the influence ° 

his acid are 


anion may be disregarded. The toxic properties of t 


199] TOXIC EFFECT OF DELETERIOUS AGENTS 319 


therefore to be attributed almost wholly to the un-ionized mole- 


a 
ail, CH, COOH, which proves to have a toxic value of 2.8.H. 
Penicillium showed a marked specific resistance to this acid, 


requiring ; for 48 hours to kill. This observation was accepted 


‘mily after repeated trials. Sterigmatocystis, so resistant to the 


16 
Dotrytis was particularly susceptible, being killed by ee which 


mineral acids, succumbed to —, and - inhibited germination. 


8 but one eighth the strength of HCl required for the same 
result, Although so much more fatal to Sterigmatocystis and 
Botrytis, germination and early mycelial development was much 


less retarded than with the mineral acids; so retarded the 


mination of CEdocephalum, but in the other forms this con- 
“aration had little effect. 

That acetic acid should prove so much more toxic to fungi than 
fe mineral acids was not anticipated (Migula’g90). Heald (96) 
und that it had but one eighth the toxic value of i on Zea, and 
"te fourth on Pisum. Kahlenberg and True (’96) found about the 
“me relation with Lupinus. The great variation in protoplasmic 
Stance to this acid is well shown by the following data: the 


Mar eel, Rhabditis aceti, thrives in a — solution, which is the 
. 2 
os ‘oncentration for Penicillium. Aspergillus spores are killed | 


qn 

4 j) those of Sterigmatocystis and CEdocephalum by “zs and 
ee of Botrytis by ~, jis fatal to Zea, and _” to Pisum. 
. 6 200 1600 

— a interest to note in this connection that not only ge 
la to be expected between different penne 2 
Same nce to deleterious agents, but different indivi png 
St pow, “pg =a even of the same species may have very ‘ ae 
irvamect, resistance, depending largely no doubt on Pr® itive 
Ned; - Pfeffer (95) grew Aspergillus on a nutri 
ining 8 per cent. dextrose and 1 per cent. acetic 


oo t 
) “td found that the fungus assimilated a far larger — 


320 BOTANICAL GAZETTE [NOVEMBER 


of acid than the dextrose. It will be noted that ; (0.7 per cent.) 


acetic acid proved fatal to the spores of Aspergillus used in this 
study, and in a 0.17 per cent. solution less than 1 per cent. ger- 
minated. 

The chloracetic acids—These acids are formed by the replace- 
ment of one, two,and three atoms of H, respectively, in the 
acetic acid radical by the element Cl; thus — 


pa th Li of 
S-G.00H = =-C-COH o-C-COH  GsGgue 
| | é 
CI Cl 


Acetic acid Monochloracetic acid Dichloracetic acid Trichloracetic acid 


It is a rule that the halogen substitution-products of carbon 
compounds have a toxic value which bears a close relation to the 
number of H atoms in the organic radical or hydrocarbon which 
have been replaced by the halogen. To quote from Davenport 
('97): ‘ Beginning with methane, CH,, we find this substance 
—marsh gas—innocuous when mingled with air. As the 
atoms become replaced by one or more Cl atoms, the poisonous 
properties increase,— 

CH, Cl is slightly anesthetic, 

CHCl1, = chloroform, 

CCl, is very dangerous, stupefying involuntary m 
Many such examples might be quoted, establishing this seo : 
was a matter, then, of great surprise to find what, at first sight, 
seemed to be a direct exception to this rule in the action of the 
chloracetic acids on the mold fungi. Their critical points wer’ 
determined as follows : 


uscles.” 


(Acetic acid - - - - 25.6 83 314) 
Monochloracetic acid - - 8.8 58 64 
Dichloracetic acid - - = 10.4 4 64 
Trichloracetic acid - - 37 sa oe 

lutions 


As soon as the experiment was complete the stock © d the 
used were placed in the hands of a chemist who aa 
potassium salt of each by just neutralizing with KOH. + 

fo 
salts are all quite highly ionized, the cathion being go a 
the anion the acid radicals of the respective acids. The 


1899] TOXIC EFFECT OF DELETERIOUS AGENTS 321 


values of these anions, as nearly as could be determined with 

Aspergillus and GZdocephalum, were as follows: 
Monochloracetic - - - —CH,CICOO, x 
Dichloracetic - - - ._ CHC], Coo; aT 

| Trichloracetic - 3 . ~ Sa os ar 


it s+ e+ 


Here the toxic values, although not high, are in the expected 
order, 
Let us next glance at the toxic properties of these acids in a 
practically completely ionized condition. Kahlenberg and True 
(96) found that with Lupinus 
6400 ‘ 
sowth. The same concentration of dichloracetic was evidently 
im the line, one plant being killed by it while the check survived. 


monochloracetic permitted 


q 
fygo ttichloracetic proved fatal. Ionization being practically com- 
Hee at this great dilution, we have to deal with ionic i and the 
Mons only. The toxic values of such solutions should be equal 
‘Uthat of HCl+ the value of the anion. This is exactly what 
ws find, Monochloracetic with its non-toxic anion has the same 
oa Value as HCl for Lupinus. The others are somewhat more 
‘ne: due doubtless to the influence of the anions, which are 
/ Pparently telatively more toxic towards the higher plants ge 
ards the molds, 

At the concentrations at which they are effective towards the 
°adadae we have an entirely different condition as reget 
_ ~~ ®8, as may be seen from the following: 

Monochloracetic inhibits at % and is 20 per cent. ionized. 


35 
Dichloracetic ‘i ” 


: “ 70 “c “ 
: Trichloracetic « a « ggg « “ 
2 Thi i : 22.8 i ; 
tion in the ionization {Ostwald ’89) is the key e 
* Planation of this apparent deviation from the rule referre 
above, 
“Stinet 
ee changes in the chemical and physical properties 
oniz A great increase in the toxic properties 
ed molecules, (2) A great increase in the 10nl? 


© introduction of Cl into the acetic radical np 2 a 
of tne 


of the 
ation of 


322 BOTANICAL GAZETTE [ NOVEMBER 


the acid in aqueous solution of given concentration. That the 
great ionization often masks the effect of the increase in the 
toxic properties of the un-ionized part is easily understood when 
we recall that the toxic value of the ionized portion of an 


acid is not greater than the toxic value of the ionic H + the 
toxic value of the anion. 
Putting all together, we get in brief the following: (1) The 
replacement of one H in the acetic radical by Cl doubles the toxic 
properties of the un-ionized portion, and increases the ionization 


in a ~ solution from 2 per cent. to 20 per cent.; the resultant 
of these two factors is an increase in the toxic value of about 40 
per cent. in a ” concentration. (2) The replacement of two 
atoms of H by Cl more than trebles the toxicity of the un-ionized 
molecule and causes ionization to advance from 2 per cent. t0 


70 per cent. ina oa solution. This gives a net increase of about 


30 per cent. in toxic properties at this concentration. (3) a 
all three H atoms are thus replaced by Cl the un-ionized mo g 
cule has a toxic value of over five times its original value an 


ionization advances from 2 per cent. to 88.8 per cent. in a 558 


i ; : joniza- 
solution. At this concentration the effect of the greater ion : 
tion is more than can be made up by the increased toxic i 
ties of the 11.2 per cent. remaining in molecular form, hen 


4 : ; n ion as 
fall in the toxic properties of this acid at [3 concentratl 


compared with the original acetic. t in 

The values of the un-ionized molecules as worked i an 
table I, p. 325, show that these acids are very far from — 
exception to the rule that the toxic properties of ee 
increase with the introduction of Cl into the organi sa 
These values are as follows: 


+ 
Acetic acid, 2.8 times the value of ionic H. 
Monochloracetic acid, 4.7 “ ““ “ 
Dichloracetic acid, 9.5 “ “ " 


Trichloracetic acid, 14.1 r 


1899] TOXIC EFFECT OF DELETERIOUS AGENTS 323 


The addition of any highly ionized acid to a solution of a 
weaker acid throws back the ionization of both, but more par- 
ticularly that of the one having the low ionization. This method 
ofdecreasing the ionization, and thereby increasing the toxic 
properties, was used with di- and trichloracetic acids. To solu- 
tions of these acids HCl was added — molecule for molecule— 
_ md the resulting mixture was tested with the five molds. The 
 taltulated increase of toxic properties due to the forcing back 
if the ionization of dichloracetic acid was found to be 168 units. 
The experimental test gave an increase of toxicity for the mix- 
| lure of 147 units more than the additive toxic properties of the 
| lWo acids mixed. In the case of trichloric acid the experi- 
) mental test gave a slight excess (13 units) over the calculated 
—«*WMcrease.3 
Krénig and Paul’s (97) work on anthrax spores is of especial 
terest here. A preparation of anthrax spores immersed in a 


-; “olution of trichloracetic acid for 2 hours and afterwards 


cultured in a favorable medium proved to be entirely sterilized, 
tot a spore Surviving. In another test when a similar prepara- 
. = of spores was immersed in the same concentration for 20 
‘nutes, comparatively few survived. Tests were also made 
with the same acid at = concentration. Spores immersed in 
aes I 

- ths “encentration for 56 hours showed much less injury than 
those immersed for 209 minutes in ~ solution, thus showing 
; : I 

ag that the efficacy of the acid as a disinfectant was more 
J Yeduced than could be accounted for by dilution only. Thew 
~ "the other acetic acids, although not extensive, is quite 
: oa with the results here recorded for the molds. 
Yarocyanic acid J: 20. This poison, so 
dead : HCN 0.46.7, 


. toxic to less highly organized structures. esse 
iy, . ‘ven in minute quantities, the more pay Bt uk 
q Piysicay Pisciis, Physiological action of acids, see article by the writer 

. "Y 33263. May 1899. 


324 BOTANICAL GAZETTE [NOVEMBER 


Ascaris resists a 3 per cent. solution for 75 minutes. A myria- 
pod (Fontaria) excretes HCN when irritated! The accumula- 
tion of many such data has led to the general acceptance of the 
theory that HCN acts chiefly or wholly upon the aldehydes of 
the nerve centers (Loew ’93). This is no doubt quite satisfac- 
tory from the point of view of the animal physiologist, but it 
leaves us without any explanation for its violent toxic properties 
toward plants, which have no nerve centers. 

Little work seems to have been done with it on plants. 
Kahlenberg and True (’96) found that toward Lupinus it had 


a . 
double the toxic value of ionic H proving fatal. To the 


Li 
’ 6400 
molds, however, it is relatively a much more powerful agent, 
415 : : 
having 76.6 times the value of H, thus ranking as one of their 
most fatal poisons. The data on the ionization of this acid are 


meager. At as it gives about one sixteenth the electrical con 


ductivity of acetic acid at the same concentration (Ostwald 85). 
From this we would judge that the ionization is practically zero 
at the concentrations with which we have to deal. 

The value of the CN ion was determined by means of the 
potassium salt, which is quite highly ionized (Kohlrausch 79): 
Were 


It was found to be approximately 8H _ for the molds. 
a toxic 


HCN fully ionized we would expect its solutions to have 


value of about oF. The fact that the practically un-fonit® 
solutions with which we deal have a value of over eight Humes - 
calculated for the entirely ionized acid tells for the extreme!y 
toxic influence of the un-ionized molecule, HCN. a 

Aspergillus showed a high specific resistance to this age™ 
= being necessary to kill all the spores. CEdocephalum 4 
"inhibited by 3943’ 
32768’ 


particularly sensitive, being injured by 


and killed by ne 


i of the 
In table I the toxic values of the un-ionized molecules 


325 


TOXIC EFFECT OF DELETERIOUS AGENTS 


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109] TOXIC EFFECT OF DELETERIOUS AGENTS 327 


different acids, as shown by their inhibiting and killing powers 
_ toward the spores of the five molds, are approximated. Too much 
importance must not be attached to the exact numerical value 
tere expressed for the different molecules, for several reasons. 
We do not know the reaction of the acids toward the nutrient 
medium in which we grow our plants ; we do not know the exact 
liect of the salts and sugars present on the ionization of the 
acids; and were it possible to eliminate every factor causing 
doubt or error, we should undoubtedly find the relative toxic 
operties of the molecules varying with almost every plant 
tested. Let me repeat: the exact numerical values here given 
uenot significant. The order and general proportions of these 
talues ave significant. The emphasis is laid on part I of this table 
for reasons already given, 
Line 1 gives the strength of the various acids required on an 
 Ntrage to inhibit germination. Line 2 is developed from line 1. 
It gives the relative toxic properties of the acids expressed 
/Merically, HCl being taken as 100 for a basis of comparison. 
iM ionized and un-ionized portions are considered separately, 
: ines 4, 5, and 6 being devoted to the former, and 7, 8, and 9 to 
the latter, Line 4 gives the toxic value of the cathion in units 


E ionic i; line 5 that of the anion. Line 6, being the total of 
: ne 5, Sives that portion of the total toxic value of the acid 
" eto be attributed to ionized portion. Line 7 gives the 
tl units.” In other words, that part of the total toxic 
se be accounted for by the un-ionized portion. Line 8 gives 
_,, Ptcentage of such un-ionized molecules present at the inhib- 
_ SPoint. Line 9, the quotient of the residual units divided by 


{itrentage of un-ionized molecules, gives in terms of ionic H 
2 ‘oxic value of the different acid molecules. 

Il is worked out similarly, and has reference to the 
ear “ the acids towards the molds, as shown by their power 
ey the Spores 


: [ Zo be concluded. | 


THE DEVELOPMENT OF THE MICROSPORANGIUM 
AND MICROSPORES IN CONVALLARIA AND 
POTAMOGETON. 

KARL M. WIEGAND. 
(WITH PLATES XXIV—XXv) 


Durinc the past decade perhaps no one portion of the field 
of botany has been worked upon so much as that which deals 
directly with the organs concerned in the sexual process. Espe- 
cially is this true of the higher plants, but a careful survey of 
the present condition of our knowledge on this very point shows 
that some of the most vital questions have as yet received no 
solution. The researches of such men as Hofmeister, Stras- 
burger, Guignard, and Warming have discovered facts in regard 
to the ovule, the embryo-sac, and the cell which have already 
become so universally known as to need no further mention here. 
But we still know almost nothing about the essential significance 
of some phenomena of most common occurrence, and ary 
questions have as yet been investigated only in connection with 
so few plants that generalizations are extremely unsafe, It — 
principally with the hope of increasing, if only by a few species, 
the range of observations that the present studies were under- 
taken. 

The choice of subjects signifies very little. 
much more by the necessity of using plants obtaina 
times rather than by an idea that they all represente 
types of structure. 


It was influenced 
ble at certain 
d different 


METHODS. ‘al 
The methods employed in this work differ in no gies 

way from those so often described in recent cy tological i 

consequently it is scarcely necessary to repeat them pate 

detail. The following general statements are intended ape 

for those who wish simply to know the stains, fixing @ 

chosen for the work. 


gents, 


: [NOVEMBER 
328 


1899] DEVELOPMENT OF THE MICROSPORANGIUM 329 


The Flemming chrom-osmo-acetic acid solution has of late 
years proved of such value that it must now be regarded as the 
very best fixing agent for cytological work. For the following 
studies material from no other fixing agent was used, although 
_ Some was put up in alcohol, sublimate, and picricacid.. None of 
the latter gave satisfactory results. The secret, if there is any, 
in the use of the Flemming solution seems to lie in obtaining 
pid penetration. To accomplish this the material cannot be 
subdivided too much, and when possible even the anthers them- 
selves should be cut open. From six to twelve hours is sufficient 
for complete fixation. The black discoloration caused by the 
_ tsmic acid was removed by the aid of hydrogen peroxid either 
pon the sections themselves or preferably upon the material 
Mm toto, 

For clearing, cedar-wood oil gave the best results. It was 
ways added with great care in order to avoid too rapid change 
of density, otherwise collapse of the cells often occurred. The 
Mrafin used possessed a melting point of 54°. This also was 
added with the same degree of care. The sections ranged in 
thickness from four to six and two thirds microns, and were all 
ton a Minot-Zimmerman revolving microtome. 

: Considerable experimentation was necessary before a suitable 
4 “aining method could be devised. Among others, Rosens’ 
‘chsin-methy lene blue method,! iron-hamatoxylin and the Flem- 
mg ‘afranin-gentian-violet-orange combination were the most 
: “portant. The latter was at length almost exclusively employed, 
; “thy found very satisfactory. Asa general nnclear re 
| *t, better results were obtained by the omission of the 
“nin, and at the same time a large amount of time was saved. 
~ Mange G was always used in a very dilute solution, and 
f i.” Short time, from 15 sec. to 2 min. giving a 
Wace a ora chromosome stain the gentian-violet was 
a short time only, but for differentiating the sp 

r results were 
he stock 


- obta; md Kinoplasmic radiations, much bette 
tained 


‘3 With a very weak solution (2-3 drops of t 
ttrage zur Kenntniss der Pflanzenzellen. Cohn’s Beitrage 5+ 443: 1892. 


330 BOTANICAL GAZETTE | NOVEMBER 


solution made according to Lee in 100° of water) acting for 
from 12 to 24 hours. After washing out in absolute alcohol, 
further differentiation was obtained with clove-oil, or in the case 
of Potamogeton, preferably with anilin oil. The action was 
then stopped with bergamot oil before mounting in balsam. 


CONVALLARIA MAJALIS L. 

The Liliacee have so far furnished some of the very best 
subjects for cytological study. To those already studied must 
now be added another good type, namely Convallaria. This 
plant has unfortunately up to the present time received very 
little attention, although the large nuclei and long rod-like 
chromosomes almost equal Lilium in the ease with which they 
may be studied. The published observations are at present 
limited to those of Strasburger on the pollen of C. Polygonatum, 
and on the endosperm of C. majalis3 The observations on both 
of these are very brief. 

The material for this study was obtained from plants gr 
in the University greenhouses. They were here under a constant 
though moderate condition of forcing, which thus enabled one 
to obtain an unusually large proportion of dividing nuclei. The 
progressive development of the flowers in the raceme makes it 
possible to find many stages on a plant all at one time. +° 
insure rapid penetration of the fixing agent the uppet and er 
ends of each bud were cut away, thus exposing directly the cells 
of the anther. 

In order to determine if there might not be a relatio 
the nuclear division and the environment of the plant, ©sP' 
as to the amount of light, humidity, etc., several experime " 
were made. In the case of Convallaria only the effect of Bit 
could be studied, since the other conditions in the ear ht, 
were practically the same. In order to test the effect of oe 
material was collected at various times during the day and at 
The results, however, were wholly negative. Spindles 
found in about the same proportion in every collection. 

* Befruchtungsvorginge bei den Phanerogamen 171. 1884. 

3 Theilungsvorgange der Zellkerne 43. 1882. 


own 


n betwee? 


especially 
nts 


1899] DEVELOPMENT OF THE MICROSPORANGIUM 331 


THE DEVELOPMENT OF THE MICROSPORANGIUM. 


In 1873 Warming’s important work on the development of 
theanther apeared.* In this paper we find for the first time a 
correct description of the succession of cell divisions result- 
ing in the formation of the archesporium and the anther wall. 
| It was not followed up, however, by other investigators, and 
en in late years nothing of importance has been done along 
this line. The most important of the subsequent papers is 
indoubtedly that of Engler in which Orchis is taken up in 
detail, 


Warming established the fact that the archesporium arises 
from the daughter cells resulting from the division of the hypo- 
dermal layer at each corner of the anther. If the hypodermal 
lls form a true layer, then the archesporium will usually also 
tein the form of a layer; but in some cases the hypodermal 
cals may be reduced to one, and the resulting archesporium in 
that case is simply a vertical row of cells. In any event, there is 
most no subsequent division in the archesporium, growth being 
‘onfined entirely to an increase in size of the existing cells. Of 
) 3 Wo original daughter cells of any hypodermal cell, the inner 
Yes tise to the archesporium, the outer to the anther wall. By 
"0 or three periclinal divisions progressing in a centrifugal 
Tanner a radial row of cells is formed, the inner cell of which 
Mes the tapetum, the outer the endothecial layer, and the 
= finally disintegrate. Although many of the divisions are 
a a linal, some are radial and others transverse, by which epee 
fe genmttion is provided for the growth of the archesporium. 
a nds also that cell division in the epidermis is almost entirely 
yd transverse, and this structure remains always one-cell 
“Me oo His subjects for investigation, however, included a 
Aleagl "cotyledon; and Engler,’ who attempted to show mor 
: Y the application of Warming’s laws to the monocotyledons, 
4 


pau ay “9 nt. 
: ti. cL achungen liber pollenbildende Phyllome und Caulome. Hanstein’s Bot 
: te ANd Se 


Beitrag i t. 10: 
Abe age zur Kenntniss der Antherenbildung. Pringsh. Jahrb. f. wiss- Bo 


332 BOTANICAL GAZETTE [NOVEMBER 


used Orchis as the only type. At present, therefore, our knowl- 
edge of the details in this group is very meager.® 

In Convallaria the early stages in the development of the 
anther are not so easily understood as are those of the dicotyle- 
dons; but after the investigation of a large number of prepara- 
tions it seems probable that the following is the proper inter- 
pretation, not alone for Convallaria but also for many other 
-monocotyledons. The earliest stages obtained show in cross- 
section a four-angled anther with a radial row of cells at each 
angle, which apparently result from the division of a single 
hypodermal cell. 

The condition at this stage is represented in fig. I. The 
innermost cell, the primary archesporial cell, very soon divides 
in various directions until a considerable mass of tissue 'S 
formed. This division takes place very early, so that the final 
number of archesporial cells is formed even before the anther 
has become obviously lobed. At this stage it is only with com 
siderable difficulty that the archesporium can be distinguished 
from the wall (fig. 2). The original radial row of cells, 
descendants of the primary hypodermal cell, may often be rec- 
ognized for a considerable time after they are first formed, vo 
in some cases one or two cells on either side may also divide 
several times in a radial direction. The greater portion of the 
wall, however, is derived from a few irregular divisions of o 
cells at each side of the archesporium, while the epidermis #6 
the same time increased by a few anticlinal divisions. In sat 
to this the cells between the archesporium and the connect 
may also undergo a few radial divisions. As 4 result the 
cells in the older anthers do not stand in distinct radial se 
as in the dicotyledons, simply because they were not all web 
by radial division; but, notwithstanding this, there are 4 

6 While in press the following papers have appeared: © ar cas pil 

GUIGNARD, L.— Le développement du pollen et la réduction dans le Nava 
Arch. d’anat. Microscop. 2:455. 1899. . an. 1899- 

CALDWELL, O.—- Life history of Lemna minor. BOT. Gaz. 27 +37: ‘erospores of 

FULLMER, E. L.— The development of the microsporang!a and mi 
Hemerocallis fulva. Bot. GAz. 28:81. 1899. 


1899] DEVELOPMENT OF THE M1CROSPORANGIUM 333 


regularly three or four layers of cells differentiated entirely 
wound the sporogenous tissue. These stages were found in the 
autumn previous to the time of flowering, and all subsequent 
growth, both in the archesporium and in the wall, is due 
eitirely to the increase in diameter of the cells already formed. 
This was determined by an actual count of the cells in a great 
many cases, 

Inmost dicotyledonous anthers the archesporium becomes: 
distinct at a very early period. No such sharp demarcation 
exists, however, in the monocotyledons. In all of the cases 
‘tudied by the writer, and in those treated by Warming and 
Engler, the transition from the wall to the archesporium is so 
sradual, especially in the younger conditions, that only the most 


/ teotyled 
the 


r of 


‘ Ata slightly later period the fourth or innermost laye 
fea and 


ay . begins to enlarge. The cells grow considerably, 
\ in be mistaken for archesporial cells if it was not for 
: a. in the cytoplasm, This layer is the tapetum 


The |: : 
i history of the tapetal nucleus has been studied by 


334 BOTANICAL GAZETTE [NOVEMBER 


Strasburger in Malva, but not in detail.?_ In this plant two nuclei 
were found in each cell. Guignard-also figured two nuclei in 
the tapetal cells of Lilium, and it now seems probable that the 
phenomenon is quite general, as Strasburger in the above cited 
work states. It was originally believed that these two nuclei 
were formed by direct division; but Strasburger*® has shown that 
this was really a process of fusion, and that the two nuclei were 
formed at an earlier date by the ordinary indirect method. 

In Convallaria, at the period just preceding synapsis, the 
already enlarged tapetal cells contain only one nucleus; but 
during synapsis, and even up to the first pollen-mother-cell divi- 
sion, the nuclei one by one divide by the mitotic method. Such 
spindles are seen in fig. 4. It is probable that every tapetal 
nucleus finally undergoes division, but this could not be accu- 
rately determined because of the subsequent fusion. The 
different stages in the fusion of these nuclei were sufficiently 
frequent after the first division of the pollen-mother cell (jig. 5): 
Cases could often be found even in anthers where tapetal spi 
dles also occurred. It is doubtful if all the pairs of nuclei fuse 
—in fact, it is probable that they do not, since many remain 
distinct even after the tapetum shows signs of disintegration. 

The wall of the mature anther in Convallaria presents no 
new features. It is composed of a conspicuous sae 
well-developed fibrous layer (endothecium) with beautiful spit 
markings, and the remains of the other wall layer (fig. 6). é 
remnant of the tapetum is now left. The time of disappearance 
of this structure was interesting because it occurred in some eget 
as early as the pollen-mother-cell stage, while in cae oi 
until after the pollen grains were mature. The time is seid 
the same even in anthers from the same bud. 

THE NUCLEUS OF THE ARCHESPORIUM. ae 

The very earliest stages of the archesporial nucleus oes 
vallaria differ only slightly in appearance from trite 7 ne 
nuclei ( fig. 7). The linin thread is exceedingly thin 2° 

7 Ueber den Bau und das Wachsthum der Zellhaute 89. 1882. 

*Theilungsvorgange der Zellkerne 99. 1882. 


1899] DEVELOPMENT OF THE MICROSPORANGIUM 335 


aswell as extensive. Careful examination shows that it is not 
acontinuous ribbon at this. period, but rather an anastomosing 
aetwork in which large granules of chromatin of unequal sizes are 
imbedded, giving it thus a more or less knotty appearance. 
Ata period quite early in the development of the anther the 
chromatin network contracts into a dense and ultimately spher- 
_ttalmass, which in most cases is in contact with the nuclear 
fembrane, but may also lie free in the nuclear cavity (fig. 8). 
The ball becomes so dense that it is ordinarily impossible, even 
with the weakest stains, to distinguish the separate threads of 
Which it is composed, except at the periphery where they pro- 
ket into the nuclear cavity outside. This condition is synapsis 
|Convallaria. The nucleolus does not lie inside the mass and 
adjacent to the membrane, as has been claimed for some other 
Plants. On the contrary, it always lies outside so far as could 
te determined, but usually in contact with the mass and at the 
‘ide away from the wall more often than toward it. 
Judging from the number of preparations obtained, the 
. tucleus must remain in the synapsis condition for a considerable 
ime, The first indication of the return from synapsis is found 
| the gradual Separation of the outermost linin threads, and 
Tally the whole mass becomes more open (fig.9). Buta vey 
Mange thing now occurs. At the moment when the chromatic 
2 ad is Spreading out, between its meshes and in the cavity 
de are to be seen large granules, or more properly speaking 
pe of various sizes which are decidedly chromatic, in fact 
: o “xactly like chromatin. The nucleolus meanwhile still 
ous intact, Where these bodies come from or where they 
Mt, at th: determined. They were always eins ber 
: Meleolus ig Stage. The masses were often as po 5 as 
c. a more often smaller. It might be ee ual 
Wy the Se the nucleolus becomes fragmented either n at 
Sport pate of the reagents, but the preparations 
Mote aa In very weakly stained nee - caG 
si. €olus with its central vacuole coul - ap 
: in the same nucleus the expelled masses were stat 


336 BOTANICAL GAZETTE [NOVEMBER 


dark like the chromosomes at the time of division. Again itis 
possible that the chromatin had left the linin thread and col- 
lected in the above manner before the material was fixed. Or 
perhaps the reagents caused such a change in the chromatin. 
The investigation of fresh material alone can decide whether the 
process is normal or artificial. 

A remarkable change has also taken place in the chromatin 
thread. Before synapsis it was a network containing very large 
irregular granules, now it is a spirem with the granules much 
reduced in size and more uniform. For the most part they are 
but slightly broader than the scarcely thickened linin. These 
above mentioned peculiarities were found not in one case alone, 
but in a great many preparations, in fact in every preparation 
that contained the right stage. 

Synapsis has now been found in many plants, in all of which 
it always seems to be a natural condition, and it is quite proba- 
ble that it will prove to be a universal phenomenon, occurring a 
a certain period previous to each heterotypic division. Besides 
this, it has now been shown to occur in at least a few animals. 
Among plants it has been found in the Liliacea by Strasburger, 
Sargant, and others; in the Hepaticae by Farmer; and in Pota- 
mogeton and Acorus by the writer, in addition to several other 
‘plants, the studies on which have not yet been published. There 
now seems to be little doubt that the condition is a natural - 
for the following reasons. It may be found in the fresh mate 
rial, at least in Convallaria and Lilium. Moreover, it always 
occurs at the same stage in the development of the anther. 
Various structural changes in the chromatin thread always 
accompany it. In addition to this, many preparations show 1 
the mass may be deposited at any side of the nuclear cavity 
without reference to its position in the anther. 

Considerable uncertainty has always existe 
synapsis really is. The term was introduced by Moo 


year 1895, but the description was so brief that it is s° 
and plants 


das to just what 
re? in the 


Essential similarity of the chromosome reduction in animals 
Annals of Botany 9: 435. 1895. 


1899] DEVELOPMENT OF THE MICROSPORANGIUM 337 


dificult to understand just what the author had in mind. It 
| ems probable, however, that the condition found by him was 
the same as that described above for Convallaria. He was able 
- to demonstrate the appearance of the same phenomenon also in 
animals, especially in Triton, thus emphasizing the fact that the 
sleps in the maturation of the sexual gametes is to a certain 
‘fxtent similar in both plants and animals. 
During the past few years synapsis has been described by 
sveral authors, most of whom now consider it to bea natural 
condition of the cell, although it seems probable that the phe- 
_fomenon described is not the same in all cases. 
Farmer found a contracted condition of the chromatin thread 
Wthe spore-mother-cells of the Hepaticae, but the figures do 
lot show as great a contraction as is found in Convallaria. He 
Mites that the nuclei at this stage are difficult to fix, often show- 
|S signs of fragmentation, and that there is usually a chro- 
matic change in the cell. 
Convallaria has so far shown no case similar to that described 
- by Miss Sargant*™™ for Lilium, in which two rows of dots were 
found on the thread before synapsis. A double row in the for- 
| oy Plant is found only very late in the history of the spirem ; 
aaa does the nuclear membrane disappear or even become 
F “distinct during Synapsis. Otherwise the description and figures 
quite Similar to Convallaria. In some preparations an extru- 
oa of granules from the contracted mass was found, but was 
ore 2 fragmentation of the nucleolus rather than 
ie Separation of portions of the chromatin. 
€ Conflicting results obtained are probably due, in part at 
eed ys 
these A ea have been referred to ya . eee 
. rmal condition, and to this the term synap 
: te ye formation and nuclear division in the Hepaticae. Annals of Botany 


a The 
Bick . t n 
Wi g50 _ of the sexual nuclei in Liliwm Martagon. Annals of Botany 


» Oa 
*ryokinetische Probleme. Pringsh. Jahrb. £. wiss. bot. 28: 158. 1895- 


338 BOTANICAL GAZETTE | NOVEMBER 


properly belongs. The others are probably caused by the action 
of the reagents. 

Mottier’s*3 figures representing synapsis in Lilium do not show 
nearly as much contraction as in Convallaria. From the descrip- 
tion it would seem also that they represent a much later stage 
in the development of the spirem than the one in which synapsis 
occurs, either in Convallaria or in Potamageton. 


FIRST NUCLEAR DIVISION OF THE MOTHER-CELL. 


The spirem stage preceding division is very well marked in 
Convallaria (fig. 0). The much coiled wire-like thread found 
immediately after synapsis gradually increases in thickness, and 
the chromatin granules become less prominent. One large 
nucleolus and usually two or three smaller ones are present at 
this period. The longitudinal splitting of the chromatin thread 
is accomplished so quickly that it was found impossible to 
observe the successive steps in the process. At the same time 
the thread becomes considerably thicker than immediately before 
the division (fig. rz). It is difficult to understand just how this 
doubling takes place, because at the very first indication of such 
a condition the threads are already separated. They lie parallel 
with each other and slightly coiled. The further development 
indicated a more or less complete subsequent fusion of the parts, 
so that before passing into the nuclear-plate stage the dual 
nature is entirely lost except for an occasional lobing at either 
end (fig.12). Not even the granules are longer visible, and the 
chromosomes are at this stage apparently homogeneous. ae 
is probably only apparent, however, and due really to the density 
of the stain and the close proximity of the parts. 

After the chromatin thread has become double, beside i 
thicker than before, it also possesses fewer coils, which 8 sr 
ably due to a longitudinal contraction of the whole spirem: 
about the time when the nuclear membrane disappears the thic 


: ; pager a: somes 
ened chromatin band segments into the individual chromo 


sides being 


3 Beitrage zur Kenntniss der Kerntheilung in den Pollenmutt 
Dikotylen und Monokotylen. Pringsh. Jahrb. f. wiss. bot. 30: 175- 1897. 


; 
q 
: 
i 


1899] DEVELOPMENT OF THE MICROSPORANGIUM 339 


ig. 12). The successive stages in the process of segmentation 
could be easily traced. The constrictions gradually become 
deeperand deeper, while at the same time the chromatin is with- 
trawn from the constricted regions. The resulting segments 
were always of the same length as the mature chromosomes 
appearing on the nuclear plate. Repeated examination of these 
stages failed to reveal a V-shaped bending back of the segments 
‘omesponding to that described by many investigators for 
lium. Moreover, the chromosomes do not have four lobes at 
the end, as would be more likely the case if they were formed 
by the folding-back process, but only two. It seems, therefore, 
that in Convallaria at least the chromosomes, although double 
nature and hence sometimes showing a longitudinal split, are 
always straight and formed simply by the transverse fission of 
the chromatin thread. 

The changes which the chromosomes pass through while on 
the nuclear plate are very difficult to make out, and conse- 
| Mently little that is definite can be said about them. The 
“gments seem to be straight or slightly curved and lie mostly 
4 radial manner on the plate (figs. rg, 79, 20). The earlier 
‘tages show a nearly cylindrical chromosome, but very soon this 
comes changed into the characteristic +-like structures which 
a very Commonly seen at this stage. These structures seem 
eg formed, as Belajeff'* and Strasburger' have shown, by a 
ada Splitting at each end of the cylinder, but in peepee 
Staal Planes, The inner forks are drawn apart by, the spindle 
ie the two outer ones separate in the plane pe oe 
aa €. In many cases a fissure may be seen ape 
j - Pex of one long arm directly through the mi 
os ne (fig. 20). oo 
“the i ‘ei V-shaped segments resulting Ons the separa i 
le ves of the + pass to the poles in the ordinary man 

WES. 14, 75, 22). The daughter segments proceed with the 
3 _ Kenntniss der Karyokinese bei den Pflanzen. Flora, Erganzungsb. 79 ' 434- 


ee 
*yokinetische Probleme. Pringsh. Jahrb. f. wiss. Bot. 28: 183. 1895. 


340 BOTANICAL GAZETTE [ NOVEMBER 


angle of the V in front, as one might expect if the spindle fibers 
are assumed to be exerting a pull upon them. They are always 
more slender than the parent chromosomes, which would be the 
case if a division of the original substance had taken place. 
Occasionally a V will straighten out and lie along the spindle 
fiber, reaching almost from the equator to the pole. 

The spindle in Convallaria is formed in the same way as in 
Lilium. The disappearance of the membrane is exactly coin- 
cident with the appearance of the kinoplasmic threads which 
immediately penetrate the nuclear cavity, and also extend 
outward into the cytoplasm (fig. 72). Very little light, how- 
ever, could be thrown on the fate of the nucleolus, and its dis- 
appearance was very sudden. The multipolar spindle is not 
very distinct in this plant (fig. 73), and the poles scarcely ever 
extend much beyond the limits of the old nuclear membrane, 
and are often difficult to distinguish at all. The bipolar spindle 
is usually truncate at the poles, but unlike that of Potamogeton, 
it is broad and barrel-shaped, possibly due to the large number 
of chromosomes (jig. 74). In this respect it is similar to 
Lilium. In no case was there even so much as a granule present 
at the pole, or any thing that could be mistaken for a centro- 
sphere. A strong nuclear plate follows the division, resulting 
in a cross wall separating the cell into two hemispherical parts. 


SECOND NUCLEAR DIVISION OF THE MOTHER-CELL. 
The resting stage between the first and second divisions 9 
Convallaria is very short. One may often find the nuclei at 
one end of an anther in the cell-plate stage, while those at bas 
other end have formed the nuclear plate for the second han? 
The daughter nuclei do not seem to pass entirely into 4 nowt 
resting condition. So far as could be determined, no saan 
ever appears, nor does the nuclear membrane become well et 
oped, however a very delicate membrane may often be amr 
The chromosomes apparently retain their identity fete 
this stage. From the pole view it can be determined that t* 
still possess their V-shaped form (fig. 16). 


1899] 


DEVELOPMENT OF THE MICROSPORANGIUM 341 


The transition from the resting stage to the second spindle is 
very abrupt. Two poles are formed in the cytoplasm, which 
move farther apart, so that a bipolar spindle is very quickly 
formed (fig. 77). In the preparations examined, no polar radia- 
tions or multipolar spindles were found. The already distinct 
chromosomes merely move together toward the center in order 
0 form the nuclear plate. Even now most of them retain the 
Vform, and only an occasional one becomes nearly straight. 
They are very irregularly arranged, so that the long arms of 
Some project outward toward the poles, giving a ragged appear- 
ace to the plate, thus distinguishing it immediately from the 
plate formed in the heterotypic division. 

It was found impossible to determine absolutely the nature 
of the segmentation, but from a study of all the stages obtain- 
ible it seems probable that the V is divided transversely. This 
@ course would not mean a transverse division of the original 
ttromosome, if the processes described for the first division are 
he true ones, but rather the completion of a second longitudinal 
‘ltting. No figures were found in any of the spindles that could 
‘interpreted as a case of undoubted longitudinal splitting of 
: Vafter coming on to the nuclear plate of the second spindle. 
: ie other hand, the nearly straight segments moving along 
ee toward the poles are all much shorter than the Vs, but 
ee stely the same diameter instead of narrower, as one 
: expect if longitudinal splitting had taken place. 

* The later Stages are all perfectly normal. The chromosomes 
; | 8¢ themselves in the daughter nuclei and appear at length 


© into a continuous chromatin thread. The nuclear mem- 


‘8° Of the segments to the poles, is much less distinct 
“ring the first division, and it is composed of fewer fibers. 
the = daughter nuclei are formed, a cell-plate is deposited in 
ion ¢ unet- The spindle now disappears and the tetrad 
VOF the pollen-mother-cell nucleus is complete. 


342 BOTANICAL GAZETTE [NOVEMBER 


The number of chromosomes in Convallaria is quite large. 
A count in the nuclear plate stage showed eighteen segments as 
the reduced number. The same number may be counted during 
the subsequent resting stage and also after the second division. 


THE MICROSPORES. 


After the second division of the mother-cell nucleus the 
young pollen grains do not separate immediately, but remain a 
short time inclosed in the thickened walls of the parent cells. 
With little difficulty one can follow all the steps in the process 
of dissolution which these walls undergo. First the increasing 
sponginess of the already thick wall; a simultaneous differentia- 
tion of its inner layer destined to become the wall of the spore; 
and finally the complete solution of the outer part, leaving the 
young pollen grains united only by the intervening walls. These 
apparently split at once into two layers, thus freeing the mem- 
bers of the tetrad. : : 

The pollen grains at first are quite small, and possess thin 
purple-staining walls, finely granular cytoplasm similar in ane 
sistency to that of the somatic cells, and a highly chromatic 
nucleus which occupies about one fourth of the cell-cavity 
(fig. 23). The further changes are mostly normal. The pollen 
grains, which from the first are elliptical, gradually increase in 
size until their volume is more than doubled. The wall increases 
in thickness, and the whole grain assumes a bluer tinge with 
gentian-violet. 

A short time before the flower opens, the nucleus undergoes 
division, whereby a generative cell is cut off (fig. 24): shee 
cell is lenticular in form, and separated from the gencral ie! 
of the grain by a distinct cell wall. The generative nucleus 15 
exceedingly chromatic, so much so in fact that it stains ere 
homogeneous dark purple with the gentian-violet. The wren 
tive cell in Convallaria seems to differ from those in most of ne 
monocotyledons described by other writers in not sep 
an early period from the wall of the pollen-grain. It app y 
remains in all cases attached until the time of pollination. 


1899] DEVELOPMENT OF THE MICROSPORANGIUM 343 


The division of the generative nucleus into the two sperm 
wclei must take place in the pollen tube, good stages of which 
were not obtained. Except in the one point above mentioned, 
| the microspores of Convallaria do not differ in any essential way 
ftom those described by Strasburger”® for a large number of the 
higher monocotyledons. . 


POTAMOGETON FOLIOSUS RAF. 


The numerous investigations recently made upon plants 
belonging to the orders Alismacee and Naidacee have shown 
that many peculiar conditions are to be found among these 
soups of monocotyledons. During the past summer the writer 
Was able to procure excellent material of Potamogeton foliosus in 
the ponds about Ithaca - and it was decided to make a detailed 
Study of this plant for comparison with the studies already made 
‘Yothers. No one seems to have investigated this genus from 
| ‘cytological standpoint. 

f the papers on nearly related plants must be mentioned 
. that on Naias by Magnus,?7 on Naias and Zannichellia by Camp- 
ll® and on Alisma” and Sagittaria° by Schaffner. 

S The material was collected during the months of July and 
August, at which time the oldest flowers are just producing fruit. 
The floral Spikes mature in succession as the plant branches, so 
by the very youngest flowers and also the fruits may be found 
‘nthe same individual. There being no cutinized layer sur- 
utding the bud, the latter is especially easy to penetrate with 
mg agent. The material used for this study was therefore 
the “eptionally good condition. As in the case of Convallaria, 
“collections Were made at certain times during the day and 
the result also was exactly the same. 

: age tttssvorginge bei den Phanerugamen 22. 1884. 
= Fage zur Kenntniss der Gattung Naias. Berlin. 1870. 


A 
yy, P'sical study of Naias and Zannichellia. Proc. Calif. Acad. Sci. 11. 
“The emb 
% 


TY sac of Alisma plantago. Bot. GAZ. 21:123. 1896. 
‘ontibution to the life history of Sagittaria variabilts. Bot. GaZ. 23252+ 


344 BOTANICAL GAZETTE [NOVEMBER 


THE DEVELOPMENT OF THE MICROSPORANGIUM. 

To trace the development of the anther, and especially the 
differentiation of the microsporangial archesporium in Pota- 
mogeton, is no less difficult than in Convallaria. Oddly enough, 
the only monocotyledonous type studied by Warming was one 
of the Naiadacez, namely Zannichellia. Warming thought that 
in this plant the process was essentially identical with that in 
the dicotyledons, but these results may be questioned, owing to 
the apparent paucity of material at his command. 

The present study of Potamogeton seems to throw a little 
more light on the problem. In the young anther, which at 
maturity is always two-celled, there is found at each of the two 
more prominent angles of the quadrangular cross-section a single 
hypodermal cell, which at this stage is slightly larger than the 
surrounding cells and richer in protoplasm. This presently is 
divided by periclinal walls into two, and later into three daughter 
cells, each produced probably in centrifugal succession (jig. 25). 
The innermost of this series now immediately begins to enlarge, 
and becomes at once the primary archesporial cell. This cell 
undergoes rapid division, resulting at length in a number of cells, 
all formed from this one original archesporial cell. The irregu- 
larity in arrangement, and the gradual decrease in size from the 
center toward all sides, nevertheless suggest that some may owe 
their origin to the division of the surrounding tissue. The 
same gradual decrease in size takes place also on the side of the 
archesporium toward the connective. 

It will be seen from the above account that the tapetum here 
and in Convallaria is not a morphologically distinct structure 
until at a comparatively late period in the development of the 
anther. It is not until the stamen is half mature that the arche- 
sporium becomes distinct from the wall. It can always be gat 
nized at this period by the finely granular contents of the = C 
just as was the case in Convallaria. No tapetum can be ae 
guished for some time. The cells of the inner layer of the wall, 
which from the first are smaller than the central cells, ace 
‘take on a dense and partially disorganized appearance. There 


1899] DEVELOPMENT OF THE MICROSPORANGIUM 345 


mw longer any doubt but that the tapetum is differentiated from 
the wall, rather than from the archesporium, as a hasty inspec- 
tion would seem to indicate. 

The wall of the anther at this stage is composed of three, or 
mely four, layers of cells (fig. 26). The outermost of these 
ayers remains almost unchanged until the anther is mature, and 
‘Sindeed the true epidermis. The other two or three layers 
lave some of their cells arranged in more or less distinct radial 
Ws, Suggesting, as in the first case, that each row is the deriva- 
tive of one hypodermal cell. The greater portion of the wall, 
lowever, is formed, as in Convallaria, from the cells of either 
side of the archesporium, and these are not necessarily deriva- 
tives of a hypodermal cell. Indeed, so far as could be deter- 
tined, the growth was brought about exactly as in Convallaria. 

ing the maturation of the anther the behavior of the cells is 
‘mal, The third layer undergoes disintegration, as does also 
‘he fourth, Which is the tapetum. The epidermis remains nor- 
mi,while at the same time the second layer becomes thicker 
walled than the rest, acquires spiral or reticulated thickenings, 
‘itis indeed a true endothecial layer ( fig. 27). 

\ Campbell considers the anther of Naias to be a so-called 
/Gulome” Structure, in which are early differentiated plerome 
be periblem, the upper cell of the plerome cylinder becoming 
~* Mtchesporium, This in itself does not preclude a process 
ad to that described above for Potamogeton ; although 
. Pbell himself js quite certain that the origin of the arche- 
& i Naias is not traceable to a single cell. He found 
tee of only two layers besides the pigs 
A tacin ree. In Zannichellia the same difficulty was fo 

‘ym, the development, but this was probably because the 
thy are “ay Not counted as a wall layer. The satu 
et cat ere also at first scarcely distinguishable from ae a a 
®hesporis In this plant there are three layers surroun ing 

py Mall of which finally become disintegrated. 

plete disintegration of the tapetal cells in Potamogeton 
o Soincident with the divisions of the pollen-mother-cell. 


ane COm 
Salm 


346 BOTANICAL GAZETTE [NOVEMBER 


When the young pollen grains are free in the anther, therefore, 
only a disorganized mass of protoplasm is in the position for- 
merly occupied by the tapetum. This substance is very soon dis- 
tributed among the pollen grains, where it possibly serves as 
nutriment. The tapetal cells of Potamogeton never contain two 
nuclei. In this respect, therefore, they differ decidedly from 
Convallaria. 


THE ARCHESPORIUM AND MOTHER-CELLS. 

Division in the primitive archesporium ceases at an early 
period, after which the development is confined to growth and 
constitutional changes in the cells already formed. The defini- 
tive archesporial cells are at first quite small, but during the 
long period of growth that now commences they double or even 
triple their original size. The mature pollen-mother-cell con- 
tains a very large nucleus surrounded by abundant cytoplasm. 
Unlike most monocotyledons, the cell wall here remains very 
thin, and does not become irregularly thickened, as in Conval- 
laria and Lilium (fig. 35). A similar condition has = 
observed also in Naias and Zannichellia. A very short me 
therefore, is required for disintegration, which undoubtedly 
accounts for the almost immediate separation of the yous 
pollen grains. 

THE ARCHESPORIAL NUCLEUS. 

Potamogeton belongs to an entirely different class ht 
Convallaria so far as the nuclei are concerned. The Lilium 
type, to which the latter plant belongs, possesses the wel a 
dense spirem and the large oblong chromosomes. The aon 
of Potamogeton are apparently very poor in chromatin. 
few chromosomes are small and spherical and the spirem ted 
meager. A detailed comparison with Convallaria, therefore, 
will be especially interesting. at 

The very young archesporial nuclei in Potente 
scarcely different from the surrounding vegetative nuclei. 
are surrounded by a definite membrane, have a large — 
like body, and a very poor linin network, which lies close t° 


|-known 


1899] DEVELOPMENT OF THE MICROSPORANGIUM 347 


wall (fig. 25). As the cell gradually expands, the nucleus also 
increases in size. The nucleus just before synapsis has already 
aquired nearly its full size, and the linin network composed of 
very slender threads is plainly visible. In it are irregularly 
distributed a few large and small granules of chromatin. Both 
the linin and the granules are exceedingly meager as compared 
with Convallaria. The nucleolus is a gigantic body, much larger 
than those in most plants, and takes the gentian-violet stain 
readily, making it thus a very striking object in the cell. 
Attached to it on one side is a small wart-like body only slightly 
lager than the largest chromatin granules. Rarely two of these 


&. JY Pressed against it. With the highest magnification, 
_ «tral part of the mass still appears too dense to distinguish 
Y structural characters. On the periphery, however, the free 
: of the network may be seen easily. The large nucleolus, 
: “panied by the wart-like body, remains in its central posi- 
™ the nucleus throughout the synapsis stage. It always 
pyre deeply than the linin. The mass of linin, therefore, 


Ween the nucleolus and the nuclear membrane. The later 


348 BOTANICAL GAZETTE | NOVEMBER 


stages of synapsis are marked by the same striking peculiarities 
that were met with in Convallaria (fig. 30). In just the same 
way large globules of some deeply stainable matter accumulate 
on the outside of the contracted mass, appearing as if expelled 
from it. They seem, however, to be distinct from the nucleolus, 
as there is no apparent fragmentation nor budding of the latter. 
The nucleolus throughout the whole process remains of exactly 
the same size and regular contour, and with the little wart-like 
attachment undisturbed. 

The spirem stage is much shorter than in Convallaria. The 
contracted linin network gradually begins to expand until the 
threads are again spread out beneath the membrane. Here 
again we notice a decided change in the structure of the linin 
thread, just as was the case in Convallaria. It is no longer s0 
slender, and provided with such large granules, nor is it so con- 
spicuously in the form of a network. The spirem is composed 
of a few rather thick linin threads extending in various direc- 
tions around the nucleolus, and crossing each other occasionally. 
In them appear small chromatin granules, which, however, af 
much smaller and more regular in size than those present before 
synapsis (fig. 37). The whole process, therefore, is exactly 
comparable with that in Convallaria. 


FIRST NUCLEAR DIVISION OF THE MOTHER-CELL. 


The stages in the nuclear development preparatory to the 
first nuclear division are not nearly so marked as in Convallana. 
The first indication of division is found in the gradual massing 
together of the chromatin into a number of irregular masses 
simulating those found just before synapsis. The difference a 
in their larger and more equal size. Just at the time when ef 
nuclear membrane is disappearing the number of these gee 
of chromatin may be determined approximately. Proba if 
fourteen or sixteen is the correct number. They S007 ae 
lie together in pairs, in which case two may be easily mist : 
for one (fig. 32). During the later stages the two pan 
each pair seem to lose their identity, so that when on the sp! 


1899] DEVELOPMENT OF THE MICROSPORANGIUM 349 
itis not possible to count more than seven or eight chromo- 
somes. 

Whether a fusion takes place here one cannot determine, 
since the small size renders it impossible to follow the process 
accurately. The exact manner of segmentation upon the spindle 
is also still in doubt. Seven chromosomes are found in the 
daughter nuclei, which leads us to infer that each of the origi- 
ual masses splits into two, but the preparations show no indica- 
tions of any such division farther than that in many cases 
under high magnification, it seemed as if the chromosomes pos- 
“ssed a +-like structure similar to that in Convallaria, but 
the figures were not distinct enough to allow of any definite 
conclusions, 

Inregard to the formation of the spindle a few notes may 
be given, although the results do not differ essentially from 
those obtained by Mottier in Lilium. The kinoplasm is at first 

ited to a thin felt-like coat surrounding the nucleus (fig. 32). 
On account of the very large space occupied by the nuclear 
| QD, it is easy to observe the entrance of the kinoplasm into the 
; tuclear cavity. This takes place apparently before the entire 
tisappearance of the membrane. The latter sometimes is still 
Mable after the nuclear cavity is nearly filled with kinoplasm. 
a first thought it seems impossible to conceive of a substance 
_ Pissing through the nuclear membrane in this way. But Mot- 
- has shown that the membrane itself is probably nothing 
Be close weft of kinoplasm. We have then merely 
~ “sume that the inner threads of this weft separate from the 
_* traverse the nuclear cavity instead. Finally the whole 


.." membrane. They are from the first a part of it. At 
7% € spindle is multipolar (figs. 33,34)» but the poles are 
i. amber. and very soon disappear, thus giving ae 
|. Normal bipolar type. In its mature condition the spindle 
. a = and the poles are very acute (jig. 35). The fibers 
: Of. cit, Pringsh. Jahrb. f. wiss. bot. 30: 176. 1897. 


350 BOTANICAL GAZETTE [NOVEMBER 


are few in number, probably not exceeding the number of chro- 
mosomes. 

In many cases the point at the pole. toward which the spin- 
dle fibers converge was occupied by a granule both in the first 
and second division spindles. This granule in well-stained 
preparations was always dark, but its inconstant occurrence 
was decidedly against its being considered a permanent struc- 
ture. The cell plate forms before the spindles of the second 
division (jig. 36). 

The body, which looks so much like a nucleolus, disappears 
previous to the first division at almost the same time as does 
the nucleolar membrane. At this period it presents a more of 
less irregular and lobed appearance, but vanishes so quickly that 
it was impossible to determine whether the process was one of 


fragmentation or solution. 


SECOND NUCLEAR DIVISION OF THE MOTHER-CELL. 


Before the second division there is a distinct resting stage 
An indistinct membrane is formed, and even a nucleolar body 
may appear. This latter, however, never becomes so large as in 
the archesporial nucleus, and often seems to be entirely absent, 
or at least indistinguishable from the chromosomes (fig: 3): 
A thick linin thread is usually formed, but the chromosomes 
remain distinct. In this character Potomageton agres® ” 
with Convallaria. During this resting stage it is again possible 
to count the chromosomes, when the number is Still found to ae 
seven or eight. This resting nucleus can be distinguished ea 
by its much larger size from the one formed after the secon 
division. 

The origin of the spindle could not be trac 
preparations showed it in the mature condition 
These spindles are smaller and more slender 
described above. Like the latter, they have very P 
and in both cases the poles are almost if not quite in con 
the cell wall. 

The chromosomes are closely aggregated in t 


ees well 


ed, but many 
(fig. 37 ). 
than those 
ointed poles, 
tact w! 


he nucleat 


1899] DEVELOPMENT OF THE MICROSPORANGIUM 351 


plate stage, so that it was impossible to determine just what 
happened to them at this point. The daughter chromosomes 
_ move to the poles very evenly, that is, with the same degree of 
tapidity (fig. 38). A count here showed the seven segments 
present. Although the daughter nuclei are quite small, the 
chromosomes remain distinct for some time, and can be again 
counted. After a time the nucleolar bodies also reappear, and 
the division is then complete. 


THE MICROSPORE. 

Our knowledge of the internal structure of the pollen grain 
_teally dates from the time of Hartig.? In this author’s work is 
the first mention of the discovery in Tradescantia and several 
other plants of two nuclei in the pollen. This important dis- 
every seems not to have been noticed by subsequent investi- 
fators until Strasburger’s exhaustive work appeared in 1877.73 
twas not until then that the fact was generally recognized that 
Wo nuclei are to be found sooner or later in the development of 
‘very angiospermous microspore. This author was also able to 
‘emonstrate that the larger of these two nuclei is to be con- 
‘dered as a vegetative or prothallial nucleus and the smaller a 
| eacrative nucleus.** Strasburger found also that a second 
tivision takes place regularly in angiosperms, either in the spore 
"Self or in the pollen tube just before fertilization. Two sperm 
_ bells ada thus formed from the one generative cell. 

& Since that time many investigators working upon widely 
ferent Plants have found two nuclei, and these observations 
_ fad to one result, namely the confirmation of Strasburger’s 
ations in every essential particular. The time of forma- 
"2, ultimate Shape of the generative cell, and the time when 
latter divides were indeed not always the same in different 
“ats; on the contrary, all degrees of variation were found, 
om of Which are noted below. 

5 ag Untersuchungen aus der physiologische Lab. Land. Lehrung. Ber- 
* Sarsten 3: 294. 1866. 


Befruchtung und Zellth 
Befry 


y 
« : eilung 18, 1877. f 
entungs Vorginge bei den Phanerogamen 5, 1884. 


352 BOTANICAL GAZETTE [NOVEMBER 


The microspores of Potamogeton become separate immedi- 
ately after the second division of the mother-cell.. The anthers 
are at this time still quite small, the subsequent growth being in 
reality for the purpose of accommodating the increase in size of 
the pollen grains. The microspores at first have a thin, although 
distinct and homogeneous cell wall surrounding the cytoplasm, 
and a very large nucleus. The latter fills at least one fourth of 
the whole cavity of the cell (fig. go). The limited amount of 
cytoplasm present at this time is decidedly much more homo- 
geneous than in the mature pollen, and stains with the gentian- 
violet a uniform pale violet similar to that of the cell wall. For 
a short time after the wall of the mother-cell disintegrates the 
pollen grains are still held together by the remains of these 
walls. In fact they are as if imbedded in a ground mass of 
some viscid matter. 

The young grains very soon begin to increase in size, but 
the cytoplasm does not keep pace. As a result, the latter at 
length is confined to the parietal layer, but with a considerable 
increase in thickness on the side where the nucleus is located. 
These stages occur when the embryo-sac is yet one-celled, and 
of course while the spike of flowers is still enclosed within the 
bud. 

Just before the nucleus begins to prepare for div 
the following conditions: The cytoplasm is decidedly “cay 
granular, and stains more deeply with the orange. The large 
vesicular nucleus possesses a very distinct membrane. ce ie 
close against this is the linin thread which is rather extenst" 
for Potamogeton. The thread however is nearly destitute ° 
chromatin. The nucleolar-like body is smaller than usual, af 
in some cases more than one may occur, tare 

The first division of the microsporial nucleus takes P : 
much earlier than in Convallaria, and while the whole je 
yet enclosed in the bud. The spores reach their full size 
the division, and it is at this time that the exine first ae 
show signs of the thickening which produces the very gl 
roughened surface of the mature spore. Owing to me 


ision we find 


1899] DEVELOPMENT OF THE MICROSPORANGIUM 353 


aumber of chromosomes present, the spindles in the pollen grain 
ae exceedingly minute and slender (fig. gz). Fhe process of 
division here does not seem to present any new features. When 
preparing for division, the primary nucleus moves toward one 
side of the cell, so that the resulting spindle has one pole in con- 
tact with the cell wall. This pole unlike the free one, is not 
pointed; on the contrary, it is usually quite broad, so that the 
spindle fibers are attached to the wall over a considerable area. 
The spindle is quite dense and stains readily, but is composed 
olfew fibers. After the chromosomes pass to the poles a dis- 
tinct cell-plate forms, and is later followed by a definite 
| Membrane (fig. 42). The latter is arched in such a way as to 
cut off one daughter nucleus in a small lenticular cell, of which 
oe wall is the wall of the spore itself. This is the so-called 
seerative cell, 

The cytoplasm henceforth occupies the greater part of 
Be cell cavity. It gradually becomes filled with large bodies 
/Mhich stain purple with gentian-violet, and blue with iodine. 
‘y are in reality starch grains. A similar occurrence of 


| in in the pollen grain has been described in Naias by Camp- 
fe 


Slsely attached to the wall. The elongation produces an 
‘Mong cell, and is the first step in preparation for the second 
Ot. The chromosomes for this division are formed early. 
They ‘an often be seen to occupy nearly the entire nuclear 
“vity, and So distinct are they in many cases that one may 
om them. The number here again is uniformly seven. The 
; a were found in considerable numbers, one of ge : 
“tly aa 43. The spindle fibers are very coarse an — 

7 arcely more numerous than the chromosomes. Gu 
tan 'o stain more easily than is ordinarily the case in this 
a. The chromosomes are during this division seeniey 


% s 
Prog, Calif, Acad. Sci. III. +: 16, 1897. 


354 BOTANICAL GAZETTE [NOVEMBER 


The cell plate is soon deposited, and divides the generative 
cell into two parts. The two daughter cells do not separate, 
but. remain connected as a two-celled body during their entire 
stay in the spore. A more or less prominent constriction often 
occurs at the middle, but this does not seem to be constant 
(fig. 44). The pollen grain is always somewhat flattened, and 
since the generative nucleus is usually adjacent to the flat side, 
it is not possible to tell whether the latter remains attached to 
the wall after division. The difficulty was increased since the 
pollen escapes from the anther very soon after the second 
division. 

An examination of the literature relating to the pollen grain 
of the monocotyledons furnishes some interesting facts. Stras- 
burger found the division of the generative cell to take place 
within the spore only in Juncus and Arum; while in all other 
cases the division was in the tube. All these cases belong 
either to the Liliaceew, Orchidacee, Amaryllidacee, or Iridacez. 
Schaffner found the division occurring in the spores of Typha, 
Alisma, and Sagittaria,?7 and Campbell found the same to be the 
case in Naias, while the writer finds the same phenomena ' 
Acorus and Potamogeton. In all cases among the monocotyle- 
dons, where division occurs in the pollen grain, with the — 
tion of Alisma and Sagittaria, the generative cell is - vai 
enclosed by a wall, and always becomes two-celled after cen 
although Campbell claims that the two cells in Naias Ga 
before passing into the tube. Schaffner was not able to sees : 
any walls around the generative cell in the two above a oe 
species. From this it appears that the division of the et 
nucleus in the tube is mostly confined to the ee 
orchidaceous groups among the monocotyledons, while t os 
sion within the spore characterizes the spadiceous and 0 
ceous groups. 

*° Befruchtungsvorginge bei den Phanerogamen 22. 1884. 


77 A contribution to the life history of Seg#tlaria variabilis. BOT: 
1897. 


1899] DEVELOPMENT OF THE MICROSPORANGIUM 355 


SUMMARY. 

The following brief summary may aid in bringing together 
the results reached in the foregoing pages. 

The experiments with regard to the effect of external condi- 
tions on nuclear division both in Convallaria and Potamogeton 
gave no results for light and humidity, which were the only con- 
ditions tested. 

The material illustrating the younger stages in the develop- 
ment of the microsporangium shows that the process is slightly 
different in Convallaria and Potamogeton from the normal 
method as given by Warming and Engler. The archesporial 
tells arise by the division of a hypodermal cell at one corner of 
the anther. Therefore, instead of the archesporium arising from 
alayer of hypodermal cells, as Warming describes for dicotyle- 
dons, it arises from one or rarely two hypodermal cells. The 
primary archesporial cells divide only a few times, but there is 
‘siderable subsequent growth in size of each cell. The next 
outer cell in the original row forms part of the tapetum, and the 
tmainder are wall cells. Most of the wall and tapetum, how- 
"Wer, is formed from the tissue at either side of the archesporium 


| “amine seem to show that this is more likely the normal pro- 
Die for the whole group. 

: The tapetal nuclei of Convallaria show nicely the process of 
pear fusion which has been described by Strasburger and 
: for many other plants. After the division of the primary 
a nucleus by the mitotic method, the two daughter nucle! 
Fi many cases fuse again, and all stages of the process may be 
: a piten in the same anther. It is probable that not all the 
% divide, and also that not all of those that do divide fuse 
“ae before disintegration. It seems that in Potamogeton no 
pet the tapetal nuclei takes place. 


© structure of the wall in the mature microsporangium 


356 BOTANICAL GAZETTE [NOVEMBER 


was found to agree with the monocotyledonous type in general. 
The sequence in development was centrifugal and resulted ina 
well-defined endothecial layer, together with two or three inner 
wall layers in addition to the epidermis. At maturity only the 
epidermis and endothecial layer are present. 

The development of the archesporial nucleus shows some 
very important features. The coatracted condition called syn- 
apsis is without doubt a normal process accompanied by 
radical changes in the chromatin thread. The latter, which 
before synapsis was in the form of a network in which were 
imbedded large irregular chromatin masses, after synapsis is 
thicker, coil-like, and with the chromatin in smaller more equal 
masses. The spirem therefore begins at the close of the synap- 
sis stage. In both plants studied irregular dark masses were 
apparently expelled from the chromatin thread at the close of the 
synapsis, and in Potamogeton at least it was plainly evident that 
these had no connection whatever with the nucleolus. The ult 
mate fate of this chromatin-like matter was not determined. 
Whether this phenomenon was artificial or natural could not be 
determined from the material at hand. 

The growth and segmentation of the spirem in Convallaria 
almost identical with that in Lilium as described by oe. 
The longitudinal splitting of the ribbon is especially noticeable. 
In Potamogeton the process is different but could not be morse 
out satisfactorily owing to the minute size of the nuclei. * _ 
plant sixteen chromatin masses were counted just before division, 
but later there were only about seven, seeming to indicate a fusion 
of the widely separated primary segments to form the pn 
somes. The number of chromosomes after reduction was ©18 
een in Convallaria and seven in Potamogeton. This last nu™ 
ber is one of the smallest so far recorded for the phaser 

Spindle formation in all three plants agrees in every eee 
particular with the process described by Strasburget and pee 
for Lilium. The multipolar condition was evident in cace 
but was less distinct in Convallaria. 

The splitting of the chromosomes in the heteroty 


a is 


1899] DEVELOPMENT OF THE MICROSPORANGIUM 357 


| 
| 
: 
| 
| 


was in Convallaria exactly similar to that in Lilium. All the 
stages were especially clear. The + formation, however, began 
insome cases during the early multipolar condition. The pro- 
ves in Potamogeton was probably also normal. 

Nothing new could be determined in regard to the segmenta- 
tion during the second division. It seemed to be absolutely 
impossible to determine in these plants with any degree of 
cetainty whether the division was transverse or longitudinal. 
All the phenomena, however, seemed to indicate a transverse 
‘father than a longitudinal division in both plants. 

The walls of the mother-cells in Potamogeton were thin as in 
Naias and Zannichellia. 

In the microspores of Convallaria the generative nucleus is 
‘Yey chromatic, and is cut off by a distinct wall, but does not 
‘ecome detached from the wall of the spore until just previous 
‘0 the time of passing into the tube. The division of the nucleus 
‘Sprobably in the tube since it was not found within the spore. 
As in Convallaria, so also in Potamogeton the generative cell is 
tut off very early, but in the latter plant the two sperm cells are 
‘mediately formed. The two male nuclei are inclosed each 
Within its own cell wall, but they both still remain attached to 
& wall of the spore. The two-celled body then passes down 
‘te tube and even into the egg without separation of the two 
‘ils. The spindles in each case are very small and the chromo- 
Somes very minute, 


Corner, University. 


EXPLANATION OF PLATES XXIV-XXV. 


: PLATE XXIV. Convallarta majalis L, 
tr 4 '. One of the angles of a young anther in cross-section, showing the 
‘ * cells derived from the primary hypodermal cell; the two inner are 
eB “S archesporial cells ; the third will form part of the tapetum. 

Ee fig, : : ; : 
as 2. An anther cell at a later stage; the two archesporial cells in jig. 
oo 


tls at Vided several times forming a small mass of tissue; the row of wall 
— ‘op is probably equivalent to that in fig. 7, the tapetum is not yet 


358 BOTANICAL GAZETTE [ Novemser 


Fig. 3. A section of the anther wall at a still later stage ; on the extreme 
outside is the epidermis; then two layers the inner one of which will later 
disintegrate ; the fourth layer is the tapetum the cells of which often contain 
two nuclei ; and farther inside are shown a few archesporial cells. 

Fic. 4. A portion of the anther wall at nearly the same stage as in fig. }, 
showing the division of the tapetal nucleus. 

Fic. 5. A slightly later stage of the same in which the daughter nuclei 
formed in fg. ¢ are in the process of fusion. 

Fic. 6. A portion of the mature anther wall; the outer layer is the 
epidermis, the next below is the endothecium with spiral markings on the 
walls; and farther inside a disorganized mass composed of the tapetum and 
the one or two layers just outside. 

Fic. 7. An archesporial nucleus in the resting stage ; the linin network 
contains granules of chromatin of various sizes. 

Fic. 8. Synapsis, the projection at the left is the nucleolus. 

Fic. 9. Last stage of synapsis ; the spirem ribbon is opening out, and 
between its meshes are chromatin masses of various sizes ; the nucleolus at 
the left. 

Fic. 10, The spirem ribbon with chromatin granules imbedded in the 
linin. 

Fic. 11. The spirem ribbon after longitudinal segmentation, and cut in 
lengths by the section knife. 

1G. 12. Chromosomes still showing the \double nature; dissolution “ 
the nuclear membrane. 

Fic. 13. The multipolar spindle with chromosomes in various views. 

Fic. 14. The bipolar spindle and nuclear plate; the chromosomes 
the end appear -+-shaped, from the side view more elongated. 

Fic. 15. The + separates into v-shaped segments which are seen -_ 
ing toward the pole. 

Fic. 16. During the resting stage before the second division, 
viewed from the pole; the chromosomes remain distinct. 

Fig. 17. The nuclear plate of the reducing division ; 
so numerous that their form can be determined only with difficulty. 

Fic. 18. A chromosome on the nuclear plate, side view. 

Fig. 19. Pole view of a chromosome which is bent y-shaped. 

Fic. 20. End view of fg. 78. 

Fic. 21. A rare case, where fig. 20 has opened out along ¢ 
forming a ring. 

FiG. 22. v-shaped segments ready to pass to the poles. 


from 


the nucleus 


the segments ar 


Ny 
i 
s 
> 
: 
N 
S 
S 
= 
iS 
: 
N 
S 
& 


= 
be 
gy 
NS 
te 
a, 


BOTANICAL GAZETTE, XXVIT 


1899] DEVELOPMENT OF THE MICROSPORANGIUM 359 


Fic. 23. The microspore before the division of the primary nucleus. 

Fig. 24. Same after the generative cell has been cut off ; the generative 
wucleus is very dense. 

PLATE XXV. Potamogeton foltosus Raf. 
| Fic. 25. The row of cells derived from the primary hypodermal cell at 
} ach corner of the young anther. 
Fig. 26. A portion of the anther wall in a later stage; the outer layer is 
the epidermis, the second becomes the endothecium, the third disintegrates, 
the fourth is the tapetum, and farther inside are some archesporial cells. 
Fig. 27, The structure of the mature anther wall; the endothecium with 
isindistinctly spiral markings is limited on the outside by the epidermis, and 
mithin bythe disintegrated wall-cells and tapetum. 
Fig. 28. An archesporial nucleus in the resting stage; the indistinct 
{nin network contains a few pale-staining chromatin granules ; the large dark 
ass is the nucleolar-like body ; and the black bud at the side is possibly the 
tucleolus, 
Fig. 29, Synapsis; there is very little chromatin in the contracted mass. 
‘Fig, 30. Later stage of synapsis; granules are expelled and the linin is 
‘ginning to spread out. 
Fig. 31. The spirem; contains very little true chromatin. 
la The nucleolar body has disappeared and at the same time the 
€s were formed ; the membrane is here spreading out. 
Fig. 33. The multipolar spindle and chromosomes. 
Fig. 34. Same becoming bipolar. 
Ube 35. The bipolar spindle and nuclear plate; the chromosomes ase 
be double, 

Fig. 36. Daughter nuclei after the first division; the chromosomes 
“ain distinct, ; 
- The second or reducing division, nuclear plate stage 
- Same showing the daughter segments moving toward the poles. 


- The reducing division ; chromosomes in the nuclear plate. 


A microspore with its primary nucleus. 
- Same showing the mitotic division of this nucleus. 
me .. with a free vegetative nucleus, and a small dense genera- 
oe ed by a convex wall. 
: 43. Same showing the division of the generative nucleus. 


IG, : 
Sie A mature microspore in which the two dense€ sperm nuclei ar€ 
| sed by a cell wall. | 


BRIEPER ARTICLES. 


AN HERMAPHRODITE GAMETOPHORE IN PREISSIA 
COMMUTATA..’ 


(WITH ONE FIGURE) 


THE reproductive organs of most of the Marchantiacez are borne on 
specialized receptacles known as gametophores. So far as I can learn 
there has been no instance recorded of the two sex organs being found 
onthe same gametophore. In a recent study of Preissia, however, I 
discovered this phenomenon. 

The normal archegoniophore of Preissia is hemispherical in shape, 
the archegonia being situated on the lower surface. The tissue 
adjacent to the archegonia is compact and is made up of small cells 
rich in protoplasm. The tissue in the upper part, however, is muc 
looser and the cells are larger. The antheridiophore is discoid, with 
the upper surface slightly concave, in which the antheridia are sunk. 
The tissue, particularly that surrounding the antheridia, is much looser 
than that of the female receptacle, and the cells are somewhat larger. 

The shape of the hermaphrodite gametophore from which the 
accompanying illustration was made clearly indicates that it is primarily 
an archegoniophore, but it is modified to adapt it to its peculiar condi- 
tions. The upper part is more strongly developed than normally, and 
is irregular in form. The archegoniophore at maturity has a long 
stalk, but as the material from which this section was made was put up 
in the fall the stalk had not yet elongated. On the under surface os 
_ Shown a portion of one archegonium, the neck having been cut off in 
sectioning. In the upper portion of the gametophore are two well- 
developed antheridia, corresponding almost exactly in shape and sizé 
to those occurring in normal plants. Their structure is also the same- 
Their position in the gametophore is very similar to that which they 

*Miss Townsend discovered this interesting case of an hermaphrodite ere 
phore while engaged in the regular course of advanced work in comparative morp’ : 
ogy, and it was at my suggestion that she has prepared it for publication.— GEO. ** 
ATKINSON. : 


360 ecient 


- thep] BRIEFER ARTICLES 361 


- occupy in the normal antheridiophore. The tissue surrounding them 
resembles that of the latter receptacle, while the tissue of the lower 
portion is that of the ordinary archegoniophore. 
There is a question whether this is merely an abnormal condition 
o whether it is a reversion to an earlier type. In the Ricciacee the 
_ two organs may either be produced together or, as is often the case with 
Riwia glauca and Riccia hirta, they may be found on the same part of 


4 
POL ) 


Q 
Y 


ASU Or 
Ny, 
We 


Reet 
oe” 
sean 
ata 
Ra 


by 
ae 
84 


CTT Sarees eS Ws et, LER, Wik 
Ra < DOR pn 
i coome ee en Se LF. 
Sei 


5 
oa 


eee 
or 
Lo 
on 
i 
2e5 
Bae 
gece 


{? 


ies 

ue 
Qos 
arts 
Ns 
S 


erm 


Fic. 1.—Section of hermaphrodite gametophore of Preissia commutata. 


e organ being produced 
of the 


“ed and As the reproductive organs became 
devel Specialized, the gametophore might natura 
ott and specialization, and in time have co 


Me 
# 


362 BOTANICAL GAZETTE | NOVEMBER 


to the functions of the organ borne onit. A suggestive case inthe alge _ 


occurs in Vaucheria geminata and V. terrestris, where an unspecialized 
branch, or gametophore, bears both antheridia and oogonia. It is pos- 
sible that the original type of gametophore was hermaphrodite. In 
Vaucheria terrestris there are occasional branches which bear but one 
organ, showing that a unisexual gametophore might be developed 
from the hermaphroditic.— ANNE B. TowNsEND, Cornell Oniversity. 


SOME PLANTS RECENTLY INTRODUCED INTO FLORIDA. 


Two years since? I called attention to some South American species 
found by Mr. A. H. Curtiss in ballast vr about streets in various parts 
of Florida. A package of specimens recently sent for study by Mr. 
Curtiss contains several South American and southwestern species 
apparently not before reported from Florida, most of them unrecorded 
from the North American continent. The plants from Pensacola were 
collected, in the words of Mr. Curtiss, “from lands lying inside the 
wharves, such as presumably consist more or less of ballast earth.” 
This fact suggests that Jusstaea suffruticosa, and Hydrocotyle bonar we 
formerly reported from Pensacola, are perhaps introductions to be 
classed with Alternanthera pungens, Solanum cleagnifolium, and An- 
themis mixta (recorded in the article referred to), and that they have 
reached the Florida coast through the same channel as the following 
species : 

Ipom@aA PALMATA Forsk. Fl. Aigypt. Arab. 43.—This Egypia® 
plant has been recently introduced into Florida where it has made 
itself at home, growing vigorously in waste ground about St. Augus: 
tine, Jacksonville, and Pensacola (no. 6496). At Pensacola Mr. 
Curtiss states “I found it growing rampant over bushes on the bay 
shore and fruiting freely.” } 

SOLANUM GLAUCUM Dunal in DC. Prodr. 13°: 
species which was first noted at Pensacola in 1897 whe 
abundant, though now thoroughly at home (no. 653°) 

SALPICHROA RHOMBOIDEA Miers in Hook. Lond. Jo 
1845.—This delicate solanaceous plant, native of Argenti 
was recently collected by Mr. Curtiss ( Oct. 3, 1899) im 
at Jacksonville (no. 6542). 


100.—A_ Brazilian 
n it was not 


ur. Bot. 4: 326 


ne Republic, 
low ground 


? Bot. GAZ. 24: 433-436. 1897. 


{ 
{ 
| 
| 


1809] BRIEFER ARTICLES 363 


_~ Asrer pivaricatus Torr. & Gray, var. GRAMINIFOLIUS Baker in 
- Martius, Fl. Bras. 63: 22.—This common South American form has 
heen found for three years about wharves at Pensacola where it seems 
permanently established (no. 6497). 

| ERIGERON BONARIENSIS L. Sp. Pl. 2: 863. 1753.—Growing luxuri- 
-atly(6 or 7 feet high) about the wharves at Pensacola (no. 6499 : 
- Introduced from South America. 

_ Pascatia GLauca Orteg. Hort. Matr. Dec. 39. This Chilian species 
has been introduced within a year or two at Pensacola (no. 6492 ). 
PECTIS PROSTRATA Cav. Ic. 4: 12. pl. 324. Very recently intro- 
duced at Pensacola (no. 6531) : abundant from the southwestern states 
South through Mexico.—M. L. FERNALD, Gray Herbarium. 


CURRENT LITERATURE. 
MINOR NOTICES. 


LIEFERUNGEN 190-192 of Engler and Prantl’s Pflanzenfamilien have 
recently been published. They are devoted to a continuation of the Poly- 
podiacez by L. Diels.—C. R. B. 


THE PAPERS of botanical interest printed in abstract or in full in the 
Proceedings of the Indiana Academy of Science for 1898 (published 1899) 
are as the following. THomAS: Some desmids of Crawfordsville; MOTTIER: 
Nuclear division in vegetative cells; The centrosome in cells of the gameto- 
phyte of Marchantia; Endosperm haustoria in Lz/ium candidum ; RISLEY: 
Absorption of water by decorticated stems; ARTHUR: Indiana plant rusts, 
listed in accordance with latest nomenclature; SNYDER: The Uredinew of 
Madison and Noble counties, with additional specimens from Tippecanoe 
county; GOLDEN: Asfergillus oryz@,; CurTIss: A red mold; OLIVE: Affi- 
nities of the Mycetozoa; CUNNINGHAM: Morphological characters of the 
scales of Cuscuta; CouLTER: Notes on the germination and seedlings of 
certain native plants; BRANNON: Some Indiana mildews.—C. R. B. 


STATISTICAL methods have come into greater prominence « see 
study during recent years. Dr. Charles B. Davenport has prepared a use ; 
little handbook,* in which, after some preliminary definitions, he sets at 
the proper methods of measuring and counting organisms, of 
plotting of data, describes the constants of plotted curves and probable 
in their determination, and enumerates the classes of plotted curves: | 
chapter is devoted to correlated variability and the methods of peers: 
the degree of correlation and heredity. Galton’s, Pearson’s and Dunc 
methods of determining the coefficient of correlation are given. ae 7 
rules, and ten tables useful for the various calculations make up the 
the handy volume. , r 

A short chapter on the applications of statistical biological study ee 
remark. That these methods are of great value for a study of pasion ; 
heredity admits of no doubt. That they will improve our rege toe 
species and varieties is not at all clear. That “by the use 0° | 
method biology will pass from the field of the speculative sciences name 
the exact sciences” is surely a vain hope. We shall see the ai ical 

*DAveENporT, C. B.: Statistical methods with special reference M og 
variation. I2mo. pp. viiit-148, figs. 28. New York: John Wiley and 
$1.25. : [ NOVEMBER 

304 


eS ee eee On Ae ee en an ee 


ST ee eC ee ee 


1809] CURRENT LITERATURE 365 


swing far toward the side of quantitati 
titat : 
nearer equilibrium.—C. R. B i ive study and then back toa point 


T 
.. va poanieny issues as one of its bulletins the 
MR ie 4 drime e book, by the Chief of the Division of Forestry 
igaple Be csre (1) rv of forestry2 (n four chapters Mr. Pinchot presents 
Gy individual: (2) the a account of the sinstae and activities of a tree as 
oe in light ia ations of trees in @ forest, including their require- 
discussing the origin of si and their reproduction ; (3) the life of a forest, 
a e sete the struggle between the trees, their death, 
Be teowsing, and Suiee ; (4) the enemies of the forest, including man, 
fre ’ roping animals, insects and fungi, wind, snow, and 


‘The : ‘ 

 eaienames ela called for and is an excellent introduction 
og accurat reading. The physiology and anatomy are very ele- 
figures and plates whi : as such general statements can be. The numerous 
ee ee we almost all half tones from photographs, consti- 
might be spared ares: so much overdone, however, that half of them 

pared, as far as the text is concerned.—C. R. B. 

National Herba- 


THE LA 
: st of the Contributions from the United States 
e of the series. 


tium (x: 
It a 78-64. 1899) is one of the most notabl 
tal American flora ie i the most recent studies upon the Me 
} 40d in the “Sead J. N. Rose, whose untiring labor both in the field 
Mexican and  salean oak resulted in large accessions to our knowledge of 
Oe batts, vis. taxc merican plants. The Contribution consists of two dis- 
} 4 notes on eel ra studies, illustrated by text cuts and ten plates ; 
q a DB iice plants of Mexico, illustrated by thirty-seven plates of 

e : 

ee in viich studies are as follows: a rearrangement of the suborder 
| Mopsis of the Santtge genera are recognized, Pseudobravea being new; 
of them new: n merican species of Nissolia, including twelve species, 
‘Rotes on Pies -_ on Rutacez, with two new species of Xanthoxylum ; 
of Clitoria, wit ez, with a new species of Turnera notes on Mexican species 
Mbaces, with descripti notes on Malvacee and 
0G new oe of ten new species; notes on Passiflora, with 
Neluding ten s Sc bae oes of the North American species of Waltheria, 
s pecies, three of them new; notes on some Mexican species of 
anish cedar, with descrip- 


xican and Cen- 


h ee : 
descriptions of two new. species; 


a m, : 

7 tions of — of them being new; Cedrela, or Sp 

| Pecies bein ea Species; notes on new or rare Leguminose®, 
| Peing described; descriptions of twelve miscellaneous ne 

The forest. 12M» pP- 88, 


Ww species. 


*PINC 
HOT, Gir : 
,GIrForD: A primer of forestry. Part I. 


aan 
.& 4, figs, & j 
a 3 Washington: Dept. of Agric. 1899- 


/ 


366 BOTANICAL GAZETTE [NOVEMBER 


In addition to these Mexican studies a new genus of Commelinacez, Tre/easea, 
is established with three species, to include certain Texan and Mexican forms 
heretofore referred to Tradescantia; and three new species of Tradescantia 
from the United States are described. A new genus of Umbellifere from 
Mt. Ranier, Washington, Hesferogenia by name, is described by Coulter and 
Rose; and Mr. L. F. Henderson describes a new Aster and a new Angelica 
from Idaho. 

The part devoted to a description of the useful plants of Mexico is based 
upon the personal observations of Dr. Rose during a visit of four months in 
the summer of 1897. It is full of interesting information and photographic 
illustrations, and is very suggestive of lines of economic investigation.— 


NOTES FOR STUDENTS 
M. J. GOLDBERG’S experiments lead him to the conclusion that during 
the germination of wheat in darkness proteid substances are produced in 
the embryo in considerable quantity,3 although Godlewski in 1897 thought 
this to be impossible.s—C. R. B. 


COPELAND AND KAHLENBERG, by a series of carefully conducted experi- 
ments show that the injury to plants from solutions of pure metals (Nagell’s 
oligodynamic effect) is due to the toxicity of the compounds (salts) which the 
dissolved metals form and not to any peculiar or toxic action of the elemental 
metal.s—C, R. B 


M. W. PAaLLapiNE has determined that alternations of temperature 


accelerate the respiration of severed tips of etiolated shoots of Vicia Faba 
cultivated in 10 per cent. cane sugar. The increased energy of seapiree 
does not depend on the quantity of active nitrogenous foods, but the 7 
Cause is not yet determined.*—C. R. B. 

R J. W. HARSHBERGER has observed a distinct a curvature 
of leaf blade and petiole in Rhododendron maximum 1.7 cold the blades 
are revolute and the petiole arcuate downwards. On meen a beat 


a warm room erection and flattening were complete within five minutes. 4 
curvatures in a reverse direction are slower. Turgor variations are 
cause.—C. R. B 

(Proc. 


ITEMS OF TAXONOMIC INTEREST are as follows; GERRITT S. MILLER um 
Biol. Soc. Wash. 13: 79-90. 1899) has discussed the species of ieee 

3Rev. gen. de Bot. 11: 337-340. 1899. 

* Anzeiger Akad. Wiss. Krakau, March 1897, fide Goldberg. 

5 Trans. Wis. Acad. of Sci. 12: 454-474. 1899. 

° Revue gen. de Bot. 11: 241-257. 1899. 

7 Proc. Phila. Acad. Sci. 1899: 219-224. fig. 3. 


1899 | CURRENT LITERATURE 367 


inthe District of Columbia, recognizing seven, three of them being described 
asnew, two having been recently described by Professor E. C. Greene, and 
the remaining two being the well-known species of Linnaeus.— WILLIAM 
PALMER (Proc. Biol. Soc. Wash. 13: 61-70. 1899) has published a list of the 
ferns of the Dismal Swamp, Virginia, sixteen in number, one of them being 
described as a new variety.— J. N. Rose (11th Ann. Rep. Mo. Bot. Gard. 1- 
j. 1899) has described a new species of Agave and critical notes on other 
species, accompanied by four plates.— E. P. BICKNELL in his further studies 
of Sisyrinchium (Bull. Torr. Bot. Club 26: 335-349, 445-457, 496-499. 1899) 
added eleven new species to the already long list of forms.— AVEN 
NELSON in continuing his publication of new plants from Wyoming (#did. 
350-358, 480-487) describes twenty-three new species, one of which repre- 
sents a new genus, WVacrea, related to Anaphalis —C. L. PoLLarD (zdid. 365- 
372) has revised the genus Achillea in North America, recognizing ten species, 
thtee of which are new.— K. M. WIEGAND (2ézd. 399-422) presents ten species 
of Bidens found in the United States and Canada, describing one new species 
ad five new varieties ANNA M. VAIL in continuing her studies of Ascle- 
Pladaceze (2béd. 423-431) discusses the types of Gonolobus and describes 
three new Species of Vincetoxicum.—P. A. RYDBERG (zbid. 541-546) has 
described twelve miscellaneous new species from the western United States. 
~A. A. HELLER in continuing the publication of his new and interesting 
ag from western North America (zbzd. 547-552) describes ten new species, 
_ Our of which are species of Mertensia.— J. M. C. 
SOME CURIOUS experiments by A, Pagnoul® on transpiration are reported 
the Experiment Station Record 11: 118. 1899. They are difficult to 


oa without more light than the brief summary gives ; but to call atten- 


in 


Pt first period, 33 days, the plants in the poor soil t 

_ ater per gram of dry weight; as compared with a transpiration of 5 5 ba 
Water In the second period the figures were 1053 and cae : 
F Stam of dry weight, and for the last period 1084 and 555°; 

h pot was deter- 
he product of the 


and it was found that for each gram of nitrogen int 
18™ of nitrogen 


of water was transpired, while in the rich soil 
€ach kilogram of water given off. 


ay l 
mh Sta. Agron, Pas de Calais 1898: 10-15. fg: J. 


NEWS. 


PROFESSOR DR. FRANZ VON HOHNEL has left Vienna on a journey to 
Brazil. 


SYNTHETIC cassia oil, introduced by Schimmel & Co. of Leipzig, is gradu- 
_ ally displacing the ordinary cassia oil of commerce. 


A PLANT of Cypripedium insigne giganteum from the collection of the 
late Major Mason, recently sold in London by auction, brought £147, and 
other varieties from £30 to £76. The total amount realized at the sale was 
over £3000. 


THE HERBARIUM of the Geological Survey of Canada has recently ray 
enriched by Professor Macoun’s collections from Sable Island, by sie A. ; 
Low’s from northern Labrador and the coasts and islands of Hudson's adi 
by Mr. J. B. Tyrrell’s from the Yukon district ; and by Mr. N. B. Sanson’ 
from the vicinity of Banff. 


A course for teachers, consisting of studies in ecology and plant spss 
ology with excursions and practice, was conducted during the autumn Rie 
fessor C. Stuart Gager, of the New York State Normal College at Albany: 
The syllabus shows a refreshing appreciation of the modern aspect of ag 
“ considering plants from a dynamic rather than a static pomt of view. 


THE ACREAGE in peppermint in Wayne county, New York, one 
chief centers of the crop, has fallen from 3340 acres in 1890 to Me : 
on account of low prices of the oil and the better returns to farm a 
growing sugar beets. In Michigan the exports of peppermint on ae: am 
80,225 lbs. in 1894 to 162,492 lbs. in 1897, but are now falling, Be ‘i 
Ibs. in 1898. 


WITH THE CLOSE of volume II Messrs. Willard N. Clute & Co. ¢ 
publish Tze Plant World. \t will hereafter be publish umber © 
World Company, 321 4% street, Washington, D.C. The: first’? october f 
volume III will be that for January 1goo, thus leaving a gap prs editorial 
to December 1899 inclusive. The journal still remains under t fe jhe 
direction of Dr. F. H. Knowlton. The new volume will be see tion pri . 
trated and nearly doubled in size, but with the same page iler success 
The journal has proved very useful, and we bespeak for it mua _ 
it deserves, 


368 [wovensin 19? 


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Vill DECEMBER 1899 _ No. 6 


THE 


EDITORS 

M. COULTER, Zhe University of Chicago, Chicago, Ill 
CHARLES R. BARNES, Zhe University of Chicago, Chicago, Ill 
J. C. ARTHUR, Purdue University, Lafayette, Ind. 


ASSOCIATE EDITORS 


MIR DeCANDOLLE FRITZ NOL 
Geneva Ceri Ke Bonn 
ONT VOLNEY M. SPAL na 
University of Padua Un ert 3 F Michigan 
ENG ROLAND THAXTE 
University of Berlin Harvard icaect 
VIGNARD WILLIAM TRELEASE Zs 
LE tole de Pharmacie Parts Mi rig ze sie Garden % 
A. HARPER H. MARSHALL 
<i 3. of Wisconsin Univer ra 
SUMURA EUGEN. WARMIN ah 
aes University, Tokyo ew of Copenhagen 


VEIT WITTROCK 
Royal Academy of Sciences, Stockholm 


CHICAGO, ILLINOIS 
“Pudtisne bp the Gnibversity of Chicago 


Che Aniversityp of Chicage press 


COPYRIGHT 1899 BY THE UNIVERSITY OF CHICAGO 


ae Cambridge Ae 


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Hotanical Gazette 


 Montbly Journal Embracing all Departments of Botanical Science 
n per year, $4.00 Single Numbers, 40 Cents 


subscription price must be paid in advance. No numbers are sent we the expiration 
of the time paid for. No reduction is made to dealers or agen 


FOREIGN AGENTS: 


fain— Wm. WESLEY & Son, 28 Essex Germany — GEBRUDER BoRNTRAEGER, Berlin 
on don 18 Shillings. SW. 46, consincaceaas 17a. 18 Marks 


XVI No.6 | - Issued January 10, 1900 


CONTENTS 


SKY MOUNTAIN CHRYSOTHAMNI. Aven Nelson : 369 
OXIG EFFECT OF DELETERIOUS AGENTS ON THE GERMINATION AND 

FELOPMENT OF CERTAIN FILAMENTOUS FUNGI. (Concluded). J. F. Clark 478 
MN CRATAGUS. I. C.D. Beade - y : : : . - 405 


b Pecurariries IN PUCCINIA TELEUTOSPORES Pi six FIGURES). H. Harold Hume 418 
Tis Prunus Insrrit1a? P. A. Rydberg - fe 
ON THOREA (WITH PLATE xxvI). George G. Hedgcock and Abel A. Hunter - 425 
ORE ON Corn Smut. A. S. Hitchcock - ' : Me 
OTANICAL ART GALLERY. Conway MacMillan - - : . ~ 430 


C. W. Hyams - - - - - - 431 
LITERATURE. 
VIEWS é ‘ - 3 = " . 432 
PROPAGATION OF MOSSES. 
OR STUDENTS - : ‘ : - - - - - 435 
i * - : ; ‘ 5 ; ; - - - 444 
of any article (not less than 50 copies) will be printed only upon special order in advan 
Twenty sha pies (witha cower) will be furnished graézs to authors of leading ptrien; 
wing ra 
Number of copies 50 100 150 — 
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€ (paper like Geberce cover)... $1.00 $1.50 $2.25 
pe a ae 


ith sone nigh 


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$ to follow the form shown in the pages of the GAZETTE. M 
The University of Chicago, Chicago, 
Pamphlets for Review should a sent to the same addre ue har eon kh 
“ae will be replaced free only when claim is made within thirty day: 


should be addressed to The University 


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; y Press Division, Chicago, Ill. 
de: and drafts se be made eae to bd lating iby 
Orric 


] 
(Enren 7 Cuicaco, ILL., A “ie eae eS 


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a7 3 rT a0) SE’ ko. LWMi¢,\Ss AR SS SS Ss 
ores (I eu Ss Sxl es sy = y= Sa > Kes mesa Tie eS SE. 
GSW us See DO mane es a as DO Leake 


My SS ZZ phy SSS ZSPN Meo SX, uy eet Vy 
A = (a>: Su mes on ESKEs ay ey Kes mS a= 
yy Fd LS BPD Lee Ds = Nae SBS Ps LL SS 408 eon 


Novelty in Literary and Art Features. 


OLIVER 
CROMWELL 


By the Right Hon. John Morley, M. P. 


Tne conductors of The Century take especial pleasure in an- 
nouncing Lae as bag leading historical serial of the magazine 
1900. Nom ore competent than John Morley, who was 
acted by Mr. “Gladstone's family to write the biography of Glad- 
stone, to treat sab ell in the spirit of the en \ 
if fj century. His w as a historian, as seen in the ne 

Edmund Burke poe Richard Cobden, is well known. 
Swine: it has movement, variety,— above all things, the strong pulsation of inner passion 


THE ILLUSTRATIONS OF THE CROMWELL HISTORY 
( will be — Besides original drawings by well-known artists, 
QZ lished portraits, permission to reproduce which has been given r Majesty the Que 
Universities or ‘Oxford and Cambridge, the British and South Lcunnetan Museums, etc 
: ERNE EST SETON-THOMPSON’S pict getce FROM MARK TWAIN'S 
‘« Biography of a Grizzly,” delightfully meni ANDONED AUTOBIOGRAPHY. 


an 

portant literary work of the author i cWild 

Animals I Have Known,” begins in November. ITERARY REMINISCENCES. ning 

paint accounts of Te 5 SON yhittier, ond 

PARIS, ILLUSTRATED BY CASTAIGNE. Lowel Saeet 

.A series of papers for the ee year, b en ee 

Richard Whiteing, author of ‘‘No. 5 John MILITARY PREPAREDNESS. 

sp : co as ws dly illustrated with more than By Theo yah Roosevelt. An imp 

pictures the st | wet i i 
including-views of the the Paris  Eapectiicn ie paces suggestions 


IES 
LONDON, ILLUSTRATED BY PHIL MAY. |p. prodinnt Wit ot ae 
by sir Walter Besa ty East End of London | Pomas Bailey Aldrich, Professo 
magn ay Srbcoh Perot ,» with pictures by Phil | \ijcon , John Burroughs, and othe: rs 


SAILING ALONE AROUND THE WORLD. AMERICAN SECRET HISTORY: 


11 ol Ae subscription with Novembey,— 
a tant number, beautifully printed in tint. It opens the 
a year, Subscribe rte age Ab pet agents or remit “Girectly to fo the 


VY, 
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The record of a voyage of 46,000 miles under- A series of papers of commanding interest. 
taken single- -handed and alone in a 40-foot boat NTURY: 
SNe F THE CE 
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Sees Cre reece Wes farish a short serial | has led the world in art. Timothy Cole’s vt 
of re cilickueie pay chological interest, “The Auto- | and beautiful wood blocks will continue f vet 
biography of a Quack. feature, with the work of many other eng 
eye w nave e the American scho A wood 
OTHER STORIES ane fine half-tone plates—reéngrate ie distio- WW 
include a short serial of California by Mary Hal- | engravers — for which the magazine f printing Ni) 
lock Foote, stories by Harry Stillwell Edwards, guished, will appear W with new met thods © 
Seumas MacManus, and many others. | and illustrating. 


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E XXVIII NUMBER 6 


i 
" 
i 


MTANICAL GsAZETTE 


i 


DECEMBER 1899 


SOME ROCKY MOUNTAIN CHRYSOTHAMNI, 
AVEN NELSON, 

\FTER an experience of several years as a collector and as a 
tof this group both in the field and in the herbarium, I 
teached the conclusion that the genus Chrysothamnus, as 
Sin the Rocky mountains, is still to an unsuspected degree 
Own and unappreciated. 

he plants of the far west became known to the botanical 
orld at an earlier day than those of the interior west, and it 
Sto have been assumed that the species of the two regions 
Ktobe the same. As a result of this assumption we have 
Toducing a confusion in the herbaria that will be cleared 
difficulty. Dr. Greene’s* papers on this genus teprescnt 
attempt to do justice to the species of the range in ques- 
In this genus, as in many others, Nuttall’s knowledge 
d in the field has been discounted, but, like other attempts 
rdinate his Species, it has simply resulted in confusion. 
following notes it is hoped may be serviceable not only 
ent students, but to all who are interested in this charac- 


| others believed to represent species not heretofore col- 
Only such species are included in this paper as belong 
“43392-96; 107-115. 1895. 

3 


370 BOTANICAL GAZETTE | DECEMBER 


to that natural botanical region of which Wyoming, with its high 
arid plains and mountain ranges, is the center. Localities men- 
tioned are in Wyoming unless otherwise stated. 

CHRYSOTHAMNUS GRAVEOLENS Nutt.— For the literature of 
this species and a thoroughly usable characterization see Erythea 

.3: 108. 1895. Its range as there given, and the habitat, ‘‘ denu- 
dated soils,” given by Nuttall, will prevent confusing with this 
certain other species that have been too often included. Thor- 
oughly typical are the following collections: George E. Oster- 
hout, New Windsor, Colo., September 8, 1898; the writer’s nos. 
503 and 2758 from the Platte cafion, in 1894 and 1896. 

CurysotHamnus PLatrensis Greene, Pitt. 4:42. 1899. C 
speciosus Plattensis Greene, Erythea 3: 111. 1895.—This is the 
representative in this range of the far northwestern C. spectosus 
Nutt.2_ I know of no specimens except from the eastern base of 
the Rockies. Well represented by the following collections: 
C. S. Crandall, Fort Collins, Colo., September 17, 1898; George 
E. Osterhout, New Windsor, Colo., September 8, 1898; E. L. 
Greene, LaSalle, Colo., September 10, 1896; the writer, Chey- 
enne, Wyo., August 27, 1896. 

Chrysothamnus pulcherrimus, n. sp.—Shrub 0.5- 
tree-like in form (a short trunk-like base much branched above); 
main stem and branches with grayish bark, the season's stems 
yellow but under the lens minutely lanate-puberulent, rather 
slender-virgate, terminating in an ample, compact, paniculate 
cluster: leaves moderately numerous, narrowly linear, rather 


‘=> high, 


lax and spreading, 5-8 long, from white-tomentose to green: | 


ish-glabrate: involucral bracts only two or three in each row, 
oblong, acute, nearly glabrous, ciliate on the margins: corolla 
tube longer than the throat into which it gradually ¢€ 
rather closely short-hairy : anthers well exserted, the append- 
ages of the style exceeding these: akene softly pubescent. 

A handsome shrub, its arboreous habit, numerous, slender pranchlets, 
large trusses of bright yellow flowers make it one of the strongly ore 
ized and conspicuous species. It is to be distinguished from C. graven" 
but I believe 


xpands, 


and 


2 This and its var. albicaulis is sometimes reported from this range, 
cannot be authenticated. 


1899] SOME ROCKY MOUNTAIN CHRYSOTHAMNI 371 


Nutt., to which it is allied on the one side, by its less glabrate condition, its 
narrower leaves, yellow stems, paniculate (not corymbose) inflorescence, and 
proportionately shorter style appendages. From C. sfeciosus Nutt., to which 
it must be compared on the other side, by its stout yellow (not whitish) 
branchlets, its longer less filiform leaves, glabrate involucral bracts, pubes- 
cent corolla tube, and more open-paniculate inflorescence. 

Under no. 2066 it has been distributed as C. Speciosus, from Cummins, 
August 10, 1896. Collected near the same locality (Wood’s Landing), 
August 9, 1897, no. 3477, the latter taken as the type number. Specimens 
from Gros Ventre river, August 23, 1894, no. 966, is a form with more virgate 
and glabrate branchlets. 

CuRYSOTHAMNUS PULCHERRIMUs fasciculatus, n. var.— size and 
habit of the species; the season’s branchlets short, numerous, 

forming a brush-like fascicle at the ends of the woody branches : 
leaves numerous, short (2-3), somewhat rigid, green or yel- 
lowish-green, with a thin tomentum: inflorescence similar to that 
of the Species but smaller, terminating the numerous branchlets. 

The fascicled branchlets and short rigid leaves suggest C. co/linus 
Greene, but the size, habit, inflorescence, and floral characters are those of 
C pulcherrimus. C. collinus, in its pubescent long acuminate bracts, is quite 
N contrast to this. 

First collected in 1894 on Boulder creek, August 26, no. 1120; in 1897 
at Creston, August 28, no. 4419 and distributed as C. sfeciosus. 

_ Curysornamnus FRIGIDUS Greene, 1. c.— What is typical of 
| this species is not quite clear (see note under var. concolor), but 
it seems plain that it was intended to include that superabundant 
but variable species of the high plains, popularly known as 
_“fabbit-brush.” It is illustrated by Dr. Greene’s specimens from 
famie, August 10, 1895; the writer’s nos. 2787, 5280, 5282, 
ay 
Curysornamnus FRIGIDUS concolor, n. var.— Yellowish-green 
| throughout except for the bright yellow flowers, 3-4°" high from 
. barely shrubby base, the numerous stems rather slenderly 
“itgate, terminating in a narrow thyrsus (sometimes peicete 
_“*tymbose): leaves linear, erect or irregularly spreading, eA 
6: t-2™ wide: bracts mostly acute, the outer with a light 
‘omentum like that of the stems and leaves, all with a thin, 
- atious-ciliate margin: corolla tube short-pubescent. 


>) 


372 BOTANICAL GAZETTE | DECEMBER 


frigidus so long as 1 am unable to say what is typical of the latter. Ever 
since its publication (E7ythea 3: 112. 1895), based upon specimens (one of 
which I have) collected at Laramie by Dr. Greene, I have collected freely in 
this genus. Though in the center of distribution for C. /rigidus, and in spite 
of the large series representing it, I have never secured duplicates of the 
specimens distributed by Dr. Greene, nor any quite typical, judged by the 
description. Assuming that the plant, so common on the Laramie plains, 
of which there seem to be several forms, must stand as C. frigidus, the 
preceding may at least be ranked as a variety, no matter what may be typical 
of the other. Nor may this be connected with C. Plattensis. Some forms of 
that seem to connect very closely with C. /rigzdus, but typical specimens of 
the two are really widely different. C. Plattensis one at once associates 
with C. sfeciosus on general appearance; not so with C. frigidus and the 
variety now proposed. 

Collected on the banks of Hutton lakes, in rather sandy, but more or less 
alkali impregnated soil, September 6, 1898, no. 5300. 

: dm 

Chrysothamnus pallidus, n. sp.—A small tufted shrub, 2-5 
high, with a close felted tomentum which persists even on the 
old stems: stems rather scraggy branched, somewhat rigid, the 
season’s twigs very short: leaves mostly confined to the season s 
growth, the tomentum looser than on the stems, ascending, irreg- 
ularly spreading or reflexed, linear, acute, 2-3™ long; shorter 
and crowded at the base of the inflorescence: heads small, in 
thyrsoid panicles or somewhat corymbosely clustered cymes; 
involucre short, sub-campanulate, its bracts short, oblong, sub- 
acute, ciliate-pubescent especially on the margins, about three 
in each row: tube of the corolla closely covered with short, 

: € 
clavellate hairs, the throat longer than the tube and cleft as 
third its length: style divisions exserted, the stigmatic asa 
about as long as the appendages: akenes densely pubescen 
about equaling the corolla tube. 

Allied to C. frigidus but distinguished by its more rigid, scrag 
which are leafy only on the branchlets ; by the close, persistent, 
ment; by the shorter bracts, florets, akenes ; by the exserted sty 
and the denser hairiness of the corolla tube. ie. Sep- 

Seemingly a rare plant; collected on an alkaline flat near aint a 
tember 24, 1898, no. 5347. Also on Bacon creek, August 15, 1894, B® 


dm hj xh, 
Chrysothamnus Wyomingensis, 1. sp.—Tufted, 2-4 ae 
bushy-branched from the base, the branches with asce? ing 


gy branches 
white indu- 
le divisions 


Ia 


1899] SOME ROCKY MOUNTAIN CHRYSOTHAMNI 373 


erect, yellowish-green branchlets, witha thin inconspicuous tomen- 
tum throughout: leaves rather numerous, especially above, 4- 
6" long, narrowly linear, sharp-pointed, plane or somewhat cana- 
liculate, viscidulous as are also the branchlets: inflorescence a 
narrow thyrsiform-panicle, rather leafy, at maturity barely sur- 
passing the uppermost leaves: heads about 12™" high; bracts 
few (10-14), not in strict vertical ranks, mostly acute or acutish, 
glandular on the greenish keel, from glabrate to ciliate-pubescent : 
corolla tube and throat scarcely distinguishable, but slightly 
expanded upward, the lobes about one eighth of the whole 
length, obscurely short-pubescent below: style branches exserted, 
the appendages longer than the stigmatic portion. 

This species has been secured but twice, both times on strongly saline 
soil, viz., at Buffalo, July 25, 1896, no. 2495, and on Vermilion creek, July 24, 
1897, no. 3590. Its nearest ally in habit and some other characteristics is 
C. frigidus, but in form of inflorescence it approaches C. Parry? (Gray) 
Greene. It differs from C. frigidus in being almost devoid of tomentum, in 
its yellowish branches, its green leaves, narrow leafy inflorescence, and its 
*xserted styles. It is also an earlier plant, one of the earliest of the several 
Species of this genus in this region. 

CurysorHamnus PARRYI (Gray) Greene, |. c.—The habitat and 
fange of this well-known species is always given as “parks of the 
Rocky mountains in Colorado,” but certainly similar parks in 
_ Southern Wyoming, at least, must be added. Typical specimens 
tte, J. H. Cowen, Breckinridge, Colo., August 1896; the writer's 
Nos. 2617 and 3495, Lincoln gulch, August 1896and Big creek, 
August 1897. 
___ Curysoruamnus Howarot (Parry) Greene,? l.c.—The habi 

of this, like the preceding, is given as “ parks of the Rocky moun- 
tains,” but it should be stated that the parks are very different 
| ‘Mtheir character, C. Parryi inhabits moist open ground, known 
Sparks, occurring at intervals in the timbered mountain ranges. 
C. Howardi inhabits that other class of parks, viz., extensive, 
Nigh, dry table-lands like North Park, Colo, and the Laramie 
‘Plains. It occupies the dry foothills and ridges and I doubt not 
that 


tat 


ith “Dr. Greene’s papers cite the literature of all the well-known species so fully 
| “ete Seems necessary only to call attention to this fact. 


374 BOTANICAL GAZETTE [DECEMBER 


may occur on the eastern base of the Rockies, possibiy extend- 
ing into eastern Nebraska, as given in Britton & Brown’s Flora, 
a statement questioned by Dr. Greene. It seems probable that 
some confusion on this point has arisen by the distribution of a 
somewhat similar plant occurring in situations much like those 
in which C. Parryi occurs. This plant I think should stand asa 
species, and may be named as below. 

Chrysothamnus affinis, n. sp.—Scarcely shrubby, the persistent 
base hardly more than a much-branched woody caudex: the sea- 
son’sstems very numerous, simple, 1-2" high, yellowish, glabrate: 
leaves crowded, narrowly linear, acute, erect or spreading, dark 
green, nearly glabrous, 3-4™ long: inflorescence a crowded 
spicate thyrsus which at maturity distinctly surpasses the leaves : 
bracts glabrate, arachnoid-ciliate on the margins, somewhat 
thickened-coriaceous, about three in each row; the outer with an 
ovate base, contracted in a usually spreading acumination ; the 
inner linear-oblong, abruptly acuminate, shorter than the pappus: 
corolla tube slender, bearing only a few, minute, scattering clav- 
ellate hairs, shorter than the expanded, tubular throat which 1s 
cleft about one fourth its length: style appendages tardily but 
at length wholly exserted: akene linear-cylindric, about 6™ long. 

Allied to C. Howardi (Parry) Greene, but clearly distinct by its seat 
shrubby habit, its greenish glabrate aspect, and its crowded yellow inflor- 
escence which surpasses the leaves. C. Howardi has cinerous leaves 4” 
stems, a dirty whitish-yellow inflorescence, and the leaves overtop the com 
paratively few and large heads whose bracts are distinctly arachnoid. 

Excellent specimens were collected by J. H. Cowen, Jefferson, ae 
August 1896, and distributed as Bigelovia Howardi Gray. Typ ait 
University of Wyoming. . 

CHRYSOTHAMNUS AFFINIS attenuatus, 0. comb. (Bigelovia 
Howardi attenuata Jones, Proc. Cal. Acad. Sci. II. 5+ 691: 1695" 
The specimen of Mr. Jones’ type number, 591% 

vdi. ts 
approach C. affinis much more closely than it does C. Howd an 
more persistent stems and longer branchlets, its very 1ong 
nate bracts, and long exserted styles will readily distinguish : F 

CHRYSOTHAMNUS COLLINUS Greene, Pitt. 3° 24. ate ge 
distinct and clearly characterized species I think has not 


$ 


OS EERE TO erg Cae PEI Oe CE OU a ee 


mat 
- i 
= . Se 
ig gp 
y 


2-4" high: stemgseveral to many, from athick woody base, strictly 
erect and so mi What fascicled, grayish with a thin tomentum, the 
annual twigs a rising from near their summit, these also fascicled- 
erect, slender and yellowish-green: leaves erect, linear-filiform, 
very acute, canaliculate, green and glabrate, 3-5” long: heads 
small (about 1™ high), in small fastigiate cymes: bracts oblong, 
abruptly sub-acute, only two or three in each vertical row, the 
scarious margins ciliate-pubescent: corolla sparsely short-hairy, 
divisible into three equal regions (tube, throat proper, and a tran- 
sition region); lobes more than half as long as the tube proper, 
distinctly glandular-thickened at apex: pappus rather sparse: 
style appendages longer than the stigmatic portion, at length 
exserted : the short akene finely pubescent. 

The erect habit of stems, the twigs and leaves, the greenish aspect, and 
numerous but small flower clusters mark this as peculiarly distinct from the 
other species of this range. It is abundant on stony slopes in the Bear river 
hills, near Evanston. Type number 4105, July 27, 1897. Represented also 
by M. E. Jones’ no. 6040, distributed unnamed. 

CurysoTHAMNus puMILus Nutt., Trans. Am. Phil. Soc. 7: 323: 
1840.—The recharacterization by Dr. Greene,’ after every poss! 
ble effort has been made to settle what was the original of the 
species, is of noticeable service to us all. This common and 
Somewhat variable species is now recognizable. It is well repre= 
sented by the following numbers from various parts of Wyoming : 
617, 903, 1121, 1 197, 2883, 3524 and 5398. The last, from 
Hutton’s lake, September 7, 1898, is typical so far as I am able 
to judge. 


- 


CHRYSOTHAMNUS PUMILUS varus, 0. var.— Smaller than the © 
aos: only 1-3 high, the shrubby base divaricately scragey 
branched, the season’s branchlets slender, very numerous, 5—15 
long, with a whitish or straw-colored bark: leaves glabrous 
xcept for an obscurely scabrous margin, linear, almost filiform, 


‘Erythea 3: 93. 1895. 


376 BOTANICAL GAZETTE [ DECEMBER 


one-nerved, somewhat involute and usually more or less twisted, 
irregularly and widely divaricate, very numerous on the new 
wood and often fascicled on two-year old branches, short, rarely 
exceeding 3, usually much shorter: inflorescence and bracts 
much as in the species. 

This is what has often been called in this range Bigelovia Douglasit 
stenophylla Gray. I have distributed some specimens under that name, but 
I am now satisfied that that is a very different plant and belongs to a more 
western range.’ It occurs mostly on dry ridges and stony or sandy slopes. 
Type of the variety is no. 1847, Centennial valley, August 26, 1895. Another 
collection is no. 4434. 

CHRYSOTHAMNUS PUMILUS acuminatus, n. var.— The habit of 
the species: numerous slender stems from a woody base 2-3" 
high, with whitish bark: leaves numerous but early deciduous 
below, crowded toward the inflorescence, nearly filiform, ascend- 
ing, somewhat twisted, 2-3™ long: inflorescence more paniculate 
than in the species; bracts lanceolate, long-acuminate with more 
or less spreading tips, nearly equaling the 5-6 flowered disk. 

This variety I have from La Veta, Colorado, only, collected by Professor 
C. S. Crandall, August 21, 1897. 

CHRYSOTHAMNUS LANCEOLATUS Nutt. 1. c.—I see no reason 
for reducing this to a variety. Numerous collections of it show 
as much constancy in the essential characters as most of the 
recognized species. Its low tufted habit, its uniformly scabro- 
puberulent surface, and its either plane or twisted lanceolate 
leaves make it not hard to recognize. Specimens by the writer, 
nos. 889, 905, 2672, 2793, 5294 and 5314 are representative, 4 
is also Professor Crandall’s from Walden, Colorado, July 1894. 

CurysoTHAMNus GLAUCUS Aven Nelson, Bull. Torr. Bot. Club 
25: 377. 1898.-At the time this was published only meager 
material was at hand, but it has since been secured in abun 
The characters as given are well borne out, except that in cl 
specimens the leaves are less glaucous. ‘ 

5I think it should be noted that C. viscidiflorus Nutt. l.c. (Bigelovia D om glass 
Gray) occurs probably in a range to the northwest of that now under oe 
and that several of its varieties, though often attributed to the eastern peur 
not in this range at all. Among such may be named vars. latifolius, serrulatus, 
tortifolius. 


1899] SOME ROCKY MOUNTAIN CHRYSOTHAMNI 377 


CHRYSOTHAMNUS LINIFOLIUs Greene, Pitt. 3: 24. 1896.-— This 

I think has no tybeen secured except in south-central Wyoming, 
s fs on the banks of strongly saline creeks. Its halo- 
phytic anddhy drophilous nature is very marked, as it often grows 
‘feet’ in water so strongly impregnated with salts as to 

be whol unfit for any use whatever. Its ae collection was 


= 
—e 
or 
=> 
ss 
cr 
n 
= 
pa 


CurysoTHAMNUS VasEyI (Gray) Greene, Erythea 3: 96. 
1895.—As pointed out by Dr. Greene, this has an akene quite 
at variance with the species that have preceded. The range 
given for it is too limited, for certainly very characteristic speci- 
mens are at hand as follows: G. E. Osterhout, North Park, 
| Colorado, September 1897; by the writer, Big creek, August 11, 
1897, no. 3494, and near Laramie, September 1898, no. 5331. 


UNIVERSITY OF WYOMING, 
Laramie, Wyo. 


ON THE TOXIC EFFECT OF DELETERIOUS AGENTS 
ON THE GERMINATION AND DEVELOPMENT OF 
CERTAIN FILAMENTOUS FUNGI. 

jah. CLARE. 
(Concluded from p. 327.) 
Att acids tested retard germination and early mycelial 
development of the mold fungi. In the case of the mineral acids 


this retarding action is usually evident in oa concentration. 


The acetic acids do not have any perceptible influence at this 
dilution. Cultures which are only slightly retarded almost 
invariably take on new vigor a few hours after germination, and 
overtake and surpass the checks in development of mycelium. 
This excessive mycelial development is usually accompanied by 
retardation of fruiting, and usually reaches its maximum in 
cultures two or three removes from the inhibiting concentration. 
Cultures in the acetic acids show a greater stimulation of mycelial 
development than in HCl or H,SO,. Cultures in HNO; 
resembled those in the acetic acids in development of mycelium 
but were not so greatly retarded in fruiting. 

Smaller quantities of acid on an average proved injurious to 
20” 

2048 ; 
distinctly detrimental. In this respect the others came 11 ©” 
following order: Botrytis, 23; Penicillium, a8; Aspergillus, 425 
Sterigmatocystis, 64. 

CEdocephalum was also the most easily inhibite 
inhibiting coefficient being 72; followed by Botrytis, 
lium, 100; Aspergillus, 104; and Sterigmatocystis, 200 

Botrytis, however, was the most easily killed. T 
in this respect was: Botrytis, 100; (Edocephalum, 1373 
gillus, 272; Sterigmatocystis, 369; and finally Penicillium, ' ‘ 
spores showed by far the greatest resistance, represented by 
378 [DEcEMBES 


being on the average 
the 


(Edocephalum than to the other forms, 


d, its relative 
76; Penicil- 
he order 


Aspet- 


1899 ] TOXIC EFFECT OF DELETERIOUS AGENTS 379 


coefficient 498. This high resistance of Penicillium in regard to 
the death-point may be partly due to the tendency of the spores 
to adhere in bunches in making the inoculation, a difficulty never 
fully overcome. 

In table II a comparison of the chemical affinities of the 
different acids tested by seven methods is given. 


TABLE II. 
CHEMICAL AFFINITIES OF ACIDS. 
HCN |” Tri. Di. Mono, | Acetic | HS o,| HNO, | HCl Acid 
=) 02.3 | 25.3 4.9 Tua? G52 99.6 100 | Relative ionization in = 
sol. 
| 68 23 4.3 ~34\.. 74 g2 100 | Catalysis of methyl ace- 
e 
63 18 Ce He eae | 67 110 100 | Catalysis of calcium 
xalate ; 
e | 82 34 9:21 184 8 102 100 | Neutralizing hydroxids 
po 19-4.| 27 4.9 4:\ 93 100. 100 | Invertion of cane sugar 
= 199.7 | 62.4 | 17.2 | 4.7 | 72-9 | 99 100 | Multirotations of dext- 
rose 
200+/200 100+ |100 25 100 100 100 Physiological action on 
upinus 
7966 [255 |359 |396 |277 | 112 163 100 | Ditto, on molds 


Line 1 gives the relative ionization of these acids (except 
HCN) in normal solution. HCl being the basis of comparison 
here as in the succeeding tests is given 100 units. Line 2 gives 
the relative powers of the different acids to promote the well 
: known catalysis of methyl acetate in aqueous solution. Line 3 
= gives their relative activity in decomposing calcium oxalate. 
Line 4 gives Ostwald’s (’g!) determinations of their relative 
affinities for hydroxids. Line 5 gives their relative activity 
Minverting cane sugar. 
: The correspondence of the results presented on lines is 
_ With the ionization data on line 1 is certainly quite striking. 
Arrhenius (’83) was the first to point out this close numerical 
‘greement. Since the publication of his work in 1883 the ee 
as taken firm hold on many chemists that the ionized — 
ofan acid and that only is chemically active. Whetham (’95) 


380 BOTANICAL GAZETTE [ DECEMBER 


expresses concisely recent views when he says (p. 165): “We 
may take it, then, that only that portion of a body is chemically 
active which is electrolytically active —that ionization is necessary 
for chemical activity just as it is necessary for electrolytic 
conductivity.” 

What applies to chemical activity must also apply to physi- 
ological activity, for in its ultimate analysis the latter is doubtless 
due to the former. Kahlenberg and True (’96) remark (p. 35) : 
‘It has always been taken as axiomatic that the physiological 
action of any substance is due to its chemical character.” 

The first work which deviated irreconcilably from the theory 
that all acids have ‘specific coefficients of affinity . . . . based 
on the fact that the relative affinities of different acids are the 
same, whatever the nature of the action by which they are com- 
pared” (Whetham, ’95, p. 162) was that of Levy (’95). It will 
be seen (line 6) that acetic, mono- and dichloracetic give coeffi- 
cients of activity which are in round numbers 200 per cent. in 
excess of that called for by the theory. The physiological 
activity of the acids towards phanerogams (Kahlenberg and 
True ’96) (line 7) is equally out of harmony with the theory, 
when we find the almost un-ionized HCN much more active 
than the ‘‘strong” mineral acids. The climax, however, 18 
reached in the data recorded on line 8, where we are dealing with 
concentrations which contain in many cases a very large propor 
tion of un-ionized molecules. : 

The chemical reactions involved in physiological investiga- 
tions are doubtless vastly more complex than in the case of the 
earlier studies recorded on lines 2-5. In the data recorded on 
lines 6-8, derived from the action of the acids on the more com 
plex carbon compounds, and the highly complex aldehydes, 
albuminoids, etc., found in the protoplasm of living cells, Lb 
surely find a great exception to the alleged law that the relative 
affinities of different acids are the same whatever the nature of the 
action by which they are compared. These affinities, indeed, appeat 
in some cases to be almost the converse of that required by !° 
theory above noted. 


1899] TOXIC EFFECT OF DELETERIOUS AGENTS 381 


HYDROXIDS. 


As it is quite impossible to handle solutions of hydroxids in 
an ordinary atmosphere without a greater or less loss by neutral- 
ization by the CO, of the air, the following toxic values of 
potassium hydroxid and ammonia must be somewhat less than 
their absolute toxic value. This source of error was reduced as 
much as possible by rapid handling in making the cultures. 
Potassium hydroxid, KOH; 77 (?), 166, 282. In no other 
case was it found so difficult to determine where to place the 
- coefficient of injury, pe retarded germination in all cases, and 
| Ld Hn 

sane and even eee: 
an injurious influence. 

At 24 and 36 hours, however, the cultures presented a very 


with some forms 


(CEdocephalum) showed 


n 


different appearence. Cultures in concentration showed a 


2048 
heavier mycelium than the checks, and with stronger concen- 
trations this stimulation of mycelial development was more 


n n : 
— or — concentration. 


Marked until the climax was reached in ba as 


. . . . . . 
As 7p Proved the average inhibiting point, it will be seen 


that, as with the acids, the maximum stimulation of mycelial 
! development occurred about two removes from the limit of ger- 
: Mination, or, in other words, in solutions containing one fourth 
: the Concentration of the agent inhibiting the germination of the 
‘Pores. The retardation of fruiting in the stimulated cultures 
_ Was very marked, and suggested the query as to whether they 
Were both due to the same cause, or whether one was a result of 
E the Other. It is well known that with the higher plants vias 
_ Pression of fruiting tends to force the energies of the plant into 
Vegetative lines, and it is not apparent to the writer why this 
Pout not be trué: of the fungi also. On the other hand, 
Strytis, which did not fruit, showed nearly or quite as great a 
‘mulation of mycelial development as any other form. 


382 BOTANICAL GAZETTE [ DECEMBER 


The toxic properties of KOH are probably largely due to 
the OH ion. It is about 94 per cent. ionized at = (Ostwald 
’86), its inhibiting point. Just what proportion of the toxic 
properties is to be attributed to the remaining 6 per cent. 
un-ionized KOH we have as yet no means of knowing. _Inas- 
much, however, as KOH is more highly ionized than HCl at the 
inhibiting point and is distinctly more toxic, we may safely con- 
clude that the OH ion is somewhat more toxic for fungi than 
ionic H. 


Ammonium hydroxid, NH ,OH; 29 (?), 57,83. This hydroxid, 
in contrast with KOH, is but slightly ionized, oe the inhibiting 


concentration, being about 8 per cent. ionized (Kohlrausch 85.) 
Its high toxic value is then doubtless due very largely to the 
un-ionized molecule. 

Like KOH, although to a less degree, it caused a marked 
stimulation of mycelial development in many cultures. It is 
worthy of note that Sterigmatocystis, which is particularly resis- 


: aie an ; 
tant to both ionic H and OH, proves quite susceptible to both 
acids and hydroxids in the molecular form. 

These results with hydroxids are not in harmony with those 
obtained by Krénig and Paul (’97) with anthrax spores,, They 
found that the bases KOH, NaOH, LiOH and NH , OF ‘disinfect 
in direct proportion to their degree of ionization,” NH,OH being 
practically non-toxic. Anthrax spores have evidently a great 
specific resistance to this agent, and perhaps even 4 general 
comparison would be unfair. Be this as it may, ammonia !> 
without doubt one of the more violent poisons for fungi, far sur 
passing the mineral acids, copper, cobalt, etc., in toxic proper 
ties, and comparing favorably with KCN. 

Data as to the effect of hydroxids on the higher plants * 
rather meager. Kahlenberg and True (’96) found Lupinus 0 


. ‘ ; plete 
survive in — KOH. As ionization is practically comple 


ij : : ‘ 0 to: 
here, Lupinus is evidently much more resistant to OH than 


1899] TOXIC EFFECT OF DELETERIOUS AGENTS 383 


ionic H. Bokorny (’88) found that ammonia in os concentra- 
tion, in common with other basic substances, caused the produc- 
tion of granules in the protoplasm of Spirogyra cells, but failed 
to modify otherwise the normal activities of the cell. Detmoor 
(94) found that a 10 per cent. solution of ammonia at first 
energetically excited the protoplasm of Tradescantia hairs, later 
producing anzsthesia. Washing with water, however, restored 
the original characters of the protoplasm. From these data it 
would seem that hydroxids are more fatal to the molds than to 
the higher plants. 
Formaldehyde, HCHO; 0.553, 7.43, 2. Formaldehyde, as 
Was anticipated, proved to be one of the most deadly agents 
tested, being surpassed in this respect by mercury, silver, and 
the two chromates only. Chemically considered, formaldehyde — 
isa very unstable compound intermediate between methyl alco- 
hol and formic acid, being in fact the intermediate step in the 
oxidation process by means of which the latter is derived from 
the former. It is both a reducing and oxidizing agent, ‘and this. 
together with its great instability may account for its extremely 
‘oxic properties toward fungi. 
To many kinds of protoplasm, including that of the higher 
animals and perhaps the higher plants, formaldehyde is non- 
toxic. Instances are on record of persons having drunk a I per 
cent. solution without inconvenience (Arthur ’97, p. 21). To 
the lower animals, however, it is more toxic, > being fatal to 
Yorms, mollusks and isopods in two hours (Loew ’88). Acton 
89), in experiments on the assimilation of organic compounds 
by Steen plants, found that while they could use glucose, sac- 
charin, glycerin, etc., they failed to use aldehydes or their 
detivatives. Cohn (’94) found a I per cent. solution very fatal 
0 Spirogyra. 

For the fungi, however, there is no doubt that in formalde- 
Yde we have one of our safest, most energetic, and most ser- 


: "iceable poisons, —— proved fatal to Aspergillus and Penicillium, 
3 512 


384 BOTANICAL GAZETTE [ DECEMBER 


an to Sterigmatocystis and Botrytis. Cédocephalum was 
inhibited by 5648, and killed by yy This will perhaps be 


better appreciated if stated in another way. One part by weight 
_ in 273,066 parts beet infusion proved fatal to GEdocephalum, and 
I part in 4,369,066 permitted the germination of but Io per 
cent. of the spores in eleven hours (as compared with 95 per 
cent. in four hours in the checks) and greatly injured the 
mycelial development. In regard to the other forms, I part to 
273,066, although greatly retarding germination, caused a dis- 
tinct stimulation of mycelial growth on the second day. These 
stimulated cultures resembled those growing in media contain- 
ing alcohol. 

Very interesting in this connection are the theories regarding 
the synthesis of starch in green plants, and of the proteids in 
the fungi (Kozlowski ’99), in both of which formaldehyde has 
long been regarded as forming a very important step. These 
theories, particularly that in regard to the synthesis of proteids 
in the fungi, challenge further careful investigation. It seems, 
prima facie, inconceivable that a compound markedly injurious 
to a plant when present in the almost infinite dilution of one 
part of weight in 4,369,0Co parts nutrient medium, as is the case 
of CEdocephalum, should be formed by the protoplasm of that 
plant and be used again in the synthesis of its proteids, as must 
be the case if our theories be correct. 

In regard to the nature of its toxic action we have few data. 
It is believed to act upon the propeptones and the albumins, 
affording compounds which are not readily soluble (pave 
port ’97). 

Ethyl alcohol, C,H ,OH ; 717, 3686, 8602. Alcohol when ie 
trasted with formaldehyde, is apparently non-toxic. One molecu : 
of formaldehyde has powers of inhibiting germination of aie * 
spores equal to those of 2600 molecules of alcohol. The Bes | 
in their killing powers is even greater, being as I : 4300! . 
alcohol is a distinct poison to the protoplasm of plants eee 
moto ’95), being used, in fact, quite extensively as 4 xing 


1899] TOXIC EFFECT OF DELETERIOUS AGENTS 385 


agent for tissues, and even more widely asa preservative. It 
inhibits germination in the mold spores used at an average con- 


nm RN ae 
centration of rae and is fatal to all except Penicillium in *. 


Five per cent. of the spores of Penicillium survived immersion in 
this concentration (17.6 per cent.) for 72 hours at 28° C. All, 


} ; 8 ; 

_ however, were killed by = which was the greatest concentra- 
tion required to kill in the case of any agent tested. 

| The presence of alcohol in “ and = concentration distinctly 


fetarded germination of Sterigmatocystis and Botrytis. No 
retardation was noticed with Aspergillus or GEdocephalum, The 
latter, indeed, showed some evidence of acceleration of germina- 
tion in = and = concentrations, but it was not sufficiently 
marked to be certainly stated. The stimulation of mycelial 
development and retardation of fruiting in ” was very marked 
with Aspergillus and Sterigmatocystis. Cdocephalum and Peni- 
tillium showed some stimulation of mycelial development and 
‘slight retardation of fruiting in this concentration. Botrytis 
Produced its heaviest mycelium in = and “ . 

_ The change that a plant may undergo as it grows older, 
‘lM the character of its election of foods is of great interest to 
Physiologists (Davenport ’99, p. 333). Duclaux (’89) found 
that while alcohol restrains or arrests germination in mold 
‘pores, it is made use of almost as abundantly as sugar by the 
/Mdult plant. My results would seem to support this view. It 
pay be possible, however, that alcohol acts as a stimulant rather 
Man as a food, as is the case with zinc sulfate, and other non- 
Nourishing compounds. Richards (’97) found that the addition 
0035 per cent. Zn SO, to aculture medium in which Aspergillus 
Meer Was growing doubled the dry weight of the mycelium. 008 
‘Pt cent. Zn SO , Similarly added to a flask culture of Botrytis 
“tsed a production of quadruple the normal weight of myce- 
fom, We cannot suppose that the small amount of zinc present 


386 BOTANICAL GAZETTE [DECEMBER 


in itself caused the greater growth by supplying nourishment, 
Zn not being a necessary or desirable element in a nutrient 
medium for fungi. Richards interpreted the function of the 
- zinc to be that of a stimulant rather than a food. May it not be 
that alcohol performs a similar stimulating function, rather than 
that it produces an acceleration of growth by nourishment? 
Whatever may be the correct explanation of the influence of 
alcohol on the development of the mold fungi, it seems to be 
demonstrated that the protoplasm of the molds is more sensitive 
in the conidial stage to the influence of this and most other dele- 
terious agents than at any other stage in their development. 
Potassium cyanid, KCN ; 2.2, 25.6, 77. Potassium cyanid in 
aqueous solution is very unstable. A solution of 24 per cent. 
KCN in pure water was prepared by the chemist, and on being 
used within three hours of titration gave the following critical 
points with Aspergillus and Penicillium : 
2 


e failed. 


Aspergillus in — grew, in 
Penicillium in 5 grew, in - failed. 
Ten days later this stock solution was again tested, having 
been kept in a dark cupboard at ordinary laboratory tempera- 
ture in the meantime, with the following result: 


n n 
i in — in. — failed. 
Aspergillus in a grew, in — aile 


nt . 
Penicillium in — grew, in — failed. 
64 32 
5 ‘ Ree 2) ee then made 
A solution in beet infusion containing 3 CN was 
k at a constant 


up from the stock solution and placed in the dar 
tested. 


temperature of 28° C. for 10 days longer and again 
The critical points were then as follows: 
a 


76 Brew: in ~ failed. 


Aspergillus in 3 


: nN . 
Penicillium in “ grew, in failed. 
I 


1899] TOXIC EFFECT OF DELETERIOUS AGENTS 387 


From these data we learn that a 24 per cent. aqueous solu- 
tion of KCN deteriorates so that at the end of ten days it has 
but little more than one fourth its former toxic value. Made up 
Wt 
re 
retains but one eighth of its original toxic value. All cultures with 
KCN, reported in this paper, except those noted above, were 
made up within four hours of titration of the stock solution. 

It will be seen by reference to the charts that KCN in 68 


in beet infusion at = concentration and kept ten days longer it 


tion has almost exactly nine times the toxic effect of ionic H. 
KCN in the concentrations used is quite highly ionized (Kohl- 
fausch ’79), but in trying to approximate the toxic value of the 
CN ion, the fact that a certain amount of hydrolysis takes place 
in aqueous solutions of this salt, with a corresponding formation 
of the deadly HCN, must not be overlooked. According to the 
data worked out by Shields (’93) and what we already know of 
the properties of HCN, approximately 15 per cent. of the total 
oxic value of these solutions must be attributed to the HCN 
besent. This would give something under 8H as the value of 
the CN ion. 
KCN retarded germination and early development in all forms. 
o|h this, as with many other agents, those cultures not greatly 
injured soon overcame the effect of the poison and grew and 
i ttuited normally. No marked retardation of fruiting nor unusual 
-tevelopment of mycelium was noted. Sterigmatocystis, in so 
Many cases highly resistant, proved equally sensitive hs (Edo- 
~“Phalum, both being inhibited by —~; and killed by ¢ - 
Kahlenberg and True (’96) found that towards Lupinus it 
tas the value of 1H only. They also show that in the cases of 
Potassium ferro- and ferri-cyanid the iron and CN radical form 
‘“mplex ions, the toxic value of which is far less than that of 
| Ne CN ions, 
Mercuric chlorid.—HgCl, ; 0.0258, 0.287, 0.331. This proved 
. Most fatal compound tested, leading silver nitrate by a narrow 
_argin, 


388 BOTANICAL GAZETTE | DECEMBER 


It is of interest to note that although only very slightly 
volatile at ordinary temperatures, and doubtless less so in 
aqueous solution, mercuric chlorid is sufficiently volatile even 
in dilute aqueous solution at 28°C. to be distinctly toxic. This 
was demonstrated by placing a few drops of a very dilute solu- 
tion in the bottoms of cells containing hanging drops of pure 
beet infusion inoculated with mold spores. The germination of 
the spores was inhibited. Mycologists have frequently reported 
failure to germinate spores in cells which had been sterilized by 
rinsing in a dilute solution of HgCl,. These failures were 
doubtless due to the volatile properties of this agent together 


with its extremely deadly character. 
Hn 
Botrytis proved particularly sensitive to this agent, 65530 


proving fatal. Penicillium failed to show its usual high relative | 
LA 

4096’ 

proved fatal to Aspergillus and Sterigmatocystis. Toward bacteria 


the same concentration as 


resistance, being killed by 


it is also extremely fatal, in nutritive bouillon preventing 


Li 
70000 
the development of the splenic fever bacterium (Davenport 97, 
p. 14). The data regarding its influence on the higher plants are 
n 


fatal to Lupinus, 
6400 


meager. Kahlenberg and True (’96) found 
n 
12800 
but double the toxic value of HCl. For the molds its averag® 
value will be seen to be over 800 times that of HCl. : 
Silver nitrate, AgNO, ; 0.0125, 0.375; 0.375. Almost, ater! 
altogether, as violent a poison as mercury, silver stands with itat | 
the head of the list of toxic agents tested. Among the poison? 
for molds tested, it is comparable with mercury alone gee 
the metals, and with the chromate and. dichromate anions 8 h 
formaldehyde only among the other agents. As is the case on 
bacteria (Davenport ’97, p- 14) toward the molds silver 15 2 
quently a more violent poison than mercury. Of the five a to 
used CEdocephalum and Penicillium proved more susceptible 


while it survived in 


This is unexpectedly low, being in fact 


1899] TOXIC EFFECT OF DELETERIOUS AGENTS 389 


silver, while Aspergillus and Botrytis were more susceptible to 
mercury. The fifth form, Sterigmatocystis, had an equal resist- 
ance to both. The extraordinarily small quantity of mercury 
required to kill Botrytis, however, left the honors with mercury 
as the more deadly agent. 

A very striking contrast in specific resistances is afforded by 
Botrytis and Penicillium in solutions of these agents. One six- 
teenth the amount of HgCl, necessary to kill Penicillium was 
fatal to Botrytis. With AgNO, one fourth the concentration 
_ fequired to kill Botrytis was fatal to Penicillium. Or, putting it 
another way, AgNO, has eight times the toxic value of HgCl, 
toward Penicillium, while the exact converse is true with Botrytis, 
HgCl, being eight times as effective as AgNO, to this form. 


The low resistance of Penicillium to silver is quite striking, 5 
Proving fatal. In one other case only (H,O,) did Penicillium 
show a lower resistance than Botrytis. 

Toward splenic fever bacteria gold is the only other metal 
‘omparable in toxic properties with mercury and silver (Daven- 
port '97, p. 14). Toward phanerogams silver is much more 


to 


a 
a 

i Zea, and ———— 
see being fatal to 204800 


Lupinus (Heald ’96, p. 152). At the concentrations used the 
AgNO, would be practically entirely ionized (Kohlrausch, ’85). 

Cadmium nitrate, Cd(NO,) 4; 0.075, 6.7, 24. Ranking closely 
With silver as a poison for the higher plants, cadmium proves 
tobe very highly toxic to the mold fungi. Perhaps the most 
Marked feature in the cultures with cadmium was the very wide 
ange between the killing point and the point where develop- 
ment was only noticeably injured. It will be noticed that the 
: Tatio of its coefficient of injury to that of HgCl, is less than 
_'Lwhile the ratio between their death points is 75:1. A sec 
ond str iking feature of the toxicity of cadmium is the great varia- 


toxic than mercury, 


‘ * ‘ = 
Fon in the specific resistances of the different forms. 4096 


Moved fatal to Botrytis, but — was required to kill Sterig- 
: 32 


390 BOTANICAL GAZETTE [DECEMBER 


matocystis. Penicillium spores failed again to show their usu- 
7. 
256 
the usually sensitive CEdocephalum actually germinated 95 per 
cent. and matured a few fruits. 
Molisch (’94) was one of the first to record the toxic prop- 


ally high resistance, proving fatal, a concentration in which 


erties of cadmium to plants. He found = fatal to Aspergillus 
when experimenting with various metals in an endeavor to find 
a substitute for calcium in nutrient media. vi proved fatal 
to the form of Aspergillus used by the writer. 


At its average inhibiting concentration cadmium nitrate 
would be about go per cent. ionized (Grotrian, ’83). 


OXIDIZING AGENTS. - 
Potassium dichromate, K,Cr,O,; 0.094, 0.3, 1.25. 
Potassium chromate, K,CrO,; 0.156, 0.4, 2.25. 
These salts at the dilutions at which they are effective are 
doubtless practically entirely ionized (Ostwald, 88). 
As poisons for the molds they rank, as already mentioned, 
with formaldehyde, silver, and mercury. The anion of the 


dichromate, Cr,O, has a toxic value of about 770 H; that of the 


chromate 575 EH. This may indicate some relation between 
their oxidizing powers and their toxicity. 
The effects of these salts in concentrations permitting 
development of the fungi were very similar, and resembled that 
of H,O,. Retardation of germination in the cultures approach- 
ing the inhibiting point was noticed in all forms with both 
agents, but it was not nearly so well marked as it is in wanes 
Another feature was the fact that every culture that germinate 
any spores developed some conidia within forty-eight hours. 
Toward the higher plants these anions seem to be relatively 
” _ solution of HgrOw 


the same concentration as permitted growth with HCI ( 
93), howevel, 


berg and True, ’96). Toward the alge (Loew, 


much less toxic. Lupinus survives in a 


1899] TOXIC EFFECT OF DELETERIOUS AGENIS 391 


both these anions are strongly toxic, oa K,Cr,O, being fatal 
to Spirogyra in a few hours. : 
Hydrogen peroxid, H,O,; 38, 205, 127. This agent had 
an effect on the molds very similar to that described for the 
chromates. With it, however, the characteristics given for the 
chromates were somewhat intensified. Germination was but 


slightly retarded in most forms in ar concentration, and 


(Edocephalum actually showed a higher percentage germinated 
in these cultures at four hours than in the checks. The differ- 
ence, however, was not sufficient to establish the conclusion that 
the H,O, accelerated germination. That this concentration 
accelerated early mycelial growth with this form was undoubtedly 
established. At four hours the average length of the germ tubes 


i : . . . 

in aa concentration was 120“ as compared with 4op in the 
checks. At seven hours they were 3104 and 115m respectively. 
: . 
156° the limiting culture for this form, made the best mycelial 


development and matured the heaviest crop of conidia in the set. 
The characteristic already mentioned for the chromates regard- 
ing fruiting in the cultures was even more marked with this agent. 
Every culture that produced even the scantiest mycelium pres- 
ently developed at least two or three all but normal conidiophores. 

In regard to the action of this agent on other organisms, 
the data are meager and conflicting. Miquel (’83) places it third 
ss disinfecting properties of all agents used by him. In his 
results it is rated above both HgCl, and AgNOs, being agian 
antiseptic in dilution of 1 part to 20,000. This is certainly too 
igh an estimate. Sternberg (’92) found it to have a com- 
_ Paratively low toxic value for bacteria. One in 1000 kills ordi- 
nary water bacteria, cholera, and typhoid (Altehofer ’90). This 
Would be about —, or a 3 to 4 per cent. solution of the ordi- 


‘Naty Io-volume commercial article. It will be seen that this 


: n 
agrees fairly closely with its toxic properties for the molds, =) 


392 BOTANICAL GAZETTE | [DECEMBER 


being fatal to Sterigmatocystis and es to CEdocephalum. The 


others are more resistant. It is, however, more toxic to alge 
(Bokorny, ’86) than to molds, and Ciliata are even more suscep- 
tible. Paneth (’89) found a .005 per cent. solution to be the 
limiting line for the latter. 

Commercial preparations of H,O, vary very greatly in the 
amount of H,O, in solution. A true ten-volume solution 
should yield, when fully decomposed, ten volumes of O, and 
should contain by weight 3.04 per cent. H,O,. The preparation 
used in this study although “ fully guaranteed,” etc., contained 
but 2.59 per cent. H,O, on being tested. 


SULFATES OF THE STRONGLY-TOXIC METALS. 


These salts are arranged in the order of their toxic properties 
towards molds in the following list : 

Nickelous sulfate, NiSO,; 4.8, 33.6, 1155. 

Cobaltous sulfate, CoSO,; 6, 57.6, 389. 

Ferrous sulfate, FeSO, ; 14.4, 775, 2150. 

Copper sulfate, CuSO, ; 8.4, 7377.2, 582. 

(Copper nitrate, Cu(NO,),; 8.4, 234, 634.) 

Zinc sulfate, ZnSO, ; 26.4, 602, 3072. 
_ The data regarding the ionizations of these salts are rather 
‘meager. They are, however, not greatly different in ionization 
at similar concentrations. This is about 40 per cent. to 44 P® 


n ‘ 
cent. at ee concentration (Whetham, ’95, pp. 218-276). 


Nickelous sulfate—The different molds exhibited more vari 

tions in their specific resistance to this agent in regard to the 
: n 

death point than was observed with any other. 3 proved fatal 
to Botrytis, while Aspergillus failed to lose its vitality 1n 4 
normal solution (containing over 13 per cent. ant 
NiSO,) for 48 hours. Much less variation was shown 10 . 
inhibiting powers. Aspergillus and Penicillium germinated . 


Mi Woe: : 
aa 266 inhibited Botrytis. The fact that 32 times the strength 


1899] TOXIC EFFECT OF DELETERIOUS AGENTS 393 


which inhibited the spores of Aspergillus failed to kill them was 
nowhere else paralleled with this form and but once surpassed by 
Penicillium. 

Cobaltous sulfate stands in second place in this group as an 
inhibiting agent, but, as will be seen, it is relatively much more 
powerful as a disinfectant. Penicillium, however, showed its 
usual high powers of resistance in this respect. Inhibited by 
n nu 


> was required to kill. 

Ferrous sulfate-—Iron, a necessary element for the nutrition 
of the molds (Molisch, ’94) in common with all other plants, in 
excess proves to be a very strongly toxic agent, surpassing in 
this respect that king of modern fungicides, copper. With the 
exception of nickel as noted above, iron showed a greater differ- 
ence between the average concentration required to inhibit the 
spores and the concentration required to kill than any other 
agent. Botrytis, as usual, showed less variation in this respect 
than the other forms, but even with it one eighth the fatal con- 
fentration inhibited germination. (CEdocephalum showed the 
§teatest resistance to this agent both as regards inhibition of 
sermination and killing of the spores. This was the only 
agent with which it had a higher specific resistance than any 
other form, 

Copper sulfate and nitrate.—These salts proved to be quite 
Similar in toxic properties, as may be noticed by a glance at the 
. diagrams, p. 312. The nitrate, however, is much more highly 
| ionized at the critical concentrations ; hence we judge that the 
| “N-ionized molecule CuSO, has a toxic value not greatly differ- 
it from ionic +Cut. : 


Hn 
Penicillium, although inhibited by se Cu(NO,), and = 
CuSO, required a ” concentration in both cases to kill the spores. 


: This certainly shows great resistance to these agents as compared 


_ With the other molds. It, however, appears insignificant when con- 
. n 


- tasted with many of the results gotten by other workers. [53 


394 BOTANICAL GAZETTE [DECEMBER 


CuSO,, which effectually inhibited germination in the form used 
by the writer, contains about 0.1 per cent. CuSQ,.. DeSeynes 
(’95) reports growing cultures of Penicillium glaucum, gotten from 
different sources, in solutions containing 2 to 9.5 per cent. 
CuSO,. Cultures grown on the stronger concentrations bore red 
spores. Pfeffer (81) reports finding Penicillium growing ona 
concentrated solution of CuSO,. Manasein (fide Loew 93) 
finds from his experiments that this salt must be present in a 
.25 per cent. concentration before it has any appreciable effect on 
this fungus. Others might be quoted, but sufficient has been 
said to indicate the possibilities yet to be investigated of the 
acclimatization of fungi (and other plants) to chemical agents 
(Davenport and Neal, ’96). 

Zinc sulfate—Inasmuch as zinc chloride is used very exten- 
sively for impregnating railroad ties to prevent attacks of wood- 
destroying fungi (Roth, ’95), it was a surprise to find it having 
so low a toxic value, particularly when it is recalled that one of 
the molds tested, Penicillium, is one of the enemies of the wooden 
ties (Ward, ’98). Koch (’81) finds the chlorid and the sulfate 
to have practically the same disinfecting power. Towards 
Aspergillus we may say that zinc is non-toxic, the spores surviv- 


4 E m a 2n 
ing an immersion of 48 hours in a on (27 per cent. anhydrous 


ZnSO,) concentration. In a - concentration (7 per cent.) 25 
per cent. of the spores germinated and grew slowly. The 
mycelium produced was very irregular and closely septate. 
Strychnin sulfate, CoH UNO HGSO:. This alkaloid, the 
only one experimented upon, was dissolved in the slightly acid 


beet infusion until a saturated solution was obtained. This was 
found to have a content of 4.31 per cent. strychnin sulfate. This 
| solu- 


s é 2 
in terms of a normal solution would be about = a norma 


tion of this substance requiring over 30 per cent. on account of 

its very large and heavy molecule. 
Sterigmatocystis grew and fruited normally i 

solution, although germination and early growth were fr 


n this saturated 
etarded. 


1899 | TOXIC EFFECT OF DELETERIOUS AGENTS 395 


Aspergillus and GEdocephalum also grew and fruited, but the cul- 
tures were much behind the checks. Penicillium was inhibited 


by = but was not killed by the saturated solution. Botrytis 
28 


was also inhibited by = and was killed by - That the solu- 
tion used was completely saturated was shown by the appearance 
of numerous microscopic crystals in some of the hanging-drops 
which were exposed for a few moments to the air, the evapora- 
tion from the culture medium causing some of the strychnin to 
crystallize out. The molds, however, continued to thrive in these 
cultures, their hyphe growing among the crystals. 

Of its effect on plants in general we have few data. Daven- 
port (’97) mentions that it kills the protoplasm of Drosera 
tentacles, and hinders the development of peas, corn, and lupines. 
The injurious concentrations, however, are not mentioned. 
Much interesting work has been done on Protozoa (Schiirmayer 
-'90) by various workers. The results of these studies as well as 
those presented here for the molds are in harmony with the 
theory of Loew (’93) that the action of alkaloids is chiefly con- 
fined to the plasma of the ganglion cells. Fungi and bacteria 
having no differentiation of nerve protoplasm are practically 
unharmed by this agent. 

Potassium iodid, bromid, and chlorid—These salts proved to 
have a very low toxic value. A complete series of cultures with 
the five molds was made up with the iodid only. Its coeffi- 
cients were determined to be 384, 2457, and 4915. 4 inhibited 
all except Aspergillus, — was fatal to all except Penicillium. 
As the potassium salts of the haloid acids are all quite highly 
lonized, an attempt was made to determine the relative toxic 
Properties of the zonic halogen elements. Aspergillus and CEdo- 
ephalum were used. To these molds ionic I proved doubly 
toxic as compared with Cl. Br occupied an intermediate post 
‘ion, being very slightly more toxic than ionic Cl. 


Sodium salicylate GyH ere we 24, 162,182, Tt was 


396 BOTANICAL GAZETTE [DECEMBER 


thought desirable to test this salt on account of its wide use as a 
preservative in laboratories and elsewhere. It proved somewhat 
more toxic than HCl, but not so fatal as HNO,. Cdocephalum 
was quite susceptible to its influence, —s being fatal. It is not, 
however, of much value as a disinfectant, over I per cent. being 
necessary to prevent the development of molds. 

In the following table the various agents are arranged in the 
order of their toxic properties as shown by their powers of inhib- 
iting germination of the spores of the five species of molds used. 
The fifth column gives in round numbers the number of mole- 
cules of each substance required to produce an inhibiting effect 
equal to that of one molecule of mercuric chlorid. The coefficients 


us n 
have the usual value of x in the expression, ——; of -. 
2048 I 
TABLE IL 
Agent Formula psig eae of Ratio 
injury | inhibition |death-point 
~Mercuric chlorid  - . HgCl, .026 281 -331 te 
Potassiu Cette K,Cr,O, .094 1.25 va 
Silver nitrate - AgNO, 013 375 275 1.3 
Potassium hindiate : K,CrO, 156 4 2.25 1.4 
Formaldehyde - HCHO S531. hae be 5: 
ydrocyanic acid HCN 965. 3, 20. oa 
Cadmium nitrate - Cd(NOg)s 075| 6.1 24. se 
Potass yanid KCN 2.2 25.6 | 77: oe 
Nickelous sulfate -  - NiSO, 4.8 | 733-6 ss 8 
monium hydroxid NH,OH 8 51. 83 A 
Cobaltous sulfate - CoSO, 6 57.6 389. ae 
Monochloracetic acid - CH,CICO,H 8.8 58. 64- lle 
ia ic acid - - CHCI,CO,H 10.4 64 64. ob. 
Ac cid “ CH,CO,H 25.6 a3. 314- ce 
Pi chloracetié acd - “ CC1,CO,H Ree go. 115 Eur 
Hydrogen peroxid— - ,0, 38. 105. 127. a 
Ferrous sulfate - FeSO, 14.4 IIS. 2150. i. 
Copper sulfate - - CuSO, 131 82. 4 9 
Sopper nitrate a Cu(NO,)» 8.4 134. 634 sas 
N d ee HNO 48. r4t. 384 
Potassium hydroxid KOH { 19- | 166. | 282+ 593- 
Sodium salicylate a Oo: Reape Na 24. 182 182+ oe 
_Sulfuric acid - —- - %H,SO, . 61.. | 205. 589% oe 
H eauaehione acid - HCl 70. Sa pis: 2150: 
Zin ulfate - . ZnSO, 26.4 | 602 3072. 
Strychnin sulfate : C,2Hgg.N,0,H,SO,4| 179 8775+ 
Potassium iodid : K 384. |2457- [495° | oy, 
Ethyl alcohol — - - C,H, OH 717. |3686. peniebte  ° 


~ { Bik wa a | 
fon [He oe TS Os BE 
Litiiils 
ons Cott EQ® 
WM ee Be H'OO'1ID9 
tort 
ce Es a OR A 
ttt bt 
5 EE AE | H'OD I9HO 
‘BERS REE 
ee GR MA TE 
teetet+e z & 
SEaeeeal H*OOID" HD 
{ coor 
t Corey 
ceeeeee| , 
t EE RSES OS°D 
t Trt 
t 2 mw oF | 
| Het toed re me 
~ secur ured ahaa ana HO’HN 
a: Cee 
| ee a 
6 Dosages: eee 
ies t Petri OSIN 
t 2: BH | 
i jE oe we ee | 
= + coe ttee 
<i See NOW 
rect 
faa (Sa F Sawa e 
S HoH ace 
Coot ("ON)PD 
< Corre 
pt coe 
(an : 4 
NOH 
} 
4 
3 44 OHOH 
| 
1 71 
Sh 
¥, & 
010° 
1 
e 
ON3V 
| ch 
1 aD 4 ‘ 
7S SS BS BN I Too ; 
it if i bi i et tt i i i] it i i i Lilies 40%9*y 
| co PL LL i er ee mI CONC 
| tits [HSER PARRA SRRRR RT ARRAVS RA RRRK RE RNERERRRRAREESEPABR GRRE AME 
| 1 fe ial ERASERS REDPHRSATK Ge HSO Se HSL SKS EVAR SAS SS SSS LY www oo oe ew 
PITTI EL eee ee ee a ee 
— Ff co 103H 
a i 
g ae 
= CD 


| (DECEMBER 


DIAGRAM X. 


398 


@ 


pt} t + ji} 


ta 


Ot ft 
tt 4+ + 4+ 


i 
sassserzaawange 


*osuz: 
DH 
"OS*HA 


®NOOD oo HPD 


Fosn 
*O°H 
H*O 7199 
H*0O°HO 
H*OO*IOHO 
H*OOID*HO 
*oseo 
HO'HN 
SoM 


NOH 


1899] TOXIC EFFECT OF DELETERIOUS AGENTS 399 


Diagrams IX and X are simply a graphic representation of 
the more important features of this table. The values of the 


abscisse in each case =z in the expression 


of a normal 
2048 
solution. 


SUMMARY. 


rious agents than the higher plants. In the case of the mineral 
acids a concentration of from two to four hundred times the 
"strength fatal to the higher plants is required to inhibit the 
germination of mold spores under favorable conditions. 

2. Different species of fungi present great differences of 
tesistance to many agents. Of the agents tested in this-study, 
NiSO, permitted the greatest specific variation and dichloracetic 
acid the least. 

3. Particular forms of the same species present very different 
powers of resistance, depending probably on previous environ- 
ment. 

4. Individual spores taken from the same pure culture often 
Present considerable variation in resistance. 

5. The five forms used were found to be increasingly resistant 
to the toxic action of acids in the following order: Cédo- 
fephalum, Botrytis, Penicillium, Aspergillus, and Sterigmato- 
| ¢ystis. 


tum, Sterigmatocystis, Aspergillus. 

7. GEdocephalum and Botrytis, although on the average the 

Most susceptible to the various agents, have great specific PESES: 

“ices to certain agents. See FeSQ,, KI, and alcohol. 

8. Tests made with media not well suited for the normal 

development of the fungi tested will give a correct value for he 

: ling concentration, but the data regarding the point of inhibition 

*f germination are not of value. 

_ 9. Tests of the toxic value of solutions are unreliable when 
‘Made in hanging-drop cultures where water was used in the 


I, Fungi are in general much more resistant to most delete- 


i 6. Toward all the agents tested they proved increasingly 
_ Tesistant in the following order: Botrytis, Edocephalum, Penicil-, 


r 


400 BOTANICAL GAZETTE [ DECEMBER 


bottom of the cell instead of a solution similar in composition 
to that forming the hanging-drop. The variation in the toxic 
value indicated from the actual value will depend on the vapor 
pressures of the solutions used. Volatile—especially highly 
volatile—and hygroscopic solutions will show the greatest error. 

10. Many deleterious agents which at certain concentrations 
retard germination and early growth, afterwards cause a great 
acceleration of mycelial development in these retarded cultures. 
This abnormal development of mycelium is usually accompanied 
by retardation of fruiting. 

11. In the conidial stage the protoplasm of molds is in gen- 
eral more sensitive to the action of deleterious agents than at 
any other stage in their life history. 

12. The effect of the different deleterious agents on the 
appearance of the mycelium is very varied and often quite 
characteristic. 

13. One is not justified in drawing any conclusions as to the 
killing powers of an agent from its effect in inhibiting the ger 
mination of the spores. 

14. The hydroxyl group OH is rather more toxic to molds 


than ionic H. 
15. The toxic value of the halogens, Cl, Br, and I, in the 
ionic state, increases somewhat in the order of increasing atomic 
weight. 
16. The cyanogen radical is a very powerful poison to fungh 
KCN having nine times the toxic value of HCl. 

17. Mercuric chlorid and silver nitrate are about equally 
toxic to molds; and are followed in toxic properties by potas- 
sium dichromate and chromate, and formaldehyde. 

18. Strychnin and hydrocyanic acid, both extremely fatal 
poisons to the higher animals, and both supposed to act 0” the 
protoplasm of the nerve cells, react very differently toward 
fungi. To the molds strychnin is practically non-toxic, whereas 
hydrocyanic acid is a very violent poison. 

19. Nickel, cobalt, iron, copper, and zinc inhibit mo 
in the order named. Zinc is much less toxic than the oth 


ld spores 
ers. 


1899] TOXIC EFFECT OF DELETERIOUS AGENTS 401 


20. That an element is necessary for the nutrition of a plant | 
does not indicate whether it would or would not bea poison in 
greater concentration. See iron, oxygen, etc. 

21. That an element is not necessary for the normal devel- 
opment of a plant does not imply that it would be injurious even 
in relatively great concentration. See chlorin, calcium, etc. 

22. The ionization of the molecule of electrolytes in aqueous 
solution has a very important bearing on the study of the 
physiology of poisons. It is of especial value in determining 
the element or group of elements in a compound to which its 
toxic properties are to be attributed. 

23. In this study no new evidence has been adduced sup- 
porting the theory that the chemical activities of a substance are 
due wholly or chiefly to the ionized portion. 

24. Evidence has been adduced to the effect that in the 
case of several acids ionization /essens the chemical activities 
toward the substances involved in the life processes of the plant. 
25. In the case of the eight acids investigated six were found 
tobe much more toxic in the molecular form than after ioniza- 
tion. The toxic properties of the un-ionized molecules vary from 


épproximately 2.8 times that of ionic H in the case of acetic acid 


t f : 
to 76.6 times that of H in hydrocyanic acid. 
26. The substitution of Cl for H in the acetic acid radical 
has a double effect. In the first place it increases the toxicity 
of the un-ionized molecules to a greater or less extent depend- 
Ing on the number of H atoms so replaced. In the second place, 
_ it increases the ionization of the acid. The amount of the ioniza- 
tion is also dependent on the amount of H so replaced, being 
- Steatest, as are the toxic properties of the whole molecules, 
When all three H atoms have been replaced by Cl. 
i __ 27. These two factors to a great extent counterbalance each 
: other, Which has the greater influence in any given solution 
depends altogether on the concentration, the increased toxicity 
, % the molecules having the predominating influence at the 
: Steater concentrations, and the ionization being more effective 
: at the greater dilutions. 


402 BOTANICAL GAZETTE | DECEMBER 


28. At the concentrations inhibiting fungus spores mono- 
and dichloracetic acids are more influenced by the increased 
toxic properties of the molecule, and trichloracetic by the ioniza- 


F i. ee n ; ‘ 
tion. The former in —— and — concentration are respectively 
increased in toxicity 40 per cent. and 30 per cent. over the origi- 


nal acetic. Trichloracetic, on the contrary, at ae suffers a 


reduction of 10 per cent. in toxic properties as compared with 
the original acetic. 
29. The anions of the mineral acids, HCl, HNO,, and 
—fi,SO,, have a low toxic value for fungi, having less than one 
thirty-second that of ionic H. 


In conclusion I wish to acknowledge my indebtedness to Dr. 
B. M. Duggar, instructor ian plant physiology, and Professor 
George F. Atkinson, professor of botany in Cornell University, . 
for much valuable advice and assistance, and constant encourage- 
ment. My best thanks are also due to Professor W. D. Bancroft 
and Mr. A. L. Knisely of the chemical department for much 
help and information on the chemical aspects of the work. 


BOTANICAL LABORATORY, CORNELL UNIVERSITY. 


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Wesen. Jena, Zeitschr. 24: 402-470. 1890. 


SHIELDS, J. (’93): Uber Hydrolyse in wasserigen Salzlésungen. Zeit. phys- 
Chem. 12: 167-187. 1893. ee 
STEVENS, F. L. (’98): The effect of aqueous solutions upon the germination 


of fungus spores. Bot. Gaz. 26 : 377-406. 1898. 
STERNBERG, G. M. (’g2): Manual of Bacteriology, 1892. 
-TsuKAMoToO, M. (’95): On the poisonous action of alcohols u 

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Wuetuay, W.C. D. (’95): Solutions and electrolysis. Cambridge. 1895: 


Ann. of Bot. 


STUDIES IN CRATAGUS. 1. 
C. D, BEADLE, 

Few genera so widely distributed inthe United States have 
been so poorly interpreted by American botanists as the genus 
Crategus. In Europe there exists a better understanding of a 
number of American forms, a knowledge gained almost wholly 
from cultivated specimens, but it must be evident to any student 
of these interesting plants that all of the descriptions published 
up to the present time fall short of embracing the forms grow- 
ing in almost any section of the country. My attention was 
first drawn to the thorns about ten years ago, when I attempted 
to raise many thousands of young plants from seeds gathered in 
the southern Alleghany region. Making no attempt to propa- 
gate other than thrifty plants of the well-known and widely 
recognized species, I was perplexed to find a nursery of the most 
diverse forms under almost every label. Low and bushy plants 
With dark foliage stood in bold contrast with tall and fastigiate 
individuals with leaves of different tint and outline. No expla- 
Nation seemed more reasonable than that some careless gardener 
had mixed the seeds so painstakingly collected from fine, healthy 
individuals, and with a determination that more care should be 
- €xercised in succeeding attempts, the matter was dropped. The 
 ‘fext autumn the sowing was most carefully done, and while the 

fruits and seeds did not all look quite alike, it was easily demon- 
 Strated that the species recognized in our botanical field-books 
Were not confounded. The results of this seeding were even 
More confusing than the first, as the seeds were gathered from 
Wider fields, And thus the evidence accumulated each year, 
Until, almost unconsciously, I commenced to separate the forms as 
"ey Srew and the seeds as they were gathered. This acration 
: as complete, and as the oldest seedlings are now bearing the 
nds of fruits earlier recognized as strange, the publication of 
1899] we 


406 BOTANICAL GAZETTE ~ [ CECEMBER 


new species is fully justified. In this, and other papers which 
will appear as suitable material accumulates, I propose to deal 
with new forms coming to my attention, and to characterize the 
published species, with which are now confounded others with 
constant and widely different features. 

Crategus Biltmoreana, n. sp.—A branching shrub, 1-5™ tall, 
growing in dry or rocky woodlands: flowers, which appear 
when the leaves are almost fully grown, white, 2-2.5™ in diam- 
eter when expanded, produced in simple, pubescent, 3-7- 
flowered corymbs with lanceolate, pectinately-glandular, cadu- 
cous bracts; they are borne on strict, pubescent pedicels which 
vary from 7™"-2.5™ in length and open in the vicinity of Bilt- 
more, N. C. (type locality), about the twentieth of May: calyx 
obconic, pubescent or tomentose on the outside, with lanceolate, 
dentate or pectinate, glandular lobes about 5"" long which are 
reflexed after anthesis: petals broadly-obovate or nearly orbic- 
ular in outline, 8-12™, contracted near the base into short 
claws: stamens 10, shorter than the petals; the anthers pale 
yellow: styles 3-5, shorter than the stamens, surrounded at the 
base with pale hairs: fruit, which ripens and falls at the end of 
September or early in October, yellow, greenish-yellow or some- 
times orange, the parts exposed to the sun being rosy-cheeked 
or diffused with red, containing from 3-5 nutlets; they are 
depressed-globose, bluntly angled, 10-15" broad, 10-12" 
high, the cavity broad, 3-5™", and deep, 2-3", surrounded vy 
the calyx lobes and portions of the filaments: nutlets thick 
walled, 5-7" in greatest diameter, 3-5" measured dorso- 
ventrally, a volume of 125° containing about 1858 thoroughly 
cleaned, dry seeds; they are deeply grooved on the back and 
display a prominent ridge near the middle, the inner faces being 
nearly plane: leaves 2-6™ wide, 3-10™ long including the 
petiole, or occasionally larger, ovate, or round-ovate, acute at 
the apex; rounded, truncate or on vigorous shoots subcordate, 
but usually wedge-shaped at the base and prolonged into 4 
slightly winged or margined petiole 1-3 long and bearing, 45 
does the extreme base of the leaf, a few dark-colored stalked 


| 
| 


39] STUDIES IN CRATEGUS 407 


glands; borders acutely incised, or slightly 5-9-lobed and 
sharply and irregularly serrate to near the base: they are thin 
and membranaceous at flowering time, becoming firmer and 
thicker with age, harshly, though rather inconspicuously pubes- 
cent on both surfaces throughout the vegetating season, bright 
green on the upper surface, pale beneath, the prominent veins 
being disposed in 4-6 pairs: spines stout, 2-5™ long on the 
branches, slightly curved, dark chestnut-brown on the new and 
Stay on the old wood: bark of the main stem reddish-brown, 
slightly fissured and broken into many small, persistent, ashy- 
gray scales; that of the branchlets chestnut- or reddish-brown, 
or gray, sprinkled with small, pale lenticels: buds almost globu- 
lar, bright reddish-brown. 

Crategus Biltmoreana is distributed from North Carolina, 
northern Alabama, and eastern Tennessee to Virginia and Pennsyl- 
Vania. It has been usually confounded with and preserved in 
herbaria under the name C. mollis (T. & G.) Scheele,* from 
Which, or any form now resting under this name, the Biltmore 
thorn may be known by its smaller size, simple corymbs, later 
time of blossoming, and by the color and texture of the fruit. 
The type material is preserved in the Biltmore Herbarium. 

Crategus Sargenti, n. sp.—An intricately branched tree, sel- 
dom more than 6" tall, or more frequently a large shrub 2-5" in 
height, with one or several stems: the bark of the trunk ashy- 
Stay or light brown, slightly fissured and broken into many thin 
plate-like scales, or nearly smooth with. scattered patches of 
“PPressed, small scales: branches spreading or ascending, armed 
eh straight or curved, simple or branched, dark chestnut-brown 
gray spines 2.5-7™ long; they are intricately divided — 
- Mmerous, short branchlets which are clothed with dark reddish- 
brown bark and marked by small round or elongated pale oa 
ticels, forming a narrow, or occasionally round or sagt 
head: buds globular, bright reddish-brown: flowers, 
‘ppear about the first of May in the vicinity of Valley . ‘ 
| Alabama, and when the leaves are almost fully grown, dispose 


* Linnea ar: 569. 1848. 


408 BOTANICAL GAZETTE [DECEMBER 


in a few- (mostly 3-) flowered, more or less pubescent, simple 
corymbs: lateral pedicels longer than the intermediate ones, 
1.5-3.5 long, more or less pubescent or pilose: calyx obconic, 
pubescent, the segments glandular-serrate, 6-9" long, persistent 
or nearly so: corolla white, the divisions nearly round or a little 
broader than long: stamens normally 20, 5—7™" long: pistils 3-5, 
surrounded at the base with pale hairs: fruit, which ripens and 
falls after the middle of September, globose or depressed-globose, 
10-13™ broad, to—12™ high, yellow, orange-yellow or flushed 
with red, the flesh thin and firm ; cavity 3-5" broad and nearly 
as. deep, surrounded by the remnants of the stamens: nutlets 
3-5, but usually 4, hard and bony, the walls thick, 7-9™ long, 
4-6™" measured dorso-ventrally, with the back ridged and 
grooved and the lateral faces nearly plane: leaves thin to sub- 
coriaceous, sparsely pubescent when young, soon glabrous, 
yellowish-green on the upper surface, paler below. and displaying 
5-7 pairs of prominent veins; they are ovate, ovate-lanceolate 
or round-ovate, 2.5-12™ long, 1-6™ wide, or occasionally larger 
on vigorous shoots, acute at the apex, rounded or abruptly con- 
tracted at the base into a margined or winged, slightly glandular 
petiole, 5""-3.5™ long, the borders irregularly and doubly ser- 
rate and incisely lobed and the serratures minutely glandular- 
tipped: stipules linear to linear-lanceolate, glandular, or on 
strong shoots foliaceous, lunate, glandular-serrate, caducous. 

Crategus Sargenti is a remarkably distinct and showy species; 
especially in the autumn when the foliage assumes many lively 
tints of red and yellow. It inhabits the rocky woods and bluffs, 
or occasionally the rich, deep soil of the mountainous regions of 
northwestern Georgia, northern Alabama (extending 4s ag 
south as Birmingham), and southeastern Tennessee. The species 
belongs to an interesting and very natural group of several dis- 
tinct species of which no type has, so far as I have observed, 
been published. Many specimens of the related forms are aha 
served in herbaria under the names C. rotundifolia, C. glandulosa, 
C. coccinea, etc., titles which are, when correctly applied, asso- 
ciated with widely different plants. 


| 
| 
| 
1 
| 


1899] STUDIES IN CRATA:GUS 409 


I take pleasure in naming the species in honor of Professor 
C.S. Sargent, Director of the Arnold Arboretum of Harvard Uni- 
versity, who first called my attention to specimens collected by 
him near Rome, Georgia, in April 1899. The type material, 
which I had the opportunity of selecting from thousands of 
examples near Valley Head, Alabama (type locality), is pre- 
served in the Biltmore Herbarium. 

Crategus Boyntoni, n. sp.—A tree seldom more than 6” in 
height, or more frequently a large branching shrub, 2-4” tall, 
frequenting the banks of streams and even the shallow, dry soil 
of old fields and upland woods: flowers, which expand about 
thetenth of May in the vicinity of Biltmore, N.C. (type locality), 
and when the leaves are almost fully grown, 1.75-2.25™ in 
diameter, produced in short, glandular-bracteate, 4—10-flowered 
corymbs: pedicels 7™"—1.5™ long, glabrous, bearing one or two 
glandular or pectinately-glandular bractlets : calyx obconic, 
smooth, the divisions acute, glandular serrate, 4-6"" long: petals 
White, nearly orbicular, or a little broader than long, with a short 
and broad claw at the base, g-12™" in diameter: stamens 10, 
6-9" long, the anthers light yellow: pistils 3-5, surrounded at 
the base with pale hairs: fruit dull, yellowish-green, flushed with 
lusset-red, depressed-globose, angled, 10-14™ high, 12-16™ 
broad, ripening and falling early in October: nutlets 3-5, hard 
and bony, 6—8™™ long, 4-5™™ measured dorso-ventrally, the back 
tidged and grooved and the lateral faces nearly plane, a volume 
of 125° containing about 1293 thoroughly clean and dry seeds: 
leaves at first membranaceous, becoming subcoriaceous with age, 
Yellowish-green on the upper, paler on the lower surface, glab- 
_ Tous, or with a few scattered hairs along the midrib and larger 
Veins, which are disposed in 4~—7 pairs; they are broadly ovate 
*Foval in outline, acute at the apex, rounded or narrowed at se 
base and prolonged into a margined, glandular petiole 1-2.5 
‘ong, or on vigorous shoots deltoid-ovate and truncate or sub- 
_ *©rdate at the base; the borders are sharply and ir regularly woul 
Tate, doubly serrate or incisely 5—7-lobed, the serratures minutely 
- Sland-tipped: stipules linear, glandular, caducous, or on strong 


410 BOTANICAL GAZETTE [DECEMBER 


shoots foliaceous, lunate, glandular-serrate : trunk, which is 2-3" 
long and 1-2" in diameter, and occasionally armed with gray, 
branched spines, covered with ashy-gray bark, not infrequently 
tinged with brown, or in the shade of the forest dark-gray, 
slightly fissured and broken into many small, plate-like scales: 
branches stout, ascending, armed with dark, chestnut-brown or 
gray, straight or slightly curved spines, 3-7™ long, or larger, the 
bark close and smooth, dark-gray or brown: branchlets smooth, 
the bark dark reddish-brown, sprinkled with small, round or 
elongated, pale lenticels, the whole forming a narrow ofr occa- 
sionally round or flat-topped tree, or a much-branched oval or 
pyramidal shrub. 

Crategus Boyntoni is distributed from Georgia, Alabama, and 
Tennessee to Virginia, Pennsylvania, and Delaware, and will 
doubtless be found to extend over a broader area when better — 
known. It is closely related to C. Sargenti, above proposed, but _ 
may be separated by the numerous-flowered, glabrous corymbs, 
shorter pedicels, and fewer stamens, and by the different habit of 
growth. Many specimens are preserved in herbaria, the greater 
part of which are also labeled C. coccinea, C. glandulosa, or ©. 
rotundifolia, My attention was first directed to this form by Mr. 
.F. E. Boynton, of the Biltmore Herbarium, for whom the 
Species is named, by the exhibition of plants loaded with the 
characteristic and distinct fruit, and of branches displaying the 
glandular and brightly colored characters of the unfolding leaves 
and shoots. To this species I refer, in part, the material repre- 
senting the southern distribution of C. rotundifolia of the Illus- 
trated Flora’, and the note under this name published by me !” 
the Boranicat Gazetre3, The type material is preserved in the 
Biltmore Herbarium. 

CRATAGUS TRIFLORA Chapm. Flora Southern United States, 
Suppl. II. 684. 1892 [Ed. 2].—A large shrub or small tree, 
2-7" tall, growing on limestone bluffs near Rome, Ga. (tyPe 
locality), and in similar situations near Birmingham, Ala. Main 

* Ill. Flora Northern U. S., etc., 2: 243. 1897. 

3Bor, Gaz. 25: 446. 1808. 


1899 | STUDIES 1N CRATA:GUS 411 


stem single, or branching near the base into several spreading 
shoots, armed with numerous branched spines: bark light gray, 
or with a decided tinge of brown, either close or slightly fur- 
towed and broken into many small, plate-like scales: branches 
ascending, clothed with light gray, smooth bark and bearing a 
few simple or branched, straight or slightly curved, chestnut- 
brown to gray spines, 2.5-6™ long; they are intricately divided 
near the summit into many short, pilose or pubescent, brown or 
dark reddish-brown branchlets, forming an oblong or occa- 
sionally round or flat-topped head: buds globular, bright 
reddish-brown, the terminal one on strong shoots displaying 
~ several spreading scales: leaves, which are full grown at flower- 
ing time, thin at first, becoming subcoriaceous, dark green above, 
paler below, ovate, elliptical, or slightly obovate, acute at the 
apex, rounded or abruptly contracted at the base into a winged 
or margined petiole, 8""-3.5™ long: they vary in size from 
those on the fertile branches, which are 2-10" long including 
the petiole, 1-5 wide, to large, broad leaves on vigorous 
Shoots, 13™ long, 8™ wide, the borders being sharply and 
irregularly serrate, or doubly serrate and incisely lobed and 
bearing, as does the petiole, a number of stalked, black-tipped 
glands near the base; the upper surface harshly and rather 
inconspicuously pubescent throughout the vegetating season, 
below more densely and permanently pubescent, especially 
along the 5-7 prominent veins: stipules lanceolate or oblong or 
on the stronger shoots foliaceous and lunate, densely glandular 
or glandular-serrate, caducous: flowers, which open about the 
first of May, disposed in few-, mostly 3-flowered, pilose corymbs : 
pedicels 1~3.5™ long, the lateral much longer than ne rae 
mediate one, bearing a densely or pectinately glandular, decia- 
Yous bractlet: calyx obconic, densely pilose, the Stee 
Persistent, pubescent, glandular-serrate, ee as i 2-4 ae ee 
acute: corolla 2-3 wide, the divisions. orbicular, 10-13" 1 
diameter: stamens normally 20, 5-7" long: styles eee 
founded at the base with pale hairs: fruit, which ripens oat 
the middle of September, globose, 12-15" in diameter, brig 


412 BOTANICAL GAZETTE [DECEMBER 


or deep red, pubescent, the cavity 5—6™ broad, 3~57": deep: 
nutlets 3-5, hard and bony, 7-9™ long, about 5"™" measured 
dorso-ventrally, a volume of 125° containing about 1633 clean 
and dry seeds. 

The type material is preserved in the Chapman Herbarium 
at Biltmore. 

Crategus austromontana, n. sp.—A straggling shrub, 1-4” in 
height, frequenting rocky woods and banks: main stems 1-3, aris- 
ing from large coarse roots or horizontal rootstocks, or frequently 
a group or clump more or less united, occupying a surface of 
5-10 square meters: bark close, or slightly broken into numer- 
ous small plates, gray, or with a decided tinge of brown: 
branches unarmed, or an occasional spine on a young plant or 
vigorous shoot, the bark smooth, gray or nearly brown: branch- 
lets pilose-pubescent or tomentose, covered with dark chestnut 
or reddish-brown bark which is marked by round or elongated, 
pale lenticels, buds globular, bright reddish-brown: flowers 
large, borne in simple 2~5- (usually 3-) flowered cymes, opening 
in the vicinity of Valley Head, Ala. (type locality), the first 
part of May: pedicels stout, pilose or tomentose, I-2.5™ long, 
bearing a linear, glandular, deciduous bract near the summit: 
calyx. broad, . obconic, pubescent, the divisions lanceolate, 
6-10" long, 1-3™ wide, pectinately-glandular or glandular- 
serrate, pubescent: stamens 10, 4—7™ long: pistils 3-5, sur 
rounded at the base with pale hairs : fruit, which ripens and falls 
during the latter part of September, large, globose, 12-15™ 11 
diameter, bright red, pubescent, and frequently punctate, con 
taining 3-5 hard, bony nutlets which are 8—10"™ long, so 
wide, measured from the back to the inner edge, bluntly angled 
on the back and with lateral faces narrow and nearly plane: 
cavity broad, 4-6" wide, surrounded by the persistent calyx 
lobes and remnants of the filaments: leaves orbicular, broadly 
oval or round-ovate, 3.5-12™ long, including the pubescent oF 
tomentose petiole, which, like the extreme base of the leaf, bears 
a number of stalked, black-tipped glands, 2.5-7.5™ wide, oF 
sometimes larger on strong shoots; they are harshly, though 


ee aT eT ee 


1899] STUDIES IN CRATEGUS 413 


rather inconspicuously, pubescent on both surfaces, the pubes- 
cence being more pronounced on the lower side and along the 
principal veins, which are disposed in about 5-7 pairs; acute 
at the apex, contracted at the rounded, truncate or sometimes 
subcordate base into a margined or winged petiole 1-4™ long, 
the borders very sharply and irregularly serrate, or frequently 
doubly serrate or incisely lobed, the serratures tipped with 
minute dark colored glands. 

Crategus austromontana is distributed throughout the Sand 
mountain region of Alabama, and has also been collected at 
several stations in the Cumberland mountains and hill country 
of eastern and middle Tennessee. The new species is closely 
associated with C. triflora Chapm., but may be recognized by its 
smaller size, broader leaves, fewer stamens, and by the larger 
and coarser seeds. The type material is preserved in the Bilt- 
more Herbarium. 

Crategus Harbisoni, n. sp.—A tree 5-8 meters in height, 


frequenting rocky slopes and ridges: leaves obovate, oval, or 


broadly ovate, 3-12 long including the petiole, 2-9 wide, 
acute at the apex, narrowed at the rounded or tapering base into 
margined or winged petioles; they are harshly and rather 


| Inconspicuously pubescent on the upper and more densely coated 


on the lower surface and along the 5-7 principal veins, sub- 
coriaceous, dark green and lustrous above, pale below, the 
borders doubly and irregularly serrate to near the base, or fre- 
quently incisely lobed: petioles 6™-2™ long, bearing, = does 
the extreme base of the blade, a number of stalked, black-tipped 
glands : stipules glandular-serrate or pectinately-glandular, 
deciduous, foliaceous on the stronger shoots, acute or lanate: 
°wers, which appear in early May in the vicinity of Nashville, 
Tennessee (type locality), produced in broad, pubescent, or 
Pilose, divergently-branched corymbs, the lower branches from 
the axils of leaves: bracts subtending the branches of the 
orymbs large, 7-18™ long, 2-4™ broad, pectinately or 
glandular-serrate, caducous: pedicels pubescent or pilose, stout, 
3.5" long, bearing a pectinately-glandular, elongated, 


414 BOTANICAL GAZETTE [DECEMBER 


deciduous bractlet : calyx obconic, pubescent, the lobes lanceo- 
late, glandular-serrate, persistent : stamens normally 20, 4-6™" 
long: pistils 3-5, surrounded at the base with pale hairs: fruit 
large, red, or orange-red, globose, 10-13”" in diameter, pubes- 
cent or smooth, punctate, ripening in October: nutlets 3-5, 
6-g™ long, 4-5"" from back to inner angle, either furrowed 
on the dorsal side or with a blunt ridge and two grooves, the 
lateral surfaces nearly plane, a volume of 125° containing about 
1490 clean and dry seeds; cavity 4-6"" wide: bark of the 
trunk, which is from 1—2°" in diameter and I-—2 meters long, 
close or slightly fissured and scaly, ashy-gray or darker in color 
and frequently armed with simple or branched spines: branches 
clothed. with smooth, gray or light brown bark, and_ bearing 
straight or curved chestnut-brown or gray spines 3-6™ long, 
the growth of the season reddish-brown in color and marked by 
small, pale lenticels: buds nearly round, bright reddish-brown, 
the terminal one displaying several large, spreading, acute 
scales. 

Crategus Harbisoni was discovered by Mr. T. G. Harbison of 
the Biltmore Herbarium, for whom the species is named, on the 
limestone hills and ridges near Nashville, Tennessee, where 
numerous examples were observed at intervals during the past 
summer. 

The new species probably belongs to a group of which C. 
triflora Chapm. and the C. austromontana previously proposed are 
types. From the former it may be separated by the compound, 
many-flowered corymbs, and from the latter by its greater size, 
numerous stamens, spiny branches, large flower-clusters, and the 
different habit of growth. The type material is preserved in the 
Biltmore Herbarium. 

Crategus silvicola, n. sp.—A tree, attaining in low moist 
woodlandsa height of 6-10”, or in upland forests a shrub with 
one or more stems, 1—s™ tall: trunk, which is sometimes 2 
in diameter, covered with a close or slightly fissured and scaly, 
gray or reddish-brown bark, and armed with stout, branched 
spines: branches, which are spreading or ascending and form 4 


= 


1899] STUDIES IN CRAT&GUS 415 


round or flat-topped head, armed with straight or curved, chestnut- 
brown or gray spines 2—6™ long, and clothed with gray or light 
brown bark: branchlets chestnut- or reddish-brown, sprinkled 
with small pale lenticels: buds globular, bright reddish-brown: 
leaves ovate, round ovate or on vigorous shoots deltoid, acute at 
the apex, rounded and narrowed at the base, or occasionally 
truncate or subcordate, 3-10 long, including the petiole, 
2-6" wide, the borders sharply and irregularly serrate, or 
doubly serrate and incisely 5--7-lobed, the serratures minutely 
glandular-apiculate; they are bright or yellowish-green and 
minutely roughened, or occasionally scabrous-pubescent on the 
upper surface, paler below and generally smooth, or with a few 
hairs along the larger veins, which are disposed in 3-5 pairs : 
petioles slender, 5""-3™ long, glandular: pedicels strict, 7°” 
-1.5"long: calyx obconic, the divisions short, entire or glan- 
dular-serrate, 3—-4™" long, acute: stamens 10: styles 3-5, sur- 
rounded at the base with pale hairs: fruit globose, 10-11™ in 
diameter, red or greenish-yellow with ruddy cheek, ripening and © 
falling the last of September in the vicinity of Gadsden, Ala- 
bama (type locality): nutlets 3-5, hard and bony, 5-6"" long, 
3-4" measured dorso-ventrally, the back ridged and grooved 
and the lateral faces nearly plane: flesh thin and firm. 

Crategus silvicola is abundantly represented in the ‘‘ flat- 
Woods” of northern Alabama and northwestern Georgia, and 
®ccasionally ascends into the poorer and drier woodlands of the 
‘Surrounding country. The lower leaves and those from young 
Plants are much rougher to the touch than foliage from the 
upper branches of the larger and older trees, but the fruit, which 


When ripe falls from the trees at the slightest interference, iS. 


‘onstant in form and color. I presume the proposed species 
Tepresents one of the several forms of the South Atlantic region 
heretofore referred to C. coccinea, L.1 Taking the original 
description and a copy of a tracing of the type specimen of the 
Scarlet thorn, C. silvicola may be distinguished by the rough 


; i ine,. 
: leaves, which are less incised and broader and longer in outlin 


‘Sp. Pl. 476. 1753. 


416 BOTANICAL GAZETTE [DECEMBER 


and by the short, strict, and stout pedicels. The type material 
is preserved in the Biltmore Herbarium. 
Crategus Mohri, n. sp —A tree 6-10” tall with a slender 
trunk 1~2 in diameter, unarmed, or sparsely furnished with 
long, simple or branched spines, and covered with thin, scaly, 
ashy-gray or reddish-brown bark; or in unfavorable situations a 
large, erect and branching shrub: branches ascending or nearly 
horizontal, spiny, forming an oblong or occasionally round com- 
pact head; the bark close and usually gray: branchlets and 
smaller branches zig-zag, armed with slightly curved or straight, 
chestnut-brown or gray spines 2-5™ long; the growth of the 
season at first pubescent, clothed with light brown or gray, lus- 
trous bark which is marked by small, pale lenticels: buds nearly 
round, or the lateral occasionally compressed, bright reddish- 
brown: leaves cuneate-obovate, or on vigorous shoots varying 
from oval to orbicular, 2~7™ long including the petiole, ak 
wide, or larger on the shoots, acute or rounded at the apex and 
contracted below into winged or margined petioles in 
long, sharply and irregularly serrate to or below the middle, 
entire or nearly so at the base, and sometimes, especially on 
vigorous shoots, doubly serrate or incisely lobed ; they are more 
or less pubescent along the veins when young, dark green and 
lustrous above, pale below, becoming thick and coriaceous with 
age: stipules linear, glandular, 5-10" long, caducous: flowers, 
which appear when the leaves are of full size, disposed in slen- 
der, elongated and often flexuous, many-flowered, bracteate 
corymbs, which are more or less pubescent or pilose at flowering 
time, and open in the vicinity of Rome, Georgia (type locality ) 
about the first of May; they are 1.5-1.75™ in diameter an 
borne on more or less pilose, slender, often flexuous pedicels 
8™—2.5™ long, which bear one or two minute, subulate, cadu- 
cous bractlets: calyx narrow, obconic, glabrous or occasionally 
a little pilose; the segments linear-lanceolate, 2-5"" long, entire 
or slightly glandular serrate,. reflexed after anthesis; corolla 
white, the divisions round-ovate or nearly orbicular, 6-8"™ long, 
5-7"" wide with undulate or erose borders: stamens normally 


1899 ] STUDIES IN CRATA:GUS 417 


20, 3-5" long: styles 3-5, surrounded at the base with pale 
hairs: fruit globose, 8—-9"" in diameter, dull, dark-red or green- 
ish-red, or frequently covered with black spots and blotches, 
tripe about the first of October and hanging until early winter, 
the cavity 2-3"" wide and deep, bordered by the remnants of 
the calyx lobes and stamens: nutlets 3-5, thick-walled, 5—7"" 
long, about 3°" wide measured dorso-ventrally, with a promi- 
nent ridge and two deep grooves on the back, the inner faces 
nearly plane. 

Crategus Mohri is distributed from Georgia westward through 
upper and central Alabama and Mississippi, and northward to 
middle Tennessee. It reaches its best development in the rich 
and moist soil of the ‘“flat-woods” of central Alabama, though 
not infrequently it ascends into the poorer and drier soil of the 
hills and mountains. The species has been usually confounded 
with Crategus crus-galli L.,3 or more recently with C. collina 
Chapm.6 From the former it may be recognized by the pilose 
corymbs, smaller and globular fruit, more numerous and smaller 
nutlets, habit of growth, and by the outline of the leaves; while 
from C. collina it may be separated by the later time of flower- 
ing, smaller fruit and nutlets, and by the luster of the leaves. 

I take pleasure in associating with this beautiful and most 
distinct hawthorn, the name of Dr. Charles Mohr, of Mobile, 
Alabama. 

The type material is preserved in the Biltmore Herbarium. 


BILTMORE HERBARIUM. 


5Sp. Pl. 476. 1753. 
°Flora S. U. S., suppl. II. 684. 1892. [Ed. 2.] 


BRIEPER ARTICLES. 


SOME PECULIARITIES IN PUCCINIA TELEUTOSPORES.' 
(WITH SIX FIGURES) 


Tue distinguishing characteristics of many Puccinia teleutospores 
are very slight, while on the other hand such species as P. podophylli, 
coronata, pruni, etc., are, in typical specimens, at once set aside from all 
others by the markings of their epispores. In many species, variations 
in the number and position of the septa are characteristic. Generally, 
however, the shape and size of the spores are fairly constant, and this is 
particularly true of P. graminis Pers. and allied gramineous rusts. 
Occasionally one-celled spores are found, due perhaps, as Professor 
Burrill* suggests, to insufficient nutrition. Eriksson and Henning’ 
call attention to the so-called mesospores and other peculiarities found 
among spores from closed sori of P. graminis, attributing them to the 
pressure of the epidermis, and Bolley* in regard to the anomalous 
shapes found among spores says “certain it is that pressure within the 
crowded sorus is capable of producing an almost unlimited number Of 
irregularities in the spore forms.” J. A. Warren’ has written regard- 
ing some very striking variations in the spores of P. Windsorie Schw. 
while Dietel,° writing on peculiarities in Puccinia spores, refers to the 
finding of a well-developed four-celled spore of P. graminis, and habe 
time to time the odd shapes of rust spores have been noted by various 
writers. : 

Puccinia heterospora B. & C. is a species showing many interesting 
variations, giving as they do an indication of the close relationship © 

*Contribution from Botanical Dept. Iowa State College of Agriculture - 
Mechanic Arts, no. 16 

* Parasitic fungi of Illinois 1: 171. 1885. 

3Die Getreideroste 129. #/. 4. 

‘Sub-epidermal rusts. Bot. Gaz. 14: 139-144. p/."75. Je. 1889. 

5 Notes on the variations of Puccinia Windsoriae, Am. Nat. 32: 779-781: 4 kes 
1898. 


: Jatt 32 * 
*Beitrige zur Morphologie und Biologie der Uredineen. Bot. Centralblatt 3 
86-88. 1887, 


418 [pacer 


1899] BRIEFER ARTICLES 419 


the Uromyces and Puccinia genera of the Uredinee; in fact, this 
species must be regarded as one of the connecting links between the 
two. The spores are of two kinds, one and two-celled, the one-celled 
being globose to subglobose, and measuring 19 X 28. 


Maha avs 


Fic. 1.—Teleutospores of Puccinia heterospora. 4, one-celled; 4, two-celled. 


The two-celled spores are of three kinds : some have the septum 
transverse, in some it is oblique, while in others it coincides with 
the axis of the spore. The measurements do not differ materially from 
those given for the one-celled spores. The one-celled forms are much 
more numerous and the epispores of all are thick and smooth. 

_ In 1884, Dr. Trelease’ described a species occurring on Bromus 
ciliatus and called it P. tomipara. He says, “this species is remarkable 


Ins sgesnd 


. Fic, 2.—Teleutospores of Puccinia tomipara. 4, one-celled; 4, two-celled; ¢, 
_three-celled; ¢, four-celled. y 


e or four-celled with 
tly parallel to the axis 
than one row of 
s of this species 


ftom the fact that the spores are commonly thre 
the uppermost septum oblique or not infrequen 
of the spore, which is thus made to consist of more 
cells.” To this comment on the very variable spore 
Nothing can be added. 
~~ Puccinia irregularis E. & T. is another species 
teleutospores, one, two, and three-celled spores bein 


showing variable 
g found. The 


’ Preliminary list of parasitic fungi of Wisconsin 22-23. N. 1884. 


420 BOTANICAL GAZETTE | DECEMBER 


septa are always transverse. In addition to the variation in the number 
of cells, the spores are peculiar because of the position of strongly 
developed papillae. Usually a single papilla is found at the apex of 
the spore, but often the spores are truncate, when two papillae appear, 


cero 


Fic. 3.—Teleutospores of Puccinia irregularis. a, one-celled; 4, with papilla 
below septum; c, with side thickening ; ¢, two- and three-celled. 


one on each side. In some the papilla is found just below the septum. 
One spore was observed having the whole of one side thickened instead 
of the apex. The spores measure 50-75 X19~-28m, the one-celled 
spores being somewhat smaller. ‘ 

An examination of the material in the herbaria of the Missoum 
Botanical Garden and the Iowa State College leads to the belief that 
this is undoubtedly the species referred to P. Solidaginis Pk. by Dr. Tre- 
lease, no. 169, Preliminary list of parasitic fungi of Wisconsin. 


AE@Tey 


Fic. 4.—Teleutospores of Puccinia Montanensis. a, normal; 4, three-celled; % 
four-celled; d, five-celled. 


A specimen of Puccinia occurring on Elymus robustus Was collected 
at Ames, Iowa, by G. W. Carver, October 14, 1895. The ee 
number of the spores were quite normal in shape and just what mig 


i, 5 


eStart etn een eee 


set 


Pe Ss crac hid ee ee AN A ty acme MRD PN ge see erat G Tome MS 


1899 | BRIEFER ARTICLES 421 
be expected in a Puccinia, but among them were found a few spores of 
more than two cells. The multicellular spores were in some cases not 


unlike those of P. ¢riarticulata B. & C., but owing to their scarcity and 
the fact that the two-celled spores agree in size and shape with those 


Wag 


a, normal; 4, three-celled; c, 


Fic. 5.—Teleutospores of Puccinia Rubigo-vera. 
four-celled. 
of P. Montanensis Ell. it is best to regard it as that species. The posi- 
tion of the septa in some of the spores would be very hard to des- 
cribe. Some spores were quite regularly three and four-celled, while 
in others oblique, transverse, and perpendicular septa were present, all 
in one spore. The three-celled forms were of two kinds, regular ones, 
and those in which two cells were found at the apex with one at the 
base, but occasionally this was just reversed. The measurements varied 
from 33 X 25m to 5533p 

A specimen of Puccinta Rubigo-vera (DC.) Wint. on Agropyron 
tenerum, collected at Ames in 1896, showed a considerable number of 
spores having three and four cells. The normal two-celled ge 
were greatly in excess of the others, being found in the proportion 0 
about fifty to one. The three-celled forms were usually somewhat 
irregular, though occasionally one was found in which the septa eee 
the spore at the right angles to the lateral walls. The four-celle 
forms were fewer in number than the three-celled, only a half dozen 
being found in three mounts. The septa were so placed as to divide er 
Spore in different planes. ‘The measurements of all were confine 
Within the usual limits for this species, 13-20% 28-55 ee 

On November 24, 1898, the writer picked up a piece of Ane 
Oat straw (Avena sativa L.) on the road in front of the von * = . 
at the Iowa State College. The rust sori presented all the noms % 
the sori of P. graminis Pers., appearing on the sheath as black 


oval confluent patches. 


422 BOTANICAL GAZETTE [DECEMBER | 


“ID 22) 


& <@ 


QD) => =D SO 


UC 


mmmioe ne mm e aa ‘3 my 
a ifr’ stn, a see “Si. 3 
aes ess 
“ Nowe ye sabe 8 


| 


e 


ae ; rmal; ¢-4 
Fic. 6.—Teleutospores of Puccinia graminis. a-a, one-celled ; See ee, four- 
three-celled, septum horizontal; ¢-d, three- and four-celled, septum obliq 
celled, septum horizontal. 


1899] BRIEFER ARTICLES " 423 


A microscopical examination made sometime afterward revealed 
some noteworthy peculiarities in the shape, size, and number of cells 
in the teleutospores. The usual two-celled spores were present but 
accompanied by others having one, three, and four cells, 

The different forms were about equal in number but differed con- 
siderably in size, the four-celled variety being the largest, as one would 
naturally expect. Of each form ten measurements were made, giving 
the following extremes: one-celled, 27-36 X15-20n; two-celled, 30— 
45X 15-21; three-celled, 45-54 15-21; four-celled, 52-66 15— 
20m. 

The one-celled spores might easily have passed for the teleuto- 
spores of some Uromyces such as U. graminicola Burrill. The two- 
celled ones were quite normal. in size and shape, except that in some 
the pedicels were much stouter than are usually found in P. graminis, 
more closely resembling the pedicels of P. emaculata Schw. The three 
and four-celled forms were of three kinds, some having the upper septum 
horizontal, some oblique, and others vertical, as though the upper cell 
had been formed as a sort of afterthought, by the division of the second 
or third cell as the case might be. 

In these the evolutionary development of several genera of Ure- 
dinae could be plainly traced, passing from the lower Uromyces 
through Puccinia and Triphragmium to Phragmidium. The spores, 
aside from the number of cells, were not likely to be mistaken for 
those of Triphragmium or ‘Phragmidium, as they were quite different 
in general appearance. ‘These genera have undoubtedly a common 
origin, and must be looked upon as being more highly developed, 
more specialized, in direct relation to the number of cells in the spores, 
as itis quite apparent that a larger number of sporidia can be pro- 
duced with less effort in those having the larger number of divisions 
in the spores.— H. HaroLp Hume, /owa State College, Ames. 


WHAT IS PRUNUS INSITITIA? 

In the June number of the BoranicaL GazeTTE there appeared 
‘n article under the heading given above and written by Frovessor 
P. A. Waugh. The conclusion to which the author qr: his own 

Words, is “that there is no such species as Prunus insiitia. 

: n 

To me this seems rather strange. I happen to have a 
the land of Linneus and received a large portion of my botan 


424 - BOTANICAL GAZETTE [ DECEMBER 


education at the Royal Gymnasium at Skara, Sweden. As a boy, I 
used to pick and eat the fruit of what there is known as Prunus insititia, 
and as a young botanist I made herbarium specimens theréof. I know 
that the tree which goes under that name is more distinct from P. 
domestica, as well as from P. spinosa, than P. hortulana, or P. nigra, or 
even P. angustifolia is from P. Americana. I know that there are at 
least three species of plums in Sweden, for I have seen them myself. 

The latest catalogue of the plants of Scandinavia, published in 1897, 
also gives the following plums: “ Prunus spinosa L., P. spinosa coata- 
nea W. &. Gr., P. insititia L., P. insititia rustica Hn., and P. domestica 
L.” Of these the first three are recognized as being natives of Sweden, 
while P. domestica and P. insititia rustica are regarded as only escaped 
from cultivation. So far as I know, P. cnsititia L. has always been 
regarded as a good species in Sweden; but let us see how botanists of 
other countries have treated it. 

As Linnzus in the original description stated that P. cusititia is a 
native of England and Germany, it will suffice to see how the botanists 
of those countries have treated the species. In almost every German 
flora P. insititia is regarded as a good species. Koch, the acknowl- 
edged authority in Germany, recognized it, and in Thomé’s elaborate 
work there is an excellent description. 

It is true that Bentham put P. domestica, P. insititia, and P. spinosa 
into P. communis Huds.; but Hooker, who has always been known for 
his conservatism, recognized all three as distinct species, not to men- 
tion other less important English botanists. It is figured in Sowerby’s 
English Botany 12 : 841. 

With these facts in view, it is surprising that one who has not studied 
the native plums of Europe in the field, with the few specimens found 
in the American herbaria, undertakes to settle the existence or non- 
existence of P. ¢nsititia, and can state positively ‘that there is no such 
species as P. insititia.”’ 

If Professor Waugh had said that P. insititia is the same as P. domes- 
“ica Damascena, or that P. insititia is not found in America, I should 
have been the last to criticise. I have not the means to disprove the 
former, and I am more than willing to accept the latter. /?. domestica 
Damascena \. was based upon “ Pruna majora dulcia et parva cain 
caerulea, Bauh. pin. 443, 0.23,” and P. insititia L. on “ Pruna sylvestria 
praccocta, Bauh. pin. 444.” Apparently, therefore, they seem to 
two different things. For that matter they might well belong to 


1899] BRIEFER ARTICLES 425 


the same species. Koch states in the older edition of his Flora 
that Linnzeus included several forms in P. domestica, which rightfully 
belonged to PP. insititia. Even this question has to be settled in 
Europe. 

As to the non-existence of P. ¢nsititia in America, I agree fully 
with Professor Waugh, for the following reasons: If Dr. Gray had 
had what is known as FP. insititia in Sweden, I doubt that he would 
have made it a variety of P. spinosa. Dr. Gray’s statement that it is 
“adventitious in hedgerows” made me very suspicious when I saw it 
in his manual a year or two ago; for P. inszti#ia, so far as 1 know, is 
never used for hedges. I think that P. msifitia should be erased from 
the list of American plants.—P. A. RypBerc, Wew York Botanical 
Garden. 


NOTES ON THOREA. 
(WITH PLATE XXVI) 


On October 1, 1898, Mr. A. A. Hunter, collector for the botanical 
department of the University of Nebraska, found specimens of Zhorea 
ramosissima Bory in Rock creek, a small stream near Lincoln, Neb.’ 
The plants were floating from a gravelly bottom in swift running 
water at a depth of half a meter and were surrounded by a mass of 
other algee, principally Vaucheria. Subsequent search for Thorea in 
this locality has thus far proved unavailing. 

So far as we know, Thorea has been found toa certainty ay but 
three other localities in North America. E. Hall collected a specimen 
of Thorea ramosissima Bory in the Sangamon river, Illinois, in 1866, 
and this, with specimens of other fresh water alge, was afterwards sent 
to the Botanical Museum of Berlin, where it is still preserved.” 
Francis Wolle found a mere fragment of Thorea in a lake at wintes 
Park, F lorida, date not given.’ Professor De Alton Saunders,’ in 
December 1898, found Thorea in abundance in running water from 
springs in Texas, the stations being San Marcos (Hayes county ee 
Braunfels (Comal county), and San Antonio (Bexar county). 

*See notice in Bor. Gaz. 27:71. 1899. 

Us: Thorea ramosissima Bory bei Belgrad in Serbien un 
- Hedwigia 38:114. 1899. 
*WOLLE: Fresh water algze of the United States, 58. 1887. 

n formalin. 


d denen weitere 


- ®Macn 
Verbreitung 


‘Communicated in a letter, accompanied by specimens ! 


426 BOTANICAL GAZETTE [ DECEMBER 


Thorea is widely distributed over the world, having been reported 
from France, Germany, England, Denmark, Austria, Venezuela, Ecua- 
dor, Java, and the Marianne islands. In Ecuador it is said to be espe- 
cially abundant. 

Our description of the Nebraska plant does not differ essentially 
from that of Schmidle in his excellent monograph of Zhorea ramosis- 
sima Bory In our plant the body consists of long cylindrical 
branches, originating near the base, and these again have occasional 
branches. The color is an olive green, rather than the black or brown 
color mentioned by Schmidle. When dried, the plant retains its olive 
green color, becoming somewhat brownish. ‘The whole plant is about 
5°" long, and 2 to 3™ wide, when floating in the water. Each branch 
consists of two distinct portions, viz., an outer covering of several- 
celled hairs or ramelli, and a denser axial portion of interlacing 
cellular filaments, which are held more firmly together by a mucilagi- 
nous matrix which sheaths every fiber and extends outwards as far as 
the first two or three basal cells of the hairs. The surrounding zone of 
hairs. has a width of from 400 to 6oop, being of nearly the same 
diameter upon all portions of the plant body, except at the base. 
The axial portion has a varying diameter, ranging from 700p at the 
base to less than 1oop at the growing point of the branch. 

The hairs grow at right angles to the axis and constitute two quite 
distinct belts; an outer belt of quite evenly distributed long hairs, 
having an average length of soom; and an inner belt of clustered, 
short hairs of an average length of 70 to gow. The long and short 
hairs are intermingled, and both kinds spring from the same basal 
cell. The short hairs are protected by gelatinous sheaths, which 
are extensions of the central gelatinous matrix. The cells of the long 
hairs are rectangular in shape, and quite uniform in diameter. The 
short hairs have shorter cells, which are also nearly uniform in diam- 
eter, yet in some cases they taper slightly towards the apex. In the 
older portions of the branches the short hairs are more numerous, 
while in the younger region the long hairs predominate. As the plant 
matures, the apical cell of the short hairs often develops into an 
asexual spore (aplanospore). Among the short hairs, and often from 
the same basal cells, there may arise narrow hairs which develop 4 
small cluster Of similar asexual spores, rarely over five in number. 

SSCHMIDLE: Untersuchungen iiber Zhorea ramosissima Bory. Hedwigia 
35 :1-33. 1896. 


1899 ] BRIEFER ARTICLES 427 


In both cases the spores when young are spherical, and when mature 
the former are oval, the latter pyriform. 

The axis consists of three structural parts: (1) a more or less dis- 
tinct outer portion, of irregular basal cells from which the hairs origi- 
nate; (2) a belt of interlacing cellular fibers most of which run 
longitudinally in the axis; these fibers are connected either to certain 
hair clusters by basal cells, or in some cases to a single long hair; 
there are frequent oblique or transverse fibers among the longitudinal ; 
(3) the innermost portion of the axis consisting of an interlacing mass 
of cellular fibers running in all directions; these are connections or 
continuations of the outer longitudinal and transverse fibers. 

Each hair cell contains greenish, disk-shaped chromatophores, 
and a distinct nucleus. There is also a distinct protoplasmic connec- 
tion between the cells through the center of each cell partition. 
The fibers near the outer edge of the axis, especially those directly 
connected to the hairs by basal cells, contain chlorophyll bodies more 
or less irregular in shape, and show in many Cases intercellular proto- 
plasmic connections similar to those in the hairs. The intercellular 
walls of the internal fibers are often oblique, but are always transverse 
in the enlarged portions. Towards the center of the axis chlorophyll 
bodies become rarer, and in some fibers entirely disappear. Se 

We found, also, in the outer portion of the axis certain longitudinal 
fibers, which show no chlorophyll bodies, and whose protoplasmic oon 
tents seem to be homogenous. Cell partitions in these ar 
lacking or at considerable distances apart.’ 
sionally, and are connected in a few cases to basal cells of hair clusters. 
Others are united to the oblique or transverse fibers. In sections 
which had been treated on the slide with acid alcohol to remove the. 
gelatinous sheaths, and first stained for two or three hours with a 
orseillin, then with methyl green or echtgriin for one — 
fibers were differentiated from the others, the cell contents Se 


We are unable to assign any particular function ; 
than that they a eer of the assimilative mee em the 
Plant. The hairs are both vegetative and reproductive He - : 

According to Schmidle, Thorea has three distinct gee . 
Stages of growth. The first, or prothallium stage, consists : vie 
less branched cellular fibers which develop directly from the sp ens 
We found plants in this stage, but found.neither spores nor tetrasp 


428 BOTANICAL GAZETTE [ DECEMBER 


developing from them. Schmidle found what he thought might be 
tetraspores, but questions their existence. The second, or Chantransia 
stage, develops directly from the former. The plants assume a Chan- 
transia-like form, growing up in little dark green or brown tufts on 
the surface of stones, etc., at the bottom of running streams. In this stage 
Thorea bears asexual spores in abundance. The third and most highly 
developed form of Thorea is the branching plant body described above. 
It develops from a union and growth of a number of Chantransia-like 
plants into a branching thallus of greater size, yet possessing all the 
forms of structure found in the preceding stages, with the addition of 
carpogones. 

The position of Thorea was long in doubt. Moebius in 1891-2 
placed it among the Floridez.’ Schmitz in 1892 placed it among the 
Phaeophycez,’ but changed his mind in 1894° and left it between this 
group and the Floridez, giving preference to the latter. Schmidle, 
who has devoted more time to the study of Thorea than any other 
botanist, is certain that it properly belongs to the Floridee for the 
following reasons: (1) in containing phycoerythrin, like the red sea- 
weeds; (2) in having cystocarps resembling those of Batrachosper- 
mum; (3) in that the hair cells contain intercellular protoplasmic 
connections typical of many of the lower Floridez ; (4) in developing 
from Chantransia-like forms in much the same manner as Batracho- 
spermum. 

The Nebraska specimens of Thorea agree in points of general 
structure with the published. descriptions of Zhorea ramosissima Bory, 
with a few exceptions. The plants have a decided olive-green color 
which persists in the herbarium specimens, rather than the purple 
tinge of the dried specimens from Worms and Paris. Our Thorea 
branches very sparingly, the longer branches often attaining a length 
of 3°" without side branches. On the contrary, the specimens from 
Worms and Paris are much branched, the diameter of the zone of hairs 
also being two or three times greater than that of the central portion 
while in ours the zone of hairs has nearly the same diaineter as the 
central portion. There is a marked difference in the hairs as found in 
our material and that obtained by Professor Saunders in Texas. In 
the former both the long and short hairs are of nearly equal diameter 

*Mogstus: Ber. d. deut. bot. Gesell. 10: 333-344. 1891; 11: 266-270. 1892. 

7SCHMITZ: Ber, d. deut. bot. Gesell. rr tII5-141. 1892. 

*ScuMitz: Nuova Notarisia 5°705-720. 1894. 


mn See eee 


_ BOTANICAL GAZET1E, XXVIII ay ae eer 


We 
Oe 


HEDGCOCK and HUNTER on THOREA 


1899] BRIEFER ARTICLES 429 


throughout their length, while in the latter a great many of the longer 
hairs as well as many of the shorter ones taper almost to a point. 

The description and measurements of structural parts are as 
follows : 

ZONE OF HAIRS (300—500y in diameter). 

Hairs: (a, pi. XXVTJ) long hairs consisting of 18-20 (more or less) cells 
outside the gelatinous sheath, counting from the basal cells; diameter of 
outer cells 5-8u, of inner 5-8; length of middle and outer (apical) cells 
18-40, of basal 13-20n; shape of basal cells elliptical to oval, of outer mostly 
rectangular, of apical usually rounded at tip. 

(6) Short hairs consisting of 1-6 cells outside the basal cells; diameter 
of basal cells 6—8y, of apical 4-6u (rarely wider than the basal); length of 
cells 13-164; shape of basal cells round to elliptical or oval; of the others 
rounded to rectangular (apical often pyriform). ’ 

Sfores-(c), single or in clusters, 10-15 in diameter, 15-264 long, spherical 
when young, pyriform when mature. 

INTERNAL FILAMENTS. 

Basal cells (d) 8-18 in diameter (broader in a few cases), 10-40m in 
length (rarely larger), oblong to ovoid, often very irregular. 

Cells of longitudinal fibers (e) 3-15u in diameter, very variable and often 
indefinite in length, irregularly cylindrical and often tapering. 

Cells of transverse fibers (f) 4-12y in diameter, much extended and 
variable in length, irregularly cylindrical. 

___ GEORGE G. Hepecock and ABEL A. HunTER, University of Nebraska. 
; EXPLANATION OF PLATE XXVI.—The plate represents a longitudinal 
radial section of a branch a short distance from the base, reaching from the 
outer edge of hairs to the middle of the axis. X 450. @, long hair; 4, short 
hair; ¢, spore formed from a single end cell; d, basal cell; ¢, longitudinal 
fiber of central portion ; f transverse fiber of central portion. 


NOTE ON CORN SMUT. 


_ A FEW years ago the per cent. of smut on corn in the vicinity of 
Manhattan was investigated with considerable care." 


cent. of smutted stalks. This year the amount of smut was <— 
SNe field giving go smutted stalks in 840, or 10.7 per cent. Another, 


"A. S, Hircncock and J. B.S. NorTOoN, Bulletin 62, Kansas Experiment Station, 
December 1896, E 


430 BOTANICAL GAZETTE | DECEMBER 


250 smutted stalks in 1250, or 20 per cent. These observations were 
made in order to compare the results with those obtained from the 
third field, where 140 stalks showed 38 to be smutted, or 27 per cent. 
This field was an experimental plat in which a number of crossed 
varieties were being self-pollinated. The tassels to be used for this 
purpose were enclosed in sacks, but the remainder were pulled out when 
young. At the time the pollinating was begun, several ears were 
beginning to silk. These were cut off with a corn knife. In some 
cases the entire ear was cut off; in others it was cut above the base. 
After a period the ears were allowed to grow as they appeared. In 
this last plat 117 stalks had ears upon them of which 10 had been 
cut. Of the ro cut ears 9 were smutted, or go per cent.; of the 107 
uncut ears 5 were smutted, or 4.7 per cent. The cut ears were growing 
at the time of mutilation. These observations serve to show that corn 
smut is greatly increased by mutilation which exposes the growing 
issue.— A. S. Hircucock, Manhattan, Kan. 


A BOTANICAL ART GALLERY 


Durinc the past season the University of Minnesota has taken steps 
to found a photographic exhibit of the vegetation of the state, and 
several hundred dollars have been expended for experimental work. 
The results are so far gratifying that the writer feels justified in giving 
the outlines of the plan for the benefit of other institutions that may 
care to develop similar exhibits. 

Considerable time was spent during the summer in securing nega- 
tives of vegetation. A photographer has been continuously employed, 
and about 300 8 X 10 negatives have been obtained. For the present 
the efforts have been limited to (a) plant portraits in their habitats and 
(2) ecologic groups. Many of these have been enlarged and framed. 
A commodious and well-lighted room has been chosen for the hang- 
ing, and at present twenty enlargements, 30 X 40, and several of smaller 
size, have been hung as the nucleus of the gallery. The pictures are 
numbered and framed in the ordinary manner, and promise to have 


much educational value, not only to undergraduates, but to the public . 


generally. 

I find that a picture 30 X 4ocan be produced, properly framed and 
hung, at a minimum expense of about $17. Higher prices are, how- 
ever, demanded for the best work in framing. It is important to 


PT ee ee Oe TT 


1899] . BRIEFER ARTICLES 431 


command the services of very expert photographers and skillful enlargers 
to obtain proper results. A good negative can be produced at an aver- 
age price of $2.50. Enlargement costs about $6, and the frame cannot 
be secured for less than $8.50. This price is trifling when the beauty 
and value of the whole exhibit is taken into account. 

Among the subjects of study that I have used might be named por- 
traits of Verbascum, Euphorbia, Helianthus, Solidago, Laciniaria, Cas- 
talia, Pteris, Quercus, Cuscuta, Pyrola, and groups of shore-lines, 
shade-plants, mat-plants, wand-plants, forest-floor coverings, swamp 
vegetation, etc. Such an exhibit when fairly extended would give a 
very adequate and pleasing idea of the vegetation in the region that 
has been selected for analysis. Conway MacMILLan, Zhe University 
of Minnesota. 


A NEW LILIUM. 


Lilium Masseyi, n. sp.—Bulb 12” in diameter or less, composed of 
fleshy scales: stem 1.5 to 3 high, with two distant scales below: 
leaves linear, acute at both ends or the lower obtuse, 12 to 25" long, 2 
to 4™" wide, in whorls of 3 to 8, the central ones generally alternate, 
glaucous, the margins revolute, prominently three-veined: flowers 1 to 
3, erect, 2.5 to 5" high; perianth reddish-orange, its segments spatu- 
late, obtuse, slightly pubescent, the blade 6 to 12™ wide, gradually 
narrowed into the claw, purple spotted below: capsule obovoid, 12 to 
250" high 

High mountain meadows of North Carolina, July-August. Named 
in honor. of Professor W. F. Massey, Horticulturist N. C. Agr. Exp. 
pion. —C. W. Hyams, Agric. Exper. Station, West Raleigh, N. C. 


GURRENT LITERATURE. 
BOOK REVIEW S. 
Flower ecology. 


AS PREVIOUSLY stated in the pages of this journal (26: 358), the first volume 
of Knuth’s Handbuch der Blitenbiologie’’* gives a general introduction to the 
subject of the pollination of flowers, with the bibliography. 

The second volume gives an account of the investigations which have 
been made upon the European and Arctic floras. The work being profes- 
sedly based upon Hermann Miiller’s “ Befruchtung der Blumen durch Insek- 
ten,’’ even more closely resembles the English translation by Thompson, 
“The Fertilization of Flowers.” The resemblance here extends to the intro- 
duction of the general results of other authors and in the arrangement of the 
families, part 1 including the Ranunculacee to Compositae, and part 2, 
Lobeliaceae to Gnetacee. As an example of the treatment of the particular 
cases we may take that of Malva silvestris. There is a citation of authors, an 
account of the mode of pollination, accompanied by a reproduction of 
Miiller’s figures, and a combined list of visitors observed by Miller and 
the author. Then follows the visitors observed by Alfken, Schletterer, 
Schenck, Loew, Scott-Elliot, and Smith. The lists are often quite fragmen- 
tary, observed in different places, and under varying conditions, so that they 
will hardly form a homogeneous combination, but it is a good thing to have 
them collected together. The fact that the most scattered and inaccessible 
writings are usually the most worthless does not keep anyone from rejoicing 
that he does not have to waste his time hunting forthem. Knuth draws upon 
the writings of pure entomologists. Their writings are very important for 
the insects they especially collect, but, as a rule, they do not notice all of the 
visitors, Their lists are far better than those made out by persons who make 
you wonder if they can tell a hive-bee or bumblebee from an Eristalis. 

The first part contains 697 pages, 210 figures, and asa frontispiece a 
portrait of Herman Miiller. Fora frontispiece the second part contains 2 
portrait of Darwin, surrounded by Severin Axell, Federico Delpino, F. Hilde- 
brand, and Fritz Miller. This part contains another 210 figures. The 


*KnuTH, PauL: Handbuch der Sree hntaiy unter caaeitsanen gaits? 
Ba 


Hermanns Miiller’s Werk “ Die Befruchtun ng der en durch Insekten.’ 
Die bisher in Europa und im arktischen Gebiet Sane ee Beob- 
achtungen. Teil 2: Lobeliacee bis Gnetacez. 8vo. pp- iv-+705. pl. 7. figs. 210 


Leipzig : Wilhelm Engelmann, 1899. J/ 18. 


432 [DECEMBER 


1899] CURRENT LITERATURE 433 


volume closes with a list of the flower-visiting insects, each species being 
followed by a list of the plants on whose flowers it has been taken. 

Facts appear here which are quite interesting in connection with state- 
ments made on page 114 of the first volume, as well as in connection with 
some recent writings of mine on oligotropic bees (BOT. GAZ. 28:32). Andrena 
florea, mentioned on the page cited as an exclusive visitor of Bryonia dioica, 
appears in 2:606 as a visitor of Bryonta alba, also, as well as Sisymbrium 
officinale, Stellaria media, Rubus Sruticosus, Cirsium arvense, and Carduus 
nutans. Andrena cettii (=A. marginata F.), mentioned in 1:114 as an 
exclusive visitor of Scadiosa (Knautia) arvenszs, in 2:607 is shown to visit 5S. 
columbaria and S. suaveolens, also, as well as Succtsa pratensis, Onopordon 
acanthium, Leontodon autumnalis, Flieractum pilosella, Jastone montana, 
Andrena nasuta visits Melilotus alba also, and Bombus gerstaeckeri visits 
three species of Aconitum and Gentiana asclepiadea. So all of the exclu- 
sive cases are shown to be merely erroneous inferences from improbable data. 
These lists were evidently made later, and so are not connected with the first 
Statements. 

In this work the classification, although systematic within each volume, is 
geographical. The second volume contains observations made in Europe, 
excluding all others. If a native European plant has been introduced, for 
example, to America, any observations made upon the plant in America 
should be included with the observations made in Europe. This permits a 
convenient comparison of its normal relations at home with its acquired rela- 
tions abroad. The latter are clearly subordinate and cannot take the place 
of the other. 

If we enter upon close criticism of the present status of flower ecology, 
it must be said that it is a heterogeneous mixture of data derived from native 
plants occupying more or less normal positions in the original flora and 
€xposed to a more or less normal insect fauna, as well as data derived from 
_ introduced and cultivated plants which seem erroneously to be expected to 
reveal their ecological relations without regard to their surroundings. It 
Seems to me that introduced plants should be treated together in a supple- 
Mentary part, while garden flowers should be again separated as surrounded 
by conditions clearly abnormal. The objection is to the mixture, not to any 
_ kind of data. 


_ the nomenclature which is connected with it, whether they like it or not. 

: This work is by far the most important source of information on the facts 
and literature of flower ecology, and it should be in the hands of all those 

who are interested in the subject. I have done enough work with bibliography 


y 
~ AX 


434 BOTANICAL GAZETTE [DECEMBER 


to observe that too large a part of it does not show anything except that the 
writers were ignorant of the literature. Even their interest is shown to be 
fraudulent. They expect us to take an interest in their writings, when they 
do not care enough for the subject to read what has been written by others. 
CHARLES ROBERTSON. 
Propagation of mosses. 

Dr. CARL CORRENS has been at work for five years upon the vegetative 
reproduction of the true mosses, In that time he has published several pre- 
liminary papers on his researches,’ and an important collateral paper ‘‘ Ueber 
Scheitelwachsthum, Blattstellung, und Astanlagen des Laubmoosstammchen ” 
in the Festschrift fiir Schwendener (1899). These investigations are now 
brought together zz extenso to form a bulky volume of almost 500 pages. 

e gametophyte of mosses propagates itself by many methods and by 
brood-bodies which arise from protonema, or rhizoid, or stem, or leaf. The 
sporophyte produces normally no propagative organs except the spores. The 
homology of the brood-bodies of Georgia (with which the researches of Correns 
began in 1894), @Edipodium, Drepanophyllum, anda few others (all of possible 
paraphysine nature) remain undetermined. The term brood-organsis restricted 
to those structures of which propagation is the chief or the codrdinate function. 
The term cuttings is applied to those parts artificially separated by which prop- 
agation is possible. The brood-organs of one hundred and twenty-five 
species are monographically described. Forty species which can be repro- 
duced from cuttings are also elaborately treated. 

The terms applied to the brood-bodies have such resemblance in form and 
sound in German that they must prove objectionable, particularly in speaking. 
Bruchstémmcehen or Bruchdste are stems or branches which are brittle through- 
out and may break off anywhere, or even break up into pieces, each capable 
of growth. Arutdste are brittle at base only where a separation layer is formed. 
Bruchknospen are branches brittle only beneath the apex, while Brutknospen 
are short Brutdste. In such a case perhaps it is unimportant whether one 
perceives the distinction between Bruch and Brut/ In the same way we 
have Bruchblitter and Brutblatter. Bulbillen are defined as Bruthnospen 
with reduced leaves, but the organs described as bulbils in Leptobryum 
pyriforme bear not the ae trace of leaf rudiments; indeed they are else- 
where called “ root tubercles 

In the general part (pp. 325-460) Dr. Correns discusses the morphology 
and phylogeny of the brood-bodies arising on stem, leaves, and protonema. 

*Berichte d. deutsch. bot. Gesells. 13: 420. 1895; 14:94. 1896; 15:373- 18975 
16: 22. ded 

3CORRENS, CARL: Untersuchungen iiber die Vermehrung der Laubmoose durch 
Sieeloeyies et Stecklinge. 8vo. pp. xxiv-+472. figs. 185. Jena: Gustav Fischer. 
1899. M 15. 


1899] CURRENT LITERATURE 435 


He rejects Schimper’s view that the cluster of gemma in Georgia and 

dipodium is homologous with a male “flower,” the gemmez being sterilized 
antheridia ; and also dismisses as improbable Brefeld’s suggestion that they 
are sterile sporangia, like chlamydospores. 

The structure and development of the brood-bodies, their separation, dis- 
tribution, mode and conditions of germination, and the conditions for their 
formation are described. Finally the author furnishes a key to the various 
kinds of brood organs and the species in which they occur. 

A list of the literature, which consists mainly of the {systematic works 
referred to, the special literature being very scanty, and an inadequate index 
complete the work. 

The interest and value of the book lie in the exhaustive treatment of a 
subject, presumably narrow, which has shown itself broad when thoroughly 
studied. It would be interesting to have a similar study of the vegetative 
reproduction among the Hepatice, and we trust Dr. Correns wil] include in 
this thorough investigation both classes of the Bryophyta.—C. R. B. 


NOTES FOR STUDENTS 
V6CHTING, the author of the well-known work on Transplantation and of 
various papers having to do with the correlations of organs and tissues, has 
published the results of some extended investigations on tuberous plants.4 It 


other organs to take their place. In the above cases the replacing organ 1s 
€ssentially the same in kind as that replaced, but 
almost any organ, if properly stimulated, may become a tu 
tuber as a fleshy body used for storage, whether morpholo 


or leaf —a definition that the following results obviously require. 
ange themselves Into two 


uch a position that its nor- 


ber. He defnes a 
gically stem, root, 


The experiments upon vicarious organs arr 
§roups: a tuber ma hen put ins 
; y replace a stem w P ee 
mal function cannot be performed, or a tuber may be developed sa meee 
any organ if the normal formation of tubers is suppressed. ny am 
Plastic plants studied was Oxalis crassicaulis, a plant which no y 


*Prings. Jahrb. fiir wiss. Bot. 34: 1-148. 1899- 


436 BOTANICAL GAZETTE [DECEMBER 


resembles the potato in the development of subterranean stem tubers. A tuber 
placed in an erect position, partially below the soil line and partially above it, 
develops roots and rhizomes from the buried portions and green shoots from 
the aerial portions. Instead of decaying rapidly, as a tuber commonly does 
when it has disposed of its stock of reserve food materials, the Oxalis tuber, 
placed in this abnormal position, lives and grows through the entire season. 
The tuber is obliged to function as a stem, bothin the conduction of water and 
the plastic foodstuffs and in giving mechanical support to the aerial organs. 
As a result of these new functions, the author finds a striking change in the 
anatomy of the tuber. Instead of the predominance of parenchymatous 
storage cells, there is a great increase in the area of the vascular bundles; 
the secondary phloem and xylem develop remarkably, and the new cells and 
cell fusions have a much larger cross section and more complicated structures. 
The strong development of bast fibers and wood cells materially adds to the 
mechanical strength of the tuber in its new relations. Thus the tuber has 
become like a normal stem in structure as in function. 

The suppression of normal tuber formation in Oxalis stimulates their 
development in a new position. The plant sometimes develops stolons 
whose ends bury in the soil and develop tubers from the terminal bud. If this 
bud is removed, the bud nearest the end becomes a tuber, and if all the buds 
are removed one of the stolon internodes swells up like a tuber. In some 
cases the leaves instead of the internodes become tubers, and the leaflets 
may remain as rudiments or may swell up like the petiole. The structure of 
the petiole which functions as a tuber undergoes a remarkable change. The 
ventral furrow is absent, the petiole being round in cross section. There is 
no collenchyma, or green tissue, and the bundles which are so prominent in a 
normal petiole remain in a rudimentary condition; even the vessels which 
are present may be filled up with tyloses. The changed petiole is almost 


assume the most fantastic shapes ; indeed Vichting commonly finds that the 
more abnormal the organ which becomes a tuber, the more abnormal are the 
starch grains, indicating an intimate correlation of structures of a surprising 
nature. 

Experiments similar to those just mentioned were performed on the 
potato but were less successful as a rule, showing, the author thinks, that 
Solanum is less plastic and that the metamorphosis of stem to tuber has gone 
further than in Oxalis. A root of Dahlia variabilis was planted somewhat 
emerging from the soil. The new roots, which commonly form in a cluster 
at the base of the stem, formed at the base of the parent root, since the base 
of the stem was inthe light. The next year the compound root system was 
planted with these new roots emerging from the soil, and new roots formed at 
the base of this second story of roots. This process was repeated until 


tuber formation in Oxalis, but the internal forces are 50 


1899 | CURRENT LITERATURE 437 


finally four tiers of roots were developed. Each year the plant started later 
and later, and for a long time the leaves remained yellow, showing the 
difficulty the plants had in conducting materials up through the series of root 
tubers. The fifth year the struggle was too severe, and the plant died. 
These tubers did not develop buds and were hence incapable of propagating 
the species, but experiments showed that decapitated budless tubers can 
remain alive and fresh for several years. The Dahlia tubers developed a 
stem structure like those of Oxalis. 

One of the most plastic plants employed was Boussingaultia baselloides, a 
plant with the potato type of tuber. This plant grows readily from cuttings, 
roots and rhizomes springing from the subterranean buds, and green shoots 
from the aerial buds. When cuttings are placed in the soil so that all buds 
are in the light, the base of the stem itself, z. e., the buried internode, swells up 
intoa tuber; if the base of the internode lies deep, the tuber is elongated, if 
shallow, it is shortened, showing the restricting influences of light in tuber 
formation. As in Dahlia, these tubers are budless and remain fresh and living 
fora year or two, but cannot propagate the species. The change in struc- 
_ ture is similar to that in the Oxalis petiole which becomes a tuber. A leaf of 

Boussingaultia when placed in the soil gives off roots, one of which swells 
into a budless tuber; these tubers live and function if a shoot is grafted upon 
them. (Leaves of Gloxinia root similarly but develop buds and are capable 
of growth.) The experiments on this plant and on Oxalis seem to show a 
strong inclination toward tuber formation, an inclination which must be sat- 
isfied in one way if notin another. In 7, ‘Aladiantha dubia, a tuberous mem- 
ber of the melon family, the author succeeded in setting out a tendril and 
developing a starchy tuber from its base, which remained fresh long after the 
tendril died. ; 
Véchting carried on several experiments for the purpose of showing the 
influence of external factors, especially light and darkness, upon tuber forma- 
tion. The restricting influence of light in the case of Dahlia and Boussin- 
Saultia has already been indicated. In the radish the tuber is partly root but 
mostly hypocotyl. Plants with all the hypocotyl and part of the bra oes 
the soil tended to develop elongated tubes, to which the root contribute . 
ormore ; in some cases all of the tuber was developed from the root. Etio- 
lated seedlings were darkened at various points by tinfoil, and tuberous 
swellings appeared within the darkened areas, but always ag near oi we ian 
Possible. Two or three separate tubers were sometimes developed in 


: wn to facilitate 
way on a sin mperature and drought were sho 
y ingle plant. Low temp strong that they are 


r hand, will warmth and 
In this species also light 
The 


hot effective early in the season, nor, on the othe 
Moisture greatly retard tuber formation in the fall. econ 
'S unable to prevent finally tuber formation, though it greatly © 


438 BOTANICAL GAZETTE [DECEMBER 


strong tendency to tuber formation which finally overcomes all obstacles is 
due, the author thinks, not only to the internal force which looks to the propa- 
gation of the species, but also to a demand for organic symmetry ; the occa- 
sional development of potato tubers without starch, and therefore functionally 
impotent, appears to favor this conclusion. 

The author has thus been enabled to establish upon a firmer basis than 
ever before his ideas as to the great plasticity of plants and the vicarious 
nature of their organs. Perhaps the most astounding thing of all is the 
power shown by a mature organ, like the tuber of Oxalis or Dahlia, to be born 
again, as it were, and start on a period of. secondary growth. The plasticity 
of a young organ is well known, and perhaps not so surprising, but one would 
scarcely have expected to see such evidences of life and vigor in a specialized 
organ like a tuber— HENRY C. COWLES. 

THE ORIGIN of the cilia of the spermatozoid is very briefly but clearly 
traced by Belajeff5 in Gymnogramme sulphurea and Eguisetum arvense. In 
Gymnogramme two centrosomes (the blepharoplasts of Webber and others) 
make their appearance at opposite poles of the nucleus of the grandmother 
cell of the spermatozoid. The division of this nucleus is not accompanied 
by a division of the centrosome, and consequently each of the resulting cells 
receives only a single centrosome. The centrosome, originally spherical, elon- 
gates into a narrow band lying alongside the nucleus, and the cilia arise from 
the peripheral portion of the band. 

The sequence is the same in Equisetum, but here the writer was able to 
show that the band is made up of a row of intensely staining granules and a 
less deeply staining portion. Each granule gives rise to a single cilium. 

he spherical organs which give rise to the band are regarded as genuine 
centrosomes, and Belajeff would homologize with them the blepharoplasts of 
Webber (Zamia) and Shaw (Marsilea and Onoclea), and also with the cilia- 
forming centrosomes of Hirase (Gingko) and Ikeno (Cycas). He would also 
homologize the cilia-forming band with the “middle piece” of the animal 
spermatozoon, as described by Hermann for the salamander. — CHARLES J. 
CHAMBERLAIN, 

IN A sTuDy of the influence of weather and the condition of the soil upon 
the anatomical structure of plants, W. Meyer® objects to culture experiments 
and goes for his material to nature, where plants may be found under the 
same conditions for many generations. He compares numerous members of 
the Caryophyllacez, chiefly alpine forms, and shows how species in different 
divisions of the same family have a close resemblance to one another when 
growing in similar situations. For example, species of the Silenez, Alsine, 

5 Ueber die Cilienbildner in den spermatogenen Zellen. Ber. d. deutsch. bot. 
Gesell 16:140-144. A/. 7. 1898 

° Bot. Centralb. 79: 337-350. 1899. 


1899] CURRENT LITERATURE 439 


and Paronychiee growing in deserts resemble each other and also those on 
alpine heights, for in high altitudes the sun’s rays are very powerful and 
plants need the same protection as in deserts. On the other hand, specimens 
of the same species, under various conditions, show extreme divergence. 
He also shows that many species of the Primulacew resemble those of the 
Caryophyllaceze when grown under like conditions. Only a causal depend- 
ence between situation and structure can explain such resemblance, since 
common origin cannot do it.— L. M. SNow 

SEEDS whose viability had been previously tested by samples were recently 
submitted by Professor Dewar to the intense cold of liquid hydrogen, z. ¢., 
— 250°C. for half an hour. Some of the seeds were cooled in a sealed glass 
tube, and others were immersed without protection in the liquid hydrogen. 
All the seeds in both sets germinated. —C. R. B. 

THE LITERATURE of diatoms has recently been enriched by a very 
important contribution.?. The work is not merely a guide for the determina- 
tion of the species of a limited locality, but is a comprehensive text-book of 
diatom lore. The author has departed from the usual comparatively super- 
ficial methods, and has taken into account the form and structure of the pro- 
toplast, the position of the nucleus, the number, form, and position of the 
chromatophores, the occurrence of pyrenoids, and, finally—a most important 
consideration—the complete life history of each species as far as this has 
been possible. A study of cell characters convinced the author that the 
number and position of chromatophores is the most important taxonomic 
character, and that mere frustule characters are not sufficient for determining 
the limits of species. 

The second part of the work gives a somewhat extended account of the 
diatom cell, cell division, movements of diatoms, the relation of variety of 
form to environmental factors, the auxospores, and the role of diatoms in the 
economy of nature. CHARLES J. CHAMBERLAIN, 

THE PRODUCTION of apospory by environment has been brought about 
in various ferns. Mr. F.W. Stansfield® has succeeded in producing apospory 
in Athyrium filix-femina, unco-glomeratum, an apparently barren form. In 
all cases it was noted that prothalli are produced with much more ease from 
young fronds than from adult ones. If the first fronds from a prothallus are 
Pinned down, the edges rapidly develop into prothalli. The aposporous eu 
duction of prothalli is regarded as: an atavistic trait, and the nae eter . 
made that apospory could be produced in many ferns by demise em 
Bucht. Wissenschaftliche 


STEN, GEORGE; Die Diatomeen von Kieler S 


7KarR 
Meeresuntersuchungen herausgegeben von der Commission zur Untersuchung 
deutschen Meere in Kiel und der biologischen Anstalt auf Helgoland, Abtheilung 
Kiel. Neue Folge 4: 19-295. figs. 219. 1899. 

*Jour. Linn. Soc, Bot. 34: 262-268. 1899. 


440 BOTANICAL GAZETTE [DECEMBER 


care. The fact that Mr. Druery, a few years ago, succeeded in producing 
apospory in Scolopendrium vulgare, presumably a most unlikely form for 
such an experiment on account of the smooth stri.p-shaped leaves, indicates 
that the suggestion has some weight.—-CHARLES J. CHAMBERLAIN. 


APPLE CANKER, which attacks the bark of the limbs of apple trees of all 
ages, has been traced by Mr. W. Paddock,’ of Geneva, N. Y., to the well-known 
Spheropsis malorum Pk., causing the black rot of apples. Cultures have 
been made on sterilized bean stems, and the disease produced by inoculation. 
Ina later communication” further observations are given upon the destruc- 
tiveness of the disease, which occurs, as it is discovered, in pears and quinces 
as well asin apples. Trees may be entirely killed by this disease, which in 
most cases progresses from the smaller branches toward the trunk.—J. C. A. 


WEEDS have been the subject of a number of bulletins from the agricul- 
tural experiment stations, not yet mentioned in these pages. Only the western 
states are represented. F. H. Hillman (Nev. no. 38: 1-131. 127 cuts in 
text) describes the seeds of many weeds with much clearness and detail, and 
presents one hundred and twenty five cuts, drawn by himself, illustrating as 
many kinds of seeds. These illustrations are worthy of special commenda- 
tion for their accuracy and artistic merit,and also because they are well 
printed. L. F. Henderson (Idaho no. 14: 91-136. 13 pl. and 5 cuts in text) 
discusses twelve of the state’s worst weeds, and says good things about 
the value and justice of weed laws. E. E. Bogue (Oklahoma no, 41: I-I2. 
14 cuts in text) presents information regarding seventeen weeds, of which 
those least known eastward are Solanum Ti orreyt, Acacia filiculoides and 
Croton Texensis. A. S. Hitchcock and G. L. Clothier have issued a press 
bulletin (no. 18) of two pages giving notes on weeds, and also a sixth report 
on Kansas weeds (Kans, no. 80: 113-164). A large fund of information is 
presented regarding the habits and distribution of weeds, not only of Kansas, 
but of the whole United States. Charts are used to show the distribution by 
counties in Kansas of 209 species, and by states in the whole country of 194 
species. L.H. Pammel presents a full account (Iowa no. 38: 7-24. 7 cuts 
in text) of the Russian thistle, with a bibliography ; also a discussion of the 
weeds of cornfields (Iowa no. 39: 27-52. 17 cuts in text), and of horse nettle 
{Solanum Carolinense), Convolvulus arvensis and Tribulus terrestris (lowa, 
no. 42: 130-140. 5 cuts in text), the last species having recently gained a 
foothold on Muscatine island in the Mississippi river. E. S. Goff (Wis. no. 76 : 
1-53. 39 Cuts in text) gives illustrations and information regarding the ten 
weeds mentioned in the Wisconsin weed law, with notes on eight others.— J. 
(A. 


9 Science 8: 595. 
© Science 8: 836. 


) 
| 
. 
| 
F 


1899 | CURRENT LITERATURE 441 


THE RUST FLORA of California, according to E. W. D. Holway in the 
October Evythea, embraces 122 species of Puccinia, 42 of Uromyces, and 73 
of other genera.—J. C. A. 


ANOTHER ARTICLE™ has recently been added to the valuable series of 
physiological papers already so auspiciously inaugurated by Dr. Klebs. The 
same ingenious accurate experimentation which characterized the earlier 
papers of the series is evident. The presentation is masterly. The purpose of 
the research is to determine the chemical and physical factors which incite or 
alter the various modes of reproduction in Saprolegnia mixta. It is found 
that this species will grow indefinitely without either sexual or asexual repro- 
duction if nourishment be abundant; but at any time the extensive formation 
of zoospores can be incited by simply starving the hyphe, e. £., by placing 
them in water. By noting the maximum concentration at which various foods 
induce the formation of zoospores an idea was obtained as to their relative 
food value. Albumens are rich; amido-acids can furnish C as wellas N ; in 
general the food value rises with the carbon content; glucosides vary from 
toxic to indifferent or even favorable; inorganic acids ani their salts are of 
but little value. 

By varying the nutritive value of any medium the fungus can be made at 
will to assume a purely vegetative condition; to produce rudimentary spo- 
fangia ; to form sporangia which bear zoospores that do not escape 3 and to 
produce functional zoospores. All of these phenomena depend for their exist- 
€nce upon the concentration of the medium, not upon the total quantity of 
nutriment. 
strong solutions the formation of zoo- 


It happens, however, that even in 
led Dr. Klebs to infer the pres- 


Spores is often eventually suppressed. This 
ence of an inhibiting agent formed in the medium by the growth of the fungus. 
One such substance he finds is ammonium carbonate. If the mediuts be 
rendered weakly acid zoospore formation can be resumed. Starvation, if very 
§tadual, causes the mycelium to become too weak to build zoospores. Poisons 
inhibit their formation as does also. high osmotic pressure. 
very clearly what are the necessary relations and also the responses, but the 
Teasons for both are totally obscure. Zoospores are never found unless oe 
tips of the hyphz are in contact with water. Oxygen, light, and heat are 0 
little importance. 

If a well nourished mycelium be placed in a poor m 
ditions render the formation of zoospores impossible, ¢. om 
Sexual organs will soon appear in abundance. These, however, are eee . 
to heat (their maximum being 26°, that of sporangia 32°) and fastidious a 

“Zur Physiologie der Fortpflanzung einiger Pilze: Jahr. 
1899. Reviews of earlier papers may be found in this journal 23 : 21 
77. 1899. 


edium where the con- 
ina solid medium, 


f, wiss. Bot. 33: 71- 
4. 1893, and 27: 


442 BOTANICAL GAZETTE [DECEMBER 


their inorganic food, seeming quite dependent upon the presence of some 
form of potassium phosphate. This is particularly true of antheridia, and by 
varying the medium a filament may be obtained which bears no sex organs, 
or one bearing only oogonia, or one with oogonia and a few antheridia, or, 
finally, one with many antheridia some of which form fertilizing tubes. In 
this connection it should be recalled that specific distinctions have been based 
on the abundance of antheridia. In general the relation between oogonia 
and antheridia is such that support is given to the view of DeBary, viz., that 
the presence of oogonia induces the formation of antheridia. Dr. Klebs 
thinks this is due either to chemotaxis when proper inorganic salts are pres- 
ent, or that these salts render the twigs sensitive to the chemical stimulus 
emanating from the oogonia. It is evident, however, that normal oogonia 
can exist without inducing antheridial formation. 

While as conclusively proved in this research, there is no dominating 
inherent tendency toward an alternation of generations, nevertheless the con- 
ditions are such that in nature an alternation is usually brought about through 
the exhaustion of the nutriment afforded by each newly attacked host. 

Previous to oosphere formation the incipient oogonium may revert toa 
vegetative condition, but after the oospheres are differentiated the power to 
vegetate is irretrievably lost. This, the author thinks, is due to nuclear 
changes possibly to a chromosome reduction. 

An interesting chapter is given to the consideration of gemmee and the 
author concludes, apparently with ample ground, that they are of no signifi- 
cance in phylogeny. They are special structures whose function is to tide 
over times when the formation of other spores is precluded. They behave in 
general as do hyphae, and develop into oogonia or sporangia according to 
environment. Dr. Klebs closes by saying that an acquaintance with mere 
morphological marks does not constitute sufficient knowledge of a species. 
To meet his high ideal the systematist must hereafter determine, both quan- 
titatively and qualitatively, the life relations of the plant, its limits of varia- 
tion, and the stimuli that cause these variations.— F. L. STEVENS. 


WHETHER THE Saprolegniacez are exclusively apogamous or not is a 
question that has been argued fro and con in pre-cytological days by DeBary, 
Pringsheim, Cornu, Zopf, Ward, Humphrey, and others. Four years ago 
Messrs. Hartog and Trow almost simultaneously published papers expressing 
quite opposite views regarding fertilization in this group. Trow has recently 
made extended researches on Achlya® and arrives at conclusions in harmony 
with his earlier paper, He describes a karyokinetic division of the oogonial 
nuclei and a degeneration or digestion of the supernumerary ones, so that only 


* Trow, A. H.: Observations on the biology and cytology of a new variety of 
Achlya Americana. Ann. Bot. 193 1a1. 


te lg ee Dh Oa ies, Peel Ts), » 


1899] CURRENT LITERATURE 443 


one remains to function in each oosphere. Trowis convinced that true fertil- 
ization, a fusion of sexual nuclei, does occur. It is to be regretted, however, 
that his technique was not improved to such a point of efficiency as to insure 
more unequivocal evidence than he presents. The final impression that is 
left with the critical reader is that Trow has seen some things which make a 
fertilization seem possible, or even probable; but that it is far from being 
proved. A really valuable feature of Trow’s work consists in the observa- 
tions on live material, by which he has followed the growth of the organism 
from the zoospore to complete maturity, including oC ripening, and 
germination of the oospores. 

An article which bristles with caustic but mainly petty criticism regarding 
Trow’s conclusions and theories appears in the September Anmads of Botany. 
This criticism, like Hartog’s criticism of Trow’s earlier paper, while it 
increases the literature by several pages, sheds no light on the perplexing 
questions.— F, L. STEVENS. 


3HARTOG, MARCUS: The alleged fertilization in the Saprolegniacex. Ann. 
Bot. 13: 447 


NEWS. 


Mr. ALBERT H. Trow has been granted the degree D.Sc. by the Uni- 
versity of London. 


PROFESSOR ERNST EBERMAYER has resigned the professorship of forestry 
in the University of Munich. 


Mr. W. L. JEPSON has been promoted from the instructorship to be asso- 
ciate professor of botany in the University of California. 


Mr. R. K. Beatrigz has been appointed instructor in botany in the 
Agricultural College of Washington, located at Pullman. 


Dr. JoHN M. CouLTER will be away from his post at the University of 
Chicago for nine months. He is spending the winter at Washington, D.C. 


MR. GRANT ALLEN, author of several popular books and papers on botani- 
cal subjects, and more lately widely known as a writer of acknowledged 
fiction, died October 25, in his fifty-first year. 

Mr. H. H. Hume, assistant in the departments of botany and horticulture 
in the Iowa Agricultural College, has been elected horticulturist and botanist 
of the Florida Agricultural College at Lake City. 


THE EXTRAORDINARY delay in the publication of the October number 


was due to the illness of the present managing editor from October 13 to 
November 4, and the absence of the senior editor. 


Two SCHOLARsuIPs for garden pupils in the Missouri Botanical Garden 
- will be awarded by the director, Dr. Wm. Trelease, before April first next. 
Applications must be in his hands not later than March first. 


Mr. A. A. HELLER, of Lancaster, Pa., will start for Porto Rico on a col- 
lecting trip about January 1. He expects to bring back a large and interesting 
collection of all classes of plants. The cryptogams of that island should be 
an especially interesting study. 

PROFESSOR Dr. PauL KNurtH, whose recent return from a tour around 
the world, including a somewhat prolonged stay in this country, we lately 
chronicled, died in Kiel on October 30, at the age of forty-five. His death 
will be felt as a severe loss to ecology. ; 

Mr. J. W. Duvet, of the Agricuitural Experiment Station at Wooster, 
Ohio, has been appointed to the D, M. Ferry Botanical Fellowship in the 

444 [DECEMBER 


Sap ieiiaie bee eno Sh dine ace te ae wy ae 


1899 ] NEWS 445. 


University of Michigan. Mr. Duvel has begun an investigation of certain 
factors affecting the vitality and germination of seeds. 


PROFESSOR IGNATIUS URBAN announces that the second fascicle of his 
Porto Rico plants is now ready for distribution, the price per hundred being 
M 40. A few sets of the first fascicle may still be obtained, at J/ 30 per hun- 
dred. Professor Urban’s address is Grunewald-strasse 6/7 Berlin W. 


PROFESSOR JEAN BAPTISTE CARNOY, professor of botany in the Catholic 
University of Louvain, died on September 8, at the age of sixty-three. Pro- 
fessor Carnoy was the founder and editor of the elegant periodical La Cellule, 
and the author of several works and papers of high value on cytology. 


THERE HAS come into the possession of the city of Philadelphia the dwell- 
ing and part of the grounds which belonged to James Logan (1674-1751), 
with Penn one of the founders of Pennsylvania, and a botanist of note, after 
whom Logania, the type of the Loganiacez, was named by Robert Brown. 
The property will be known as Stenton Park, the original name of the Logan 
estate, as there is already a Logan Square in the city. Mr. Thomas Meehan, 
the venerable horticulturist, has been the mover in the proceedings which 
have secured the preservation of the grounds of Bartram, McMahon, and 

ogan, 


Mr. J. B, Exxis being unable to continue the issue of Fungi Columbiant, 
has requested Mr. C. L. Shear, of Takoma Park, D. C., to take charge of it. 
Mr. Shear has acceded to this request, and seeks the cooperation of working 
mycologists and collectors. Mr. Ellis will assist in the determination of 
Material, and doubtful specimens of the various orders will be submitted to- 
other specialists. The series of North American Fungi has been discon- 
tinued, but the present series is in great part a continuation of it. Century 

is now in course of preparation. Complete sets of the first fourteen 
centuries can still be supplied. 

ON JANUARY 1 the Cambridge Botanical Supply Company will discon- 
tinue the publication of the card index of American Botany, and the commit- 
tee appointed by Section G of the A. A. A. S. is seeking to continue It under 
other arrangements, The present subscription price of $5 per year was made 
when only about 500 cards were issued and 1s inadequate to support the 
€nterprise with the increased number of titles. It has, therefore, been decided 
to make the rate one cent percard. The number of subscribers will govern 
the number of sets printed and the matter will not be electrotyped. Intend- 
ing subscribers should at once notify Professor L. M. Underwood, Columbia 
University, New York city. 


MR. Maturin L. DELAFIELD, JR., 56 Liberty street, New aes or 
having fulfilled the requirement of the constitution of the Botanical Soceity 


446 BOTANICAL GAZETTE [DECEMBER 1899 


of America, becomes a patron of that organization. At the Columbus (0.) 
meeting last August, among other amendments to the constitution of the 
society, one was pe admitting patrons. The clause relating to patrons 
reads as follow: 

The payment to the treasurer of the sum not less than $250 at any one time, ora 
bequest of such sum, shall constitute the donor a patron of the society. The names of 
patrons shall be published with the annual lists of officers and members, and the 
patrons shall be entitled to receive copies of all the pean of the society. 
Patrons’ fees shall be added to the permanent fund of the s 

Mr. Delafield, therefore, becomes the first patron a the Botanical Society 
of America.— GEORGE F. ATKINSON, Secretary. 


THE MissourI BOTANICAL GARDEN has this summer received a decision 
from the Supreme Court of Missouri which empowers the trustees of the gar- 
den at their discretion to sell unproductive endowment real estate, which was 
made inalienable by the will of Henry Shaw, the founder of the garden. 
While it is not probable that sales will be made rapidly, the purpose being to 
effect them at full market value, the power to dispose of this property promises 
to add many thousand dollars to the annual revenue of the establishment 
within a few years, and it makes available for immediate use each year some 
ten or fifteen thousand dollars which business wisdom has compelled the 
trustees to hold out of the current income thus far, as an emergency fund for 
the protection of the property when street improvements and other special 
expenses should be forced upon it. 

Immediate use is to be made of some of the money thus liberated, by the 
addition to the garden of about twenty acres of ground, to be graded and 
planted in accordancewith plans made by the landscape architects, F. L. and 
J. C. Olmsted, and to represent in synopsis the principal features of the North 
American flora. 


GENERAL INDEX. 


The most important classified entries will be found under Contributors, Cytology, 
Diseases, fone Hosts, Personals, Physiology, and Reviews. New names, and 
names of new genera, species, and varieties, are printed in bold-face type; synonyms 


in a 
A 
A. A. A. S., Botanical ee 212; officers 
of club b 283: — G2 
Abrams, Ler 
Achillea 36 “i 
Aconitum reclinatum 


lan 
Dsospore pusilla a ey 
A i a 


272 
pels Sigg Cate eo viatie 120; 
129; 


n tenerum ort 
225; edicus 231; Portulacz 231 


te ad scagpieegdd of Tilopteridacez 
213; I 425 

Allen, Eeant’ ae of 444 

Alternanthera pungens 362 

aed Aas 


the a 362 
Antheridia, of Cryptomitrium 112 
Anthoceros lzevis, spore-mother cell 89 
Apetaly, significance of 281 
Aplectrum 
Apocynum 366 
Apospory, Stansfield on 439 
Apple, disease of 440 
ea ' eataage 135 
Araga 
8 a ex 
\rceuthobium pusillum 287 
\rchegoni ia, of Cryptomitrium 115 


~~ 


oO 
oe 
Ton 
Pay 
= os 
#5 FF og mt 
Qe 
ae 
“Oo 
Wp 
if 
° 
$ 
> 
_ 
-_ 


3,3 
phyllum meander 252 


Aspergiv pis 291, 378 
ter 366; divaricatus 36 

conde G. F. 1, 445 Ss 

Avena sativa 421 


Bacillus Betz 1 
79; aoe 65 
pacterial disease of beet 177 

+, personal oe 

.» “ Botanizing ” 281 
December 1899] 


Baker, J. G., personal 286 
Ball, C. R., personal 146 
Barnes, C. R., 71, 74, 140, 142, 207, 210, 
218, 276, 278, 281, 284, 364, 365, 366, 
367, 434, 439 
Batrachospermum Bohneri, fertilization 
of 282 


Ke; aobageage 444 


eattie, W. 
REE roots, disease 0 £75 
Bees, aie opic 27, 215; influence on 
flow 8 


Beet, barkerial disease of 177 
Belaje 


ney 
oe N., personal 285 
., personal Bek work of 281 
Beser, Er nst A., perso 8 
a beans ed of iteriean sags 445 
Bic l, E. P., work of 144, 367 


Bi 367 
Bienria; Douglassii stenophylla 375; 
Howardt 
Biological station 286; fei Winona Lake 
223; at Wood 


Hole 
Birds, porte peatlens soabaina of 44 


Blight of Sorghum 65 
e, E. E., work of 440 
Bombac 


zw 305 
Bo peel Society of America, Columbus 
meeting 210 
Botanical — Kew 286; Missouri 
6; P. 268; U. S. National 287 


Boubier, 

Hoasainaanttta ‘paselloides 437 
cach emum verticillatum 132 
Brissoni 

1 

] 


Bromus cilia tus 419 
ate L., “ The plant baby and its 


Brown, Ki 
ds’ 


friends” 7 
Bryologists, letter to 275 
Burt, Edward A., personal 79 


c 

Calcium oxalate, in buds of Prunus 282 
Caldwell, a Mae 73; pers 
Callitham Baileyi, 
Borreri, “noldfas s 256 


rsonal 7 
holdfasts 257; 


447 


448 BOTANICAL GAZETTE 


Calypso 
— hee ee work of 74 
noy, J. B., death of 445 
Cassa oil 368 
Cedrela 365 
Cell, ae centrifugal force 222 
Moet rook axicola 2 the oe peda 274 
Centrosomes, in Albugo 165 
oa 220 
ramium, meee iy holdfasts 258; stric- 
, holdfasts 258 
in. C.J. a 44; 279, 282, 438, 439 
parvula, holdfas 248 
Chloroplasts, of Anthoc is 
Chondria Awe Linp ila, holdasts 249; tenu- 
issima, ‘holdf 
ago arta in P Albugo 165; in Arisema 
Convallaria 338; 
in isactne a in Lilium 1 45; in Naias 
144; in Podophyllum Pee. in Potamoge- 
ton 349; in Trillium 
ee nes, affini 
$ 374; co 


374 affinis atten- 
i 


Pp i 

ficulatus 371; pumilus 
atus 376; pumilus varus 375; 

speciosus 370, 371; Vaseyi 377; Wyo- 


mingensis 372 
Cilia, Belajell 0 on origin 438 
ladophora oe 
Ps rk, J. F., 289, 9, 378% sae 79 
lements, F, spo 285 
Clitoria 36 a 
lothier, G 
Collins, G. " rscual 2 287 
Contributors: Abrams, LeRoy 110; Ar- 
ur, J. C. 139, 440, 43%; Ashe, W.W. 


ps Atkinson, Geo. 445; Barnes, 

C. R. 71, 74, 140, 14 i: see 210, 218, 

276, 278, 281, 284, 364, 365, 366, 367, 

434, 439; Caldwell, OW. 73: Cham- 

berlain, C. J. 144, 279, 282, oa : 1395 

Clark, f} F. 289 » 378; Cow GG. 
c 


: 4295 
ay, E. si D, 273: 
. M. 275; Hume, H. H. 

418; Hunter, A. A. (Hedgcock and ] 


[| DECEMBER 


425; Hyams, C. W. 431; Lamb, F. H 


e B. 360; Trelease, W. 

280, a Wiegand. K. M. 328 

Convallaria pesuicoan microspores of 330 

Co a tee F., personal 287 

Copela d, E. personal 223; work of 366 

Cosatcthizn 4 

orn, pears of 4 

pis re E; personal sf “Vermehr- 
aubmoose ” 

Coulter Joka | DH. 40, 72, Pr tas ait. 281, 

a 3003 personal ay 444; “ Plant 

7 366 


ea 278; w of 
Coulter, J. G., teckel! 9 be 
Coville, F. we fa) 
Cowles, H. C., E 4353 persona 


A irginiana 196; Virwinia na holosericea 


Se ieee sg bbe 20 
metremoatiie 412, 414; Bilt- 


Crategus, 
06; Boyntoni 409; Chap- 
271; coccinea 408, 410, 415; 
collina 271, 417; crus-galli ; glan- 
dulosa 408, 410; Harbisoni 413; Mohr 
416; mollis 407; rotundifolia 408, 4105 
Ss i 407; Sauratonae 270; silvi- 
col 14; tomentosa Ch ant 271; 
triflora 410, 413, 414; uniflora 271; 
Vailiz 271 
ob laseoriay tenerum I10 
Cunn am, C. A. 


177 
Cyprian insigne giganteum 368 
Cytas' 
Ctolowy, Abugo 149, 225; Arisazema 1; 
Anthoceros 89; Convallaria 330; Fu- 
n 343; tech- 
nique 279; ‘Trillium 1 


D 


Dahlia variabilis 436 
Darnel, cause of poisonous effect 136 


1899] 


Dasya elegans, holdfasts 253 
Davenport tC. Bs, paraded 146; ‘‘ Statis- 
oat peede a io 


282 
Decanisle, Ma grasa 285 
Seg 


DeToni, J. B. 268; personal 285 
Dewar, work of 439 
iatoms, Karsten on 439 
Diels, L. wo i of 364 
e 440; beech roots 75; beet 

Piaccach” 7 blig ht of Sorghum 65; 
bocednus (* peckiness”’) 
: e (Tilletia) Ta5: 
)74 


is 
“ianlg (and Wildeman), work of 74 
1, J. 


Duy W., personal 4 
Earle, F. S 
Eaton, D. C., 


03, 215, 2775 
ti ; 


ect of yids and alkalies on 


‘apt 

Babryo sac, of Composit 76 
Endocon idium temulentum 136 

Engler, A., “ Nat Pflancenfamilien Maly 


F 
Fairchild, D. G. 
Farmer (and Williams), es of 75 
erments 218 
Fernald, M. L. 1 62 
Fertilization, in Albugo 170, 225; in 
“ Batrachospermum 282; in Lilium 145 
oral W.,, personal 285 
Ischer, A., “ Fixirung, Farbung und Bau 
Protoplasmas 279 


246 
Fo S and insects rth 141, 215, 280 
btestry, primer of 365 


INDEX TO VOLUME XXVIII 


449 


Fothergilla rss piss take 
Fraxinus profun 

enti 7 tology of We serratus 253 
Fullm 


Fu ipa he a ani 445; compound o 


cies of Albugo 149, sae tertilization 
of Albugo isi 225; hypogeeous 281; 
mary of light on respiration 142; 
species ara: 273; toxic action of 
dcisieioas agents 289, 378 
G 
Gager, C; Si, tee 368 
Glade ‘acanlis 1 
Gale, G. P., perso 87 
alega, améigua 201; hispidula 199; 
paucifolia he se iscatoria 202; spicata 
: he ; Virginica 19 
Galera crispa pres 
moa rbense ses iid 216; caraca- 


216; hispi 
Gametophore, of Preissia 360 
Ganong, F., “Teaching botanist” 


a Lp Botanical Gardens] 


he Ye 
MI. ork o 

per Mathilde, gioie 3 . 

Gonolobus 367 

Green, J. R., “ Ferments and fermenta- 
tion” 21 

Greene, a L., personal eh 

Greenm ‘ B., rk o 

Grifithsia ’Bornetiana, eyo 255, 259 


sam 
Guignard, oi of 144 
Gyimnogracnine ne A 438 
H 


Barthold, work of 284 
Hansteen, Bar Sienchll ai 


Haplospora globosa ae 
Ha it E M., eld, forest, and 
wayside flowers , bs 
Ha ss, ork of 2 
Harshberger, J. W., work f 366 
Hart, J. W., ie : = 
artog, M., 
Hasselbring, ise i personal 79 
Hastings, G. J., per’ al 79 


s rson 
Hedgcock, G. G. (and Hunte r) 425 
Heller, A. A., personal 444; work of 144, 


307 
Hemerocallis fulva 81 


450 


Henderson, os F., work of 366, 440 

Hepatice 144; Cryptomitrium 110, 
Preissia 

Herbarium, Geol. Surv. Canada 368 

Hesperogenia 36 

eis spora Vidovi 213 

oria eomgy teunoaiie en 271 

Hil, Beji2 

Hillman, F. tL. , work of 440 

Hiltner, L., persona al 285 

Hitchcock, A. S. 429; work of 440 

Hohnel, F. aby personal 368 

Holm, vem 

Holway, E D. il work of 441 

Holanger I M. 

Hormann, G., “Cham ee sss crag of 
the living canteens 

Hosts : A.gopogon 273; Pa earns 233° 
Andropogon pa Cenchrus 274; Chloris 
273; Fimbristylis 274; Hilaria aS 
Hordeum 274; Oryza 138; Pani 


273,274; Paspalum 274; Rhyn ncospor 
274; Rumex 274; Setaria a 273; Tri 
cum 273; Triticum 274; 4 
ume, H. H. 418; personal 444 
Hunt, L. E., personal 288 
Hunter, - A. (Hedgcock and), 425; per- 
Pattee 
Symapernn 
Hydrangea opuloides, and insects 141 


Gaia 
Ue draseryie: icasrienis 362 


I 
Indiana Acad. Sci 


Insects and Sesick ay 141, 215, 280 
Ipomcea palmata 362 


J 


ay ees 288, 444 
al 78 


Jesup, H. G., pers 
Jussiza salicabenie 362 
K 

Kahlenberg, os Sey of 366 
Karsten, G., k of 439 
Kennedy, Bs personal 146 

ew Gantens 
Klebs, G., 


rk ee 441 
Knobel, E., “ Grasses, rip a and rushes 
of the northern U. S,” 
Knowlton, F. H., pena. 368 


BOTANICAL GAZETTE 


[ DECEMBER 


Knuth, P., death of 444; “‘ Handbuch der 
Bliitenbiologie ” 280, 432; personal 285 

rena Risirteaai I 32 

Kol , R., work of 142 


ls 
Lamb, F. H. 6 
Lang, W. H. , work of 2 
Lawson, A. A. , personal 288 


Leguminose q 
Lemna minor, ss desponive of proteids 284 
Lesquere emoration of 207 


» Leo mm 
Le acobryacke, clabsihoatt tion of 74 
Libocedrus decurrens, “ peckiness ” of 74 
Lichens 74 
Lignin i in buds e See 282 
Lilium Masse 
Lindau, work a sy 
Loeb, J., work of 74 
Logan’s fons e€ 445 
Lolium temulentum, cause of poisonous 


effect 136 
Lomentaria uncinata, holdfasts of 248 
Longyear, B. O. 272 
Lounsberry, ee “Guide to the wild 


flow 
Lahboek, J., “ Buds and stipules ” 277 
Lyco opodium, prothallia of 282 
Lyon, Florence M., personal 78 


M 


- Macbride, T. ae Spake nh 79 - 
D7. 


MacDouga 1, 
MacMilla n, C. 1430 
Makitios metal 144 
Malvacez eu 
Mar 


Merrell, W. D., 76; personal 78 

Mertensia 6 

saccet of pure 366 

Mex ico, eco sary oe of 366 
yer, W., work of 43 

Microsporangia, of Coneiiacis 330; of 
Hemerocallis 81; of Potamogeton 343 

Microspores, of Arisema 1 ; of Conv allaria 
330; of Fears soa 81 ; of Potamo 
geton ot ha Sagal 

of 36 66 


Mimosa fae 135 
Minnesota, photographs of re of 430 
446 


Missouri Bo ania Garden 
Miyake, te ., work of 143 
Montanoa 


rheatowy tT. personal 78, 223 


1899] INDEX TO VOLUME XXVIII 451 


‘ Moseley, E. L., “Sandusky flora” 139 Cowles, H. C. Daven 
Mosses, letter to bryologists 275 146; De Gandolle, C. 285 Deal, 
Mottier, ar M. Pee wes work of 222 M. Li, jr. 445; DeToni, 285; 
Mycorhia 220 avis Duvel, J. Ws ee; Bhenpayet: R444; 
Be Pre. ae Eriksson, A. I. 286; Figdor, W. 285 ; 
p rsonal 79 Fitzpatrick, T. J. 146; Gager, C. S. 368; 
Gale, G. P. 287; Gardiner, J. 146; 
N Giesenhagen, K. 285; Glatfelter, N. 
a 36 M. 286; Greene, E. L. 286; 
aias major, reduction in 144 W.146; eee H. ce Hestings 
id ska, Thorea n 425 G. J. 79; Heller, A. A. 444; Hiltner. 
Nelson, Aven ae 369; personal 223; L. 285; Hume, H. H. 4443 Hunt, L. 
hel of 144. 367 E. 288; Hunter, A. A. 285 ; Janczewski, 
Nelson, Elias, — of 1 “i ob ng J. e Gs : of saa Ww. 
Kiecs . 288,444; Jesup, H. G.78; Kennedy 
ae mbe, F. » personal 79; work of |B 146s Knowlton, F. H. 368; Knuth, 
Nisso lia 365 hide pelea Bagh ropa 
‘ Nomencl orence M.78; Macbride, T. H. 79, 
ate i: pp of cultivated plants 264 286 Martin, G. W. wat: Mbekes. 
Nylan os. 445; Merrell, E. D. 146; Merrell, 
Bet, Ws biography ot, 146 W. D. 78; Moore, G. T. 78, 223; . 
O Mottier, D. M. 223; Murrill, W. A. 79; 
Myers, P. C. 79; Nelson, Aven 223; 
(Edocephalum lok hte 291, 378 Newcombe, F. C. 79; Orton, W. A. 
Oils, in Albugo 147; Pollard,C. L. 286; Pollock. J 
Becshores. of Albugo Bliti 149, 225 79; Pound, R. 285, 286; Prillieux, E 
q “Nae of Albugo 225 78; Pringle, C. G. 285; Ramaley, F 
Fh fees Se personal 147 140; Rimbach, A. 148; Roberts, H 
. Ox a itibeed ath 435 F. 79; Rolfs, P. H. 78, 223; Rose, J 
tk Oxytropis 144 148; Savage, T. E. 79; S hrenk, H 
| P von 147; Setchell, W. A ee 
Shear, C. L. 445; Shimek, B. 70% 
Paddock, W., work of 440 Smith, W. R. 79; Snow, Julia W. 79; 
-adua, botanical garden 268 Spalding, V. M. 793 Swingle, W. T. 
Pagnoul, A., work of 367 224, 287; Timberlake, H. G. 78; 
?alladin . W., work of 366 Treub, 78; Trow, A. = 4443 
a ria ied , work a Underwood, L. M. 224; Waite, M. B 
am ern ork ¢ of 4 1473 Webber, H. I ras Wettivels, R. 
— n 286; Wiegan <7 


ru, desert Las ig ak See 
assiflora 365 ! Ferien 


hlo 
Photoaeibn collection of ecological 430 


irce, G. J., work of 74 Physiology, effect of pure me etals 366; 
benieiilinas ei 291, 378 effect of temperature on respiration 366; 
eppermint, _ sbeljeonasy of 368 ferment d fermentation 218; woe 
pereee grates sl rminating eat ; 
ersonals hia: Grist 444; Bailey, L symbio aprophyt ; 


: s 
H. 147; Baker, J. G. 286; Ball, C. R. synthesis of proteids 284 ; 
i i is force : 
366; toxic action on fu 
Be rst A. 289; P iaapiretinl 3 
a Burt, Edward A. 79; Caldwell, O. Picradenia macrantha 130; Richardsonii 


W.78; Carnoy, J. B. : fee 
79; Clements, F. E. sett Cokiinn: G. N. Pine t, G., ‘Primer of cameeg 365 
287; Cook, O. F. 287; Copeland, E. se echinata root sucker 
B. 223; Correns, C. E. 285; Coulter, Plant World 3 
J. G. 78; Coulter, J. M. 146, 444; lope work of 141 


452 


Podophyllum 20 
Pollard, C. L., personal 286; work of 
6 


307 
Pollen, of Arisz heat Sra 
30; of Pemcrocaite 81; of Pot 
eto of Trillium 1 
Pollock, J. B. personal 79 


Polysiphonia ee hora of 252; 
violacea, holdfasts of 2 

Potamogeton foli feet microspores of 343 
a a scoe, coq: 285, 2 

Preiaai commutata, hetaphrodite 


Prothali of Botrychium 282; of Lycopo- 


Pusndabins a 365 

Pseudo ota ‘axifotia, root suckers of 69 

Prunus a, buds of 282, insititia, 
what rs re 


oe emaculata 423; irregularis 419; 
raminis 418, 421, 422; het terospora 


— 
Lol 


] 
tomipara 419; triticulata 421 
i I 
pire saloer lanceolatum 132; mtilicum 
132; Torreyi 133; verticillatum 130 
Pyrenoids me 


Quercus ellipsoidalis 215; Texana 204 


R 


Radais, Maxime, 65 
Ramaley, F. personal 146 
paperioone in Arisema 1; in Fucacez 753 
aias ae in Bocepheylluin 20; in 
Trillium 1 
Reineria 19 
nities effect of temperature 366; of 
ungi 142 
— Bailey’s “ anpreiarsc 281; 
Brown’s 


“The ‘plant bald andoas 
friends” 73; Correns’s ‘“ Vermeh hrung 
d patina aR 434; Coulter’s “ Plant 


sg: - Pflanzenfamilien” 217; Fisch- 
rele “Fixirung, Farbung und Bau 

Protoplasmas” 279; Ganong’s 
ng « Teachnig botanist” 276; Green’s 


BOTANICAL GAZETTE 


[ DECEMBER 


“Ferments and fermentation” 218; 
Hardinge’ Ses Field, forest, and wayside 
flowers” 72; Hérmann’s “Chemical 
continuity of the living substance” 
141; Kno bel’s_ ho sedges, and 
rushes’ 72; ** Handbuch der 
Sane ape Le ate +432; Lounsberry’s 
“Guide to the wild flowers ” 72 & co ~ 
bock’s “Buds and es 277; M 
ley's “San ied flora’ 139; Pinchot’ s 

Pl aeals of fo city 365; Sargen 
“The plants, etc.” 73; Solereder's s 
“Anatomy of the dicot tyledons”” 140; 
Ver 


n’s “General physiology” 71; 
Wi idem n & Du rand’s “ Illustrations 
de la Floré du Congo” 74 


Ribes rubrum 146 
Rice, Tilletia on 138 E 
Ricinus communis, synthesis of proteids 


Rimbach, A. L., person 
Sar calh teniera, vanes af 247 : 
Span ag Catawbiense 272; maxi- 


Rob nal 79 
ce, Chases a 215, 4 
gps B. L. 134; 193; ie work of 


Rol SP: Te tig oe ae 223 
Sia disease of bee LP 
Root suckers, on 1 Douglas r 69 

Rose, J. N., personal 148; an of 281, 


36 5, 3 
Rusts, Holway on 441 
Rutaceze ee 
Rydberg, 423; bp of 467 
Panel ithe 


S 
a tn ita alegre 


362 
Salvi m, and insects 141 
Sarolc. ares on 443; Klebs on, 
01; Trow on 442 


ob is corn plants, ete.” 73 


€," tr. E. 28 erso 
anes speciosa 213 
Schlechter, R., pater of 143 
Schmidle, W., k of 282 


Schrenk, H. ey Taringa 147; work of 
be 

Seedl 

Seeds Pcl by fish 142; effect of in- 


nse cold on 43 
Seana sempervirens, root suckers ot 69 


1899] 


Setcheil, W. A., personal 78, 288 
Btiaality, in Tilopteridaceze 213 


7 
Silene rectiramea 4 verecunda 134 
Si isy rinchium 144, 367 


144 
mith, J. ws work of 144 
1% personal 79 


y, Julia ww, ees 79 
ow, Letitia M 
Solanum Tease ae 362; glaucum 362 
iain ee es “Anatomy of the dicoty- 
edon 


srg, ight of 65 
Sow “English Botany” 78 


S fae, . M., personal 79 

Spermatozoids, Belajeff on 4 

Spermoth ium ‘Turneri, *roldtaats of 
255, 2 


_ 2555 ag 
Spirogyra 
Sporophyte, i Cryptomitrium 117; origin 


eafy 4 
tet Bee: ae apt of 257 
ane fie 


Swingle, » personal 224, 287 
Symbiosis 220. 
Ss a in Anthoceros 96; in Convallaria 


336; in Potamogeton 347 
Synthesis of proteids in green phanero- 
284 


g i 
Tainionema I 
Taxodium, ‘ hee 
seed effect on respiration 366; 


199; flexuosa 


Mloteriiee a leiocarpa 195; lepto 
Lindheim 


I 
exuosa 200; Virginiana 196; Virgin- 
lana holosericea 196 


INDEX TO VOLUME XXVIII 


453 


Tetraneuris 126; acaulis 126; acaulis 
cespitosa 127; incana 128; lanata 129; 
Henenone 129; simplex 127; Torrey- 

x 


Thalietr m 365 

Thladiantha abs 437 

Thorea ramosissima 425 

Tilopteri aaaee iene | in 213 
Tilletia corona 138; horrida 138 
Timberlake, H. G., personal 78 
Townsend, Anne B. 360 

Toxic action on ‘ha 289, 378 

bh lepe S 367 

Zs ii e, W. eh 366 

Tre b, M. , personal 78 

Teil grandifloram 10 

Tro .. pe Aapoay 14443 work of 442 
r ate seis 

Tuber formation, Vochtin ng on 435 
Tumion Cali fornica, root backers of 69 
Turnera 365 


U 
Un ood, L. M., personal 22 
Unireniity Chica ago 78; init 79; lowa 
79; Michigan 79; Nebraska 79; Yale 
79, 147 : 
Urban, I., Porto Rican plants 444; work 
oF 


143 
ces ane 


nis 27 Hs ® andropoginis. 


27 P: 
Panici. ‘prolifer 274; Parlatorei 2748 


Rabe ein a 273; Tritici 274; Ulei 
273; Z 
Vv 
ail, A. M., work of 367 
Vargasia caracasana 216 
Venezuela = 
Verbesina I . 
Verworn, iiss, “ General peer = 


i Faba 366; synthesis of prote 


284 
Vilmo in, H. L. de, death of 287 
3 


Vochting, work of 435 


W 


Waite, M. B., personal 147 
haha L., work of 282 

Walteene sas 

Webber, H. J., personal 147 


(454 BOTANICAL GAZETTE [DECEMBER 1899 


Weeds, bulletins on 440 ye 

hetapes R. von, personal 286 

Hides egand, K. M. 328; Perea 79; work Xanthoxylum 365 

Wildeman ane Du rand), “ Tllustrations Z 
de a du Congo” Zaluzania 144 

Williams (and Farmer), work of 75 Zea Mays 222 


Wullschlegelia 220 Zostera marina 255 


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