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BOTANICAL GAZETTE
ee eee ee ee ee ee oe ee
THE
BOTANICAL GAZETTE
EDITORS:
JOHN MERLE COULTER anp CHARLES REID BARNES
VOLUME XLI
JANUARY—JUNE, 1906
WITH SIXTEEN PLATES AND SIXTY-TWO FIGURES
CHICAGO, ILLINOIS
PUBLISHED BY THE UNIVERSITY OF CHICAGO
1906
Mo.Bot Garagen
1906
PRINTED AT |
The University of Chicago Press
CHICAGO.
EE OE ee VTE ee ees
Oe TE EN FR
a a ee en
TABLE OF CONTENTS.
: PA
The nodes of grasses (with plates I and II) - Mintin Asbury Chrysler
The bogs and bog flora of the Huron river valley
(with sixteen figures) - Edgar Nelson Transeau
Nuclear division in Zygnema (with Ebiten Ii ai IV) Mabel L. Merriman
Effect of certain solids upoch the growth of seedlings in
water cultures (with four —— E = ‘J. F. Breazeale
Chemotropism of fungi - - - Harry R. Fulton
The embryology and development of Riccia fee
and Riccia crystallina (with plates V-IX) - - Charles E. Lewis
A morphological study of Sargassum fili pendula. Con:
tributions from the Hull Botanical Laboratory.
L
II (with plates Xand XI) - - - Etoile B. Simons
Chromosome reduction in the microsporocytes of
Lilium tigrinum (with plates XII and XIII) - John H. Schaffner
Cytological studies on the Entomophthoreae. I. The
morphology and development of iiss a
plates XIV and XV) - Edgar W. Olive
Cytological studies on the Pica phtbouee Il.
Nuclear and cell division of seis ore (witht gay
XVI Kee Edgar W. Olive
Biological relations of ae nabe Il. Aor
tion of water by leaves - V. M. Spalding
New species of Californian plants (with two ro figures) Alice Eastwood
New and noteworthy western plants. III. A, D. E. Elmer
Some littoral spermatophytes of the Naples region - J. Y. Bergen
New and noteworthy North American me of Tri-
folium (with twelve figures) —- - Homer Doliver House
Some studies regarding the biology of site and twigs
in winter (with eight figures) - - Karl M. Wiegand
The life history of Polysiphonia ioc Contribu-
tions from the Hull Botanical matoeg
LXXXIII - - Shigeo Yamanouchi
The structure. and Sevaoplnsli of the bark in the
Sassafras (with nine figures) - - - Howard Frederick Weiss
BRIEFER ARTICLES—
Notes on ak American Grasses. V. Some
Trinius Panicum types CS - A. S. Hitchcock
‘
GE
I
IIo
373
425
vi CONTENTS [VOLUME XLI
PAGE
oe in Pallavicinia = - - - - J.B. Farmer 67
- - Andrew C. Moore 69,
Notes on she relation between — of roots
and of tops in wheat. Contributions from
Hull Botanical Laboratory. LXXXI (with
five figures) - - - - Edward Burton Livingston 139
New normal icicles for use in = sa aad
ogy. III (with two figures) - W. F. Ganong 209
Notes on North American 2 et VI. $55:
nopsis of Tripsacum_— - A. S. Hitchcock 294
The basidium of Amanita bsporiger vith seven-_
teen figures) - - Charles E. Lewis 348
The distribution and 1 habits of some common
oaks = e ss 3 - - Picks Hill 445
CURRENT LITERATURE - - 71, 144, 214, 299, 353, 448
For titles of books et: see bisdete ar
author’s name and Reviews.
Papers noticed in ‘‘Notes for Students” are
indexed under author’s name and subjects.
News 65. eS a OE Oo ig ond ae a
DATES OF PUBLICATION.
No. 1, January 26; No. 2, March 3; No. 3, March 31; No. 4, April 28;
No. 5, May 31; No. 6, June 30.
ERRATA.
10, line 19, for (10) read (11).
14,.line 2 from bottom, for nternodes read internodes.
15, line 4 from bottom, for Contribution read Contributions.
75, line 16 from bottom, for East read West.
76, line 4 from bottom, for Perti read Petri.
. 167, line 19, for stand read stain.
168, line 13 from bottom, for comma read in.
77, line 1, after was insert in.
. 178, line 12 from bottom, for its read their.
- 179, line 11 from bottom, after paraphyses insert Ste anameneee
- 180, line 3, for The read Two.
. 340, line 12, for heller read Heller.
- 353, line 3 from bottom, for Petrostemonaceae read Podostemonaceae.
- 379, line 13, for Alluim read Allium
-
lea MaDe a a aa a ea a
Bete
Paeee pear een
Che Botanical Gazette
HA Montbly Fournal Embracing all Departments of Botanical Science
Edited by Jonn M. CovLtEer and CHARLES R. BARNES, with the assistance of other members of the
botanical staff of the University of Chicago
Vol. XLI, No. J Issued January 26, 1906
i MR EN Ti TE i a a el
.
CONTENTS
THE NODES OF GRASSES (wirH PLATES I AND It). Mintin Asbury Chrysler - I
THE BOGS AND BOG FLORA OF bei BRON, REVERS VALLEY Cigars SIXTEEN
FIGURES). Edgar Nelson Transeau
: NUCLEAR DIVISION IN ZYGNEMA (with PLATES I AND Iv). Mabel L. Merriman
i
|
VC
EFFECT OF CERTAIN SOLIDS UPON THE GROWTH OF SEEDLINGS IN WATER
CULTURES (witH rour FiGuRES). J. F. Breazeale - - - - 54
BRIEFER ARTICLES
NOTES ON Nor ae ae VY. Some Trinius PAnicum Types. A. S.
Hitchcoc - - . - - - - - - - - 64
AC eat IN PALLAVICINIA. J. B. Farmer - - - - - - - 67
REPLY. An C. Moe -- - - - “ -—" - - - -
CURRENT LITERATURE.
BOOK REVIEWS - - + - - ee ee A EF UR HE Be
THE ALGAL VEGETATION OF THE FAEROESE COASTS. ‘
PLANT DISEASES. REGENERATION. PLANT HISTOLOGY.
BIBLIOGRAPHICAL INDEX OF NORTH AMERICAN FUNGI.
MINOR NOTICES - - - es Snel men, Sew ae Maa ESE 76
NOTES FOR STUDENTS ~- - - - - - - - . - : - - 76
: NEWS - - . . - - - - - - - - - - - - - 80
6 Fal 42 tne tha Wats 1 oe 2 ae Cees + sho FTe ity of Chicago, Chicago, ll.
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VOLUME XLI NUMBER 1
BOTANICA (GAZETTE
JANUARY, 1906
THE NODES OF GRASSES.!
MINTIN ASBURY CHRYSLER.
(WITH PLATES I AND II)
ALTHOUGH the stems and leaves of grasses have received a good
share of attention from anatomists, and the bundles of the internodes
are perhaps sufficiently well known, the nodes have been largely
neglected. The reason for this may be the supposed difficulty of
unraveling the tangle of bundles found in»the node, or the obstacles
which the sclerified tissues offer to the preparation of satisfactory
sections. Yet the nodes are probably the most interesting regions
of the grass stem, for they lack the comparative uniformity of the
internodes. Since the application of the celloidin method to hard
tissues the difficulties of cutting the necessary serial sections have
been removed, so that we are now in a position to know intimately
the structure of these critical regions of the stem.
The object of the present account is to trace the course of the
bundles of the grass stem, and to discuss the significance of certain
structures which make their appearance at the nodes, in particular
the amphivasal bundles and cambium. The investigation has been
confined to forty-five genera, but since these represent the eleven
largest tribes and there is a considerable degree of uniformity in
structure, the account is believed to represent the family fairly from
the standpoint chosen.
The salient features may best be brought out by the description
of types selected to illustrate certain points. The first to be con-
sidered represents a medium condition as regards both taxonomic
position and ecological relations.
« Contributions from the Phanerogamic Laboratorics of Harvard University.
No. 3.
I
2 BOTANICAL GAZETTE [JANUARY
Avena barbata.—At a short distance above a node the stem pre-
sents a simple structure compared with that of many grasses, showing
just two circles of collateral bundles surrounding a central cavity.
The inner circle of bundles lies at the periphery of the central cylinder,
which in this genus is not clearly marked off from the cortex. The
bundles of the outer circle lie between groups of cells, which above
the leaf-sheath bear chlorophyll; they are considerably smaller than
the bundles of the inner ring, and from comparison with correspond- |
ing bundles in other genera must be regarded as cortical bundles.
In the upper part of a node these assume an amphivasal structure
and immediately anastomose with one another and with bundles
of the inner circle. But just at this level the structure of the stem
is further complicated by the entrance of bundles of the leaf-trace,
the course of which must now be described. The leaf-sheath in
this species extends a little more than 360° around the stem, and
contains, as do most of the genera examined, bundles of two distinct
sizes: larger ones, alternating with others which are less than Nalf
as great in diameter. These two kinds of leaf-trace bundles have
a different course in the stem. The larger bundles undergo a pro-
found modification as they enter the stem, as may be seen by com-
paring figs. r and 2, in which the magnification is the same. As
such a bundle enters the stem it rapidly increases in size, owing to
increase in the number of xylem elements. Most of the added
elements are tracheids with more or less suberized scalariform or
reticulate thickenings, but some parenchyma cells are also present.
These elements gradually extend around the sides of the phlcem
until this is surrounded by xylem, except a small area on its outer
side, which is generally occupied by sclerotic fibres belonging to the
group which is so well marked im the sheath (fig. 1). In certain
species, e. g., Lolium perenne, these fibres disappear, and the phlcem
is completely surrounded by xylem. These bundles evidently must
be placed in the amphivasal class. As fig. 2 shows, the phloem —
enlarges very little, but the xylem increases so much that the bundle :
may be five or SIX times as broad in the middle of the node as it is ~
in the leaf-sheath. These bundles are by far the most conspicuous —
objects in a cross-section of the stem at this level. They have 2
spindle shape which is not due to their oblique course, for they slant —
1906] CHRYSLER—NODES OF GRASSES i.
very slightly from the vertical and do not dip deeply into the central
cylinder. The xylem consists so largely of tracheids running irregu-
larly and mixed with parenchyma, that the mass has a considerable
resemblance to transfusion tissue. Apparently, this region in a
bundle forms an important water-storing organ. A further peculiarity
of the bundle at this level is the presence of a distinct bundle-sheath
or endodermis,? consisting of cells whose walls are reticulately thick -
ened and suberized. As these larger leaf-trace bundles descend through
the node, branches from the anastomoses mentioned above extend
outward between the leaf-trace bundles, and probably anastomose
with these, though the fusion is not so plain as in A. sativa. Below
this level the bundles gradually resume the ordinary collateral shape,
lose their endodermis, and run down in a single circle through the
internode as already described. The smaller leaf-trace bundles
also undergo some expansion as they penetrate to the boundary of
the central cylinder, but throughout their course they may be dis-
tinguished from the larger bundles, not only by their size but by
their early turning outward into the cortex and running down to
the next node as the cortical bundles described above.
Though it is not plain in A. barbata that the larger leaf-trace
bundles are joined, soon after their entrance into the stem, by other
bundles of the node, in A. sativa and in many other grasses it may
clearly be made out that on each flank of the leaf-trace bundle another
strand applies itself, swinging through an angle, so that its phloern
first joins on, then its xylem. In some species, e. g., Arundo Donax,
two or more bundles join on each flank of the leaf-trace bundle.
Certain features of the cortical strands are more clearly seen in
Panicularia americana. The cortical nature of these strands is
unquestioned, for they run quite outside-the central cylinder, in a
wide area of lacunar parenchyma (jig. 7). As they reach the upper
part of a node they anastomose with one another so as to form a
transverse ring or girdle (jig. 8), which, at a slightly lower level,
sends branches to the bundles of the central cylinder, forming nearly
‘or quite amphivasal bundles, though some of them very soon resume
2 The term endodermis is here used in the general sense employed by VAN
TieGHEM, rather than in the histological sense proposed by other writers.
4 BOTANICAL GAZETTE [JANUARY
the collateral structure which is characteristic of all the bundles of
an internode. Further down in the node, the bundles of the leaf-
trace enter the central cylinder and the smaller of these anastomose_
with branches of the nodal complex, and then turn outward to run
down through the internode as cortical bundles. The larger leaf-
trace bundles behave as in A. sativa.
In Leersia oryzoides it may plainly be seen from a series of sections,
that as each of the smaller leaf-trace bundles enters a node, it is
joined by two small bundles from the nodal complex, and this rein-
forced bundle proceeds downward through the sclerified cortex.
Whether the smaller leaf-trace bundles run down through the
cortex, or in the outer region of the central cylinder of an internode,
cannot in all cases be determined with certainty, for the boundary
of the central cylinder is often poorly marked, and the cortex may
be a very narrow zone. Or the boundary of the central cylinder
may be marked by a narrow sclerenchymatous ring, and the bundles
may lie along this, projecting either towards the inside or the outside.
But the position of these bundles inside or outside the central cylinder
appears to be a matter of indifference, and in either case they pursue
a different course from that of the larger bundles. In a general
way it may be stated that the smaller bundles of the leaf-trace, after
fusing with bundles from the nodal complex, run downward through
the next succeeding internode in the cortex or at its inner border,
and at the next node below join with the bundles of the central
cylinder. Species to which this statement applies are: Zizania
aquatica, Leersia oryzoides, Avena barbata, A. sativa, Panicularia
americana, P. nervata, Agropyron caninum, Elymus americanus,
Triticum sativum.
The course of the leaf-trace bundles in the grasses, as here de
scribed, differs in several respects from the course of such bundles
in other families, even in so closely related a family as the sedges,
recently described by PLowMAN (11). VaN TIEGHEM’s second class
of cortical bundles (6, p. 751) corresponds the most closely, and is
thus described: “Le faisceau médian de la feuille, qui en prend
trois, entre directement dans le cylindre central, tandis que les deux
latéraux descendent dans l’écorce pour n’entrer dans le cylindre
1906] CHRYSLER—NODES OF GRASSES 5
qu’au noeud suivant.” See further his remarks on the monocoty-
ledons.
Phalaris arundinacea, like Avena, has the bundles of its internode
crowded into an annular area surrounding the fistular pith. As
these reach the node they anastomose extensively, and at the same
time assume the amphivasal condition, which is shown with especial
clearness in the variety variegata, figs. 3 and 4, the latter more highly
magnified in order to show the tendency for bundles to form nests
of three or more, enclosed by an armor of sclerified fibres. These
amphivasal bundles, though abundant in the nodes, are absent from
the internodes.
Arundo Donax may be mentioned as typical of species having
several circles of bundles surrounding a central cavity. As the leaf-
trace bundles enter the stem they swell out, though not to so great
an extent as in Avena. The xylem completely encloses the phloem,
and the usual suberized sheath of cells with reticulately thickened
walls becomes visible. Farther down in the node the leaf-trace
bundle is joined on each flank by one or more cauline bundles.
The bundles of the latter class are provided with a sheath of heavily
thickened cells, and some of them appear to pass through the ncde
without anastomosing with other bundles, though this condition is
rare in the members of the family which have fewer bundles. In
accordance with the greater thickness of the solid part of the stem,
the leaf-trace bundles penetrate more deeply into the central cylinder
than in such genera as Avena, making their general course conform
more nearly to the palm type of von Mont. The number of circles
of bundles in an internode appears to be dependent on the size of
stem characteristic of the species, and to have little value in estab-
lishing relationships.
Grasses with a solid stem conform even more nearly to VON Mount’s
type, for the largest leaf-trace bundles penetrate nearly to the center
of the stem, before curving outward and downward toward the
periphery of the central cylinder. STRASBURGER (4) has given an
excellent account of the course of the bundles in Zea Mays. He
distinguishes leaf-trace bundles of five different ranks, and finds
that the largest of these penetrate most deeply into the stem, while
the smallest merely reach the periphery of the central cylinder.
6 BOTANICAL GAZETTE [JANUARY 4
The increase in complexity of the leaf-trace system over the condi-
tion found in Avena, seems to be associated with the greater size
of the leaf-sheath in Zea; just as a large stem generally has several
circles of bundles, so a heavy leaf has a better developed bundle 4
system. Most of the bundles in an aerial node of Zea are collateral,
leaving out of consideration the swollen leaf-trace bundles. STRAS-
BURGER (4, p. 348) finds amphivasal bundles at the point of origin
of axillary buds and adventitious roots; I have confirmed his obser-
vation in the former case. Much larger and more numerous amphi-
vasal strands are however to be found in the nodes of the axillary
branches bearing the ear of corn. One of these bundles is represented 5 1
in fig. 5. It is only in the leafy part of the branch that these occur, j
for in the “cob” the bundles are collateral, with an exceedingly well- .
developed phloem, doubtless associated with the transfer of elabo-
rated food. The amphivasal bundles of these branches are as usual
bundle fusions, and their occurrence in the reproductive axis of a
plant showing few elsewhere, seems to be a point of some significance,
especially when we consider that Zea is probably a highly organized
member of the family.
Zizania aquatica merits special attention on account of certain
features which may be regarded as primitive, e. g., the six stamens.
In an aerial internode a narrow cortex surrounds the hollow central
cylinder, and the two are separated by a ring of sclerified cells.
Partly imbedded in this ring are a number of small bundles, some
of which project into the cortex and on aceount of their origin must
be regarded as cortical bundles. All the bundles are collateral,
and those inside the sclerotic ring lie at different depths within the
central cylinder. As would be expected from the aquatic habitat
of the plant, the xylem is reduced; in fact, in some instances, it is
represented only by a cavity, and in all cases it has its vessels very
slightly lignified. The phloem does not share in this reduction.
As the node is approached the bundles at the periphery of the central
cylinder anastomose, at the same time becoming amphivasal, and
a number of transverse strands join up some of the inner bundles
of the stele with one another. At this level the leaf-trace bundles
enter the stem; they are of at least three ranks, and of these the
largest penetrate into the central cylinder, enlarging on the way;
ee eee ee Se te ee ee ee
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1906] CHRYSLER—NODES OF GRASSES 7
owing to increase in the xylem elements, which come to enclose the
phloem more or less completely, as already described for Avena.
A little further down they are joined on the flanks by several bundles
from the internode above, each leaf-trace bundle with its tangle of
contributing bundles forming a complex bunch of vascular tissue.
A suberized endodermis surrounds the leaf-trace bundles in the node.
The second rank of bundles of the leaf-trace also enter the central
cylinder, where they are joined by other bundles, but soon return
to the periphery of the central cylinder, where they give rise to some
of the bundles which run through the next internode below on the
border line between the sclerotic ring and the cortex. The smallest
bundles of the leaf trace go no deeper than the sclerotic ring, and
here fuse with other bundles found in this zone. Thus, the course
of the bundles of different rank agrees with what STRASBURGER
found in Zea. In the lowest part of the node, the very numerous
bundles of the sclerotic ring anastomose and proceed downward,
greatly reduced in numbers, while the bundles inside the central
_cylinder also become much fewer, owing to completion of the fusion
of the large leaf-trace bundles with cauline strands.
The basal region of the stem has a cortex which differs from the
aerial parts in being much broader and more spongy, on account
of the large intercellular spaces. At any of the basal nodes the
central cylinder is bounded by an endodermis, consisting of a single
layer of rounded cells with suberized walls (fig. 6). Inside this is
a narrow zone of vascular tissue, whose elements run circularly;
then a wide zone, consisting of small bundles running vertically,
and so closely packed together that it is generally impossible to dis-
tinguish their limits. Bounding the two vascular rings on the inside
is a band of sclerified cells which are in contact with the pith. As
the photograph shows, leaf-trace bundles make their way into the
central cylinder through wide gaps in this four-layered ring, and
it may be clearly seen, even in unstained sections, that along the
edges of such a gap the external suberized endodermis is continuous
with the inner sclerified layer. All of the large leaf-trace bundles
pass through such gaps, but the roots leave the central cylinder
without causing a gap, as has been observed in plants of various
groups. In the pith of the central cylinder are scattered many
8 BOTANICAL GAZETTE [JANUARY
bundles, nearly all of which are amphivasal, and where the nodes
are crowded, as they are at the very base of the stem, the amphivasal
condition is retained by bundles from one node to another, though
in the more elongated internodes, found a little higher in the stem,
only collateral bundles occur. The contrast between the upper
and the basal nodes is indeed striking, for the former show no amphi-
vasal bundles running longitudinally in the pith, except the enlarged
leaf-traces. This feature of the aerial node may be partly accounted
for by the thinness of the diaphragm in which run the anastomosing
bundles, which are generally amphivasal.
AMPHIVASAL BUNDLES
_ Although these have been repeatedly reported as occurring in
the subterranean stems of monocotyledons (see STRASBURGER, 4,
p. 348, footnote; DEBARY, 2; SCHULZE, 5)3, the only references to
their occurrence in grasses that have been found are by STRASBURGER
(4) and Jerrrey (9). Duvat-Jouve (1) figures the rhizomes of
many grasses, but shows no amphivasal bundles. Yet an cxamina-
tion of the nodes of some of the same species shows that the bundles
in question occur here. Two sorts must be distinguished in this
family: (1) the swollen portion of a leaf-trace bundle, the xylem
consisting largely of a mass of tracheids running irregularly; (2) the
type usually figured, the xylem forming simply a ring of vessels.
The features of the first class have been described under Avena.
Such a bundle is always enclosed by an endodermis which generally
has pitted or reticulated walls, and shows, in addition to the ordinary
metaxylem elements, a large number of reticulated tracheids, which
almost or quite enclose the phloem. All the species examined show
these bundles, from hydrophytes such as Zizania to xerophytes such as
Ammophila, and there appears to be no relation between the size
which a bundle attains at its widest part and the condition under
which the species grows. If, as already suggested, these bundles
serve to store up water, it might be expected that they would be
poorly developed in aquatics, but the only peculiarity of the bundles
found in such plants is the slight lignification of the xylem, a char-
3Since the above was written, Horm has reported the occurrence of amphivasal
bundles in the rhizome and also the aerial stem of Croomia pauciflora (Amer. Jour.
Sci. 20:50-54. 19¢5).
Be ee
1906] CHRYSLER—NODES OF GRASSES 9
acter shared by all the bundles of such plants. Quite commonly,
the bundles are surrounded by a layer of parenchyma rich in chloro-
phyll. This suggests that the node is an active assimilating organ.
Bundles of the second class—amphivasal bundles as usually under-
stood—are found in the nodes of the great majority of the grasses
examined, but are especially numerous in the following species:
Coix lachryma, Paspalum stolonijerum, Panicum sanguinale, Sor-
ghum halepense, Leersia oryzoides, Phalaris arundinacea, Calama-
grostis canadensis, Avena barbata, Panicularia americana, Panicularia
nervata, Festuca arundinacea, Triticum sativum. The fact that they
are practically confined to the nodal regions, suggests that they are
associated with bundle fusions, and this assumption has been amply
borne out by observation. Further, since the bundles which fuse
are usually traces of leaves which come off higher up, it appears
that the occurrence of such bundles is to be referred to the leaves.
The closed mode of venation, prevalent in the monocotyledons,
involves that a large number of bundles shall run down parallel to
one another through the petiole or sheathing base of the leaf. In the
grasses the numerous bundles are accommodated in the leaf-sheath,
which frequently encircles the stem for somewhat more than
360°. The large number of bundles cannot at once find room in
the vascular ring, which we may believe constitutes the primitive
stele in both dicotyledons and monocotyledons, according to the
results of JEFFREY (8), and CHRYSLER (10). Hence the leaf-trace
bundles, or some of them, pass into the inside of the central cylinder,
and sooner or later join other bundles. It will be readily seen that
a bundle lying in the pith has a better chance to orient itself with
regard to some other bundle which it may join, than has a bundle
which merely fits itself into a gap in a vascular ring (e. g., the leaf-
trace of a fern such as Adiantum). Hence it is not surprising to
find that before two bundles of a monocotyledon fuse, they swing
around, so that phloem fuses with phloem, and the xylem accordingly
surrounds the compound bundle. How far beyond the point of
fusion of the bundles this amphivasal condition persists, is a feature
which varies greatly. In most of the grasses the collateral structure
is soon resumed, but the example of Zizania shows that, at the base
of the stem where the nodes are crowded, the amphivasal condition
10 BOTANICAL GAZETTE [JANUARY
may continue through several internodes. This probably accounts
for the prevalence of these bundles in the rhizomes of monocotyle-
dons, where they were first observed. A comparison of the aerial
and subterranean nodes of the grasses under study has not yielded 4
results of great significance; most species show no noticeable differ-
ence in the number of amphivasal strands in the two cases. But
in Andropogon jfurcatus, A. scoparius, Chrysopogon avenaceus,
Zizania aquatica, and Phleum pratense, the amphivasal strands are
distinctly more numerous in the basal nodes. No examples of the
opposite condition have been found. Querva found in Gloriosa
(7) that the amphivasal bundles are connected with the origin of
a branch. Among the grasses, Phalaris arundinacea, Paspalum —
stoloniferum, Sporobolus Wrightii, Coix lachryma, and Zea Mays :
show these bundles at the point of origin of branches, but in other
species only collateral bundles could be discovered at these places.
Too much importance should not be attached to the few cases named,
in view of the fact that the sedges uniformly show amphivasal bundles
associated with leaves and not with branches, as PLOWMAN has _
shown (10). This is one reason for considering the grasses a more
specialized group than the sedges; in fact it may be premised that
the amphivasal condition originally connected with leaf-traces has
in the Gramineae spread to the branches. The occurrence of amphi-
vasal bundles in the leafy reproductive axis of Zea, while they are
rare in the main stem, deserves emphasis. While many of the
grasses show amphivasal bundles in all the nodes, in this highly |
developed genus the bundles in question have nearly disappeared
from the ordinary nodes, but .have persisted in the conservative
region named. STRASBURGER proposes (4, p. 348) a physiological
explanation for the occurrence of these bundles, viz., that the amphi-
vasal structure is favorable for the taking up of reserve materials
stored in a rhizome, but this explanation is not in accord with the
accepted view that elaborated food is carried not by the xylem, but
by the phloem. Examination of serial sections leads to the opinion
that the mechanical necessities of bundle fusion rather than con-
siderations of absorption of food have been the determining factor
in producing these bundles.
If we accept the view advanced above, that these bundles are to
1906] CHRYSLER—NODES OF GRASSES Ti
be associated with the large number of leaf-trace bundles of the
monocotyledons, their phylogenetic significance is considerable.
The ferns and dicotyledons have a comparatively small number of
leaf-trace bundles; amphivasal bundles are absent in the former
and rare in the latter group, but are widely distributed in the mono-
cotyledons, which accordingly appear to represent a more recent
and highly specialized group. In the most highly organized members
of the Gramineae, such as Zea, is shown a tendency to reduce the
number of amphivasal strands, but even in such cases they may
linger on in the reproductive axis. JEFFREY (9) has called attention to
the fact that in highly organized families, such as Tridaceae and
Orchidaceae, these bundles disappear even from the reproductive axis.
CAMBIUM.
A feature of the vascular bundles of Avena barbata, not so far
mentioned, is represented, in fig. 9, which shows a bundle from the
stem at a distance of about 1™™ above one of the upper nodes. That
the tissues are immature is shown by the presence of protoplasm
and a nucleus in certain of the vessels. The shrunken protophloem
is represented by the dark band at the outer edge of the bundle, and
the metaphloem as usual has its elements arranged irregularly.
Between these elements and the vessels are a number of rows of
flattened cells, radially arranged, corresponding well to the cambium
of dicotyledons. This feature is not confined to the young stem,
as is seen in jig. 10, which represents a bundle from above one of
the lower nodes of the same plant. A few tangential divisions are
to be seen in the leaf-trace bundle shown in fig. 1, though cambial
activity here is slight. Toward the node and farther up in the
internode and sheath, this peculiarity is not shown by the bundles,
but at the regions mentioned most of the bundles have a more or
less clear indication of cambium.
In the leaf-sheath of Andropogon argenteum (fig. 11), at a distance
of 1-2™™ above its insertion on the stem the larger bundles show
an unmistakable cambial layer. From the small amount of phloem
or xylem showing radial arrangement of its elements, it appears
that the cambium is functional for only a short time. Sections.
through the leaf-sheath, cut 5™™ above the one shown in the figure,
still exhibit a layer of radially arranged cells exterior to the xylem,
12 BOTANICAL GAZETTE [JANUAR
but the cells have thicker walls and a rounded outline, indicating
that in that region the period of activity of the cells is past. The
stem of this plant does not show good examples of cambium, nor do
A. scoparius or A. jurcatus show the feature, even in the leaf-sheath.
A further example is shown in fig. 12, which represents a bundle
from the stem of Erianthus Ravennae 1-2™™ above the level of inser-
tion of the leaf-sheath. A similar appearance is presented by the
bundles of the sheath. The leaf-trace bundles of Zizania frequently
show a large amount of their phlcem radially arranged, in spite of
the fact that dicotyledoncus aquatics generally show a marked
reduction of their cambium. The examples so far cited include
only the more striking instances of cambium found in the family.
More or less plain evidences of cambium have also been observed
in the following species: (1) In both stem and leaf-sheath; Coix
lachryma, Panicum crus-galli, Avena sterilis, Lolium italicum, L.
perenne. (2) In stem; Tripsacum dactyloides, Miscanthus sinensis,
Pennisetum longistylum, Panicum sanguinale, Leersia oryzoides,
Sporobolus Wrightii, Calamagrostis canadensis, Arundo Donax, Avena
sativa, Briza maxima, Panicularia americana, Bromus inermis,
Triticum sativum. The occurrence of a cambium in the region
just above the nodes in grasses recalls the well-known power which
members of this family possess of bending upward at these regions
if the stem is by any means laid horizontally. In this connection
it is of interest to note that PLowMAN (11) has found only traces of
cambium in the sedges, and in line with this, the stems of sedges
are unable to right themselves if bent over into a horizontal position.
Miss ANDERSSON (3) has reported the occurrence of a more or less
plain cambial zone in the young plants of representatives of many
monocotyledonous families. She calls attention to the similarity
between the bundles in the seedling of Lilium and those in Ranun-
culus. The only grass referred to is Zea, in which the mature _
bundles often show a radial arrangement of the cells between xylem
and phloem, as is illustrated by the well-known figure in SACHS’
Text-book. From the occurrence of cambium in the tuberous stems
of Gloriosa, QuEvA (7) has already concluded that the mono-
cotyledons are derived from dicotyledons. In the case of the
grasses it would seem that the cambium possessed by the ancestors
i
4
1906] CHRYSLER—NODES OF GRASSES 13
has been retained in the regions where it is of use. On the other
hand, it may be argued that we have here the first appearance of
a feature which in the dicotyledons becomes prevalent. But why
should the cambium appear only at the nodes, where it is of use ?
It may be regarded as almost axiomatic that the need for a struc-
ture is not a sufficient cause for its appearance. So it seems more
reasonable to read the evidence in the way first suggested, viz.,
that we have here a relic of a structure which was present in the
ancestors of the grasses, but has disappeared from most parts of
the plant and from most families of monocotyledons, and_ is
retained above the nodes of grasses in connection with their power
of bending at these regions: Thus the evidence favors the derivation
of the grasses from ancestors having a cambium.
The stele of the grass stem has evidently departed widely from
the primitive protostele or siphonostele. It has been repeatedly
shown that reproductive axes are able to retain ancestral characters.
An examination of this region in seventeen species of grasses belong-
ing to seven of the tribes has failed to disclose any instances where
the stele presents the primitive type described by PLOwMAN (11).
This result seems to confirm the opinion derived from other con-
siderations, that the Gramineae represent a more specialized group
than the Cyperaceae. These considerations may now be stated
categorically:
(1) The grass family has adapted itself to every habitat, from
salt marsh to pampas, and shows every gradation in habit from
the bamboo downward. The sedges are prevailingly hydrophytes,
and few of them attain a considerable size.
(2) The hollow stem characteristic of most grasses has probably
been derived from a solid stem such as is present in the sedges and
most monocotyledonous families.
(3) Amphivasal bundles are not found in so large a proportion
of species nor are. they as numerous in an individual in the grasses
as in the sedges.
(4) In practically all of the grasses the leaf-trace bundles are of
at least two ranks, while the sedges show no such distinction.
(5) The floral axis of the grasses dees not present the simple
type of stele shown by some sedges.
14 BOTANICAL GAZETTE [JANUARY
On the other hand, the open sheath of the grass leaf may be con-
sidered to be more primitive than the closed sheath of the sedges.
Further, the cambium found in the grasses is here considered to be
a primitive feature. But on the whole the Gramineae are to be
_ regarded as a more highly specialized family than the Cyperaceae,
though the families are evidently very closely related.
Certain anatomical features of the grasses, such as the distribu-
tion of the amphivasal bundles, seem to have an important bearing
on the phylogenetic position of the family among monocotyledons,
but since the anatomy of the group is as yet very imperfectly known,
a discussion of this point would be premature.
SUMMARY AND CONCLUSIONS.
1. The grasses depart considerably from the scheme propcsed
by von Mout for the course of the bundles, chiefly owing to the
stem being hollow in most cases. The leaf-trace bundles are of at least © 4 j
two ranks; of these the largest penetrate most deeply as they enter
the central cylinder, generally receiving one or more bundles on
each flank as they pass downward to the lower part of the node;
the smaller leaf-trace bundles do not penetrate deeply into the central
cylinder, but after anastomosing with other bundles pass downward
either in the cortex or at the inner border of this. At the next node
lower, these cortical bundles anast mcse with one another, and
then with the bundles of the central cylinder which have come from
fusions ‘with leaf-trace bundles at the next node above.
2. The leaf-trace bundles, especially the larger ones, undergo
a marked change as they enter the stem. This consists in the appear-
ance of a distinct endodermis, and in an increase in the xylem,
leading to the formation of a greatly swollen amphivasal bundle.
Below the node these bundles resume the collateral type.
3- Amphivasal bundles of the ordinary type, though absent in
ihe ce internodes, are very commonly found in the nodes, and
arise by fusion of collateral bundles which are generally leaf-trace —
bundles. In some species they are more numerous in the nodes
at the base of the plant, and where such nodes are crowded, the
bundles may retain the amphivasal condition through successive
internodes. The presence of amphivasal bundles in reproductive
branches of plants in which these bundles are scarce in ordinary —
aa
1906] CHRYSLER—NODES OF GRASSES 15
nodes, points to their being an ancestral feature, which, in highly
organized members, has disappeared from most parts of the plant,
but is retained in the conservative flowering axis. It appears that
the amphivasal bundles so characteristic of monocotyledons, in all
probability made their appearance in connection with the entry
of numerous leaf-trace bundles into the nodes, but that secondarily,
in certain instances, they are found to be related to branching.
4. A well-marked, though generally short-lived, cambium occurs
in the bundles just above the node or near the base of the leaf-sheath
in certain grasses. This fact is considered to lend support to the
view that monocotyledons have been derived from some group
possessing a cambium, probably the dicotyledons.
5. The anatomical features of the grasses point to their being a
more highly specialized family than the sedges.
This investigation has been carried on in the Phanerogamic
Laboratories of Harvard University. I am indebted to Professor
G. L. GoopateE for material, and to Professor E. C. JEFFREY for
material and for advice during the progress of the work.
HARVARD UNIVERSITY.
LITERATURE CITED.
Norte.—Practically nothing bearing immediately on the subject of this
research has been found in the older literature, so it is not cited here. References
to it may be found in DEBAry (2), and the text of Kny’s Wandtafein.
1. Duvat-Jouve, M. J., Etude anatomique de quelques Graminées. Mém.
Acad. Sci. Montpellier 7: 309-406. 1
2. DeBary, A., Comparative anatomy of the phanerogams and ferns (trans.).
1884. ;
3. ANDERSSON, S., Ueber die Entwickelung der primaren Gefassbundelstrange
der Monocotylen. Review in Bot. Cent. 38:586, 618. 1889
4. STRASBURGER, E., pce den Bau und die Verrichtungen der Leitungs-
: nen. Jena. 1
5. Scuutze, R., sieht zur vergleichenden Anatomie der Liliaceen, Haemo-
doraceen, Hypwicidoak: und Velloziaceen. Bot. ‘Jahrb. 17: 295-394. 1893
6. VAN TiecHEM, Ph., Traité de Botanique. Paris. 1891.
7. Queva, C., Contribution & l’anatomie des monocotyledonées. I. Tra-
vaux et Mém. Univ Lille VII. 22:1-162, pls. 1-11. 1
8. Jerrrey, E. C., The morphology of the central cylinder in the angiosperms
rans. Can. Inst. 6:1-40. pls. 7-11. 19920.
16 BOTANICAL GAZETTE [JANUARY
9- re E. C., A new key to the phylogeny of the monocotyledons. Science
N. 5. 37: eee. 1903.
10. CHRYSLER, M. A., The development of the central cylinder of Araceae and
Liliaceae. Bor. GAZETTE 38: 161-184. pls. 12-15. 1904.
11. PLowMan, A., The comparative anatomy and phylogeny of the Cyperaceae.
Annals of Bot. (ined.) vol. 20.
EXPLANATION OF PLATES I AND II,
PLATE I.
Fic. 1. Avena barbaia; leaf-trace bundle in the leaf-sheath 1™™ above its
insertion on the stem. 130.
2. Same; swollen and amphivasal condition of a leaf-trace bundle in
the middle region of node. X 130.
Fic. 3. Phalaris arundinacea; section through upper part of a node; near
the outside are the leaf-trace bundles, alternating large and small; just internal
to these are the nests of amphivasal bundles. X30
1G. 4. Same; one of the nests of amphivasal bundles more magnified; on
three ie are leat bic bundles. X 150.
Fic. 5. Zea Mays; one of the large amphivasal bundles from a node of x
reproductive branch. 115.
Fic. 6. Zizania aquatica; part of stele and cortex in basal region of the stem.
Two gaps with their leaf-trace bundles are visible. 35.
PLATE II.
Fic. 7. Panicularia americana; section 1™™ above node, showing two
cortical sae X45.
ame; section from upper part of the node, just above insertion of
Kskakoa. The cortical bundles are connected by a ring-shaped anastomosis.
x45.
Fic. 9. Avena barbata; bundle from a young stem 1™™ above insertion of
the leaf-sheath, showing cambium. X 150.
IG. 10. Same; bundle from the same position in a mature stem. 150.
Fic. 11. Andropogon argenteum; bundle from the leaf-sheath 1-2™™ above
its insertion on the stem, showing cambium. X 150.
Fic. 12. Erianthus Ravennae; bundle from the stem a short distance above
anode. X150.
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THE BOGS AND BOG FLORA OF THE HURON RIVER
VALLEY.
EDGAR NELSON TRANSEAU.
(WITH SIXTEEN FIGURES)
[Concluded from p. 448.]
IV. The ecological characteristics of the bog flora and their causes.
The plants occurring in the bog habitat are almost all perennials.
In the case of the herbaceous vegetation, the winter is passed by
means of subterranean rootstocks. The shrubs are in part evergreen
and in part deciduous. The tamaracks and the two birches are
deciduous, and the black spruce and pine are evergreen.
Most of the herbaceous and shrubby forms multiply abundantly
by vegetative shoots of one form or another. The length of the
underground stems of the shrubs is proverbial, but is best appreciated
by one who has attempted to dig up one of them entire. In con-
nection with the competition between species for space in the habitat,
this is of the greatest importance. A luxuriant growth of cassandra
furnishes the most :favorable situation for the development of sphag-
num in this vicinity. Its profuse branching affords a framework
for the upbuilding of the sphagnous layer, its shade properties do not
interfere with the photosynthetic work of the moss, and it protects
‘it from the drying effects of wind and direct insolation. Where such
associations occur, the difficulties presented for the germination for
most seeds, and the efficiency with which competition is combated,
are evidenced by the fact that among the tree species only the tama-
rack, spruce, and pine are successful invaders. All of these plants
send out adventitious roots from the stems and branches, and so keep
pace with the upward development of the moss. The absence of
poplars, willows, red maples, and elms in such undisturbed situations
must be in part attributed to the completeness with which such terri-
tory is controlled by the cassandra-sphagnum association.
ECOLOGICAL ANATOMY.
Aside from the purely aquatic forms which have received much
Botanical Gazette, vol. 41-] [17
18 BOTANICAL GAZETTE [JANUARY
ecological attention, it is of interest to look at the anatomical char-
acteristics of certain of these plants.
Eriophorum virginicum may be taken as a type of this group, and
also of the sedge zone vegetation in general. The culm is very
slender and erect, leaves flat, and very narrow, perennial by root —
stocks. Stem: epidermis very thick-walled and cuticularized. As
development proceeds, certain radial rows of the primary cortex
cells have their walls thickened, and served to connect the tissues of
the central cylinder with those of the three-or four outer layers of
hypodermal cells which also become thick-walled. Between these
radial groups of cells lysigenic air cavities are formed. Root: epi-
dermal cells in part thin-walled and in part secondarily thickened,
no definite arrangement of the thick-walled cells apparent; internal
structures closely resemble those of the stem; no mycorhiza present.
Leaj: outer epidermal cell walls strongly thickened and cuticularized,
radial and inner walls less so; lysigenic air spaces traverse the leaf
longitudinally; a very thick layer of stereome adjoins the leptome,
decreasing to one or two cell layers on the hadrome side of the
bundle; chloroplasts massed among the outer layers of the cortex, but
occur throughout.
Sarracenia purpurea.—Well known for its insect-capturing
pitchers. Stem: epidermis and first hypodermal layer thick-walled;
lysigenic air cavities throughout pith and cortex; resin deposits
confined to the epidermis and one or two hypodermal cell layers, but
where wounded heayy deposits of resin take place in the exposed
and underlying cells. Root: cell walls firm, resinous bodies present —
throughout, but especially prominent in the two outer cortical layers,
in which the cell walls are also strongly thickened. Leaj: epidermis
thick-walled and, slightly cuticularized; stomata on both sides of
the lamina, with guard cells strongly cuticularized and slightly
protuberant; resinous deposits throughout; inner face of lamina
with strong downward pointing bristles.
Oxycoccus macrocarpus.—Stem: pith thick-walled, with resinous
bodies; a thick layer of broad-celled bast forms a complete cylinder
within the epidermis. Leaf: margins revolute, upper epidermis
without stomata, heavily cuticularized, radial walls thick, wavy;
hypodermal collenchyma of two or three cell layers on leptome side
1906] TRANSEAU—BOGS OF THE HURON RIVER VALLEY 19
of midvein, one or two cell layers on the side of the hadrome, develop-
ment of the stereome cells also smaller on hadrome side; palisade
of two cell layers; lower epidermis covered with wax, especially at
* the stomata, guard cells slightly elevated. Mycorhiza present in the
larger roots, wanting in the hairlike branches, no root hairs. -
Andromeda polijolia—Leaf: margins revolute, upper epidermal
cells thick-walled, radial walls undulate, no stomata; lower epidermis
supplied with unicellular short stiff hairs, and covered with wax,
stomata slightly protuberant, strongly cuticularized beneath mid-
vein; palisade of three layers of long narrow cells; stereome strongly
developed above and below vascular bundle; on the ventral side
this adjoining three layers of large thin-walled air cells and a one- |
layered hypoderma. Root: resinous deposits throughout, no mycor-
hizal fungi found.
Chamaedaphne calyculata.—Leaj: margin slightly revolute, epi-
dermis thick-walled, heavily cuticularized, cuticle rough, no stomata
on upper surface; ventral epidermis covered by shield-shaped multi-
cellular hairs, and a deposit of wax; cuticle unusually thickened
beneath the midvein, guard cells sunken, subsidiary cells protuberantr
palisade tissue of four or five layers. Root: inner and radial walls
thickened, cortical tissues thick-walled; resin deposits in vascular
bundle and cortex; no mycorhizal fungi found.
Chiogenes hispidula—Lea}: margin revolute, epidermal walls
very thick, cuticle present, papillate, palisade not strongly developed;
mesophyll cells in part thick-walled and in part thin-walled; resinous
bodies in the epidermis; stomata slightly protuberant. Stem: resin
Present in cortex; mycorhizal fungi in the epidermis of the smalle;
Toots and throughout the cortex of the larger.
Vaccinium corymbosum.—Leaj: cuticle present, epidermal walls
hot thickened, palisade of one layer, mesophyll tissues with resinous
bodies, cuticle of ventral surface papillate; abundant unicellular
irs on lower epidermis few on upper; leptome side of mid-vein
adjoined by three layers of stereome and two or three layers of hypo-
dermal collenchyma,.on the hadrome side reduced to two of stereome
and two of collenchyma, cuticular papilli usually developed beneath
the midvein and at edge of leaf. Root: cortical tissue with resin,
mycorhiza present. No resin deposits found in stem.
20 BOTANICAL GAZETTE [JANUARY
Salix sericea.—Leaf: upper epidermal cells small, strongly
cuticularized; mesophyll compact, palisade of two layers of long
narrow cells; stomata on under surface, guard cells sunken beneath
the slightly protuberant companion cells; hypoderma of five- or
six-cell-layers on hadrome side, and eight layers on leptome side of
midvein. Root: resinous bodies present in medullary rays and
cortex, the latter consisting of thick-walled cells; no mycorhiza.
Ledum_ groenlandicum.—Leaj: upper epidermis rugose, with
scattered unicellular hairs, margins strongly revolute, cuticle present,
cell walls thickened, the radial walls being broadly undulate; lower
epidermis covered with a thick cuticle and a felt of long multicellular
and short unicellular hairs, glandular hairs usually present near the
small veins, stomata protuberant; palisade of three or four layers
of broadly oblong cells; beneath vascular tissue of midvein and
between the mestome bundles occur large air.cells which may form
lysigenic air cavities in the older leaves. Root: mycorhizal.
Larix laricina—Leaf: bifacial, deciduous; epidermis thick-
walled, slightly cuticularized, guard cells sunken beneath the com-
panion cells; palisade tissue developed toward the dorsal surface,
two layers thick showing a radial tendency, stereome reduced to a
few cells beneath the leptome; two resin ducts near edges of leaf.
Root: composed of mycorhiza, resinous deposits throughout, cortical
tissues early destroyed by fungus. When grown in culture solutions
and well aerated, normal roots with root hairs are produced.
Picea Mariana.—Plants in bogs are stunted. Leaj: epidermal
cells thick-walled, cuticle present, guard cells sunk beneath the
companion cells; mesophyll cells compact, of a more or less radial
palisade type. Root: mycorhizal, resin deposits throughout, cortical
tissues destroyed by ieee Normal roots are developed under
culture conditions.
Pinus Strobus.—Plants very much stunted-in the bogs, leaves
shorter and thicker. Leaj, epidermal walls so greatly thickened
that scarcely a lumen remains, beneath this a hypodermal layer of
thick-walled cells; mesophyll cells compact and of the usual lobate
type. Root: mycorhizal, cortical tissues traversed by the fungus
hyphae; resinous deposits throughout. Stem: annual rings narrow
Sr
1906] TRANSEAU—BOGS OF THE HURON RIVER VALLEY 21
and distorted, resin bodies throughout cortex and meristematic
tissues of the wood.
To summarize these characteristics, it is evident (1) that epidermal
and hypodermal tissues are thick-walled; (2) that for the ccnserva-
tion of water these are reinforced outwardly by a heavy cuticle, by
coverings of wax and air containing hairs; (3) that resinous bodies are
found in the roots and leaves of many of the plants; (4) that there
is a general reduction in the size of the leaves, and that these are
frequently revolute-margined; (5) that palisade tissue is quite uni-
formly developed; (6) that mycorhizal fungi are present in the
roots of most of the plants; (7) that, when compared with the xero-
phytes of dry sand plains (25, 6), they show a similarity in respect
to the reduction in size of the foliage, in the development of external
protective coverings of the sub-aerial parts, and in the presence of
palisade tissues, but are very different in the matter of root develop-
ment and character of root structures.
To account for the peculiarities of the bog vegetation various
theories have been brought forward. KraiMman (28), in accounting
for the xerophilous character of the plants of arctic swamps, which
include several species common to American bogs, lays stress upon
two factors: (1) the low temperature of the .moist substratum, and
(2) the presence of drying winds. The former influences the plants
by decreasing the power of absorption, the latter increases the rate
of transpiration. The plants of such habitats must therefore be
protected against the loss of water by the subaerial parts.
SCHIMPER (44, p. 11) in classifying the natural habitats in which
xerophytes occur mentions among others “peat, bogs, because of the
humous acids in the soil.” On page 18 he says: _
The xerophilous character of the vegetation of peat moors has hitherto been
considered an incomprehensible anomaly, and yet the rich supply of humous
acids in the soil furnishes a condition for its occurrence as cornprehensible as it
is necessary. The presence of Scotch pine-and heather on both dry sand and on
Wet peat is thus not more remarkable. than is that of Ledum palustre, Vaccinium
uliginosum, and other peat-plants on the cold dry soil in the polar zones.
Further (p. 124) the statement occurs that ‘‘on the very acid humus
of moors the vegetation assumes a decidedly. xerophilous character,
because . the. humous,.acids impede the. absorption of water by the
22 BOTANICAL GAZETTE [JANUARY
roots.” However, in describing the arctic vegetation (44, pp- 11,
715), he follows the suggestion of KrHLMAN, a conclusion to which
he had come independently. GANONG (16) also accepts KIHLMAN’S
explanation for the xerophilous nature of the raised-bog flora of
New Brunswick. |
In the study of the structural adaptations of these plants and the
causes of their occurrence in bog areas, several questions arise. Are
these two factors, cold substratum and acidity, efficient causes of
xerophily ? Do they act, in the case of the bogs of this region, with
sufficient strength to cause xerophilous modifications in the plants
there found, or to permit the growth of only such forms as are xero-
philous ?
The last question may be answered from field observations.
They indicate that most low-ground plants grow quite as well on the
bog substratum as on the ordinary swamp soils, and that the swamp
species of this vicinity may all be found at one place or another grow-
ing on bog soils. It would seem that here the bog substratum is no
more efficient as a selective agent than are the swamp soils.
The only cases which have come under my observation in south-
ern Michigan which will throw light upon the question of the effect-
tiveness of the temperatures and acidity in the production of xero-
philous adaptations is in the case of Picea Mariana? and Pinus
Strobus. These two plants both show reduced size of stem and
leaf, in the Oxford bog, when compared with plants growing on the
margin. But to what extent this may be due to sterility of the
bog substratum rather than to temperature and acidity is indeter-
minable at this time.
. EXPERIMENTS.
To answer the question of the efficiency of a cold substratum
and soil acidity to produce xerophily, experiments have been in
progress for approximately two years. The difficulties in the way
of experimentation along these lines are numerous. The means for
controlling soil temperatures in bodies of soil sufficiently large for
experimentation with the larger bog plants are practically beyond
the possibility of a university laboratory. When it is further realized
ons The so-called P. brevifolia Pk. This form is certainly no more deserving of a
distinctive name than is the bog form of the white pine.
1906] TRANSEAU—BOGS OF THE HURON RIVER VALLEY 22
that the experiments should extend over a series of years in the
case of the shrubby forms, the problem becomes still more com-
plicated.
Cold bog water Cold nutrient solution
Warm nutrient solution Warm bog water
Pic. 12.—Average plants from the several cultures of Indian corn, From photo-
graphs. ‘
In order to test the relative effects of humous acids (of the con-
centration found in the bogs of this vicinity) and low substratum
temperatures, experiments were made in the form of water cultures
24 BOTANICAL GAZETTE [JANUARY
and with a peat substratum. All of the bog water used was brought
to the plant house from the First Sister Lake. The acidity of the
water varied from .o005 to .co23 normal acid, as measured by 7 / 100
KOH solution.
WATER CULTURES.—(1) The plants were grown in four-liter battery
jars covered with a plaster of Paris plate, having five one-inch open-
ings for the passage of the plants and one of smaller size for a ther-
mometer. Four such jars were employed in each experiment, two
containing a 0.2 per cent. Knop’s solution, and the others bog water.
One of each was further maintained at a lower temperature. The
cooling was accomplished by passing tap water through 15 feet of
quarter-inch (4.5"xX7™™) glass tubing, arranged in a coil within
the jar, somewhat below the surface of the liquid. The sides and
bottoms of the jars were covered with black paper, and those which
were to be cooled were further surrounded by white paper and
sphagnum. Daily readings of the temperatures of the air, warm-water
solutions and cold-water solutions during the warmest period of the
day were recorded. In this way the maximum differences between
substrata and air were obtained. As these temperatures were not
constant they exaggerate, to a slight degree, the average differences
in temperature. Thus, four conditions were obtained which are
comparable: (1) warm nutrient solution (temperature approximat-
ing that of the air of the plant-house), (2) warm bog solution, (3)
cold nutrient solution, and (4) cold bog solution.
» Fig. 12 shows the results of one of these experiments with corn.
The photograph was taken eighteen days after the experiment was
started. When the cultures were set up, the plumule had developed
to a length of 2 inches (5°™). The air temperatures during the period
of experimentation averaged 18.8° C., that of the warm cultures 18.7°
C., and of the cold cultures 10.8° C.
- It is to be noted that under these conditions the best growth of
the leaves and roots occurred in the bog water. But a reduction of
8° in the substratum temperatures caused a diminution in the devel-
opment of both leaves and roots; the plants in the nutrient solution
and the bog water being equally affected. When all of the plants
had developed five leaves, it was noted that in the case of the cold
cultures the two lower leaves had withered. This experiment was
1906] TRANSEAU—BOGS OF THE HURON RIVER VALLEY 25
repeated with corn, white lupine, and bean under similar conditions,
with similar results. The greater development of roots in the case
of the warm bog water may be due to the presence of a poison in very
minute quantities; but this I have been unable to prove.
(2) A third culture was then made in which five plants of corn
were grown in each of the four water culture conditions, and in
addition in four similar conditions, using a mixture of sphagnum
and peat for the substratum. Wooden boxes 2 feet long, 1 foot
wide and a half foot deep (60 X30 *15°™) were constructed, and two
were lined with galvanized iron. The bottoms of the unlined ones .
were perforated so as to allow of easy drainage. The lined boxes
served for the undrained conditions. Further, in one of the drained
and in one of the undrained boxes, 40 féet (12™) of glass tubing,
bent into coils, the joints: ‘being connected» by rubber tubing,were
arranged so that a constant flow of cold water, for lowering the
temperature, could be maintained. The water level in the undrained
bog substratum was kept just below the surface. The water was
obtained from the bog at First Sister Lake, but occasionally all were
watered with distilled water. The amount added to each box was
practically the same. In order to keep the solutions in the water
culture jars at the same acidity as in the undrained boxes, the water
was siphoned off and transferred once a week. Care was taken in this —
transfer to aerate the water in the boxes as little as possible, while
that of the jars was aerated at irregular intervals by means of a
bulb. There were thus produced eight conditions, in which it was
possible to test the effect of the acidity of the bog water, of aeration
(drainage) of the substratum, and of low temperatures. As a result,
it was found that the growth of roots and leaves was best in the
warm bog water, in the warm nutrient solution, and in the drained
warm peat substratum. Reduction in size of both roots and leaves
occurred in the cold bog and nutrient solutions, and in the drained
cold and undrained warm and cold peat substrata. But the plants
in the undrained cold peat showed the most marked reduction in
size. The conclusion was reached (1) that humous acids (acidity
varying from .coo5 to .0023 normal acid) have no effect upon corn
in the matter of leaf and root development; (2) that low temperature
and lack of aeration of the substratum both cause reduction in size;
26 BOTANICAL GAZETTE [JANUARY
and (3) that when low temperature is combined with poor aeration
the effect is very marked.
This experiment was repeated with peas, and the same result
was obtained, although the effects were not so marked (jig. 13).
The roots in the undrained substrata were killed when they attained
a depth of a half inch (12™™) below the surface.
(3) In order to test the effects of drainage and of low temperature
on bog species, another set of cultures in peat-sphagnum substrata
was made. The apparatus used consisted of two flower-pots and
. two glass dishes aproximately a foot in diameter and three inches
Fic. 13.—Effect of the several conditions upon the development of pea seedlings.
All are average specimens. From photographs.
deep (30X7.5°™). <A flower-pot and a glass dish were kept cool by
passing cold water through fifteen feet of glass tubing arranged in
coils, as in previous experiments. Three species were tested in these
conditions: two-year-old Larix laricina, Rumex acetosella, and
Prunella vulgaris. The first cultures were made in the spring of
1903 with the Rumex and Prunella. The air temperature averaged
about eighteen degrees. The cold substratum was maintained about
ten degrees lower. In the case of Rumex it was found that the
largest leaves were produced in the drained peat-sphagnum substra-
tum. Lack of drainage and low temperature both caused a reduction
in leaf area, and when combined produced leaves which were less
than half as large as those of the drained warm substratum.
The Prunella under the same conditions showed the same results.
: RON RIVER VALLEY 27
1906] TRAN SEAU—BOGS OF THE HU
ual
7
F.
E. Dry sand
Fic. 14.—A, B, C, D, E, camera drawings of leaf sections resulting ia 2 7
in the Pee te sia *135- F, diagrams showing average :
breadth of leaves,
28 BOTANICAL GAZETTE JANUARY
Fifteen plants were grown in each condition. At the end of the
experiment each had produced six to eight mature leaves. The
leaves were measured as to length and breadth. An index was
obtained by multiplying these two numbers together and averaging
for each culture. Following are the indexes of leaf area thus derived:
drained warm substratum 1268.3, drained cold 682.6, undrained
warm 518.5, undrained cold 421.8.
In the spring of 1904 the experiment with Rumex was repeated.
The results correspond with those of the preceding year. The
structure of the leaves, resulting in the several cultures, was investti-
gated, and found to show marked variations (§6). Fig. 14 represents
the cross-sections and average leaf areas produced (seventy-five
leaves being measured in each case). When grown on a warm
drained substratum, the leaves are large, and the cells are exceedingly
loose and turgid. The epidermis is composed of large thin-walled
’ cells, having a thin cuticle outside. The mesophyll consists of a
single layer of palisade and three layers of spongy tissue. No resin
bodies are present. The plants grown in the undrained substratum,
whose temperature was reduced about 8° C. below that of the air,
show marked xerophilous characters. THe leaf is reduced in area,
increased in relative thickness, and the margins become revolute;
the epidermal cells are smaller and outwardly. thick-walled; a well-
marked cuticle is present; the mesophyll is very compact and made
up of two or three layers of well-developed palisade cells and three —
layers of spongy tissue; and in the epidermal cells and those adja-
cent to the bundles there are marked accumulations of resinous
bodies.
For the purpose of comparison, a corresponding set of plants
were grown on sand kept just sufficiently moist to allow the plants
to live. As will be seen in fig. 14, the xerophily is not more marked
than that of the undrained cold bog substratum. Fig. 15 shows the
relative appearance of the plants produced by the different con-
ditions.
In the case of the plants grown in the undrained warm and the
drained cold substrata, these same effects were noticeable, but to a
less marked degree. That, in the case of the undrained cultures,
. these effects are not due to the acidity of the bog water is shown by
Riana. . re * - ba my 2 POOR eS} Cae ee es
Bid dd ies RT
1906] TRANSEAU—BOGS OF THE HURON RIVER VALLEY 29
the fact that plants grown in bog-water cultures develop normally.
The light conditions in the several cultures were the same, direct
sunlight being avoided by a cloth screen. It is evident that in this
case there is no response to strong light in the development of the
palisade tissue (49). It would seem rather to be a response called
forth by a reduced transpiration current (44, p. 7). As to function,
it may aid in the transfer of food materials as suggested by HABER-
LANDT (20, p. 260).
Fic. ee plants showing effect of surrounding “conditions. From
photographs.
This plant proved to be the most plastic of all of the species used
in the experimentation, and was the only one which showed marked
variation in the internal structures. Ecologically the results indicate
(t) that an undrained peat substratum may cause xerophilous struc-
tures, but that the effect is to be correlated with lack of aeration of :
the substratum rather than with the acidity; (2) that the same effect
may be induced by lowering the substratum temperature (the air
temperature remaining the same), and thus impeding the rate of
30 BOTANICAL GAZETTE [JANUARY
root growth and absorption; (3) that a cold undrained bog sub-
stratum is analogous to a dry warm soil in that both produce physio-
logical drought; (4) that resin bodies, which are characteristic of the
bog plants, may be produced by this environment in a plant which
under favorable conditions is without them.
‘Undrained cold
. : ce |
Drained cold Undrained warm
16.—Relative effects of drainage and reduced substratum temperature, on
Fic.
Larix. From photographs.
The seedling tamaracks, ten of which were cultivated in each of
the four conditions just described for the Rumex, also showed con-
siderable variation. Their relative development at the end of forty-
1906] TRANSEAU—BOGS OF THE HURON RIVER VALLEY 31
four days is shown in jig. 16. The leaves of the drained warm sub-
stratum have an average length of 12.6 ™™, of the drained cold 10 ™™,
of the undrained warm 11.4™™, and of the undrained cold Ga,
Internally, the leaves show a reduction in the intercellular spaces and
in the size of the cells in the case of the plants grown on the undrained
cold substratum, when compared with those of the warm drained
condition.
(4) In another series of experiments with plants of Larix four to
five years old practically the same results were obtained. There
were the greatest number and length of leaves and branches produced
in the case of the drained warm substratum. The smallest and
shortest leaves and branches were produced by the undrained cold
substratum.
Experiments with Ledum Groenlandicum, Chamaedaphne caly-
culata, Andromeda Polifolia, Betula pumila, and Oxycoccus macro-
carpus have failed to produce satisfactory results. This is believed
to be due to the shortness of the time under which they were under
cultivation. The plants were brought from the bogs in the late
autumn and placed in cold frames over the winter. About the
beginning of March they were brought into the greenhouse, and
after a few days planted in the warm and cold, drained and undrained
boxes, previously described. They have grown vigorously, but the
differences noticeable may not be correlated with the four conditions.
The cranberry has shown the greatest amount of plasticity, but this
could not in all cases be correlated with the environment. If these
plants can be kept under known conditions for two or more years, it
is probable that they will yield valuable results.
(5) In order to test the effect of mineral soils, and the ability to
withstand the presence of large quantities of calcium and magnesium,
specimens of andromeda, cassandra, and cranberry were grown in
Sandy loam and sand. They were watered daily with tap water.
The cultures were started in the autumn of 1902, and produced
vigorous vegetative shoots during the summer of 1903. They failed
to bloom, however, and although they are growing well at this time
(June 1904), they have again failed to bloom. This may be in part
due to the warm plant-house conditions. The experiment was
originally started to observe the changes in the roots, and in so far
a3 BOTANICAL GAZETTE [JANUARY
¥
have been of value. In a sphagnum substratum all three of the
plants produced hairlike roots which attain a length of 5-7°™. The
roots are commonly several times branched, very little difference in
thickness being shown by the several branches. When grown in
sand the roots are still slender, but the frequency of branching is
enormously increased. Usually the branching occurs just back of
the growing tip. The older root ceases growth as the lateral root
develops. The branch continues for 2-3 ™™, and it also stops growth
with the formation of a second lateral root. The result of this pro-
cess is a zigzag root showing root branches which have been succes-
sively the main root tip. Occasionally several lateral roots develop
and the main axis is divided.
(6) The statement that waters containing lime and other mineral
salts are unfavorable to the growth of sphagnum has gained wide
circulation in ecological literature. Because of the great abundance
of lime and magnesia in the waters of this vicinity, I was led to test
this fact by growing the sphagnum in tap water and solutions of
CaCO,. In one experiment the water in a battery jar was saturated
with CO,, CaCO, was added in excess, and the CO, was again
allowed to pass through the water for thirty minutes. In this solu-
tion sphagnum was placed, and it has been growing vigorously for
three months, although watered daily with water containing over 100
parts of CaCO, to the million. Some of the sphagnum cultures have
been running for ten months, and show no signs of deterioration.
Whether the sphagnum of this vicinity has become accustomed to
the presence of lime, owing to the nature of the soil waters, or whether
sphagnum is generally able to withstand such conditions, remains to
be proved. Since the above experiments were performed, I have
found an account of somewhat similar experiments by WEBER (58),
the results of which are the same. It would seem, therefore, that the
presence or absence of sphagnum is not to be correlated with the
presence or absence of lime.
(7) Among the plants growing in the bogs of this vicinity the fol-
lowing have been found to possess mycorhiza: Larix laricina, Pinus
Strobus, Picea Mariana, Betula lutea, Betula pumila, Oxycoccus
macrocar pus, O. Oxycoccus, Chiogenes his pidula, wi te corym-
bosum, Ledum Groenlandicum, Populus tremuloides.
1906] TRANSEAU—BOGS OF THE HURON RIVER VALLEY 33
In order to get at the conditions which favor or cause the develop-
ment of mycorhiza, cultures of Larix were made in loose sphagnum,
sand, undrained sphagnum, etc. The roots in the many other cul-
tures previously noted were also carefully watched. It has been
found without exception that where the plants were grown under
properly aerated soil conditions, normal roots developed in place of
the mycorhiza. That the acidity of the bog water has nothing to do
with the production of mycorhiza is shown by the fact that in water
cultures of the same acidity as the solution in the undrained peat, the
plants develop normal roots. In the case of roots developed in loose
sphagnum, sand, and moist air, an abundance of root hairs were pro-
duced. The normal roots in Larix have a diameter about three
times that of the mycorhiza, so that when they begin to develop they
appear like white pendants from the dark brown mycorhiza. That
mychoriza will not develop in a well-aerated substratum was further
tested by the following experiment: Two 30°™ test tubes were set
upright, and 8°™ of glass beads were poured into the bottom of
each. Into onea glass tube, at whose end were several small open-
ings, was passed to the bottom. The upper part of the tube was
connected with a gasometer. Upon this foundation of beads, three
plants of Larix were planted in a 5°™ layer of peat in each tube.
The water level in the two tubes was kept just at the surface, bog
water being used throughout. Air was then forced from the gas-
ometer to the bottom of the one tube and allowed to pass slowly
through the beads and peat. When the experiment was started, all
of the plants possessed only mycorhiza. In the course of a week the
aerated plants began to develop normal roots. The experiment was
continued for six weeks. The unaerated plants developed only
mycorhiza, while those which were aerated developed normal roots.®
The growth of mycorhiza is exceedingly slow, and the fungus grows
with the root. The development of the above ground parts cor-
responds to the root development. The plants which produce normal
roots have longer shoots, and longer, thicker leaves.
It seems evident, in the case of Larix at least, that (1) the mycor-
hizas develop only in poorly aerated siento (2) their growth is
8 In the case of a number of the plants of Larix grown in the undrained peat
in previous experiments, one or two normal roots were developed just at the surface
of the substratum
34 BOTANICAL GAZETTE [JANUARY
exceedingly slow, the fungus developing along with the root; (3) the
acidity of the substratum is not a factor in their development; (4) in
a naturally well-aerated soil or in an artifically aerated substratum
normal roots develop; (5) when the roots are not surrounded by
water, root hairs develop abundantly. Mycorhiza therefore appears
to be an abnormal root condition. Whether the fungus is of advan-
tage to the root under these poorly aerated conditions cannot as yet
be stated.
(8) In order to determine whether the zone of tamaracks follows
the shrub zone because of the occasional submergence of the sedge
zone, the following test was made: Ten Larix seedlings averaging
7°™ in height were placed in a crystallizing dish with the roots
imbedded in 2°™ of sphagnum. Over this a layer of bog water 4°™
in depth was maintained for six weeks. The plants grew quite as
well as those in a peat substratum. Stem and root submergence is
therefore not a factor in preventing the growth of seedlings tamarack
in the sedge zone. The liability to submergence in the bogs I have
studied would not extend over nearly so long a period of time.
V. Summary.
The Huron River basin shows three well-marked physiographic
divisions which differ in forest covering and the possibilities for bog
development. These are (1) the region of the Saginaw-Erie inter-
lobate moraine; (2) the Erie morainic belt; and (3) the lake plain.
In discussing the meteorological conditions of a region as affecting
the flora, attention is called to the fact that the significance of the
data is not apparent unless the temperature and rainfall phenomena
are compared with those of the optimum region of dispersal of the
plant societies involved. In the case of the bog plant societies the
temperature of the region under discussion averages several degrees
higher during the summer months than the eastern maritime prov-
inces of Canada (the optimum region of dispersal for the bog plants),
while the rainfall during the same period averages about three-
fourths as much. This is believed to account for the general differ-
ence in character and development of bog societies in the two regions.
Bog and lake basins are here associated with deposits of glacial
drift. The most frequent causes of these basins are (1) the melt-
_
4
|
iS)
1906] TRANSEAU—BOGS OF THE HURON RIVER VALLEY 35
ing of stagnant bodies of ice in old glacial drainage channels after
their abandonment; (2) the differential settling of fluvio-glacial
deposits; and (3) unequal deposition of glacial material in moraines
and till plains.
Marl and peat deposits are commonly associated. The former
are of interest in so far as they aid in the filling of the lake basins.
Both are formed through plant agencies.
Peat deposits may be classified under two general heads: (1) those
connected with glaciation, and (2) those associated with coastal plain
phenomena. In North America the bulk of the deposits come under
the first head. Their geographic distribution approximates that of
the Pleistocene glaciers. Near the southern border the peat areas are
scattered, but they become more nearly continuous and more inde-
pendent of depressions as we go northward. The same effect is
brought about in mountainous regions by increased altitude. In
the tundra, peat accumulates because of the low temperature and in
spite of the scant vegetation. In temperate regions a vigorous vege-
tation and areas of stagnant water render peat accumulation possible.
In the southern coastal plain swamps, peat is formed in stagnant
water because of the luxuriant vegetation and in spite of the high
temperature.
During peat formation two processes are involved: (1) erema-
Causis and (2) putrefaction. The former is essentially an oxidizing
process, brought about in the presence of air by certain fungi and
bacteria. Its products are of direct value as food materials for
plants. Putrefaction is carried on in the absence of oxygen and is
essentially reduction; the organisms involved are anaerobic bacteria,
and the products are of no value to the higher plants as food materials.
The accumulation of peat depends upon the scarcity of oxygen below
the water level, the acidity of the ground water, and the occurrence
of low temperatures.
Peat varies in color beneath the various plant societies, being light
brown in the youngest (bog sedge) and dark brown in the oldest, the
darkest and most thoroughly decayed form being known as “muck.”
As disintregration proceeds it brings about a decrease in water capac-
ity, a decrease in volatile combustible matter, and an increase in
the amount of ash.
36 BOTANICAL GAZETTE JANUARY
The bog as a habitat for plants differs widely from the other plant
habitats of the region in that its substratum has been built by fore-
runners of the present vegetation. Owing to the influence of the
wind in the production of waves, the bogs are largely wanting on the
eastern shores of lakes, and in the case of basins which have been
almost completely filled with peat, the open water lies toward the
eastern margin.
It is well known that bog areas are more liable to late spring
frosts than adjoining uplands. ‘This is due to the topography as it
affects air drainage, and to the low conductivity of the substratum
covering. Under natural conditions it has been found that the areas ~
of cassandra and tamarack dominance are more exposed to late frosts
than other societies.
Observations in bog areas show that the soil temperatures beneath
the several plant societies differ markedly in range. The records
indicate that the areas of bog sedges have temperatures correspond-
ing closely with those of the upland and approximating those of the
atmosphere. The willow-sedge (swamp) and maple-poplar areas
have slightly lower temperatures during early spring. When the
trees leaf out, however, the shade produced causes the maple-poplar
area to have the lowest temperatures recorded. The bog shrub and
tamarack societies show the lowest average temperature throughout
the spring months.
Low soil temperatures retard chemical action, diffusion, solution,
and osmosis, and render the substratum unsuited to soil bacteria.
When coincident with higher air temperatures, plants having a low
transpiration ratio are favored in the competition between species.
In so far as southern Michigan is concerned, the substratum
temperatures prevailing in bog areas do not seem to be adequate to
account for the presence or absence of bog plants or their xerophilous
structures. Experiments suggest, however, that farther north this
factor is of prime importance.
In texture the bog substratum shows every gradation from the
coarse fibrous peat of the bog-sedge zone to the black powdery muck
of cleared land. Bog soils in general do not afford as good a foothold
for trees as do the mineral soils.
Peat is very resistant to the diffusion of mineral salts, hence bog
|
;
:
1906} TRANSEAU —BOGS OF THE HURON RIVER VALLEY 37
areas have a very different soil solution from that of the mineral
soils adjoining. The high water capacity of peat is detrimental to
plants, in so far as it prevents proper aeration of the substratum.
Bog waters have no higher osmotic pressure than ordinary soil
waters.
The absence of sphagnum from local bogs cannot be explained
by the presence of calcium salts, as shown by observation, chemical
analyses, and experiments.
The acidity of local bog water varies from .ooors to .00258
normal acid. The lowest values are found in areas covered by bog
sedges and swamp plants, and they are approximately the same.
The highest occur under the tamaracks. The variations in acidity
are related inversely to the temperature. As shown by experiment,
this is because of increased oxidation at the higher temperatures.
It is suggested that we should find increased acidity as we go north.
There is no apparent relation between color and acidity, except that
light colored waters usually show slight acidity. The acid nature
of the soil solution is a factor in the competition between different
Species for the occupancy of bog areas.
Bog soils are notably deficient in potassium and available nitrogen.
Nitrifying bacteria are prevented from carrying on their normal activ-
ities by the acidity of the soil solution, by the lack of oxygen, and by
the lower temperature of the substratum.
With few exceptions bog plants are light-demanding forms; hence,
in their competition with one another, size and shading ability
are prime factors.
That the conditions in the Huron valley are at present not as favor-
able to the bog plants as to the swamp plants, is shown wherever the
two societies come into competition. This fact must be contrasted with
the situation in the optimum region of the distribution of bog plants,
where the opposite relation has been shown to exist.
An examination of all the physical and chemical data now avail-
able fails to account for the differences in flora of bog and swamp
areas in this region. The most important factor is believed to be
their physiographic history. Where the habitat dates back to Pleis-
tocene times and has remained undisturbed, we find today the bog
flora. Where the habitat is of recent origin or has been recently dis-
38 BOTANICAL GAZETTE [JANUARY
turbed, we find the swamp flora, or mixtures of swamp and bog
species.
The nature of the bog plant societies of the Huron basin is shown
by the description of several local bogs, selected to show both the
local bog flora and the variation in societies, and arranged to present
the genetic changes in a bog flora as_a basin filled by peat accumu-
lation. It is shown that during the early stages of bog development,
bog sedge, bog shrub, and conifer societies follow each other in the
invasion of the basin. These several societies may vary considerably
in composition, but they are closely related and show every gradation
in a definite order of succession. The bog conifers, however, show
no relationship to the surrounding broad-leaved forests of the upland.
On the other hand, where clearing has occurred, swamp sedges,
swamp shrubs, and swamp trees gain the ascendency, and these not
only show an order of succession among themselves, but are genetically
related to the broad-leaved trees of the region. The bog societies
are part of the northeastern conifer forest formation, while the swamp
societies are related to the southeastern broad-leaved forests.
An anatomical study of the bog plants shows that epidermal and
hypodermal tissues are thick-walled, that a heavy cuticle is present,
frequently supplemented by wax and hairs. Resinous bodies are
to be found in the roots and leaves of many of the plants. The leaves
are usually small and revolute-margined. Palisade tissue makes
up a large part of the mesophyll. Mycorhizas are present in most
of the plants. Bog plants resemble the plants of dry sand plains
in reduction of foliage area, in development of protective coverings
for above-ground parts, and in palisade tissues, but differ from the
latter in the matter of root development and root structures.
Experiments indicate that the local bog water itself has no tendency
toward the production of xerophilous modifications. Low soil
temperatures and lack of soil aeration, however, cause a reduction
in the development of the several plant organs. When these two
factors are combined, the effect is very marked.
Experiments with Rumex acetosella are of especial interest in
that nearly all of the characteristics of bog plants may be developed
either by lowering the soil temperature, as compared with the air
temperature, by preventing proper soil aeration, or by growing in
1906] TRANSEAU—BOGS OF THE HURON RIVER VALLEY 39
dry sand. Palisade tissue was developed in the leaves of these plants
in diffuse light, and it is shown that palisade tissue is to be correlated
with physiological drought. An analogy between the bog habitat
and the dry sand habitat is established.
Experiments with Larix indicate that mycorhizas develop only in
poorly aerated substrata; their growth is exceedingly slow; the
acidity of the substratum is not a factor in their development; a
naturally or artificially aerated substratum favors the development
of normal roots, and these roots when not surrounded by water
develop root hairs abundantly. Larix seedlings can withstand
prolonged submergence. When exposed to low substratum tem-
peratures and poorly aerated soil conditions, Larix produces more
xerophilous leaves.
Further field work on the bog plant societies needs to be carried
on in the region extending from Winnipeg to New Brunswick. Data
on the soil and air temperatures, the acidity, the chemical composition
of the soil solution, and the plants associated in bog areas throughout
this region will go far toward solving the problems of the distribution
of bog plants. Experimentation on the production of xerophilous
structures by bog conditions should be continued on a larger scale
than is possible in the ordinary university plant-house.
To Professor V. M. SpaLpineG and Professor F. C. NEWCOMBE,
of the University of Michigan, under whose direction this work was
planned and carried out, I desire to express my sincere thanks both
for helpful suggestions and the facilities of the institution which were
freely placed at my disposal. Many thanks are also due Professor
I. C. Russexz for criticism of the physiographic part of this paper.
I wish to acknowledge the kindness of Mr. FRANK LEVERETT, of
the U. S. Geological Survey, whose intimate knowledge of the glacial
geology of this region has been most helpful to me in the prosecution
of my own field work. To Mrs. N. L. Brirron I am indebted for
the determination of the mosses. Finally I take this opportunity
to express my appreciation of my friend and former instructor, Dr.
H. C. Cowxes, to whose writings and lectures I owe my interest
in ecological botany.
UNIVERSITY OF MICHIGAN.
40
2
>
ut
BOTANICAL GAZETTE [JANUARY
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U. S. Geol. Sub. 22:635. Igor
43. ———, Glaciers of North Aneticn. Ginn & Co. 1gor.
44. ScHIMPER, A. F.-W., Pflanzengeographie auf physiologischer Grundlage.
Gustav Fischer, tac. 1898.
45. SErtey, D. A., The temperature of the soil and surface of the ground.
Monthly Weather Rev. 29:501. 1go1.
46. SHater, N. S., The freshwater morasses of the United States. Ann. Rept.
U. S. Geol. Surv. 10:261. 1
47. SiTensky, F., Ueber die Tsteeon Béhmens. Arch. der Naturw. Landes-
durchforschung von Béhmen 61: 228. 1891.
48. Snyper, H., Report on composition of muskeag soils. Bull. 81, Minn.
Agric. Exper. Sta. 1903.
49. Sraut, E., Ueber den Einfluss des sonnigen oder schattigen Standortes auf
die Ausbildung der Laubblitter. Jenaische Zeitschrift fiir Naturw.
1883: 16.
BOTANICAL GAZETTE [JANUARY
. StocxpripcE, H. E., Rocks and soils. J. Wiley & Sons, New York. 1895.
. Srupart, R. F., The climate of Canada. Scot. Geog. Mag. 14:73. 1898.
~ Tarr, R.5., The A es geography of New York state. Macmillan Co.
New Mark,
~ ravior, F. B:, pea ee of Erie-Huron beaches with outlets and moraines
in southern Michio. Bull. Geol. mer. 8:31. 1897.
Topp, J. E., The moraines of southeast South Dakota and their
attendant evades Bull. 158, U. S. Geol. Surv. 1899
. TRANSEAU, E. N., On the geographic distribution and ecological relations
of the plant societies of northern North America. Bot. Gaz. 36:
401. 190
, The development of palisade tissue and resinous deposits in leaves.
Sejm N. S. 19:866. 1904.
WAGNER, G., Observations on Platygonus compressus LeConte. Journal
Geol. 11:777. 1903.
. WesER, C. A., Ueber die Moore, u. s. w. Jahresbericht der Manner vom
Morgenstern 3:1-23. Review, Bot. Cent. 88:17
WELD, L. H., A peat bog and morainal lake. ak GAZ. 37:39.
1904
- WHEELER, H. J., Results of many experiments on “acid upland soils” are
to be found in the 6th, roth, and 12th Ann. Rept. of the R. I. Agric. Exper.
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go and 95 of the same station.
- Wottny, E., Die Zersetzung der organischen stoffe. Heidelberg. 1897.
NUCLEAR DIVISION IN ZYGNEMA.
MABEL L, MERRIMAN.
(WITH PLATES III AND Iv)
THE species of Zygnema chosen for this investigation possesses
a nucleus unobscured by chromatophores, and hence one in which
division stages can be easily followed. No zygospores were found
in the material, so the species could not be identified with any degree
of accuracy. The number of pyrenoids are normally two, one on
each side of the nucleus. The material was gathered from the same
locality, the margin of a brook, during the months of August and
September of two successive years. The filaments were studied in
a living condition to make sure of the presence of dividing nuclei,
and were then killed in chromacetic acid and the weaker solution of
Flemming for later study. The greater part of them were killed in
the evening, as it was also desired to secure division stages of other
Conjugatae, which grew in great abundance in the locality and have
been reported by investigators as dividing more actively at night.
Of these, three species of Spirogyra and two of Mesocarpus will
furnish the material for a later contribution.
As nearly all the literature upon the cytology of the Conjugatae
relates to forms of Spirogyra, its consideration will be deferred until
the completion of further studies in the nuclear division of the group.
It is hoped then to bring into accord all the observations as to the
character of chromatin and nucleoli.
Filaments of Zygnema treated with the combination stain of
safranin and gentian violet, were found upon examination to have
retained the violet only in the cell sheath, while the nuclear structures
and pyrenoids retained the safranin. Various results were obtained
with those treated with Heidenhain’s haematoxylin in combination
with iron alum and eosin. As the same length of exposure to the
stain did not suffice for Spirogyra and Mesocarpus growing entangled
with the Zygnema, the material was allowed to remain in the staining
fluids for a shorter or longer time. Filaments show pyrenoids stained
black by the haematoxylin, the nuclear structures retaining the
eosin; or the pyrenoids may be stained red by the eosin, and the
43] [Botanical Gazette, vol. 4x
44 BOTANICAL GAZETTE [JANUARY
nuclear structures black by the haematoxylin; or finally both may
appear stained red by eosin. Such differences are shown in the
drawings from the different preparations; the parts shaded in black
represent portions stained by the haematoxylin, as in jig. 33, those in
gray the portions stained by the eosin, as in jig. 13.
Within a quiescent nucleus situated between the two pyrenoids
thus stained, there can be seen a central body stained somewhat
redder or blacker, as the case may be, than the peripheral network
of granules. This network of granules, ordinarily scarcely distin-
guishable from the cytoplasmic reticulum, was found in some cases
to be quite conspicuous.
If an examination is made of a nucleus in process of reconstruc-
tion from the telophase, within the forming membrane can be seen
a conglomerate mass of substance, very evidently non-homogeneous
both in surface view and as seen in outline, figs. 1, 39. Around
this smaller bodies can be seen in the meshes of a delicate network.
The staining capacity of the larger mass and the small bodies varies
in the different preparations; in some instances they are sharply
defined from one another, at other times they retain the same kind
and amount of stain.
It cannot be denied, however, after a careful examination of
stages preceding the appearance of these bodies, that the substances
in both came from the chromosomes of the metaphase. Bearing in
mind, then, that the large mass and the smaller granules have the
same origin, it would hardly seem correct to discriminate between
the two, terming the one nucleolus and the others chromatin granules.
Neither method of staining nor study of their history yields evidence
other than that they are of similar substance, differing only in position ._
and aggregation. It is as if in the revolutions going on within the
cell some of the chromatin granules had been drawn to the center,
there incompletely cohering, while others were left at the periphery.
In describing, then, the quiescent nucleus of Zygnema it seems prefer-
able to say that the larger portion of the chromatin granules cohere
to form a central body analogous in its position to the nucleolus of
higher plants.
The division of the nucleus is presaged by granules collecting in
the region where the cell wall will form. The activity of these vibrat-
19c6] MERRIMAN—NUCLEAR DIVISION IN ZYGNEMA 45
ing granules in the living cell renders the nuclei about to divide easily
distinguishable from the remainder in the filaments. Owing to the
activity of these granules, changes going on within the living nucleus
could not be easily followed, but changes in the form and position
of the nucleus together with those of the pyrenoids were followed
throughout division. Accordingly the history of changes in the
chromatin is all deduced from comparison of dividing nuclei stained
by haematoxylin or safranin as outlined above.
If haematoxylin in combination with iron alum could be considered
as an infallible criterion for distinguishing chromatic from achro-
matic material, and stages could be selected from material stained
by one of the methods only, it would be an easy matter to trace the
history of this central body originating from the chromosomes of the
metaphase. Often, as in fig. 2b, numerous deeply stained bodies
are to be seen lying in the space surrounded by a membrane, with
no trace of chromatin bodies without. In the nucleus represented
in fig. 5, in place of the central body several smaller bodies can be
seen marked off from the eosin-stained bodies by the blackness of
the stain. Passing to fig. 9, where the beginnings of an intranuclear
spindle are manifest, and where there are several more deeply stained
bodies, and then to jig. 12, where six discrete bodies distinctly form
an equatorial plate, the natural conclusion, based wholly upon simi-
lar staining properties, would be that the central mass of chromatin
alone furnishes the chromosomes for the equatorial plate. Such
was the conclusion reached during the first year of this investigation,
but further study of the material shows it to have been premature,
or, if applicable at all, only to a few cases. The conviction that
_ difference in staining of nuclear structures is more often a matter of
manipulation than of chemical reaction, and that difference in the
shade produced by the stain is merely due to the density of the body
and time given for penetration, renders necessary in interpretation a
great degree of caution.
The following account is derived from a comparison of parallel
Stages in all the preparations.
As the nuclei pass from the quiescent to the active state, the cen-
trally lying mass disintegrates into small bodies (figs. 2, 3); at the
same time the granules lying at the periphery increase in size. The
46 BOTANICAL GAZETTE [JANUARY
space within the nucleus becomes gradually clearer (jig. 5), the nuclear
sap probably reinforcing the substance of the granules. As the
result of the disintegration of the central body and the growth of the
other granules, there may be seen lying within the nucleus twenty
or more granules (jigs. 4, 5, 6). In a few cases these bodies may
slightly cohere, but in the majority of cases they lie free. No cases
were found here or in later stages of the formation of a spirem.
In many instances all the bodies within the nucleus retained merely
the eosin stain (fig. 6), and hence were entirely undifferentiated from
each other. In a few cases, like jigs. 5, 7, 8, some of the bodies
retained only the black stain from the hematoxylin. In one instance
(jig. 11), a faintly stained larger body, with one or two smaller ones
of similar shade, can be seen lying within the nuclear space, sur-
rounded by numerous more deeply stained granules. If the other
stages mentioned had not been observed, the latter faintly stained
body might have been interpreted as a nucleolus like those in higher
plants, now in the act of becoming dissolved in the cytoplasm.
Extended comparison, however, of parallel stages justifies the view
that this body is only a portion of the central mass of the quiescent
nucleus, about to undergo still further disintegration into chromo-
somes.
The many chromosomes thus resulting approach one another
(jigs. 6, 9), presenting in many cases an appearance analogous to
the synapsis stage described as occurring in the higher plants. Finally
they become arranged in a circle concentric with the short axis of
the cell. In one case (fig. 10), such an arrangement was observed
before the nuclear membrane became dissolved. Fig. 14 shows this
massing of granules in the equatorial plane after the dissolution of
the nuclear membrane. The chromosomes in this cell were all
stained black, but some were drawn in lighter tint to show that they
were lying in three different planes. Fig. 15 also represents a
similar stage, and fig. 18 one somewhat further advanced. The
chromosomes now appear to be denser than in previous stages, an
interpretation based upon the circumstance that the hematoxylin
stain does not as readily become washed out.
After having formed the ring they appear to be drawn inward,
becoming denser and undergoing a process of fusion. By this draw-
f
'
1906] MERRIMAN—NUCLEAR DIVISION IN ZYGNEMA 47
ing-in process they come to lie in two closely adjoining parallel rows.
As no case of a single row of isolated granules in the same plane was
found, there is no evidence that such double row was produced by
the division of a single row. Fig. 25 represents two rows of chromo-
somes lying in the same plane. In fig. 12 fusion has taken place to
such an extent that only three chromosomes are present in each row.
Many of the chromosomes presented a tetrahedral appearance, -
as in jigs. 16 and 20, thus pointing to the conclusion that the fusion
of the condensing granules may take place in fours. In some cases
the fusion has gone so far as to result in only four groups of tetrads
(figs. 22, 23, 21). Fig. 24, of a more highly magnified group, shows
especially distinctly this grouping of the granules. Careful focusing
on this stage indicated the presence of another underlying group.
As many cases were found of such grouping of the chromosomes in
fours, it does not seem that it could have been purely accidental.
When the maximum amount of fusion and condensation is reached,
the limit apparently varying in different cells, each half of a group
becomes dissociated from its adjoining members and gradually draws
away, as in jig. 27. In process of separation each group becomes
broken up into smaller groups, in the meantime all becoming again
arranged in two rings concentric with the short axis of the cell (fig. 26).
Thus numerous chromosomes are arranged in a circle in stages
preceding and immediately following the stage of the equatorial plate,
in which commonly four to six chromosome groups may be seen.
It seems difficult to believe that six chromosomes (jig. 12) could have
resulted directly from condensation and fusion of thirty or more
chromosomes (fig. 14). A comparison of chromosomes as to size
and staining qualities in the two adjoining cells (figs. 12, 13), drawn
with Abbé camera, would certainly indicate that each chromosome
must suffer a loss of its more liquid substance in the process of being
drawn into the equatorial plate, or that a few must be entirely dis-
solved. Whether all condense to form a few, or whether only a few
are chosen to transmit the chromatin to daughter nuclei, the remain-
der becoming dissolved in the cytoplasm, cannot be stated with
certainty, as the staining process does not solve the problem as to
the fate of the individual granules. When all the preparations are
48 BOTANICAL GAZETTE [JANUARY
examined and not a selected few, there seems to be more evidence of
the first being the true account of events.
It was thought at first that this difference in number of chromosomes
might be due to difference of species, as none of the Zygnema exam-
ined had zygospores, and hence it is possible that two or more species
might have been growing together. The discovery of cells like those
_ in figs. 6, 12, 13, 15, 23, 22, in the same filament is indisputable evi-
dence that in the same individuals the number of chromosomes de-
creases from thirty or more down to six or eight, and then increases to
thirty or more. This change in number occurs in a few moments, as
determined in living cells by the changes in the position of the nucleus.
All the filaments were examined in surface view, so it cannot be
maintained that the number of chromosomes had been increased by
sectioning.
As the rings of chromosomes approach the chromatophores, the
cytoplasm is condensed on the side nearest the chromatophore.
The explanation of this might be that a large part of the cytoplasm
which is not diverted to the region of the formation of the cell plate
was streaming in toward the center, as in jigs. 14-18, while in figs.
20-30 it was streaming out towards the chromatophores; that the
chromosomes are forced together by the inflowing streams and in
the vortex of opposing currents become dissociated. The word
“dissociated” is used in preference to the word “splitting,” as there
appears to be no evidence of splitting and hence of equal distribution
of homogeneous bodies. The chromosomes being heavier than the
cytoplasm, the condensation appears on the side nearest the chro-
matophore (figs. 28, 29). |
It is to be regretted that in the living cells chromosomes could
not be distinguished from actively vibrating granules in the cytoplasm.
Nothing could be discovered which in any way resembled spindle
fibers, although streams of granules and the alternating space of
nuclear activity was easily traced.
The number of chromosomes finally arriving at the chromatophore _
may be fifteen to twenty in each ring, as in fig. 30. The cytoplasm,
being somewhat arrested in its flow by the chromatophore, causes a
change in the position of the chromosomes. The majority, as they
undergo still further dissociation, are drawn to the center, incom-
1906} MERRIMAN—NUCLEAR DIVISION IN ZYGNEMA 49
pletely cohering, while a few appear lying in delicate strands about
them (jig. 31). In some cases all the chromosomes may cohere to
form the central body. The nuclear membrane now emerges from
the condensation of cytoplasm (fig. 32). As the chromosomes are
now shut off from the influence of currents in the cytoplasm they
generally remain unchanged in position, fusing either to form one
mass (fig. 33), or three or more smaller masses (fig. 34), or rarely
(jig. 35) all the chromatic material may be diffused in the nuclear
plasm, forming numerous more or less tetrahedral granules.
It is to be noted that not until the chromatic rings have separated
and have approached the chromatophores do the pyrenoids ordinarily
show any evidence of division. Tviis observation was easily con-
firmed from the study of living cells. Fig. 29 represents the only
one seen, out of many filaments examined, in which the pyrenoids
divided before the formation of the nuclear membrane. As the
newly divided nuclei approach their respective chromatophores,
one or both plastids begins to show a constriction. This deepens
until when the nuclei come to lie directly over, only a narrow band
of less dense substance resembling linin connects the two daughter
pyrenoids (figs. 37, 38). This becomes gradually reduced until it
appears only as a thread (fig. 39). Later the nucleus sinks down
and the separation is complete. The constriction of the plastids
forming the center of the two pyrenoids takes place synchronously,
as is the case with the stages in the daughter nuclei. One instance
only was observed in which one plastid suffered division when other
plastids had just begun to elongate (jig. 32).
Although division of the pyrenoid may be influenced by division
of the nucleus, that it is not wholly dependent upon it was demon-
strated by leaving actively dividing filaments of Zygnema for one
hour in a watch crystal containing 10°° of water to which two drops
of chloroform were added. There were but few visible signs of plas-
molysis in filaments killed and stained as in other material, but while
a majority of the nuclei had ceased to divide, a majority of the pyre-
noids were dividing as in normal filaments. That this division was
not merely fragmentation was shown by sequence of stages and the
presence of the band connecting the plastids. Fragmentation of
the pyrenoids took place in filaments in stronger solutions of water
Mo. Got.Garacn
1906
50 BOTANICAL GAZETTE [JANUARY
and chloroform in which plasmolysis occurred to a much greater
extent.
Hence, cytoplasmic streams, nuclear structures, chromatophores,
and pyrenoids take an active part in the division of cells in Zygnema.
The streams of granules, collecting where the cell plate is to form,
marks the beginning; the nuclear changes then proceed, followed
by division in chromatophores and pyrenoids, while all are correlative
with the formation of the cell plate.
It cannot then be said that division of the nucleus, the chromato-
phores, and the pyrenoids are synchronous. Rather is it true that
the center of activities of the cell shifts, and with this shifting division
of the bodies lying in the vicinity occurs. As regards the nuclear
structures in Zygnema it is apparent that there are no bodies analo-
gous to the nucleoli found in the higher plants. A large portion of
the chromatin, or in a few cases possibly all, fuses in the anaphase to
form one or more bodies corresponding in appearance and position
to that of nucleoli of higher plants. Instead of waste products of
chromatin condensing to form one or more bodies in the nucleus,
the waste products are not separated from the chromosomes, but
retained in them until after the nuclear membrane disappears in the
next division. The substances which make up chromosomes and
nuclear waste products, if such we may rightly regard the nucleoli
of higher plants to be, are in Zygnema morphologically indistinguish-
ble.
The history of chromatin before the formation of the equatorial
plate may be summarized as consisting of growth, association, and
condensation of chromatin bodies in groups. These groups may
be partially coherent, but in no case forma spireme. After equatorial
plate formation, dissociation into groups follows, continuing until
the chromosomes reach the chromatophores.
Although the term chromosome has been used in this account,
researches as yet incomplete make it exceedingly doubtful whether
the chromatin bodies in any of the Conjugatae are to be rega
as at all homologous with chromosomes of higher plants. If we
restrict the term chromosomes to segments of the tubular spirem,*
*See MERRIMAN, Vegetative cell division in Allium, Bot. GazETTE 37:178-207-
pls. 11-13. March 1904.
1906] MERRIMAN—NUCLEAR DIVISION IN ZYGNEMA 51
then the chromatin bodies seen in jigs. 14 and 15 of Zygnema cells
are homologous not with the chromosomes of Allium but with the
granules seen in the earliest stage of the spirem, while groups in
figs. 16 to 23 are directly comparable with the groups or rings of
tetrads, which in Allium fused to form the tubular chromosomes.
Zygnema possesses a mechanism of nuclear division less elabo-
rated than that of the higher plants, inasmuch as dissociation of
chromatin bodies occurs immediately after their association into
primary groups without the intervention of a spirem. From this
point of view appearances observed in Zygnema support the inter-
pretation suggested in my account of nuclear division in Allium,
namely, that the chromosomes are formed by fusion of bodies in
groups, and that when a longitudinal splitting appears it is not to
be considered a true splitting of a homogeneous substance but rather
a dissociation of bodies which from the first were discrete.
If this be true, then doubts may reasonably be entertained as to
the validity of the conception held by Roux, and successively by
many other investigators, that the complex apparatus for indirect
division of the nucleus exists for the purpose of enabling each
chromatin body to furnish its quota to the daughter nuclei.
The essential feature of indirect division, and therein its advan-
tage over direct division, appears to be the dissolution of the nuclear
membrane. Thus is made possible a free interchange of nuclear
and cytoplasmic substances and a renewal of the vitality of the cell.
Zygnema, then, may be considered as furnishing additional evi-
dence of interchangeability of nucleoli and chromatin bodies, of
variability in their number, and negatively as furnishing no
evidence that equal distribution of chromatin is effected by either
transverse or longitudinal splitting of homogeneous bodies. Nuclear
structures, cytoplasm, pyrenoids, and chromatophores are trans-
ferred in equal amounts to the daughter nuclei and by a process
differing not fundamentally in the result from that which would
_ have been attained by direct division.
NORTHFIELD, Mass. -
52 : BOTANICAL GAZETTE [JANUARY
EXPLANATION OF PLATES III AND IV.
The figures were drawn with the aid of an Abbé camera.
PLATE III.
Fic. 1. Daughter nucleus from a cell -where the cell plate is not yet com-
pleted. The nuclear structures in this cell retained the eosin stain, the pyrenoids
black from haematoxylin. x 1750.
Fic. 2a. Nucleus preparing to divide, showing growth of bodies in the periph-
eral network before breaking up of the central body. Pyrenoids and nuclear
structures in this cell retained only the eosin stain. 1750.
uclear material stained black by the haematoxylin, all the chro-
matic material being apparently condensed in the ee occupied by the central
body. X 1750.
. Fic. 3. Nucleus showing the breaking up of chromatin body and increase
in size of the peripheral bodies. The pyrenoids retained the eosin stain; all the
nuclear structures are stained black, several of them somewhat darker than the
others. X 1750.
IG. 4. Nucleus showing the beginning of the massing of the chromosomes,
the nuclear membrane as yet undissolved, the granules in the region of the cell
plate formation being conspicuous. Chromosomes black, pyrenoids red
Fic. 5. Later stage, showing the clearing of the nuclear interior, recalling the
synapsis stage described in higher plants. Pyrenoids red, several chromosomes
black, remainder red. X 1
Fic. 6. Similar stage, very frequent; chromosomes numerous, massed
together, all stained red. 1750.
Fics. 7, 8. Similar stages where there is no massing of the chromosomes.
In 7, a chromosomes were stained black, others red. In 8 those stained black
are grouped in one corner of the nucleus, those red are scattered. 1750.
Fic 9. A stage where distinct lines of granules connect chromosomes with
nuclear membrane. Four chromosomes black, others red. X 1750.
Fic. to. A rare stage with numerous chromosomes arranged in circle within
the sitenk before the nuclear membrane becomes dissolved. All chromosomes
lack. X 1750.
_ Fic. 11. Another rare stage; nuclear membrane dissolved, remains of central
body still in the cytoplasm, retaining a lighter eosin stain than the other chro-
mosomes. X 1750.
Fics. 12, 13. Two adjoining cells in same filament showing disparity in
size and number of chromosomes. Pyrenoids red, in fig. 12 chromosomes stained
sere by haematoxylin; in fig. 13 nuclear structures stained red. The line
of granules marking the region of cell plate formation shown in both figures.
x1
Fic. 14. All chromosomes black, but some drawn lighter to indicate that
they were lying in three different planes. x 2440.
Fic. 15. Chromosomes black, showing indefinite arrangement as hey are
being drawn to the center. 2440.
ta
Fa ea aalicialtn ani Y ee oe
4 os
poems
. +s
PLATE. Tif
Fa
——e
Bes
MERRIMAN on ZYGNEMA
BOTANICAL GAZETTE, XLI
E.IV
PLAT.
BOTANICAL GAZETTE, XLI
MERRIMAN on ZYGNEMA
ip
1906] MERRIMAN—NUCLEAR DIVISION IN ZYGNEMA 53
Fics. 16, 17, 19, 20, 21, 22, 23 show successive stages in condensation of
chromosomes. Chromosomes all black. X 1750.
Fics. 18, 24, 25, 26. Chromosomes and pyrenoids black. X 2440.
Fics. 27, 28. Chromosomes becoming dissociated into smaller groups.
Chromosomes black. x 1750.
An ‘unusual case of division of-pyrenoids before formation of
FIG. 29
Chromosomes black, both in central mass
membranes of daughter nuclei.
and in the periphery, pyrenoids red. X 1750.
Fics. 30, 31, 32. Chromosomes black, pyrenoids red. 1750.
Fics. 33. Pyrenoids and central body of nucleus black, peripheral bodies
red. X1750.
Fic. 34. Pyrenoids red, all the nuclear bodies black. < 1750.
Fic. 35. Nuclear bodies red, pyrenoids black. 1750.
Fics. 36, 37, 38. All nuclear bodies red, pyrenoids black. 1
. Nuclear bodies red, pyrenoids black, showing vestiges of connect-
ing substance. X 1750.
EFFECT OF CERTAIN SOLIDS UPON THE GROWTH OF
SEEDLINGS IN WATER CULTURES.'
J. F. BREAZEALE.
(WITH FOUR FIGURES)
In certain investigations on the growth of wheat seedlings in
aqueous extracts of soil, it was observed that the growth of these
plants was greatly accelerated by the presence in the medium of |
undissolved calcium carbonate. That the observed acceleration was
not due to an increase in dissolved calcium was apparent from the
fact that the presence of other slightly soluble salts of this element
failed to produce any response. It appeared possible that the effect
of calcium carbonate might be due to its taking up some injurious
substance present in the extract. This was suggested by NAGELI’s
well-known discovery? that water, which is toxic to algae because of
minute traces of metals, can be improved by placing in it such insoluble
bodies as graphite, paraffin shavings, or torn filter paper. It was
determined to try other slightly soluble compounds which might
remove from solution small amounts of solutes, either by chemical
action or mechanically. The results of this investigation make up
the present paper.
The Russian variety of wheat known as “Chul,” obtained from
Arizona, was used in most of these experiments. The seedlings were
germinated in sand and then grown in water cultures in large-mouthed
black bottles of about 60°° capacity. They were fixed in cork stop-
pers, four in a bottle, in the manner described by WHITNEY and
CaMERON: for cultures of this kind, so that the roots were submerged
in the solution while the seeds were just above its surface. The
solutions were always aerated by violent and repeated shaking before
the cultures were started. During the growth of the plants the bottles
were weighed in groups of three at intervals of three or four days,
and the water lost was replaced with distilled water. The manner
I Published by permission of the Secretary of Agricu
2 Nageli, C. von, Ueber ip SPL E Sitar ees in lebenden Zellen.
Denkschr. Schweiz - Naturforsch. segs 33:2:
3 Wuitney, M., and Cameron, F. K. geese in soil fertility. U. S. Dept
Agric., fina of Seite Bull. sa sand
Botanical Gazette, vol. 41] : [54
Tas
1906] BREAZEALE—EFFECT OF SOLIDS UPON GROWTH 55
of fixing the seedlings practically prevents water loss, except through
the transpiration of the plants. The work of Lrvincston? indicates
that total loss by transpiration for a period of two or three weeks
furnishes a fair criterion for comparison of the growth of different
cultures of wheat grown in this manner. The transpiration figures
are used in this way in the experiments here given. The work was
carried on in a greenhouse with a temperature of 15 to 25° C.
For Experiments I to III a soil extract from poor Leonardtown
loam, collected near Leonardtown, Md., was used. It was prepared
by stirring the soil for three minutes with water in amount equal to
twice its air dry weight, allowing it to stand twenty minutes, and then
filtering through a clean Pasteur-Chamberland filter tube, in the man-
ner described by WHITNEY and CAMERON.5
In Experiment I the solids used were calcium carbonate, tri-
calcium phosphate, ferric hydrate and aluminum hydrate. Ferric
hydrate was prepared by precipitation from the chlorid with ammonia,
followed by thorough washing with hot water. It was transferred
moist to the culture media. Aluminum hydrate was prepared in a
similar way from the sulfate. Data for this and the three following
experiments are given in Table J. The percentage increase in transpi-
ration for each solid is computed by considering the transpiration
from the untreated extract as unity.
All of the solids accelerated growth, as is shown by the transpira-
tion figures. But in the case of the ferric hydrate the root growth
was accelerated to a much greater extent than that of the tops. The
roots of the culture with this substance were much longer than those
of the other cultures. It is evident here that root growth was acceler-
ated without a corresponding increase in transpiraticn.
Experiment II comprised, besides calcium carbonate and ferric
hydrate, carbon black (prepared from burning petroleum, and
thoroughly washed), magnesium carbonate, and barium carbonate.
The small amounts of water transpired are due to cloudy weather.
The plants of this series are shown in fig. 1, the numbers in the
figure corresponding to the culture numbers given in parentheses
4 Lrvincston, B. E., Relation of transpiration to growthin wheat. Bort. GAZETTE
40:178~-195. 1905.
s Witney, M., and Cameron, F. K., The chemistry of the soil as related to
crop production. U.S. Dept. Agric., Bureau of Soils, Bull. 22:16 ff. 1903.
56 BOTANICAL GAZETTE [JANUARY
in the table. It will be noticed that carbon black shows the same
tendency to produce abnormal root growth as does ferric hydrate, but
to a less marked degree.
Experiment III included very finely pulverized quartz flour, as
well as ferric hydrate and carbon black. The two last named bodies
showed here the same abnormal acceleration of root growth as was
previously observed, but quartz flour, although it improved the
general growth of the plants, produced no such effect.
iG. 1.—Experiment II; 24 wheat plants grown 19 days. 1, Extract of Leonard-
town foals 2, the same with calcium carbonate; 3, the same with ferric hydrate;
4, the same with carbon black; 5, the same with magnesium carbonate; 6, the same
with barium carbonate.
The experiment with ferric hydrate and carbon black has been
repeated many times with extract of Leonardtown loam, as well as
that of other soils, and always with the same result. In some cases
acceleration of root growth is more marked with carbon black than
with ferric hydrate, but usually the reverse is true.
Experiment IV was carried out with an aqueous extract, prepared
as above, from Miami silt loam collected at the Rhode Island Experi-
ment Staticn, at Kingston, R. I. This soil had been in hoed crops
1906] BREAZEALE—EFFECT OF SOLIDS UPON GROWTH 57
for ten years without fertilizer, and was acid to litmus paper. Te
make absolutely sure that the effect of carbon black was not due to
any substance added with it, the distilled water for the soil extract
was shaken with the solid carbon black and filtered through a
Fic. 2.—Experiment IV; 36 wheat plants grown 13 days. 1, extract of Miami
silt loam; 2, the same with ferric hydrate; 3, the same with carbon black; 4, the
Same with ferric hydrate, carbon black, and calcium carbonate.
Pasteur-Chamberland tube before being used. Ferric hydrate, car-
bon black, and a mixture of these two bodies, together with calcium
carbonate, were used in this case. The plants are shown in jig. 2.
From this experiment and others of similar nature it seems clear
58 BOTANICAL GAZETTE [JANUARY
that the effect of the carbon black is not due to any stimulating sub-
stance which it carries into the medium. Other experiments have
shown that no acceleration of growth is obtained with the addition
TABLE I
DATA FOR EXPERIMENTS I TO Iv.
ExPerIMENT I | Expermwent II | Experiment III) Experment IV
24 PLANTS 24 PLANTS PLANTS a
GROWN I9 DAYS | GROWN 19 DAYS | GROWN 17 DAYS | GROWN 13 DAYS
MEDIUM
Total
Total Per | Total Per Bit Per Total Per
trans- cent. trans- cent. cent. trans- cent.
piration| increase| piration| increase dav er z increase| piration| increase
cultur’s)
anresire Bon extract:.% |. 84:1. cc's ;. }{2) 33 tne 51 Preis,
o.+tri-calc. phosphate..} 182 | 18.1 i tele ieee : + ates spe tans
Decicaiiis carbonate..} 191 | 24.0 (2) 8 TAG Al hes corns eee
Do.+ ferric hydrate. ..... 184 | 19.5 Bae 233.3 | 187 |266.6 |(2) 176} 158.8
Do. +-sitminwin Wydiate..6 2 487: eta ian be | ek Pe op eee ee
’ Do.+carbon black ...... ..ee |... | (4) 69/109.0 | 154 [201.9 | (3)139] 104-4
Do.+magnes. carbonate..| .... | ....-|(5) 73|121.2 ced i Baers ey ones er
Do.+ barium carbonate...| .... | .... | (6) 78/136.3 aie wee
Do.+ quartz flour........ pa ee ear es heel Sete 87 Fors
Do.+ ferric hydrate, car-
crt and calcium
etic Ap a ors ae See Wee Py crag cre tk mr en deren es bP pee
of iron salts to the extract of poor Leonardtown loam. Therefore,
a slight increase in dissolved iron cannot be the cause of the accelera-
tion noted in the case of this hydrate. It seemed probable that ferric
hydrate and carbon black had their effect through an active removal,
perhaps by mere adsorption, of some injurious substance occurring
in the culture medium. Such a substance might have been in the
soil extract originally, or might be produced by the plant roots, or
both suppositions might be true together. The third alternative
proved to be the correct one.
To obtain evidence in this regard, Experiment V was carried out.
Four different soil extracts were shaken with carbon black, filtered,
and then used as culture media, comparison being made with controls
in untreated extracts. The four soils-were of two types, a good and
a poor soil of the Cecil clay type, and a good and a poor of the Leonard-
town loam type, the former from Statesville, N. C., and the latter
from Leonardtown, Md. Chemical analysis both of the aqueous
extracts and of the solution obtained by digestion with hydrochloric
1906] BREAZEALE—EFFECT OF SOLIDS UPON GROWTH 59
acid, fails to show any material difference between the good and the
poor varieties of these soils, although they are agriculturally quite
different. These cultures were grown under the direction of Mr.
F. D. GARDNER, in charge of the Division of Soil Management, of
this Bureau. They consisted of forty-eight plants and were grown for
1, extract of Cecil
.—Experiment V; 48 wheat plants grown 15 days.
clay, ey 2, the same, filtered from carbon black; 3, extract of Cecil clay, poor;
4, the same, filtered from carbon blac
fifteen days. The results are given in Table II; percentage increases
are given for each treated extract compared with the same extract
untreated considered as unity.
It will be seen from this table that the extract of good Cecil clay
60 BOTANICAL GAZETTE [JANUARY
EI.
DATA FOR EXPERIMENT V.
Culture no. Meatum ao) aos
on ek er : aapedel st = mi sie POO cr. ne ote ewe ASG ian a |! —codnape
ee Mise sachs wean pete Do. ROC oe tee ae 471 = 200
Bcad mops tae Extract 3 Cet clay, POOR sx ccsta vec P srs WOT scan Wy eed ieees
AD cess ae ney th Sica] Oe UE DORN LEE OTOO 5. ooo. ssa Jonnie oh aces Sesh maee 477 “ize?
[Ser eS eee Extract of Leonardtown loam, good.. a! ee eee ee
irae, ogee Ceara SN TRMOTEE S Scw re emo ope eee 407 +13.4
ype eres Ea ract of ie te loam, poor.. DRL ea «tte average
Buea sae Do carbon filtered si. i465 sien ees 349 + 28.7
was not improved by treatment with carbon where the latter is filtered
out. All the others were im-
proved, although the improve-
ment in the case of good Leon-
ardtown loam was not as marked
as that in case of the poor. The
leaf development and _ general
appearance of the plants were
essentially in proportion to their
transpiration. The roots showed
__ the same acceleration with carbon
~ treatment and filtering that had
been observed in experiments in
which the solid was left in the
solution. Fig. 3 shows the roots
of cultures nos. 1, 2, 3 and 4.
es
part at least ‘of the injurious
matter which is removed by car-
bon black is in the soil extract
at the start.
: A number of experiments were
: glee pede igi ee grown carried out, using water twice
Ae ee be ed water; 2,thesame, redistilled, first from potassium
dichromate and sulfuric acid, then
from alkaline eens permanganate, both in glass boilers, con-
1906] BREAZEALE—EFFECT OF SOLIDS UPON GROWTH 61
densation being carried on in a platinum tube. Upon this water the
effects of carbon black and ferric hydrate were tested, the solids
remaining in the water during the growth of the plants. The former
gives little or no increase in transpiration, the latter a moderate
increase, but both solids produce’a marked acceleration in root growth.
Twelve seedlings grown fifteen days in redistilled water with and
without ferric hydrate are shown in fig..4. No. 1 shows those from
the untreated water, no. 2 those with the solid. The water here used
probably did not contain injurious substances, and therefore the effect
of the solids is most probably due to the removal of some injurious
exudation arising from the plants. Further, distilled water in which
plants have been grown for a number of days is found to give less
growth upon replanting than does unused water, and the injurious
effect of the used water is corrected by shaking with carbon black or
ferric hydrate and filtering off the solid. Thus it seems that wheat
seedlings do give off bodies from their roots which are toxic to them-
selves.
When the work so far recorded was practically completed, the
appearance of TRUE and OGLEVEE’s paper® on somewhat similar
experiments made it seem advisable to withhold publication until
some further tests suggested by that paper could be made. These
authors find that by the introduction of sand, filter paper, parafiin,
or potato starch into solutions cf copper sulfate in which seedlings
of Lupinus albus are growing, the toxicity of the solute is remarkably
decreased. By this means the killing concentration of the salt may
_ be effectively reduced, according to the amount of the insoluble body
present, either to a stimulating concentration or to one in which the
physiological effect is not apparent. They. reasoned that the solid
absorbed the salt from the solution and in this way produced an effect
closely paralleling that of simple dilution.
It was determined to test the effect of solids in solutions of sulfuric
acid. Maize was here used instead of wheat. First, the strength
of this acid necessary to prevent the growth of maize seedlings was
6 True, R. H. and Octever, C. C., The effect of the presence of insoluble sub-
stances on the toxic action of poisons. Science N. S. 19:421-424... 1904.
Bot. GAZETTE 39:1-21. 1905. In this connection see also DANDENO, J. B., The
relation of mass action and physical affinity to toxicity, etc. Am. Jour. Sci. 17: 437-
458. 1904.
62 BOTANICAL GAZETTE [JANUARY
determined as lying between 2/2750 and /3250.7. Then a series of
varying concentrations of this acid on either side of the toxic limit
was carried out, placing clear sand, quartz flour, filter paper, and
paraffin shavings in the solutions. In no case was the apparent death
limit modified by the presence of these substances. The death limit
was also determined for a solution of sulfuric acid saturated with
calcium sulfate, and for the same solution with an excess of calcium
sulfate, but the solid again had no apparent effect.
With copper sulfate solutions carbon black was found to decrease
the toxic effect, just as the authors above cited found to be true for
the solids with which they worked. In order to bring out the effect
of the carbon black, copper sulfate in the proportions of one and five
parts per million of copper was added to the nutrient solution above
described; a portion of the solution thus prepared was shaken with
carbon black and then filtered, and wheat seedlings were grown in
the treated and untreated solution. This series constitutes Experi-
ment VI. Twenty-four plants composed a culture and the experiment
lasted twelve days. The results are given in Table III.
‘TABLE: Tf.
DATA FOR EXPERIMENT VI.
Culture no. Medium — eff eign 2 aoe
Be vp te wee Nutrient solution........... 220-2
PEGs Lees Do.+1 p. an vas Tenaya ee 67.
Lege seen Decks pms Cu. Si... 41
per emner sey eater As 2, a pdt Siig TE 140.2
sl i a ce toate As 3, but carbon-treated. . T7875
The growth of the plants was proportional to their transpiration.
It is evident that the carbon removed sufficient copper to render
carbon-treated solutions much less toxic than the untreated ones.
This is a direct corroboration of the work of TRUE and OGLEVEE,
but with another solid, and in this experiment the filtering out of the
solid removes any possibility of its having any effect directly upon
the roots. The explanation of these authors seems to be correct,
as far as copper sulfate is concerned. The failure in the present
7 This or aeromine had been made previously, but was repeated for the present
work. See Cameron, F. K., and BRreazEAate, J. F., Toxic action of acids and salts
on seedlings. wee aie aia: 8:1-13. 1904.
i
1906] BREAZEALE—EFFECT OF SOLIDS UPON GROWTH 63
instance to get the same effect with sulfuric acid may be due to the
fact that to be injurious this substance must be at a much higher con-
centration than is needed in case of the copper salt, while the relative
amount of solute absorbed by solids is much greater in dilute solutions
than in more concentrated: ones. TRruE and OGLEVEE suggest this
explanation for their failure to get marked improvement by the use
of sand in solutions of phenol and resorcinol.
On account of the presence of toxic substances in distilled water
as ordinarily prepared, from copper boiler and tin condenser, most
workers with toxicity problems have used water redistilled in glass.
In these laboratories the distilled water is often quite toxic to wheat
seedlings, but its injurious effect is prevented if it is first shaken with
carbon black or ferric hydrate and the solid filtered off. It is found
that water so treated produces as good a growth of seedlings as does
the most carefully prepared redistilled water.
It appears from these experiments (1) that extracts of certain soils
are toxic to wheat seedlings in water culture, and that this toxicity is
removed wholly or in part by carbon black, calcium carbonate, ferric
hydrate, and other solids; (2) that the toxic substances of ordinary
distilled water may be removed by ferric hydrate or carbon black;
(3) that the roots of wheat seedlings give off substances which are
toxic to themselves and that these substances can be made inactive
by the presence of the last named solids in the culture medium; (4)
that the presence of ferric hydrate and carbon black in the solution
seemingly accelerates to a marked degree the development of roots,
causing them to surpass the tops in growth.
The work here reported was done chiefly in the laboratories of
the U. S. Department of Agriculture, Bureau of Soils, Washington,
D.C. It was finished at the Rhode Island Agricultural Experiment
Station, Kingston, R. I. I am indebted to Dr. F. K. CAMERON, and
to Dr. B. E. Lrvrncston, of the Bureau of Soils, for much valuable
suggestion and advice.
U. S. DEPARTMENT OF AGRICULTURE,
Bureau of Soils,
Washington, D. C.
PRrehe ek AR LICLES.
NOTES ON NORTH AMERICAN GRASSES. V.
SOME TRINIUS PANICUM TYPES.
BELOw are given a few notes upon certain species of Panicum described
by Trinius from American material. The herbarium of Trrntus, the
father of modern agrostology, is deposited in the St. Petersburg Academy
of Sciences. The curator of the herbarium, Dr. D. I. Lrrwrnow, very
kindly sent, for deposit in the National Herbarium, portions of the speci-
mens of the species mentioned in this article. It will be observed that
three of the seven species had been described by earlier authors, while four
have been recently described as new species. It must be said, however,
that it would be impossible to identify any of these four species from the
description alone. This is true of many of the earlier descriptions of the
group Dichotoma of the genus Panicum, and on account of the difficulty
of consulting the types of these species, it is in a measure excusable, on the
ground of convenience, to describe unidentified species as new, without
determining the application of the older names. Such painstaking and
laborious comparison should not, however, be shirked by a monographer.
P. CHAMAELONCHE Trin. 1826, Gram. Pan. 242.
are et minus. Panicula (semipollicari) lucidissima; Spiculis pusillis,
obovato-oblongis, obtusiusculis, glabris; pedicellis hispidulis; Gluma inferiore
flosculis triplo breviori 1-, superiore eosdem aequante 5-nervi; Hermaphrodito
elliptico, mucronulato, laevi, neutrum aequante.
V. spp. Am. bor. (TRATTINICK, ex. coll. Enslini).
Plantula caespitosa, stricta, culmo tenuissimo, ramuloso. Folia lanceolata
1. lineari-lanceolata, strictiuscula, semipollicaria, glabra; culmea et juniora
subinvoluta, acicularia, breviora. Panicula pauciflora, tenera, prodit e summa
vagina elongata, cujus folium eandem fere aequat. Flosculus neuter univalvis ?
Hermaphroditus albescens.
Label accompanying type specimen: ‘‘In Am. bor. s. dat. sine nom.
ex. hb’° Enslini cl. Trattinick.”
Specimen =P. Baldwinii Nutt. 1898. Scribn. Div. Agros. Bull. 11:43-
P. pemissuM Trin. Spec. Gram. 3:319. oe
The type specimen is from Rio Janeiro. I have not seen this species
from North America. It is mentioned here because the name occurs
occasionally in the literature of North American grasses, and has been
doubtfully applied to certain of our species.
Botanical Gazette, vol 41] [64
1906} BRIEFER ARTICLES 65
P. Ensiint Trin. 1826, Gram. Pan. 2
Pedale. Panicula (subdigitali) ude lucida; Spiculis subultra-
linealibus, ellipticis, acutiusculis, pilosulis: pedicellis glabris; Gluma inferiore
flosculis sub 4-plo breviori 1-, superiore eosdem vix ae 7-nervi; Her-
maphrodito oblongo, acutiusculo, laevi, neutrum fere aequa
An Panicum tenue Miihlenb. (quod Pan. liton ne Mant 2. 'p. 250)
quaerit ill. N. AB Esenp. in litt.
V. spp. Am. bor. (TRATTINICK, e collect. Enslini.)
Culmus basi ramosus, ad paniculam usque vaginatus: vaginis arctis, fissura
pubescentibus. Folia glabra: radicalia lanceolata-oblonga 1. ovata, sesqui-
pollicaria, lineas 4-5 lata; superiora lineari-lanceolata, duplolongiora, patentia.
Panicula e summa vagina prodit radiis subcapillaribus, parum patulis. Gluma
inferior epilis, superior mucronulata cum flosculi neutrius valvula inferiore pilis
adspersa. Hermaphroditus albescens.
Label accompanying type specimen: “(an Pan, tenue Muhlb. quaerit
NEEs AB Es.) ab Enslino in Am. bor. ]. dt. sine nom. cf. Trattinick Wiennae
1820.”
Specimen =P. equilaterale Scribn. 1898, Div. Agros. Bull. 11:42.
Characterized by having the spikelets of P. commutatum but the leaves
elongated and widely spreading.
P. FLORIDANUM Trin. 1835, Mem. Acad. Petersb. VI. 37: 248.
The type specimen from Georgia is Paspalum racemosum Nutt.=P.
bifidum (A. Bertol.) Nash, as has been generally recognized.
P. JEJUNUM Trin. 1836. Bull. Acad. Petersb. 1:76.
The type specimen from Louisiana sent by Hooker in 1835 is Panicum
hians Ell. =Steinchisma hians (Ell.) Nash.
P. LANCEARIUM Trin. 1822, Clavis Agros. 234.
1179. Gr. miliaceum americanum, minus, panicula parva. Pluk, Phyt. p.
176. Tab. 92 f. 6.
Mor. p. 197. no. 15. Panicum lancearium m. (de quo alio loco).
Label accompanying specimen: ‘“Plukn. Tb. 92 {. 6.2? In Am. bor.
ab Enslino I. dt. cl. Trattinick.”
Specimen=P. Nashianum Scribn. 1897, Div. Agros. Bull. 7:79.
The specimen matches Curtiss 4029 from Florida, the first specimen
cited in the original description of P. Nashianum. Both have glabrous
spikelets. The second specimen cited by Scripner, Nash 466, from
Florida (the type on account of the specific name), has pubescent spikelets.
Since Trinius gives a binomial to a plant described by PLUKENET and
by Morrison under a polynomial designation, PLUKENET’s plant is the
type.
In Kew Index P. ancearium is cited as Agrost. Bras. 246. The name
66 BOTANICAL GAZETTE [JANUARY
is mentioned in that work (NEES, 1829, Agrost. Bras. 226) in a note
appended to P. parvifolium Lam., where it is referred to P. angustijolium
Ell. It is quite distinct from ELLiort’s species, however.
Trintus himself describes the species | later as follows (1826, Gram.
Pan. 22
Spit beaes maeum. | Panicula (vix pollicari) lucidissima; Spiculis parvis, obovatis,
glabris: pedicellis scabriusculis; Gluma inferiore foarailia triplo breviori enervi,
superiore eosdem aequante 7-nervi; Hermaphrodito oblongo, acutiusculo,
laevi,*neutrum aequante.
V. spp. Am. bor. (TRATTINICK ex hbi°. Enslini).
Ima basi in ramos discedit simplices, tenues attamen firmos, satis multi-
(sex-)folios. Folia lanceolata, pollicaria, sensim breviora, praesertim basin
versus ciliatula. Panicula e summa vagin ae satis pauciflora, lineari-
oblonga. Flosculus neuter bivalis. Hermaphro
Since PLUKENET’s figures of this and the next cannot be identified,
TRINIUS’ spectmens should be taken as the substitute type of these two
species.
P, LEUCOBLEPHARIS Trin. 1822. Clavis Agrost. 234.
1177. Gr. miliaceum americanum, majus, panicula minore. Pluk. Phytogr.
p. 176. Tab. 92. f. 7. Mant. p. 95. (excl. Syn. Sloan. ut. ipse Sloaneus monet).
Citatur a Gronowio (Virg. p. 12.) ad Pan. paniculatum floribus muticis; sed
quid illud? Figura bene convenit cum Panico quodam herb. notsr. ex Amer. bor.
(Pan. leucoblepharis m.) praeter cilia foliorum elegantissima, rigidiuscula.—
Synon. Recchit ap. Pluk. admodum dubium
Label ereenenyte specimens: ‘“‘ab Enslino in Am. bor. 1. dt. cl.
Trattinick.”
Specimen =P. ciliatum Ell. 1816. Sketch 1:126.
Like P. lancearium the name is founded upon a figure in ‘’PLUKENET
and is further described by Trintus in Gram. Pan. 219. 1826.
Spithamaeum et minus. Panicula (ultrapollicari) lucida; Spiculis sub-
rvis, obovato-oblongis, pilosulis: pedicellis scabriusculis; Gluma_ inferiore
flosculis plus duplo breviore 3-5, superiore eosdem aequante 7-nervi; Her-
maphrodito oblongo, obtusiusculo, scrobiculato-punctato, neutrum aequante.
V er. bor. (TRATTINICK e plantis Enslini).
asi ramosum. Folia cordato-lanceolata, amplexicaulia, pl. min. pollicaria,
pallide viridia (plerumque elegantissime), pectinato-ciliata. Vaginae superiores
elongatae. Axis pilosus. Gluma inferior epilis. Flosculus neuter bivalvis.
P. ciliatum is characterized by the ciliate but otherwise glabrous leaves
and pubescent spikelets a little less than 2 ™™ long
P. UNCIPHYLLUM Trin. 1826, Gram. Pan. 242. :
Spithamaeum-pedale. Panicula (1-sesqui-pollicari) lucidula; Spiculis mini-
1906] BRIEFER ARTICLES 67
mis, oblongis, pilosis: pedicellis glabriusculis; Gluma inferiore flosculis triplo
breviori 1-, superiore eosdem aequante 7-nervi; Hermaphrodito elliptico,
laevi neutrum aequante.
Panicum laxiflorum Spreng. in Mém. de St. Pétersb. II. p. 291.
Panicum heterophyllum Miihlenb. teste N. ab Es.
V. spp. Am. bor. (TRATTINICK).
Culmus tenuis, adscendens, basi ramosus. Folia, quorum plura basi plerum-
lata: superiora angustiora, dissita. Panicula ovata, axis radiisque glabris.
Flosculus neuter bivalvis. Hermaphroditus albescens.
Label accompanying type specimen: “Pan. heterophyllum Muhl.
(Test. Nees) an Pluckn. Tab. 92 f. 8? ex herb Enslini, spmna Am. bor.
Trattinick.”
Specimen =P. columbianum Scribn. 1897. Div. Agros. Bull. 7:78.
In recent works this name has been applied to a species of the Janu-
gimosum group having rather stiff foliage and the leaf blades hirsute on
_ both surfaces. The true P. unciphyllum is easily recognized by the short
crisp pubescence and the very short ligule, characters not mentioned in
the original description.—A. S. Hirccock, U. S. Dept. Agric., Wash-
ington, D. C
SPOROGENESIS IN PALLAVICINIA.
Tue August number of the BoTANicaAL GAZETTE contains a paper
by Mr. A. C. Moore on Sporogenesis in Pallavicinia. 1 regret again to
ask for space on this matter, but Mr. Moore has so completely (though
of course inadvertently) misrepresented my own position with regard to-
the nature and the significance of the quadripolar spindle in the Junger-
mannieae, and further, the grounds on which he founds his criticism
appear to me to be so open to objection, that I venture to ask for an oppor-
tunity of replying to his strictures.
Firstly, then, as to the significance attached to the quadripolar spindle
in 1894-5.
From Mr. Moore’s account it would seem that I regarded, as the
Most essential feature of its importance, the simultaneous distribution
of the chromosomes of the dividing nucleus of the mother-cell to the four
spores that are finally produced.
I certainly believed that in Pallavicinia decipiens such a distribution
occurred, and that it resulted from the suppression of the period of rest
normally intervening between the first and second maiotic divisions. In
this I may be right, or further investigations may show that, in the species
In question, I missed the binucleate stage, But this is really not the
68 BOTANICAL GAZETTE [JANUARY
essential matter at all. The result of my work published in 1895 went
to show that in most forms there are two consecutive mitoses, the second,
following more or less rapidly on the first, and I believed that in P. decipiens
the brief interval might be so shortened as to have become practically
obliterated.
But the circumstance that quadripolar spindles were shown by me
to be plainly visible in properly fixed material of forms in which no such
extreme telescoping of the normal sequence of events takes place, clearly
proves that, whatever the significance of the quadripolar spindle may
be, it certainly is not essentially related to a simultaneous distribution
of the chromosomes amongst four daughter nuclei, and I never thought
it was.
What I believed in 1895 (and I have seen no reason to materially
alter my view), was expressed as follows: ‘“‘The quadripolar spindle, then,
is only a special case of ordinary karyokinetic phenomena; instead of
two relatively large masses of protoplasm there are four distinct aggre-
gations, one in every lobe, each exercising an independent strain, and the
direction of the strains may continue separate to the very end of the
process or not, according to the form and special circumstances of the
cell.”* I may perhaps add, that the principal importance of the phe-
nomenon, in my view, lay in its bearing on the permanence of the
centrosomes, at that time a widely accepted doctrine.
In the second place, Mr. Moore seems to think that his observations
on P. Lyellii vitiate the conclusions based on a study of P. decipiens.
I venture to think they do nothing of the sort. It is clear that the two
species differ in the form of their spore mother-cells to a marked degree,
and also that this difference is exactly of a nature to account for the unequal
persistence of the peculiarities of the spindle in the two cases. For the
lobing of the spore mother-cell is so much less in P. Lyellii than in the
other species, that it would be a matter for surprise if the quadripolar
character of its spindle were so long retained.
I confess, however, that I should have expected centrospheres to be
present at the stages represented in pl. III, figs. 1-3 of Mr. Moore’s
paper. They are so obviously demonstrable in Ameura pinguis and in
Fossombronia pusilla, the spore mother-cells of which resemble in their
lobing those of Mr. Moore’s plant.
One feels a little difficulty in repressing a suspicion as to the successful
fixation of his material, a suspicion not dispelled by the further contem-
plation of figs. r2 and 13. They so faithfully depict preparations I have
t Annals of Botany 9: 508.
.
7
1906] BRIEFER ARTICLES 69
myself very often obtained when the fixation had been imperfect. It
is, of course, easy in these plants to secure admirable preparations of the
stages preceding and following on the maiotic divisions, but I am sure
Mr. Moore will agree with me as to the great difficulty encountered in
successfully fixing the cell contents at this critical period. Personally,
I have not found chromacetic acid (the fixative used by him) very suc-
cessful, but obtained far better results with Flemming’s solution and,
if due precautions are taken, with acetic alcohol. The latter, in par-
ticular, has yielded results of especial excellence, owing partly, no doubt,
to the relative rapidity with which it traverses the somewhat impervious
cell wall—J. B. Farmer, Royal College oj Science, London.
REPLY.
PROFESSOR FARMER acknowledges that in 1894 he believed in the
simultaneous distribution of the chromosomes to the four spores in Pal-
lavicinia decipiens. His description stands as the only account of a pro-
cess without parallel in the plant kingdom, and he must have realized its
exceptional nature. The account became all the more remarkable when
Professor FARMER’S own studies on a number of liverworts, published in
the following year, showed two successive mitoses in the spore mother-
cells as in other groups of plants. He acknowledges now that he may
have missed the binucleate stage. This is precisely what I believe he did,
but since I have not investigated P. decipiens I cannot assert that he did
so. Now he states that this simultaneous distribution is really not the _
essential matter at all. Apparently the essential matter to him is his
observation that several liverworts conform to the normal sequence of
nuclear division during sporogenesis. Yet these conclusions, bearing as
they do on Pallavicinia decipiens, served to emphasize the peculiarities of
that account, and I feel confident that most, if not all, cytologists would
pick out the description of a simultaneous distribution of chromosomes as -
the most essential feature of his paper of 1894.
I venture to think that botanists are not so much interested in the
explanations which Professor FARMER may make of what he did or did
not believe in 1894 and 1895 relative to the quadripolar spindle (which
opinions they can form for themselves), as in the facts of sporogenesis in
the liverworts. My study of Pallavicinia Lyellii is plainly a challenge of
his account of P. decipiens, and together with Professor Davis’s work on
Pellia, leads us to believe that the ‘‘quadripolar spindle” in all liverworts is
a phenomenon of prophase followed by spindles of two successive mitoses,
in essential agreement with the events of sporogenesis in other plants.
vic) : BOTANICAL GAZETTE [JANUARY
The reader must judge for himself whether it is at all likely that two
species in the same genus should differ from one another so fundament-
ally as would appear from Professor FARMER’s description of sporogenesis
in Pallavicinia decipiens and my own account of P. Lyellii.
Respecting the fixation of my material, I may say that I have no reason
to think the penetration was not sufficiently rapid to fix the cell contents.
Even with imperfectly fixed material my main conclusion is easily demon-
strable, viz., that in P. Lyellii there are two successive mitoses in the
spore mother-cell. Let us not lose sight of the main point at issue—
ANDREW C. Moore, South Carolina College, Columbia.
CURREN T-LELERA TURE.
BOOK REVIEWS.
The algal vegetation of the Faeréese coasts.
BORGESEN’s extremely interesting account of the algal associations on the
coasts of the Faerée Islands,‘ is one of the most important contributions to the
cS eran ll «Sey
ecological side of marine botany. The work is a description of conspicuous algal
associations along a varied rocky coast line, particularly favorable to algal vege-
tation, and is illustrated by more than thirty very excellent plates and figures
from photographs. The factors affecting the algal vegetation are discussed;
such as temperature and salinity of the water, tides and currents, wave action,
temperature and humidity of the air, and light. The littoral and sublittoral
floras are described, both for exposed and sheltered coasts, and also the floras of
tide-pools and caves. A great many algal associations and formations may be
clearly recognized in the Faerdes, some of them very conspicuous, as the Chloro-
phyceae formation, the Porphyra association, Fucaceae formations, Laminariaceae
formation, and Alaria association. A particularly interesting chart plots the
position of these associations in their position above and below the mean sea level.
It is extremely interesting to note that the cave flora is composed of forms of
the sublittoral flora, which in the dim light are able to grow near the surface, or
they are types which have the habit of growing in shaded situations outside.
Littoral forms which grow in the brightest light are only found near the entrance
of the caves. On entering a cave a condensed picture is obtained of the vertical
: distribution of algae from above downward. The forms in the deepest shadows
f are all red algae and some of them species which are usually found at great depths
in the open sea. It is clear that light is the most important factor affecting the
position of algal associations along a coast.
There is a detailed comparison of the algal flora of the Faerée Islands with
neighboring countries, Scotland, the Orkney and Shetland Islands, Norway, and
Iceland, preliminary to a discussion of its origin. The flora had its origin from
a mixture of Atlantic and Arctic species, which wandered northward with the
retreat of the ice. Some of the arctic forms remained, adjusting themselves to
the warmer waters, but there are many peculiarities of the algal flora which
demand special explanations. BORGESEN does not believe that there were post-
glacial bridges of land which made possible the migration of forms, but holds
that factors now operative might have brought to the islands many algae from
neighboring countries.
* BORGESEN, F., The pee peta of Faerdese coasts. Im
834. pls. 13-24. Meads en: . Thiele. 1905. [Reprinted ible ‘Rtens a
the Faerdes. See Bor. GAZETTE ty 392. 1903-]
1906] | 71
ye BOTANICAL GAZETTE [JANUARY
Sea currents are regarded as of greatest importance. The pronounced cur-
rents from the nearest land do not bathe the islands, but experiments have proved
that heavy winds and storms will drive floating objects out of the main currents,
and BORGESEN believes the general conditions to be favorable to the introduction
of algae from the west and north coasts of Ireland, the west coast of Scotland,
and the Hebrides, while the currents from east Iceland run straight to the Faerées.
It is also possible that algae may be introduced from the west coast of Norway. -
Fragments of the algae may drift for many days, especially such as are provided
with bladder-like floats, or their spores may be so carried, and floating pieces
of timber covered with algal growths are known to travel long distances. Smaller
algae of the littoral flora are very likely to be introduced with mud upon the
feet and bodies of birds. Finally BORGESEN believes that algae may be introduced
through the shipping which visits the islands.
These are merely some of the most striking conclusions in an account that
is full of interesting observations on the life conditions and habits of marine
algae.—B. M.
Plant diseases.
FREEMAN has produced a finely illustrated volume on plant diseases,? the first
part of which is devoted to a discussion of fungi in general, while the second
special part treats of specific fungous diseases of plants. The object of this book,
as set forth in the preface, is “rather educational than immediately practical.”
It is an attempt to give a general account of the nature of fungi, for the purpose
the work becomes rather broader than would be indicated by the title, Minnesota
plant diseases.
t part comprises a discussion of the morphology, physiology, and
pee of fungi; but, while this part contains much excellent material, the
arrangement lacks the logical sequence of first importance in a book of an edu-
cational character. It consists rather of a series of interesting pictures without
due regard to pedagogical principles. This is likely to leave the mind of the
reader confused. The sub-headings of the first chapter on nutrition are as fol-
lows: What the fungi are; The fungus method of obtaining nutrition; How
the nutritive method is expressed in structure; Parasitism and saprophytism;
Storage organs; Fungus shoestrings or strands; Physiology of the mycelium.
Then, in chapter III, Fungus life methods, we have as the first subhead, again,
Parasitism and saprophytism, the rest of the chapter dealing with habits or rather
habitats of different aati 00 great an effort is made to avoid scientific pass
Thus we have such “‘spore-like swimming-spore-cases,””
“‘Sac-spore-capsule.” It would seem that the reader who can comprehend the
allusions to the phylogenetic relationships between fungi and algae would not
find it too difficult to comprehend a few scientific terms.
? FREEMAN, E. M., Minnesota plant diseases. Imp. 8vo. pp. xxiii+ 432- figs,
211. St. Paul: Rape of the Survey. Bot. Ser. V. 1905.
2 ey ciel apn ert age mar ani muratety 2m Cero eo eT a
1906] CURRENT LITERATURE 73
The second part of the book is devoted to descriptions of special diseases.
These are classified according to the nature of the crops on which they occur,
as follows: Timber and shade trees; Field and forage crops; Garden net
Orchards and vineyards; Greenhouse and ornamental plants; Wild pla
Under those heads the groups of fungi, as rusts, smuts, mildews, etc., are
together —H. HaAssELBRING.
Regeneration.
WITH THE TITLE Studies in regeneration NEMEC? has published in rather
voluminous form the results of his investigations on the regeneration of root-tips.
The general conclusions may be briefly summarized as follows. Cutting a trans-
verse section just at the tip results in the regeneration of a new tip in a radial
manner. The dermatogen-and outer part of the periblem takes no part in this,
the new tissue arising from the inner part of the cortex and the plerome. There
is first of all the formation of a callus of hypertrophied cells, between which and
the meristem arises the group of initials by which the new root-tip is organized.
This group is radial from the beginning, the majority of its cells arising from the
plerome, only the peripheral ones coming from the periblem. Proceeding back
from the tip, the se tesane for regeneration diminishes from the periphery inwards,
soon disappearing from the periblem. As long as the central cells of the plerome
still possess this capacity the regeneration is radial. Farther back it is confined
arises. When the capacity of the inner cells of the periblem and the outer cells
of the plerome to take part in regeneration is lost, the replacement of the removed
root-tip occurs only through the origin of lateral roots, which arise in the peri-
cambium.
When the root is cut through obliquely, the regeneration of the new root-tip
occurs at the part of the cut surface nearest the tip. When the tip is slit lengthwise
each half re-forms a new tip. If a tip is slit lengthwise for about 1™™, and then
one of the halves is removed by a transverse incision, the remaining half regene-
rates a new half, and also, at the surface formed by the transverse cut, a new tip
is developed. Lateral incisions to produce new roots must go at least half way
through the plerome. Unless such an incision is made just back of the tip a
new tip is soon organized immediately above the cut. The original tip is pushed
to one side and finally is displaced entirely. When the incisions are made on
two opposite sides of the root at different levels, new root primordia arise at each
place, but only the one nearest the original tip continues to develop. If two
incisions are made, on opposite sides and at the same level, a new root arises at
each, but one is soon suppressed, while the other develops and finally replaces
the original tip. About forty-eight hours after the wounding, starch usually
appears in the cells of the periblem just above the cut. The grains are not yet
3 NEmec, B., Studien tiber die Regeneration. Imp. 8vo. pp. 387. figs. 180.
Berlin: Gebriider Borntraeger. 1905. M 9.50.
2
74 BOTANICAL GAZETTE [JANUARY
mobile, and are aggregated about the nucleus. In about twenty-four hours more,
however, they become statoliths and fall to the bottom of the cells. During this
time the original tip has been losing its starch, and there is a period of from forty-
eight to seventy-two hours in which the old tip has lost its starch and the new
tip has none in a movable form. During this period the roots are ageotropic.
In ferns the root-tips do not regenerate. Tips cut off transversely just back
of the apical cell are unable to organize a new one, though they may continue
growing for several weeks. As the statolithic starch is in the root-cap, and this
does not regenerate, such roots remain ageotropic.
Besides the discussion of the experiments, a number of chapters are devoted
to a discussion of such topics as the influence of external conditions on regener-
ation, polarity and regeneration, growth and regeneration, purposefulness of
regeneration, relation between geotropism and the presence of statocytes, and
other interesting topics connected with regeneration.
As the root-tip regenerates from so many kinds of injuries that could never
occur in nature Némec considers that at least in the great majority of cases the
capacity could not have arisen because of its utility. The immediate stimulus,
he thinks, does not lie among nutritive changes, or arise from the wound, but is
a phenomenon of correlation, due to the breaking of the connection between the
vegetative tip and the root meristem.—W. B. McCattium.
Plant histology.
CHAMBERLAIN has revised and rewritten much of his Methods in plant
histology,+ adding several new chapters, elaborating and in many instances
shortening the processes. Several new formulae are given for killing and fixing.
e paraffin method has been notably improved and the celloidin method has
been treated at greater length. A method for embedding in soap is also given.
The new chapters deal with microchemical tests, free-hand sectioning, special
methods, the use of the microscope, and micrometric methods involving the use
of the camera lucida. A very important new chapter deals with methods of
staining filamentous algae and fungi and mounting them in Venetian turpentine.
An abstract of the methods of PFEIFFER and WELLHEIM is given, together with
such modifications as have been found to give successful preparations. Delicate
forms like Vaucheria can be carried through the stains and finally mounted in
Venetian turpentine without showing the least trace of plasmolysis, and even if
slight plasmolysis should occur it can be corrected by manipulation of the mount-
ing medium. Preparations made by this method are exceedingly brilliant and
show a wealth of detail not possible with other methods. For example, the two
nuclei in zygospores of Spirogyra can be readily seen with a low magnification.
The Venetian turpentine method, which gives preparations requiring no sealing
and as hard and durable as balsam mounts, should almost entirely replace the
glycerin method.
4 CHAMBERLAIN, CHARLES J., Methods in plant histology. pp. x+262. 7gs- 88.
Chicago: The University of Chicago Press. 1905. Net $2.25; postpaid, $2-39-
1906] CURRENT LITERATURE 75
Much attention is given to collecting and keeping material alive in the lab-
oratory. K1EBs’s method of securing reproductive phases in algae and fungi is
presented in a practical manner. Specific directions are given for making such
preparations as are needed by teachers and by those who wish to get a compre-
hensive view of the plant kingdom from the lowest to the highest forms. The
book will be useful to those who wish to keep in touch with modern microtech-
nique.—W. J. G. LAND.
Bibliographical index of North American fungi.
THE compilation of a bibliographical index of North American fungi by
FARLOWS is one of the most serviceable tasks ever undertaken in the interests of
American systematic mycology, and the publication of it by the Carnegie Institu-
tion one of its best contributions to the promotion of botany. The work is the
' outgrowth of an effort to bring together references to all North American species
in the form of a card catalogue. This was begun in 1874, at a time when there
was no complete record of the species known from North America. Within a
few years of its inception Mr. A. B. SeyMour was entrusted with the details of
this herculean labor, under Dr. Farlow’s direction, and his painstaking fidelity
is worthy of recognition.
It is the aim of the work to include all references having any bearing on the
taxonomy of fungi occurring in countries north of the Isthmus of Panama, the
scope of the original plan (which was restricted to the region north of Mexico)
having been greatly extended, on account of the close connection of species from
our southern border with those of Mexico, Central America, and the East Indies.
References to works of purely morphological, cytological, and sui
interest have been excluded; so have purely popular accounts, unless Ww
of use in giving divination of the species or in furnishing good inns.
In nomenclature the work is conservative. The principle of adopting the oldest
specific name has been generally followed. Where the vagueness of older descrip-
tions has made it uncertain to what stn they applied the writers have had no
scruples in rejecting the older n
The index itself is arranged ‘aiphaetically The names are printed in bold-
face type, synonyms and cross references being in italics. The citations, arranged
in chronological order under each name, follow the form adopted by the Madison
Botanical Congress in 1893 and by Section G, A.A.A.S. in 1894. In many cases
of confused synonomy, critical examinations were made of authentic specimens
and the related literature. Notes of interest obtained thus are added under the
Species in question. The present part, which is part I of the first volume, includes
names from Abrothallus to Badhamia.—H. HASSELBRING.
S Fartow, W. G., Bibliographical index of North American fungi. Vol. J,
part 1. 8vo. pp. xxxv+312. Washington: Carnegie Institution. 1905.
76 BOTANICAL GAZETTE [JANUARY
MINOR NOTICES.
Japanese vegetation —Professor Mryosut, of the University of Tokyo, has
begun the publication of photogravures of Japanese vegetation,® to represent
wild and cultivated plants and plant societies. Each picture is on a separate
sheet of cardboard 20.5 27°™, the size of the print being 16X23°". Accom-
panying the illustrations is a descriptive text in both English and Japanese.
The author has not yet determined the number of plates to be issued. So far,
two parts have appeared, part I containing eight plates of cultivated and semi-
cultivated an and part II containing eight illustrations of the vegetation of
the island o
The illustrations are well chosen and well made. Among the most effective
and characteristic are the long avenues of giant mountain cherry trees, gorgeous .
with their spring blossoms, the graceful bamboos bending beneath their burden
of winter snow, and the forest vegetation around the Hannya waterfall. The
descriptive text is precise, and interspersed by interesting remarks which show
that the ae has an eye for color and setting.
e hoped that the series may be continued to give us many more
Testes of the flora of this interesting country.—F. C. NEWCOMBE.
A botanical cyclopedia—-An illustrated German dictionary of botanical
terms has appeared under the editorship of Camitto K. SCHNEIDER,’ with the
assistance of a number of other German botanists. This volume of almost 700
pages presents much more than a list of definitions, for there are illustrated
descriptions of the morphology and minute structures of organs, of the sort one
would expect to find in a cyclopedia. The terms, of course, are those em-
ployed in the German language, and the work will not take the place, for the
English or American botanist, of Jackson’s excellent Glossary of botanic terms.
—B. M. Davis
NOTES FOR STUDENTS.
Chemotaxis of spermatozoids.—The chemotaxis of the spermatozoids of Isoetes
has been studied by Surpata.8 In Jsoetes japonica, which was used for the
study, the sporangia ripen in autumn. Microspores, sown in tap water in Perti
dishes late in November, begin to germinate about the middle of January. The
duration of the swarming movements of the spermatozoids is shorter than in the
ferns, vigorous movements lasting only about five minutes; some movement of
6 Miyosur, M., Atlas of Japanese vegetation. With explanatory text. Tokyo:
Maruzen Kabushiki Kaisha. 1905
7 SCHNEIDER, C. K., Illustriertes Handwoérterbuch der Botanik. Imp. 8vo.
pp- 690. figs. 341. Leipzig: Wilhelm Engelmann. 1905. M 16.
8 SHIBATA, K., Studien tiber die Chemotaxis der Isoetes-Spermatozoiden. Jahrb.
Wiss. Bot. 41:561-610. 1905.
SE ae a ee
a
y
a
‘
1906] CURRENT LITERATURE 77
the spermatozoid, however, may continue for ten or fifteen minutes, and of the
cilia for five minutes longer. PFEFFER’s capillary method was used in the experi-
ments. The principal headings are: position chemotaxis, relation between the
intensity of the stimulus and extent of the reaction, repulsion by free acids and
alkalis, negative chemotaxis with the ions of heavy metals, repulsive effect of
alkali salts, behavior with osmotically acting substances, repulsive effect of ions
of certain organic acids, the action of narcotics, theoretical, and review.
Malic acid acts as a strong topochemotactic stimulus and may be regarded as
the specific stimulant for the spermatozoids of Isoetes, although certain other
substances also exert some topochemotactic influence. Free malic acid in weak
solutions exerts a positive chemotactic influence, but in stronger solutions a nega-
tive one. The salts of various metals act as negatively chemotactic stimuli, as do
also the anions of di- and tribasic organic acids, including malic acid. The
positive chemotaxis with malic acid is of a typically topotactic nature. The
reaction consists in a turning of the body axis of the spermatozoid and a movement
toward the source of stimulation. Whether the structure for the perception of
chemotactic stimuli consists of the whole body of the spermatozoid or only of
localized portions of it is not yet determined.—C. J. CHAMBERLAIN.
Tuberization.—The causes of tuberization still furnish a field for study.
BERNARD first supposed that Fusarium Solani was the endophytic fungus of the
potato; this has since been disproved by GALLAUD and by BERNARD himself, but
the identity of the fungus is still undetermined. H. Jumette® has been con-
ducting experiments on Solanum Commersoni, a tuber-bearing species related to
the potato, but as yet his results are largely negative. The chief interest attached
to his studies are occasioned by the fact that S. Commersoni has small slowly
developing tubers placed on long stolons; these are the very characters which the
potato is said to have had when first introduced into Europe, before the endo-
phytic fungus became sufficiently abundant. S. Commersoni was infected by
fungi from S. tuberosum, but, as stated above, with negative results. JUMELLE
thinks that with suitable infection, it may be possible to secure tubers like those
of the potato, and further experiments are in progress. It should be said that
GALLAuD thinks that BERNARD has not yet isolated the true tuber-forming
fungus.—H. C. Cow es.
Two parasitic fungi—KirBaHNn'? has worked out the life histories of two
common species of the so-called Imperfecti group. The first of these is the
common elm fungus, Phleospora Ulmi (Fr.) Wallr. This is connected with an
ascomycetous form, which appears on the infected dead leaves during the
winter and ripens in spring, when the spores are ejected and infect the young
9 JuMELLE, H., Del’influence des endophytes sur la tubérisation des Solanum.
Rev. Gén. Bot. 17: 49-59. 1905.
to KLEBABN, H., Untersuchungen iiber einige Fungi im perjecti unddie ea
Aivtalapeles Rais I. u. Il. Jahrb. Wiss. Bot. 41: 485-560. figs. 75.
™
78 BOTANICAL GAZETTE [JANUARY
elm leaves. The form is named Mycosphaerella Ulmi Klebahnt'. Both conidia
_and ascospores produced identical mycelia in cultures. The Phleospora was
produced by sowing ascospores on the under side of elm leaves. No infec-
tion took place from spores sown on the upper surface. The study of Gloeo-
sporium nervisequum (Fckl.) Sacc. revealed a rather complicated series of forms
belonging to this fungus. An ascogenous stage develops on the dead leaves,
asin Phleospora. This is Gnomonia Veneta Klebahn. Beside the usual conidial
form and the ascogenous form, the fungus assumes thc form of a Myxosporium
on the young branches, and there produces the twig wilt always noticed on
sycamore trees affected with the Gloeosporium. A fourth form develops on
the dead leaves. This is a conidial form of the Fusicoccum type. Proof of the
connection of all these forms rests mainly in the similarity of the mycelia produced
in pure cultures from the various spore forms. Infections could not be pro-
duced readily, but a few cases of inoculation with ascospores were successful.
The various spore forms have been described under different names, which are
given in the synonymy.—H. HAssELBRING.
Shore formations in Denmark.—WarmIN«, in collaboration with WESENBERG-
Lunp and others in an interesting paper, has correlated the work of plants and
animals in the shore formations of western Denmark.t? A “vad” is a shallow
coastal lagoon, cut off from the sea proper by a line of islands, and bare at low
tide; the bottom may be of sand or clay, the latter type prevailing in the more
tranquil places. The sandworm, Arenicola marina, is the most characteristic
animal of the sandy “‘vader”’ or shallows, and the excrements of this worm are
found there in great abundance. Hence it has commonly been thought that
these animals have a soil function, similar to that of the earthworm, and that they
help to build up the shallow into a marsh. The authors, however, find that
Arenicola is very sedentary in its habits, swallows only from surface layers, and
that it retards rather than furthers soil enrichment. The waves wash any fine
particles landward, leaving the Arenicola shallow as sandy as before and hence
well adapted for the continued prosperity of Arenicola. In shallower water
closer to the shore, where the bottom material is finer, the amphipod, Corophium
grossipes, dominates; here the bottom is characteristically red-brown in colo1
and presents a riddled surface appearance. e Corophium shallows teem with
animal life, and here the influence of the animals is such as to convert the area
somewhat rapidly into a marsh. To this work blue green algae contribute some-
thing, but animals are much more important land builders than plants in the
Corophium shallows. Large areas of land have been gained from the sea in this
way in Denmark, and one case is cited where a fertile meadow has been developed
from a barren sandy shallow within two hundred years. In the argillaceous
tt Preliminary note in Zeitschr. Pfl. krankh. 12:257. 1902.
12 WARMING, E., Bidrag til Vadernes, Sandenes, og Marskens Naturhistorie. In
collaboration with Wesbrnies: LuND, OsTRUP , et al., with French résumé. Ko ngl.
Dansk. Vid. Selsk. Skrift. VII. beak 1904.
,
1906] CURRENT LITERATURE 79
shallows, in contrast to the above, plants are the more efficient land-builders;
the developmental processes in such places are well known and need not be
recounted here. ARMING also speaks of sandy plains that are subject to occa-
sional inundation. Here algae play a great part in soil-making; it is common
for a layer of Phycochromaceae to penetrate for three to five centimeters into the
sand, cementing the grains together, and giving a greenish appearance to the
ground. Many of the diatoms characteristic of such places are listed by habitats.
The peculiar depressions of salt marshes, called ‘‘pans”’ by OLIVER and TANSLEY,
are thought by WARMING to be formed where heaps of decaying vegetation have
lain; the consequent destruction of the vegetation makes it easy for the waters to
wash out the soil in such places.—H. C. Cow es.
Periodicity of sexual organs in Dictyota.—WILtAqs, in the third contribution
to his series of Studies in the Dictyotaceae,'3 discusses the remarkable periodicity
in the formation of the sexual cells in Dictyota. The sexual organs are produced
during the fruiting season in fortnightly crops, synchronous with the spring Sa
and a general liberation of the gametes takes place on a particular day, at a
interval after the highest spring tide, varying, however, in different localities
Of the factors (temperature, pressure, aeration, etc.) which fluctuate with the
alternation of neap and spring tides, the one which seems to account most satis-
factorily for the facts of periodicity is the increased illumination of the plant
during the low water of spring tides. The times of initiation and liberation may
be slightly accelerated or retarded by exceptional meteorological conditions, as
when wind causes a difference of two or three feet in the height of water, or a
rise of one inch in the barometer accompanies a depression of six or seven inches
in the tide—B. M. Davis.
Brown pigment of algae.—The generally SE view that the color of the
chromatophores of the brown algae and diatoms res the presence of
a brown pigment, phycophaein, in addition to " Madipbyll is challenged by
Mottscn,'+ who believes that the phycophaein garcons from these algae is a
post mortem product. He holds that the brown pigment is a + tie substance,
phaeophyll, which passes readily over to chlorophyll ates poabole with hot water,
alcohol, and other fluids. A similar brown pigment is found in the orchid,
Neottia nidus-avis. Beside the phaeophyll, 20 brown pan “ag diatoms contain
carotin and a bluish-green pigment, leucocyan
racts of Bennettites —LicNier's from a reexamination a his preparations
of che involucral bracts of Bennettites Morierei concludes that in all the sterile
scales, superficial or otherwise, which enter into the composition of the strobilus,
the terminal enlargement is due to hypertrophy and does not result from a reduc-
tion of the bract.—W. J. G. Lanp.
*3 Witiiams, K. L., Studies in the — tik; The origami of the
sexual cells in Dict yota dichotic’. Ann. Botany 19:531-560. figs. 6. 1905.
™4 Motiscn, H., Ueber der braunen behead der Phaeophyceen und Diatomeen.
Bot. Zeit. San. sone pl. O- 1905:
1SLIGNIER, O., Notes aaamacet ines sur la structure du Bennettites Morierei.
Bull. Soc. Linn. Norwaidie V. 8:(pp. 7.) 1904-
NEWS.
PROFESSOR EDUARD STRASBURGER traveled in Egypt during part of December
and January.
Dr. OskaR BREFELD, professor of botany at Breslau, has retired owing to
failing eyesight.
Mr. WALTER FISCHER has resigned his position as Assistant in Botany at
the Ohio State University, and has taken up work in the United States Department
of Agriculture, Bureau of Plant Industry.
Dr. A. F. BLAKESLEE, who is spending the winter in investigations at Halle,
was awarded the Bowdoin prize of Harvard University ($200) last spring for his
work on sexual reproduction in black molds.
In November and December Dr. Joan W. HARSHBERGER, of the University
of Pennsylvania, delivered a series of five lectures on “Weird and. marvelous
plants” in the Ludwick Institute courses of free lectures, Philadelphia.
Dr. Ernest A. Bessey has been transferred from Washington to the Sub-
tropical Laboratory of the U. S. Department of Agriculture at Miami, Florida,
where he may be addressed in future. Professor P. H. Rotrs, formerly of the
Sub-tropical Laboratory has accepted the directorship of the Agricultural Experi-
ment Station of Florida.
A RECENT circular gives an account of the Royal Hungarian Central Institute
of Viticulture, the buildings of which were completed in 1904. This institute
was initiated in 1896 by a law decreeing the establishment near Budapest of an
institution for studying the problems of viticulture and wine-making, and giving
scientific and practical instruction in these subjects. In 1898 Dr. IstvANFFI,
then professor at the University of Kolozsvar, but perhaps better known from
his association with BREFELD during part of the latter’s extensive investigations,
was called to organize and direct the new institution, whose first work was done
in quarters rented until the completion of the new ones. The present buildings
are five, each of three stories. The main hall, 66%21™ contains the library,
museum, offices, and lecture room. The four smaller halls, all similar in con-
struction, are devoted respectively to the four sections: (1) physiology and path-
ology, (2) chemistry, (3) zymology, and (4) practical viticulture and oenology.
The institution is excellently equipped for carrying on these different branches
of work. Its primary object is that of an experiment station whose field is restricted
to the wine-growing interests alone; secondarily, provision is made for instruction
to advanced students in the practical aspects of viticulture and oenology.
[1906 80
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L/1C GO PIANOS38 23222
| Vol. XLI A
Se
BOTANICAL GAZETTE
| February, 1906
| THE /00
)
Editors: JOHN M. COULTER and CHARLES R. BARNES
A gl _
CONTENTS
Chemotropism of Fungi Harry R. Fulton
a eee
The Embryology and Development of Riccia lutescens
and Ricca crystallina (with plates V-IX) Charles E. Lewis
Briefer Articles
Note on the Relation between Growth of Roots and of Tops in Wheat
Burton Edward Livingston
Current Literature iad
News
The University of Chicago Press
CHICAGO and NEW YORE
William Wesley and Son, London
Purity and Pears
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“All rights secured.” ai APS nin red OTTO OF ROSE IS THE BEST. —
Che Botanical Gazette
A Montbly Fournal Embracing all Departments of Botanical Science
Edited by Joun M. CouLter and CHartes R. BARNES, with the assistance of other members of the
botanical staff of the University of Chica
Vol. XLI, No. 2 Issued March 3, 1906
CONTENTS
CHEMOTROPISM OF FUNGI. Harry R. Fulton -— - es et gine eae 81
THE EMBRYOLOGY AND DEVELOPMENT OF RICCIA LUTESCENS AND RICCIA
CRYSTALLINA (wirH PLATEs V-Ix). Charles E. Lewis - - 110
BRIEFER ARTICLES.
OTES ON THE RELATION BETWEEN GROWTH OF ROOTS AND OF TOPS IN WHEAT. CON-
TRIBUTIONS FROM THE Hutt Botanica, LAporaAtory. (LXXXI WITH FIVE
ston
FIGURES). Edward Burton Livings n : : . - | 9
CURRENT LITERATURE.
BOOK REVIEWS eee ; i icneuk te tc Si SY LOSS 8 Gs
THE SWISS MOORS. REPRODUCTION OF MILDEWS,
MINOR NOTICES - - - - - - - - - - - tl < - 146
NOTES FOR STUDENTS - - - - - - - . - - - ~ - 149
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VOLUME XLI NUMBER 2
BOTANICAL GAZETTE
FEBRUARY, 1906
CHEMOTROPISM OF FUNGI!
HARRY R. FULTON.
HISTORICAL.
De Bary (5) advanced the supposition that the oogonia of
certain Phycomycetes not only attract the antheridia-producing
branches, but determine the formation of these branches as well.
The same writer (6) later raises the question as to whether the bend-
ing of a germ-tube toward the epidermis of its proper host, but not
toward every membrane or moist surface, may not be brought about
by a specific reaction in the parasite, induced by physical or chemical
stimuli which may be supposed to operate through unknown secre-
tions from the host plant. This was written, though not published,
before PrEFFERr’s first studies (25) on the reactions of motile unicel-
lular organisms to chemical stimuli. During the progress of his
studies with Saprolegnia swarm-spores, he observed that the hyphae
of the fungus turned toward the nutrient substances, and he supposed
that chemical agents might in many cases determine, the direction of
growth of these and other hyphae.
KIHILMAN (16) observed that if a germinating ascospore of Mel-
anospora parasitica lies at a distance of not more than four or five
spore-diameters from a growing hypha of its host, Jsaria farinosa,
the direction of growth of the latter is deflected toward the spore of
‘the parasite until there is contact with its germ-tube.
According to BREFELD (1), the fact that neighboring sporidia of
Ustilaginaceae conjugate in pairs, the connecting tubes taking the
shortest course between two sporidia, may be explained by a
a directive chemical influence.
* Contribution No. 4 from the Botanical Laboratory of the University of Missouri.
81
82. BOTANICAL GAZETTE [FEBRUARY
WorTMANN (36) observed the turning of young germ-tubes of
Saprolegnia sp., and concluded that these are very sensitive to chem-
ical stimuli, especially to those concerned in nutrition; a most
energetic turning was observed toward flies’ legs.
MARSHALL WARD (32) mentions two factors as mainly influen-
cing the direction of growth of “‘lily-botrytis,” namely, the contact of
hyphae with one another or with solid bodies, and the direction in
which food lies in relation to the growing hyphae. |
Worontn (34) holds that it is through chemical influence that
the conidial germ-tubes of Peziza baccarum reach the wounds of the
host plant.
STRANGE (29) made careful observations on growing hyphae of
Saprolegnia ferax for the purpose ‘of confirming their reported turn-
ing toward nutrient substances. That the hyphae turn toward a
region of diffusing nutriment was regarded as very questionable;
there was noticed, however, a stronger growth of the hyphae in the
nutrient region, and the hyphae, by their branching, became more
abundant here than elsewhere. The germinating conidia sent their
tubes in all directions, provided enough nutriment for growth was
present, and not markedly toward any area containing a greater
amount of nutriment. Similar results were obtained with germ-
tubes from Penicillium spores; these showed no chemotropic turn-
ing, but a much better growth when they reached the diffusion
region around the openings of the capillary tubes containing the
test solutions.
REINHARDT (28) found that the direction of growth in Peziza
sp. is affected by chemical influences. Gelatin containing sugar and
spores of Mucor sp. have an attractive effect. Spores of Aspergillus
niger, A. flavus, and Penicillium glaucum, as well as colonies of
various bacteria, cause a cessation of growth, which is followed by a
reversal of the direction of growth.
BUSGEN (2) observed chemotropism in the case of Botrytis cinerea,
and supposed that a chemical stimulus might cause the germ-tubes
in the dew-drop to seek the host epidermis; but he considered that
penetration is brought about by contact influence. He asserts the
possibility of chemotropism in the case of germ-tubes from uredo-
:
F
Pi
i
Be
_*
it
Ma
E
‘@
‘ae
1906] FULTON—CHEMOTROPISM OF FUNGI 83
spores and from the conidia of Peronosporaceae, and of chemotaxis
in the case of Cystopus swarm-spores.
Many of the foregoing are merely opinions or passing observa-
tions, made in the course of investigation of other phenomena.
Mryosur (20), however, made chemotropism of certain fungi the
subject of systematic and extensive experimental study. The -prin-
cipal fungi used by him were Mucor stolonijer, M. mucedo, Phy-
comyces nitens, Penicillium glaucum, Aspergillus niger, and Sapro-
legnia ferax. The tests were made with the aid of perforated mem-
branes, such as strips of epidermis, celloidin films, and mica plates;
of capillary tubes; and of injected leaves and petioles of Tradescantia
sp. He concludes that, in the case of the fungi enumerated, mole-
cules of many substances diffusing from the openings cause diversion
of the hyphae from their original direction of growth, the turning
being either toward the diffusing substance (positive chemotropism),
when the substance is attractive to the fungus, or away from the
substance (negative chemotropism), when the substance is repellent.
Some substances are wholly or almost wholly neutral. The direc-
tion and amount of turning are dependent upon the concentration.
Chemotropism is most marked at an optimum concentration, which
varies for the substance and the fungus. The concentration just
sufficient to cause turning is very low for most attractive substances.
Repellent substances are acids, alkalis, alcohol, certain neutral salts
and toxic compounds; also very strong solutions of substances that
are neutral or attractive at lower concentrations. Generally attract-
ive substances are ammonium nitrate, ammonium chlorid, ammon-
ium malate, ammonium tartrate, potassium phosphate, sodium
phosphate, ammonium phosphate, meat extract, peptone, sugar,
asparagin, etc. For chemotropic phenomena Weber’s law holds.
The effect of an attractive substance may be overcome by the pres-
ence, in sufficient quantity, of a repellent substance.
The same investigator (21), in connection with his study of the
penetration of natural and artificial membranes by fungi, found that
the hyphae of Botrytis cinerea and Penicillium glaucum would grow
through a rhembrane only when they were placed on nutrient substrata;
there would be, however, no penetration through the membrane to
84 BOTANIGAL GAZETTE [FEBRUARY
the substratum if the fungi were grown in a nutrient medium, although
the mycelial growth was more vigorous.
SWINGLE (31), in explanation of the effects of Bordeaux mixture,
advanced the hypothesis, which he based on the studies of REIN-
HARDT, BiscGEN, and Mryosut, that copper hydroxid may prevent
the germ-tubes of parasitic fungi from entering the host plant through
negative chemotropic action.
NORDHAUSEN (24) accepted MryosuHt’s conclusions, and investi-
gated the biological bearing of chemotropism upon the penetration
of plant tissues by certain fungi, without bringing forward any
additional evidence in favor of chemotropism.
No further investigation seems to have been made with relation
to fungi, until CLARK (4), in his investigation of SWINGLE’s hypoth-
esis, had occasion to test the chemotropic reactions of certain species,
especially Mucor stolonifer, to toxic substances. For the most part
he followed Mrvyosut’s methods closely. In every case it was found
that the hyphae would turn from a nutrient medium and grow into
media containing such toxic substances as salts of copper, cobalt,
nickel, zinc, etc., until a concentration sufficient to cause death was
reached. The hyphae turned toward non-nutrient media and dis-
tilled water as readily as toward nutrient media. His conclusion
is that Mucor stolonijer is negatively chemotropic to some secretion
of its own mycelium, and that this negative chemotropism is much
greater than any positive chemotropism it may have for food sub-
stances or oxygen.
MASSEE (19) found that fungi are attracted to their hosts by
specific stimuli from substances in the cell sap. In the case of
saprophytes and facultative parasites, the attractive substance is
saccharose; the facultative parasites, however, may be repelled by
more potent negatively chemotropic substances in the cell sap. In
the case of obligate parasites, the cell sap of the host plant is the
strongest positive chemotropic agent; malic acid is the specific attrac-
tive substance for the germ tubes of Monilia fructigena, and the enzyme
pectase for Cercospora cucumis. Immune plants owe their immunity
to the absence of the chemotropic substance.
Other factors affecting the direction of growth of fungous hyphae
have received but little attention, while the causes of the bending —
}
F
é
Ey
ca
a
f
,
1906] FULTON—CHEMOTROPISM OF FUNGI 85
of sporangiophores, especially of certain Mucorineae, have been
carefully studied.
A negative hydrotropism for the sporangiophores of Phycomyces
nitens was first experimentally established by WoRTMANN (35), and
was later confirmed by the more extended studies of- ERRERA (10)
and of STEYER (30). MotiscH (22) showed that the sporangio-
phores of Mucor stolonifer and Coprinus velaris are negatively hydro-
tropic; while Kress (17) made similar observations for Sporodinia
grandis, which FaLcK (12) has failed to confirm.
WorTMANN (35) observed what he supposed to be negative
hydrotropism in the case of the mycelium of Mucor sp., which would
grow towards water, but would turn aside and branch profusely
on reaching its surface. The conditions of experiment were such
as to make his explanation of the phenomenon doubtful. STEYER
(30) concludes that moisture plays an unimportant réle in deter-
mining the growth and spreading of the mycelium of Phycomyces
nitens.
JONsson (15) grew mycelia of Mucor stolonijer, Phycomyces
nitens, and Botrytis cinerea on sloping filter-paper strips having
their two ends dipped in vessels of water at different levels. Phy-
comyces and Mucor showed negative rheotropism under these con-
ditions, while Botrytis showed positive rheotropism.
HorMEISTER (14), WORTMANN (35), Dietz (7), KLEBs (17),
and STEYER (30) have established for various Mucoraceae a negative
reaction of their sporangiophores to gravity and to strong light;
but there is a positive reaction to contact and to moderate intensi-
ties of light.
Kwny (18) holds that gravity has no effect upon the growth of
the mycelium of Mucor mucedo, M. stolonijer, Trichothecium roseum,
and Eurotium repens. Muyosut (20) concludes from his tests that
neither gravity, light, contact, nor moisture affected the turning
of the six fungi used in his investigations. STEYER (20) in his study
of the reactions of Phycomyces, found that the mycelium is indifferent
to light, contact, and gravity.
MATERIALS AND GENERAL METHODS.
To a greater or less extent fourteen species of fungi have been
used; these, with respect to nutritive adaptations, may be classed
as follows:
86 BOTANICAL GAZETTE [FEBRUARY
STRICT PARASITES STRICT SAPROPHYTES
Uromyces caryophyllinus Mucor stolonifer
FACULTATIVE SAPROPHYTES Mucor mucedo
Sphaeropsis malorum Phycomyces nitens
Cercospora apii Penicillium glaucum
Monilia fructigena Monilia sitophila
FACULTATIVE PARASITES Sterigmatocystis nigra
Botrytis vulgaris Coprinus micaceus
Daedalia quercina c Agaricus fabaceus
With exceptions as noted below, spores of the various fungi from
pure cultures one to two weeks old were used in making inoculations.
S phaeropsis malorum was not ready in pure culture as soon as needed,
and inoculations for the first experiments with this form were made
directly from infected apple twigs; the spores were found to germi-
nate quickly and the hyphae grew rapidly, so that the observed
bacterial and mold contamination in these cultures was slight.
Cercospora apii, obtained from celery leaves, was grown in artificial
culture on pieces of sugar beet; spores were not produced, but
satisfactory inoculations were made with small portions of detached
mycelium. Inoculations in the case of the three Hymenomycetes
were made with portions of mycelium from pure cultures, which
had been made from sporophores by the “tissue-culture” method
(DuccaR, 9). Spores of Uromyces caryophyillinus, taken directly
from carnation leaves, were used to some extent. The germination
of these was not certain under all conditions, and the growth was
limited; the use of the fungus was soon abandoned. All other
spores gave perfectly satisfactory germination in gelatin and agar-
agar media. Even such species as Penicillium glaucum and Ster-
igmatocystis nigra, which have been found (DuceGar, 8) not to ger-
minate in distilled water, gave a germination of practically 100 per
cent. in gelatin and agar made up with distilled water.
Precautions were taken to have all apparatus chemically clean and
thoroughly sterile. Glassware and mica plates were boiled in alkali
and in acid, and again, after a thorough rinsing, in distilled water-
Covers were rinsed in 95 per cent. alcohol, then wiped with a sterile
cloth. Heavier glassware and mica plates were sterilized with dry
heat at a temperature of 140° to 150° C. Celloidin films were steril-
ized by being boiled in redistilled water; strips of epidermis, by being
tikes ee
1906] FULTON—CHEMOTROPISM OF FUNGI 87
rinsed in alcohol and afterwards soaked in two changes of sterile
redistilled water. The media used were sterilized, whenever possible,
under 15 Ibs. pressure of steam, or by fractional sterilization at 100° C.
Except in those experiments in which capillary tubes were used,
contaminations were of rare occurrence; whenever contamination
was apparent, the experiment was disregarded in tabulating results.
With few exceptions the chemicals used were the chemically pure
_ preparations of reliable manufacturers. The water used in making
up test solutions was redistilled in glass apparatus. All experiments
were made in duplicate, and were repeated when occasion demanded.
A fairly constant temperature of 28-29° C. was maintained for the
cultures. Beet decoction, made by boiling 450%" fresh weight of
sugar beet root in 1,000°° tap water, was the usual basis of nutrient
agar and gelatin media. The stock decoction was diluted two to
ten times for use.
TESTS FOR CHEMOTROPISM.
The capillary tube method.—In the first tests PFEFFER’S (26)
method with capillary tubes was used. These tubes were filled
with the chemical solutions under the air-pump, were rapidly rinsed
in sterile distilled water, and were placed on the cover glasses in
drops of the culture medium, while it was still liquid. The cover
glasses were then inverted over Van Tieghem cells made up in the
way described by CLarK (3), and were sealed to the cell rim with
vaseline. A small quantity of liquid, the same in composition as
that used in making up the culture medium, was placed in the bottom
of each cell. Observations were made when the hyphae were 40-75
spore-diameters in length, and again three or four hours later when
growth had become indefinitely great. In estimating the effects
of the chemicals, regard was had for the hyphae from spores lying
within a radius of one lumen diameter from the opening of the tube,
and such other hyphae from more distant spores as entered this
area. Only those were held to be chemotropically affected that
showed a turning out of their former courses toward or away from
the tube opening. In recording the observations Mryosxt’s method
was used; to denote a turning away of 12 per cent. to 37 per cent.,
the symbol r was used; for a turning away of less than 12 per cent. and
an attraction of less than 12 per cent., the symbol 0; for an attraction
6888 BOTANICAL GAZETTE [FEBRUARY
of 12 per cent. to 37 per cent., @,; for an attraction of 37 per cent.
to 62 per cent., @,; etc. An interrogation point indicates a value
near the lower limit of the class indicated.
This method was used in the preliminary testing of a large number
of representative chemical substances with Sterigmatocystis nigra
and Mucor mucedo; Botrytis villgaris was used with a few that gave
decided effects. The series included the sulfate of sodium, of
magnesium, of calcium, of ammonium, of potassium; the normal
phosphate of ammonium, of potassium; the dibasic phosphate of
ammonium, of potassium; the monobasic phosphate of potassium,
of sodium, of calcium; the chlorid of potassium, of ammonium, of
calcium, of magnesium, of sodium, of lithium; the nitrate of potas-
sium, of ammonium, of calcium, of magnesium, of sodium; the
oxalate of magnesium, of potassium; the tartrate of magnesium,
of potassium, of sodium-potassium; the bitartrate of potassium;
the lactate of magnesium; the malate of magnesium, of ammonium;
the acetate of potassium; acetic, lactic, tartaric, malic, oxalic acids;
cane sugar, dextrose, galactose, maltose, lactose, starch; glycerin,
ethyl alcohol; active pepsin, boiled pepsin, trypsin, peptone, mannit
agaricin, casein, asparagin, urea; meat extract, beet decoction, bean
decoction, distilled water; mercuric chlorid, copper-sulfate, copper
acetate, lead nitrate, zinc nitrate, iron sulfate, iron chlorid, potassium
ferrocyanid. In the case of such of these compounds as were tested
by MrvyosuI, concentrations were used that would give the maximum
effect of attraction or repulsion. With nearly all compounds two,
and with many three, concentrations were tested.
The hyphae showed a tendency, in small numbers, to turn toward
the tubes in the majority of tests; but in only a few instances were
more than about 37 per cent. of the hyphae in the observed area
TABLE I
A se S S $ in > = s, Ss} tid “J
Pe ra S| dé le le le BS) BS [gael gS
SPECIES Eacl S| | 3 FaaclZacl“ac| “nel Sas] J |. les
a} O |O hse Fa|.8 Mo a\e8 g au 4
SSIS [g lg Bs) & geige
o ae < a BAW l<
Sterigmatocystis nigra.......... Fire . Po a ee ps r
Mucor mucedos. Yoav eeu: jvc) Qe fest 4 ae lect YF | Bel ae) Ax] Bat Oa) Se
Botrytis vulgaris... cic. es cok fla [ocebessdeks
r ily te SEAS Bi nee banca lear ae
ee : Pe RSS ae x bigest uted
a a eI Te fee IB ine hy, S He ne REN é i
1906] FULTON—CHEMOTROPISM OF FUNGI 89
affected. The repellent effect was much less marked, never affecting
more than one-fourth of the hyphae, and even in the most marked
cases some of the hyphae would grow toward the tubes and enter
them. Table I gives the most marked instances of attraction and
repulsion noted in the series; the symbols have been explained above.
In Table II is given a comparison of the effects of a number
of representative compounds as observed by Miyosnr and in the
present investigation. Leaving out of consideration Saprolegnia
jerax, the extreme effects recorded in Mryosut’s table were obtained
TABLE It
Aspexott- |__ STERIG-
Mucor MUCEDO LUs NIGER | MATOC¥STIS
REAGENT NIGRA
I i iil IV
onobasic sree phot, 2 Joss. a3 ae ar ar
Monobasic sodium phosphate, 2 %....... ae a2? ed bi
utral ammonium oles phate, 2%...... ay az a2 as
al ammonium phosphate, 0.2% ee ax ee ar
Neutral ammonium phosphate, 0.1 %.. ay os ciate
Pirate, By eee es eae G a3 az? az r
Potassiuin nitrate, ¢ 0... 2... 2 es seca r ay r _
Calcium nitrate, 29%... 60 0c eee e e ar Tr .
Magnesium sulfate, 2%.........++...+- r fc) r r
€ ice Mike eo se we se ew eeee a3 ° ar °
Potassium chlorid, 2%... 02.02.0000 r ° r 2
odi OM Bek usivi'g on aees os r ° r °
um-potassium tartrate, Oe Nee eee r a2? r °
Cane sugar, 2 A epee avccste sey Garg wks ta ae ay ° ar °
2a ree tetera we cae a eae a —
= ee Epeloe Sat ee bs eee ae ° ae °
us iy Boy Pape ee ae eg a3 ar a2 ar?
ad = pe Paine © pe ere ay *e -
Lactose, 2 2% Oe ence es a, ay? vee
Oey kk MESON chivas ESET ok week ay ay?
Malice’ 2 %, Rohe Wire hw eee e Wie ay ete ay ° see
ER 2 Sear ee Ey tee ro fs) ax?
Dextrose, $0 Geet ee a, aa ae
ies OR een gr ea er ee ° ay
a 2 %, WOKS Ste Big ce ee a ae eee Bae ag a3 5h
1% ee ee a3 r ? ay °
Ethyl alobol, Wists paSiy penne aces z a2? r ay?
— i eee ele ao a kee a eee we we ay 3° ay ? ay Fi
a SEE SS SRUREN, Manian eae tae pe ag a2? ax ar?
2 oF EEE SEE acer pene eas te Sees ° ay? ° ay?
. Peptone, re eae ° aot az?
J M. - % Peer w ere e sere ress eerteonese a3 a2 mae
leat extract, ta ee RES re ere ay ag ag ?
1% a3 ar ag ay
es eS aries ay ° as af
Rs ae eee ay ° ° ar?
go BOTANICAL GAZETTE [FEBRUARY
with Mucor mucedo and Aspergillus niger; the values given by him
for these forms are to be found in columns I and III below. It was
impracticable to use Asperillgus niger in the present study, and
' Sterigmatocystis nigra was substituted with the understanding that
the two forms are not always distinguished. The present results
for Mucor mucedo and Sterigmatocystis nigra are given below in
columns II and IV respectively. '
In the control cultures, where distilled water was used in the
tubes, the effect was the same as for the majority of chemical sub-
stances; that is, 10-30 per cent. of the hyphae turned toward the
tubes. The same amount of positive turning was observed in the
case of all the strongly toxic compounds used. Even with a 0.05 per
cent. solution of mercuric chlorid and a 1 per cent. solution of copper
sulfate, which completely inhibited germination within a radius of
eight to twelve tube diameters from the openings, the hyphae not
only grew across the diffusion areas, but 10-30 per cent. of those
approaching the openings turned toward them and grew for a con-
siderable distance into the tubes.
Although four concentrations of cane sugar ranging from 20 to
0.1 per cent. and four concentrations of meat extract ranging from
10-0.01 per cent. were used, no definite relation between the strength
of stimulus and that of response was apparent.
Two corresponding series were made with ten representative
compounds; in one series sugar-beet agar was the culture medium;
in the other, distilled-water agar. No difference in the behavior
of the hyphae due to a difference in the media in which they grew
could be observed.
Tests with mica plates —Thin sheets of mica were cut into pieces
about 25X16™™; these were perforated with a needle, the holes
being 0.1-0.15™™ in diameter, and about 2™™ apart. Covers of
suitable size were cut from glass 1™™ thick. A layer of gelatin or
agar was placed on a cover, a mica plate was placed on this just
before it hardened, and a second layer was placed above the plate.
The chemical to be tested was made up in double strength solution
in redistilled water, and one volume of this was added to one volume
of gelatin or agar, also made up double strength in redistilled water.
It was usually found convenient to have the layer containing the
ice Ce Seen: ce
.
1906] FULTON—CHEMOTROPISM OF FUNGI QI
chemical next to the cover, the spores being distributed in the outer
layer. The cover was inverted over a stender dish of 36™™ diameter
containing distilled water, and was sealed with a coating of vaseline
around the rim of the vessel.
This method seemed to possess distinct advantages over the one
with capillary tubes: the numerous perforations made it possible
to make a large number of observations from a single preparation;
TABLE III
5 s S g 2) E 8 4 &
z g z : Ba<| & 2 Ae |p E
4 i] no
ae ea =
i = a im
Neutral potassium phosphate, 2 2% ecesas 12 19 24 22 33 ee
A a s3 20 18 25 36 25
Monobasic potassium phosphate, 2 2 fe =} ag 22 24 26 36 23
Ds| 20 26 23 20 25 21
Dibasic ammonium eS hate, 2 % 13 20 2I 20 15 23
ane ee 2% ie saped eweaes 49” | 41 34 31 35 Bd
Phosphorie acid, 6.9 9.06054 less cn ces 48’ | 30 35 39 23 3°
Malic acid, r 1% AP Serra ali, le te ee x x 24 x 38 x
e me | day aie et teh tare Oe 21 21 26 21 17 x
Tartaric acid, 170 per ne Re eae Ne ae pe * 20 43 22 19 18
5 7 eicaie ee el 25 27 34 25 24 22
Oxalic acid, 1 es eet er aan x 17 23 x 17
0 Rcainintt em aaianad Ges 23 22 23 34 II 40
Cane sugar, 5 $ Yor. iver sSa mew ea eas waa nee 38 32 24 31 a0 48
BE el. ca pene oe 30 37 20 30 20 55
. Sp ee ee ere eee ra ee 30 35 24 a2 23
Glucose, 5 o. Rea Segre Grape igs Dog oa are a 39 39 20 34 30 37
et Gnic aGere sete ee ce un 40 24 23 39 20 3°
POPRONE BT Senne 14 28 21 13 20 19
: eee ais ws Son cs Wee eas ous 26 23 20 21 13 22
Pepsin active Nig ae ae SSE 24 20 19 42 23 30
te ; 3 Sey MCLE as pee 25 30 22 27 31 22
Potced Sopa 2 %.. eee isn neers a6 t.. 37 ah 33) 4 | ee
Aspa: dy OR analaeee eireeterer acer tty 19 23 30 23 30 15
be ey ot as 15 23 30 20 30 24
es sulfate, 0.00 % Nees ae wae aie 9 21 21 25 34 31
Dibasic sodium phosphate, 2 2% ors uaa? 35 25 26 26 15 20
mew wees 38 Ki) 27 20 26 a3
Starch vag Mek. WORE ROT OE Tee 24 20 21 27 25
te a see ge tree cee . 27 22 20 26 15
Maltose, 2 2%, Sree ee earn Reena 33 30 22 21 25 14
pee eect aire e sss 36 29 28 31 30 28
Control (lestilbad wate @elatin) ©... 5 60s 28 30 29 24 22 18
Note: gana X indicates that germination was inhibited. Uromyces
caryophyllinus was tested with the majority of these substances, in the few
instances in which growth was sufficiently great for a determination, the —
was practically the same as for the other fungi.
g2 BOTANICAL GAZETTE [FEBRUARY
the medium containing the chemical and that containing the spores
could be more nearly equalized in amount and in consistency; fewer
hyphae would take a course that would lead them through the open-
ings without apparent turning; better sterilization could be secured;
and there was less difficulty in making up the preparations.
In making the counts, hyphae within a radius of one opening
diameter from the margin of each opening were considered; the
hyphae within such an area were classed in the counts as those
turning toward the openings, those turning away from the openings,
and those apparently indifferent. After an examination of the
entire preparation in each case, those holes were selected for the
counts which represented the average condition. In calculating
the percentages from the counts, the difference between those attracted
and those repelled was made the dividend, and the total number
within the observed area was made the divisor. The results are
shown in Table ITI.
If the percentages of turning toward distilled-water gelatin as
determined by the control experiments, be deducted from the per-
centages of turning toward the various chemical compounds, it will
be found that in only three instances would the difference, which
would be supposed to indicate the amount of turning due to chemical
influences, approximate 25 per cent., or about the average of the
lowest of the several degrees of positive chemotropism recognized
by Mryosui. Most of the values, even for supposedly highly attract-
ive substances, are very near the controls.
Evidently, the results thus far have not been favorable to the
theory of chemotropism. But it was thought that the fungi, all of
which grew perfectly normally in gelatin made up in distilled water,
might turn more strongly from some other medium in which there
was less of available nutriment, to one having an abundance. In
agar made up with distilled water, the fungi germinated after a
greater length of time and grew more slowly; but the turning from
this non-nutrient agar toward a nutrient salt solution favorable for
fungous growth was no more marked than from agar containing the
same proportions of the nutrient salts to the nutrient salt solution;
nor did the nutrient solution seem to attract from either medium
more strongly than ‘did distilled water. This test was made with
1906] FULTON—CHEMOTROPISM OF FUNGI 93
Sterigmatocystis nigra, Mucor mucedo, and Mucor stolonijer by the
capillary tube method. |
It is possible, although there is ‘evidence against it in some of
the previous experiments, that the diffusion of the solutes might
be rapid enough to bring about practical uniformity in the media
before the germination of the spores. To determine whether the
time at which the stimulus is applied has an influence upon the
reaction, series were made up with two of the more slowly growing
species, Botrytis vulgaris and Monilia fructigena, the spores being
distributed in non-nutrient gelatin placed above the mica plates.
Four duplicate series were arranged. In one the layers of gelatin
containing the substances to be tested were placed below the plates
at the time of making up the cultures; in another series these layers
were added just as the spores began to germinate; in a third, after
the most vigorous hyphae had attained a length of 15 to 20 spore-
diameters; in a fourth when the length equaled 40 to 50 spore-
diameters. The final counts were made a little later, when the
growth was about 75 spore-diameters. The results are given in
Table IV for the five compounds tested and the control. The
numbers in columns I indicate the average number of hyphae in
the area around each hole; those in columns II, the percentages of
turning toward the holes.
TABLE IV
Non-
Hi Ox KH,PO (NH,)NO
sve Beat] Paar | SSG [mua | MEO | ON | come
Cres | Appli-
cation ‘
i | ue | I Ir | I | n | I | It I | II I It
8 of} ar | 35 | xz 38 | 6.3 4-9} 37 | 5-5 | 36 | 9-2 | 34
a3 £0. 9.6). 38° 1 Gio 1238 Te. | 36.1 §4.) 30 | 72} 36-1 7-61 34
e6| 13 a) ge fee Tae tat 95.1 6.4.) 3t $14: 33.4 9.21 38
AE! x6 || 3.3| 30 | 7-6| 34 | 5-5] 3015-8] 35 | 5-0| 32 | 5-6 | 36
<
Sa0.0 UG.g1 384-260-140 1 tr] 98 4 3.7) -3t.| 8:2 | 34 | 4-4 [06
Be 10 9-5 | 36 1 3x ao 4 Es ao. }.3,7 | 38--1.6.8 | 25) 21 -| 32
S8| ts | 13 | 33 |7-s| 27 | 10 | 32 | 4-3| 30 | 5-4] 27 | 33 | 27
i} 20 fT 32 ay 16.47 8615-5 1. 20 1 8-1] 26 | 8-3 | go
It is evident that the time of application has little or no effect upon
the amount of turning. It will also be seen by comparing these
94 BOTANICAL GAZETTE [FEBRUARY
results with former ones, that it is immaterial, as far as the per-
centage of turning is concerned, whether the spore-containing layer
is above or below the one containing the test substance.
Other culture media were used, such as 10 per cent. and 25 per
cent. glycerin, gelatin made up to contain 10 per cent. of glycerin
and to contain 5 per cent. of cane sugar. With none of these were
there more distinct indications of chemotropism than in former
tests, in which non-nutrient gelatin was the culture medium. This
would indicate that the available nutriment and the concentration
of the medium have no effect.
The final test along this line was made with silica jelly, a medium
free from organic matter and of suitable consistency. ‘The method
of preparation was that used by Moore (23), except that dialysis
was accomplished satisfactorily with parchment paper. In order
to secure the proper coagulation of the medium, it was necessary
to add mineral salts to all media used. A nutrient salt solution,
containing 18“of ammonium nitrate, 0.5%" of monobasic potas-
sium phosphate, 0.25%" of magnesium sulfate, a trace of ferric
chlorid, and 58" of glucose, in a volume of 1o0°*, was made the
basis of the work. It would seem that if fungi show chemotropism
under any conditions it would be by turning from a medium lacking
some one or more of the elements necessary for full development,
toward the diffusion centers of compounds supplying the missing
element or elements, or toward a full nutrient solution.
In the tests each compound in turn, except ferric chlorid, was
omitted from the silica jelly containing the spores, and in each case
jelly containing the omitted compound in proper proportion on the
one hand, and full nutrient jelly on the other were used on the opposite
side of the mica plate from the above-mentioned spore-containing
layer. Control tests were carried on with jelly lacking one and the
same compound on each side of the plate, and also with full nutrient
jelly on each side of the plate. To reduce evaporation, the lower
layer was covered with an unperforated mica plate. The results
are given in Table V, in percentage of response by turning from _
the first-named medium in which the spores were sown to the con-
trasted one.
There is a very striking uniformity in the percentages, and this
1906] FULTON—CHEMOTROPISM OF FUNGI 95
TABLE V
ne =<
a Ho < &
B 5 e a| < F B : E
CONTRASTED CULTURES IN SILICA JELLY “u& Bog 4B ao #65
opegen waa: 2 at ER
Oe |aea Ze ae Ze
pa 3 on 5 om
a A a a
Full nutrient : full nutrient .....2...7........ 35 32 32 33 20
acking glucose : aesiie pete ce Poe 32 36 37 3° 33
Lacking glucose : containing glucose........... 27 3° 34 oO 26
Lacking glucose : full nutrient................ 3 36 33 3° 3°
Lacking KH,PO, : lacking KH,PO,.......... 92.09 40 Sy On ae Ae
Lackin aPO, de pein me PO, eran 36 36 31 30 28
Lacking KH,PO, x fall watrient. toca: ewer 32 a0" 3g 36 | 28
Lacking (NH,)NO, : lackin ae “CNET Ale Saree as ee a ae Ge we 6 | 28
Lacking (NH,)NO, : ati aes )NO, eto a 32 32 33 29
Lacking (NH,)NO, e full nutiaent is. escras ere. 34 33 33 She 3?
Lacking MgSO, : lackin ng MgSO er ee 33 ar 33 a3 27
Lacking MgSO, : containing MgsO, setae ade is 39 34 33 33 29
Lacking MgSO, : full nutrient................ 29 33 32 35 3°
under conditions that would be presumed to be most favorable for
chemotropic reaction.
Tests with epidermis and cabuan jilms.—To test the effect of
physically different perforated sheets as well as effectually to repeat
the methods used by former investigators, use was made of celloidin
films which had been perforated, and of strips of epidermis of Yucca
aloifolia. This gave, with reference to physical properties, a range
from the wholly impermeable mica plates on the one hand to the
semipermeable celloidin films on the other.
The tests with epidermis were made with Monilia fructigena,
Sierigmatocystis nigra, Botrytis vulgaris, Sphaeropsis malorum, and
Mucor stolonijer, the spores of which were distributed in non-nutrient
8 per cent. gelatin above the epidermis in its final position; gelatin
layers containing 5 per cent. cane sugar, 4.5 per cent. dextrose,
0.01 per cent. copper sulfate, 0.1 per cent. oxalic acid, 0.2 per cent.
phosphoric acid, and non-nutrient gelatin, were spread below the
epidermis. Under these conditions the penetration of the stomates
or turning toward the stomates was practically zero. When no
culture medium was used, the spores being merely spread on the
under surface of the epidermis, hyphal growéh was good. A few
hyphae of each species grew through the stomates; but there was
no evident turning toward them, and in no case was there pene-
tration of more than one or two per cent. of the stomates.
96 BOTANICAL GAZETTE [FEBRUARY
In similar series with celloidin films, the turning from one gelatin
layer to another was about equal to that obtained with mica plates.
When the spores were spread on the film without a culture medium,
very few of the hyphae grew through the holes, the percentage of
turning being negligible. The hyphae in these cases were sur-
rounded by a distinct film of condensed moisture.
TESTS FOR OTHER FACTORS.
The tests thus far have failed to give evidence of the existence
of any marked chemotropism. There has been at the same time a
considerable and fairly constant turning of the hyphae from a medium
containing spores to a sterile medium, when these were separated
the one from the other by any one of several partitions. Since this
turning is apparently unaffected by the chemical relationships of
the media employed, the cause of the turning must be sought in
other factors. Two possibilities at first present themselves; the
mechanical partitions may have a thigmotropic or other influence,
or the germinating spores themselves may affect the direction of
owth.
Tests without mechanical partitions —A slight modification of
the method employed by CiarK (4) was used. A large drop of
agar, 8™™ in diameter, was placed in the center of a sufficiently
large square of glass; this drop was surrounded by four drops of
about 5™™ diameter, equidistant from the first, and with a space
of about 3™™ between each smaller drop and the larger one. Non-
nutrient agar and to per cent. sugar-beet agar were used for the
drops, and were arranged in four combinations: the central drop
was of nutrient agar and two small drops diagonally opposite each
other were of nutrient agar, the other two being non-nutrient; the
central drop was of nutrient agar, two small drops adjacent to each
other of nutrient agar, and the other two of non-nutrient agar; two
similarly arranged combinations had central drops of non-nutrient
agar. The fungi used were Monilia sitophila, Mucor stolonijer,
and Botrytis vulgaris. A few spores were sown with the platinum
‘needle at the center of the large drop in each preparation; the cover
was inverted over a Stender dish containing distilled water and was
sealed to its rim. The growth was watched until the hyphae had
grown about two-thirds of the distance from the center to the margin
1906] FULTON—CHEMOTROPISM OF FUNGI 97
of a large drop; the preparations were then opened, and with a
sterile needle the small drops were pushed up until their edges came
in contact with the larger drops. Later observations showed that
the hyphae of the three fungi grew readily from either medium into
a similar or a dissimilar medium, and with the same percentage
of turning. An equal amount of turning toward the agar drops was
observed in the case of those hyphae which had grown beyond the
bounds of the larger drops on the moist glass; whether the agar was
nutrient or non-nutrient seemed immaterial. The turning was
apparent at a considerable distance from the surface film in so large
a percentage of cases as to negative the supposition that the physical
condition of the film has an influence.
A further test without mechanical separation was made by placing
small crystals of cane sugar, copper sulfate, oxalic acid, monobasic
potassium phosphate, and ammonium nitrate in the center of layers
of non-nutrient gelatin on cover glasses. Spores of Monilia jructi-
gena, Botrytis vulgaris, Sterigmatocystis nigra, Mucor stolonijer, and
Monilia sitophila were used; in some instances they were evenly
distributed in the gelatin, in other instances the gelatin was inocu-
lated by being touched with the needle at several points varying
in distance from the crystal. In no case was there any distinct
turning toward or away from the diffusion center.
The effect of hyphae upon the direction of growth—CLARK (4)
explains his results by supposing that the fungus secretes some
substance to which the growing hyphae are negatively chemotropic.
While this hypothesis would very well explain his results, he seems
not to have made it the subject of experimental study.
It may be reasonably assumed that if a fungus is negatively chemo-
tropic to its own secretion, the stimulus to turn away from an area
containing the fungus would, in early stages of growth, be in some
degree proportional to the amount of mycelium in that area.
To determine whether the amount of mycelium has an effect,
inoculations were made with differing numbers of spores; the growth
was from non-nutrient gelatin and gelatin containing M/4 solution
of dextrose, and was toward similar as well as different media as
indicated in Table VI, where the results are given. The direction
of growth is from the first-named medium to the second. Columns
98 BOTANICAL GAZETTE [FEBRUARY
I give the average number of spores per hole, and columns II give
the percentages of turning toward the holes.
TABLE VI
S x Mucor
uatoorssis| MO Bomaceens-| “yea |: eee
CoNnTRASTED CULTURES IN GELATIN NIGRA ee ee GENA LONIFER
2 : : AVE] S4. [15 (Ol 24 | O.8) 27 || 5.0) 207) Essig
Non-nutrient : M/4 dextrose..... re hoe 26 | ae | we | er [5 gh ee eae
: $.5] 24.| 4.2| 25 | 8.3] 29 | 4.8] 26 | 5.0) 25
M/4 dextrose : M/a dextrose..... ae | ae | 2g \-au.} 2 | 47. | a | 96 | oe
M/4 dextrose : non-nutrient...... 8.0] 25 | 7.7| 26 | 6.7) 28 | 6.0] 28 | 6.3) 31
BY 35 | 7-1 36 33 37 | 1 ae
xt ; . e ; 3-01 33° | 5:0) 30 oO 3h | 4: 22 I
Non-nutrient : non-nutrient...... Bel got c6 | ad | x8 | aot ep | oe] ee
Soe 3.01 30:|. 464] 27-|-6.3| 20 |-7.0|-28-| 27.729
M/ 4 glycerin : M/4 dextrose..... ag ely oe er ers re ee 3% pf
i Kine: Ss 0rad t-§. 3) 28%) 6.3) 28° | 5.0) 27) 83
Non-nutrient : 0.01% CuSO,.... sod an bas ae|ar | ga} 2) oe ee
There is indication that the number of hyphae in a given area
and the amount of turning from that area are correlated. It may
be said of this, as of succeeding tests, that it is at best only relative.
It is manifestly impossible to eliminate growing hyphae from the
experiments, and their effect is the very factor to be tested.
A test was made by comparing preparations in which only one
layer of gelatin was inoculated, with preparations in which both
layers were inoculated. Two parallel series were prepared: in one
the spores in the lower layer were about twice as numerous as in the
other series. An examination of Table VII, which gives the results,
shows that the percentage of turning toward a layer containing
hyphae is less than toward a sterile layer; and in either case the
abundance of mycelium in the layer influences directly the turning
TABLE VII
Montnta STERIGMATOCYSTIS | Mtcor sToLo-
NiORA ahies Mucor MUCEDO
I II Til I II Itt I I Ii I II Ill I Il ul
6.5 | 0. |: 26:4 6.6) ot 92 beet Ot ee) are oh ge o | 28
taj 0: | 35. .45 | @y go | tA} oO 1 ae 8 4540) oe) ee te
4-5} 7 | 9615.01) 8 | os 14.51: > 4 36-1 47 | 45-1} 98 @ | 10 4 me
9 G4 oa) 19 8 ae be tS 24-0. 30.135 1.40 fF 29
1906] FULTON—CHEMOTROPISM OF FUNGI 99
of the hyphae from that layer. Column I gives the average number
of hyphae per hole in the layer from which the turning takes place;
column II, the average number of hyphae per hole in the layer toward
which turning takes place; column III, the percentage of turning.
To determine whether this negative turning is due largely to
chemical changes in the culture medium, beet decoction in which a
fungus had grown was tested with that fungus. The sugar-beet
decoction was diluted to one-third the strength of the stock solution.
When a good amount of mycelium had been formed, the liquid was
filtered under sterile conditions at room temperature. The filtrate
was replaced in the incubator in the case of those species which had
fruited, and was allowed to remain for twenty-four hours in order
that any spores that had passed through the filter might germinate;
it was then refiltered. In this way a practically sterile medium
was obtained. One volume of this decoction was added to one
volume of double-strength gelatin, and tests were made with this
TABLE VIII
* a B
GrowTH TOWARD GELA-} GROWTH TOWARD GELA-
tin Mape UP witH Tin Mane Up witH
Bret DecocTIon IN _| FrEsH Beet DECOCTION.
Wuicu Founeus Hap RO
Grown
I Il I It
1, Doteytls vulgadis. 6.5002 si: 6.2 20 5-3 33
; 15 3% 15 35
II. Sterigmatocystis nigra............ 10 29 g-o 34
19 27 17 37
IiI. Penicillium MAUCUE ys 23a heen 8.0 23 12 3r
18 22 28 23
IV. Monilia sitophila........0.2+...- 8.0 22 9-0 26
: 23 25 20 33
¥.. Mucor-stolonifer. >... 2... 0. 33s 19 -32 22 38
VI enricus fabaceus.. < 5c ys neces 49 24 50 32
VIt. Coprinus micaceus.............- 31 24 43 30
VIII. Daedalia quercina............+-- wo 468 35 31
in comparison with control beet decoction in which there had been
no fungous growth. The results are shown in Table VIII. Column
I gives the average number of hyphae per hole; column II, the
percentage of turning toward the holes.
. The percentages for all of the fungi in this table, except Mucor
100 BOTANICAL GAZETTE [FEBRUARY
stolonifer, have been computed from four distinct series. In this
way erratic results have been nullified, and the difference in effect,
although not marked, may be regarded as constant. There is
certainly a lessened attractive influence in the case of the decoction
in which the fungi have grown; this might be due to the mere abstrac-
tion from the decoction of nutrient substances, or to the conversion
of compounds occurring in the decoction into compounds which are
repellent in their effect, or to the secretion of products by the fungus
which have a repellent effect. That the first is probably not the
case is to be inferred from previous tests, in which these fungi have
been found to grow as readily toward distilled water and other non-
nutrients as toward nutrients; a mere decrease, therefore, in the
amount of available nutriment could hardly have a pronounced
effect. We must conclude, therefore, that a medium in which a
fungus has grown may become less attractive, or more repellent,
to the fungus through the agency of some undetermined substance
or substances, which are secreted or otherwise formed by the growing
fungus; this reaction would -be a special kind of negative chemo-
tropism.
Miss FERGUSON (13) found that germinating spores of Agaricus
campestris, or bits of older growing mycelia, have a very marked
effect in causing the germination of spores of this species; at the
same time there seems to be a retarding effect upon the growth of
the protruded germ-mycelium. Mycelium that is not growing,
masses of ungerminated spores, or growing mycelia of other fungi
do not have the same influence over germination. Her observations
lead her to suppose that oxygen or carbon dioxid is not the deter-
mining factor, but that some secretion is formed which stimulates
or makes possible the emission of the germ-tubes. Other observa-
tions relative to the influence of germinating spores upon the growth
of fungi have been made by KrHLMAN (16) and REINHARDT (28);
to these reference has already been made. :
Numerous instances have been recorded of the influence by
various plant cells upon the direction of movement and of growth
of other cells of the same or a different kind, and the general terms
cytotaxis and cytotropism have been applied to this peculiar sort
of chemical influence. In the cases enumerated by PFEFFER (27,
#
1906] FULTON—CHEMOTROPISM OF FUNGI IOI
sec. 155), the effects seem to be due either to the excretion of a hypo-
thetical specific substance which furnishes the stimulus, or to changes
in the relative proportions of oxygen and carbon dioxid through
respiratory or photosynthetic activities. Since these phenomena
seem to be analogous in a general way to the above-described turning
of fungous hyphae, and since the term cytotropism indicates nothing
as to the exact nature of the ultimate stimulus-substance, this term
might be found a convenient one for designating the present cases.
The effect of moisture-—The circumstance that a number of
preparations in which the culture medium had become evidently
dry, gave large percentages of turning, suggested that the hyphae
might react to a hydrotropic stimulus. Layers of agar containing
spores of several fungi were spread on cover glasses, and sterile
strips of filter paper were placed in contact with the agar and allowed
_ to dip into the water of the Van Tieghem cell. The average per-
centage of turning toward the strips for those spores within a distance
of 0.35™™ from their edges was 4o per cent. for Penicillium glaucum;
50 per cent. for Mucor stolonijer, Monilia sitophila, and Sterigmato-
cystis nigra; 55 per cent. for Mucor mucedo; 57 per cent. for Botrytis
vulgaris; and 60 per cent. for Monilia jructigena and Sphaeropsis
malorum.
The percentages given are less than they would be if a smaller
area about each strip had been considered; this may be due in part
to the circumstance that the spores were rather thickly sown, and
the hyphae from those nearest the strip, being numerous, exerted
an effective repellent influence on those more distant, causing them
to grow away from the strip. Notwithstanding this, the evidence
of positive hydrotropism for these fungi was quite conclusive.
As a further test, mica plates were cut to fit Van Tieghem cells
and were perforated; a drop of non-nutrient gelatin was placed on
each and covered with a perforated plate small enough to fit inside
the cell; upon this was placed another layer of non-nutrient jelly
containing the spores; another perforated plate was added, and a
third layer of gelatin, sterile like the first; the lower surface of this
was left uncovered. The mica cover was inverted over a cell con- ’
taining water, during the time required for the proper growth of
the fungi, evaporation took place from the now uppermost layer to
s
102 BOTANICAL GAZETTE [FEBRUARY
the surrounding atmosphere, as was apparent from the dry condition
of the gelatin around the holes in the uppermost plate; water diffused
from the lower layers to supply the deficiency. In this way it came
about that the middle layer, which contained the spores, was moister
than the uppermost layer, but drier than the lowermost. There was
observed a very decided turning of the hyphae toward and through
the openings in the third plate, which separated the middle and
lowermost layers, while comparatively few grew toward the upper-
most layer. The estimated percentages are shown in Table IX.
TABLE IX
=
< 4s = a “| a
< n 2
S8/S2/$2)Ee| 88/68) 38| 8 ge
DrrEcTIONs PO|Bo|s4|/eea1f5/2#/8e|82)| 28
z Ze|°o iH < blagipYipdl ge
S0).09)3n/80) 82) a2 seisei se
AB\SE/BR/RB| Ae/Be |" 4/28 | BA
& = ae > a a a) ey
From middle layer to lowermost layer..| 43 | 70 | 80 | 55 | 68 | 65 | 73 | 63 | 65
From middle layer to uppermost layer..| 20 | 20 | 20 | 30 | 18 | 15 | 18 | 1 18
Experiments were set up in which very firm gelatin (16 per cent.)
containing the spores was covered with mica plates having a few
perforations. The plates were sealed to the covers by an application
of vaseline around their margins.. The covers were then inverted
over stender dishes level full of sterile distilled water. In this way
the water came in contact with the gelatin only through the perfora-
tions, and diffused from these through the gelatin layer. Hyphae
of Mucor stolonifer grown under these conditions showed a tendency
to grow toward the openings from a distance of 1.5™™, but on coming
within 0.5™™ of the openings, the course was changed, and the
hyphae circled the openings in lines more or less concentric with
their margins. The majority of those nearer the openings than
0.5™™ grew in a radial direction away from them. Ina few instances
hyphae grew into the water. Mucor mucedo showed a quite decided
turning toward the holes, and about 65 per cent of the hyphae within
a radius of three hole-diameters turned through them and grew into
the water. With Botrytis vulgaris about 40 per cent., and with
Penicillium glaucum 85 per cent. of the hyphae within a correspond-
ing area were affected positively. In every case the growth
—EEO
1906] FULTON—CHEMOTROPISM OF FUNGI - 103
in the water was in all directions, directly downward, as well as
radially in a horizontal plane. The value of the control cultures,
which were duplicates in all respects excepting that the dishes were
only partly filled with water, was vitiated by the accumulation of
condensed moisture in comparatively large drops about the openings
in the plates. This caused an unmistakable turning toward the
holes, which was not so decided, however, as in the test cultures.
It is evident from these results that all of the fungi tested in this
regard are, under the conditions of experiment, positively hydro-
tropic; but Mucor stolonifer may under certain conditions show a
negative hydrotropism. This response to a hydrotropic stimulus
probably accounts in large measure for the constant turning toward
protected layers from those more exposed, which latter may have
become drier through evaporation.
A sharp distinction between bedwieeneis and Pacareries on
the one hand, or between hydrotropism and osmotropism on the ’
other, cannot in all cases be made, although these phenomena in
typical cases are quite distinct. The phenomena here reported are
probably due primarily to differences in the moisture content of
the layers, and not to water currents, either molar or molecular.
For this reason the term hydrotropism has been applied, which is not
in agreement, however, with the current view (PFEFFER, 27, p. 592),
that in the case of the fungous mycelia heretofore studied, osmo-
tropism and rheotropism, but not hydrotropism, have been estab-
lished. It is further recognized that chemical rather than other
properties of water furnish the effective stimulus, in which event
hydrotropism would properly be regarded as a special kind of chemo-
tropism. :
Aerotropism.—Under the conditions prevailing in some of the
experiments above described, there was doubtless an inadequate
supply of oxygen, as when a medium poor in oxygen was enclosed
between impervious plates. There was then a very decided tendency
for the hyphae to turn toward the edges of the plates. The obser-
vation of this phenomenon from time to time suggested that the
fungi might show an aerotropic sensibility, either as a positive
reaction to oxygen, or as a negative reaction to carbon dioxid.
In order definitely to test the matter, experiments were arranged
104 BOTANICAL GAZETTE [FEBRUARY
in which the growth toward holes in mica plates could be observed
when the plates separated normal non-nutrient gelatin from non-
nutrient gelatin saturated with carbon dioxid on the one hand, and
from normal non-nutrient gelatin on the other. The fungi used
were Penicillium glaucum, Sterigmatocystis nigra, Mucor mucedo,
Botrytis vulgaris, Monilia fructigena, Monilia sitophila, and Phy-
comyces nitens. In no case was the percentage of turning toward
the carbon dioxid gelatin greatly different from that toward the
_control gelatin.
As a further test, a layer of gelatin containing spores was placed
below a perforated mica cover for a Van Tieghem cell, and a per-
forated mica plate, small enough to fit inside the cell, was placed
below the gelatin. A layer of normal gelatin was spread below this
last plate, and a layer of carbon dioxid gelatin above the cover.
This preparation was sealed to the cell rim, and the whole placed
* under a bell jar practically filled with carbon dioxid and kept at
a room temperature of 21-24° C. Efforts were made to have the
moisture conditions equal within and without the cells; and the
exposed gelatin layers, which served very well as indicators, showed
no difference in this respect until after the observations on the majority
of the preparations had been made, although there was drying of
the outer gelatin layer by the time the more slowly growing fungi
had reached the proper stage. The same fungi were used in this
experiment as in the preceding one, with the addition of Mucor
stolonifer. In most instances the turning toward the gelatin con-
taining carbon dioxid and exposed to an atmosphere of carbon
dioxid, was as great as toward normal gelatin; the growth, however,
was less vigorous in the former case. In those preparations in
which there was less turning toward the carbon dioxid gelatin, this
gelatin had become evidently rather dry.
It is to be concluded, therefore, that the observed turning toward
the edges of preparations is not due primarily to aerotropic sensi-
bility. The experiments also negative the supposition that the
observed repellent influence of growing hyphae may be due to the |
consumption of oxygen or to the production of carbon dioxid by
the fungus, or to both.
oO Z rah Al ck
&
tt RE Fe Te he
g the fact that osmotropism is
1906] FULTON—CHEMOTROPISM OF FUNGI 105
intimately associated with chemotropism, and that many of the tests
for the latter are in equal measure tests for the former, direct tests
were made by growing the fungi in media of higher osmotic pressure
and of lower osmotic pressure than the test media, as well as in an
isosmotic medium; glycerin, a good nutrient substance, and yet a
substance reported by Mryosut to be neutral in its chemotropic
effect, was used to give the desired concentration to the culture
media. The series failed to show that the concentration of the
culture medium has an effect upon the amount of turning. How-
ever, no excessively high concentrations of mineral salts were used.
Other tropic phenomena.—Under conditions that would favor a
manifestation of geotropism and of thigmotropism, there was no
indication that these are concerned in determining the direction of
growth of the fungi. The effects of light and of heat were in no
way tested, but they probably do not enter as factors.
Biological significance—The conclusions reached in these studies
may be found to have a somewhat important bearing upon the
biological problem of infection by parasitic fungi. In the absence
of any experimental investigation, nothing definite can now be said.
It would seem, however, that the drying of dew and other surface
moisture in which spores had gerimnated, might be a condition
favoring the hydrotropic turning of the germ-tubes toward the
stomates, especially if the cells within are over-distended with water,
which has frequently been observed to be a condition favorable
for infection; if the germ-tubes are numerous in the vicinity of a
stomate, the repellent influence of these upon one another would
Cause some to seek the unoccupied region within the stomate. At
all events the phenomenon of the entrance of germ-tubes, whether
by way of the stoma or through the cuticle, is a complex one, of
which many factors remain undetermined. That mere entrance is
probably not due to specific peculiarities, either of host or parasite,
is evidenced by the recent work of Miss Grsson which has been
mentioned by MarsHatt Warp (33). Miss Grsson found that
Spores of various members of the Uredineae sent their germ-tubes
readily into the stomates of plants widely different from their hosts,
and which they were unable to infect. MARSHALL WARD met with
numerous instances of the same phenomenon. This would indicate
106 BOTANICAL GAZETTE [FEBRUARY
that the entrance of a hypha into a stoma is merely a preliminary
act, distinct from infection proper, and controlled. by general con-
ditions, while the fate of the hypha after its entrance is determined
by complex reactions between parasite and host, which are largely
specific in their nature. In the light, then, of known facts, no simple
explanation, such as the theory of chemotropism due to the presence
of specific chemical compounds, is adequate. Chemotropism may
possibly be one factor in the complex phenomenon, but it is certainly
not the predominant factor.
CONCLUSIONS.
Various tests upon a number of fungi failed to indicate the exist-
ence of any definite chemotropic sensibility to nutrient substances
or other chemical compounds in solution. If positive chemotropism
exists, it is less prominent than other tropic phenomena involved,
and was obscured by them.
Those substances which furnished nutriment to the fungi caused
a decided growth, often with thickening of the hyphae and an increased
branching; but they did not cause a more marked turning of the
hyphae toward the diffusion centers than did non-nutrient and
toxic substances.
All of the fungi tested show a tendency to turn from a region in
which hyphae of the same kind are growing toward one destitute
of hyphae, or in which the hyphae are less abundant. The turning
toward a medium in which mycelium has grown, but from which
the mycelium has been removed, is less marked than that toward
a medium in which no mycelium has grown. This may be regarded
as a negative reaction to stimuli from chemical substances, which
owe their origin in some way to the growing fungus.
Various fungi show a positive hydrotropism; but an over-
abundance of moisture may cause a negative reaction in certain fungi.
The changing of the direction of growth of fungous hyphae is
a complex phenomenon in which at least two factors, cytotropism
and hydrotropism, are concerned. Since the complete elimination
of neither of these factors is possible, all tests must be relative, and
to that extent unsatisfactory.
It would seem that the reactions of mycelium to various stimuli
1906] FULTON—CHEMOTROPISM OF FUNGI 107
are not necessarily the same as the reactions, under similar conditions,
of sporangiophores, gametophores, and other specialized parts.
The writer wishes to acknowledge his indebtedness to Dr. B. M.
Duccar for his very helpful suggestion and cirticism, and to Dr.
Witt1am TRELEASE for the opportunity to consult the library of
the Missouri Botanical Gardens.
BoranicaL LABORATORY,
University of Missouri.
LITERATURE CITED.
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CrarK, J. F., On the toxic effect of deleterious agents on the germination
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Now
"
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Dietz, S., Beitrige zur Kenntniss der Substratrichtung der Pflanzen.
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Unters. Bot. Inst. Tiib. 2:4 88.
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Ba Sea ee ee ee ee
BED ang a dr gS ET at Sel iar SE
THE EMBRYOLOGY AND DEVELOPMENT OF RICCIA
LUTESCENS AND RICCIA CRYSTALLINA.'
CHARLES E. LEwis.
(WITH PLATES V-IX)
In June 1903, while collecting liverworts in the vicinity of Ithaca,
N. Y., an abundance of material of Riccia lutescens was found growing
around the edges of dried-up ponds. In some cases the plants
formed beautiful rosettes, but usually they grew in irregular clusters,
often being so closely crowded together as to cover the ground for
several square centimeters.
The individual plants vary greatly in shape and size. The
younger light green plants consist of a narrow, thin, ribbon-shaped
thallus which has a longitudinal median groove. In the older plants
the fore part of the thallus is thickened, very large air cavities being
formed: The thallus is attached to the soil by numerous rhizoids
from the older part, the apical end being free. On the under side
are numerous colorless lamellae.
As the fruiting plant is unknown there is doubt as to the rela-
tionship of this species, authorities differing widely as to its status.
LINDBERG (21) claimed that it was merely a sterile terrestrial form
of Ricciocarpus natans. UNpdERWwooD (30) says of it: “approaches
certain terrestrial forms of Ricciocarpus natans, and possibly derived
from that species, but better kept distinct.”” SrEPHANI (28) states
that it is probably not a Riccia but a sterile marchantiaceous hepatic.
For the purpose of determining the true relationship of the species,
Professor ATKINSON suggested the desirability of following the
development of the plant through the summer and autumn, and
of securing fruiting specimens if possible. He had found young
antheridia in plants collected several years before, but had not traced
the development further. It also seemed desirable to study the
embryology and cytology of the plant if material could be obtained,
because comparatively little has been done on these phases of the
life history of Riccia.
* Contribution no. 106, from the Department of Botany, Cornell University.
109] [Botanical Gazette, vol. 4:
110 BOTANICAL GAZETTE [FEBRUARY
BIsCHOFF (2) investigated a number of species and settled beyond
a doubt the function of the sexual organs. His work was followed
the next year by LINDENBERG’S monograph (20) which added little
that was new.
- The study of the development of Riccia really begins with Hor-
MEISTER (15), who gave an account of the development of the thallus,
sexual organs, and fruit of Riccia glauca.
Kwy (19) made a careful study of the apical cells and the method
of growth of the thallus. He did not secure plants developing from
spores but used delicate thalli which had grown crowded together
and did not bear sexual organs in the younger parts, so that the
regular order of cells was not disturbed. He discovered the origin
and manner of growth of the ventral scales and described the develop-
ment of the sexual organs. Although HormeEIsTER believed that
young antheridia and archegonia could not be distinguished, Kny
points out that they are distinct after the first walls are formed.
LEITGEB (22) gives a complete account of the method of growth
of the thallus in the Ricciaceae. His study of the sexual organs
and fruit was in many cases incomplete on account of insufficient
material.
BIOLOGY OF RICCIA LUTESCENS.
The account of the biology of Riccia lutescens given here is based
on field observations extending through two years, together with
experiments and observations upon plants kept growing under
favorable conditions in the greenhouse and laboratory.
The first observations were made late in June. At that time
the plants were growing-upon the mud around the edges of ponds.
Some of the thalli were very small and delicate, appearing merely as
green specks on the mud, while others, which seemed to be older, had
the ribbon-shaped form and thickened apical end already described
(figs. 1-3).
Material was collected and examined from time to time during
the summer and autumn, with the expectation of finding plants
bearing the sexual organs, because the statement is usually made
that the species of Riccia fruit in summer and autumn when growing
on the soil. The plants continued to grow well vegetatively through-
out the summer, when they were in such a location that they were
1906] LEWIS—DEVELOPMENT OF RICCIA T1y
supplied with sufficient moisture. In some cases the mud became
so dry and hard that the plants were killed, but whenever they were
sheltered by a stone or other object, or were growing on the sides
of holes, such as cattle tracks, they grew well.
In October all but the youngest and most crowded plants showed
the typical Riccia lutescens form. At this time young antheridia
were found. Material was now collected and fixed from time to
time for the purpose of studying the development of the sexual
organs. In very few cases were archegonia found in plants collected
in autumn. A few young stages were found in plants collected
late in November, at which time the older antheridia were almost
mature. No further development took place out of doors until
spring, because the plants became covered with snow, or with water
by the filling up of the ponds, and remained so until April. A
quantity of the plants were kept growing on the soil in the green-
house through the winter, and developed mature sexual organs
long before spring. Plants taken from under water in March, just
as the ice was going out of the ponds, showed exactly the same form
as in November, and little or no further development had taken
place. So it seems that the development depends to some extent
on temperature, and might be expected to vary with different con-
ditions of climate. A warm winter, in which some growth might
take place, would in all probability hasten the development of the
sexual organs. Another point of interest is that the submerged
plants did not seem to have been injured.
A quantity of material still attached to the soil was taken from
under water late in March, and was kept growing in shallow pans
in the laboratory so that it could be kept supplied with a sufficient
quantity of water for growth but not enough to flood the plants.
This was done in order to determine whether the plants would con-
tinue the development of sexual organs and fruit in the same way
when supplied with a limited amount of water and growing on the
soil, as when supplied with a large amount of water which would
tend to cause them to break loose and float. It was found that the’
Plants growing on the soil did produce fruit abundantly and at
the same time as those growing under natural conditions. The
archegonia begin to develop in April in the same thalli which have
hz BOTANICAL GAZETTE [FEBRUARY
produced antheridia, and all stages are found by May 1. About
this time fertilization takes place, and by May 25 all stages of sporo-
phyte are found. The arrangement of the sexual organs in the
thallus is shown by figs. 7-11.
The vegetative growth is very rapid during April and May, the
thallus becoming broad and branched by the increase in the number
of growing points. The narrow older part by which the thallus is
attached decays, and the younger part bearing the sexual organs
and sporophytes is set free and floats upon the water. When the
plants are supplied with a large amount of water changes take
place in the lamellae. They grow to great size and become purple.
In the floating thallus decay of the older part continues; the part
bearing the antheridia first disappears, then the part bearing the
sporophytes, and finally the growing points may be separated, one
thallus thus giving rise to several new individuals. In most cases
observed the decay of the older parts in floating plants did not advance
so far. The plants were carried up around the edge of the pond
by the waves, and as the water went down were left stranded upon
the mud. When the thalli settle down upon the mud, the large
ventral plates wither, and rhizoids are put forth which in a few
days attach the thallus to the soil. Growth now continues at the
growing points, so that new branches are produced which form
rosettes.
When the thallus is injured at this time new plants are imme-
diately produced from the cells of the apical region. This was
first observed in plants injured by being covered with mud, in which
case slender delicate outgrowths were produced (fig. 4). Other
plants injured by snails soon developed long slender plants (fig. 36)-
Thalli were cut into pieces to determine whether other cells would
show the same plasticity, but new plants were produced only from
cells near the growing point. V6cHTING (33) found in Lunularia
that regeneration takes place from cells in various parts of the thallus,
but this does not seem to be true of Riccia natans under the conditions
in which I have studied it. Large numbers of the plants which
were left upon the mud when the water went down were injured
by cattle coming down to the ponds to drink. Later in the season
the cattle tracks were lined with young, green, ribbon-shaped plants
1906] - LEWIS—DEVELOPMENT OF RICCIA It3
which were outgrowths from the growing points of the older plants.
The cattle tracks serve a good purpose, as the young delicate plants
are shaded and protected to some extent during the dry season.
Two forms of the thallus are produced by the different methods of
propagation. In the one case the thallus after it becomes attached
to the soil continues its growth, branches and forms a rosette, while
in the other case the thallus is injured, and very delicate forms are
produced. When large numbers of the floating thalli are deposited
near together and are then injured, we find the irregular clusters
of plants which have been described in the first paragraph.
The thallus of this plant during the floating period bears such
a striking resemblance to Ricciocarpus natans that one is led to the
conclusion that Riccia lutescens is only a ground form of Ricciocar pus
natans. Since the beginning of this study and after it was well
under way, a paper was published by GARBER (11) which dealt.
‘with the life history of Ricciocarpus natans. Several points in the
biology of the plant as given by GARBER differ from those found
to obtain at Ithaca, and since the structure of the thallus as well
as the embryology is conclusive proof that the two forms are the
Same species, it seems proper to call attention to these differences
and then to give briefly the embryology before taking up the other
phases of the study.
The greatest difference in our observations lies in the relation
of the supply of water to sexual reproduction. GARBER states that
Ricciocarpus natans as it grows at Chicago spends its entire life,
from the germination of the spore to the production of spores, in
the floating state, and that the occasional fruiting plants found
upon the soil in summer are plants in which the sexual organs devel-
oped and the sporophytes began their development while the plants
were floating. He observed no case in which sexual organs were
produced on plants growing upon the soil and states that Ricciocar pus
natans has not yet acquired the power to reproduce sexually when
growing upon the soil. The sexual organs develop in April.
The plants at Ithaca, however, spend the greater part of their
life upon the soil and only float upon the water for a few weeks at
the fruiting period. The sexual organs begin to develop in autumn
while the plants are on the soil and plants kept on the soil and sup-
114 BOTANICAL GAZETTE [FEBRUARY
plied with a limited amount of water developed fruit. At the time
when the antheridia begin to develop the gametophyte is under
favorable conditions for vegetative growth, but is not supplied with
an abundance of water. The soil is moist and the conditions are
such as would favor the growth of a terrestrial form like Marchantia.
The plants seem especially adapted to spend the winter submerged
and do not perish under such conditions. The fore part of the
thallus contains very large air cavities and thus the tissue is aerated.
It is well known that certain higher plants which grow in wet situa-
tions have large air spaces in the tissue, and GANONG (12) calls
attention to the fact that those marsh plants which are submerged
for a portion of the year are able to survive on account of their capacity
for air storage. About May 1 the older part of the thallus, which
is narrow and thin, has decayed, and the younger parts, bearing
the sexual organs, is set free and floats. GARBER points out that
when land forms are placed upon the water only a small portion -
of the apical end remains above the surface, while the older part
of the thallus extends into the water and decays. This is true of
plants taken from the soil in summer, but in the spring when the
free part of the thallus is thick and contains large air cavities, it
floats readily. The length of the floating period depends of course
upon the conditions of the pond. In some cases the plants may
very soon be carried up around the edge of the pond and deposited
on the mud, but floating forms are usually found until the ponds
are almost dry. In the case of ponds which do not become dry
in summer, both forms would be found. The floating period affords
an excellent means for distribution.
When the plants grow upon the soil and are not protected during
the winter by a covering of snow or water, they are usually killed
by freezing, but in some cases plants which were brown and seemed
to be dead produced new thalli from the growing point. The young
delicate thalli are well adapted to tide over the dry season, because
they can live with a less supply of water than would be needed by
the older plants.
RELATIONSHIP OF THE SPECIES.
The form usually described as Riccia lutescens should be regarded
as a ground form of Ricciocarpus natans. Both in the field and in
1906] LEWIS—DEVELOPMENT OF RICCIA ars
cultures in the laboratory the forms have been observed showing
the transition. There can be no doubt that the plant which I have
described is the true Ricciocarpus natans, and the description of
the ground form as a distinct species came about naturally from
the conditions of its growth. In such ponds as have been observed
here, the water is high in April and May, so that the floating plants
are carried up around the edge and left on the soil. In June or
July the water has entirely disappeared from the pond and the only
plants found are the slender ribbon-shaped ones which have developed
from the floating form. In my first summer’s collecting, when
the ponds were dry by the last of June, I saw not a vestige of the
old Ricciocarpus natans, and felt sure that the plants collected were
Riccia lutescens. It seems possible that the plant was first described
as a distinct species from material collected under similar conditions,
because it is said to occur in dried up ponds and ditches. If in
summer and autumn some water were present, so that some of the
typical Ricciocarpus natans would be found floating, the origin of
the ground form might readily be seen, but in such a case there
might be failure to associate the ground form with Riccia lutescens.
Only by following the development and observing the transition of
one form into the other under different conditions of growth can
the true relationship be determined. My observations have con-
vinced me that Riccia lutescens is only a ground form of Ricciocar pus
natans and should not be regarded as a distinct species.’
The plant now known as Ricciocarpus natans was formerly
‘regarded as a Riccia. In the structure of the thallus Ricciocarpus
is more complicated than the species of Riccia. The most important
taxonomic characters, however, have been the arrangement of the
sexual organs and structure of the sporophyte.
Hooker first found fruiting plants in dry specimens sent to him
by Torrey from New York in 1824. BrscHorr found fruiting
plants in the autumn of 1829 near Heidelberg, and describes anthe-
ridial plants, but his figure of the antheridium is not very convincing,
? Having determined the ground form as Riccia lutescens, specimens were sent to
Professor A. W. EVANS in October 1904. He considered that we were right in refer-
ring the plants to that species, but stated the views of different authorities in regard
to the status of the species.
116 . BOTANICAL GAZETTE [FEBRUARY
as it looks more like the mass of tissue which projects up as a ridge
into the median groove, the cells being quite too large for those of
an antheridium.
Although Hooker considered that the plant should remain in
the genus Riccia, Corpa placed it in a new genus, Ricciocarpus,
on the basis of Hooxker’s description and figures which were taken
from dried material. Corpa’s figures are copies of HOOKER’S.
BiscHorrF held that there was no real basis for the change, as the
mature sporophyte does not differ from that of other Riccias, the
separation being based on the mistaken notion that the capsule
walls disappear entirely at maturity, and that the genus Riccia
should not be divided on account of differences in the thallus brought
about by the different conditions under which the plant grows, since
the method of fruiting is the same in all the species.
LEITGEB regarded Ricciocarpus as a distinct genus, on account
of the more complex structure of the thallus and the grouping of
the sexual organs. He thought that the antheridia were collected
into groups similar to those in the Marchantiaceae, but GARBER’S
results and my own show that LerrcEB was not correct, and that
the antheridia actually form only one group. The archegonia are
also arranged in a definite part of the plant in one group.
The question now arises whether this is a more advanced con-
dition of development than is found in species of Riccia. In the
lower species of Riccia, the sexual organs are said to be indiscrimi-
nately scattered over the surface of the thallus, while in Riccia fluitans
a regular alternation of single antheridia and archegonia is described.”
CAMPBELL, in discussing the arrangement of sex organs in Riccia,
says that in the two forms which he studied, Riccia hirta and Riccia
glauca, he found as a rule that several of one sort or the other would
be formed in succession. I have observed the same in Riccia crystal-
lina, although the older sporophytes appear scattered in the thallus.
LINDENBERG described the fruit of Riccia crystallina as scattered,
but the antheridia are described and figured as being in a group
along the middle part of the thallus. He described and figured
the fruit in Riccia glauca as being sometimes in rows and sometimes
scattered. Most of the figures show them in more or less perfect
rows along the longitudinal axis.
1906] | LEWIS—DEVELOPMENT OF RICCIA 1i7
In Riccia minima, LINDENBERG (20, p. 429) describes and in
pl. 20 figures the antheridia as arranged in two rows, one on each
side of a median groove. In Riccia bulbosa the antheridia are along
the median groove for its entire length, sometimes in pairs and
sometimes far apart. Riccia Bischoffiit has the antheridia in two
or three rows in the thallus.
It seems highly probable that a careful study of a large number
of species of Riccia by modern methods would show that in many
of them there are produced groups of antheridia and archegonia
in distinct parts of the thallus.
Since the characters upon which the genus Ricciocarpus has
_ been based, with the single exception of the structure of the thallus,
have been found wanting, it seems to me that there is not sufficient
reason for retaining the genus. The thallus varies in form according
to the supply of water, and when growing on the soil has been called
a species of Riccia. Many plants assume quite different forms
when growing under different conditions, but the different forms
are not regarded as species. |
We should then write:
RICcIA NATANS L. Syst. Veget. 956. 1774.—Bischoff, Nova Acta
Acad. Caes. Leop. Carol. 17: 2. 1835.—Lindenberg, Nova Acta
Acad. Caes. Leop. Carol. 18: ease —Sullivant, Gray’s Manual
2ed. 1856.
Ricciocarpus natans Corda, Opiz Naturalischentausch. 1829.—Leitgeb, Die
Riccien, Unters. Lebermoose 4:1879.—Lindberg, Revue Bryol. 9:82. 1882.
(Includes Riccia natans L. and Riccia lutescens Schw.)—Schiffner, Engler and
Prantl. 1893.—Campbell, Mosses and Ferns. 1895.—Underwood, Systematic
Botany of oe America. Hepaticae. 1895.—Garber, Bot. GAZETTE 37:101-
177. pls. 9-10. 1904.
Riccia dso Schw. Specimen Fl. Amer. Sept. Crypt. 26. 1821.—Linden-
berg, Nova Acta Acad. Caes. Leop. Carol. 18: pl. 26. 1836.—Sullivant, Mem.
Amer. Acad. II. 4: pl. 4. 1849.—Sullivant, 2d ed. Gray’s Manual 684. 1856.
—Underwood, Systematic Botany of North America, Hepaticae. 1895.
Riccia velutina Hooker (in part) Ic. Pl. pl. 149: founded on sterlile thalli of
Riccia lutescens and fertile thalli of Riccia crystalline, —s to Sullivant,
Gray’s Manual, 1856.
EMBRYOLOGY.
Material for study was collected during the autumn and spring,
and fixed very satisfactorily in 1 per cent. chromacetic acid or in
chromosmacetic.
118 BOTANICAL GAZETTE [FEBRUARY
The large air cavities prevent the penetration of the fixing fluid,
to overcome which the pieces were submerged by means of cotton
plugs. After dehydration the material was passed through chloroform —
into paraffin. Sections were stained with the triple stain of Flem-
ming or with Heidenhain’s iron-alum haematoxylin.
SEXUAL ORGANS.
Young antheridia were found in October. They begin to develop
while the plants are young and growing on soil not supplied with a
large quantity of water, although the conditions for vegetative growth
are good. At this time the thallus is ribbon-shaped, with a thick-
ened apical end and a longitudinal median groove, the thallus in
cross-section having about the shape of an inverted Y with a ridge
of tissue between the arms (fig. 9). Very few plants are found
which do not produce antheridia. The archegonia develop later
in the same thallus. At first there seemed to be in this a distinction
between Riccia lutescens and Ricciocarpus natans, because Ricctocar-
pus natans has been described by SCHIFFNER, LEITGEB, and CAMpP-
BELL as being strictly dioecious, but the work of GARBER shows
conclusively that it is monoecious. The earlier observers state that
Ricciocarpus fruits in autumn, so it seems probable that their material
was collected after the older portion of the plant had decayed, leaving
only the portion bearing sporophytes.
The antheridia are produced in acropetal succession in three to
five rows (figs. 10, IT).
The antheridium develops as has been described for other species
of Riccia. A superficial cell on the floor of the dorsal furrow just
back of an apical cell protrudes above the surface and is cut off by
a horizontal wall. The outer cell increases in size, and is divided
by three or four cross walls, then a longitudinal wall is formed divid-
ing the young antheridium into two equal parts: this is followed
by a second longitudinal wall perpendicular to the first. Then
periclinal walls are formed which cut off the single layer of cells
which form the wall of the. antheridium. The cells in the center
now undergo repeated divisions until a very large number of cells
is formed. Each of these cells is almost. cubical in form and in
Riccia has been described as producing a single spermatozoid,
1906] LEWIS—DEVELOPMENT OF RICCIA 119
Kwy (19). The mature antheridium’ is a short stalked oval body
with a conical apex.
As the antheridium develops, the vegetative tissue grows up and
surrounds it so that it is enclosed in a cavity which opens into the
dorsal furrow. This cavity is formed in the same way as the air
spaces of the thallus. The apex of the antheridium is a little below
the floor of the dorsal furrow and the sperms escape through the
neck formed by the surrounding tissue. Although the antheridia
begin to develop in autumn, they are not mature until the following
spring, because the growth is checked by the cold. Plants kept in
a warm place produced mature antheridia during the winter.
A series of archegonia is developed which is a continuation of
the series of antheridia (fig. 7). The archegonium is at first super-
ficial on the floor of the dorsal furrow. Later it becomes enclosed
in a cavity by the upward growth of the vegetative tissue as in the
case of the antheridium except that the neck of the mature arche-
gonium protrudes above the bottom of the furrow. The origin of
the archegonia side by side at the bottom of the dorsal groove is
Shown in figure 9. In this way three to five rows are formed and
later a large number of sporophytes are found in each thallus.
The archegonium develops in general as has been described by
JANCZEWSKI (17). My observations confirm the account given by
GARBER for Ricciocarpus natans, as a comparison of the figures
will show, so it is unnecessary to describe the development here.
About the time when the archegonia are mature, cross-sections
of the thallus show numerous, delicate, almost hyaline, club-shaped
hairs extending up from the floor of the median groove. Each
hair consists of a stalk of two or three short, narrow cells with a
much larger cell at the free end. These hairs bear a striking resem-
blance to paraphyses(jig. 78). LEITGEB (22, p.31) describes ‘‘papillae’”’
which grow up from the bottom of the groove and regards it as
highly probable that it was the dried remnants of these which LINDEN-
BERG observed when he wrote: “Sporangium vor aussen mit kleinen
unregelmissigen braunen Schuppen bedeckt ist, die Fragmente
einer zersprengten friiheren Hiille zu sein scheinen.”” As the hairs
become older they become brown and break down so that they
120 BOTANICAL GAZETTE [FEBRUARY
would give much the appearance described by LINDENBERG. We
know now that the sporophyte has no Hiille or sheath.
SPOROPHYTE.
The development of the sporophyte agrees with the account given
by CAMPBELL (3) for Riccia and by GARBER for Ricciocarpus natans.
The first division is usually transverse but may be oblique (jig. 21).
The next wall may be perpendicular to the first so as to form a quad-
rant (fig. 22), or parallel to it, producing a row of cells. Divisions
take place in all directions after this until an almost spherical mass
containing several cells is formed. Then the amphithecium becomes
distinct. as a single layer of cells enclosing the spore producing cells.
The growth takes place rapidly but the divisions of the cells are not
simultaneous, usually only a few dividing cells being found in a
sporophyte. 7
The sporophyte continues its growth until a solid mass of three
to four hundred cells is produced. Then the calyptra and amphi-
thecium expand and the spore mother-cells becoming free separate
from one another and become rounded. From the surrounding
cells, which are richly stored with food, there is secreted a large
amount of nutritive material which fills the space around the mother-
cells, giving them favorable conditions for growth (figs. 25, 206).
The spore mother-cells increase rapidly in size and again fill the
cavity. That part of the nutritive material not absorbed by the
spore mother-cells is pressed into thin plates between them. This
material takes .a deep blue stain with gentian violet. A fuller dis-
cussion of the spore mother-cells and of their division to produce
the spores will be given in another place.
Before the spores are mature the inner layer of the calyptra
collapses. The amphithecium is distinguishable until the spores
are almost mature. The outer layer persists but the cells are usually
shrunken. The contents of these cells is no doubt used up to supply
the growing spores with nourishment. All of the spore mother-
cells produce spores, there being no sterile tissue in the form of elaters.
In discussing the simple form of sporophyte of Riccia, GARBER
considers that the absence of sterile tissue is to be associated with
the habit of the plants; since there is not much chance for the attach-
ment of an independent sporophyte, there is no sterile tissue in
1906] LEWIS—DEVELOPMENT OF RICCIA
the form of a foot. When we consider the fact that some other
Hepaticae which have the foot well developed grow on very wet
-soil and require as much moisture for their development as do some
of the species of Riccia, this theory does not seem entirely convincing.
The sporophyte develops during May and June. A given sporo-
phyte requires about three weeks for its development.
SPOROGENESIS.
Usually the most favorable cells for the study of cytological details
are the spore mother-cells. Their large size, abundant contents
and active growth at the time when divisions are taking place, permit
good results in fixation. Riccia crystallina has furnished the most
satisfactory material.
In July, 1903, an abundance of fruiting Riccia crystallina was
found growing on the mud on the bottom of a dried up pond not
far from the ponds where the form known as Riccia lutescens was
growing. This species had never been collected in this region
before. Having so determined the plant, I referred specimens to
Prof. A. W. Evans who confirmed my determination. He says:
“Apparently this species represents an addition to the hepatic flora
of New York. I find no mention of it in local lists of New York
plants and there are no specimens of it from your state in my
herbarium.” » |
These plants had been growing under favorable conditions, as
the pond had not contained much water at any time during the
spring. The thalli formed rosettes growing so close together as almost
to cover the ground. The number of fruiting plants was very
striking, as it seemed impossible to find a single sterile plant. All
Stages in the development of the sporogonium and spores were
easily obtained, and some stages in the development of the sexual
organs, but changes were taking place very rapidly and the younger
Stages were of comparatively rare occurrence. The development
of the sexual organs and fruit agrees with that of other species of
Riccia. Each thallus produces several sporophytes which are easily
recognized when mature as small black spherical bodies imbedded
In the tissue.
These plants continued.to develop and produce sporophytes for
only a short time after they were discovered. The month of July
522 BOTANICAL GAZETTE [FEBRUARY
was the most favorable time for the collection of material showing
karyokinesis in the spore mother-cells. During August, the spores
became mature and the thalli broke down. No good specimens
could be collected after August 25. This differs from what has
been observed for some other species of Riccia, which are described
as withstanding long periods of drought, the thalli continuing their
growth again when supplied with moisture. (CAMPBELL, 3.)
During the following winter this pond became filled with water
and did not become dry until late in the summer, so that only a few
plants were found as compared with the large number of the pre-
ceding year. This made a difference in the time of fruiting. In
September the sporophytes were in about the same stage of develop-
ment as in July of the preceding year. This may explain why
different authors give different seasons for the fruiting of Riccia.
It seems that conditions of temperature and water supply exert such
an influence that in the same species and locality the time may vary
considerably from year to year. In general, I think it may be said
that good conditions for vegetative growth will hasten rather than
retard the fruiting of Riccia.
The thallus of Riccia crystallina is small and thin; its surface
presents a series of wide depressions separated by thin lamellae;
and there are no ventral scales. The fixing fluid easily penetrates
and the spore mother-cells are usually well fixed.
The development of the spore mother-cells agrees with the account
given for Riccia natans, but there is not such a large number produced
in a sporogonium. When the spore mother-cells come to lie loosely
in the sporogonium, they are surrounded by nutritive material.
The mature spore mother-cells are then generally spherical, but
they may be elliptical or so angular by crowding as to look like a
tissue. The contents of the spore mother-cells of Riccia has been
described as granular by CAMPBELL (3) but the structure of the
cytoplasm in Riccia crystallina is a fine reticulum with the granules
occurring usually at points of intersection of the fine threads of the
network. The older spore mother-cells as well as the mature spores
contain considerable oil.
In the nucleus of the spore mother-cell the chromatin is scanty
and is irregularly scattered on a fine linin network. No nucleolus
1906] LEWIS—DEVELOPMENT OF RICCIA 123
has ever been observed (fig. 34). When the nucleus is preparing
for division, the chromatin leaves the linin network and collects into
several bodies which soon move together to form one irregular mass.
I regard this as the synapsis stage. Such bodies of chromatin have
been found often and in cells which seemed to be well fixed so it
Seems to represent a stage in the preparation for division and not
to be a result of shrinkage as has been suggested by certain authors
for other plant cells in which the same condition has been ‘observed.
The body of chromatin occupies a position at one side of the nucleus,
and the rather large nuclear cavity appears hyaline. There can
be little doubt that the body described by CAMPBELL (3) as a nucleolus
is really the’entire mass of chromatin in the synapsis stage.
From this mass of chromatin a short thread develops which later
Segments to produce the chromosomes (jig. 35). The small amount
of chromatin present here makes the details very difficult to deter-
mine. The four chromosomes, easily counted here as well as in
the nuclear plate and on the way to the poles, are very small and
appear almost spherical when on the spindle although they are
short, thick, curved rods.
The development of the spindle is not easily observed. Divisions
take place almost simultaneously in all the cells of a sporogonium
and the changes are very rapid. By far the commonest stage of
division is that in which the chromosomes are in the nuclear plate
(fig. 38). Neither centrosomes nor centrospheres occur in the spore
mother-cells of either Riccia crystallina or Riccia natans. Around
the nucleus preceding the formation of the spindle, there is an accu-
mulation of material, apparently composed of fine fibres. The
nucleus elongates, becoming somewhat elliptical but not sharp.
pointed. The fibres about the nucleus do not give the appearance
of centrospheres but are like the weft of kinoplasmic fibres described
for certain pollen mother-cells (jigs. 36, 37). It has been impossible
to find any nucleus which showed anything resembling a multipolar
spindle. The poles of the spindle are probably determined by the
elongation of the nucleus at an early stage in the spindle formation.
The spindle is composed of very fine fibres, some of which extend
from pole to pole, while others extend from the poles into the
cytoplasm, reaching almost to the nuclear plate (fig. 38). The
124 BOTANICAL GAZETTE [FEBRUARY
mature spindle has very broad poles and its formation does not
seem to have been controlled by a centrosome or a centrosphere,
as a comparison of the spindles of the spore mother-cells with those
of the cells of the antheridium makes clear.
The minute chromosomes separate, four going to each pole, after
which a cell plate is formed in the usual way (figs. go, 41). The
daughter nuclei do not come to a true resting stage. The chromatin
is scattered in almost spherical bodies in the hyaline cavity of the
nucleus, which do not represent the individual chromosomes, as
their number and size vary considerably (jig. 42).
The second division takes place in much the same manner as
the first. The spindles are arranged with their long axes parallel
to the first cell plate, so that the cell plates formed in these spindles
are almost perpendicular to that formed in the first division (figs.
43-47). The latter does not disappear during the second division
but remains and the walls separating the spores are laid down here
(fig. 47). The walls separating the cells of the young tetrad are
thin and delicate, but in the mature spore the outer layer of the
wall becomes thickened and folded. The mature spore is almost
black, and its contents are largely oil. When carried through chloro-
form into paraffin and sectioned, the spores seem to have only scanty
. granular contents, due to the fact that the oil has been removed in
the process. The nucleus is very small.
During the winter and spring following their development, unsuc-
cessful attempts were made to germinate the spores. It may be
that they had been allowed to remain dry too long before they were
moistened, for in nature they would not be dry very long even in
tiding over a dry season. ;
The spore mother-cells of Riccia natans do not furnish such
satisfactory material for study as do those of Riccia crystallina,
because-the large air cavities of the thallus prevent the penetration
of the fixing fluid and so the spore mother-cells often shrunk.
sufficient number of good preparations was secured, however, to
show that the process of division does not differ from that of Riccia
crystallina.
|
1906] LEWIS—DEVELOPMENT OF RICCIA 125
SPERMATOGENESIS.
The development of the spermatozoids has been treated by a
number of investigators, among them, CAMPBELL (4), LECLERC DU
SABLON (23), GUIGNARD (14), SCHOTTLANDER (27), and StTRas-
BURGER (29). It will be observed that most of these papers were
published before methods of preparing material for study were so
well developed as at present. The work of BELAJEFF (1)-confirmed
by that of STRASBURGER (29) shows that the spermatozoid in the
Hepaticae consists not only of the metamorphosed nucleus but also
of the cytoplasm.
IkENo (16) not only confirms the view that the spermatozoid
consists of cytoplasm as well as nucleus but also discusses the develop-
ment of the cilia and the homology of the blepharoplast and centro-
some of Marchantia polymorpha.
He finds that the body which becomes a blepharoplast in the
developing spermatozoid appears in the earlier nuclear divisions of
the antheridium and functions as a centrosome. It is, however, not
permanent, but appears at the time of nuclear division and disappears
during the process, so that it is not found in the daughter cells until
about the time for the formation of the spindles of their division.
After the last division which gives rise to the cells that develop into
the spermatozoids, the body does not disappear but remains and
becomes a blepharoplast. IKENo argues from this that the centro-
Some and blepharoplast are homologous. He has good grounds for
Such an argument in the case of Marchantia polymorpha, because
centrosomes have been reported also in the vegetative cells of that
plant, by Morrrer (24) and by VAN Hook (32). In other plants
which have the blepharoplast, centrosomes are not found, and the
body appears in only one or two‘divisions before the formation of
the cells which produce the spermatozoids.
Morrtier (26) in discussing IKENO’s Paper raises the question
whether the bodies which Ikeno has figured as are in some
Cases more than ordinary granules such as appear in the cytoplasm
of other cells in which centrosomes are known to be absent. IKENO
has pointed out, however, that the cytoplasm of these cells is very
finely granular, there being no other bodies in the cell which bear
any resemblance to the ones figured as centrosomes. He also calls
126 BOTANICAL GAZETTE [FEBRUARY
attention to the fact that centrospheres have been described in dividing
spore mother-cells of Pellia epiphylla, by FARMER (6, 7, 8, 10) and
by Davis (5). The occurrence of centrospheres here has been
questioned, however, by GREGOIRE (13). In a recent paper, FAR-
MER (Q) reports centrospheres and occasional centrosomes in the
spore mother-cells of Aneura pinguts.
In order to get good results in Riccia natans it is necessary to
fix the material when growing rapidly. About equally good results
were secured with chromacetic acid and with Flemming’s weaker
solution. The sections were stained with anilin safranin and gentian
violet. It was found best to stain deeply in gentian violet and then
to wash out carefully. In this way all details can be brought out
clearly, although IkENo did not find it good for Marchantia.
The development of the antheridium has been described. When
almost mature it consists of a large central mass of cubical cells
surrounded by a wall one cell in thickness (fig. 33). In preparations
from plants in which some antheridia are mature, one finds several
stages in the development. The nuclear divisions do not take place
simultaneously throughout an antheridium but usually all the cells
of one of the segments marked out by the first walls dividing the
antheridium, show the nuclei in the same stage of karyokinesis.
In the most favorable preparations, therefore, one may find several
stages of division in the same antheridium.
The cells of the young antheridium are almost cubical, with finely
granular cytoplasm. The nucleus is rarely exactly spherical and
has a rather thick membrane. The chromatin is in an irregular
central mass, made up of a number of pieces. A nucleolus cannot
be distinguished. The cavity surrounding the chromatin is large
and hyaline (figs. 53, 54). In some cases a large number of small
bodies of chromatin were found scattered irregularly in the nuclear
cavity. The number of chromosomes is four. It seems that the
nuclei in the young rapidly growing antheridium rarely come to a
typical resting stage.
The question of the presence or absence of centrosomes in the
cells of the young antheridium was taken up carefully, because
previous observations on the karyokinetic figures in the sporophyte
cells and spore mother-cells have convinced me that no such body
0
1906] LEWIS—DEVELOPMENT OF RICCIA 127
appears there. On the other hand centrosome-like bodies appear
in the cells of the older antheridia at the time of nuclear division.
There can be no doubt that these are distinct bodies, and they cannot
possibly be interpreted as accidental granules in that position. In
Some of my preparations hundreds of cells showing them are found
on a single slide, and they are so distinct that the preparation could
easily be used for class demonstration. These bodies appear in
the cells of the antheridium in early stages of its development. I
have been unable to determine whether they appear in the earliest
cell divisions but they appear in the antheridia which consist of only
a few cells. They are not permanent, but disappear and arise anew
with each division.
IKENO regarded it as highly probable (though unable to state this
positively) that in Marchantia these bodies were of nuclear origin.
He figures a small spherical body inside the nuclear membrane,
which in a later stage is found outside the membrane. This body
then divides into two, which arrange themselves on opposite sides of
the nucleus. If the bodies have their origin as one, which later
divides as described, they act as do the ceritrosomes which have
been described for other plants.
In Riccia natans, nothing has been observed to indicate that the
body is of nuclear origin, except that it stains in much the same
Way as the mass of chromatin in the nucleus. In some of my pre-
parations a single body has been observed near the nuclear membrane
(fig. 53). These bodies have never seemed so distinct as the ones
Which appear at the opposite ends of the nucleus and in the poles
of the spindle. There is a dark central part, surrounded by a mass
of cytoplasm which is more or less irregular but does not give the
appearance of distinct radiations such as are described i in the centro-
spheres of certain plants.
When these single bodies were discovered, a careful search was
made of the same preparations and of others in which the two bodies
were on the opposite sides of the nucleus, in order to discover if
possible the intermediate stages which it would seem should appear
in such preparations. In cases in which two bodies have been
observed, they have always been on opposite sides of the nucleus,
©r so nearly opposite that -the spindle developing between them
128 BOTANICAL GAZETTE [FEBRUARY
might take the curved form shown in jig. 60. The origin of the
two bodies is of importance in determining the homology of the
centrosome and blepharoplast and will be discussed later.
Starting with the stage in which the centrosome-like bodies are
on opposite sides of the nucleus, the nuclear division takes place
in the following manner. At first the bodies are at a little distance
from the nuclear membrane, then the nucleus elongates so that the
membrane closely approaches the bodies, becoming somewhat
pointed. At the same time one observes that there is a collection
of kinoplasm at the poles of the nucleus and extending along the
nuclear membrane for some distance. At this time the bodies at
the poles do not show radiations in any direction, but are very distinct
(fig. 54). The spindle is formed from the kinoplasm which has been
described, and when formed consists of a few thick fibres which
converge at the poles, so that the centrosome-like bodies occupy the
position of true centrosomes. About the time when the spindle
develops, the chromosomes are formed from the central mass of the
nucleus and become arranged in the nuclear plate. They are closely
crowded together in this stage, and not so easily counted as when
they have moved to the poles. The photograph (figs. 75, 76) shows
the dense mass formed by the chromosomes when arranged in the
nuclear plate. It was impossible to determine how the division takes
place in the chromosomes as they move to the poles. The changes
take place so rapidly that stages are rarely found in which the chromo-
somes are on their way to the poles.
The centrosome-like bodies disappear during the division, but
it is difficult to say at just what point. Fig. 56, a cell taken from
an antheridium in which only one or two more divisions will take
place, shows the centrosome-like bodies quite distinctly when the
chromosomes are almost at the poles, but by the time the chromosomes
are at the poles and before the daughter nuclei are formed, the bodies
disappear (fig. 57).
These bodies are best seen in preparations which have been over-
stained and washed out. In some cases my preparations were
stained deeply enough to show the spindle and chromosomes well,
but only an occasional spindle showed the bodies at the poles. When
these slides were over-stained and carefully washed out, the bodies
were brought out very distinctly in all cases. ;
1906] LEWIS—DEVELOPMENT OF RICCIA 129
After a large number of divisions has taken place the antheridium
consists of nearly cubical cells, each of which has been considered
by earlier investigators to produce a single spermatozoid. SrrRas-
BURGER (30, p. 482) says of Marchantia polymorpha: “Die Spezial-
mutterzellen der Spermatozoiden sind durch fortgesetzte, sich
rechtwinklig schneidende Teilungsschnitte angelegt worden.” Camp-
BELL (4) describes and figures the spermatozoid mother-cell of Pellia
as producing two spermatozoids. IKENO (16) discovered that in
Marchantia each of the cubical cells undergoes another division in
which the spindles are arranged diagonally, in the earlier divisions
the long axis of the spindle being parallel to the long axis of the cell.
In this last diagonal division no cell wall is formed between the
daughter cells, each of which develops into a spermatozoid. Thus
each of the cubical cells produces two spermatozoids instead of one.
IKENO cites several cases in which two spermatozoids are produced
from a single mother-cell and thinks that this is probably general
in the liverworts and mosses.
JOHNSON (18) has described a diagonal division of the cubical
cells of Monoclea, but he figures a wall separating the two parts of
the cell and regards each three-cornered cell as the mother-cell of
a spermatozoid. He does not give the details of nuclear division
in the earlier stages of the antheridium nor in the formation of the
Spermatozoid mother-cells.
In the last division of the cells in the antheridium of Riccia natans
the spindles are arranged diagonally as in Marchantia. This arrange-
ment of the spindles is quite striking. They are larger than in the
earlier divisions and the bodies at the poles are very distinct. In
Some cases the spindles are curved (figs. 58-60).
No wall is formed between the daughter cells, each of which
develops into a spermatozoid. The centrosome-like bodies do not
disappear after this division (fig. 6r). They remain in the cells, at
first near the nuclei. The daughter cells are contracted, occupying
the central part of the cell cavity (figs. 62, 63). Soon the centrosome-
like body moves away from the nucleus toward the end of the cell.
- Those in the two spermatids may be at the same end or at opposite
ends (figs. 63-67). When the spermatid has become somewhat
rounded, the centrosome-like body has taken its position in contact
130 BOTANICAL GAZETTE [FEBRUARY
with the cell membrane (jig. 68). When the cilia appear they are
inserted in this very small body so that it comes to function as a
blepharoplast. Its small size as compared with that of the cilia of
the mature sperm makes it seem probable that some of the material
for the growth of the cilia must be drawn from another source than
the blepharoplast itself, although it disappears to such an extent
that in the mature sperm it cannot be recognized as the point of
insertion of the cilia.
The developing spermatozoids of Riccia natans do not remain
enclosed in the mother-cells until they are mature, but at about
the stage represented by figs. 70, 71 the walls break down and the
young spermatozoids lie free in the cavity of the antheridium. Here
they seem to undergo considerable growth. The material for this
growth is probably derived from the surrounding cells as they become
collapsed in old antheridia.
The nucleus of the developing spermatozoid takes a position at
one side of the cell and becomes homogeneous. It seems probable
that other material than the chromatin of the spermatid nucleus
must enter into this part of the spermatozoid, because it is very
evident that the body contains more material than would be obtained
from the chromatin alone. Soon the nucleus elongates, following
the outline of the cell and becoming crescent-shaped (figs. 71-73):
In some cases, a distinct vacuole occurs in the cytoplasm although
this is not always the case (figs. 71, 72). The mature spermatozoid
becomes long and slender and consists of the nucleus, the material
of which seems to have increased in amount, a small amount of
cytoplasm, and the cilia which are derived from the blepharoplast
and in all probability from a part of the cytoplasm surrounding it.
IKENO describes a spherical body which appears in the spermatids
of Marchantia before the cilia begin to develop and disappears about
the time that changes take place in the nucleus. It has been impos-
sible to find such a body in Riccia, although it would seem, judging
from IKENo’s figures, that it could easily be seen if present.
The question of the homology of the blepharoplast and centro-
some is one which it seems to me has not yet been settled. In Mar-
chantia, where centrosomes have been reported in the vegetative
cells as well as in the antheridium, the evidence seems good that the
i
1906] LEWIS—DEVELOPMENT OF RICCIA 131
centrosome and the blepharoplast are homologous, and this is the
conclusion of keno. In all other plants in which blepharoplasts
are known to occur, centrosome-like bodies are not present in any
cell divisions except those immediately preceding the formation of
the sperms. That centrosomes occur in liverworts in cells outside
the antheridium is open to question. The conflicting reports of
those who have investigated Pellia epiphylla make it clear that no
distinct body occurs there which can be regarded as a centrosome,
although aggregations of kinoplasm, called centrospheres by most
authors, do occur.
In Riccia natans, it seems very evident that centrosomes do not
occur in the divisions of the spore mother-cells. The spindle poles
are broad, and there is not even a suggestion of a centrosphere such
as has been described ‘for Pellia. In the cells of the sporophyte
GARBER reports centrospheres but no centrosomes. I have never
been able to observe them in my preparations. When the spindle
is fully formed there are no fibres radiating into the surrounding
cytoplasm (figs. 48-52).
Although the thallus of Riccia natans does not present favorable
material for cytological study, a number of cells showing nuclear
division in the gametophyte have been observed near the growing
point. The greatest difficulty here is the presence of numerous
deeply staining granules in the cell. In some cases granules resem-
bling centrosomes appear at the poles of the spindle, but they do
not differ in appearance from the other granules of the cell, and it
seems probable that their occurrence here is accidental.
Summing up, we find that in Riccia natans centrosomes are not
found in the cells of the gametophyte, sporophyte, or spore mother-
cells, but that bodies occur in the dividing cells of the antheridium
which seem to function as centrosomes. In Riccia and Marchantia,
the blepharoplasts certainly have much more the appearance of
centrosomes than in any other plants in which blepharoplasts have
been described. The bodies have every appearance of centrosomes
when at the poles of the elongated nucleus or at the poles of the
spindle. Perhaps the strongest objection to regarding these bodies
as centrosomes lies in the fact that in Riccia natans they occur only
in the cells of the antheridium, while the blepharoplasts reported
132 BOTANICAL GAZETTE [FEBRUARY
in other plants appear only in the last two generations of cells con-
cerned in the formation of spermatozoids.
Those who argue in favor of the homology of the centrosome
and blepharoplast certainly find their best evidence so far in the
liverworts, but it seems to me that this evidence is not conclusive
when the bodies occur only in cells of the antheridium.
In those plants in which centrosomes are known to occur, a single
body divides to produce two, which arrange themselves on opposite
sides of the nucleus (MoTTIER, 25). IKENO has reported a similar
condition in Marchantia. In Riccia natans, however, the evidence
seems to favor the view that the two bodies arise anew with each
division, appearing on opposite sides of the nucleus at the same
time. In this respect they behave more like blepharoplasts.
MorrieR (26) in discussing this question has called attention
to the fact that it is questionable whether we can speak of organs
as homologous which, as such, are without genetic continuity. The
question as to whether true centrosomes have genetic continuity
has not yet been decided, but it is probable that they do not in all
cases.
| SUMMARY.
1. Reccia lutescens and Ricciocarpus natans are forms of the same
plant, the former occurring on the ground in summer and autumn
when the ponds are dry, and the latter as a floating form. Either
form can be changed into the other by altering the supply of water.
Therefore, Riccia Jutescens should not be regarded as a distinct
species. :
2. The genus Ricciocarpus has been based largely on characters
which do not exist. In my opinion, the only real basis for separating
it from Riccia is the more complex structure of the thallus. BiscHOFF
did not regard this as a good character for the separation of the genus.
3. The plant is monoecious, antheridia and archegonia being
produced in definite groups in the same thallus. The sexual organs
appear in autumn when the thalli are growing on the ground and
complete their development the following April. Abundance of
water is not essential to sexual reproduction, as the plants fruit when
kept growing on the soil and supplied with a limited amount of
a
1906] LEWIS—DEVELOPMENT OF RICCIA 133
water; therefore the ground form is not sterile, as was the opinion
of LINDBERG and GARBER.
4. Plants which have been growing attached to the soil and have
been submerged by the filling up of the ponds do not necessarily
perish, but are adapted to spend the winter under water and then
to break loose by the decay of the older part of the thallus and float
upon the water in the spring.
5. The plants are propagated vegetatively by the separation of
branches of the thallus, by the decay of the older part, and also by
the growth of new plants from cells in the apical region.
6. The sexual organs and fruit of the two species studied agree
in their development with the accounts given for the other species
of Riccia. There is no rudimentary integument surrounding the
archegonium or sporophyte of Riccia natans. The sporogonium
of Riccia natans is larger than that of Riccia crystallina and produces
a larger number of spores. The only sterile tissue in either is the
amphithecium, a single layer of tabular cells surrounding the mass
of spore mother-cells.
7- Centrosomes are not present in cells outside the antheridium
nor would I interpret any structure observed in the cells of the 5 eee
phyte or the spore mother-cells as a centrosphere.
8. Bodies which resemble centrosomes, and which are con-
sidered to be true centrosomes by certain authors, occur in the cells
of the antheridium. These bodies do not have genetic’ continuity,
but arise de novo with each division. They do not disappear after
the last division of antheridial cells but remain in the spermatids
and later become blepharoplasts.
9. In the earlier divisions of cells in the antheridium, the spindle
is arranged parallel to the long axis of the cell, but in the last division,
the spindle is placed diagonally in the cell. No wall is formed
between the two cells produced by this division, each of which becomes
a spermatozoid. Thus two sperms are produced from each cuboidal
cell. ,
10. In the developing sperm, the blepharoplast takes a position
on the membrane of the cell and the two cilia grow from it, the nucleus
becomes almost homogeneous in structure and _ crescent-shaped,
almost enclosing the cytoplasm. The mature sperm consists of the
134 ; BOTANICAL GAZETTE [FEBRUARY
nucleus, the cytoplasm, and cilia which have received material for
their growth from the blepharoplast and probably also from the
material surrounding it.
11. The amount of chromatin in the nucleus is small. There
is no nucleolus present unless the masses of chromatin which are
found in nuclei which are undergoing repeated division be inter-
preted as nucleoli.
12. The number of chromosomes is four for the gametophyte
and eight for the sporophyte.
13. The cytoplasm of the spore mother-cells appears to be a
fine reticulum, in which are numerous granules usually located at
the point of intersection of the fibres of the reticulum.
14. The mature spore contains a large quantity of oil together
with a small amount of granular matter. The nucleus of the spore
is very small.
In conclusion I wish to thank Professor Gro. F. ATKINSON and
Dr. E. J. Duranp for valuable advice and assistance during the
progress of this study.
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4
an
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NI
al
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as
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136 BOTANICAL GAZETTE [FEBRUARY
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EXPLANATION OF PLATES V-IX.
All drawings, except fig. 7, were made with camera lucida. Figs. 8-52, with
Bausch & Lomb oculars and objectives, as follows: Figs. 8, 10, 11, 1 in. ocular,
% objective; 9 and 25 2 in. oc., $ obj.; 19, 21-24, 26, 32, 33, 2 im. oc., wy obj.;
12-18, 20, 27-31, 34-52, I in. on as obj.; Figs. 53-73 with Zeiss oc. 18, 2™™
apochromatic objective, 1.40 apert
The figures of plate VI were oe slightly more than one-half in repro-
duction.
All figures are of Riccia natans except the spore mother-cells (figs. 34-47)
which are of Riccia crystallina.
Fic. 1. Rosette of plants awhes on ae soil; a, natural size; b, enlarged.
Fic. 2. Land plants growing in regular clusters.
Fic. 3. Two plants growing on soil, one of which has been igaud and has
grown out in an irregular way from the growing point.
Fic. 4. New plants growing from apical cells of old thalli.
Fic. 5. a, Decay of older part of thallus of the land form to give the floating
form; b, plants collected in May. If these thalli should become stranded on the
mud and growth should continue rosettes would be formed.
Fic. 6. Plants decolorized in alcohol. The sporophytes appear as chains of
dark bodies in the thallus.
PLATE VI.
Fic. 7. Longitudinal section of thallus parallel to the dorsal furrow, showing
arrangement of sexual organs.
Fic. 8. Cross-section of thallus showing archegonia.
Fic. 9. Cross-section of thallus, showing the origin of archegonia in rows on
floor of dorsal groove.
Fic. 10.. Cross-section of thallus, antheridia.
Fic. 11. Longitudinal section of thallus parallel to surface, showing the
arrangement of antheridia. Archegonia have not begun to develop.
1Gs. 12-18. Stages in development of archegonium.
1G. 19. Archegonium in which egg-cell has not been fertilized and is
shrunken
Fic. 20. Cross-section of neck of archegonium.
Fic. 21-25. Stages in development of sporophyte.
Fic. 26. Spore mother-cells. :
Fics. 27-33. Development of antheridium. Figs. 27-31, from materia]
collected in October.
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BOTANICAL GAZETTE, XLI PLALE, 1X
LEWIS on RICCIA
1906] LEWIS—DELOPMENT OF RICCIA 137
PLATE Vii.
Fic. 34. Spore mother-cell in resting state. Chromatin on a fine linin net-
Fic. 35. The chromatin is in the form of an irregular thread.
Fic. 36. Chromosomes formed; weft of delicate fibres about the nucleus.
Fic. 37. Nucleus elongated and showing a weft of fibres.
Fic. 38. Spindle with chromosomes in plate. No centrosome.
Fic. 39. Chromosomes moving to poles of spindle.
Fic. 40. Chromosomes at the poles, thickening of spindle fibres to form cell
Fic. 41. Daughter nuclei.
G. 42. Cell plate. Daughter nuclei contain numerous spherical bodies of
chromatin which stain bright red with safranin.
Fic. 43. Daughter nuclei preparing for divi
Fic. 44. Daughter nucleus with casa in plate. Neither centro-
sphere nor centrosome.
PLATE VIII.
Fic. 45. Chromosomes moving to poles.
1G. 46. Daughter nuclei with chromosomes at the poles to form nuclei of
spores. The cell plate formed in the first division persists.
Fic. 47. Second division completed.
Ss. 48-52. Stages in the division of a sporophyte cell. No centrosome.
Fig. 48 shows slight radiation of cytoplasm from the poles of the elongated nucleus.
Fic. 53. Cells of antheridium which show a single rather irregular body near
the nucleus.
Fic. 54. Cells of antheridium which shows the distinct centrosome-like
bodies at the poles of the elongated nuclei. Compare jig. 74.
Fic. 55. Spindle with centrosome-like bodies at the poles.
Fic. 56. Centrosome-like bodies present when the chromosomes are almost
at the poles.
Fic. 57. Cell from young antheridium. Chromosomes at poles. No
centrosome can be distinguis|
Fic. 58. One cell preparing for last division, while the adjoining cel] has the
spindle formed and arranged diagonally.
Fic. 59. Diagonal arrangement of spindles in last division of cells in the
antheridium
Fic. Oa: Curved spindles. :
Fic. 61. — nuclei formed after diagonal division; centrosome-like
bodies presen
Fic. 62. Cell of antheridium after last division.
Fics. 63-67. Spermatids in mother-cells.
Fics. 68-73. Stages in the development of the spermatozoids.
138 BOTANICAL GAZETTE [FEBRUARY
PLATE IX.
G. 74. Antheridium in which the nuclei are elongated and preparing for
Fic. 75. Portion of section of an antheridium showing the spindles with the
dense chromosomes in the plate and in some cases the centrosome-like bodies at
€ poles.
Fic. 76. Two cells of the same section enlarged three times.
_ Fic. 77. Chromosomes at the poles of the spindles.
_ Fic. 78. Cross-section of thallus showing the hyaline hairs which extend up
into the median grooves.
BRIEFER R ARTICLES.
NOTE ON THE RELATION BETWEEN GROWTH OF ROOTS
AND OF TOPS IN WHEAT.
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY.
LXXXI
Srupts in the experimental morphology of plants have dealt mainly
with the subaerial portions, comparatively little attention having been paid
to variations in the growth of roots. Thus the literature of the subject is
very meager. Several authors, especially. MOELLER,’ DETMER,? PERSEKE,3
Mer,‘ Gat, and FREIDENFELT,® have studied the relation of water to
the growth of these organs, with the general result that in water itself they
grow longer and thinner, with fewer branches and root hairs than are
observed in moist soils. Curiously enough, in soils which are too dry for
optimum growth, the response is very similar. The roots are long and
slender and possess few branches excepting near their tips, which lie, of
course, in moister soil.
The only observation on the relation of root growth to that of tops, with
which the writer is acquainted, is that of MoELLER, to the effect that in a
series of nutrient solutions, of concentrations ranging from 1.0 to 0.05 parts
per thousand, the actual weight of roots produced varies generally with the
concentration, but that the ratio of weight of roots to that of tops is much
larger in the most dilute solution than in any of the others.
A number of experiments have suggested to the author that the accel-
fertierns H., Beitrage zur Kenntniss der Verzwergung. Landw. Jahrb. 13:
167-173.
2DETMER, ie Ueber den Einfluss dusserer Verhiltnisse auf die Wurzelentwick-
elung. “ Landw. Versuchst. I5: 107-113. 1872
3PERSEKE, K., Ueber die Formveranderungen der Wurzel in Erde und Wasser.
Leipzig. 1877.
4MerR, E. De Vinfluence des milieux sur la structure des racines. Comptes
Rend. 88: 1277-1280. 1879.
Recherches expérimentales sur les conditions de développement des poils
radicaux. Ibid.
ep anee de structure et de forme qu’éprouvent les racines
suivant les sie oti elles végétent. Assoc. Franc. pour l’avance. sc. Compt. rend.
de la 9° session. Rheims. 1880.
5GAIN, E., Réle ieilaae de l’eau dans la végétation. Paris. 1895.
©FREIDENFELT, F., Studien iiber die Wurzeln kraiitiger Pflanzen. I. Ueber
die Formbildung der Wurzel vom biologischen Gesichtspunkte. Flora 91: 115-208.
Tg902.
139] {Botanical Gazette, vol. 4x
140 BOTANICAL GAZETTE [FEBRUARY
erating or retarding effect of the soil upon plant growth may often be due
primarily to a response of the roots themselves, and that the ordinarily
observed effect upon the tops may be due to the nature of the roots rather
than to that of the soil directly. This question deserves thorough study ;
the results to be given here cover only a very small portion of the field.
This work was carried on at the laboratories of the Bureau of Soils
of the U. S. Department of Agriculture, Washington, D. C. The plant
used was the Russian variety of wheat known as “Chul.” The plants
were grown directly from the seed in paraffined wire baskets of the form
described by Wuitney and Cameron.’ As these authors have already
pointed out, such baskets possess the advantage over pots of producing a
root system uniformly distributed throughout the soil mass, rather than
the accumulation of roots on the inner surface of the vessel which occurs
in the case of ordinary pots.
The studies to be discussed in this paper were made upon the roots
developed in the first six series described in the author’s previous publica-
tion’ on the growth of tops, and in similar cultures. The medium used
was a very poor soil from Takoma Park, Md., and the same soil with
varying amounts of fermented stable manure added thereto.2 The cultures
of any series were placed side by side in a greenhouse, the amount of
water in all the baskets being kept practically uniform by weighing at
intervals of one or two days and adding the amount of water which was
found to have been lost by transpiration. It is thus seen that the different
cultures were all subjected to the same conditions excepting those which
depend upon the treatment of the soil.
Series I of the paper on growth of topst° will serve as an example ; the
results of all the series are in accurate agreement. The soils and culture
numbers were as follows :
Basket 8
Number - 2 3 4 5 6 7
Takoma | Do. Do.+ Do.+ Do.+ Do Do.+ | Do+
Soil Soil 5000 10,000 15,000 20,000 30,000 40,000 50,000
p.p.m.‘f | p.p.m. p-p.m. p.p.m. p.p.m. p.p.m. p-p.m.
untreated | manure Manure | manure manure manure manure | manure
Sram
7Wauirney, M. and Cameron, F. K., Investigations in soil fertility. U. 5. Dept.
Agric., tiiean of Soils, Bull. 23. 1904
SLIvINGSTON, B. E., Relation of sentation to growth in wheat. Bor. GAZz-
ETTE fhe: 78-195. figs. 21. 1905.
a description of this soil and a discussion of its properties, see LIVINGSTON,
_E, genes ON, J. C., and Ret, F. R., Studies on the properties of a sterile soil.
Ss. De ept. Apric., siren of Soils, Bull. 28. 190
104 photograph of the tops and data for their — are given in that paper.
— abbreviation p. p. m. is used to denote parts per million by weight of air
soil.
1906] BRIEFER ARTICLES 141
At the end of the experiment, which lasted seventeen days, the soil
masses were taken from the baskets and the roots removed from them
with as little injury to the latter as possible. The fresh roots thus prepared
are shown in jig. z. The numbers correspond to the culture numbers
given above. It is at once evident that the root system increases in
amount throughout the series. Closer observation shows that this is
due mainly to differences in the relative number and length of secondary
roots and succeeding branches; the primary roots are of the same number
Fic. 1.—Roots from a series of wheat cultures grown in Takoma soil untreated
and in te same with addition of 5000 p.p.m. to 50,000 p.p.m. of stable manure
in culture 8 as in culture r. They are somewhat longer and more slender
in culture 8.
Photographs of single root systems from a similar series are shown
in figs. 2-5, which bring out the last point more clearly than fig. z. In
fig. 2, which represents the roots from natural Takoma soil, very few
branches are to be seen, and these are exceedingly short; practically the
whole root system consists of the primary roots, with a few adventitious
roots developed at the extreme base of the stem shortly after germination.
ig. 3 shows roots from a soil containing 5000 parts per million of manure.
142 BOTANICAL GAZETTE [FEBRUARY
A very slight increase in number and length of branches is to be observed.
Fig. 4, from a soil containing 10,000 parts per million of manure, shows
numerous well-developed branches, while from a soil with 40,000 parts per
million of manure, as shown in jig. 5, the branches have increased so
markedly in number and extent as to make up by far the greater part of
the system.
From these facts it is seen that, for this series of soils the variation in
growth of tops is correlated with the number and length of lateral roots.
The water content of all the soils was the same, so that the variations in
—Wheat roots grown in Ta- 3-—Roots apie in Takoma
Fic. Fic.
koma ot untreated. soil with poe p-p.m. manu
growth cannot be related to this factor; therefore they must be connected
with some unknown condition in the soil itself, a condition which is related
_ to the amount of manure present.
The comparative anatomy of these roots was investigated, both by
hand and paraffin sections, with the result that in the poor soils the main
roots have a strong tendency to swell by direct enlargement of the cortical
cells, without increase in the number of these cells, while in the better soils
this tendency is not nearly so marked. Very old wheat roots from autumm
stubble in the field show this balloon-like enlargement of the cortical cells
to a still greater degree. This is apparently a phenomenon of age, sug-
gesting that roots in the poor soils age more rapidly than in the better
ones. It was also found that the zone of root hairs, which normally has
its lower limit s-1o™™ from the root apex, extends in the poor soils to
within 1-3™™ of the tip. The outgrowth of root hairs from the piliferous
layer may also be related to the age of the cells; as is well known, these
Se
1906] BRIEFER ARTICLES 143
organs normally appear only after the cells from which they arise have
passed through their period of most rapid growt
It would seem that the poor soil, by inhibits branch growth and
causing the enlargement of cortical cells, may render the root system unable
to carry on an adequate amount of absorption for normal growth, and that
this fact may be the main clue to an explanation of the stunted tops in such
cases. That the inadequacy of the stunted roots is in regard to the water
supply rather than to that of salts, is indicated by the fact that in distilled
IG. 4.—Roots grown in Takoma soil Fic. 5.—Roots grown in Takoma soil
with 10,090 p-p-m. manure. with 40,000 p.p.m. manure.
water, for the first two or three weeks, a better growth of tops is obtained
than in the natural Takoma soil.
Determination of the relation of the dry weight of the root system to
the nature of the soil was deemed advisable, but a number of tests yielded
only negative results. The dry weight of the complete system was found
to be practically the same from all of the soils. The variations were
always irregular. This may, of course, be partially due to the fact that,
while it is manifestly impossible to obtain anything like the entire root
System of one of the better cultures (owing to the extreme fineness of the
branches and their adhesion to the soil particles), yet from the poorer
cultures, where branches are short and few, a much larger part of the
System is obtainable—Burton Epwarp Livincston, The University of
Chicago, March 5, 1905.
CURRENT LITERATURE.
BOOK REVIEWS.
The Swiss moors. ~*
Fring and Scurorer have published a remarkable work on the moors of
Switzerland, and have thus placed all who are interested in bog studies under
lasting obligations. More than half of the huge volume is given over to the
discussion of general geological, chemical, physical and biological problems
connected with peat formation and the ecology of bog plants. One of the most
valuable features is the discussion and scurry of the mass of European and
foreign literature touching these problem
After defining the scope of the nveignton, the authors take up the peat-
producing plant associations of Switzerland. These are described with the
greatest detail, from the plankton to i seca! vegetation; and their relation
to peat accumulation is explained.
e moors are distinguished primarily as flat bogs, raised bogs, and bogs
of the alpine regions. The first two types are of such general occurrence that
their peculiarities may be briefly summarized here.
The Flachmoor is characterized as occurring in connection with waters rich
in mineral matter, especially lime, in both wet and dry climates. Usually they
show centripetal growth, and are dominated by species of Cyperaceae, Gram-
ineae, Juncaceae and Hypneae. Species of Alnus, Betula, and Frangula
make up the woody growth. Sphagnum, Ericaceae, and Empetrum are
entirely wanting.
n the other hand, the Hochmoor type occurs under the influence of
waters poor in mineral ee where the rainfall is abundant and the tem-
perature mild or cold. The ace is convex. The oldest portion is toward
the center; hence the BO: is ccokatiad The dominant plants are Sphag-
um, Oxycoccus, Andromeda, Calluna, Vaccinium, Empetrum, Eriophorum
vaginatum, Pinus montana uncinata, Betula pubescens, and B. nana. These
plants do not occur on the flat bogs, and are driven out by irrigation with
waters of si mineral content, especially lime.
his summary suggests the essential difference between American bogs and
the poner: of European authors. Grass marshes occur here, which are
strictly comparable. But the term “flat bog” is usually applied to areas
tFrtg, J. and Scuréter, C., Die Moore der Schweiz mit ii ir der
gesamten Moorfrage. Beitrage zur Geologie des Schweiz, herausgegeben von =
Geol. Komm. der Schweiz. naturf. Gesells. Geotechnische Sire III Lief. 4to,
pp. xviii+751. Bern: A. Francke. 1904. M 40.
144
1906] CURRENT LITERATURE . 145
having a peat substratum, dominated by Sphagnum, Cassandra, Andromeda,
Oxycoccus, Ledum, Vaccinium, etc.—-a vegetation more nearly related flor-
istically to the Hochmaor. Further, these occur in areas whose soil water is
rich in mineral salts, frequently overlying marl! The raised bogs of America
are strictly comparable to the Hochmoor of Europe, but their occurrence
appears to be localized by climate rather than by the character of the soil
r
For both the flat and raised-bog plants, the authors conclude that the
substratum is physiologically dry because of the combined influence of three
factors: (1) high water-content of the substratum, (2) consequent low tempera-
ture and (3) the difficulties in the way of root respiration (due to wet soil and
scarcity of oxygen), accompanied by a general impairing of all root functions.
The plants of the Flachmoor are noted for their ready absorption of
mineral salts, absence of mycor ni unusual development of the — nd
parts, and the high percentage of as cause of the xerophytic characters
of many of the plants is not clea They a re probably connected with the
difficulties of absorption. These at are hes hydrophilous, never occur-
ring in dry situations.
The Huchmoor plants, however, absorb mineral salts with difficulty; mycorhiza
bas
occurs in all the Ericaceae, in Empetrum, Betula, and Pinus; carnivorous plants
are common; the root systems are poorly developed; and the ash content is
low. The xesophyie character of the plants is in part due to the difficulties of
absorption, and in part to their evergreen habit. Many raised-bog plants also
occur in dry situations.
The third and fourth chapters discuss the conditions and processes involved
in peat accumulation, the chemical and physical properties of the end products,
the classification of peats, the bog minerals, and the relation of bogs to coal
deposits
The geographic distribution of the bogs of Switzerland, a geomorphic classi-
fication of the moors of the world, the relation of settlement to moor develop-
ment, the economics of the Swiss moors (with bibliography), the bog deposits
as records of the postglacial history of northern ngniariag form the other
principal chapter headings of the first portion of the wo
The second division gives a detailed description . the individual Swiss
bogs. In most instances these include not only plant associations of the
present surface but also the succession of plant remains occurring in the peat.
The location of the bogs is made clear by-an excellent topographic map, upon
which the distribution of the several bog types is shown. The bibliography
occupies seventeen pages and includes only the more important purely scientific
. Papers.
e importance of the work from the standpoint of future investigation
is undoubted. To American students it furnishes not only a key to the present
status of the subject, but also a model for the study and description of our
own bogs, marshes, and swamps.—E. N. TRANSEAU.
146 BOTANICAL GAZETTE [FEBRUARY
Reproduction of mildews.
Harper has brought together the results of several years of study of
nuclear activities in the mildews*in a lengthy and beautifully illustrated
publication from the Carnegie Institution.2 It is impossible for us to consider
more than the striking new features of his investigations. The paper contains
a résumé of much of his earlier work and a broad discussion of many
cytological principles which are of general interest and will richly repay the
reader of this very creditable contribution to American botany. The author
takes a strong stand for critical morphological analysis and classification of
the stages in the life history of thallophytes, with a clear separation of phylo-
anaes history from physiological functions.
m mportant new features of HARpER’s research, chiefly in
Phyllactinia, are @) the establishment of a “central body” within the nucleus,
which constitutes a point of attachment for the chromatic elements and gives
a clear polarity to the structure, and its continuous existence through the most
important phases in the life history; (2) the evidence for the permanence of
the chromosomes; and (3) the evidence that the triple mitoses preceded by
synapsis in the ascus constitute a double reduction of the chromosomes which
are quadrupled by the two nuclear fusions in the life history, the first fusion
at the time of the sexual act and the second fusion within the young ascus.
The central body is a permanent structure, always present in the resting
nucleus, dividing with each mitosis, and the center for an arrangement of
chromatic threads within the nucleus and for the attachment of spindle fibers
during nuclear division. Its position determines a pole in the. nucleus around
which are grouped the chromatic elements, which are thus always in connection
with the central body, both in the resting nucleus and during mitosis. This
e succession of mitoses in the life-history. HARPER has not been able to
i the different sets of chromosomes after the nuclear fusions, for the
chromatic elements and the central bodies unite very intimately. But the
second fusion in the life history, that in the ascus, is followed at once by a
period of synapsis and the triple mitoses out of which come the eight chromo-
somes characteristic of the gametophytic phase of the form.—B. M. Davis.
MINOR NOTICES.
Observations in Spitzbergen——The flora of Spitzbergen is fairly known.
Therefore, Dk. WuLFr, who accompanied the Swedish expedition for the measure-
ment of an arc of the meridian, undertook to make ecological observations on
the arctic plants,3 especially touching their transpiration, occurrence of mycorhiza
2 HARPER, R. A., Sexual ee oe and the organization of the nucleus i
certain mildews imp. 8vo. pp. . pls. 7. Washington: er Institution of
apAbae gos.
ULFF, THORILD, Observations botaniques faites au Spitzberg. Missions scien-
ious pour a mesure d’un arc de méridien au Spitzberg. Mission uédoise. ‘Tome
Il, X¢ section, Botanique. Traduit de Allemand par H. Marcet Harpy & Dundee.
4to. pp. 63, pls. 4. Stockholm. 1903.
1906] CURRENT LITERATURE 147
and anthocyan, the vegetation of the “polygonal” soils, and to make miscel-
laneous floristic notes at various stations. The transpiration he finds very feeble
and almost without diurnal periodicity or plant control. This feeble transpira-
tion he accuses of being a cause of feeble growth; instead, is not its feebleness
due to the same cause as the feebleness of growth, the low supply of energy ?
Mycorhizas, internal and external, are common. Anthocyan is found in fifty
species, about half the known higher plants. It is always lacking in plants grow-
ing on soil enriched by the droppings of wild birds, whereas the same species
growing on poor soils show it abundantly. As to the rdle of anthocyan, he holds
it for an absorber of energy, and without it no plant can become dominant in
arctic regions. For other interesting observations one must consult the work
itself —C. R. B.
Polypodiacez and edible fungi—Not that there is any connection between
them; but both are treated by CopELAND in a bulletin+ from the Government
Laboratories at Manila. The section on Polypodiacee forms the bulk of the
bulletin and is “an attempt to collect and publish descriptions of all the ferns
known to have been found in these islands.”” The author adds: “I am not per-
sonally acquainted with a large part of those ferns still known here only from
earlier collections.” Which leads us to remark that he should then have abstained
from describing a new genus and new species among them, as hedid in Dr.
PERKIns’s last Fragmenta. In reprinting here these descriptions he has neglected
to indicate that they have already been published elsewhere. He has sinned again
in adding one more new name in this bulletin. The compilation of such
descriptive floras is undoubtedly serviceable; but one who is not a taxonomist
and who confesses the absence of indispensable books and specimens, should
not take the chances of cumbering pteridology with new names which may or
may not be justified. And the same may be said regarding the brief fungus
part.—C. RB.
Genera of Mexican plants.—The flora of Mexico is so closely related to our
own that any work on it is of essential assistance to American taxonomists. So
we welcome the assembling and description of the Mexican genera, and the list-
ing of the species, undertaken by Professor ConzaTTI, director of the State Nor-
mal School of Oaxaca, of which the first volume, on Polypetalae, has recently
been published by the Ministry of Public Works. This volume 5 begins with an
artificial key covering about 50 pages*including all genera, and contains descrip-
tions of 667 genera of Polypetale, representing 71 families, and including close
to 4,500 species. This is to be followed by another on Gamopetalae and a third
4CopELAND, E. B., I. The Polypodiacee of the Philippine Islands. II]. New
species of edible Philippine fungi. Bureau of Government Labs. Bull. 28. 8vo.
pp. 146. pls. 3. 1905.
sConzatTI, C., Los géneros vegetales mexicanos. Imp. 8vo. pp. 449. Mexico:
Oficina Tip. de la Secretaria de Fomento. 1905. $3 (Mexican).
148 BOTANICAL GAZETTE [vEBRUARY
on Monochlamydeae, Monocotyledoneae, Gymnospermeae, and Pteridophyta,
embracing in all about 1900 genera.
The descriptions are very full, and though the diagnostic characters are not
indicated, this is largely atoned for “ the tae system of synoptic characters
under the tribes and subtribes.—C. R
Germs of mind in plants.—A little book,® unknown to us in the original
French, now translated into English by A. M. Sruons, well-known for his work
in Chicago along social and philanthropic lines, shows that there exists in France
the same sort of popularizers of science as in our country—writers who with a
smattering of scientific knowledge lack the fuller knowledge that forms a back-
ground and furnishes scientific perspective. The facts of plant ecology are herein
so distorted in their relation as to become caricatures; the use of words is so fan-
ciful as to convert sober ideas into grotesque fairy-tales. For this, doubtless,
the author is chiefly responsible; but the translator slips occasionally through
unfamiliarity with a technical use of some common word.
The book is interesting; but it is as little “science” as a historical novel is
history. It is difficult to see how such fiction can be “a contribution to the cause
of socialism and science.” —C. R. B
Hepaticae of France.—LAcouture has prepared a helpful series of
descriptive analytical keys to facilitate the identification of French liverworts by
amateurs.’? The keys are arranged in a convenient bracket fashion, which is
easy to use but makes the form of the thin volume rather unhandy and pre-
cludes its use as a field manual. The description of each species is accompanied
by an excellent figure illustrating tl t essential features described. The keys,
in the form of tables, are arranged in three series, of which the first, consisting
of tables 1 and 1m gives the characters of the tribes; the second, tables 11I-Ix
the characters of the genera; and the third, tables xm-xxxIx, the characters
of the species and the illustrations. No attempt is made to exhibit the natural
classification —C. J. CHAMBERLAIN.
Index Filicum.— The fourth and fifth fascicles of CHRISTENSEN’s important
work® were issued respectively in October and December last. They carry the
references from Cyat/:ea lanuginosa to Gleichenia cryptocarpa. The huge genus
Dryopteris alone takes fifty-two pages, which indicates something of the compre-
hensiveness of the work. Let colleges and libraries hasten to support by their
6FrANcE, R. H., Germs of mind in plants. Trans. by A. M. Smmons. 12mo.
pp. 151. Chicago: C. H. Kerr & Co., 1905. 50 cts
Lacouturr, CH., Hépatiques de la France. Tableaux synoptiques de.
caractéres sedllatite des tribus, des genres, et des espéces. 4to. pp- 78. /#gs- 200-
Paris: Paul Klincksieck. 1905. /r. 10.
8CHRISTENSEN, C., Index Filicum, etc. Fasc. 4,5. Copenhagen: H. Hagerups
Boghandel. 1905. Each 3sh. 6d.
1906] CURRENT LITERATURE 149
orders the stupendous and too thankless task which the author has undertaken.
The employment of the American system of citation is notable-—C. R. B.
Das Pflanzenreich.°—Of this work parts 22 and 23 have lately appeared,
including respectively the Primulaceae by Pax and Knurs, and the Halorrha-
gaceae by SCHINDLER. The rate at which these monographs are appearing is
remarkable, and shows something of the energy of the editor and his sagacity
in the selection of his collaborators. The publisher’s part, too, is admirably
done—C., RB,
Eucalyptus.— Marpen’s revision’® has now reached part 7, which includes
EE. regnans, vitellina, vitrea, dives, Andrewsi, and diversijolia, and is illustrated
by four plates.—C. R. B
NOTES FOR. STUDENTS.
Items of taxonomic interest —ZAHLBRUCKNER lists (Beihefte Bot. Cent. 19?:
75-84. 1905) the lichens collected by Professor D, H. MreveEr in the Ecuador
highlands in 1903, describing six new species.—Carpor (idem 85-148. figs. 39)
enumerates 125 species of the mosses of Formosa, collected by Abbé Faurre in
1903, bringing the total known species of this island to 130, of which 39 are
new. Herpetineuron (C. Mill. as Anomodon §) is raised to generic rank.—
ENGLER describes (Bot. Jahrb. Syst. 37: 95, 96. 1905) a new genus of Araceae,
Ulearum, and in his tenth contribution to a knowledge of the Araceae, (idem)
adds to the family nearly a hundred new species, chiefly from Central America,
the subequatorial andine province, the Philippines, and East Indies——Drere1,
in his sixth paper on Japanese Uredineae (idem 97-109) describes 16 new species,
and in one on Japanese fungi (dem 156-160) ten others.—RADLKOFER (idem
144-155) describes 8 new species of Serjania and 8 of Paullinia (Sapindaceae)
from Peru, Brazil, Bolivia, and Columbia.—StTEPHANI (Bull. Herb. Boiss. IT.
5: 885-900, 917-946. 1905) in his Species Hepaticarum concludes the treatment of
the genus Plagiochila, describing 26 new species, a number of them from equa-
torial America.—Domn (idem 947, 948) describes 2 new species of Koeleria from
Asia, and BEAUVERD (idem 948) a new Burmannia from Brazil and (g90-991
a new Hesperantha from the Transvaal.—FERNALD characterizes (Ottawa Nat.
19: 156. 1905) a new variety of Antennaria nevdiica Green from E. Quebec.—
SCHNEIDER, in a prodromus to a monograph of Berberis (Bull. Herb. Boiss.
II. 5: 139 ff. 1905) recognizes 159 species, among them a number of new ones
of his own creation, which he divides into 21 sections. The regions of their
9 ENGLER, A., Das Pflanzenreich. Heft 22, ere by F. Pax and R.
KnurH. pp. 386 Ae. 75 (311), maps 2. M 19. 20.—Heft 23. Halorrhagaceae by
ANTON K. SCHINDLER. pp. 133, figs. 36 ae M 6. 80. Leipzig: Wilhelm
Engelmann. oer
10 MAIDEN, J. H., A critical revision of the genus Eucalyptus. 4to. pp. 183-205, ,
pls. 33-36. Sydney: Government N. S. Wales. 1905. 2sh. 6d.
150 BOTANICAL GAZETTE [FEBRUARY
dominance are South America and E. Asia ——HeEtier describes (Muhlenb. 1:
124) a new Veratrum from Idaho, and (idem 125) a Linanthus or Gilia from
California.—McALPrnE adds a new genus, Uromycladium, to the Uredineae (Ann.
Mycol. 3: 303-323. pls. 6-9. 1905). It is based on 7 Australian species occurring
on Acacias, and is intermediate between Uromyces and Ravenelia.—VUILLEMIN
shows the identity of Hartigiella with Meria (idem 340-343).—SCHMIDLE found
eases ; .
proposes an extension of the moss family Pterobryaceae to include five other
families, in whole or in part, and gathers from various genera some 25 species
to swell his genus Pterobryopsis. He establishes a new monotypic genus
Miillerobryum on an Australian moss already referred to 3 separate genera.
Trachypodaceae is a new family, and Trachypodopsis its characteristic new
genus, for both of which he has “gathered of every kind,” and Teil I is
only begun!—Prck (Rept. N. Y. State Botanist 1904) describes new fungi;
Boletus (3), Clavaria (2), Cortinarius, Lactarius (2), Pholiota—CARDOT
finds 35 new species of mosses in SkoTTSBERG’s collections made on the
Swedish antarctic expedition (Bull. Herb. Boiss. II. 5: 997-1011. 1905).—
Hieronymus has studied (Bot. Jahrb. Syst. 36: 458-573. 1905) the Compositae
collected by JretsK1 in Peru, among which he finds 58 new species—— DIELS
idem Beiblatt 82: 1-138) makes hundreds of additions to his flora of central
yee eee ng many new species and three new genera, Giraldzella Dammer
), Pleroxygonum Dammer and Diels (Polygonaceae), and Biondia
per (Necletiadarena), —NELSON describes (Proc. Biol. Soc. Wash. 18: 171-
776. 1905) new species from Nevada in Cleomella (2), Sphaerostigma, Zauschneria,
Rhamnus, Polemonium, Artemisia, and a new genus of Solanaceae, Bosleria—
ENAULD and Carpor in their tenth paper on Musci Evxotici (Bull. Soc.
Roy. Bot. Belgique 41: 7-122. 1905) describe, among many others, largely
Mascarene and East Indian, 9 new species from Porto Rico, 3 from Costa
Rico, one from Guadeloupe, 3 from Cocos Island (Pacific Cent. Am.), 1 from
Mexico, and 1 from Hawaii. They also establish as a new genus of Hypnaceae
Miiller’s section of Hypnum, Dimorphella. The same authors (idem 123 ff.) in
their third article on Musci Costaricenses describe 22 new species—-HELLER
has found some new species in his collections for 1905 in California and
describes them (Muhlenb. 2: 1-6. 1905), under Eriogonum (3), Montia, Del-
phinium, Ranunculus, Thysanocarpus (2), Lithophragma, Ribes, and Amelan-
chier.—Howe adds several algae to our flora (Bull. Torr. Bot. Club 32: 563-586.
pls. 23-29. 1905) from the Bahama-Florida region; Halimeda, Avrainvillea,
Sarcomenia, Dudresnaya, and a new genus Cadoci pati (Codiaceae), besides
changing several names.—UNDERWOOD (idem 587-596) maintains the genus
Alcicornium Gaud. as valid, gives a synopsis of the species, and describes
A. Veitchii as a new species—R¥DBERG, about to publish a Flora of Colorado,
makes (idem 597-610) what he considers necessary changes in names, and
describes new species of Deschampsia, Eatonia, Poa (9), Festuca (2), and
1906] CURRENT LITERATURE 151
Elymus (2).—OsrerHout proposes from Colorado (idem 611-61 3) new species
of Allionia, Aster, Senecio, and Carduus (2), which are respectfully referred to
Mr. RYDBERG.—SARGENT adds (Rhodora 7: 192-219. 1905) 24 new species of
Crataegus, all from New England.—Rostnson describes (idem 219-222) a new
Ranunculus from Gaspé and Labrador.—C. R. B.
Fossil gymnosperms.—Two trunks of Cycadoidea have been found in the
Portland beds of Boulogne, to which MM. Fricue and ZEILLER give the specific
name C. pumila on account of their small size."* Another Cycadoidea is des-
cribed without attribution of a specific name. An interesting and important dis-
covery is a cone of Sequoia of the S. giguntea type, which is named 5S. portlandica.
The oldest well authenticated cone of Sequoia previously known is Heer’s S.
lusitanica from the Wealden beds of Portugal, which belongs to the type repre-
sented by the living S. sempervirens. It thus is demonstrated that Sequoia existed
in its two living types as far back as the Jurassic period and must thus be very much
more ancient in its first appearance. Other important discoveries are pine-cones
representing the two main series of the present day, viz., the sections Strobus and
Pinaster. The cone of the Strobus type is very much flattened and does not
yield any definite information as to its internal organization, so the authors
include it under the provisional fossil genus Pinites, with the specific appellation
P. strobijormis, which would appear to be too close to our western Pinus strobi-
jormis to stand as a permanent name. The other cone is exceedingly well pre-
served and resembles very closely, as the authors point out, small cones of the
living P. Luricio. This cone is referred to Pinus as P. Sauvagei. ‘These obser-
vations are of very special interest because they establish that Pinus too must be
a very old genus, since examples of both the hard and soft pine series existed
already in the Jurassic.
GoTHAN calls attention to the somewhat Puts TEA condition of X ylopa-
laeontolgie at the present time and by comprehensive study of fossil and living
woods, including many type-specimens of the former, reaches a number of con-
clusions of greater or less importance.t? The proposition of FELrIx to divide
fossil woods presenting tracheary structure resembling that of living Araucarineae,
into Cordaioxyla for the palaeozoic woods, which may be supposed to be those
of Cordaites, and into Araucarioxyla for mesozoic and later woods, is rejected,
since in the author’s opinion no distinction can be made histologically between
the two. For these w ENDLICHER’s name Dadoxylon is retained. Cedroxylon.
Kraus and Cupressinoxylon Goeppert are separated from each other, not on the
basis of the presence of resiniferous parenchyma in the latter genus and ts
absence in the former, but on the character of the medullary ray-cells, since many
Cedroxyla and even Pityoxyla have resinous parenchyma. This distinction has
LICHE, P., et ZEILLER, R., Note sur un florule portlandienne des environs de
Roper Me Bull. Soc. Geol. de la France IV. 4: 787-812. 1904.
2 GoTHAN, W., Zur Anatomie lebender und fossiler a
ABhasdl. ag pilcaed: geol. Landesanstalt, Neue Folge, Heft 44.
152 BOTANICAL GAZETTE [FEBRUARY
already and previously been clearly made by PENHALLOW. The author also
attempts to separate the woods of the Podocarpez from those of the Cupressineae
in the larger sense, on the basis of the structure of the pits in the ray-cells. The
success of this distinction may be judged from the fact that it results in putting
Sciadopitys with the Podocarpéae. Pityoxylon of Kraus is broken up by this
writer into two genera, Piceoxylon and Pinusoxylon. The latter genus repre-
sents the wood of Pinus, and seems somewhat unfortunate, since it is doubtful if
the mesozoic pines had the wood structure which is found as characteristic of
that genus in Tertiary and modern times. There are also disquisitious on spiral
_Striation in the wood of the nosperms and on the value of annual woody
rings as diagnostic of geologic formations. The work closes with two tables for
the determination respectively of living and fossil gymnospermous woods. There
is likewise an index and an alphabetical list of the living woods investigated by
the author.—E. C. JEFFREY
Injury by smoke.—F requent controversies and law suits, arising from damage
to agricultural crops by the smoke produced by manufacturing establishments
in Germany, have made the recognition of this form of injury extremely impor-
tant. In order to furnish a basis for distinguishing smoke-injury from injuries
due to other factors, SorAvER'S has made a comparative anatomical study of
various kinds of injury commonly occurring in the more important grains, oats,
wheat, and barley. The paper contains detailed comparative descriptions of
changes in the cell walls and cell contents which cannot be severally noted here.
The general plan followed in each case is represented by the following heads:
e behavior of the normal plant in its gradual, natural dissolution; abnormali-
ties in smoke-free regions; the phenomena in plants injured by chlorin and by
hydrochloric acid fumes; experimental tests of the influence of hydrochloric
acid fumes; phenomena confused with smoke injuries. In natural death the
cells lose a large part of their contents and finally (except the epidermal cells)
collapse completely. This process first involves the tip and edges of the leaves.
In cases of death resulting from other causes, as drought, the cells do not collapse
so completely, since the contents are not fully resorbed. In injuries due to acid
fumes from smoke, the contents of the mesophyll cells contract into an irregular
greenish lump, while the cell walls partially collapse.
The most striking feature about this form of injury is the collapse of the
epidermal cells. The accompanying changes of the cell contents and cell walls
in these and in many other forms of i injury are minutely described. The recogni-
tion of smoke injury in general is based on the fact that the cells, dying rapidly,
collapse partially without being emptied of their contents, the epidermal cells
showing the same phenomena. The author continually emphasizes the fact,
however, that no clearly defined symptoms for the absolute and certain recogni-
tion of smoke injury can be given, but that in all cases a comparative study of
. 13 SORAUER, P., Beitrag zur anatomischen Analyse rauchbeschadigter Pflanzen.
Landw. Jahrb. 33:585-664. pl. 15-18. 1904.
.
*
1906] CURRENT LITERATURE 153
plants growing under the immediate influence of the acid fumes and others
growing under similar conditions but not within the smoke zone, must be made.
—H. HaAssE.princ.
Viticulture—Recent publications from the Royal Hungarian Central Insti-
tute of Viticulture are as follows: Volume III, part 2, consists of chemical analyses
of the stems and shoots of American species used for stocks in Hungary.'4 The
points determined were the moisture content, ether extractives (oils, fats, waxes,
ms, and organic acids not further determined), alcoholic extractives (tannin,
pashan, vanillin, and organic acids), nitrogen, starch, cellulose, and pro-
teids. e paper contains a large number of analyses made at different seasons,
but no general results have yet been reached, and it is difficult to see what may
be expected. Part 3 of this volume is a small paper by IstvANFFI'S in which he
describes a disease of the vine caused by Phyllosticta Bizzozeriana Massal. The
disease is not of great importance, but has been mistaken for the black rot, one
of the most dangerous vine diseases. In the part 4 IstvANFrFI’® gives the results
of his investigations on the gray rot, caused by Botrytis cinerea. The first part
of this paper is taken up with the effects of various kinds of poisons and other
treatments as cold, drying, etc., on the spores of the fungus. One of the most
striking results is the unusually high resistance which the spores are said to have
to copper. Spores were kept twenty-four hours in different strengths of Bordeaux
mixture ranging from 1 to 10 per cent., to which was then added must containing
I per cent. of tartaric acid, so that the resulting solutions contained the equivalent
of 0.3 per cent. CuSO,. Of the spores from the lowest strength mixture 38-40
per cent, germinated, of those in the highest 10-12 per cent. germinated. Spores
sown on berries in 3 per cent. Bordeaux mixture germinated and penetrated the
epidermis. Spores, kept one hour in a 2 per cent. solution of CuSO,, which
was then diluted with ten times its volume of must, germinated. Many other
similar experiments are given. The second part of the paper deals with the
development and life history of Botrytis cinerea and methods of control. Ve
little new is added to the life history of the fungus. For treatment, spraying
with a 5 per cent. solution of calcium bisulfid is recommended.—H. HasseLBRING.
Endotrophic mycorhiza.— The long and important paper of GALLAUD'? on
this subject merits brief summary, as his conclusions are quite revolutionary.
He has described for the first time the anatomical and cytological characters of
™ Gaspar, J., Analyses des sarments américains. Ann. Inst. Cent. Ampél.
Roy. Hongrois 3; nen pls. 4-12. 1905.
5 IsrvANrFFI, Gy. de, D’une maladie de la vigne causée par le Ph-yllosticta Bizzo-
zeriana. Idem, 167-182. ap 1}. 1906:
‘6 IsTvANFFI, ae de, Etudes scbitnistegiaiinn et mycologiques sur le rot gris
de la vigne. Idem 183-360. pls. 14-21. 1995.
17 GALLAUD, L, Etudes sur les mycorhizes endotrophes. Rev. Gén. Bot. 17:
pls. 4. 1905.
154 BOTANICAL GAZETTE [FEBRUARY
a large number of endophytes, and his study enables him to distinguish four
types: (1) type of Arum maculatum, hyphae intercellular after traversing the
outer cells, their growth arrested by formation of simple terminal haustoria
which penetrate the cortical cells; (2) type of Paris quadrifolia, hyphae intra-
cellular, of indefinite growth, with complex lateral haustoria arising at definite
points; (3) type of Hepaticae, hyphae intracellular, of indefinite growth, enter-
ing via rhizoids and bearing haustoria transformed into sporangioles; (4) type
of Orchideae, hyphae intracellular, of indefinite growth, forming tight pellets
which are sometimes permanent and sometimes undergo more or less complete
digestion.
There is a remarkable uniformity in the constitution of the cell walls and
in the cytological structure.’ Repeated attempts to isolate the fungi by direct
extraction and by inoculation were unsuccessful. The first method failed, prob-
ably because the fungus already in was already too much altered by the digestive
action of the host, and the second leads the author to distrust utterly the identi-
fications of previous authors. The endophytes, he holds, are saprophytes internes,
which by their highly differentiated haustoria borrow some non-living nutritive
material from the cells in which they live. These cells react very rapidly on the
fungus, killing its haustoria, digesting and absorbing them in part; then they
resume normal life, momentarily disturbed. It cannot be said that there
is a harmonious symbiosis between the two plants, but rather a conflict between
the caotine but little harmful, fungus and the cells which defend themselves
by their digestive power—C. R. B
Sexual reproduction of Stigeoclonium.—PascHeER in an account of the sexual
reproduction of Stigeoclonium fasciculatum,*® touches briefly on the formation
and behavior of the zoospores (macrospores), which in general agrees with that
of other forms, but in a few cases the sporelings developed into filaments of a few
cells only, which then formed in each cell a single four-ciliate zoospore (macro-
spore) that developed like other zoospores. The microspores are four-ciliate and
long motile; after losing their motility they become spherical and either form
resting-cells, or (rarely) conjugate and form zygotes. The development of the
latter was not followed, but from hasty observation he concludes that their germi-
nation does not depart from that of the zoospores or the resting-cells. After an
indefinite period the resting-cells germinate like the zoospores. Some, however,
(akinetes or palmella stage), grow into a few-celled filament, each cell giving rise
to four biciliate zoospores, resembling the microspores in size and activity, except
that they will not conjugate but germinate at once like the zoospores
Phylogenetically he claims for Stigeoclonium fasciculatum a position midway
between Ulothrix and Draparnaldia, the three kinds of spores indicating that it
is on the border-line of sexual reproduction. The same position was long ago
claimed by Dopet-Porrt for Ulothrix zonata. But such generalizations will bear,
8 Pascuer, A., Zur Kenntnis der geschlechtlichen Fortpflanzung bei Stigeo-
clonium. Flora 95: 95-107. figs. 2. 1905.
RS Ae NO ENR
1906] CURRENT LITERATURE 155
revision, and investigations of the cytological phenomena involved are especially
needed. PascHer’s observations were microscopic to be sure, but he has appar-
ently attempted no cytological observations at all—R. THrESsEN.
Sigillarian stems.—Owing to the rarity of sigillarian stems showing structure
the description of new specimens is of particular interest to paleobotanists.
IDsTON’® has given a well-illustrated and adequate description of Sigillaria
elegans, which differs from the historic S. Menardi in that the primary wood of
the former is continuous instead of broken up into bundles. The protoxylem is
external to the metaxylem, and both are composed of scalariform tracheids.
The secondary wood is about equal in thickness to the primary, and shows medul-
lary rays which are mostly one cell thick and one to nine cells high. The outer
margin of the primary wood is crenate, and from the furrows arise the leaf traces,
of which there are about twenty-eight in a cross section; these do not seem to
possess any secondary wood. As is usual in sigillarian stems the pith, phloem,
and inner cortex have perished, and the outer cortex contains a broad zone of
periderm. . elegans, with a continuous ring of primary xylem, S. spinulosa,
with a mixture of continuous and discrete xylem, and S. Menardi, with separate
bundles, form a good series, and judging from the scanty data available it seems
that this series represents a sequence in time. The features of S. elegans support
the view that the genus sprung from forms more like Lepidodendron.—M. A.
CHRYSLER.
Mycoplasmic propagation of grain rust—Erixsson has published another
instalment of his studies on the demonstration of the propagation of grain rust
by means of mycoplasm, this time dealing with Puccinia graminis.?° Four means
are recognized by which the uredo stage of the rust may possibly arise in spring
time in winter wheat: (1) from spores of the barberry aecidium, which in turn
arose from the resting teleutospores that had remained dormant over winter;
(2) direct infection of the wheat plant from the resting teleutospores (homoecism);
(3) uredo infection from mycelium remaining alive in the wheat plant over winter;
and (4) from endogenous germs of disease (mycoplasm) which pass the winter
in a resting condition in the live wheat plant. He marshals a large array of data,
drawn from his own observations and experiments and from a wide range of
literature, to show that the first method, although it exists, is by no means uni-
versal, that the second is highly probable, that the third never occurs in northern
regions, if anywhere, and that the fourth is the most common method everywhere.
Although the conclusions of the author will not be accepted by most investigators
of this difficult problem, yet the array of data is interesting. Two clearly drawn
gans of Brongniart’s
19 Kipston, RosBeERT, On
Histoire des végétaux jossiles. Trans. mele Soc. as 4r: 533-550. pls. 1-3.
1905.
a9 > ERIKSSON, JaKop, Ueber das vegetative Leben der Getreiderostpilze IV:
nia graminis Pers. in der heranwachsenden Getreidepflanze. Kungl. Sy. Vet.-
tay Handl. 395:1-41. pls. I, 2. 1905.
156 BOTANICAL GAZETTE [FEBRUARY
colored plates are used to show the author’s interpretation of the transformation
of the resting mycoplasm into the mycelium condition of the rust.—J. C. ARTHUR
Light relations at high altitudes—Wirsner’s study of the Lichtgenuss of
plants, already comprehensive for varying latitudes, has now been extended?!
to include high altitudes. During a period of thirty days from Aug. 16, photo-
metric observations were made in the Yellowstone territory at eight altitudes
ranging from 515 to 2210™ above sea level. The investigation shows that the
behavior of plants with advancing latitude does not agree with that manifested
under increasing altitude. The relative amount of available light appropriated
by arctic plants increases inversely with the distance from the pole. This relation
holds with increasing altitude only to a certain limit, above which a smaller and
smaller share of available light is appropriated. The cypress habit of growth
is evidently intended to protect from increased intensity of light, whether this
accompanies low latitudes or high altitudes. This seems all the more probable
heat, which is manifested by other species that do not show it at lower levels.—
Raymonp H. Ponp.
Tomato rot.—Von Oven”? has recently described a disease of tomatoes caused
by Fusarium rubescens Appel & Von Oven. This fungus causes a rotting of
the tomato fruit, and evidently does not belong to the fungi in this group producing
stem rot or wilt disease, although in cultures the pink and violet shades char-
acteristic of the latter are also produced by this new species. As it is impossible
to separate the species of Fusarium on morphological grounds, von OVEN has
attempted to distinguish this species at least from several disease-producing
fusariums by their physiological characteristics. It is thus distinguished from
F. Solani, F. putrefaciens, and F. rhizogenum. In cultures on sterilized potato
small sclerotia were formed, which produced conidia after being exposed during
December and January. The author concludes that this is a hibernating stage
of the fungus, although he does not mention finding them in nature—H. HassEL-
BRING.
Axillary scales of aquatic monocots.—As aquatic monocotyledons are by
some held to be modern representatives of the more primitive angiosperms; 4S
these forms may have been genetically related to some such type as Isoetes; and
as he regards the ligule as an important phylogenetic organ, Grsson?3 has made
a study of the vestigial structures of the following families: Potamogetonaceae,
21 WIESNER, J., Untersuchungen iiber den Lichtgenuss der Pflanzen im Yellow-
— und in anderen Gegenden Nordamerikas. Sitzungsber. Kaiserl. Akad.
Wiss.
. Wien, Math.-Naturw. 2 lee II4':(pp. 74.) figs. 2. ; :
22 ne E. von, Ueber eine Fusariumerkrankung der Tomaten. Landw.
Jahrb. 34:489-520. pls. 5, 6. fig. I. 1905.
23 Gipson, R. J. Harvey, The axillary scales of aquatic monocotyledons. Jour.
Linn. Soc. Bot. 3'7:228-237. pis. 5, 6. 1905.
1906] CURRENT LITERATURE 157
Aponogetonaceae, Juncaginaceae, Alismaceae, Butomaceae, and Hydrochari-
daceae. From an investigation of, adult structure and manner of development,
he has concluded that the axillary scales found at the bases of the leaves in the
plants of these genera are homologous with the more specialized and solitary
stipules of Selaginella and Isoetes. It will be recalled that Grsson regards the
ligule as a sort of specialized ramentum, protecting and keeping moist the young
leaves and growing apex of Selaginella and Isoetes.—FLORENCE Lyon.
Reserve food of trees.— NiKLEWSKI?4 confirms by macrochemical methods
the observation of Russow and of FiscHeER, that in winter the fat-content of
trees first increases and. then dimirishes. The process cannot be reversed by
temperature changes. While a rise of temperature accelerates the formation of
fat, no change affects its solution. The transformation of fat and of starch are
not related. Low temperatures promote the enim: of sugar from starch.
Complex phenomena result from a rise of temperature. So great is the loss of
reserves by the sedis a that ee seems = ba sone a other
than starch or fat share it Ca
‘ Conjugation of yeasts.—GuILLIERMOND’S has extended his studies on the
conjugation of yeasts to several additional forms of the Schizosaccharomyces
and Zygosaccharomyces. The union of the cells is followed by the fusion of the
two nuclei, after which the fusion nucleus divides and the two cells separate or
spores are formed in the fusion cell. In some forms conjugation takes place with
the germination of the spores. GUILLITERMOND regards this cell and nuclear
fusion as a sexual act, but of course chiefly on physiological grounds. Since we
do not know the history of the yeasts, it is a matter of speculation whether or
not these conjugating cells are phylogenetically gametes.—B. M. Davis.
Amphispores in Uredineae.—ArTHuR has given an account of all species of
rusts which have amphispores,”° 7. e., as defined by CARLETON, one-celled spores
which resemble the teleutospores of Uromyces in appearance, but have two or
more germ-pores, and in germination behave like uredospores, their function
seeming to be to tide the fungus over unfavorable conditions. This account
includes one species of Uromyces and eight of Puccinia, one of which, P. Garrettit,
is new. All the forms are American, for thus far no cases of the occurrence of
-amphispores have been reported from other parts of the world_—H. HassELBRING.
Photosynthesis | extra vitam.—BERNarD has again examined carefully the
24 NIKLEWSKI, B., Untersuchungen iiber die Umwandlung einiger stickstoffreier
Reservestoffe wahrend der Winterperiode der Baume. Beihefte Bot. Centralbl.
gt: 68—
UILLIERMOND, M. A., Recherches sur la germination des . et la con-
siti chez les lévures. Rev. Gén. Bot. 1'7:337-376. pls. 6-9. figs. IT. 10905.
26 ArtHuR, J. C., Amphispores of the grass and sedge rusts. Bull. Torr. Bot.
Club 32°35-42. figs. g. 1905.
158 BOTANICAL GAZETTE [FEBRUARY
question of photosynthesis im vitro, and again with negative results.27_ He repeated
Maccuiatr’s experiments (following his directions in litt.), and tried also those
of Moutscu, which lent faint support to Maccurati’s conclusions. The gas
disengaged seems due only to bacterial infection and when obtained at all does
not conform in amount to that demanded by theory. This accumulation of
negative results makes exceedingly doubtful the claims of FRIEDEL and MAc-
carati.-~-C. K.-B.
Measuring transpiration CaNNon describes? a method of studying the
rate of transpiration upon plants in place, which he calls the polymeter method,
because LaMBRecH?’s portable polymeter, a combined hygrometer and ther-
mometer is used to ascertain the increase in humidity of the atmosphere around
the experimental plant when enclosed in a bell jar. Certain defects in the
method are noted, but the most important one, that it itself produces a variable
decrease in transpiration, is not mentioned.— C. R. B.
Diastase.— KLEEMANN, finding the known methods of determining the
course of diastase formation not sufficiently accurate, proposes a new, and, as he
claims, more satisfactory one.?? Using it he has determined that the amount of
diastase formed depends, on the one hand, upon the water content of the barley,
and on the other, upon how the water is supplied and taken up, and that the
loss by respiration is greater the greater the water content.— C. R. B
The sporophyte of mosses.—TRUE finds’° that the nodding of the capsusel
of Mnium, and probably of Funaria also, is due to geotropic stimulation, while
the direction of illumination determines the plane of the curve in the seta, the
apex of the capsule sometimes curving toward and sometimes away from the
incident light. The calyptra affords important protection to the growing sporo-
phyte from mechanical injury and desiccation —C. R. B.
Chloroform a stimulant— So Miss Latham: finds it in small quantities to
Sterigmatocystis, especially at the time of germination, while larger quantities
are inimical or fatal. Less acid formation and less sugar consumption under the
stimulus indicate greater metabolic economy.—C. R. B
Chromosome reduction.—A useful collective review of the recent literature on
this subject is presented by K6rNIcKE in Bot. Zeit. 63?: 289-307. 1905.—
27 BERNARD, C., Sur l’assimilation chlorophyllienne. Beihefte Bot. Centralbl.
IQ':59-67. 1905.
28 CANNON, W. A., A new method of pes a the transpiration of plants in
place. Bull. Torr. Bot. Club 32: 515-529. 1905.
29 KLEEMANN, A., Untersuchungen iiber ie Landw. Versuchsstat. 63:
93-134- 1905.
3° TRUE, R. H., Notes on =f sanity of the sporophyte of Funaria and Mnium.
Beihefte Bot. Centralbl. I9':3
3t LatHam, M. F.., ea Z ae by chloroform. Bull. Torr.
Bot. Club 32: 337-357. 1905.
NEWS.
Dr. Enrico PANTANELLI has been appointed docent in botany at Rome.
EMILE Boupier, the eminent mycologist, has been elected director of the
Association internationale de géographie botanique for the year 1906.
PRorEssor Dr. A. RicHTER has been appointed director of the botanic
garden of the University of Kolosvar, the post recently vacated by the death of
Professor V. BorsAs.
A Portrait of Mr. Francts Darwin was lately presented to the botanical
department of the prrsbrnc hs: of Cambridge, where he was for ‘many years.an
active investigator and instruct
PRoFEssor Huco De Vries ae sail for New York about’ April 1, to deliver an
address at the bicentennial anniversary exercises in honor of BENJAMIN FRANK-
LIN to be held in Philadelphia April 17-20, under the auspices of the American
Philosophical Society. He expects to remain in this country two or three months.
Dr. D. T. MacDovaat has resigned his position as assistant director of the
New York Botanical Garden and has been appointed director of botanical
research of the Carnegie Institute. Dr. B. E. Livincston has resigned his post
as physiologist in the Bureau of Soils, U.S. Department of Agriculture, and
Professor Francis E. Luoyp his chair in the Teachers College of Columbia
University, to accept appointments as investigators on the staff of the Desert
Botanical Laboratory, with Drs. CANNoNn and SPALDING.
AFTER thirty years’ service Sir W. THIsELtoN-Dyer retired on December
15 from the directorship of the Royal Botanic Gardens, Kew, and was succeeded
by Lieutentant-Colonel D. PRAIN, formerly director of the Botanical Survey of
India, and superintendent of the Royal Botanic Gardens, Calcutta. Mr. Dyer
will remain at Kew till March 31 next, and till that date will continue to act as
botanical adviser to the secretary of state for the colonies and as technical
adviser in botany to the Board of Agriculture and Fisheries, as well as to take
charge of India Office work.
From THE Journal of the New York Botanical Garden we learn that Mr.
R. S. Wittrams has returned from two years’ explorations of the Luzon, Jolo,
and Mindinao, three of the Philippine Islands, bringing large and important
collections of herbarium and museum material, estimated at ten to twelve
thousand specimens, in spite of the loss of about three months’ collections by fire.
R. J. N. Rose with an assistant, Mr. PAINTER, spent the summer in the
arid districts of central and southern Mexico, collecting cacti, of which they
159
160 BOTANICAL GAZETTE [FEBRUARY
secured several hundred. Special arrangements are being made to study this
family thoroughly, both in living and preserved material.
me giant bamboos in the palm-house in the past season grew 65 feet (20™)
in ninety-five days, an average of about 21°™ per day.
ROM advance sheets of the seventeenth annual report of the Missouri Botani-
cal Garden, we learn what extraordinary burdens the SHAw bequest has been
carrying these sixteen years in the way of taxes, general and special, and real
estate and street improvements. This has unhappily delayed the design of
Director TRELEASE for development of the Garden as a research center, making
impossible the prompt execution of the plan to maintain a staff of specialists and
furnish them facilities for work. If the city and state were as just as Mr. SHAW
was generous they would relieve the Garden of taxes at least, since it exists
solely for the public good. Notwithstanding these unexampled inroads upon its
income the institution has not stood still; the garden has not only been main-
tained but greatly improved; a fine library and herbarium has been ‘accumulated,
and notable researches have been published annually. The grounds now embrace
65 acres, the plant houses cover 30,000 square feet, the cultivated plants number
16,000 species, noteworthy groups being the cacti (678 sp.}, bromeliads (204 sp.),
and orchids (942 sp.). The library isnow undoubtedly the best botanical library
in the United States, and the herbarium contains over half a million specimens.
We congratulate the Director and Trustees on the wise administration of their
trust in the face of serious difficulties and discouragements.
THE American Mycological Society held its third annual meeting in connec-
tion with the American Association for the Advancement of Science at New
Orleans, January 1, 1906. In the absence of the president, CHartes H. PEcK,
the vice-president, F. S. EarLe, preside e new constitution recommended
by the joint committee of the Botanical Society of America, the Society for Plant
Morphology and Physiology, and the American Mycological Society, as a basis -
for the union of the three societies, was adopted and the present officers
. ART
reasons for desiring a better classification of the Uredinales; S. M. TRACY,
Uredineae of the Gulf States; W. G. FArtow, Some peculiar fungi new to
America; F.S. Earte, North American gill fungi; Bruce Finx (by title),
Lichens and recent conceptions of species; E. M. FREEMAN, The affinities of the
fungus of Lolium temulentum; C. L. SaEar, Peridermium cerebrum Peck, and
A Nps Quercuum (Berkeley); C. L. Sear, Romularia: An illustration of
the present practice in mycological nomenclature; P H. Roirs, Notes om
cultures of Collelotrichum ‘and Gloeosporium; P. SPAULDING, The occurrence of
Fusoma parasiticum Tubeuf in this country; P. H. Rotrs, Notes on tes
cocos; P. H. Roirs, Penicillium glaucum on pineapple fruit. —C. L. SHE
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A Decade of Givic Development
By CHARLES ZUEBLIN
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thor of American Municipal Progress
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Vol. XLI, No. 3 Issued March 31, 1906
CONTENTS
A MORPHOLOGICAL STUDY OF SARGASSUM FILIPENDULA. ConrrIBUTIONS FROM
HE HULL BoranicaL LABORATORY LXXXII (wirH PLATES xX AND x1). Etoile B. Simons 161
CHROMOSOME REDUCTION IN THE MICROSPOROCYTES OF LILIUM
TIGRINUM (wirH PLATES xII AND xt). John H. Schaffner 183
CYTOLOGICAL STUDIES ON THE ENTOMOPHTHOREAE. I. THE aaa ce
D DEVELOPMENT OF EMPUSA (WITH PLATES XIV AND Xv). Edgar W. Oliv 192
BRIEFER ARTICLES.
NEw pi Aco APPLIANCES FOR USE IN PLANT PaHysroLocy. III (WITH TWO FIGURES).
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VOLUME XLI NUMBER 3
BOTANICAL GAZETTE
MARCH, 1906
A MORPHOLOGICAL STUDY OF SARGASSUM FILI-
PENDULA.
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY.
ETOILE B. SIMONS.
(WITH PLATES X AND XI)
THE family Fucaceae is less understood than its position and prom-
inence in the Phaeophyceae warrant. Many important types have
scarcely been considered at all, and, moreover, aside from the com-
paratively recent cytological studies in the family, few investigations
have been conducted with modern methods of technique. The prob-
lems of morphology and cytology in the Fucaceae center chiefly around
the sexual organs; the peculiar sunken structures in which they are
borne, termed conceptacles; the likewise sunken but sterile structures
called cryptostomata; and the sporelings.
The present investigation of these structures in Sargassum filipen-
dula Ag., a member of perhaps the most highly differentiated genus in
the Fucaceae, was undertaken with the hope of filling some of the.
obvious gaps in our knowledge of this family. It was conducted in
the University of Chicago and at the Marine Biological Laboratory,
Woods Hole, Massachusetts, under the direction of Professor BRADLEY
Moore Davis, who suggested the research to me. It gives me pleasure
to express here, both to him and to Professor JoHN MERLE COULTER,
my appreciation of valuable suggestions and assistance given me in
this work. My acknowledgments are also due the Carnegie Institu-
tion for the use of a table at the Marine Biological Laboratory during
the summer of 1904.
References to anatomical and morphological work which concern
161
162 _ BOTANICAL GAZETTE [MARCH
this subject will be given under the topics to which they belong. The
early history of the genus with its taxonomic bearing is omitted, as
having no place here, but the once credited distribution of Sargassum
which was convincingly disproved by Kuntze (’81) is a matter of his-
tory which.deserves at least brief mention.
Kuntze relates that LINNAEUS believed that a vast area of sea was
densely covered by Sargassum in active vegetative condition; Hum-
BOLDT reported that the region surpassed Germany in extent six or
seven times; Maury stated that it equaled the Mississippi valley; and
HOECKEL estimated its area to be forty thousand square miles.
That these views were generally accepted is well known. They led
to instruction regarding a ‘‘Sargasso Sea,’ whose supposed limits
were outlined upon maps of the world. Kuntze, by comparing
his own observations and those of other travelers over routes which
crossed in different places the outlined area, was able clearly to dis-
prove the existence of such a “‘sea.”” Sometimes a voyage was made
through the mapped region and little or no Sargassum was seen, and
again it appeared somewhat abundantly, but without definite limits
or fixed location. Storms which sweep tropical shores, near which
attached Sargassum grows abundantly, were found to be in great part
accountable for the appearance of the larger quantities of floating
Sargassum. Kuntze obtained no evidence to substantiate the view
that floating Sargassum vegetates. It had been believed that floating
forms of Sargassum consisted of S. bacciferum only, but Kuntze found
several species floating, and observed that the specimens in herbaria
which had been collected in mid-ocean and labeled Sargassum bacct-
ferum according to general belief, could be referred to various species.
He therefore concludes that there is no characteristic floating species.
The appearance in mid-ocean of floating masses now and then does
not seem strange when the authentic distribution and abundance of
attached Sargassum are recalled. According to KyELLMAN (’93) this
genus, which includes one hundred fifty species, over half the number
belonging to the entire family, is found attached along the coast of all
warm seas, reaches north to Cape Cod in the Atlantic, to Japan in the
Pacific, and in the south into Australian waters, where it is the most
abundant. With the extent of this distribution in mind the presence
of floating masses, especially after storms, is to be expected.
a Te a ea ach al er ae a a DEO eds WoW rh tes
1906] SIMONS—SARGASSUM FILIPENDULA 163
MATERIAL AND METHODS.
Material for this study was collected near the shores of Woods
Hole, late in July and during August. Plants both in vegetative and
in reproductive conditions were abundant. The weak solution of
chromacetic acid of Flemming (1 per cent. chromic acid 25°°, 1
per cent. acetic acid 10°°, water 65°°) proved a satisfactory killing
and fixing reagent. Microtome sections were cut from paraffin 5 » in
thickness and stained either by iron-alum-haematoxylin after the
method of Heidenhain or by safranin and gentian violet. The mucil-
age on the surface of the plant and in young conceptacles and cryp-
tostomata takes the anilin dyes readily, but is not especially trouble-
some.
GENERAL MORPHOLOGY AND HISTOLOGY.
The habit of Sargassum filipendula is so like that of other species
which have been described that it needs but slight attention. This
species grows attached to rocks below low water mark, and therefore,
unlike Fucus and Ascophyllum, is never exposed to the air. Vegeta-
tive plants and reproductive plants bearing all stages of conceptacles
are plentiful in summer. Sporelings are abundant also and easily
collected, for the discharged eggs and their products, the sporelings,
remain attached for some time by mucilage to the surface of repro-
ductive branches near the parent conceptacles.
The stem arises from a small disk-shaped holdfast and passes into
long cylindrical branches which bear spirally arranged leaves, berry-
like floats, which seem to be modified portions of teaves, as generally
stated, and short reproductive branches. This form may attain a
height of 60°™, but is commonly shorter. Cryptostomata develop
upon stems, leaves, and occasionally also upon reproductive branches
in Sargassum, which differs in this respect from Fucus, whose recep-
tacles, according to Bower, contain no cryptostomata.
KJELLMAN (’93) states that the conceptacles of Sargassum are her-
maphrodite. In Sargassum filipendula both mature bisexual and
unisexual conceptacles are formed. Some conceptacles contain only
spermatocysts (antheridia); some, more rarely, contain many sper-
matocysts and but one or two oocysts*(oogonia); and others bear
only oocysts. The appearance of a conceptacle devoted to the forma-
tion of oocysts differs decidedly from such a structure in Fucus. In
164 BOTANICAL GAZETTE [MARCH
Sargassum the oocyst has no stalk cell. It is an embedded organ,
being almost surrounded by wall cells of the conceptacle. As both the
size and contents of a conceptacle are dependent upon the activity of
wall cells (as described later), this conceptacle in Sargassum is smaller
and has fewer sexual organs and paraphyses than the corresponding
conceptacle in Fucus. The unisexual tendency in the conceptacle of
Sargassum may be due in part to the unproductiveness of the many
wall cells which abut upon the embedded oocyst.
The anatomy of the thallus of Sargassum has been studied in four
species. In 1876, REINKE reported its development in Sargassum
Boryanum from a three-sided apical cell situated at the bottom of a
pit in the apex of the stem. He stated that the holdfast is composed
of rhizoids and that a few intercellular filaments occur in the old parts
of the thallus. OxtManns (’89) in an anatomical investigation of
Sargassum linifolium and S. varians, likewise described a three-sided
apical cell, and in addition gave an account of the origin both of the
apical cell of a leaf and of a branch. He believes that the branching
in Sargassum holds no relation to dichotomy. He figures an enlarged
epidermal cell near the apical cell of the stem, and states that it
becomes a three-sided apical cell. This young cell develops an out-
growth in which a second apical cell is soon differentiated, between
the first and the stem. The first formed apical cell develops a leaf
and the last a branch. OLTMANNs agrees with Kuntze (’81) that
there are all gradations between leaves and floats, and that floats are
modified portions of leaves.
In 1892, HANSTEEN published the results of an anatomical and
physiological investigation of Sargassum bacciferum. He also reported
a three-sided apical cell, but did not trace its origin in any structure.
He described three kinds of tissues, naming them the assimilating
system, the storage system, and the conducting system. The assimi-
lating system, according to HANSTEEN, includes only the outer layer
of cells, or epidermis. Its cells are twice as long as broad, have undu-
lating walls, like the epidermal cells in higher plants, and contain
“‘phaeoplasts.” The cells of this system add to their own number by
radial, and to the cells below by tangential, divisions. The storage
system occupies a zone several cells wide between the assimilating
system and the innermost tissue which constitutes the conducting
1906] SIMONS—SARGASSUM FILIPENDULA 165
system. Most of the cells in the storage system are large. HANSTEEN
found them empty in alcoholic material of Sargassum, but he did not
doubt their function to be that of storage, because he had found much
reserve material in similar cells of living Fucus. The conducting
system consists of an axial cylinder of long cells with small diameter
and oblique end walls. These cells are believed by HANSTEEN and
others to function as sieve tubes. The cells of the three aes
communicate by pores.
HANSTEEN observed in the storage cells of Fucus serratus a
several other types, spherical grains of different sizes, which he named
fucosan. He believes that the same structures have been variously
considered as fat, proteid, and starch by other observers. The grains
do not stain blue with iodin, and are soluble in water. HANSTEEN,
who made a chemical analysis to determine their composition, con-
siders them as a carbohydrate with the formula (C;H,,O,),. CRaTo
(°92) described in Chaeto pteris plumosa spherical or elliptical bladder-
like structures which he named physodes. He reported (’93) that
they contain phloroglucin as a constant ingredient, function in direct-
ing the chemical exchange and transportation of food material within
the cell, have motion, and are independent cell organs like the nucleus
and chromatophore. Crato stated further that HANSTEEN had
confused various cell contents, and that fucosan grains and physodes"
are the same. KocH (’96) denied the presence of phloroglucin in
these bodies. Ina later paper HANSTEEN (:00) again discusses fuco-
san grains. He maintains that CRATO’s physodes are fucosan grains,
and that they are not independent cell organs but products of the
phaeoplast. HANSTEEN has made no further chemical analyses to
determine the nature of the bodies, but holds that they surely repre-
sent a product of photosynthesis. HANSEN (’95) after an investiga-
tion of several forms (Dictyota dichotoma, Taonia alomaria, Haly-
seris polypodioides, Asperococcus, Hydroclathrus, and Cystoseira),
states that the Phaeophyceae contain oil and no starch, and OLTMANNS
(:04) expresses the same view. It is seen therefore, that the character
of the reserve material in the cells of the Phaeophyceae is still some-
what problematical.
Every stem and leaf structure in Sargassum filipendula, as in other
species studied, develops through the activities of a three-sided apical
166 BOTANICAL GAZETTE [MARCH
cell. The tissue systems described by HANSTEEN are present and
each seems to have the function ascribed to it, although without
_rigidity. . Each system, too, has its origin in the group of segments
surrounding the apical cell and can be traced very near it. The cells
of every system are meristematic in the apical region, but the epider-
mal cells are apparently the only ones which retain this activity.
The cells of any one of the three systems correspond well in general
appearance with the similarly placed cells described by HANSTEEN,
but an interesting modification was observed in the cells of the con-
ducting system. All are long and of small diameter, but in respect
to thickness of walls the tissue is differentiated into two regions. The
inner cells have thin walls, while the outer ones have thick walls. The
thick-walled cells may be both supporting and conducting in func-
tion. The conducting system of a leaf blade consists only of thin-
walled tissue. No intercellular filaments, as reported by REINKE,
have beenfound. Sometimes, however, a filamentous alga creeps into
the mucilaginous walls of cells near the surface of a leaf or old stem,
and gives the appearance of intercellular filaments. As the little alga
contains true starch, its cells when stained with iodin present a
sharp contrast to the unstained cells of Sargassum. HANSTEEN
_ (792) figures pores in thin areas consisting of the middle lamella in
Sargassum baccijerum, and REINKE (’76) represents similar areas
but without pores in cell walls of Fucus vesiculosus. Such thin areas
are common between cells in the tissues of Sargassum filipendula,
but pores, though probably present, are rarely seen.
The character of the reserve material in Sargassum proved of great
interest. Sections from plants which have been preserved in forma-
lin contain much more stored material than tissues which have been
kept in alcohol. Preparations, however, which have passed through
alcohol, xylol, paraffin, the heat of the bath, etc., still contain within
the cells of the epidermis and outer cortex, many bodies which in all
probability represent reserve food material. These’ bodies, which
stain readily, vary in size and structure, but are evidently related, for
transitional stages can be found between the most extreme forms.
Judging by the appearance of the structures,some are intact and others
modified. Those which seem intact are spherical, with a diameter
which equals or exceeds the length of a chromatophore. Each con-
1906] SIMONS—SARGASSUM FILIPENDULA 167
sists of a more or less homogeneous ground substance and one or more
refractive areas which are somewhat centrally placed. The modified
structures vary from spheres, whose ground substance has been
changed only at the periphery, to swollen masses which have an
entirely modified ground substance with an irregular outline. Both the
intact and modified bodies may occur within the same cell; but the
former and the least modified are more common in epidermal cells,
whereas the most modified are in cortical cells. The occurrence of
such bodies within epidermal cells where photosynthesis is the most
active, suggests that they represent a manufactured food. The varied
modifications in the structures indicate the solvent action of the killing
fluid, or an intercellular enzyme. As the inner cells contain bodies
presenting greater modifications than the epidermal cells, the agent
producing the change is apparently applied from within the tissue. If
then within, it is probably an enzyme, for a solvent used in the process
of killing would attack the contents of epidermal cells, doubtless before
any others. The intact bodies may represent a newly formed product,
perhaps a carbohydrate, and the modified structures, the product in
process of digestion. ‘The bodies do not stand with iodin in any con-
dition. If they are carbohydrate they probably differ as much or
more from the starch of higher plants as does inulin. The presence of
many small spheres in formalin material and their absence from tissues
preserved in alcohol indicates that oil globules are present in the cell,
in addition to the structures described above. Future investigations
on living material will probably disclose the presence of both oi
and a carbohydrate in the Phaeophyceae.
THE ORIGIN AND DEVELOPMENT OF THE CONCEPTACLE,.
The conceptacle in the Fucaceae had been but little studied when
Bower (’80) gave an account of its development in four genera
and six species (Fucus serratus, F. platycarpus, F. vesiculosus, Ozon-
thallia nodosa, Halidrys siliquosa, and Himanthalia lorea). Accord-
ing to him the development of the conceptacle in every species con-
forms to one scheme with minor variations.
The “‘initial cell’? of the conceptacle, as stated by Bower, is the
terminal cell of a linear series which is produced by a modification of
the regular divisions in the segments of the apical cell of a receptacle.
168 BOTANICAL GAZETTE [MARCH
This initial cell, strangely, contributes nothing essential to the concep-
tacle. It either degenerates directly without having divided at all, or
it produces a short filament whose terminal portion degenerates. A
cortical cell below the initial is termed by BowER a “‘basal cell.”” This
cell and others which adjoin the initial cell laterally, divide and form
the walls of the conceptacle from which the sexual organs and paraph-
yses arise. The initial cell, therefore, according to BoWER, takes no
part in the development of the conceptacle, whereas the cells adjacent
to the initial produce all that is important, the walls and their products.
The prominent features of this scheme for the development of the con-
‘ceptacle are, it is seen, degeneration of an unimportant initial cell or
a part of its filamentous product, and the activity of cells adjacent
to the initial in producing the entire conceptacle.
Nearly all contributions in this field since 1880 have been in the
main confirmatory of the work of BowER. VALIANTE (’83) states that
the development of the conceptacle in Cystoseira is due to the growth of
neighboring tissue, about one or two cells. OLTMANNS (’89) describes
the walls of the conceptacle of Halidrys siliquosa, Himanthalia lorea,
and Ascophyllum nodosum, as also formed by neighboring cells, with
the one exception that in Ascophyllum the initial cell develops a mass
of tissue in the base of the conceptacle. This tissue, he reports,
shares with the rest of the inner surface formed from neighboring tis-
sue, in developing the sexual organs. As no degeneration of tissue
was observed in Ascophyllum, and as its initial cell does contribute
some important tissue the development of the conceptacle, this genus
presents an exception to a part of the scheme which Bower reports.
Although Sphlachnidium should no longer be included in the Fuca-
ceae, as shown by the Misses Mirc#ELL and WHITTING (’92), it is of
interest to note that these investigators report its conceptacle as devel-
oping by the radial division of cells adjacent to a persistent but incon-
sequential element, which they believe to be homologous with the initial
cell of Bower. GRUBER (’96) states that the conceptacle of Sezro-
coccus axillaria is more like that of Halidrys than Ascophyllum, which
means, again, that it has an initial cell which contributes nothing of
consequence to the conceptacle, whose walls are formed by cells which
are adjacent to the initial.
- Hotz (:03) reports that in the development of the conceptacle of
1906] SIMONS—SARGASSUM FILIPENDULA 169
Pelvetia jastigiata several epidermal cells cut off basal segments which
divide transversely until six or more tiers are formed. Over these
tiers, one or more epidermal cells break down and a cavity results,
which is gradually enlarged by further disintegration of epidermal and
meristematic cells. After a time this process ceases, and a “healthy
surface”’ is formed from the deeper meristematic cells. This surface,
which comprises the walls of the conceptacle with the exception of the
upper part that is formed by “cortical rows”’ of cells, produces sexual
organs and paraphyses. The prominent features which distinguish
the conceptacle of Pelvetia from others, as thus described, are the
presence of several epidermal or initial cells, the more extended disin-
tegration of tissue, and a difference in the behavior of the basal cells.
The development of the conceptacle in Sargassum filipendula is at
variance with all the prominent characteristics in the development
of the conceptacle as described by Bower. The initial cell of Sar-
gassum does not break down. It is an active cell which produces the
entire conceptacle. As the whole conceptacle is the product of this
one cell, adjacent cortical tissue takes no part whatever in the devel-
opment of the structure. The first indication of the conceptacle is a
clearly differentiated epidermal cell which lies near the apical cell of a
reproductive branch (jig. 7) and constitutes the initial cell of the con-
ceptacle. The upper portion is surrounded laterally by epidermal
tissue, whereas its central and basal regions are bounded by cortical.
The initial is much larger than any of the cells with which it is in con-
tact and differs much from them in shape. Though it may vary some-
what in length it is always flask-shaped. Its oval bowl, sometimes
slightly narrowed at the base, tapers above into an elongated neck
whose outer end is flush with the surface. The initial cell is circular
in cross section at its apex (fig. ra) and elliptical at its base (fig. 1b).
The initial cell never breaks down. On the contrary the develop-
ment of the conceptacle is initiated by its activity. Its large nucleus
divides. Then a curved wall is formed with concave surface above,
separating two very unlike cells (fig. 2), which form the two-celled stage
of the conceptacle. The upper cell, which may be designated the
tongue cell, is a long somewhat cylindrical structure; whereas the
lower is somewhat conical or wedge-shaped. The initial cell and the
two-celled stage of the conceptacle have similar outlines both in longi-
170 " BOTANICAL GAZETTE [MARCH
tudinal (figs. r and 2) and in transverse sections (figs. 1a, m, b, and 2a,
m, b).. That the lower portion of the tongue cell is surrounded by the
upper part of the cell below it is well shown in both longitudinal and
transverse sections of the two-celled stage of the conceptacle.
The lower cell of this two-celled structure divides longitudinally
into two similar daughter elements, thus producing the three-celled
stage of the conceptacle (jig. 3). The longitudinal wall reaches to the
lower portion of the tongue cell, whose basal portion is surrounded
now by two cells instead of by one. _ The relative position of the three
cells is made clearer by an examination of their transverse sections.
A cross section near the base of the three-celled structure shows two
similar cells (fig. 3b). A cross section about midway between the
apex and base shows three cells (figs. 3m and 3bm), the tongue cell
and the two lower cells which surround its base. A section of the apex
is circular in outline and consists of the tongue cell alone (fig. 3a). The
three-celled stage of the conceptacle is apparently formed occasionally
in another way. The two longitudinal sections of an initial cell are
shown in figs. 4, 5, containing three nuclei but no walls. Two nuclei
appear in one section and one in the other. It seems that the nucleus
of the initial cell in this instance divided first with its spindle perpen-
dicular to the axis of the cell, and that one of the daughter nuclei
divided with its spindle parallel to the axis.
After the three-celled stage, the development of the conceptacle is
readily followed. The two lower of the three cells divide longitudi-
nally in various planes. A condition thus results which exhibits five
cells in longitudinal median section (fig. 6). Four of the five cells are
young cells of the recent divisions, and one is the centrally placed
tongue cell. Longitudinal divisions continue as before, and a struc-
ture showing six or seven cells in longitudinal section is formed (fig. 7)-
The tongue cell is still conspicuous in this and in several succeeding
stages. Longitudinal divisions continue as illustrated in figs. 8, 9, 11;
until the walls of the entire conceptacle are formed. Some of the
wall cells begin to develop sexual organs when the conceptacle is
very small (figs. 9, 11). - This activity of the cells, however, does not
prevent them from contributing to the growth of the conceptacle.
The mouth of the conceptacle is surrounded by a marginal ring of
epidermal tissue about one or two cells deep (figs. 8, 11). As these
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1906] SIMONS—SARGASSUM FILIPENDULA 171
cells are not aggressive they may be omitted from further consider-
ation. Excluding this minor detail every portion of the conceptacle
is the product of one initial cell. Cortical tissue adjacent to the ini-
tial takes no part in its development.
The behavior of the tongue cell is similar to that of the “initial cel]”’
in other forms as reported by Bower. It may show signs of degenera-
tion (fig. 8), may remain inactive for some time (fig. rr), or may even
divide to form a filament of two or three cells (figs. 9, 10). In-no case
does it contribute to the walls of the conceptacle, but on the contrary
after its divisions resembles a paraphysis. The tongue cell is very :
conspicuous until sexual organs begin to develop, but shortly after
their appearance it cannot be distinguished. The upper and lower cells
which result from the transverse division of the initial cell (fig. 2) cor-
respond in appearance and behavior with the “‘initial cell” and “basal
cell” as described by Bower and others. It seems probable that
Bower saw both the initial cell and the two-celled stage of the concep-
tacle, but failing to observe the division in the initial cell, considered
the initial and the upper cell of the two-celled stage identical. With
this construction, degeneration of the upper or tongue cell was be-
lieved to be degeneration of the initial cell itself, and the division of
the lower cell of the two-celled stage, a product of the initial cell, was
regarded merely as the division of an unrelated cortical cell. A
conceptacle developed from cells which happen to be adjacent to a
degenerating and unimportant cell would be a very different structure
from a conceptacle developed from one active initial cell. |
THE ORIGIN AND DEVELOPMENT OF THE CRYPTOSTOMA,
The references embodied in the preceding treatment of the concep-
tacle constitute the chief source of information bearing upon the cryp-
tostoma. The structure which produces the sexual organs has com-
monly and naturally been given first attention, but investigators who
have studied both, generally agree that the conceptacle and cryp-
tostoma are homologous. Different theories regarding the signifi-
cance of the cryptostoma have been offered, but no safe generalization
can be made until a more extended investigation of both structures
has been made in a variety of forms.
Miss Barton (’91) gave an account of the cryptostoma in Turbi-
172 BOTANICAL GAZETTE [MARCH
naria, stating that an initial cell divides longitudinally, thus forming
two daughter elements which produce paraphyses. In demonstration of
this two paraphyses are figured arising from the base of a many-celled
structure. Miss Barton does not report the origin of the walls
of the cryptostoma, but as the initial cell is believed to develop directly
into paraphyses, we may assume that she believed the walls to arise
from neighboring tissue in accordance with the views of the earlier
writers.
The development of the cryptostoma in Sargassum follows step by
step the history of the conceptacle. The initial cell arises near the
apical cell of a leaf or vegetative branch. Longitudinal and cross
sections of this cell (figs. 12, 12a, m, b) show the same form and struc-
ture as the longitudinal and cross sections of the initial cell of a
conceptacle (jigs. 1, 1a, m, b). The activities of the two initials are
also identical. The initial cell of the cryptostoma divides transversely,
forming a two-celled structure (fig. 13) which is comparable in every
way to the two-celled stage of the conceptacle (fig. 2), consisting as it
does of a tongue cell and a larger lower cell. The lower cell divides
longitudinally. A group of three cells then results (figs. 14, 15) which
is entirely similar to the three-celled stage of a conceptacle (fig.3).
The two lower cells of this three-celled stage then divide longitudi-
nally in one or more planes, forming a structure which shows four
or five cells in longitudinal section (figs. 6, 17). The center of this
structure and of several which follow is occupied by the conspicuous
tongue cell (figs. 16, 17, 18, 19). Thus by the continued longitudinal
divisions of the products of the lower cell of the two-celled stage, the
walls of the entire structure are gradually developed. Paraphyses
begin to appear in the cryptostoma (jigs. 18, 20) as early as do the
sexual organs in the conceptacle (fig. g). The activity of the wall
cells in producing paraphyses, however, does not interfere with their
functioning further in developing the cryptostoma (fig. 21). Enpider-
mal cells at the mouth of the cryptostoma form here, as in the concep-
tacle, a marginal ring one or two cells deep (compare figs. 8 and 19).
The origin of the true walls of the structure, however, may be traced
as in the conceptacle to the lower cell resulting from the transverse
division of the initial.
The behavior of the tongue cell in the cryptostoma is similar to
SC
1906] SIMONS—SARGASSUM FILIPENDULA 173
that of the corresponding element in the conceptacle. Occasionally
the tongue cell of the-cryptostoma may develop a prominent filament
(fig. 20), which is clearly identical in structure with a typical paraph-
ysis (fig. 21). The young conceptacle and cryptostoma are so alike
that they can only be distinguished by their respective positions on
fruiting branches or on young vegetative structures, until the appear-
ance of sexual organs in the one and paraphyses in the other defines
their mature characters.
The development of the paraphysis is interesting for its regularity.
A wall cell enlarges, pushing into the cavity of the cryptostoma, and
then divides transversely (figs. 18, 20). The upper cell produces the
paraphysis, whereas the lower functions in the development of the
wall. The growth of the paraphysis results from the transverse divi-
sions of the cell next the wall (figs. 21, 22), a method of growth termed
trichothallic. ‘The development of a paraphysis in the cryptostoma of
Sargassum is, therefore, characteristically basipetal, as BARTON (’91)
found in Turbinaria. ;
A somewhat advanced paraphysis is composed of three regions.
That which adjoins the wall of the cryptostoma consists of the large
turgescent meristematically active basal cell (fig. 22). The middle
region is occupied by six or eight short cells which have so recently
been formed that they have not had time to lengthen much. The
upper region contains several greatly elongated cells. This region in
a mature paraphysis is partly within the cryptostoma and partly
without, for fully developed paraphyses extend far beyond the sur-
face of the plant.
A peculiar condition found in many cryptostomata deserves special
attention. Structures frequently appear between the paraphyses
which seem to bear no relation to them. These are papillae and
stalked cells, the former like the papillae which precede male organs
in a conceptacle and the latter like the male organs themselves. The
stalked cells, although slender and probably always sterile, appear to
be spermatocysts no longer functional. This surprising condition is
of great interest and importance in relation to the homology and sig-
nificance of the cryptostoma, a structure formerly believed to contain
only paraphyses, but which appears also to have sexual organs or their
degenerate representatives. That the cryptostoma and conceptacle
174 BOTANICAL GAZETTE [MARCH
are homologous cannot be doubted, since: their origin and early
development are identical in all details. The occasional appearance
of sterile representatives of sexual organs within the cryptostoma
further confirms this view of their relationship and strongly supports
the theory of Bower (’80) that the cryptostoma in the Fucaceae is
derived from the conceptacle.
The occurrence of conceptacles in special reproductive branches
only, the appearance of cryptostomata in both vegetative and repro-
ductive branches, and the development of representatives of sexual
organs within the cryptostomata, suggest a line of evolution from
plants bearing conceptacles scattered over leaf and branch struc-
tures indiscriminately, to the type now under consideration with
localization of the conceptacle upon special branches. Certain
branches were set apart to bear conceptacles as the conceptacles in all
other parts of the plant body were rendered sterile and thus changed
into cryptomostata. The presence of sexual organs or their degenerate
representatives within a cryptostoma indicates, according to these
views, that the process is not carried to its farthest point in Sargassum.
The production of conceptacles upon small special branches only,
instead of upon the entire plant, naturally results in fewer concep-
tacles upon one plant. The conceptacles, however, are much more
closely placed than the cryptostomata. On account of their com-
paratively small size the initials and young conceptacles occupy very
little space in the apex of a branch, but farther down on the receptacle
the bulging bowls of the developing flask-shaped conceptacles require
more and more space, until the mature structures nearly fill the
interior of the receptacle and there is only enough intervening tissue
to hold the conceptacles together. The cryptostomata, on the other
hand, are well scattered upon vegetative branches and mature leaves.
The contrast in the placement of cryptostomata with that of concep-
tacles is, therefore, very marked.
THE SPERMATOCYST.
The male sexual organs (antheridia), which will be called sper-
matocysts in this paper, according to the terminology of DAvis (:04),
develop from wall cells of the conceptacle in Sargassum as in other
forms of the Fucaceae. A wall cell puts forth a papilla (fig. 23) which
I a a i ti I 2
1906] SIMONS—SARGASSUM FILIPENDULA cy gs
is cut off by a transverse wall (jig. 24). The lower cell becomes a
part of the wall occupying the place of the cell from which it arose.
The upper cell enlarges for a time and then divides, forming the
sperm mother-cell or spermatocyst and its stalk (jig. 24, at the right.)
A stalk cell may have no other relation than that which it bears to
the spermatocyst which it supports, or it may function in other ways.
It may produce several spermatocysts directly, without individual
stalks; it may put forth a papilla which gives rise to a spermatocyst
and stalk (figs. 25, 26); or it may develop a hair (fig. 27). Hairs,
however, are comparatively rare within a conceptacle of Sargassum.
Owing to the variety of activities which belong to a stalk cell, the
growths within a conceptacle lack uniformity. Some structures reach
but a little distance above the wall of the conceptacle, whereas others
form conspicuous branch systems. Although these systems are prom-
inent in this conceptacle, they are considerably smaller and less dense
than the branch systems in a conceptacle of Fucus, and there is far
more unoccupied space within the cavity of a conceptacle of Sargassum
than of Fucus.
The young spermatocyst contains dense cytoplasm, a centrally
placed nucleus and deeply staining granules, the nucleus remaining in
a resting condition for a long period. The divisions of the nucleus
were not studied in detail. Sixty-four sperms are apparently formed
(figs. 27 and 28), agreeing, therefore, with the count announced by
BEHRENS (’86) for Fucus vesiculosus. The sperms within the
spermatocyst have an elliptical outline, a cytoplasmic ground mass,
and a somewhat spirally arranged band, which is probably the nucleus.
The discharge of sperms was not seen, but a rent, partly terminal and
partly lateral in empty spermatocysts, indicates their mode of escape.
THE OOCYST.
The female sexual organ (oogonium) or oocyst, according to the
_ terminology of Davis (:04), is peculiar among the Fucaceae, as far as
is known, in that it is not borne upon a stalk but is a partially embed-
_ded organ (fig. 31). The sister cell of the oocyst, instead of develop-
ing into a pedicel cell as is usual in this family, functions as one of
the wall cells of the conceptacle. The oocyst enlarges greatly, but
becomes nearly surrounded by adjacent wall cells.
176 BOTANICAL GAZETTE [MARCH
Its development is simple. A somewhat enlarged wall cell of a
young conceptacle divides transversely, forming two cells much alike
in size and contents (jig. 29). The inner cell, which is the homologue
of the stalk cell of the female organ in Fucus, cannot be distinguished
from neighboring wall cells shortly after its formation. The outer
cell, which has a free surface toward the interior of the conceptacle,
increases greatly in size and soon becomes the spherical oocyst. Fig.
30 represents a young oocyst and its sister cell, already unequal in
size. There now follows a long period of growth, during which the
oocyst attains a remarkable size, finally containing a great quantity of
reserve material, many chromatophores, much cytoplasm, and a large
nucleus. The mature organ, drawn under a lower magnification than
fig. 30, is represented in fig. 31. No trace of its sister cell could be
found.
The oocyst of Sargassum develops but one egg. The mitosis within
the wall cell whose division produces the oocyst is normally the only
mitosis in the process of oogenesis. Particular attention was given to
this point. The one nucleus of the oocyst remains in a resting
condition throughout the entire period of the growth of the cell, and
therefore becomes the nucleus of the egg. In the other genera of the
Fucaceae, as is well known, there are three mitoses within the oocyst,
resulting in eight nuclei. Each of the eight nuclei may become a
center for the development of an egg as in Fucus, or some nuclei may
degenerate and a less number of eggs be formed, as in Ascophyllum
and Pelvetia. It might be supposed from these conditions in the
Fucaceae that the oocyst of Sargassum would show similar nuclear
divisions and degeneration, but this is not the case. The mitoses
characteristic of oogenesis in Fucus are normally suppressed in Sar-
gassum. The tendency in the Fucaceae to reduce the number of eggs
produced by an oocyst reaches its culmination, therefore, in Sargassum.
It is interesting to note that Sargassum still gives proof that it be-
longs to the reduction series which has its beginning in Fucus and
allied forms that produce eight eggs in an oocyst. Out of the great
number of conceptacles examined, one oocyst was formed which con-
tained two eggs, and two oocysts which contained eight. The oocyst
with two eggs was formed in an immature conceptacle that held five
normal oocysts. The two eggs appeared fairly vigorous. One of the
1906] © SIMONS—SARGASSUM FILIPENDULA 177
two oocysts which contained eight eggs was an old conceptacle, from
which other sexual elements had apparently long been discharged.
The eight together were smaller than one normal mature egg. The
other oocyst which contained eight eggs shared a conceptacle with two
normal oocysts. It was attached in the side of a conceptacle near the
surface of the plant, which for a slight distance was modified in struc-
ture as if in response to an injury. — It is possible in this case that the
wound incited the reversion.. The appearance of an oocyst contain-
ing more than one egg in Sargassum must be regarded as a rare rever-
sion to the Fucus type.
The resting nucleus of the oocyst is always large, but varies in
structure. Sometimes it has few granules and no conspicuous reticu-
lum, whereas at other times it contains many. granules and a dense
network. The nucleolus is also large in size and variable in structure.
At the present time no suggestion can be made to account for the
changes in nuclear structure, excepting that they are the concomitants
of growth and varying nutritive conditions.
The method of discharge of the egg from the conceptacle of Sar-
gassum is somewhat unlike that reported in Fucus and other genera.
In Fucus the outer membrane of the oocyst remains attached to the
conceptacle, as explained by THuRET, and the eggs escape in a group
surrounded by a very delicate inner membrane. In Sargassum the
entire oocyst becomes freed from the conceptacle and escapes. In
Fucus the inner membrane dissolves or breaks, thereby freeing the
naked eggs which it has enclosed. In Sargassum the wall of the oocyst
‘swells, stretches, and sometimes ruptures, but it may persist for a long
time, even enveloping later a many-celled sporeling formed within it.
The inner membrane enclosing the eggs of Pelvetia is separated from
the outer as in Fucus. In Pelvetia, however, as figured by THURET,
this membrane persists about the eggs, apparently offering no great
resistance to the entrance of sperms. Whether the sperm enters the
egg of Sargassum through a break in the oocyst membrane, whether it
passes through the membrane, or whether the eggs develop par-
thenogenetically, isnot known. A study of fertilization in Sargassum
is surrounded by serious technical difficulties because both eggs and
sperms develop upon the same plant, thus making it difficult to isolate
the sexual cell.
178 BOTANICAL GAZETTE [MARCH
THE SPORELING.
Many if not all of the eggs of Sargassum on leaving the conceptacle
become fastened immediately by the mucilaginous wall of the oocyst,
which still surrounds it, to the surface of the reproductive branch. In
this position the eggs segment. ‘The first division of the egg in Sar-
gassum does not differentiate a rhizoidal region, as in Fucus and
Ascophyllum. Instead, a many-celled ellipsoidal structure is formed,
the divisions occurring with mathematical precision. Rhizoids then
develop at one end with no apparent relation to a substratum or to
gravity, so far as could be observed in fixed material. Sporelings
sometimes occur at opposite sides of a branch with rhizoids directed
toward the stem, thus showing no relation in the development of rhi-
zoids to gravity; and again, sporelings occur with rhizoids directed
away from the branch in various directions, indicating that the
parent plant exerts no special influence. It is possible that the
attachment of a sporeling upon a plant is so insecure that the
direction of its axis may be shifted in the manipulation of material.
Otherwise it is difficult to account for the conditions which were
observed.
When the many-celled sporeling has reached the condition for rhi-
zoid formation the cells at one pole elongate, thereby giving rise to a
tuft of rhizoids of approximately equal length. This mass of rhizoidal
filaments finally produces the characteristic disk-shaped holdfast of
the mature plant. Fig. 32 shows a sporeling in about the oldest con-
dition in which it remains attached to the parent plant. No apical
cells were found in these sporelings and therefore its differentiation
must occur after the sporeling has separated from the parent
plant.
The germination of the oospore deserves careful cytological investi-
gation. Many preparations have been made and studied, but further
attention will be given the subject before the observations are pub-
lished. A few conditions may be noted, however. There are numer-
ous radiations at the poles of the early spindles. The asters contain
granular inclusions which suggest centrosomes, although their origin
and relation to the processes of mitosis have not been traced. Walls
following the mitoses are developed somewhat slowly, being formed in
part at least by the membranes of contiguous vacuoles.
1906] SIMONS—SARGASSUM FILIPENDULA 179
SUMMARY.
Each stem, branch, and leaf structure develops through sic activi-
ties of a three-sided apical cell.
The thallus consists of three compact tissues, called for conven-
ience the epidermal, cortical, and conducting tissues. The latter con-
sists of only thin-walled cells in the leaves, but in mature stems con-
tains both thick and thin-walled elements. A ring of thick-walled
cells, which may have both a mechanical and conducting function,
surrounds the thin-walled conducting cells in the center of the
axis.
The tissues normally contain much reserve material, a part of
which is oil, and a part, whose nature is undetermined, appears to be
a carbohydrate.
Both the conceptacles and cryptostomata originate in a single flask-
shaped initial cell which develops the entire structure.
The first division of the initial cell results in two unlike segments: a
large lower cell which develops the walls of the conceptacle and cryp-
tostoma; and an upper cell, the tongue cell, which either remains
inactive, divides to form a short filament, or degenerates. The “‘initial
cell” of Bower is apparently the tongue cell, a product of the true
initial cell.
The conceptacle and cryptostoma are undoubtedly homologous
structures. Every stage of development in both structures is the
same, from the appearance of the similar initial cells to the develop-
ment of paraphyses in the cryptostomata and sexual organs in the
conceptacle.
The paraphyses are developed sag i by the division of the
lowermost cell in each structure.
Spermatocysts or their degenerate representatives occur in some
cryptostomata. Such conditions indicate that the cryptostomata have
been derived from conceptacles whose sexual organs have become
sterile.
The spermatocysts develop as in other Fucaceae, each finally pro-
ducing sixty-four sperms which are discharged from a partly terminal
and partly lateral rent.
The sister cell of an oocyst does not become a stalk and conse-
quently the oocyst is an embedded structure.
180 BOTANICAL GAZETTE [MARCH
The oocyst normally gives rise to but one egg. The nucleus of the
oocyst accordingly becomes the nucleus of the egg. |
The oocysts were found containing eight eggs each. These must
be considered a rare reversion to the Fucus type.
The entire oocyst of Sargassum, unlike other genera of the Fuca-
ceae which have been studied, is discharged with its enclosed egg.
The oocyst wall may break, partially freeing the egg, or it may persist
even enveloping a many-celled sporeling.
Segmentation of the egg takes place while it is attached to the sur-
face of the plant by the mucilaginous wall which surrounds it. This
segmentation results first in a many-celled undifferentiated ellipsoidal
sporeling. Rhizoids develop late at one end of the multicellular spore-
ling, with no apparent relation to gravity or other stimulus.
Asters, containing granular inclusions suggesting centrosomes,
appear at the poles of the spindles in the early mitoses of the segmenta-
tion of the egg.
THE UNIVERSITY OF CHICAGO.
LITERATURE CITED.
Barton, E. S., ’91, A systematic and structural account of the genus Turbinaria
Lamx. ‘Piers: Linn. Soc. Bot. 3: 215-226. pls. 54-55.
BEHRENS, J., ’86, Beitrag zur Kenntniss der eg ia Rarer bei Fucus
tices: lige Deutsch. Bot. Gesells. 4:92-10
Bower, F. O., ’80, On the development of the sciceplecks in the Fucaceae.
Quart. Jour. Micr. Sci. 20: 36-49. pl. 5.
Crato, E., ’92, Die Physode, ein Organ des Zellenliebes. Ber. Deutsch. Bot.
Gesells. 10:295-302. pl. 18.
Davis, B. M., :04, The relationships of sexual organs in plants. Bot. GAZETTE
GruBer, E., 96, idee Aufbau und Entwickelung einiger Fucaceen. Bibli-
otheca Bot. 38:3
HANSEN, A., ’95, Usher Stoffbildung bei den Meeresalgen. Mittheil. Zool. Sta.
Neapel rz: 255-305. pl. 12.
HANSTEEN, B., ’92, Studien zur Anatomie und Physiologie der Fucoideen.
Jahrb. Wiss. Bot. 24: 317-360. pls. 7-10.
,:00, Ueber das Fucosan als erstes scheinbares Product der Kohlensaure-
assimilation bei den Fucoideen. Jahrb. Wiss. Bot. 35:611-625 pl. 14.
Hotz, F. L., :03, Observations on Pelvetia. Minn. Bot. Studies 3:23-45-
pls. 7-12.
KJELLMAN, F., ’93, Engler and Prantl, Pfl. fam. I. 2: 268.
PLATE X
~
~
be
Sa
Ny
hy
Ry
~
S
=
1S)
~
q
&
2)
Q
1
2
Etoile B. Simons, del.
*
sIMONS # SARGASSUM
1906] SIMONS—SARGASSUM FILIPENDULA r81
Kocu, L., ’96, Untersuchungen iiber die bisher fiir Oel oder Phloroglucin
gehaltenen Inhaltskérper der Fucaceen. Inaug. Diss. Rostock.
Kuntze, O., ’81, Revision von ae und das sogenannte Sargasso-Meer.
Engler’s Bot. Jahrb. 1: 191-239. pls..1-2.
MitcHett, M. O., and Wuarrtrnc, F. G., ne On Splachnidium rugosum Grev.,
the type of a new order of algae. Phycological Memoirs Part I. pp. 1-10.
Learn F., ’89, Beitriige zur Kenntniss der Fucaceen. Cassel.
Meaghleige und Biologie der Algen. Jena
a4 L 76, Beitrage zur Kenntniss der Tange. Jahrb. Wiss. Bot. 10:317-
382. pls. 25-27.
VALIANTE, R., ’83, Le Cystoseirae del Golfo di Napoli. Fauna und Flora Golfes
Neapel '7: 1-30. pls. 15.
EXPLANATION OF PLATES X AND XI.
All figures were sketched with a camera and reduced one third in reproduc-
tion. Figs. r-30 were drawn with Zeiss apochromatic objective 1.5™™ and com-
pensating ocular number 4, magnification 1140. Figs. 31 and 32 were drawn with
dry objective, magnification 570.
PLATE X.
Figures 1-11. Development of the conceptactle.
Fic. 1. Initial cells, longitudinal section; 1a, cross section of apex; 1m, cross
section of median portion; 1b, cross section of basal portion.
Fic. 2. Two-celled stage, longitudinal section showing the slender upper
tongue cell, and a larger lower cell; 2a, cross section of the apex showing the
tongue cell only; 2m, cross section of the median portion with the centrally placed
basal region of the tongue cell surrounded by the upper part of the lower cell;
26, cross section of the basal portion showing the lower cell only. :
Fic. 3. Three-celled stage, longitudinal section; 3a, cross section of the
apex showing tongue cell only; 3m, cross section of median portion with the cen-
trally placed lower part of the tongue cell, surrounded by the upper part of its two
companion cells; 3b, cross section a little below 3m, showing the same cells;
3, cross section of the basal portion showing the two lower cells only.
Fics. 4 and 5. Longitudinal sections of a peculiar trinucleate stage of one
conceptacle. The first division of the nucleus of the initial cell must have been
with the axis of the spindle perpendicular to that of the cell. Fig. 5 contains one
of the nuclei of the first mitosis and fig. 4 the products of a division, now in late
telophase, of the other nucleus of the first mitosis.
Pb o = Oo
ag MW MW
walls Celi ~~
the tongue cell. -
tudinal section of a later stage with six similar wall cells and the
Fic. 7. Longitu
centrally placed tongue cell.
182 BOTANICAL GAZETTE [MARCH
Fic. 8. Longitudinal section of a more advanced stage illustrating the forma-
tion of the cavity of the conceptacle.
IG. 9. Pesca aes section of a young conceptacle some of whose wall cells
are ie papi he tongue cell contains two nuclei.
Fic. 10. A aaa of three cells formed from the tongue cell.
Fic. 11. Young conceptacle showing simultaneous development of wall cells
and papillae.
Figures 12-22. Development of the cryptostoma.
Fic. 12. Initials, longitudinal section; 12a, cross section of the apex; 12m,
cross section of median portion; 126, cross section of basal portion.
Fic. 13. Longitudinal section of the two-celled stage.
Fic. 14. Longitudinal section of the lateral surface of the three-celled stage.
Fic. 15. Longitudinal section of the interior of the same group of cells repre-
sented in fig. 14.
Fic. 16. Longitudinal section showing four wall cells and the tongue cell.
Fic. 17. Longitudinal section slightly more advanced.
Fic. 18. Longitudinal section of a young cryptostoma beginning to form
paraphyses very early.
Fic. 19. Longitudinal section of an older stage which has not yet begun to
develop og aa
Fic i Loaditadinal section showing five paraphyses developing from wall
cells sia one from the ton e
Fic. 21. More ‘vaiee illustrating the simultaneous development of
paraphyses and wall ce
Fic. 22. Still more iluaspach
PLATE XI.
Fig. 23. The development of papillae which will later give rise to spermato-
sts.
Fic. 24. At the left a cell which results from the separation of a papilla from
a wall cell. At the right a spermatocyst and stalk which have been formed by the
division of a cell similar to the one shown at the left.
Fic, 25. A stalk cell has given rise to a papilla, now separated by a wall.
Fic. 26. A branch system formed through the activity of stalk cells.
Ah Fic. 27. A spermatocyst containing sperms. The stalk cell has developed a
air.
Fic. 28. A mature spermatocyst, the stalk cell pushing out at one side.
Fic. 29. Very young oocyst with its sister cell, which is the homologue of the
stalk cell in Fucus
FIG. 30. Slightly older oocyst and its sister cell already unequal in size
Fic. 31. A mature embedded oocyst containing many hnshatophines and
much reserve material.
Fic. 32. A sporeling still attached to the surface of the parent plant. At one
pole rhizoids have begun to develop. The old wall of the oocyst surrounds the
sporeling.
CHROMOSOME REDUCTION IN THE MICROSPORO-
CYTES OF LILIUM TIGRINUM.?
Joun H. SCHAFFNER.
(WITH PLATES XII AND XIII)
THE progress recently made in our knowledge of hybrids has given
a new impetus to the study of chromosome reduction. Unfortunately,
there is still much disagreement in the accounts of various observers.
In order to continue my own investigations on a very favorable object,
the microsporocytes of Lilium tigrinum were selected, since material
is easily obtained in large quantities. CHAMBERLAIN has studied the
pollen grain of this plant and has also given figures of the microsporo-
cyte in the spirem stage. The chromatin granules are exceptionally
distinct, and this facilitates the correct interpretation of the complex
figures to be seen in the reduction karyokinesis.
Recently papers on the reduction division have been published by
FARMER and Moore, STRASBURGER, MONTGOMERY, WALLACE, and
others, which are in essential agreement with the interpretations of
Drxon on Lilium longiflorum and my own observations on Lilium
Philadelphicum and Erythronium. On the other hand BErcus,
REGOIRE, and ALLEN have come to somewhat different conclusions.
The observations of RosENBERG on Drosera have opened up an
important field of investigation on the individuality of the chromo-
some. These papers have been reviewed so recently by various
writers that it is needless to discuss the results here. It is sufficient
to say that it must appear to an impartial judge that the cytologist is
at present able to see in his preparations almost anything which may
be conceived of as taking place in the structures investigated. This,
however, should not hinder work in such an important field, for the
proper interpretation can be attained only by continued observation.
Little can be regarded as certain until there is a more general
agreement among competent investigators. So far as the present
research is concerned, the extent and variety of the preparations
‘Contributions from the Botanical Laboratory of Ohio State University. XXIV.
183] [Botanical Gazette, vol. 41
184 BOTANICAL GAZETTE [MARCH
studied seem to preclude the possibility of mistake. Such doubts
as were expressed in my former papers have been practically
removed. So far as the writer is concerned, the interpretation given
below is conclusive. Another investigator might perhaps come to
different conclusions by using other methods.
MATERIALS AND METHODS.
Stamens of various ages were collected in Clay County, Kansas,
during July 1904; killed in weak chrom-acetic acid (chromic acid 0.3%”,
glacial acetic acid 0.7°°, water 99°‘), passed gradually through the
grades and preserved in 70 per cent. alcohol, imbedded in paraffin,
cut 10-18 » thick, and stained on the slide. Some old slides, the mate-
rial of which had been killed in the ordinary chrom-acetic acid solu- _
tion, were also at hand. Various stains and combinations were used,
but for bringing out the chromatin network and chromatin granules
of the early stages, Delafield’s haematoxylin, when properly developed,
gave by far the best results, being superior in this respect to either
Heidenhain’s iron-alum-haematoxylin or safranin and gentian violet.
Nucleoli, both in the nuclear cavity and in the cytoplasm, are stained
very distinctly by the safranin-gentian-violet combination, but are
only slightly affected by Delafield’s haematoxylin.
_ I am indebted to my wife, MABEL ScHAFFNER, for the prepara-
tion of most of the two hundred serial slides on which the present
paper is based.
INVESTIGATION.
Before the microsporocytes are beginning to separate the promi-
nent chromatin network is being transformed into slender delicate
threads. These threads appear to be discontinuous in some places.
However, the appearance may be due to injury during the process of
cutting. The threads are small in diameter as compared with the
single chain of spherical chromatin granules (figs. 1-3). After the
spirem becomes fully developed it shows no free ends and is much
wound, looped, and twisted. In this stage it appears to be entirely
free in the nuclear cavity and is usually in the so-called synapsis stage.
Sometimes the contraction is to one side of the nuclear cavity, some-
times near the center, but often little or no contraction is evident.
Whether this is an artifact or a real stage in the process of karyokin-
=
Y?
—
1906} SCHAFFNER—LILIUM TIGRINUM 185
esis appears still to be an open question. Such contractions are so
easily produced by the ordinary killing reagents used, and have been
described for such a variety of the early stages of nuclear division,
that it seems to me no importance is to be attached to observations
which have been made on killed material. So far as any opinion is
to be expressed upon the appearances in Lilium tigrinum, I am still
convinced that the extraordinary distortions commonly figured owe
their origin to the action of the fluids used before imbedding in par-
affin. :
The spirem remains simple during the entire synapsis. The linin
thread becomes thickened and the chromatin granules are usually
more or less elongated (figs. 4, 5). At this stage the spirem has
already a strong tendency to be thrown into loops and coils (fig. 4).
After the microsporocytes have become partly separated and more
spherical, they were rarely observed to be in synapsis. The nuclear
cavity enlarges and the chromatin ribbon becomes thicker, with the
granules still more prominent (jigs. 6-8). This is an important point
to consider, for we have here a clear case of sporocytes, long past the
supposed synapsis stage, showing with remarkable clearness a simple
continuous spirem, with a single chain of chromatin granules.
Synapsis, therefore, can have nothing to do in this case with a sup-
posed longitudinal conjugation of two spirems or two networks of
chromatin before the spirem is formed. After the nuclei have passed
on to the stage represented in fig. 6, the stages are so easily followed
and the threads so prominent, that a longitudinal conjugation, if one
occurred, could not escape notice. Shortly after the stage shown in
jig. 6 the spirem begins to show double rows of elongated chromatin
granules, but the relative quantity of chromatin ribbon present in the
nuclear cavity is not much diminished (figs. 9, 10). If the amount
of spirem were diminished one-half by a longitudinal conjugation the
fact would certainly be noticeable. Often a part of the spirem appears
still single, or it will appear with a double row of granules and gradu-
ally change to a row apparently single (figs. 11-17). The appearance
would be the same whether the granules were dividing or conjugating.
The uniformity of the pairs of granules on the linin thread is remark-
able, and the pairs themselves suggest a division. If a conjugation
of the chromatin granules were established, it would certainly show
186 BOTANICAL GAZETTE : [MARCH
an almost inconceivable regularity. The granules would have to be
definite and fixed, of the same number in both the egg and sperm, and
not be subject to increase or diminution (jigs. 14-16). In the early
stages of the spirem there are, of course, numerous instances of threads
lying parallel, side by side, but these appearances are equally common
after the chromatin granules appear double (jigs. 15, 17).
The ribbon now begins to show an arrangement into definite loops
(fig. 10). It becomes much shorter and thicker (jigs. 18, 19) and finally
shows a definite twisting together into twelve loops, which have their
heads toward the nuclear wall (jig. 20). At this stage the chromatin
- granules can still be distinguished lying side by side (fig. 21). But at
this time the whole ribbon begins to undergo a change, so that it stains
of a uniform, dense color throughout, and before the loops separate
all evidence of chromatin granules is lost (figs. 22, 23). That the loops
shown in figs. 18-23 are the incipient chromosomes is self-evident.
By no manner of interpretation can such a conclusion be explained
away. By the time the twelve loops have separated, the nucleoli
have entirely disappeared from the nuclear cavity (fig. 24). The nucle-
oli break up into micronucleoli and are thrown out into the cytoplasm
(figs. 38, 51). The figures in which they do not appear were taken
from material stained in such a way that the nucleoli were not evident
or not very distinct.
The chromosomes are exceedingly interesting on account of the |
many fantastic shapes produced by the coiling of the ribbon. A series
of distinct shapes is given in jigs. 25-37. Occasionally the loop shows
evidence of its double nature (jig. 29), but usually it appears homo-
geneous throughout. Much time was spent in a study of these chro-
mosomes, and the variety of shape and coil could be extended indefi-
nitely. In all cases the chromosome is continuous, the outer end
of the loop always being closed. Occasionally the coiling takes place
in such a way as to form a double loop (figs. 26, 27, 35). The chro-
mosome is situated on the spindle with its head or closed end outward
(figs. 28-30). Sometimes it is very difficult to unravel the nature of
the coil, as in fig. 35. In other chromosomes it is an easy matter to
follow out the details of the loop, as in fig. 28.
The spindle threads are evidently attached some distance back of
the free limbs of the loop (figs. 25-28). The limbs are gradually
= Pee
Le
| Oa
. 1906] SCHAFFNER—LILIUM TIGRINUM 187
uncoiled and pulled apart. They are separated at the closed outer end.
A transverse splitting of the chromosome is thus accomplished. There
is no evidence that the two limbs of the chromosome are a male and
female chromosome joined end to end and twisted together until they
make a longitudinal pair. But theoretically such a proposition is
easily possible or even probable. Because the spindle threads are
attached some distance from the ends of the limbs, the daughter chro-
mosomes are developed as V- or U-shaped loops (jigs. 40, 43, 51).- In
favorably stained sections some evidence of chromatin granules may
be observed. These still show a distinct pairing in some cases (jig. 44),
but in others the arrangement is considerably disturbed (jig. 45).
The daughter chromosomes, as they appear in the daughter star, are
of various forms and are sometimes twisted (figs. 44-50).
The micronucleoli are gradually collected below the two daughter
skeins, and are finally all inclosed in the nuclear cavities of the daugh-
ter nuclei (figs. 52, 53). The daughter chromosomes do not appear
to form a very definite resting network, but are transformed into the
mother skein of the second division rather rapidly. Whether a con-
tinuous ribbon is formed was not ascertained. The loops are already
separate at an early stage, and it is possible that the daughter chro-
mosomes of the first division, after forming an imperfect reticulum,
break up directly into the twelve chromosomes of the second division
(fg. 54). This point, however, is doubtful. But the absence of a defi-
nite resting stage in sections having microsporocytes with loose daugh-
ter skeins in close proximity to loose mother skeins of the second
division, gives support to the above supposition. The micronucleoli
are again distributed in the cytoplasm before the mother-star stage
of the second division (figs. 54, 55). The karyokinetic figures of the
second division are easily distinguishable from those of the first.
This is especially true of the mother star (figs. 38, 55). The chro-
mosomes in the second division have their free ends directed out-
wards, as in an ordinary vegetative division. Commonly they are
more or less tangential to the spindle. Sometimes, however, they
Stand at right angles, as represented in jigs. 56, 57.
One of the most difficult points to determine was the nature of the
splitting in the second division. However, it was definitely ascer-
tained that the splitting is longitudinal. Dividing chromosomes are
188 BOTANICAL GAZETTE [MARCH
represented in jigs. 58-61. The daughter loops are completely sepa-
rated very early in their migration towards the poles (jig. 62).
They are also of various shapes. Frequently straight chromosomes
are present in the daughter star and very commonly the J shape pre-
dominates (figs. 63,64). Uand V shapes are also present. After the
chromosomes have passed into the daughter skein stage they show an
irregular outline with a row of irregular chromatin granules (jigs.
65, 66). The chromosomes develop into irregular networks, show-
ing the remains of the original loops, and thus pass into the spore
tetrad stage (jig. 67).
During the early germination stages of the microspores the figures
are again remarkable for the large nucleoli in the cytoplasm (jigs.
68-70). From the appearance of the nuclei during the several
divisions, it is evident that the nucleoli do not contribute directly to
the formation of the chromosomes, but that they are uniformly thrown _
out into the cytoplasm, in a fragmented condition, during the earlier
stages of karyokinesis.
No study of the achromatic structures was attempted, and though
some interesting points were observed from time to time, there was
nothing which calls for special mention.
SUMMARY.
1. The first division of the microsporocyte is a true reduction divi-
sion.
2. A continuous spirem is formed with a single row of chromatin
granules.
3. The spirem passes through and comes out of synapsis without
a conjugation or division of chromatin granules.
4. The chromatin granules divide but the linin thread does not
show a distinct separation.
5- The continuous spirem shortens and thickens and twists up
into twelve loops, which are the incepts of the twelve separate chro-
mosomes.
6. The chromosomes are arranged in the mother star with the
loop or head end turned outwards and the spindle threads are attached
near the ends of the free limbs or about half way between the free
ends and the head.
pe |
——
ieee Se cenilaiiiaiiiea is
1906] SCHAFFNER—LILIUM TIGRINUM 189
7. During the metakinesis stage the chromosomes uncoil and
separate by a transverse division at the middle.
8. The chromosomes of the second division appear to represent
the daughter chromosomes of the first division.
g. The division of the chromosomes in the second nuclear divi-
sion is longitudinal.
10. The nucleoli fragment and pass out into the cytoplasm during
the first and second divisions and also during the germination of the
microspore.
OHIO STATE UNIVERSITY,
Columbus.
LITERATURE.
ALLEN, C. E., Nuclear division in the pollen mother-cells of Liliwm canadense.
Annals of Botany 19: 189-258. 1905
Bercus, J., La formation des chromosomes hétérotypiques dans la sporogénése
végétale. I. La Cellule 21:171-189. 1904. II. Idem 21: 381-397. 1904.
III. Idem 22: 41-53. 1904. IV. Idem 22: 139-160. 1905.
CHAMBERLAIN, C. J., The pollen grain. Bor. GAZETTE 23: 423-430. 1897.
Dixon, H. H., The nuclei of Lilium longiflorum. Annals of Botany 9:663-
5- 1695.
FARMER, J. B. and Moore, J. E. S., New investigations into the reduction phe-
nomena of animals and plants. Proc. Roy. Soc. London 72: 104-108.
1903.
Grécorre, V., La réduction numérique des chromosomes et les cinéses de matu-
ration. La Cellule 21: 295-314. 1904.
Montcomery, T. H., Some observations and Oke upon maturation
plichomena of ooh cells. Biol. Bull. 6:137-158. 1
ROSENBERG, O., Ueber = Individualitat der {Rx minicess im Pflanzenreich.
Flora 93: aks 259.
, Ueber die ‘ecoheica aioe im Drosera. Med. Stockholms Hégs. Bot.
Tnst: (Reprint, 1904.)
SCHAFFNER, J. H., The division of the macrospore nucleus. Bot. GAZETTE
23: ay an iby.
, Acontribution to the life history and cytglogy of Erythronium. Bor.
Gazerre 31: 369-387. 1
STRASBURGER, E., Ueber Remi ciecline! Sitzb. Kénig. Preuss. Akad. Wiss.
18: 587-614. 1904.
Wattace, L. B., The spermatogenesis of the spider. Biol. Bull. 8: 169-184.
905. “> *
190 BOTANICAL GAZETTE [MARCH
EXPLANATION OF PLATES XIT AND XIII.
The plates are reduced five-eighths in reproduction. The figures represent-
ing entire cells and nuclei were studied with a Leitz no. 4 ocular and +g oil immer-
sion objective; the others with a Zeiss no. 12 ocular and a Leitz +g oil immersion
objective.
PLATE XII.
Fic. 1. Microsporocyte with chromatin network beginning to form the spirem.
Fic. 2. Microsporocyte with delicate threads showing a single row of promi-
nent chromatin granules.
' Fic. 3. Single thread of the same.
Fic. 4. Sporocytes beginning to separate. Nucleus with distinct arch
contracted spirem in the so-called synapsis stage.
Fic. 5. Single thread of the same showing a single row of chromatin granules.
1G. 6. Sporocyte some time after the synapsis stage, showing prominent
spirem with single row of chromatin granules. The free ends are cut.
Ics. 7, 8. Single threads of the same.
Fic. 9. Sporocyte with chromatin granules mostly double and flattened in
appearance.
Fic. 10. Somewhat later stage, showing the beginning of looping of spirem,
and division of chromatin granules. One nucleolus beyond the nuclear cavity.
Fics. 11-17. Single threads from the same stage, showing chromatin gran-
ules still single, some with granules partly double, and others with typical double
rows of granules.
Fic. 18. Microsporocyte with loops nearly developed.
Fic. 19. A single loop of the same stage.
Fic. 20. Chromatin loops some time before separation; somewhat cut.
Fic. 21. A single loop from the same stage.
Fic. 22. Section of microsporocyte, showing chromatin loops near the time of
separation.
Fic. 23. A single loop from the same nucleus.
Fic. 24. Microsporocyte, showing the twelve chromosomes.
Fics. 25-37. Individual chromosomes, showing various types of loops and
coils and their position on the spindle threads.
PLATE XImI.
Fic. 38. Mother star, showing position of chromosomes. Micronucleoli in
the cytoplasm.
Fics. 39-42. Chromosomes from mother-star stage, showing mode of sepa-
ration of limbs of loops.
Fic. 43. Microsporocyte with daughter stars.
Fics. 44-50. Daughter chromosomes from the daughter star, showing char-
acter of of the loops.
Fic. 51. Daughter-star stage, with micronucleoli in the cytoplasm.
BOTANICAL GAZETTE, XLI
SCHAFFNER on LILIUM
PLATE Xil
BOTANICAL GAZETTE, XLI
SCHAFFNER on LILIUM
PLATE XUil
—
1906] SCHAFFNER—LILIUM TIGRINUM IQI
Fic. 52. Loose daughter-skein stage with micronucleoli collected below the
open ends of the chromosomes.
Fic. 53. End of loose daughter-skein stage, with mnjcgemuriank collected
among the chromosomes.
1G. 54. Beginning of second division, with chromatin loops in the nucleus
and aaa ania in the cytoplasm.
. 55. Mother star of the second division, showing characteristic appear-
ance the chromosomes, with micronucleoli in the cytoplasm.
Fics. 56, 57. Single chromosomes from the mother-star stage, showing posi-
tion on the eres? threads.
IGS. . Chromosomes from the metakinesis stage, showing the nature
of the aiid splitting.
1G. 62. End of metakinesis stage.
Fic. 63. Daughter-star stage of second division.
Fic. 64. A pair of daughter chromosomes from the daughter-star stage of the
second division.
Fics. 65, 66. Pieces of chromosomes at the end of the second division, show-
ing a single row of irregular chromatin granules.
1G. 67. Tetrad at end of second division.
Fic. 68. Microspore at beginning of germination, with two large nucleoli in
the cytoplasm.
Fic. 69. Same stage as the preceding, showing three nucleoli in the cytoplasm.
Fic. 70. Microspore in germination stage, showing a number of nucleoli in
the cytoplasm.
CYTOLOGICAL STUDIES ON THE ENTOMOPHTHOREAE.
I. THE MORPHOLOGY AND DEVELOPMENT OF EMPUSA.'"
EDGAR W. OLIVE.
(WITH PLATES XIV AND XV)
THE general development of the Entomophthoreae and the exter-
nal morphology of its various members have been studied by a number
of investigators. For a detailed review of the literature pertaining to
the group, the reader is referred particularly to the papers of BREFELD
(’71,’81,’84) and of THAXTER (’88).
CoHNn’s (’55) results in his classical paper on the developmental
history of Empusa muscae have been in certain respects considerably
modified by later investigation. According to him, the fly first be-
came diseased and the fungus followed as a consequence. The first
indications of the disease which CoxN could find in the blood of
the fly were numberless minute globular or irregularly shaped bodies,
whose presence he could not explain otherwise than by the assump-
tion of spontaneous generation (p. 334).
These bodies, according to him, grow larger, become globular
or ellipsoidal, and finally grow into the filament, which, by the
formation of partitions, becomes the three-celled hypha characteristic
of the mature fungus and consisting of spore, stalk-cell, and root-cell.
This three-celled character of the hyphae of Empusa muscae was
disproved, however, the next year by LeBert (’56) and later by
BREFELD and others.
Every investigator who has attempted to infect insects artificially
has testified to the difficulties which he has encountered. COHN
(’55, P- 342), in speaking of his own lack of success, wisely empha-
sizes the caution which should characterize such experiments, noting
that one should be certain that the insects experimented with are not
already stricken with the disease, a more difficult task than would
at first appear.
I am under obligation to the Carnegie Institution of Washington for grants,
which have rendered possible this investigation.
Botanical Gazette, vol. 41] [192
~ -
=
1906] OLIVE—DEVELOPMENT OF EMPUSA 193
BREFELD (’70, "71, 77), however, was successful in transmitting
the parasite through external inoculation of spores, and he found
that in the case of Empusa muscae infection took place only through
the thinner whitish parts of the skin on the under side of the fly’s
body; whereas in another species, E. sphaerosperma, the infected
hyphae gained an entrance at any part of the skin of the cabbage
larva. This author has contributed more on this point than any
other, in that he was able to germinate the conidia in artificial
media as well as on the surface of the insect body, and to find with
the microscope the germinating hyphae actually boring through, the
skin of the host. In his series of articles, BREFELD has described
the complete course of development of two species of Empusa, E.
muscae Cohn and E. sphaerosperma Fres., which furnish quite differ-
ent types of vegetative growth. According to him, in E. muscae there
are formed from those germ-tubes which have penetrated into the
body-cavity of the insect numerous detached non-nucleate cells,
which reproduce by repeated yeast-like sprouting, and which grow
within the fat-bodies of the host. At a certain advanced stage of the
development of the fungus, the reproduction of the cells by budding
ceases, and each grows at one or both ends into a long unbranched
tube, which grows through the body-wall and produces at its external
end a single conidium. In the other species, E. sphaerosperma,
BREFELD found that the germ-tube produces, on the other hand, a
copiously branching mycelium with many cross-partitions, which
finally fills the body-cavity of the host. At the end of the vegetative
period, this mycelium sends out hyphae which grow to the surface,
branch digitately, and finally produce acrogenously at each ultimate
end a single conidium.
Em pusa sphaeros perma further differs from E. muscae in producing
resting spores. COHN (’55, p. 343) had already suggested, since he
could not make the conidiospores of E. muscae germinate, that per-
haps the conidia themselves required a year of rest. But BREFELD
(°70, ’71), proved conclusively that the spores of this form were short-
lived, living only for a few days; hence his first suggestion in regard to
the puzzling question as to the wintering of such a species was that
this form was probably heteroecious, and that resting spores were pro-
duced in some other host. Later, however (’84, p. 68), he seems
194 BOTANICAL GAZETTE [MARCH
inclined to believe that the disease is continued over winter on flies
in warmer regions, and that it migrates northward with the insects on
the return of summer; the fallacy of which theory THAXTER has
pointed out.
THAXTER (’88) in his account of the Entomophthoreae of the
United States, gives the results of morphological studies based on a
considerable number of new as well as old forms. This author dis-
agrees with BREFELD in regard to certain important points. In the
first place, he maintains that the vegetative growth in E. sphaero-
sperma is not filamentous in all cases, as is stated by BREFELD, and he
appears to be inclined to think that both the filamentous mycelium
and the broken-up, budding segments may occur in the same form
under different conditions. He asserts that the usual multiplication
of the hyphae is not by branching and continuous growth but by the
formation of ‘‘ hyphal bodies,” which “ consist of short thick fragments,
of very varied size and shape, that are continually reproduced by
budding or division, until the insect is more or less completely filled
with them.’’ But he continues further: ‘‘In cases where a direct
mycelial growth follows the entrance of the hypha of germination,
if indeed such instances occur, this mycelium must fall to pieces
into hyphal bodies, before the commencement of growth the direct
object of which is reproduction, in a fashion resembling that above
described at a similar stage for Conidiobolus’’ (p. 140). This con-
ception of reproduction by means of ‘‘hyphal bodies, ’’ however,
for reasons that are stated later in this paper, must be abandoned,
at least as a generalization.
The segments of the vegetative hyphae, or “ hyphal bodies” as
THAXTER terms them, under unfavorable conditions may each form
a thick-walled resting chlamydospore; or, when the conditions are
favorable, they may at once proceed to develop into the fructifying
state. In the latter case, according to this author, each hyphal seg-
ment sends out one or two (in some species more) hyphae which
develop into conidiophores. In the simplest case, a simple conidio- _
phore grows directly to the outer air and produces a single conid-
ium (Empusa muscae, e. g.). Or, the conidiophore may become
compound and produce a set of conidia. Or, under very favorable
conditions, ‘‘a single primary hypha may branch indefinitely, each
.
silane er
1906] OLIVE—DEVELOPMENT OF EMPUSA 195
ultimate branch becoming a conidiophore similar to those of the
more simple case just mentioned” (p. 142). A singular method of
germination of the “hyphal bodies” occurs in E. aphidis and E.
virescens, according to THAXTER’s observations. Spherical bodies,
evidently regarded as “hyphal bodies” with highly refractive con-
tents, germinate and send out a mass of hyphae in all directions. In
this condition they are said to resemble a head of Aspergillus,
although the author does not show in either of his two drawings of the
phenomenon any trace or remnant of the central cells or “hyphal
bodies” from which the radiating hyphae are said to arise.
The conidium is regarded by THAXTER as having a double wall,
and thus is to be interpreted more properly as a simple single-
spored sporangium.
MATERIALS AND METHODS.
In March 1904, the writer found in horse-dung cultures in the
laboratory a small species of fly belonging to the genus Sciara, which
was infected with an Empusa. This small fly with its attendant
disease has been propagated in horse-dung cultures since that time,
and many successive generations of the insect, the larval condition as
well as the adult, during the year and more of its cultivation, have fur-
nished a wealth of material for an almost complete cytological and
developmental study of this species of Empusa. A number of other
forms of the Entomophthoreae, most of them in the fructifying stage
of their existence, have been used for comparison, but no others have
as yet been traced through their entire life history. Enough has been
learned, however, to show the existence of a most interesting series
of distinctive variations.
My material has been killed with a variety of fixing agents, mostly
with varying strengths of Flemming’s chromic-acetic-osmic acid mix-
ture. The insect body was generally cut in two or pricked to allow
direct contact of the fixing fluid and the fungus hyphae in the body
cavity. The material was sectioned usually 3-6 » thick and stained
with Flemming’s safranin gentian-violet orange-G, or with Heiden-
hain’s iron haematoxylin.
Six species of Empusa altogether have been thus studied. These
species, determined Zaccording to the descriptions in THAXTER’S
196 BOTANICAL GAZETTE [MARCH
account of the group, are as follows: Empusa muscae Cohn, on the
common house fly; E. culicis, A. Braun, on a small green species of
Chironomus; Empusa sp., on a large fly; EZ. aphidis Hoffman, on vari-
ous aphides; E. americana Thaxter, on a blue-bottle fly; and one other
species, which is the principal one studied in this paper, on the small
fly, Sciara sp. The ovoid conidia of this last form coincide closely
with the description of the conidia of EF. montana Thaxter. They
show decided differences, however, from THAXTER’s drawings of this
species; and from £. ovispora Nowakowski and E. echinospora Thax-
ter, with which the measurements of the conidia also almost coincide,
the form is distinguished by the characteristic zygospores of the
latter species. While it is possible that, under certain conditions
unknown to me, this species on Sciara may produce zygospores, yet
the fact remains that after over a year of continuous observation, I have
failed to find any resting spores. It is therefore thought advisable to
give a new name provisionally to this species, which will hereafter
be referred to as Empusa sciarae.
Empusa sciarae n. sp.—Vegetative hyphae forming a branching,
septate mycelium, which in advanced stages is cut up into few- (gener-
ally 3-5-) nucleate cells. Radial hyphae branched; conidiophores 3-5,
bearing at each ultimate end a single ovoid uninucleate conidium with
a rounded basal papilla, 12-16 X 18-25 w. Zygospores unknown.
VEGETATIVE STAGE.
My own observations of purely vegetative stages are concerned
with two forms only, Empusa aphidis and E. sciarae, both of which
agree with the type described by BREFELD for E. sphaerosperma. In
all the other species studied, the vegetative condition had ceased
and conidiophores had grown out from the vegetative hyphae. As
observed by BREFELD, THAXTER, and others, the insect dies at the end
of the nutritive period of the fungus; but they do not seem to have
emphasized the fact that living insects alone must furnish data for the
study of the vegetative hyphae. After the initiation of the repro-
ductive period and the consequent death of the insect, the radial
growth of the conidiophores produces a mass of hyphae which might
readily be taken, in certain instances, for a vegetative filamentous
mycelial growth. The probability has suggested itself to the writer
1906] OLIVE—DEVELOPMENT OF EMPUSA 197
that the fat cells, detached in the search for the fungus, must have
been mistaken by some for stages in the development of Empusa; since
these bodies may frequently resemble closely short hyphal segments in
their fatty, granular contents, as well as in their assumption of globular
or irregular shapes, which give the suggestion of budding cells.
These detached fat-bodies, which prove in sections to be small aggre-
gates of insect cells, are particularly abundant in the adult just pre-
ceding ovipositing. An easy and certain method of distinguishing
the vegetative cells of Empusa from fat-bodies which oceur with them
in the body cavity, is to stain with a dilute methyl-green solution,
acidulated with a few drops of acetic acid, when the fungus cells
stand out conspicuously, distinguished by their relatively large and
characteristic nuclei.
In some preliminary observations on the manner of infection of
Empusa sciarae, I have not been able to make certain of this point,
but I wish to record here some notes of interest pertaining to it. The
small fly, Sciara sp., was sometimes accompanied, in the vessels in
which the dung cultures were kept, by three other species of small
flies, which hatched generally in less abundance, Psychoda sp., a so-
called moth fly, and two other undetermined forms. None, however,
other than Sciara were infected, although many times I have noted
conidia stuck on the surface of the bodies of the larvae as well as of the
adults of the other species. Successive generations of infected Sciara
larvae as well as adults have appeared with great regularity every
month or six weeks. The fly lays its eggs on the surface of the dung
or on the sides of the vessels, and the young larvae, soon after hatch-
ing, crawl below the surface of the substratum. It would appear
reasonable to assume that in this case infection would occur with ease
in this very young condition, when the larval skin is thin and deli-
cate, and before they had crawled below the surface cf the dung,
where they would obviously not be reached by shooting conidia.
My preliminary unsuccessful experiments at infecting healthy adult
larvae, confined for a week in bottles with diseased ones, at least
suggest the possibility of infection occurring in young insects only.
If the cultures in which young larvae were being nourished were
kept quite moist, perhaps the majority of individuals in this stage
were killed by the disease. When, however, the conditions were
198 BOTANICAL GAZETTE {MARCH
drier, the insect developed frequently into the adult fly, which itself,
after ovipositing, generally died of the disease.
In order to determine whether the disease could be carried from
larval to adult condition, I have examined many pupae, but only in one
or two instances have hyphae been found. In one case in which a
young insect was struggling to get out of its pupal shell, a few Empusa
hyphae were noted when the body was cut open, together with many
small eggs and globular fat-bodies. When cultures were kept at
some distance from infected ones, in an adjoining room, Sciara fre-
quently developed to maturity by hundreds and died finally a natural
death, evidently without infection.
Resting spores in this species, as in E. muscae, have not been
observed ; hence the puzzling question confronts us here also as to the
method of wintering of the fungus. An interesting suggestion is made
in this connection by the spontaneous appearance of Empusa sciarae
in laboratory cultures as early in the spring as March, in 1904. This
fact apparently still further renders useless BREFELD’s hypothesis as
to the migration of the fungus north from warmer countries. The
host, in this case, must have been breeding in the dung of the warm
stables all through the winter, and it is quite reasonable to suppose
that the short-lived fungus must have been continued at the same time
on successive generations of insects. The successful cultivation of the
form for more than a year in the laboratory gives additional reason for
this belief. It is possible that Empusa muscae and such similar forms
may have lost their sexual stage, because of their success in propaga-
ting the disease by means of conidia alone. It is well known that a
few house-flies survive the winter by hibernation or otherwise, and it is
probable that some of the individuals during the winter may continue
breeding in stables or in other favorable places, and in this way carry
over the disease, even in cold climates.
Living larvae of Sciara furnish beautiful material for a study of the
vegetative conditions of this type of Empusa. When placed on a
slide in water, a glance with the naked eye is sufficient to determine
whether the larva is infected or not, since the diseased individuals are
whitish in appearance, due to the presence of the more or less copious
mycelium, while uninfected ones are transparent. Under the micro-
scope, the diseased Jarvae show clearly the long, branched, mycelial
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1906] OLIVE—DEVELOPMENT OF EMPUSA 199
filaments, which in a young condition may have but few septa, while
in a later vegetative stage they develop numerous septa. As the
larvae crawl about over the surface of the slide, the hyphae can be
seen sometimes extending the full length of the body-cavity, some-
times copiously developed in the posterior portion only, again more
abundant in the anterior region. As the body parts move, or the
blood flows in the cavity, the mycelium shows. corresponding pul-
sations and movements. Fig. z showsa ‘portion of a branched
vegetative hypha, taken from such a larva which was crawling just
below the surface of the dung culture. Evidently the vegetative activ-
ities have here almost ceased, and the 2-, 3-, or 4-nucleate cells are
about ready to send out the radial conidiophores. Fig. 2 shows a
section of a younger hypha, in which the cells contain a varying
number of nuclei; while fig. 3 represents a still younger stage, the
- earliest condition within the body of the host which I have succeeded
in obtaining, growing in a larva in which the mycelium was composed
of but a few scattered filaments. The section of the hypha, of which
jig. 3 represents only a portion, shows in the preparation twenty-two
nuclei, but in the entire length, as far as traceable, not a single cross
partition.
While in my own investigations of the vegetative stages of Empusa
sciarae there remains as yet a small gap, from the penetration of the
infecting hypha to the production of such multinucleate mycelial fila-
ments as are shown in fig. 3, I feel reasonably certain as to the method
of procedure. Germinating conidia, growing in a sterilized decoc-
tion of cooked larvae, are shown in figs. 4-9. Figs. 4-7 illustrate
the germination of the uninucleate conidia into germ-tubes which have
grown out of the liquid decoction, thus resulting in the formation at
once of secondary conidia; jig. 7 shows the beginning of the forma-
tion of a tertiary conidium. In figs. 8 and g the germ-tube has grown
into a hypha in which all of the protoplasm appears to be in the
end cells; in the latter, sixteen apparently empty cells separate the
old conidial wall from the terminal protoplasm, which still remains
uninucleate.
‘Both BREFELD (’71, figs. 5, 29, 31) and THAXTER (’88, fig. 240)
give figures showing a somewhat further. advance over what I have
obtained in the cultivation of conidia, in that in their forms branches
200 BOTANICAL GAZETTE [MARCH
are beginning to appear. Whether an increase of the amount of
protoplasm and of the number of nuclei has accompanied this
branching is doubtful.
It is highly probable, therefore, that in Em pusa sciarae the hypha
which has penetrated into the body-cavity of the fly from the germi-
nating conidium grows rapidly at the expense of the nutrient fluids
in which it floats. After the protoplasm has increased in volume,
the nuclei increase in number by division, and from the uninucleate
condition, in the case of Empusa sctarae, the hypha finally becomes
a multinucleate branching filament, such as is shown in fig. 3.
Partition-walls in this form at first occur but sparingly; later, how-
ever, at the culmination of vegetative activity, septa are abundant and
branching becomes more frequent. Finally the body-cavity of the
insect becomes completely filled with the mycelial filaments, vegeta-
tive activity ceases, and the fructifying state begins.
Empusa aphidis furnishes a somewhat modified vegetative devel-
opment. The advanced condition, which is my only source of infor-
mation in this case, shows in sections branched coenocytic hyphae,
which appear to be but rarely divided by septa, unlike the corre-
sponding stage in Empusa sciarae. Even after the rhizoids have
grown out from the under side of the body of the insect (figs. 10, 11),
the vegetative activities appear to continue, as evidenced by the fact
that the nuclei in this instance are still undergoing division. Also
in Empusa sp. there occurs a similar prolongation of vegetative
activity, since at an advanced stage even fewer cross-partitions can
be found in the coenocytic mycelium (jigs. 23, 25).
REPRODUCTIVE STAGE.
At the culmination of vegetative growth, the body-cavity of the
larva appears to be completely filled with a mass of long hyphae.
Toward the close of this stage, the larva crawls to the surface of the
substratum or high up on the side of-the culture dish; or, in the case
of the adult Sciara, the fly seeks a conspicuous position, as is common
with such infected insects, and death ensues with the beginning of the
fructifying condition.
The initiation of the fructifying condition is marked in Empusa
sctarae by the sudden formation of radial branches from the short
ial
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1906) OLIVE-—DEVELOPMENT OF EMPUSA 201
cells which make up the hyphal filaments. These branches appear
to be put forth more or less simultaneously, in this species but one from
each cell. The cell swells up and becomes rounded off somewhat at
its ends (fig. 13). One or more vacuoles appear in the protoplasm
and a protuberance is pushed out from one end of the cell; this grows
into the radial hypha destined to become the branched conidiophore
(figs. 12, 14,15). The two forces, the swelling and consequent round-
ing off of the cells at the ends and the pushing out of the branch, com-
bine to split the partition-walls between the cells, thus causing the
hyphae to become easily broken up into one-celled segments, the
“hyphal bodies” of THAXTER (figs. 1, 12). I have seen similar
hyphal segments, forming in these instances also the origin of the
radial conidiophores, as well in my preparations of E. culicis and FE.
americana; but other species promise interesting variations from this
common type of pre-reproductive development. In Empusa sp.
and E. aphidis, for example, the vegetative hyphae remain, up to the
very initiation of the reproductive stage, either unicellular, or at least
with cross-partitions at only rare intervals. In such instances, there-
fore, no breaking up into “hyphal bodies” occurs; but the vegetative
hyphae appear to grow out directly into the conidiophores.
As BREFELD and THAXTER have pointed out, the first hyphae to
appear in the external growth of the fungus form the rhizoids, by
means of which the host is fastened firmly to the substratum. In the
common house-fly, the host is attached by means of its proboscis. . In
E. sphaeros perma, according to BREFELD, bundles of rhizoids break
out more or less irregularly from the under side of the body of the
insect and attach it to the substratum. According to THAXTER,
these rhizoidal hyphae may branch, and may terminate in a kind of
expanded sucker, which apparently secretes a viscous substance. .
In Empusa sciarae, rhizoids are developed more or less abund-
antly from the under side of the abdomen of the fly, or, in the case of
the larva, from almost any point on the under side of the body. In
certain forms of E. aphidis, groups of rhizoids break out from the
under surface of the insect and form large sucker-like hold-fasts. In
several instances, I have counted three of these hold-fasts from aphides
parasitic on Solidago. Fig. 10 shows one of these sucker-like bundles
of rhizoids in section; and fig. rz a highly magnified hypha from near
202 BOTANICAL GAZETTE [MARCH
the inner region of the hold-fast. The latter presents clearly the char-
acter of the typical rhizoid; thick, frequently yellowish walls, contain-
ing but a thin layer of protoplasm, which bounds externally the
large vacuolar cavity that almost fills the hypha. The walls of the
ordinary vegetative hyphae, on the other hand, are thin; those of the
rhizoids in this case appear to have undergone a gelatinous or slimy
modification, and the contents seems to be undergoing degeneration.
The growth of the conidiophores, in the case of E. sciarae, proceeds
more slowly than that of the rhizoidal hyphae, a phenomenon which is
probably due, in part at least, to the slower absorption of water by
those hyphae destined to bear conidia. At any rate, the vacuoles
which are formed at this time in the cells increase slowly in size, and a
conidiophore arises from near the end of each cell and grows out radi-
ally, boring its way through the tissues of the host (figs. 12, 14, 15)-
As THAXTER and others have noted, the conidiophores of certain
species remain simple and unbranched, as in the case of E. muscae
(fig. 40); or, in other species, they may become normally septate and
branched, as shown in E. sciarae (fig. 16), and Empusa sp. (fig. 23)-°
From this fact arise most interesting cytological variations in the vari-
ous conidia of these species. THAXTER has brought out clearly the
variation in size, shape, etc., of the conidia of many species, but
CAVARA (’9Q9) was the first to contrast the multinucleate condition
of the conidia of Empusa muscae with that of the uninucleate conidia
of Entomophthora Delpiniana. Of the six species studied by me,
four have uninucleate conidia (E. sciarae, fig. 27; E. americana,
figs. 36, 37; E. aphidis, figs. 42, 43; and Empusa sp., figs. 22, 26):
The conidia of E. culicis are normally two-, rarely three-nucleate
(figs. 31, 32); while those of E. muscae have a more or less
indefinite number, frequently about 15-18 (figs. 38, 39).2. The more
common uninucleate conidia arise primarily from the septation of the
conidiophores into uninucleate segments; whereas, on the other hand,
the simple conidiophore of E. muscae does not usually become septate
except at the conidium. In the last case, therefore, all of the many
nuclei of the last-formed vegetative cell, which forms the origin of the
2 While convinced of the value of CAVARA’s suggestion as to the use of nuclear
characters in the classification of the Entomophthoreae I do not think that this one
character alone would justify the separation of Entomophthora from Empusa.
ee rr we
1906] ' OLIVE—DEVELOPMENT OF EMPUSA 203
conidiophore, flow out into the single, large, bell-shaped conidium
(fg. 39).
In E. culicis, which has binucleate conidia, septation of the conidi-
ophore occurs, by which the protoplasm cuts itself off from behind;
but, unlike the case of E. sciarae, I have found no branching with
it, so that as a consequence, all of the binucleate protoplasm of the
conidiophoric hypha flows into the one terminal conidium (figs. 30-35).
The simple conidiophore of E. culicis resembles, therefore, that of EF.
muscae in being unbranched, but the origin of the conidiophores may
differ in the two cases. One may find, in fact, in E. culicis, two sorts
of “hyphal bodies;”’ either small cells which, like the corresponding
ones of E. muscae, give rise to but one conidiophore; or, on the other
hand, larger cells, which may give rise to several conidiophores by
budding from several points simultaneously, in a similar manner to
Conidiobolus. In both instances, each conidiophore remains simple
and ultimately bears, terminally, the binucleate conidiospore.
In the species with uninucleate conidia, E. sciarae, E. americana,
E. aphidis, and Empusa sp., the coenocytic conidiophore, as was
indicated above, is cut up by septa, in a manner to be described later,
into uninucleate segments (figs. 16-21). In all these cases, this results
in a branching growth and successive abjunction of the acrogenous
spores. Below the terminal cell the penultimate cell pushes out to
one side, and thence bores its way to the surface of the host, where
it abjoints a single uninucleate conidium (figs. 16, 23). From this
habit in certain forms, arises a profuse system of branching, frequently
of a digitate type (e. g., Empusa sp.) or corymbose (E. sciarae and
others), to enable the uninucleate segments to reach the surface and
to discharge their protoplasm by means of the abjointed conidia.
But the last-formed vegetative cells of E. sciarae contain only 2-4
nuclei (figs. 1, 12, 15), so that, in this instance, only a correspondingly
small number of branches are formed.
_ The process of abjection of the conidia of Empusa is apparently
similar in a general way to that described for the sporangia of Pilob-
olus, except that in these conidia there is no gelatinous collar visible.
The formation of the partition at the base of the conidium in Empusa
also is quite a different process from the formation of the columella
in the case of Pilobolus.
In Empusa, the vacuole which appears in the basal portion of the
204 BOTANICAL GAZETTE (MARCH
‘cell becomes larger and larger (fig. 27), and a small protuberance,
which has a diameter equal to about half that of the conidiophore, is
pushed out from the end (figs. 34, 41). There is now formed at the
apex of this narrowed sterigma a swelling (jigs. 27, 35, 39, 42), which,
after continued enlargement, finally receives the greater part of the
protoplasm and all of the nuclear content from the basal portion. The
process of cell-division, by means of which this apical conidium is cut
off by a wall from the penultimate cell, will be described in some detail
later, in connection with the description of cell-division in Empusa.
Certain points should be noted here, however, and among them, that
the term basidium, as applied to the penultimate cell, although in
common use in this connection, should, in my opinion, be confined to
the Basidiomycetes, where, morphologically, the true basidium is a
very different spore-bearing structure from the penultimate cell bear-
ing the conidium in Empusa.
The penultimate cell forms the explosive mechanism by which
the conidium is shot off. As the ring-formed wall which cuts through
the base of the conidium travels progressively inward, the protoplasm
passes through the narrowing opening leading from below, until at the
close of abjunction, the basal cell retains only a thin parietal layer of
protoplasm, but no nucleus. Continued swelling, due to the absorption
of water, finally results in the bursting of the basal vesicle, thereby
breaking the wall of the vesicle where it joins that of the conidium.
In some forms, a ring-shaped scar is noticeable near the base of the
conidium, marking the circle where the summit of the swollen basal
vesicle was formerly attached. The septum which separates the
conidium from the subterminal cell is at first usually pushed upward,
thus resembling a columella (figs. 28, 30, 31, 36), but when the
spore is shot off, this partition-wall reverses its former position, and
in the conidium it appears as a prominent papilla (jigs. 29, 37, 43):
When the basal vesicle bursts, its contents are thrown out of the open
ruptured end and frequently persists as a slimy covering about the
spore, serving in this case, perhaps, the double purpose of protection
against excessive evaporation and of sticking the spore to the sub-
stratum which it strikes. I have noted that the explosion, in the case
of Empusa sciarae, sometimes throws the spore a distance of 67";
while BREFELD has recorded an even greater distance in the case of E.
muscae, in which the spores are said to be sent as far as 2-3°™.
. a
—T
1906] ' OLIVE—DEVELOPMENT OF EMPUSA 205
I have not yet completely solved satisfactorily to myself the pecul-
iar method of abjection of the spores of my undetermined species of
Empusa. I am inclined to think, however, that this method, in certain
respects, is unlike that described above. In this form the conidio-
phores come to the surface, become cut up by septa into uninucleate
segments, and proceed to branch profusely (jigs. 23, 25). The cell
terminating each branch pushes out in a peculiar manner. Instead
oi forming a large basal vesicle as an explosive mechanism in the usual
manner, as described above, here the protoplasm appears to cut itself
off from behind by means of one or more successively formed walls
(figs. 22, 24, 26). A minimum of protoplasm seems to be lost in
the process, and this cut-off protoplasm soon assumes a peculiar
granular appearance. In this condition it is probably dead, for these
cut-off cells appear soon to lose their turgescence. _ It is difficult to con-
ceive of a forcible discharge of the spore in this instance, especially if
it be true that the protoplasm of the basal cells is dead and thus inca-
pable, through loss of turgidity, of functioning as an explosive mechan-
ism. The process here rather seems to be that, by means of these
successive abjunctions, the uninucleate spores are pushed off with but
little force, and that they are probably followed out of the thick, gelatin-
ous wall of the mother hypha by other cells pushed up from below. It
may be, however, that further studies on fresh material of this species
will change this impression of subterminal proliferation.
Figs. 45 and 46 represent the terminal portion of large hyphae of
E. culicis which are destined to form resting spores; and figs. 47 and
48, two fully formed, thick-walled resting spores. Such hyphae as
are shown in the first two figures are distinguished from conidiophores
by being much larger, and, further, they contain four or five nuclei,
instead of two. I have traced these hyphae in sections back to
large “hyphal bodies,” but I was unable to follow their complete
history. Whether the thick-walled resting spores of this species are
therefore true zygospores, or azygospores, as they are termed in THAX-
TER’s monograph, I am not prepared to say. I have not been able,
however, to confirm VUILLEMIN’s assertions (:00) as to the nuclear
fusions in the azygospores of Entomophthora gloeos pora.
UNIVERSITY OF WISCONSIN,
Ison.
206 BOTANICAL GAZETTE [MARCH
LITERATURE CITED.
BREFELD, O., "70, Entwicklungsgeschichte der Empusa muscae und E. radicans.
Bot. Zeit. 28; 161-166, 177-186
, 71, Untersuchungen iiber die Entwicklung der Empusa muscae und E.
radicans. Abhandl. naturf. Gesells: Halle 12: rf. pls. 4.
» > pas die Entomophthoreen und ihre Verwandten. Bot. Zeit. 35:
345-355, 368-
—— a En hes radicans. Untersuch. tiber Schimmelpilze 4:97-
Tit.
-7-
oe Conidiobolus ae und minor. Untersuch. tiber Schimmel-
niin 6:35-72. pls. 3
Cavara, F., ’99, Osservazioni a sulle certo thchone: Nuovo Giorn.
Bot. Ital. N. S. 6: 411-466. pls.
Coun, F., ’55, Empusa muscae in ae Krankheit der eset Nova
Acta head: Caes. Leop. Carol. Germ. Nat. Cur. 25:3
Lesert, S., ’56, Die Pilzkrankheit der Fivesen. Abhandl. Naturf. Gesells.
Zuri
THAXTER, R., ’88, The Entomophthoreae of the United States. Mem. Boston
. Nat. Hist. 4:133-201. pls. 14-21.
VurtLemtn, P., :00, Développement des azygospores d’Entomophthora. Compt.
Rend. Acad. Sci. Paris 130: 522-524.
EXPLANATION OF PLATES XIV AND XV.
The drawings were made with the aid of an Abbé camera-lucida, and for the
most part with various compensating oculars combined with Zeiss 2™™ apochro-
matic obj. N. A. 1.30.
PLATE XIV.
1Gs. 1-9, Empusa sciarae.
Fic. 1. A freshly killed pie fixed and stained with acetic methyl green.
% 275.
Fic. 2. Section of a younger filament.
Fic. 3. Section of a still younger vegetative hypha. 275.
Fic. 4. A germinating conidium, cultivated in a decoction of cooked larve,
oe a Ain conidium. Killed in acetic methyl green. X575.
IGS. 5, 6. Conidia with still younger germ-tubes. Killed with acetic methyl
ok *575-
Fic. 7. A germ-tube has formed a secondary conidium, which in turn has
started to form a tertiary conidium. Killed ditto. 575.
Fics. 8, 9. The conidia have produced long germ-tubes instead of secondary
conidia. From the same culture as those above. Killed ditto
Fics. to-11, Empusa aphidis
Fic. to. A cross-section of one of the sucker-like rhizoidal disks.
Fic. 11. A section of a single rhizoid. x 1080.
Ne
BOTANICAL GAZETTE, XLI ae ; PLATE XIV
pene
ae ten
; me
é
E. W. Olive, del.
PLATE XV
OLIVE on EMPUSA
.
we
del.
E. W. Olive,
ni qo ne a sees
BOTANICAL GAZETTE, XLI
1906] OLIVE—DEVELOPMENT OF EMPUSA 207
Fics. 12-21, Empusa sciarae.
Fic. 12. A section showing two cells of the old vegetative mycelium, or “hyphal
bodies,” from the upper one of which has grown out radially a conidiophore..
Fic. 13. A hyphal segment freshly killed with acetic methyl green. X 480.
Fic. 14. The same, showing the beginning of a conidiophore. X 480.
Fic. 15. The same, showing a still longer conidiophore. 4
Fic. 16. Section of a radial conidiophore which is just ees > deca the epi-
dermis of the host, showing the subterminal branching. x1
Fic. 17. Section of a conidiophore showing a palsy ERS between the
two cells. 1080.
IG. 18. A half-completed stage of cell-division in a conidiophore. X 1080.
Fic. 19. Cell-division in the vegetative mycelium. 00.
IG. 20. An almost-completed stage of cell-division in a conidiophore; the iron-
haematoxylin stain has been almost completely washed out of the preparation,
except at the innermost margin of the cleft. x 1500.
Fic. 21. A poorly differentiated construction in a vegetative hypha.
Fics. 22-24, Empusa sp. ?
Fic. 22. Section of a conidiophore, showing the peculiar method of the cutting
off of eee cells, which apparently soon die. x1
- 23. Section of a conidiophore showing a subterminal branch, and also
be hae appear to have assumed amoeboid shapes. X 1
Fic. 24. A similar section to that shown in fig. 22, but with aly one basal cell
cut off. * 1080.
Fics. 25, 26, Empusa sp.
Fic. 25. End of a conidiophore, from which has been cut off two uninucleate
segments. X 1080
Fic. 26. A re which is apparently being pushed out of the thick, slimy
wall of the mother hypha. 575.
Fics. 27-29, Empusa sciarae.
Fic. 27. A section showing two protruded terminal cells of the conidiophore,
from which young conidia are in process of formation. x1
Fic. 28. A section of a conidium in process of shetrintion, 7 in which the iron-
haematoxylin stain has not differentiated the nucleus aay has brought out some
of the metachromatic bodies. X 1500.
Fic. 29. A mature conidium.
Fics. 30-32, Empusa culicis. 1500.
G. 30. The upper portion of the terminal cell of a conidiophore, showing at
sigs side the cleft marking the ring-formed cleavage-furrow.
Fic. 31. A section showing a somewhat older stage, in which the cleft has
almost abstricted the binucleate conidium
Fic. 32. A rare occurrence, showing a ‘pontiac conidium.
208 : BOTANICAL GAZETTE [MARCH
PLATE XV.
Fics. 33-35. Empusa culicis. X1500.
Fic. 33. A thin, lightly stained section of the end of a conidiophore, showing
the usual binucleate condition.
Fic. 34. An older stage, in which the sterigma is ‘ poditins ou
Fic. 35. From a still older preparation, in which the end : the sterigma has
become swollen to form the binucleate conidium.
Fics. 36, 37, Empusa americana. 1500.
Fic. 36. A preparation in which the nucleus is poorly differentiated, but
which ina especially clearly the vacuolar cytoplasm and the columella-like wall
which finally abstricts the conidium.
Fic. 37. A mature conidiospore, showing the vacuolated cytoplasm (probably
filled vation living with oil-globules), the single nucleus, and the basal papilla
which has been formed by the reversal of the columella.
Fics. 38-40, Empusa muscae. X1080.
Fic. 38. The partition-wall which cuts off the multinucleate conidium is here
Sihirteint: The slimy enucleate protoplasm of the basal cell appears to be but
ittle shrunken.
IG. 39.. The end of a young conidiophore, just extruded from the body of the
fly, in which the conidium is in process of formation, showing the multinucleate
character of the protoplasm. :
Fic. 40. A portion of a conidiophore still within the body of the fly, which is
growing toward an opening in one of the abdominal joints.
Fics. 41-44, Empusa aphidis. X1500.
Fic. 41. A section showing at the tip of the conidiophore a young conidium in
process of formation. |
Fic. 42. An older stage of conidium-formation,
Fic. 43. A mature conidium, showing the uninucleate character.
Fic. 44. A nucleus from a conidiophore, showing two nucleoles, and portions
of the chromatic thread.
; Fics. 45-48, Empusa culicis.
Fic. 45. The end of a young azygosporic hypha.
Fic. 46. An older condition, which shows the beginning‘of the formation of a
resting spore. X 1080,
Fic. 47. A mature ‘“azygospore.”’ X 1080.
Fic. 48. A thinner section of a mature eeu ”” showing but one nucleus.
Other nuclei lie below and above in the section. X 1500
a
DRIEPER A REGLE s
NEW NORMAL APPLIANCES FOR USE IN PLANT PHYSI-
OLOGY III
(WITH TWO FIGURES)
In the two preceding articles I described several pieces of apparatus
newly devised for educational work in plant physiology and explained the
objects I have in view in their development. In brief I aim to provide for
each of the principal physiological processes such apparatus as will be
accurate in results, convenient in manipulation, and obtainable by pur-
chase from a supply company. The company to which the manufacture
has been delegated is the Bausch & Lomb Optical Co., of Rochester, N. Y.
In the earlier articles I named the appliances ‘‘precision-appliances,”
which some, though not all of them are; they are, however, more properly
normal appliances, which I shall henceforth call them.
VI. PHOTOSYNTHOMETER.
No fact in all the physiology of plants is more important, and hence
more imperatively demands complete demonstration in botanical educa-
tion, than the absorption of carbon dioxid by green plants in light, with the
equivolumetric release of oxygen. There are simple ways of demonstrating
the process in part, and somewhat complicated ways of demonstrating it
completely; but hitherto there has been no simple method of demonstrating
the entire process in one operation. This is effected, however, by the new
photosynthometer described below, and illustrated in the accompanying
fig-1. tis called by this name for the reason that it permits photosynthesis
(the quantity of the photosynthate being a function of the quantity of the
gases absorbed and released) to be measured as well as demonstrated.
The instrument consists essentially of a pear-shaped plant-chamber set
in a firm iron base, a graduated measuring tube with a small stop-cock at the
upper end, and a connecting stopper furnished with a stop-cock of con-
siderable bore. The total capacity of the apparatus when closed is exactly
102°, of which the 2° is for a shoot and 100° for the gases concerned. The
proper amount of shoot is provided by selecting a small-leaved plant and
pushing a branch down into a measuring-glass until it displaces exactly 2°¢
* Continued from Bot. GAZETTE 39: 152. February 1905.
209 | [Botanical Gazette, vol. 41
210
BOTANICAL GAZETTE [MaRcH
of water; the water-level is then noted on the stem, which is cut at this
point under water, the shoot being later, when dried, placed upright in the
chamber. (It hardly
PIGS:
shows in the figure because of irregular reflections
from the surface of the chamber.)
Taking advantage of the fact that
the shoot will carry on photosynthe-
.sis for a time in an atmosphere con-
taining carbon dioxid in demon-
strably large amount, even up to 10
per cent. or more, we add some
selected percentage of that gas to
the apparatus in this way. The
measuring tube, with stop - cock
closed, is inverted and filled with
water of room-temperature, up to a
figure of the graduation expressing
the selected percentage, for the tube
is graduated in cubic centimeters,
which are, of course, percentages of
the total gas capacity of the appara-
tus.. The stopper is then placed on
the tube, and its stop-cock closed;
its hollow is filled with water and
inverted in a pneumatic trough (or
equivalent dish of water), which has
been standing in the laboratory long
enough to take the temperature of
the air. The lower stop-cock is
then opened and carbon dioxid
from a generator is allowed to enter
the tube, either from below or, as is
most convenient, through the top of
the tube. The admission of the
gas may be perfectly controlled by
cautious manipulation of the upper
stop-cock, which is closed at the
moment when the water has been
wholly driven out to the bottom of the bore of the lower stop-cock, which
point is held exactly at the water-level.
The lower stop-cock is now closed,
and the combination, which now contains exactly the desired percentage
1906] BRIEFER ARTICLES 211
of carbon dioxid, is lifted from the water, shaken free from all adhering
drops, and placed in position on the chamber. To prevent compression,
and therefore the presence of too great a quantity, of air in the
chamber when the stopper is pushed into place, tiny holes (visible
in the figure), matching in stopper and chamber-neck, allow free
release of such pressure, and the chamber is perfectly sealed by twisting
the stopper a little. The lower stop-cock is then opened, permitting
the carbon dioxid of the tube to diffuse into the chamber, a process
hastened by its gravitational flow. The apparatus now contains obviously
2° of plant and 100° of gas, of which a known percentage (say 5, 8, or 10)
is carbon dioxid, and the remainder is air. The instrument is now placed
in a bright light (not direct sunlight) for three or four hours; then the lower
stop-cock is closed (as shown in the figure), shutting into the tube a sample
of the gas of the chamber at the close of the experiment. The analysis of
this gas can be made at leisure, and is accomplished thus. The stopper and
tube are removed and placed upright in the pneumatic trough, deeply
enough to allow the stopper to be taken off without admission of air to the
tube. The zero mark of the tube is then brought exactly to the water
surface; the upper stop-cock is cautiously opened, permitting the water
to rise slowly to the zero mark, when the stop-cock is again closed,
shutting into the tube exactly ro®° of the gas to be analyzed. First the
quantity of carbon dioxid in the tube is determined, which is accomplished
by aid of a reagent tube, of the form shown at the bottom of the figure. This
tube, of glass, is provided with an extension of soft rubber tubing closed by
a screw clamp, and it is filled completely to the clamp with a strong solution
of caustic potash. It is slipped under the water of the pneumatic trough,
with the clamp closed; the air, if any, is squeezed from the upper part; and
the rubber is slipped over the lower end of the measuring tube which it grips
firmly. The whole is then lifted from the water, the clamp is opened, the
combination is inverted and the liquid is allowed to flow back and forth
several times from one tube to the other, when it will completely absorb any
carbon dioxid present. The clamp is then closed, the combination is slipped
again under water, and the rubber tube is pulled off, when the atmos-
pheric pressure will instantly force up the water to the exact extent of the
carbon dioxid absorbed, permitting the amount, and hence the percentage,
to be read off directly. Next a determination of the percentage of oxygen
present is made. This is effected by a precisely similar method, using an-
other reagent tube containing pyrogallate of potash, freshly made up in the
usual manner. A few inversions of the combination will result in absorp-
tion of all the oxygen, and the water-level in the tube when the rubber is
212 BOTANICAL GAZETTE [MARCH
removed will give directly the percentage originally present. Some slender
vessel may then be slipped under the tube which is removed and supported
as shown by the figure.
Thus may the gas exchange in photosynthesis be demonstrated accu-
rately, completely, logically, and conveniently.
n studying the process with beginners, the demonstration is the more
striking and conclusive to them if a second instrument is set up like (and
beside) the first, but covered completely from light; while even a third, like
these two except that it has no plant, may advantageously be added. The
comparative analyses of the gases at the close of the experiment give results
leaving nothing to be desired in logical demonstration.
For all ordinary purposes, water may be used in the pneumatic trough,
its slight absorption of carbon dioxid being negligible; but for great accu-
racy mercury may be employed. Similarly the corrections for capillarity,
vapor tension, etc., may for elementary demonstration be ignored, though in
precise work they would be taken into account. Temperature and baromet-
ric pressure are compensated, obviously, by the method of use of the instru-
ment.
A much larger instrument, capa-
ble of taking large leaves, branches,
or even entire potted plants, but
operated upon substantially the same
principle, is now in advanced prep-
aration and will be described later.
VII. ALUMINUM SHELLS FOR TRANS-
PIRATION EXPERIMENTS, ETC.
In transpiration studies with
potted plants it is of course neces-
sary to eliminate evaporation from
pot and soil. There are many ways
of effecting this, of which the best,
perhaps, is the use of a tin cup or
glass jar to cover the pot, and a roof
of rubber sheeting. The advantage
of this method over those in which
the plant is wrapped in rubber,
sealed in melted wax, etc., is this,—
that the rubber roof may be readily detached from the can or jar and
lifted, permitting a complete change of air to the roots when the plant is
* watered, thus contributing greatly to the health of the underground parts.
Pies Ss.
eT ee ee ae ee
1906] BRIEFER ARTICLES 213
The aluminum shells here figured (fig. 2) are designed to provide light, neat,
clean, and easily applied covers for pots on this principle. Flower pots are
now made so nearly in standard sizes that it is possible to make the shells to
fit them closely, and shells will be made for the present in 3-inch, 34-inch,
4-inch and 5-inch sizes. To hold the rubber roof tightly to the shell, a
tightly-fitting band or strap of aluminum, resting in a groove just below the
strengthened top of the shell, may be drawn to any desired tightness by a
convenient screw-nut, shown (though dimly) on the right in the figure. The
rubber roof may be attached to the plant in any of the ways ordinarily used,
but I find upon the whole the best method to be the following. In the
middle of a proper-sized piece of medium-thick rubber-sheeting, a hole a
little smaller than the stem of the plant is made with a cork-borer, and a cut
is made with scissors from this to the margin of the piece. It is then placed
around the stem, the cut edges of the central hole are stretched to overlap
a little, sealed together with rubber cement and held clasped until this sets.
Then a line of the cement is run to the margin, sealing one edge over the
other. When fully set, the margin of the rubber is clasped to the shell,
all surplus material is cut away, and a very neat and perfectly tight roof
temains.—W. F. GANonG, Smith College, Northampton, Mass.
CURKENT EITERATURE,
BOOK REVIEWS.
Bacterial diseases.
Tuts quarto volume on bacterial plant diseases, by Dr. ERwin F. SMITH,’ is
a manual for plant pathologists, which, with the author’s characteristic care in
clearly stating details, covers the subject, from the angle at which a tube should be
held during inoculation to plans for the construction of a water still and sugges-
tions as to the proper developer for photographic plates. One of the fundamenta
reasons for the abundance of half-finished work on bacterial plant diseases which
is annually imposed upon the public has been the lack in all literature of just such
a sae? " ah areca =nitee which the author has scattered through the work
uncertainty which now pervades the subject in the minds
of many would be oan dissipated.
The Carnegie Institution is surely treading on dangerous ground in publishing
such a manual, but if such action can be justified at all, it can be in this case.
Here the need is great and the subject is so specialized as to make such a publica-
tion impossible except at a financial loss. So great was the need that a proper
presentation of the material promised for the second volume could hardly be made
without a considerable portion of the matter here given.
The volume is divided into three parts; 186 pages being devoted to the text, 16
to useful laboratory formule, and 63 to bibliography. The latter covers the gen-
eral field of bacteriology, with the exception of plant diseases, this division being
reserved for the second volume. The publication is well illustrated, many of the
cuts showing the effect of bacteria upon plant tissue. The index covers the entire
volume and will be especially useful in connection with the bibliography, which is
arranged by subjects in the body of the volume.
g the many original things in this suggestive work, the discussion of
“keeping of records” and “nomenclature and classification” are of special interest.
Any one who has attempted to keep an accurate history of the behavior of a plant
parasite in the laboratory and in its host when the work has extended over a number
‘of years, has felt the need of an improved system. The subject of bacterial classi-
fication has been fairly quiescent for some years, and we have been busy trying to
fit the forms as they are found into MicuLA’s somewhat artificial framework.
The proposition of the author to replace MicuLa’s Pseudomonas by Bacterium
and send the group represented by B. anthracis masquerading under the name
of A planobacter will come to some as a discouragement
1SMiTH, E. F., Bacteria in relation to plant diseases. Vol. I. 4to. pp- xii
285. Washington: Carnegie Institution, 1905.
214
ap eee ne
1906] CURRENT LITERATURE 215
In spite of some minor things which seem inseparable from originality, this
work is of the first quality and should be in the hands of every plant pathologist.
—H. A. Harpine.
MINOR NOTICES.
Grasses of Iowa.— As a supplementary report for 1803 the Iowa Geological
Survey issues part u of the Grasses of Iowa,? prepared by PAmmet, BALL,
ScCRIBNER, and others. This is a descriptive and geographical study of the
grasses of the state, their general and economic aspects having been treated in
part 1. Under each genus there is the generic description, with synonymy, a
key to the species, a description of the species, often a figure, a list of localities
and a map showing the distribution of each form in the state, and a statement of
distribution in North America and elsewhere. There isa chapter on physiography
and geology, with a map, a section onecology, and a partial bibliography of
works on grasses. The work seems very complete and should be especially
serviceable to Iowa botanists. It is a pity state printers are so seldom skilfuy
ook-makers.—C. R. B
Connecticut fungi.— The recently established natural history survey of Con-
necticut has begun to show results, in the publication of two bulletins listing
the Hymeniales and Ustilagineae of the state. The formers lists 375 species in
65 genera, gives analytic keys to the genera, and illustrates the commoner species
by admirable half-tones, most of which are original. The species of smuts+ are
described with lists of hosts and distribution, and notes on economic features.—
oe
NOTES FOR STUDENTS.
Photosynthesis and temperature.—The interesting results of Miss iia
on temperature as a limiting factor for photosynthesis’ have now been extended
by her work in cooperation with BLACKMAN. ey have endeavored to
interpret the quantitative variations of photosynthesis, under approximately
natural conditions, in terms of the three limiting factors thereto, viz. (1) intensity
of illumination, (2) temperature of leaf, (3) pressure of CO,. When a leaf is
- 2PamMeL, L. H., Batt, C. R., and ScriBNer, F. L. The grasses of Iowa.
Part 0, Iowa Geological Survey, co report. 1903. 8vo. pp. xiv+ 436,
figs. 270. Des Moines, Iowa. 1904
3 WHITE, E. A., A pr re report on the Hymeniales of Connecticut. State
Geol. and N. H. Survey Bulletin 3. 8vo. pp. 81. pls. 4O. 1905.
4CLinTon, G. P., The Ustilagineae or smuts of Connecticut. Idem, Bull. 5. 8vo,
Pp. 45. figs. 55. 190
5 See Bot. GAZETTE 38 : 476. 1904.
BLackan, F. F., and Matraaet, G. L. C., Experimental researches in vege-
table assimilation and reanirations. IV. A quantitative study of carbon dioxide assim-
ilation and leaf temperature in natural illumination. Proc. Roy. Soc. London B.
76: 402-460. 1905.
216 BOTANICAL GAZETTE [MARCH
exposed to diffuse daylight alone the amount of photosynthesis is a measure of the
light, and it varies with varying light only when the amount of carbon dioxid
in the atmosphere is artificially increased and the temperature is kept high. Ifnot,
photosynthesis is limited i and is constant though the light vary. Isolated
leaves may rise more than 10° C. above a bright mercury thermometer in the
sun, a result quite at variance with Brown and EscomBr’s results,7 which, how-
ever, were calculated, not observed. Further study of this point is needed.
At normal temperature leaves are not able to utilize the full amount of energy
absorbed; helianthus could reach its maximum at 29° C. with about 68 per cent.
full sunlight and cherry laurel with about 36 per cent. When light is the limit-
ing factor equal intensities produce equal photosynthesis with leaves of most
various structure and type. At low temperatures leaves as different as helianthus
and cherry laurel have similar photosynthetic maxima, but at high temperatures
these diverge. Thus at 29.5°C. the former can fix twice as much CO, as the
latter, requiring twice as much energy to do it, of course. The essential differ-
ence in the photosynthetic activity in different leaves lies, then, in that they have
different coefficients of acceleration of this function with increasing temperature.
So in nature it appears that the low pressure of CO, (entailing slow diffusion
after solution at the surfaces of the leaf cells) and the low temperatures are the
serious impediments to food making.—
Root tubercle cultures——Much interest has been excited during very recent
years by work done in the Department of Agriculture concerning soil inoculation
with various root tubercle bacteria. Widespread and rather unfortunate notori-
t
knowledge concerning the root tubercle is to be attributed to the recent investi-
gations conducted i in the Department. This popular impression is of course
erroneous. The two distinctive contributions to this subject claimed by the
workers in the Department of Agriculture were that the nitrogen-gathering
ability of the bacteria was heightened by new cultural methods, and that a
method of transportation in dried condition, upon cotton, had been devised,
whereby pure cultures could be distributed readily to farmers.
Much skepticism has existed concerning the possibility of practically height-
ening the nitrogen-gathering power of the bacteria, and in a recent bulletin ®
Harptnc and Prucwa claim to have demonstrated by an examination of
eighteen of these cotton cultures that such packages are worthless for practical
— since the organisms are unable to survive upon the cotton or survive
uch small numbers as to be practically valueless. ‘Substantially identical
sl upon six of these packages were obtained in five separate laboratories,”
and the reviewer may add that similar results were obtained in his own
7 See Bor. GAZETTE 40 : 473. 1905.
8 Harpine, H. A., and Prucua, N. J., The apa of commercial cultures for
legumes. N. Y. Agr. Exp. Sta. Bull. 270: 345-385. I
1906] CURRENT LITERATURE 217
laboratory. The inability of the cultures to live is attributed to the method
of preparation and not to any knavery upon the part “of the commercial
producers. A test conducted by the authors of this bulletin demonstrated the
inability of the organism to survive to a satisfactory degree upon the cotton.
Any intention of opposing the idea of treating the seed of legumes with living
bacteria is distinctly disavowed.
It is exceedingly unfortunate that this method should have been given such
wide publicity and launched as a commercial enterprise until the question as
to its efficiency had been thoroughly tested.—F. L. Stevens.
Streaming of protoplasm in mucors.—This go-naten although very
striking and_easily observed, has been little studied. The ment was noticed
by WorRoNIN in 1866 in Ascophanus pulcherrimus. It was pears with consid-
erable detail in a number of species belonging to different genera by SCHROTER,
writer of the latest account,? in 1897, and the conclusion was drawn that the
Movement was dependent upon osmotic conditions. A careful study was also
ment was affected only very slightly by variation in the intensity of light. The
action of ether, extremes of temperature, pressure, wounds, variation in amount
of carbon dioxid, was similar to that of the same agents when applied to the
higher plants. The streaming is found to be due to osmotic action and trans-
piration and therefore does not occur in a homogeneous substratum, as for instance
when the fungus is wholly submerged, or in a saturated atmosphere. The stream-
ing is not a rotation or circulation, as in the hairs of roots and stamens and in the
cells of Chara, Nitella, Vallisneria, etc., but a backward and forward-movement,
in which the protoplasm, vacuoles, and nuclei participate. Occasionally the
acropetal movement is somewhat balanced by a thin peripheral layer of proto-
plasm without vacuoles setting up a basipetal movement. Usually the movement
-is toward one end of the hyphae for a longer or shorter time, then stops and
starts again in the opposite direction —J. C. ARTHUR.
Germination and radium emanations.—KO6ORNIcKE’® has continued his study
of the effect of radium emanations on the germination of ungerminated seeds which
have been exposed in both the dry and wet condition. His earlier tests were made
with radium bromid contained in glass tubes. In his later study he has used a
much more powerful mixture which was contained in tubes having one side of
.
9 SCHROTER, ALFRED, Ueber Protoplasmastrémung bei Mucorineen.* Flora 95:
I-30. 1905. satis :
10 K6RNICKE, M., Weitere Untersuchungen iiber die Wirkung von Réntgen- und
Radiumstrahlen auf die Pflanzen. Ber. Deutsch. Bot. Gesells. 23: 324-332. 1905.
218 BOTANICAL GAZETTE [MARCH
aluminium, through which the emanations pass more readily. In all the trials he
finds that although the germination is not prevented there is a period of retarded
growth in the seedling. The elongation of the root or stem may be temporary or
rmanent according to the duration of the exposure. In the latter case the
injured organ persists indefinitely without disorganization, but further growth of
the seedling occurs in the form of secondary members. In the case of Vicia Faba
such a condition will follow an exposure of only one hour; yet an exposure of four-
teen days does not prevent germination. Since the retardation of growth occurs
sooner in the root than in the stem of a given seedling, the author favors the expla-
nation offered by other investigators, who have worked on entirely different mate-
rial, that organs engaged in photosynthesis are more resistant to the emanations.
The author’s experiments offer no conclusive evidence on this point. Organs of
seedlings from seeds exposed to emanations retain geotropic sensibility as long as
they are capable of growth, the two capacities being concurrent. The same is true
of heliotropic sensibility. His earlier view that radium emits enough luminosity
to induce heliotropism, which was questioned by Mo tiscu, is maintained. Im-
portant as these results are, it seems to the reviewer that their value would be much
greater if obtained under standardized conditions—RAyMoND H. Ponp.
Anatomy of Matonia.—TansLry and Luria describe the development and
mature anatomical structure of a number of specimens of Matonia pectinata
gathered by one of them on Mount Ophir in the Malay Peninsula.": The coty-
ledons in this species are bilobed as in the polypodiaceous ferns. Below the first
leaf the central cylinder of the young stem consists of a rod of xylem, surrounded
by parenchyma alone; later phloem appears on the outside of the stele and in
the center as well. Subsequently the endodermis and “oround tissue”’ likewise
appear within the stele, which becomes typically siphonostelic. By a process of
“local dilatation of the m rgin of the leaf gaps” an internal mass of fibrovascular
tissue appears, which ultimately becomes tubular and lies within the original
fibrovascular tube. This inner tubular fibrovascular bundle subsequently gives
off an internal tracheary strand, which may also become tubular, so that there
may be in Matonia as many as three tubular bundles lying one within the other.
These join each other only in the region of the nodes. The authors consider the
internal fibrovascular system asa storage tissue only, since it has no direct con-
nection with the roots, which are attached to the external cylinder, as in other
ferns of this type. The views as to the morphological nature of the complex
fibrovascular system of the stem in this species may be regarded as “ orthodox,”
since the conclusion is reached that it constitutes a single stele. The hypothesis
that the pith is intruded cortex is accordingly rejected, since the authors are of
the opinion that the only trustworthy criterion as to the morphological value of
tissues is to be derived from a study of their relation to the aera meristems
of the growing point.—E. C. JEFFREY.
_TANSLEY, A. G., and LuLHaAM, Miss R. B. J., Astudy of the vascular system
of Matonia pectinata. Annals of Botany 19:476-519. pls. 31-33. 1905-
|
|
1906] CURRENT LITERATURE 219
Chloroplasts of sun and shade plants. —LuBimENKO” refuses to accept as a
general law the statement formulated by Turriazerr that the maximum photo-
synthesis occurs under an intensity of psiaenian ome to renee one half that of
direct insolation. By measuring the rate of phot hilous plants
(Tilia and Abies) and of ombrophobous plants (Betula and Pinus) under both
artificial and natural light he finds that plants differ as to the minimum insolation
necessary to initiate photosynthesis. In this result the author finds basis for the
conception of a specific quality of the chloroplasts. Further investigation coh-
vinces him that the curve of photochemical work is determined primarily by the
specific quality of the chloroplasts and by the anatomical structure of the leaf.
The influence of the latter factor is particularly evident during periods of moderate
sunshine, but the potency of the former is manifest under insolation of high or low
intensity.
The chloroplasts of the ombrophilous plants have greater dimensions and a
sensibility almost five ne Lereaiet than that of ie ca ameaamne of ese ena
plants. Other test te that the pig
roplasts of the former. :
While auxiliary data support the author’s main conclusion, the chance of error
through imperfect technique and ignorance of all the factors is so great that final
conclusions are better withheld—RayMonD H. Ponp.
Microsporangia of Sphenopteris.—It has been suggested that the microspo-
rangia of Lyginodendron Oldhamium =Sphenopteris (Crossotheca) Héninghausii
might be found on Telangium Scotti. K1pston"s concludes from its structure that .
Telangium Scotti cannot be the microsporangia of Sphenopteris Héninghausii.
In no instance was organic connection between the two demonstrated.
He found a few microsporangiate pinnae referable to Crossotheca Zeiller in
organic connection with barren pinnae of Sphenopteris Héninghausit. The fertile
pinnule is oval, entire, and attached to the rachis by a short pedicel, which is thick-
ened very slightly upwards before merging into the pinnule, to which it seems
to be united fora short distance. The pinnules appear to be rather thick, and the
vascular bundle which enters the pinnule divides into two branches, which separate
slightly from each other. Each fertile lobe bears six to eight broadly lanceo-
late, sharply pointed, bilocular microsporangia, which in early stages bend inward,
forming a small hemispherical bunch, with their apices meeting in the center,
Later, the microsporangia spread outward and appear as a fringe hanging from the
margin of the pinnule. The microspores are either slightly oval or circular and
Measure 50 to 75 “ in diameter. The walls are roughened, being covered with
minute blunt points. The tri-radial ridge, marking the line of cleavage of the
tetrads, is sometimes apparent.—W. J. G. Lanp.
12LUBIMENKO, M. W., Sur la sensibilité de 1’ ig chlorophyllien des plantes
ombrophiles et ombrophobes. «Rev. Gén. Bot. 17: 381-415. pls. 10, 11. 1905
™3KipsTON, R., Preliminary note on the occurrence 3 microsporangia in organic
connection with the foliage of Lyginodendron. Proc. Roy. Soc. B76: 358-360, pl. 6. 1905.
220 BOTANICAL GAZETTE [MARCH
Photic sense organs.—GuTTENBERG' has demonstrated that two of the om-
brophilous species of his local flora have a photo-sensitive epithelium, whose
response consists in maintaining the leaf in the transverse heliotropic position.
The mechanism is essentially the same as was found by HABERLANDT in the so-
called velvet leaves, so abundant among the ombrophilous species of the tropical
hydrophytic forests. The épidermal cells function as converging lenses, so that
the protoplasmic membrane which covers the floor of the cell is not uniformly
illuminated. In HABERLAND?I’s studies the bright spot was centrally located,
but GUTTENBERG finds that for his species that it is excentric, because the
papillosity is not centrally located. The result is the same in both cases, for
the leaf is attuned to the distribution of interior illumination which exists when
the leaf is in the transverse position. Actual tests showed that the petiole is
not a factor in securing this position. Curiously enough the leaf assumes the
horizontal position in diffuse light, such as occurs under the open sky on a
cloudy day. In this cee, however, the internal distribution of light is the reverse
of that which exists under parallel rays, the central area of the floor wall being dark
with a margin of increasing brightness. The stimulus apparently consists in an
unequal illumination of the cell lumen.—RAyMonpD H. Ponp.
Nature of chromatophores.— MErEsCHKOWSKY'S holds that these bodies are
not organs of the plant cell and never have been, but are foreign organisms
which penetrated into the colorless plasma of the cells and live there as
symbionts. In support of this notion he adduces the facts that the chromato-
phores multiply continuously by division and do not arise de novo; that they are
in high degree independent of the nucleus; that they are completely analogous
with zoochlorellae and zooxanthellae which inhabit hydras, infusoria, etc.; that
there are organisms, (e. g., the lower Cyanophyceae, such as Aphanocapsis and
Microcystis) which can be considered as free-living chromatophores; that
certain Cyanophyceae actually live as symbionts in the cell plasma. This theory
he thinks, is the only possible explanation of the polyphyletic origin of primeval
plants, which were merely amoebae and flagellates into which Cyanophyceae
migrated; that the green, red, and brown Cyanophyceae account for the algae of
these colors; that the plant cell-wall is due to the formation, by the symbiotic
chloroplasts, of carbohydrates easily polymerized into cellulose ; which w.
makes impossible the further taking of solid food and entails the quiescent nature
and simple organization of plants, minus nerve, muscle, and psychic life. Here
is another pyramid of theory resting on its apex.—C. R. B
‘4GUTTENBERG, H. R., von, Die Lichtsinnesorgane der Laubblatter von Adoxa
Moschatellina und Cynocrambe prostrata. Ber. Deutsch. Bot. Gesells. 23:265-273-
pls. 10, II. 1905.
5 MERESCHKOWSKy, C., Ueber Natur und Ursprung der Chromatophoren im
OT en a Biol. Centralbl. 25: 593-604. 1905.
*
1906] CURRENT LITERATURE 221
Idioblasts of Cruciferae. —ScHWEIDLER"® has decided to assign a systematic
value to the peculiar idioblasts of the Cruciferae. The author at present reserves
judgment as to their generic value, though this is expected to be established by
further work. He has no doubt, however, that suborders and tribes can be accu-
rately defined. On this basis he divides the family into three suborders. The
first is characterized by the presence of idioblasts which contain chlorophyll and
which are located exclusively in the mesophyll. The idioblasts of the second sub-
order occur in the vascular tissue and differ from those of the first group in not
containing chlorophyll. The third suborder is composed.of members which have
both kinds of idioblasts. Just what would happen to the systematic standing of an
individual so unfortunate as to have had the development of its idioblasts inhibited
is certainly not for the reviewer to say, but in view of the urgent necessity of estab-
lishing systematic work upon an experimental basis rather than morphological, it
is difficult to escape the conviction that more or less futility is involved in all those
efforts of which this paper is an example.—RayMonD H. Ponp.
Araucarineae.—A preliminary note by THOMSON?’ states that in Agathis
there are many supernumerary nuclei in the pollen tube and that in Araucaria as
many as thirty were counted. The pollen tube grows along the surface of the
ligule for 22“ or more before entering the micropyle. The anatomy of the ovule
and development of the archegonia, as well as of the pollen tubes and megaspore
membranes indicate that the Araucarineae occupy a very isolated position among ©
the Coniferales
SEWARD and Forp in an abstract of a paper'® read before the Royal Society
Dec. 14, 1905, indicate the scope of an extensive investigation of the Araucarieae.
The section headings are: Introduction, distribution, diagnosis and synonymy,
seedlings, root bis stem ATONE neers 7 presi rate reproductive shoots,
fossils, and phylogen lusion
The ions ‘Annee conclusion is that the group, unlike the Cycadales, has
been derived from lycopodiaceous ancestors. The Araucarieae differ so greatly
from the other Coniferales that the — suggest the substitution of the term,
Araucariales for Araucarieae—CHARLES J. CHAMBE
Inhibitory action —ERrerA’ suggests that the non-development of lateral
branches or their growth in a particular position (e. g., of certain conifers) is
determined by inhibitory stimuli (de nature catalysatrice si l’on veut) traversing
either bark (Araucaria) or all living cells (Picea). We may conceive, he says,
the apex of the stem or root asa sort of tyrant who forbids the subjacent
16SCHWEIDLER, J. H., Die systematische Bedeutung der Eiweiss- oder Myro-
sinzellen der Cruciferen nebst Beitragen zu ihrer anatomisch-physiologischen Kennt-
niss. Ber. Deutsch. Bot. Gesells. 23:274-285. pl. 1905.
17THOMSON, R. B., Preliminary note on the Araucarineae. Science 22:88. 1905.
3SEWARD, A. C., and Forp, Sibille, O., The Araucarieae, recent and extinct.
TOERRERA, L., Conflicts de préséance et excitations inhibitoires chez les végétaux.
Mém. Soc. Roy. Bot. Belgique 42 : 27-43- 3- Aug. 1905
222 BOTANICAL GAZETTE [MARCH
branches to erect themselves or in other cases to develop, though they have the
same tendency to do so as he; their geotropism or their power of growth is held
in check by his own. Suppress the apex, let it die or become enfeebled, and the
subjugated branches lift their heads. Several could erect themselves and take
the lead, and that is sometimes observed. But ordinarily a new conflict for
precedence occurs among the branches; the one nearest the apex or the most
vigorous near one early asserts its supremacy and in its turn keepsits rivals at its
feet. Cj. the independent and almost simultaneous amas of the like idea
y McCattium, Bot. GAzETTE 40 : 262. Oct. 1905.—-C. R
Ecological survey.—P 1 PRAEGER?° have a8 another vegeta-
tion map and ecological description to the list of vegetation surveys of the British
Isles. The area discussed lies south and west of Dublin. After a historical
introduction the geology, physiography, floristics, and survey methods are briefly
explained. The vegetation is divided primarily into littoral, agrarian, hill-
pasture, and moorland zones, and the woodlands. The zones are further sub-
divided into associations. These are described in detail and as far as possible
related to the factors determining their occurrence. The text is accompanied
by a map and five excellent plates of vegetation types. The paper will prove of
especial interest to those who have followed the work of R. SmitH, W. SMITH,
and Lewis in Scotland and England.—E. N. TRANSEAU.
Alternation of generations in animals.—In criticism of CHAMBERLAIN’S paper
on this subject?? Lyon?? holds that the phylogeny of animal gametes gives no evi-
dence of their being reduced or vestigial generations, comparable with the gameto-
phytic generation in plants; similarity of cytological processes does not prove
identity of morphological value in the two cases. He refers to the alternation in
Hydrozoa, and calls attention to the earlier proposal by BEARD and MuRRAY of
a theory similar to CHAMBERLAIN’s. In reply CHAMBERLAIN maintains’$ that his
-critic fails to distinguish between a gametophytic generation and a gametophytic
plant. He holds that the generations in Hydrozoa do not alternate in the
botanical sense, and points out that although reduction of the gamete-bearing
generation has not been proved for animals, there is strong evidence for its
aving occurred in plants—M. A. CHRYSLER.
Mechanics of secretion.— PANTANELLI*4 has attempted to ascertain whether
or not true secretion of enzymes occurs. He defines secretion as “‘the emission
2°PETHYBRIDGE, G. H. and PRAEGER, R. L., The vegetation of the district lying
south of Dublin. Proc. Roy. Irish Acad. B. 25:124-180. 1905.
2tBoT. GAZETTE 39: 137-144. 1905.
22Lyon, H. L., Alternation of generations in animals. Science N. S. 21: 666-667.
05.
?3CHAMBERLAIN, C. J. Alternation of generations in animals. Science N. S. 22!
208-211. 1905.
?4PANTANELLI, E., Meccanismo di secrezione degli enzimi. Annali di Bot. 3:
i13-142. 190
line
1906] CURRENT LITERATURE 223
of substances by living protoplasm, a thing possible through a self-regulated
change in the condition of permeability of the plasmatic membranes such that the
organism is able at pleasure to reverse it.” He finds that the ferment of Roman
bread and Chianti wine truly secretes invertase, by the augmentation of the
permeability of the protoplasm during the period of fermentative activity. is
increased permeability is general, various salts escaping more freely at the same
time. In Mucor stolonijer, however, the emission of invertase seems to have the
character of a free exit of materials from dying parts, coincident with spore
formation. Whether it has a true but weak secreting power remains for further
study.—C. R. B
Respiration.— PALLADIN distinguishess three sources of the respiratory CO.
of plants: (1) nucleo-CO, produced by the action of enzymes, which, partly
soluble, partly insoluble in expressed sap, are intimately bound up with the pro-
toplasm; (2) stimulation-CO,, formed by the protoplasm itself (apparently
directly) under the action of stimuli; (3) oxydase-CO., produced various
oxidases. The process which characterizes animal and plant life consists in the
excretion of nucleo-CO, which is formed by decomposition without the partici-
pation of atmospheric oxygen. Intramolecular respiration is a primary phe-
nomenon, whose CO, is principally nucleo-CO, and in some cases also
stimulation-CO,. But alcoholic fermentation is no simple phenomenon, and,
as KostyrscHew has shown, must be distinguished from intramolecular respira-
tion.— ub
an and pear rot.— LonGyEAR?® has published the results of his study of
a rot of apples and pears due to an undescribed species of Alternaria. The
same disease has so far been found in California, Colorado, and Michigan, and
in Colorado is one of the most widely distributed and common diseases of
apples. PappocxK?7 was the first to cali attention to it. In the case of the
apple the disease attacks the fruit only, but it attacks the fruit, leaves, and young
sprouts of the pear. In the apple it appears frequently first at the blossom end
of the fruit and, in the case of sorts having a deep calyx-tube, a core-rot may
occur. The disease may be controlled by spraying with Bordeaux mixture and
plowing under or removing the diseased fruits in which the fungus is able to
pass the winter.— E. Mrap WILcox.
Reduction division.—The earliest enh were scat non- a seron, In
phylogeny, according to SCHAFFNER,”® the conjugat 8
25 PALLADIN, W., Ueber den verschiedenen Ursprung der wahrend der Atmung
der Pflanzen ausgeschiedenen Kohlensaure. Vorlaufige Mitteilung. Ber. Deutsch.
Bot. Gesells. 20: 240-247. 1905. .
26 LoncyEar, B. O., A new apple rot. Bull. Col. Agric. Exp. Stat. ros: 1-12
pls. I-4. 1905.
27 PApDock, W., A new apple disease. Rept. Col. Exp. Stat. 17: 99. 1904
28 SCHAFFNER, JOHN H., The nature of the reduction division and related phe-
nomena. Ohio Naturalist 5:331-340- 1905.
224 BOTANICAL GAZETTE [MARCH
a disturbance into the life cycle and a reduction division of some kind became an
inevitable accompaniment. The places at which a reduction division might, theo-
retically, become established in the life cycle are presented in diagram and
described. A comparison between the life cycles of plants and animals is also
illustrated by a diagram. ScHAFFNER believes that in the higher animals the
condition appears to be similar to that found in Fucus.
The significance of a transverse division of chromosomes in interpreting the
phenomena of MENDEL’s law is illustrated and discussed. — CHARLES
Migration of salts— In an extensive investigation of the content of nitrogen,
phosphoric acid, sodium, and potassium in cultivated plants, both field and pot
grown, at different periods of their development, it has been found?9 that in
different plants the maximum absorption is completed at different periods, bar-
ley, spring wheat, peas, and mustard attaining this maximum at flowering, while
potatoes reach it at maturity. These substances do not remain at a maximum,
but in the plants other than potatoes and with the exception of phosphoric acid,
migrate back, in great part, to the soil; this seems to depend on the amount of
_ a given substance available, being greater san say, potassium is lacking than
if the appropriate materials are all supplied —C. R. B.
Anatomy and affinity — Another hese SARTON, has attempted to ascer-
tain how much help is to be had from histology in determining the validity of
Jordanian species as contrasted with Linnean.3° He studied allied plants, sub-
mitted them to cultivation under diverse conditions and then examined their
structure. In some cases there were constant anatomical characters distinguish-
ing apparently closely allied forms. On the other hand the characters were as often
elusive and evidently directly adaptive. Plants long cultivated in the Jardin des
Plantes and at Fontainebleau showed no anatomical differences from wild ones
of the same species. Nor were there differences between the varieties having
different colored flowers.—C. R. B.
Scotch moors.—T he succession of plants in the moors of the Scottish southern
uplands has been studied by Lewis.3! He finds that in all the localities visited
the peat “shows a definite stratification of plant remains, indicating a swing
from woodland to heath and moss, and again to woodland. In some districts,
an arctic plant-bed is interposed between the lower and upper woodland: beds.”
The- vegetation changes are probably correlated with climatic changes at the
29 WILFaRTH, H., R6mer, H., and WIMMER, G: Veet die Nahrstoffaufnahme der
Pflanzen in verschiedenen Zeiten ihres Wachstums. Landw. Versuchsstat. 63: 1-79
pls. 3. 1905.
3° Sarton, A., Recherches expérimentales sur l’anatomie des plantes affines-
Ann. Sci. Nat. Bot. IX. 2 : 1-115. pls. I4. 1905
3t Lewis, F. J., The plant remains in the Scottish peat mosses. Pt.J. The
Scottish southern chee Trans. Roy. Scc. Edin. 413:699-722.
aintentaleteehinieeati
1906] CURRENT LITERATURE 225
close of the glacial period. He concludes also that the differences in the basal
deposits of these moors as compared with those of the higher Cross Fell district
(upon which he reported earlier) indicate the relative time of origin.— E. N
TRANSEAU.
Aberrant chromosomes.—The discovery of chromosomes of different sizes in
the same nucleus in plants suggests that the attention of botanists be called to the
terminology just proposed by MontcomeEry for aberrant chromosomes in Hemip-
tera.3? The term chromosomes is retained when ‘all the chromosomes of a nucleus
are alike; when they are unlike, the name autosoma or autosome is applied to a
chromosome of the usual form, and allosoma or allosome to an aberrant chro-
mosome. Unpaired allosomes are monosomes, and paired allosomes are diplo-
somes.—CHARLES J. CHAMBERLAIN.
Iron-algae.—After observation in the field and a study of cultures,
GaIDUKov%3 concludes that a Conferva found by him in overflow pools of the
Ocka river near Rjasan accumulates iron oxid from the waters, just as other
algae do calcium carbonate or silica. He thinks such iron secretion not peculiar
to the bacteria, but characteristic of many organisms, not as a necessary life-
process, but as an adaptive one. In the present case it seems to be protective to
the akinetes, which, eos down by the iron oxid, sink to the bottom and
so pass the winter.—C. R. B
Photosynthesis and reste. Poxiacct34 announces that electric energy,
when it does not exceed a given intensity, promotes very much the formation of
starch in leaves, and that this effect is greater with a continuous current passing
directly into the interior of the organs. Electrified leaves almost deprived of
light in some cases showed starch formation, when, in the same illumination,
unelectrified leaves did not. In view of the recent English work on photosyn-
thesis these conclusions should be received with reserve— C. R. B
_ Formation of proteids.—MonTEMARTINI’S is attacking this much investigated
problem. His first paper clears the ground, records once more a good part of
the extensive bibliography, and details two experiments, which lead to the con-
clusion that the production of proteids is greater in light than in darkness, and
greater in light and air minus CO, than in light ‘and normal air. Likewise it is
fivefold greater in the day than in the night, and he proposes to analyze the
relation of light to this result in his later experiments.—C. R. B
32?MontTGoMERY, THos. H., The terminology of aberrant chromosomes and
their behavior in certain Hemiptera. Science 23: 36-38. 1906.
33GarDUKov, N. , Ueber die Eisenalge Conferva und die ang aoa des
Siisswassers in algemeinen. Ber. Deutsch. Bot. Gesells. 23 : 250-253- 1905.
34Pottacct, G., Influenza dell’ electricita sull’ assimilazione pastels Nota
preliminare. Atti I t. Bot. Pavia II. 11: 7-10. 1905.
SIL Da L., Primi studi sulla formazione delle sostanze albuminoidi
nelle piante. Atti R. Ist. Bot. Pavia II. 10: 1-20. 1995.
226 BOTANICAL GAZETTE {MARCH
Chemotaxis of sperms of Equisetum.—Liprorss,°° to avoid anticipation by
SurBata, has made a preliminary announcement of his discovery that the sper-
matozoids of Equisetum are markedly chemotactic toward solutions of malic acid
especially, and also to maleic acid and its salts. Only indifference is shown to
solutions of fumaric acid or of its salts. The threshold concentration of malic acid
he finds to be about M/1oo00. Aerotaxis, which had been previously observed in
the case of Marchantia spermatozoids, could not be demonstrated.—RaymonD H.
Ponp. ‘
Welwitschia.—Tumboa mirabilis is so little known that any fresh observations
are welcome. PEARSON37 succeeded in securing material showing the develop-
ment of microsporangia, microspores, megasporangia and megaspores. Observa-
tions were made upon the habit, habitat, and climatic conditions. It is probable
that the plant is partially, if not wholly, insect-pollinated, and that the processes of
fertilization and maturation of the seed take place more rapidly than in other
gymnosperms.—CHARLES J. CHAMBERLAIN.
The cycadean integument.—This is discussed in a recent paper by Miss
Stoprs,38 who takes this occasion to compare the structures of the cycad ovule
with those of the fossil Lagenostoma. The single integument of the living cycads
is regarded as a double structure representing two integuments of some ancestral
fo The plane of fusion of the two integuments has been between the inner
and outer layers of the stony coat, or between the stone and the outer flesh.
—CHARLES J. CHAMBERLAIN.
A rust-resistant cantaloup.— Birnn3° finds that the Pollock strain of canta-
loups is resistant to the rust or blight which is a common and serious disease in
the Rocky Ford district of Colorado. This resistance he found was transmitted
through seed selected from resistant plants, and hence seed selection becomes a
very practical method of controlling this destructive disease wherever it may
occur. The disease is due to the fungus Macrosporium cucum2rinum E. & E.—
E. Meap Witcox.
3°LipForss, B., Ueber die Chemotaxis der Equisetum-Spermatozoiden. seg
Deutsch. Bot. Coit 23: 314-316. 1905.
37PEARSON, H. H. W., Some observations on Welwitschia mirabilis Hooker.
Abstract of a communication to the Royal Society of London, Nov. 23. 1905-
38SToPEs, MARIE C., On the double nature of the cycadean ne pee
of Botany 19:561-566. 190 5.
39BLINN, P. K., A rust-resistant cantaloup. Bull. Col. Agric. Exp. Stat. 104:
I-15. pls. I-IO. 1905.
by oe ET he OS
NEWS.
Dr. JouNn W. HARSHBERGER is delivering a course of ten lectures on North
American trees before the Wagner Free Institute of Science in Philadelphia
Dr. Pear OLsson-SEFFER has left Leland Stanford University and bis ac-
cepted a position in connection with an experiment station in Mexico, devoted
to the investigation of the growing of rubber plants. His address, which corre-
spondents are requested to note, will be La Zacualpa Botanical Station, Escuintla,
Chiapas, Mexico.
THE ANNOUNCEMENT for 1906 of the Lake Laboratory, maintained by the
Ohio State University at Cedar Point, on Lake Erie, has been issued. The only
instructor in botany for the season is Prof. MALcoim E. STICKNEY, assistant pro-
fessor of botany, Denison University. ee, the courses are limited to
one in general botany and one in ecology. Th opens June 25th, and closes
August 3d.
THE MINNESOTA SEASIDE Station on the Straits of Fuca, Vancouver
Island, opens its doors for the sixth annual session, July 8, 1906. Owing to the
low rates to the Pacific coast which will be in force, this promises to be an impor-
tant year in the history of the Station. Those contemplating marine study and
research are invited to write to Professor Conway MacMillan, University of
Minnesota, Minneapolis, for the illustrated announcement of the Vancouver
Island Laboratory-Camp.
Dr. ALBERT SCHNEIDER has resigned his position as professor of botany,
pharmacognosy, and materia medica at the California College of Pharmacy and
has been appointed pathologist and physiologist of the Spreckels Sugar Com-
pany. Dr, Henry B. Carey, formerly assistant to Dr. SCHNEIDFR, has been
elected to fill the vacancy created by the latter’s resignation. DR. SCHNEIDER
is now giving his entire time to the investigation of the so-estied California seas
beet blight, which has been the cause of great losses to California beet grow
NUAL announcement of the Marine Biological Laboratory at WL
Hole, Mass., shows that the laboratories will open on July 5th, the regular
courses of instruction extending from that date to August 16th. The depart-
ment of botany for this year will be manned by Dr. Grorce T. Moore, of
Washington, D. C., and Dr. James J. Wotre, of Trinity College, N.C. Miss
J. Macrae will act as collector. Correspondence regarding botanical
courses should be addressed to Dr. Moore at the Cosmos Club, Washington.
THE Association internationale des Botanistes has shown commendable activity
not only in the conduct of the Botanisches Centralblatt but also in arranging for a
supply of pure cultures of fungi and alge. Now it further announces a long list
227
228 BOTANICAL GAZETTE [MARCH
of places from which it is ready to supply material for demonstration or investi-
gation to members of the society. The list is too long to be republished, but it »
is evident that one can secure working materal from a wide range of localities.
Correspondence relating to such material should be addressed to the secretary,
Professor J. P. Lotsy, Leyden, Holland.
BEGINNING with January 1, 1906, the form of the publications, which in
the past have appeared as bulletins of the Bureau of Government Laboratories
in the Philippines, will be changed to a journal to be known as the Philippine
Journal of Science. This publication will include original articles by members
of the staff of the Bureau of Science, as well as scientific papers submitted for
_ publication by other officials of the Philippine government and by individuals
not officially connected therewith. The journal will include researches in
ieee zoology, chemistry (including physiological chemistry), serums and pro-
phylaxis, mineralogy, geology, paleontology, mining, and mineral resources.
The journal is to review work which is being accomplished and to present such
original results as are obtained. The subscription price is $5 (U. S.) per year.
It will be possible to secure reprints of any particular series of the articles at
reduced prices. The journal will be edited by Dr. Paut C. FREER, director
of the Bureau of Science, with Dr. RrcHArD P. Srronc, chief of the biological
laboratory, and Mr. H. D. McCaskey, chief of the division of geology and
mining, as co-editors.
IN THE SUMMER of the present year a permanent Station for the study
of arctic science will be established on the south coast of Disco Island in Danish
West-Greenland. The cost of the foundation is defrayed by a gift from Mr.
A. Hotcx, of Copenhagen, and the Danish government has promised an annual
grant of 10,060 kroner ($3000) toward its maintenance. A laboratory, equipped
with appliances and instruments especially for biological researches, will be
attached to the Station, and for the present two work-places will be furnished
for visiting naturalists. The visitors will have the free use of the instruments,
seaeay outfit, and library of the Station. Lodging will be free and a small
e will be charged only for board. The first visitors can be received in 1997,
oa notices, ,inviting application, will be issued in due course. A library of
arctic literature is to be founded at the Station and to be made as pan as
possible, but on account of the limited resources of the Station and the vastness
of the literature, only a small proportion of it can be purchased. The Director
of the Station, M. P. Porsitp, asks botanists to be good enough to come to its
assistance by giving to this library works on arctic and antarctic nature, and
especially on arctic biology. The publications of the Station will be sent in
return, and the Station will be glad to render any service in its power. Up
to May 1 Director Porsirp may be addressed at Copenhagen S., Denmark.
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The Botanical Gazette
A Montbly Journal Mea es all Departments of Botanical Science
Edited by Joun M. CovtTer and CHARLES R. BARNES, with the assistance of other members of the
botanical stat ye the University of Chica
Vol. XLI, No. 4 Issued April 28, 1906
CONTENTS
CYTOLOGICAL STUDIES ON THE ENTOMOPHTHOREAE. II. Nucrear And CELL
IVISION OF EMPUSA (WITH PLATE Xv1) Edgar W. Olive - - . - - - 229
BIOLOGICAL RELATIONS OF DESERT SHRUBS. II. AssorprioN oF WATER BY
V. M. Spalding - ~ - - See Seo ie roe = 20"
NEW SPECIES OF CALIFORNIAN PLANTS (witH two FIGURES). Alice Eastwood - - 283
BRIEFER ARTICLES.
Notes ON NortH AMERICAN Grasses. VI. A.S. Hitchcock - : ~ - - 294
CURRENT LITERATURE.
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VEGETABLE FOODS.
MINOR NOTICES - - - - - - - - - = - ps e - 300
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VOLUME XLI ; NUMBER 4
BOTANICAL GAZETTE
APRIL, 1906
CYTOLOGICAL STUDIES ON THE ENTOMOPHTHOREAE.
II. NUCLEAR AND CELL DIVISION OF EMPUSA.
EDGAR W. OLIVE.
(WITH PLATE XVI’)
THE division of the nuclei in Empusa has been found in the course
of this investigation to resemble closely that described for Amoeba,
Euglena, and other Protozoa. Such a primitive type of nucleus,
which has been regarded as the typical protozoan form, has not so far
been observed in the Metazoa, nor have any of the lower plants
heretofore revealed a type of nucleus in which the “division-center”
is permanently intranuclear. Such a type has been called by Boveri
(:00, p. 183) a “‘centronucleus,” since it contains within itself a center
of division which he assumes may be either in diffuse or concen-
trated form.
The varieties of protozoan nuclei and-the types into which they
may be conveniently grouped are discussed by WILSON (:00), by
CALKINS (:o1), and at some length by CALKmNs in a recent article
(:03); hence we may concern ourselves here mainly with those forms
which appear to show nuclear conditions nearest those in Empusa.
SCHAUDINN published in 1894 an account of the division of the
nucleolus-like body in the center of the dividing nucleus of Amoeba,
and although he recognized that this appeared to play the chief réle
in nuclear division, he reserved, till further comparative studies, his
ideas on the mechanical details of the process.
‘As in this paper I shall have to refer frequently to the figures already published
in plates XIV and XV which accompanied my foregoing paper on The morphology
and development of Empusa (Bot. GAZETTE 41: 192- -208. March 1906), I have num-
bered the figures on this plate consecutively with them.
229
230 - BOTANICAL GAZETTE [APRIL
BLOCHMANN (’94) and KEUTEN (’95) first described the division
of the centronucleus in Euglena, and the latter author gave an inter-
pretation of the function of the nucleolus, giving to it the name
“nucleolo-centrosoma” (p. 219). According to KEUTEN’s observa-
tions, the nucleolus-like body of Euglena elongates in the prophases
of nuclear division, and functions as a kind of spindle, which, how-
ever, appears to be solid and homogeneous, and not fibrillar as in the
usual type of spindle. Other spindle substance and centrosomes, as
well as “pole-bodies,” the author could not find. The chromatin
forms many chromosome-like bodies, which, after passing through
an “‘equatorial ring” stage, are finally arranged in diverging daughter
groups about the elongated axial strand of the nucleolo-centrosome.
Just what the relation is between the dumb-bell shaped nucleolo-cen-
trosome and the dividing chromosomes is not made clear in KEv-
TEN’S figures, although he asserts that this axial rod governs the entire
process of nuclear division, since it orients the plane of division and
since the chromosomes move along it. Whether this intranuclear
body functions solely as an active fibrous mechanism for separating
the chromosomes, or whether its poles have in addition the properties
of centrosomes, are matters which should be more clearly determined
before we can make comparisons with the conditions in Empusa.
BoveERI (:00, p. 182, note) suggests in this connection that the
nucleolo-centrosome of Euglena is probably a concentrated and
sharply individualized intranuclear spindle. CaLkINs (:01, p. 265)
further points out what he regards as an analogy existing between
such a connecting rod in Euglena and the true fibrous spindle seen in
higher forms. ?
SCHAUDINN (:00) has described a type of nuclear division in the
sporozoan, Coccidium shubergi, parasitic in the intestine of a myri-
apod, which resembles even more closely that of Empusa. In the
growing individual, according to this author, the nuclei divide a
number of times, and finally, by a process resembling progressive
cleavage, the body is cut up into many uninucleate individuals, which
he terms merozoites. . The division of the nucleus at this time is by a
“primitive mitosis” (p. 230), totally unlike the double division which
takes place in later stages, following the fertilization of the egg. The
division in the first instance is quite similar to that of Amoeba and
1906] OLIVE—NUCLEAR AND CELL DIVISION OF EMPUSA 231
Euglena, and also resembles very closely that in Empusa. ‘The
second kind of nuclear division is regarded by SCHAUDINN as still sim-
pler, since centrosomes appear to be wanting entirely. The close
resemblance of the latter type to certain division-figures in Empusa
suggests, however, that the differences noted by SCHAUDINN may have
been more apparent than real, and that poor fixation, due perhaps to
the thickness of the membrane about the fertilized egg, may have been
the cause for his failure to find the intranuclear centers in these cases
also.
According to SCHAUDINN (p. 229), in the first mentioned division
the chromatin granules gather in the center of the primary nucleus
about a diffuse, slightly refractive substance, which stains less with
haematoxylin than the chromatin. There results finally a globular
central body, which he calls a karyosome, made up of two substances,
plastin and chromatin. Upon the appearance of vacuoles within it,
the karyosome grows larger, and it ultimately elongates to form a
dumb-bell shaped central core to the dividing nucleus. At this stage
the chromatin strands appear to radiate from the poles of the central
body, differing in this respect from the corresponding nuclear figure
of Euglena. The continued elongation of the central core is accom-
panied by the further massing of chromatin about the two daugh-
ter-halves of the central body, and the nucleus finally assumes a shape
comparable to an hour-glass. In the slender connecting strand which
unites the diverging nuclear halves there appears a peculiar Zwischen-
kér per which ScHAUDINN regards as probably a thickening of the
fibrous strand which connects the halves of the karyosome. After
the final constriction into two, the daughter nuclei, without entering
upon a period of rest, begin immediately a second division.
While those members of the Entomophthoreae which live parasitic-
ally in the bodies of insects have attracted attention for more than a
century, only a few investigators have published observations on the
coenocytic character of the mycelium of these fungi. Maupas
('79, p. 252) records having seen many nuclei in the hyphae of
Empusa muscae; while VurtLEMIN (’86) published drawings show-
ing a similar condition in Entomophthora gloeospora Vuil. Fatr-
CHILD (’97) also mentions having noted the multinucleate mycelium
in certain species of Empusa. BREFELD, who has studied the group
232 BOTANICAL GAZETTE [APRIL
more than any other investigator, has noted also (’84; p. 41) that the ©
hyphae of Conidiobolus, which grows parasitically on Exidia and
similar fungi contain many nuclei; but he contributes no comment
on the internal structure of Empusa, whose external characters and
development he has described in great detail.
CavaRA next published (’99) some cytological observations on
Empusa muscae, which was shown to have multinucleate conidia,
and on Entomophihora Delpiniana, with multinucleate conidia; and
he showed the importance of this character in delimiting the groups
of the Entomophthoreae, a point with which I heartily agree. But
Cavara’s account of the division of the nuclei in these two forms by
simple fragmentation is without doubt incorrect, as is plain from the
complicated method described in the present paper.
Finally, GALLAUD (:05) has quite recently studied a form, Dela-
croixia, apparently similar in habit to Conidiobolus, whose mycelium
as well as conidia contain numerous small nuclei.
The application of refined technique to the study of the cytology
of these organisms has resulted in but one paper—that by FarrR-
CHILD (’97) on Basidiobolus—which deals exhaustively with the
nuclear details. Erpam (’86), who first discovered Basidiobolus and
figured its uninucleate cells, and LoEWENTHAL (:03), both working
with unsectioned material, and quite recently WoycickI (:04), have
also contributed certain cytological observations in their studies on
this form.
Basidiobolus shows, as we shall see, little resemblance cytologi-
cally to Empusa, and RacrBorskI (’96) contends that it should not be
included in the Entomophthoreae. However, since this form is gen-
erally considered in connection with the group, it seems best to review
at this time FAIRCHILD’s account of the nuclear division in Basidio-
bolus. This author has described in great detail the peculiar division
by which the two small beak-cells are cut off from the adjoining gametes.
The division of the nuclei in these beaks bears little resemblance to that
in Empusa, nor, indeed, to the process in any other thallophyte so far
described; it rather resembles, according to FAIRCHILD, that in higher
plants, in that a cell-wall is laid down through the instrumentality of
a cell-plate. During the prophases of division, the nucleole dis-
appears, and the author thinks it is probably used to form spindle
wacniae
1906] OLIVE—NUCLEAR AND CELL DIVISION OF EMPUSA 233
fibers. The nuclear membrane, as in the higher plants, appears to be
dissolved and a barrel-shaped or cylindrical multipolar spindle is
formed. Strongly staining granules terminate each of the poles of
the broad spindle, and in the early phases, the many chromosomes
gather in an equatorial plate. In the anaphases, a double row of
granules appears in the equator of the spindle, which is regarded by
the author as forming a true cell-plate, since the new cell-wall is laid
down between them. It should be noted in this connection, how-
ever, that such a cell-plate appears to lack the earlier fusion of the fibers,
which, in the higher plants, invariably precedes the splitting and the
subsequent deposition of cell-wall substance between the two new
plasma membranes thus formed. Vegetative nuclear division was
also observed by FarRCHILD, who evidently regards the process as
essentially similar to that just described, although in this instance he
did not succeed in finding a cell-plate.
WovycickI (:04), on the other hand, while agreeing with FAIRCHILD
in general as to the events of mitotic division in Basidibolus, confirms
Racrporskt’s assertion (’99) that the new cell-wall grows centripe-
tally as a ring-formed growth, like that in Spirogyra, and in this case
entirely independently of the spindle.
NUCLEAR DIVISION.
The nuclei in the coenocytic hyphae of Empusa are comparatively
large, measuring frequently as much as 7-9 # in diameter, and are
thus especially favorable for a study of the phenomena of nuclear divi-
sion. In the vegetative hyphae they are usually spherical when in a
resting condition; while in conidiophores:or in similar elongated cells
the nuclei also often become greatly elongated. In the conidiophores
of Empusa sp. (figs. 23, 257), the resting nuclei may even assume
irregular and apparently amoeboid shapes.
The resting nuclei of the vegetative hyphae of Empusa sciarae
(figs. 19, 27, 57) have no nucleole-like bodies whatever, whereas in
other forms, e. g., E. muscae (figs. 38-40) and E. culicis (fig. 48), each
nucleus possesses one sharply defined nucleole. In still others, E.
aphidis (fig. 44) and Empusa sp. (figs. 22-26), the number of nucleoles
Figures numbered 1-48 will be found on plates XIV and XV. Figs. 49-67
are on pl. XVI, herewith.
234 BOTANICAL GAZETTE [APRIL
varies, since one, two, or sometimes even four such bodies may be nor-
mally present in a resting condition. In many cases where these
structures do occur, they appear to be surrounded by a clear space,
and some,show a filamentous connection with the chromatin (figs.
22, 44). In other instances (figs. 47, 48), no such clear space is seen,
but the nucleole appears instead to be closely surrounded by a mass of
chromatin.
In optical sections, the nuclei of E. sciarae (fig. 57) show darker
granules which are apparently connected by more lightly staining por-
tions, thus giving an appearance corresponding to the common concep-
tion of the chromatin and linin in the resting condition. But careful
focusing reveals rather a more or less homogeneous, much convoluted
thread, or filamentous material. Since I cannot, in fact, see any appre-
ciable differentiation into chromatic and achromatic portions, I am
inclined to regard the chromatin in this instance as resting in the form
of a spirem thread.
I am hardly prepared, however, to accept for these nuclei the ideas
of VAN WISSELINGH (’99) and of GREGOIRE and WyGAERTs (:03),
who think that there is no distinction between linin and chromatin.
For, though it is true that in the resting nuclei of Empusa the nuclear
material appears to be homogenous, during mitosis, on the other hand,
some parts retain the stain much more tenaciously than other parts.
One may bleach out an iron haematoxylin preparation, for example,
until only that portion of the dividing nucleus immediately around the
centers remains dark. However, whether this difference brought out
by staining is due to mere physical causes, I cannot say.
- Resting nuclei take the stain readily and are thus sharply differen-
tiated; whereas those which are in a state of division stain less deeply-
Hence in searching for division-stages, one has but to find those nuclei
which are lightly stained and from which the color has been more
washed out. But this applies apparently only to those nuclei which
are somewhat advanced in the process, for such differentiation is not
readily noticeable in very young stages. The earliest stages of nuclear
division in the two species of Empusa in which I have studied.
the phenomenon, E. sciarae and E. aphidis, in fact are not altogether
clear. It is to be hoped that other species will prove more favorable
for the beginnings of the process. It is not quite clear, for example,
1906) OLIVE—NUCLEAR AND CELL DIVISION OF EMPUSA 235
just what events are transpiring in such a nucleus as that figured in
fig. 49. But from the later condition shown in fig. 50 to near the
close of the telophases, a great abundance of successive stages affords
an easy interpretation of most of the events of nuclear division.
It is highly probable that fig. 49 illustrates the early divergence of the
two centers of division present in the middle of the nucleus, although
no clew is given in the preparation toward the solution of the puzzling
question as to the origin of these centers. In this figure a clearly defined
strand connects two darker regions, where, presumably, nuclear
material is being aggregated. A clear space, probably a cavity filled
with nuclear sap, separates the two centers and encloses the connect-
ing filament. Between jig. 49 and fig. 50 is plainly a large gap. In
the latter, the two centers are large, conspicuous, intranuclear bodies,
from each of which radiate in all directions granular fibers. These
fibers appear to connect in some instances midway between the
centers with those from the opposite system of fibers; others appear to
cross over the equatorial region and to be independent of the other
system.
Figs. 51-61 record successively the phenomena attending further
divergence of the centers and the massing about them of the material
of the divided daughter-halves of the nucleus. It may be noted in
these preparations that one of the first evidences of the activity in a
nucleus leading to division is the change from a globular to an oval
form. During the progress of the internal mitotic changes, the nucleus
finally becomes elliptical or oblong and greatly elongated. It may
readily be noted also that the long axis of the dividing nucleus corres-
ponds generally with the long axis of the filament. In some instances,
however, the nucleus lies obliquely across the hypha, presumably
carried about by cyclosis.
An increasing abundance of nuclear sap is shown in figs. 49-58.
In jig. 49, the clear portion is seen to occupy the space between the
two diverging centers. From the repeated occurrence of similar nuclei
in which the middle appears to be occupied by a clear space, it seems
probable that one of the earliest manifestations of mitotic activity in
the case of Empusa is the accumulation of karyolymph in the imme-
‘diate vicinity of the intranuclear centrosomes. In fig. 50 the nuclear
Sap apparently lies both between the two centers and in the interstices
+
236 BOTANICAL GAZETTE {APRIL
between the fibers. In fig. 51 a clear space is noticed at one side of
the dividing nuclear elements, whereas, in the more advanced stages
shown in figs. 52-55, sap lies mainly between and separating the
two active centers of division. In figs. 54 and 57, a region almost
free from fibrous material separates the daughter halves and gives
the appearance of a turgid, intranuclear, vacuolar cavity. It will be
noted in these instances as well as in still later stages (jigs. 56, 58,
60), that this nuclear fluid appears to exert pressure on the chromatic
elements, as evidenced by the curved line where the massed chro-
matic material borders on the vacuolar fluid. In fig. 56, which shows
the next step in the division of the nucleus following jig. 57, the
cytoplasm has constricted in two the mother-cavity, and in this
figure as well as in the similar stages shown in jigs. 58, 60, it will be
at once noticed that the solid constituents of the young daughter
nuclei occupy a pseudosynaptic position, and that the greater part of
the cavity of the daughter nucleus is occupied by a clear space.
Whether an osmotic pressure of the intranuclear fluid causes this
appearance, or whether it is due simply to the massing or contraction
within the nuclear cavity of the chromatic elements about the polar
centrosomes, can hardly be determined with certainty, but it is prob-
°
able that both forces are thus operative.
Fig. 58 illustrates an interesting deviation from the more common
median constriction of the elongated nucleus shown in jigs. 56, 59, 60.
Here a double cytoplasmic constriction has taken place, resulting in
two daughter nuclei and between them a vacuole, which is undoubt-
edly filled with sap from the mother-nucleus. Probably in this
instance the dividing nucleus became so greatly elongated that surface
tension operated in such a way that the encroaching cytoplasm con-
stricted it into three parts instead of two, as is usual.
In fig. 59 is shown an early telophase condition in which the solid
constituents of the nucleus are being redistributed throughout the
daughter nuclei. Here the movement of chromatic material, as is
characteristic for nuclei in this condition, is opposite to that seen in
early stages of nuclear division. Whereas in the early phases this
material moves toward and masses about the polar center, in the
teleophases, it moves in the opposite direction, away from the center.
In jig. 59 the center in each nucleus is still conspicuous, although a
a
1906] OLIVE—NUCLEAR AND CELL DIVISION OF EMPUSA 237
considerable portion of the mass, especially that in the upper daugh-
ter nucleus, has moved centrifugally, towards the nuclear membrane.
We note also in the upper nucleus of this figure what appears to be .
a thickening at the outer ends of the radiating filaments, and in the
lower nucleus some of the radiations are seen to be double.
I am inclined to interpret jig. 61 as a very late telophase, and as a
near approach to a resting nucleus; between this figure, however, and
fig. 59 there is obviously a wide gap. Such nuclei as that shown in
fig. 61 are comparatively common, however, and without doubt repre-
sent a stage in which the center now exists only as a focal region for the
attachment of the chromatic fibers to the nuclear membrane. Occa-
sionally one may see at this focal point, especially in preparations
stained with the triple stain, a very dimly defined body, apparently a
remnant of the old center, lying against the nuclear membrane. But
in similar preparations stained with iron haematoxylin, the core of the
old center seems to be entirely empty, while immediately around it a
dense chromatic mass persists for some time. In fig. 61, for example,
there remains hardly any visible evidence of the old center of division;
a few conspicuous fibers and a darkly stained mass which was accu-
mulated about the center remain, however, to mark its former posi-
tion, and the fibers now serve apparently to attach the main mass of
chromatin to the nuclear membrane.
Apparently such a nucleus is ‘‘polarized,” at least in so far as there
seems to be a special and possibly permanent focal point on the
nuclear membrane for the chromatic materials. Whether this is cen-
tralized in the same sense as Euglena, or permanently polarized, as in
the case of Phyllactinia (HARPER, :05), must be settled by further
investigation.
We see that, from very early stages, the centrosomes in these divid-
ing nuclei are conspicuous bodies, which grow larger and more con-
spicuous as division progresses, and this is due, in my opinion, to the
accumulation of nuclear materials about them. Each centrosome is
lighter in the middle and has a darker rim (figs. 50-67), a phenomenon
which I am convinced is partially due to refraction. But careful wash-
ing out of the stain sometimes leaves the middle totally bleached out,
while immediately around it some parts retain the stain. Each centro-
some thus appears possibly to have a core of plastin and a rim of chro-
238 . BOTANICAL GAZETTE [APRIL
matin, as is claimed by ScHAUDINN for the similar bodies of Cocci-
dium. Fig. 61 could therefore be interpreted as showing the rim of
chromatin, but the plastin substance of the middle core has entirely
disappeared.
The division of the nucleus just described for Empusa sciarae takes
place in the later vegetative stages, when cross partitions are frequent
and when the coenocytic cells are consequently comparatively short.
Among the four or five nuclei present in each cell in this condition, we
may occasionally find two nuclei in a state of division; generally but
one, however, divides at a time. The nuclei in a certain cell do not,
therefore, divide simultaneously, but each appears to act in entire
independence of neighboring nuclei.
There occurs in earlier stages of the vegetation of the fungus an
interesting modification of the process as described above. Figs.
62-65 illustrate late stages in the division of the nuclei found in long:
coenocytic cells, in which cross-partitions are few and far apart. It
will be remembered that during the earlier vegetative activities of
Empusa sciarae, nuclear division takes place much more rapidly than
cell-division, with the result that septa occur at rare intervals, while,
on the other hand, nuclei during this period are abundant. When we
come to compare the seemingly different type of nuclear division shown
in figs. 62, 63 with that shown in figs. 50-55, we note in each the intra-
nuclear centers and the radiating chromatic filaments mentioned above.
But here in the latter type the dividing nuclei assume an hour-glass
shape, similar to those of Coccidium as shown in certain of SCHAUD-
INN’S drawings, instead of the oval or elliptical shape characteristic of
the nuclei during the division above described. A careful comparison,
however, leads to the conclusion that the only essential difference
between the two types of division is in the amount of nuclear sap. In
the latter case there is lacking the clear space filled with nuclear sap,
between the separating chromatic filaments, so conspicuous in the type
above described; or at least the fluid is much diminished in quantity.
In jig. 65 some is still present in the constricted region; but between
_ the separating daughter halves in figs. 62, 63, as well as in fig. 64,
little sap, if any, is evident. Fig. 65 shows, in fact, a transition
between the elongated, hour-glass shaped nuclei of the latter type
and the oval ones of the former.
1906] _. OLIVE—NUCLEAR AND CELL DIVISION OF EMPUSA 239
There can hardly be any doubt, especially after we make compara-
tive observations on Empusa aphidis, which has a similar type of
nuclear division, that such stretched-out nuclei as are shown in these
figures get their peculiar form from the currents of protoplasm flow-
ing in these long coenocytic hyphae. Resting nuclei, as is well known,
are plastic to a remarkable degree, and thus, in long cells, may fre-
quently become much elongated; so it seems more than probable
that these dividing nuclei may likewise become stretched out in the
_ same way.
Figs. 66, 67 represent poorly stained nuclei of Empusa aphidis in
which division is taking place in a manner evidently similar in every
respect to that described above as the second type. Here too we have
vegetative hyphae in which septa are few and far apart; hence the
general protoplasmic movements must disturb considerably the divid-
ing nuclei. Practically all of the nuclei of this species conform to the
type shown in figs. 66, 67, for I have but once found a doubtfully ellip-
tical nucleus. The fact that the second type of division alone occurs
in the long tubular filaments of Empusa aphidis points therefore to
the conclusion that the stretching out of the dividing nuclei in these
instances is brought about by cyclosis. In this second type of divi-
sion here described, we can readily imagine that the protoplasmic
currents also assist materially in the constriction and final separation
also of the halves of the dividing nucleus. We may thus conceive, in
the one case, of the protoplasm as undergoing such limited movements
on account of its confinement in a short cell, so that the dividing
nucleus is but little disturbed, and consequently, by the accumulation
of karyolymph, it assumes a short oval or rounded shape; whereas in
instances where the cells are long and the protoplasmic currents there-
fore stronger, the dividing nuclei become drawn out and elongated,
and constriction becomes very early evident.
In jigs. 64, 65, we note an interesting phenomenon. Here occurs
an infolding at the poles, giving an appearance as if some stress had
indented the nuclear membrane at this point. I have not observed
this phenomenon in the oval nuclei of the first type, but it apparently
occurs not infrequently in nuclei of the second type. It is possible
of course, that the infolding may be an artifact, caused in some man-
ner by the reagents. Such cases furnish indisputable proof, at any
240 BOTANICAL GAZETTE [APRIL
rate, that the intranuclear centers are strongly anchored to that near-
est portion of the nuclear membrane situated poleward from them.
We may summarize these results pertaining to the nuclear division
of Empusa sciarae and E. aphidis as follows. During the early
stages of division the nuclei become less stainable and slowly change
from a rounded to an oval shape. Two diverging centers of division,
or centrosomes, become conspicuous near the middle of the nucleus.
Fibers may now be seen radiating from the two intranuclear centers,
some crossing the median line between the centers, others evidently
anastomosing with fibers from the other system. The nucleus elon-
gates still more and the opposed centers, each with its system of radi-
ating fibers, diverge farther and farther apart. The centrosomes
appear to increase in size as division proceeds, probably from the
aggregation about them of the chromatic material in the radiating
fibers.
In cells which are comparatively short, a space filled with sap is
early apparent between the diverging daughter masses, as well as in
the interstices between the chromatic fibers. This sap increases in
amount until in the oval, turgescent nuclei found in such short cells,
the middle portion becomes filled with it, and we note a clearer central
part, containing at first a few scattered fibers, separating the two polar,
darkly-staining regions. On the final withdrawal of the last chro-
matic filaments to the daughter-poles, the middle of the elongated
nucleus becomes perfectly clear and transparent. The cytoplasm
now encroaches on the median sap-cavity and, by constriction, cuts
the mother-nucleus in two. In some instances, a double cytoplasmic
constriction may take place, so that a vacuole filled with nuclear sap
is cut off and left between the two daughter-nuclei.
In long cells, on the other hand, or in filaments with few, far-sepa-
rated septa, the nuclear sap does not accumulate in the manner just
described; hence the nucleus, instead of becoming turgid with the
liquid secretion, becomes early in the process of division constricted in
the middle and greatly elongated, thus assuming the shape of an hour-
glass. A few connecting strands in the constricted portions remain
for some time, while the active polar regions, with their dense accumu-
lation of chromatic material, become separated farther and farther,
with the result that the two daughter-halves are finally pulled apart-
1906} OLIVE—NUCLEAR AND CELL DIVISION OF EMPUSA 241
The lack of accumulation of nuclear sap in the latter type of nuclear
division constitutes the only difference between this type and the one
described above. ©
The accumulation of sap in the nucleus in the first instance is prob-
ably due to the lack of disturbance of the process by the restricted
protoplasmic currents in the short cells. The lack of accumulation
of sap in the second instance is probably due to the disturbing
influences of the stronger protoplasmic movements which undoubtedly
take place in the long tubular filaments. In the first type the
chromatic substance in the newly formed daughter-nuclei comes to lie
in a mass at one side of the nuclear cavity, thus resembling somewhat
a synaptic condition. In the other, the nuclear materials of the young
daughter-nucleus, massed about the centrosome, are closely enveloped
by the surrounding cytoplasm, and not until later in the reconstruc-
tive processes which follow, does the nuclear sap appear.
Towards the close of division, the center in each nucleus comes to
lie close to, if not actually on, that portion of the nuclear membrane
nearest the pole. Its attachment and anchorage to the nuclear mem-
brane is proven by the frequent indentation of the membrane at this
point. In the young nucleus the center remains conspicuous for some
time, but finally, with the resumption of a resting condition, it becomes
entirely lost to view. In the resting nucleus, the nuclear materials
appear to be distributed more or less evenly on a much convoluted,
seemingly homogeneous, filamentous thread which resembles a spi-
rem.
We have now to emphasize, before entering upon a discussion of .
the general bearing of these facts, certain peculiarities at once notice-
able in this primitive mode of nuclear division. In the type first de-
scribed, the nuclear membrane plainly persists throughout the whole
process of division; also in certain nuclei of the elongated type, it
undoubtedly persists (figs. 65-67), although it is here not so conspicu-
ous. I am inclined further to regard a membrane as present around
the chromatic fibers in jigs. 62-64, notwithstanding the fact that in the
preparations it cannot be seen. The iron haematoxylin stain is prob-
ably accountable for the failure to bring out the membrane clearly in
this instance. Hence we may record at this point that in the case of
Empusa, the nuclear membrane is at least usually persistent through-
242 '~ BOTANICAL GAZETTE [APRIL
out the whole of nuclear division, and that, consequently, the entire
process is intranuclear.
Secondly, we note the absence of any definite chromosomes in this
peculiar division; and equally noticeable is the failure of the chro-
matic material to become aggregated into an equatorial plate, as well
the want of a definite achromatic spindle. Careful counts, however, of
the fibrous strands radiating from the centrosomes indicate the prob-
ability of a constant number of these chromatic fibers. I have in
many instances counted about sixteen of these radiations from the
polar view (jig. 55), but it is perhaps impossible to determine exactly
the correct number, on account of the great confusion of threads. I
believe, nevertheless, that these fibrous strands of chromatic material
represent the chromosomes, and further, that the two daughter nuclei
each receive an equal number.
There seems little evidence for the existence of a differentiated
achromatic spindle, but further study in related species may possibly
assist in determining what here may correspond to such a structure.
It is true that in fig. 64 is shown an indefinite, intrafibrillar substance
which might be taken for a spindle, but I am convinced that the thick-
ness of the section in this instance is responsible for this misleading
appearance. Careful observation reveals chromatic fibers in a lower
plane of focus and it is their great number and close proximity in the
background that probably causes the indefinite, washed-out appear-
ance between the sharply defined filaments. In fig. 54 also there is
shown a similar substance between the radiating fibers, whereas in
fig. 55 this is not so noticeable. Fig. 50 as well shows but little
nuclear substance other than that in the sharply defined chromatic
fibers radiating from the two centrosomes. -
Since all the dividing nuclear substance outside the centers is
apparently confined to the two systems of filamentous structures radiat-
ing from the centers, we must therefore conclude that there is no intra-
fibrillar spindle-substance. And, since we see also that these radia-
ting strands appear to be chromatic in their staining reactions and
not achromatic, the only conclusion which seems possible is that
there is no substance in the dividing nuclei of Empusa which can cor-
respond to an achromatic spindle. I am not prepared,. however, for
such an extreme belief, which would obviously much belittle the
1906] OLIVE—NUCLEAR AND CELL DIVISION OF EMPUSA 243
importance of a fibrous mechanism for the accomplishment of mitotic
division. _ ;
I should prefer to believe that the achromatic spindle substance,
probably present only in small amount, is a part of, and inseparable
from the deeply staining radiations. Should this be true, then we
may conclude that the kinoplasmic spindle-mechanism is bound up
closely with the radiating parts corresponding to the chromosomes.
Possibly the chromatin is here more nearly a liquid substance than is
usual, hence it may diffuse more readily throughout the linin basis, so
as to be indistinguishable from the latter. At. any rate, I should
regard the chromatic filaments radiating from the centrosomes as cor-
responding in part to the fibers of the more differentiated spindle of
higher organisms; and, further, since these mark the paths of the
chromatin, they must also correspond to the mantle fibres. In the
case of Empusa, so far as studied, there is obviously nothing which
can correspond to the central spindle of more complicated nuclei.
CELL- pst irra
Cell-division in Empusa, as in many other jade plants, takes
place in entire independence of nuclear division, and also apparently
remote from nuclear control. There is concerned in the process no
such fibrous structure as a cell-plate; since, in fact, no cell-plate is
ever formed at the close of the nuclear division described above.
Further, cell-division may not take place till long after all division of
the nuclei has ceased; hence coenocytic hyphae result.
The branched conidiophores of E. sciarae (figs. 16, 18), as well as
conidia in the process of abstriction (figs. 28, 30, 31, 36) furnish
especially fine material for the study of cell-division. Examples are
also occasionally met with in sections of vegetative hyphae (figs. 79, 21).
A striking feature of the process as seen in conidiophores and young
vegetative hyphae is the fact that in the cleavage of the cell, the new
ring-formed partition-wall grows across a wide vacuolar space. In
the case of the abstriction of the conidia, on the -other hand, and
probably as well in older vegetative stages, although I have not as
yet seen the phenomenon in the latter instance, the new wall
grows through a mass of cytoplasm. Fig. 18 shows clearly the
method of growth progressively inward of the ring-formed septum.
244 BOTANICAL GAZETTE [APRIL
The plasma-membrane which bounds externally the thin primordial
utricle has evidently been infolded at this point, thus forming a deep,
narrow furrow. The young partition-wall which is being deposited in
this groove can not be seen in the figure. We note further in fig. 18
that the two nuclei which are shown are in a state of rest; in fact,
nuclear division does not occur at all during the pre-fructifying.
period characterized by the formation of conidiophores. And in the
same figure we also see that the nuclei are separated by a wide
vacuolar space from that part of the cell in which division is pro-
ceeding, and that they are joined to the active region only by a nar-
row cytoplasmic connection. It seems reasonable to suppose that
cell-division, in this instance, is a cytoplasmic phenomenon and is
merely remotely or indirectly subject to nuclear control. In fig. 18
it will be noted that the stain is deepest at the inner margin of the
cleft, showing that in this innermost region in which the new wall is
being laid down, the cytoplasm is densest and most active.
Fig. 19 shows a similar ring-formed septum partly across a young
vegetative hypha, at a slightly advanced stage of growth. A bridge
of cytoplasm is next thrown across the vacuolar space before the wall
is completely formed, as is seen in fig. 20. This figure brings out
most clearly the region of greatest activity. In the preparation, the
stain (iron haematoxylin) was well washed out, so that the cytoplasmic
bridge as well as the ring-formed wall are left unstained except at the
innermost part of the furrow, where a small black granule is con-
spicuous. In this dark region the new wall is evidently being depos-
ited. Immediately on the throwing across of the cytoplasmic bridge,
the greater turgor of the cell below ordinarily causes the partition to
bend outward toward the outer end of the hypha (fig. 20). This
bending is also quite noticeable after the final completion of the par-
tition wall ( jig. 17).
A study of these figures might lead to the conclusion that we have
here a process exactly similar to that already described for certain
other fungi (see HaRPER, ’99, p. 506), in which the cleavage furrow
first cuts across the cell and the wall follows later. One would in fact
naturally come to this erroneous conclusion, since every one of the
drawings mentioned above, except perhaps figs. 19, 21, shows clearly
the circular furrow, but no sign as yet of the ring-formed septum.
1906] OLIVE—NUCLEAR AND CELL DIVISION OF EMPUSA 245
In these preparations, however, the thin, delicate walls are not at all
easy to differentiate. I am convinced that, unlike the cases just re-
ferred to, in Empusa a delicate wall grows simultaneously with the
cleavage-furrow and not later. The figures which show abstriction
of the conidia furnish sufficient evidence for this conclusion. In this
case, the process of abstriction takes place essentially like the cell-
division described for conidiophores, except that here the cleavage-
furrow grows through a mass of cytoplasm instead of through a cen-
tral vacuolar space. In jig. 38, the completed wall separating the
conidium from the basal cell of the conidiophore may be plainly
seen, since the protoplasm is shrunken away on both sides. But in figs.
28, 30, 31, 36, although the cleft itself at the base of the conidium is
brought out with diagrammatic clearness, the wall which accompanies
it is not so evident. Two reasons may be noted here, however,
which are not so apparent in the case of conidiophores, for the con-
viction that the partition-wall is also present in these instances. The
wall which cuts off the conidium, when completed, as was noted
above in the case of the newly formed septa in conidiophores, is forced
upward by the greater turgor of the basal cell, and here forms a kind
of columella within the conidium. While it is possible that the
cleavage-furrow itself might be stretched and forced upward in this
fashion, yet it is more than likely that the unsupported plasma-mem-
branes bounding the cleft could not withstand the considerable
pressure which is developed. A further reason for the belief in the
necessity of the cooperation of a ring-formed wall in these instances
is seen in the shooting off of the conidia immediately on the comple-
tion of their abjunction. In figs. 29, 37, 43, are shown conidio-
spores which have evidently just been shot off and in which the tur-
gescence of the protoplasm has now reversed the position of the cross-
wall, making a papilla at the base instead of an indentation. We
see clearly in jig. 37 the delicate wall shrunken away from the spore-
plasm. An uncompleted wall at the time of the discharge of the
conidium would evidently allow the escape of the protoplasmic
contents. . |
GENERAL DISCUSSION.
_ It is clear that the division of the nuclei of Empusa which has
just been described, although apparently resembling in some respects
246 BOTANICAL GAZETTE [APRIL
amitosis, is certainly much more complicated than a mere mass divi-
sion such as occurs in the latter process. In the division of the cen-
tronucleus of Empusa, we have, as was seen, intranuclear centers
of division, or centrosomes, which function as active agents in
nuclear division. Centrosomes, when they do occur, are, on the
other hand, supposed to take no essential part in amitotic division.
In the dividing nuclei of Empusa, we have also, besides active cen-
trosomes, an arrangement of the chromatin in radiating fibers
comparable to chromosomes, and, further, a simple spindle-appara-
atus. I should therefore separate the process in this form far from
amitotic division, although still regarding it as an extremely sim-
ple type of mitosis.
In the division of the nucleus in Euglena, the resemblance of the
phenomena to amitosis was regarded by KEUTEN as so striking that
he called the process in this organism a simple intergradation
tween direct and indirect division. In the case of Coccidium
SCHAUDINN remarks that the division of the nuclei takes place by a
“primitive mitosis.” Should ScHAUDINN be able to find, further,
as is probably possible with improved fixation, the division-centers in
his second kind of division, which occurs in the stages following the
fertilization of the egg, he should come to the conclusion that he has
here also not, as he concludes in his paper, a still simpler type than
the first, but a primitive mitosis essentially like the first. For in
the event of similar intranuclear division-centers occurring in both
cases, he would have two types of division somewhat comparable to
the two types mentioned above in Empusa, which, as we have seen,
differ from each other only in the amount of nuclear sap present,
and in the earlier constriction and elongation of the second type.
In Empusa, Coccidium, and Amoeba, the absence of an arrange-
ment of the chromatin during the prophases of nuclear division in an
equatorial plate, attests the extreme simplicity of the mitotic process
in these instances. The absence of this equatorial arrangement
leaves us, in fact, unfortunately in doubt as to the manner of the equal
distribution of the chromatin between the two daughter nuclei. If
we accept, however, the commonly accepted doctrine that “the
daughter nuclei receive precisely equivalent portions of chromatin
from the mother nucleus”? (WILSON, :00, p. 70), we must conclude
1906] OLIVE—NUCLEAR AND CELL DIVISION OF EMPUSA 247
that this equal division of the chromatin occurs somewhere in the
obscure prophases; in Empusa, e. g., probably long before the appear-
ance of the conspicuous centers seen in jig. 50.
The absence of the arrangement of the chromatin into an
equatorial plate prior to the divergence of the two daughter masses
possibly results from the poor development of the achromatic -spin-
dle, due to the small amount of linin present in the nucleus. To
this same cause is probably due also the failure to form definite chro-
mosomes in these simple organisms. In Amoeba, according to
SCHAUDINN’s observations (’94), there are apparently no radially
arranged chromatic filaments; while in Coccidium (SCHAUDINN, : 00)
and Empusa, evidently a still higher type obtains, since in both these
instances we have formed, rather late in division, filaments of chro-
matin, which undoubtedly correspond to the chromosomes, and are
radially arranged about the centrosomes.
The formation of an “equatorial ring” in the nuclear division in
Euglena, and of a more compact equatorial arrangement of the chro-
matin in Euglypha (ScHewiakorr, ’88), Actinospherium (HERT-
WIG, ’98), Paramoecium (HERTWIG ’95), Aulocantha (BoRGERT, :00),
and other Protozoa, certainly indicates the presence in these forms
of a more highly differentiated mechanism for the halving of the
chromatin. In all these cases, we note the early formation of
chromosomes, which are usually very clearly defined, and generally
a well developed spindle, consisting of both central spindle as well
as polar mantle-fibers; so that we are justified in the conclusion that
in these more highly differentiated figures there is a greater amount
of intranuclear achromatic substance present than in the nuclei of
Empusa and Coccidium.
We may compare at this point the degree of differentiation of the
intranuclear spindle in these organisms. KEUTEN regards the dumb-
bell shaped nucleolo-centrosome in Euglena as probably serving as
a spindle mechanism; and Boveri (:00) and CALKINS (:01,:03) also
think that the strand of connecting substance in this constricted
nuclear body corresponds to the central spindle of higher organisms.
CALKINS (: 01, p. 265) points out in this connection that Paramoecium
furnishes a clew to the relationship of such connecting strands in
Euglena to the fibrillated central spindle, since in Paramoecium the
248 BOTANICAL GAZETTE [APRIL
“central portion of the division-figure is a single strand which widens
and becomes fibrillated at the ends.”” SCHAUDINN (:00, p. 229) eVi-
dently does not so regard the corresponding portion of the dividing.
nucleus in Coccidium, since he calls this connecting strand simply
“Verbindungsfaden der Tochterkarysome,” and says that “von
Spindlefasern und Poldifferenzirungen ist keine Spur wahrzuneh-
men.” I am also inclined to believe that no part of the constricted ©
nucleolar body in Euglena and Coccidium is homologous with the
central spindle of more complicated nuclei, since in all cases where a
structure occurs which can be positively referred to the central spin-
dle, it consists of usually distinct fibers which extend between and
connect the diverging chromosomes. In these instances, the con-
necting portion of the dividing nucleolar body bears no such relation
to the chromatic filaments, but instead it lies simply as a slender core
in the axis of the mitotic figure. Further, in the centronucleus of Em-
pusa, which is undoubtedly similar in every respect except this one
to that of Coccidium, such a connecting body does not occur at all,
unless, indeed, it be represented in fig. 49. - Therefore, the strand
connecting the constricted nucleolo-centrosome of Euglena and
Coccidium, in my opinion, does not represent, phylogenetically, the
central spindle, nor in fact any structure of the higher nuclei, but is
a structure which is confined, so far as yet known, to these two Pro-
tozoa. It is just what ScHAUDINN calls it, viz., simply a drawn-out
filament connecting the daughter centrosomes, which has no appar-
ent function. On the other hand Paramoecium, as shown in HERT-
wic’s figures, shows a true central spindle, and the final median
constriction of this spindle and the consequent aggregation of the
bers of the middle portion into what appears to be a single strand,
does not present a figure which can be in the least compared, as
CaLkrns claims, with the nucleolo-centrosome described above. If
there be any indication at all of central spindle in these simpler cen-
tronuclei, then, in my opinion, it must be looked for in the dimly
defined, continuous, bluish substance, for example, shown in the
drawings of Coccidium (see SCHAUDINN’s figs. 31, 32), which lies
between the daughter chromatin masses. SCHAUDINN himself, how-
ever, evidently believes that these are not spindle fibers. In the case
of Euglena, the central spindle is probably represented by the dim
1906] OLIVE—NUCLEAR AND CELL DIVISION OF EMPUSA 249
achromatic substance remaining between the separated chromosomes,
e. g., in KruTen’s fig. zz. But in Empusa, there is no appreciable
achromatic substance in the corresponding equatorial region of figs.
62, 67. There is, therefore, according to my interpretation, in the
simple cases where no equatorial arrangement of the chromatin takes
place, practically no development of a central spindle; but whether
these two facts are related somehow as cause and effect must await
further investigation. Hence we may regard the intranuclear figure
in the case of Empusa and Coccidium, as an extremely simple
apparatus, which consists merely of the two opposed centers of divi-
sion, each with its system of polar radiations. Further, these polar
rays must all correspond in function to the mantle-fibers, instead of
in part to the extranuclear polar asters of the higher animals, since
they all mark the paths of movement of chromatin material. As
seen in figs. 50, 54, 55, 59, for example, the fact is quite apparent that
the intranuculear centrosomes lie some distance from the nuclear
membrane, and that there is no appreciable differentiation in the
radiations which extend in all directions from them. All appear
alike to consist, at least in part, of chromatin material. In later
stages, represented in figs. 64, 65, the centers appear to have been
pulled to the periphery so that they come to lie against the nuclear
membrane. I am inclined to think that this peripheral position
represents the ultimate fate of all of the centrosomes, since the very
last stages (e. g., fig. 6r) almost invariably show the old centers lying
at one side against the nuclear membrane. Such figures lead us to
believe that after all there may be a slight differentiation in the astral
radiations, since those fibers which attach the centrosome to the
nuclear membrane may be mainly concerned in this peripheral move-
ment of the centers, forming in these instances a sort of “antipodal
cone” of fibers. At any rate, while there may be, in such a spindle,
certain polar structures which appear to have a special function and
thus to form an “antipodal cone,” there is no such striking differen-
tiation of the aster into a “principal cone” and “polar rays”’ as was
described by VAN BENEDEN.
In those more complicated centronuclei in which the chromatin is
gathered during nuclear division into an equatorial plate and in which
definite* chromosomes are formed, as in Euglypha, Paramoecium, and
250 BOTANICAL GAZETTE [APRIL
other Protozoa, a more or less clearly defined, fibrous, central spindle
is found in addition to the mantle fibers. The absence of the central
spindle in the simpler type of intranuclear division seen in Amoeba,
Coccidium, and Empusa, and its meager development even in more
complicated cases, clearly suggest that the central spindle-fibers, when
present, play only a minor réle in nuclear division as maintained by
HERMANN (’91), viz., that they are non-contractile supporting ele-
ments, which form a basis on which the movements of the chromo-
somes take place. The chromatic structures in Empusa are undoubt-
edly moved poleward without the assistance of such connecting
fibers, and they seem to be supported entirely by the surrounding
nuclear sap.
These facts may be interpreted as thus furnishing a strong argu-
ment against the acceptance of the ‘“‘pushing theory” of DRUNER
(’95), who supposes an active growth or elongation of the central
‘spindle, thus pushing the spindle-poles farther and farther apart;
and at least in part against the suggestion of MorriER (:03, :04) ,who
thinks that the chromosomes may be conveyed to the poles both by a
pushing and a pulling action of the spindle-fibers.
No clear explanation of the mechanism which accomplishes these
primitive divisions has yet been marked out. As pointed out above,
there are in Empusa no specially differentiated mantle-fibers, since
the radiating astral rays of the intranuclear figure themselves mark
the paths of the chromatin-movement. Whether the movements
which take place in these radiations are ienieeel to wee which ue
in the aster of the more highly di
I cannot say, but this seems quite probable. In Empusa the radiations
extend in all directions from the centrosome and some are anchored
firmly to to the persistent nuclear membrane at its nearest point,
while others project into the nuclear cavity, apparently ending free in
the karyolymph. Now, a contraction of the radiating fibers would
undoubtedly accomplish just the phenomenon which we see takes
place. The fibers seem to shorten and to thicken, and an appearance
suggesting an accumulation of darker staining material immediately
around the centrosome results. The distal indentation of the nuclear
membrane which we see occasionally (figs. 64, 65) should also be
regarded as strong evidence that a pull of some sort or a contraction.
1906] OLIVE—NUCLEAR AND CELL DIVISION OF EMPUSA 251
of fibers in this region has taken place. But I can see no evidence in
this instance of a using up of any of the material which has accumu-
lated about the poles, as has been suggested by STRASBURGER (:00)
to explain the shortening of the mantle-fibers in certain cases.
It may be pointed out in this connection that the fact that the
fibrillar radiations in Empusa appear to be almost homogenous, and
further, chromatic in their staining reactions instead of achromatic,
does not seriously detract from the reasonableness of the contractile
hypothesis, as applied to this form, since it is only necessary to assume
that contractile linin is also present in small amount in the fibers,
along with chromatin. WHuLSON (’95), in fact, maintains that in the
case of echinoderm eggs, the fibers are derived not merely from the
linin-substance, but also from the chromatin.
As in the telophases of mitotic processes in general, in the ‘Nite
stages in Empusa a centrifugal movement of the chromatin sets in,
which may sometimes begin even before the two daughter-nuclei are
separated by constriction from eachother. Fig. 59 shows such a late
condition, in which the chromatin-movement seems to be of the
nature of an active outward growth, since we now note at the distal
ends of the fibers accumulations of darker and apparently denser
material. Should we assume that the aggregation of chromatin
about the centrosome in the first instance is brought about by the
contractility of the kinoplasm in the radiations, then we must sup-
pose that later some subtle change occurs in the body of the
centrosome itself, or else in the fibrillar rays, to stop contraction
and to set up an opposite growth of the fibers. But I should regard
it as not an impossible assumption that the centrifugal movement in
the latter instance might be brought about simply by a loosening
up the chromatin in the increasing nuclear sap by which it is sur-
rounded—a phenomenon which would probably follow as a mere
mechanical consequence the final cessation of the forces which
caused the centripetal movement.
The suggestion that these alternating centripetal and centrifugal
movements of the chromatin are of the nature of flowing movements
appears to gain some support in the case of Empusa. MONTGOMERY
(:01, p. 352) concludes that this flowing movement of the nuclear
materials is automatic; but I fail to see how this author can retain,
252 BOTANICAL GAZETTE [APRIL
even in part as he does, the idea of the contractility of the secondary
linin-fibrils, in addition to the above theory, since an automatic move-
ment such as he conceives to take place should be regarded as an
amoeboid movement in response to chemotropic stimuli. WILSON
(:O1, p. 575) also regards the chromatin as ‘a liquid substance which
may be absorbed or given off by an achromatic basis such as plastin
or linin, and may thus flow from one part of the nucleus to the other.”’
The latter author appears to adopt to a certain extent the ideas of
BUTSCHLI (’92), in that in his studies on Toxopneustes he has become
thoroughly convinced that the astral radiations are in part the result
of centripetal currents, or diffusion-currents, of hyaloplasm converg-
ing on the centrosphere.
While it is quite possible that the chromatin in Empusa is a liquid
substance which may flow or diffuse about through an achromatic
linin basis, as WILSON suggests, this, in my opinion, does not preclude
the idea of a contractile linin substance serving as the mechanism of
mitotic division. I must say, however, that while entirely convincing
evidence is lacking that the primitive mitosis in Empusa is accom-
plished by means of a contraction and a growth of the fibrillar, kino-
plasmic radiations, there is, on the other hand, even less evidence in
favor of other theories; for example, that the movement of the chro-
matin is automatic, due to chemotropic forces which are supposed to
emanate from the centers; or that this movement is due to diffusion-
currents induced by the chemism active at the centers; or that it
results from magnetic or electrostatic forces, an idea which has been
recently revived by LILLIE (:05).
In the primitive mitotic division characteristic of Empusa, we see
but little resemblance to the corresponding process as described for
other low plants. In all these cases, even in the Myxomycetes
(HARPER, :00), a well-defined spindle and chromosome, and an
equatorial arrangement of the chromatin may be observed. It is
apparently very common among the thallophytes that the nuclear
cavity and membrane persist during a large part of the mitotic pro-
cesses; see, for example, figures of Erysiphe (HARPER, ’97), of Albugo
(STEVENS, :01), of Dictyota (Mortrer, :00). But in all of those
thallophytes in which centrosomes occur, the latter are extra- and
not iira-nuclear bodies. Empusa is therefore in this respect unique
1 je eee EE en
i
1906] OLIVE—NUCLEAR AND CELL DIVISION OF EMPUSA 253
among the thallophytes and may be regarded as a primitive form; and
further, the fact that it possesses intranuclear centers of division may
perhaps be regarded as adding another point in favor of HERTwic’s
(’98) view as to the intranuclear origin (by the extrusion of centro-
some) forming substances from the nucleus of the extranuclear
centrosomes of the more highly differentiated organisms.
It has been already pointed out above that Basidiobolus, which
has been generally regarded as a member of the Entomophthoreae,
shows in its mitotic features (its broad, multipolar spindles, and its
formation, according to FAIRCHILD (’97), of a cell-plate), as well as in
other morphological aspects, wide differences from Empusa.
Cell-division by means of the growth inward of a ring-formed wall
is apparently a common type of division among the filamentous thal-
lophytes. Such a constriction of the cell has so far been shown for
Beggiatoa (Hinze, :o1), the blue-green algae, Ulothrix (D1rPet ’65),
Spirogyra (STRASBURGER,’80), Cladophora (Davis,’04), the red algae,
and a few-other forms. WovycickI (:04), contrary to FAIRCHILD’s
(’97) assertions, contends that the cell-wall in Basidiobolus also is a
centripetal growth. The gametes of Sporodinia and the conidia of
Erysiphe are cut off in a similar manner, except that, according to
Harper (’9Q), the ingrowth here is simply a deep "narrow furrow and
not the growth inward of a ring of fungus cellulose. The wall in this
case is deposited later between the two plasma-membranes.
As has been shown in this study of Empusa, the ring-formed cleav-
age-furrow starts at a definite region of the plasma-membrane, some-
times remote from the nuclei; and further, the nuclei at no time ap-
pear to be concerned, directly, in the process. ‘TOoWNSEND’S (’97)
observations on nucleated and enucleated fragments of protoplasm
leave no doubt, however, as to the ultimate necessity of the presence
of a nucleus, in order to initiate the cytoplasmic activities in Empusa
which lead to cell-division. Whether the localized stimulus in this
case results first in a deposit of a ring of cellulose-substance on the
inner surface of the wall of the mother-cell, which might then by its
growth progressively inward be regarded as the agent of cleavage;
or whether there first occurs in this region an infolding of the plasma-
membrane, thus resulting in a circular furrow, to be soon followed by a
deposit of wall-substance in the cleft, Iam not able to state. In either
254 BOTANICAL GAZETTE [APRIL
case, at any rate, the importance of the plasma-membrane as a factor
in such a cell-division should be emphasized. The process in Empu-
sa, in fact, would seem to furnish an argument in favor of NOLL’s (:03)
view that in Bryopsis the controlling factor of embryonic growth is
located in the Hauischicht. Apparently in Empusa a definite region
of the plasma-membrane is stimulated to action, a ring-formed infold-
ing of the membrane occurs, and at once in the cleft thus produced, the
new wall begins to be deposited. A darker accumulation, presum-
ably of kinoplasm, may now be seen at the inner margin of the cleft
(jigs. 18, 20), where the activities leading to the cleavage of the in-
pushing plasma-membrane and to the ingrowth of the partition-wall
‘are evidently greatest. But the plasma-membrane does not alone
seem to be the active agent for these phenomena, for fig. 18 shows a
darker portion, having appreciable thickness, which apparently marks
a more or less broad region of concentration of kinoplasm. This fact,
therefore, may be regarded as an argument against the plasma-mem-
brane itself being the sole controlling factor in this case. Further,
MortieR’s (’99) experiments on Spirogyra and Cladophora, in
which, by reason of the disturbance due to centrifugal force, the cel-
lulose-ring, when once begun, was never brought to completion, not-
withstanding the fact that the plasma-membrane was still intact, fur-
nishes very convincing evidence against the acceptance of the theory
that the Hautschicht alone is the controlling factor of wall-formation
in these instances.
The fact that the cleft and ring-formed wall are finally carried
across a wide vacuolar space (figs. 17-21), will not permit of the
application to this case of SWINGLE’s (:03) explanation for the mechan-
ism of the cleavage in Rhizopus, Phycomyces, and other forms. For it
seems impossible to conceive how local contractions of the cytoplasm
could cause the constriction of the cell in the case of Empusa. We
could perhaps think of such a contraction as initiating the process,
but that these forces could obtain after the narrow diaphragm of cyto-
plasm had begun to be pushed across the central vacuole, seems to
me inconceivable.
In certain instances in Empusa, as, for example, when a germ-
tube is formed (jigs. 8, g), the end cell of a filament keeps cutting
itself off from behind, thus enabling the body of the protoplasm to
1906] OLIVE—NUCLEAR AND CELL DIVISION OF EMPUSA 255
travel forward, so to speak, and to seek a favorable environment in
which to grow. BREFELD (’83) seems to think, in the case of the
similar phenomenon in Ustilago, that the cells which are thus cut off
behind are empty, and that in this way no protoplasm at all is wasted
in the process. If this were true, a cell-wall would be formed from a
single plasma-membrane, thus differing from the division described
above, in which the membrane is split so that the wall is deposited
between the two. But the apparently empty cell retains its turges-
cence for a time before collapsing, thus proving that there is at
least a film of protoplasm present. Further, sections of similar con-
ditions in which conidia are cut off from a basal shooting-cell (jigs.
28, 36, 38), show clearly the thin primordial sheath of enucleate
protoplasm in the lower cell. The fact that the protoplasm of these
lower cells seems to undergo speedy degeneration contributes another
point in favor of the idea of the vital importance of the nucleus
in nutrition.
I am at a loss to understand why the conidium should be regarded
by THaxTER (’88, p. 143) as a one-spored sporangium, since in all
the sections of conidia which I have examined there is no sign of a
second inner wall. It may be that the plasma-membrane of the plas-
molyzed contents of a conidium may have been mistaken for a wall;
or, again, it is possible that this author’s figs. 320, 321 represent con-
idia still surrounded by the slimy protoplasm which is sometimes dis-
charged from the ruptured basal cell.
I wish in conclusion to express my hearty thanks to Professor R.
A. Harper for the privileges afforded in his laboratories; to Professor
W. S. Marsua tt, for assistance in the determination of insects; and
to the Carnegie Institution of Washington, for a research assistancy
under which this work has been done.
SUMMARY.
1. Life history.—The life history of Empusa sciarae may be
summarized as follows: The disease attacks both larvae and adults
of the host, Sciara, causing ultimately their death. The young,
uninucleate germ-tubes, after they have entered the body-cavity of
the insect, grow there at the expense of the nutrient fluids. After the
protoplasm has increased in amount, a branching, coenocytic myce-
256 BOTANICAL GAZETTE [APRIL
lium is produced, which in early stages is few septate; later, how-
ever, at the culmination of vegetative activity, septa are abundant and
branching becomes more frequent.
Finally, the body-cavity becomes almost completely filled wii the
mycelial filaments, vegetative activity ceases and the death of ‘the
insect ensues with the beginning of the fructifying condition.
Radial branches, which mark the beginnings of the conidiophores,
are put forth from the short, 3-5-nucleate cells which make up the
mature mycelium; in this species, but one branch grows from each
cell. These radial hyphae bore their way out through the body-wall of
the insect; some form the rhizoids which attach the host to the sub-
stratum, while others grow into branched conidiophores. Each conidi-
ophere is cut up by cell-division into uninuclueate segments, each of
which pushes out beyond the surface of the host and cuts off from its
tip a single uninucleate conidium. The basal cell below the conidium
comes to possess but a thin, enucleate primordial utricle, and it finally
becomes greatly swollen from the absorption of water. Ultimately
this swollen basal vesicle bursts in a ring at the top where it joins
the conidial wall, or the columella-like wall may be split in some
instances, and the conidium is thus shot violently away, the slimy
protoplasmic contents of the lower cell being frequently carried along
with the conidium and serving to stick the latter to the substratum.
The partition which cuts off the conidium is at first curved upward by
the greater turgescence of the vesicle; but when the spore is shot off,
this reverses its former position, and in the conidium it appears as
a prominent papilla.
a. uclear division—The nuclei of Empusa are “centronu-
clei,” since the centrosomes which are active during division are
permanently intranuclear.
The division of the nuclei which takes place during the vegetative
stages appears to be of the nature of a primitive mitosis, similar in
many respects to that described for certain of the simpler Protozoa. .
The nuclear membrane generally persists during the whole process.
A simple intranuclear figure is formed, which in later stages consists
of the two opposed centers of division, to each of which converges from
all sides a system of fibrous radiations. The many radiations which
converge at the two poles correspond to the chromosomes; and,
1906] OLIVE—NUCLEAR AND CELL DIVISION OF EMPUSA 257
although they appear to be chromatic in their staining reactions, they
probably are made up principally of chromatin, and a small amount
of linin. The chromatin at first concentrates about the centrosomes,
which thus appear to have a darker rim about a lighter center. The
centripetal movement, as well as the later centrifugal movement
characteristic of the telophases, may be regarded as of the nature of
the flowing or diffusion of a liquid chromatin through a contractile
linin basis.
In the nuclear division of Empusa sciarae, the chromatin does
not appear to pass through an equatorial plate stage.
We may distinguish two shapes of the dividing nuclei in Empusa:
one found in short cells of the vegetative hyphae, in which the nuclei
in the later stages of division assume an oval or ellipsoidal shape;
and another found in long cells, in which the nuclei become them-
selves greatly elongated and early assume a constricted, hour-glass
shape. In the oval nuclei, the nuclear sap accumulates so that the
cavity becomes turgescent; while in the elongated nuclei, the liquid
does not accumulate, at least not to such an extent as in the first
instance, so that the consequent encroachment of the cytoplasm
between the two daughter-halves results in an early constriction.
In the long cells, cyclosis is doubtless stronger than in the short
_cells, thus bringing about in such instances a greater disturbance of
the mitotic processes.
3. Cell-division—Cell-division in Empusa sciarae is accom-
plished by means of the growth inward, from the wall of the mother-
cell, of a ring-formed partition. In a majority of cases, the new cell-
wall is carried across a wide, central, vacuolar space; when the older
cells become filled with cytoplasm, however, and later when the
conidium is abstricted, the wall cuts through the protoplasm which
fills the cell. A ring-formed cleavage-furrow starts at a definite
region of the plasma-membrane, and a wall is at once deposited in
the cleft. A region of some thickness at the inner margin of the
cleft, where the processes are most active which lead to the cleavage
of the in-pushing plasma-membrane and to the deposition of the
Partition wall, stains darker than the surrounding cytoplasm. This
fact is made the basis for the conclusion that the split plasma-mem-
brane is not the sole active agent of cell-division, although it may be
258 BOTANICAL GAZETTE [APRIL
a controlling factor in the process. Cell division, in this instance, is
regarded as a cytoplasmic phenomenon, since the nuclei may be
remote from the place of constriction; and, further, they appear to
have nothing directly to do with the process. The ultimate necessity
of the presence of a nucleus, probably as a controlling factor of nutri-
tion, is proven, however, by the early death of the enucleate basal
cells cut off from the conidia and from the end-cells of germ-tubes.
UNIVERSITY OF WISCONSIN,
Madison
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260 BOTANICAL GAZETTE [APRIL
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EXPLANATION OF PLATE XVI.
The drawings were made with the aid of an Abbe camera lucida, together
with the Zeiss 2™™ apochromatic obj. N. A. 1:30, combined with compensating
ocular 12; except fig. 16, which was drawn with compensating ocular 18.
Fics. 40-65, Empusa sciarae. All X 1500, except fig. 64, which is X2250-
Fic. 49. Probably an early prophase of division.
Fic. 50. A considerably advanced stage of division.
Fic. 51. A poorly differentiated preparation in which the karyolymph has
accumulated at one side of the dividing nucleus.
Fic. 52. A thin section of an anaphase.
Fic. 53. A similar preparation.
BOTANICAL GAZETTE, XLI
PLATE XVI
- eee SS > eh
: =* pt * we ~ Jes a
: na ° * 32am.
orn. f aa a
63 65 67
E. W. OLIVE, DEL,
HELIOTYPE CO., BOSTON.
OLIVE on EMPUSA.
1906] OLIVE—NUCLEAR AND CELL DIVISION OF EMPUSA 261
Fic. 54. A stage in which the daughter-halves appear to be pressed upon
by turgescent, vacuolar middle portion
55- A partly polar view of an aidiaidy placed nucleus, showing about
16 fibrillar radiations extending from the centrosomes.
1G. 56. A late condition in which the daughter-nuclei have just been sepa-
rated from each other by a cytoplasmic constriction. The nuclear contents
occupy a pseudo-synaptic position.
Fic. 57. Showing in the upper portion of the hypha a resting nucleus and
in pi gi a late stage of nuclear division
G. 58. A double cytoplasmic constriction cm taken place so that a vacu-
ole a with nuclear sap is left between the two daughter-nuclei.
Fic. 59. A telophase condition.
Fic. 60. Another telophase condition.
Fic. 61. Probably a very late telophase, in which the dark rim of the center
has survived, whereas the achromatic centrosome-portion has disappea
1G. 62. A late condition of nuclear division which is characteristic of ate
gated cells.
Fic. 63. A somewhat later stage, also from an elongated cell.
Fic. 64. A younger stage, in which the nuclear membrane has been
indented at the ends of the nucleus distally from the poles.
Fic. 65. An elongated nucleus in division, in which one pole shows an inden-
tation.
Fics. 66, 67, Empusa aphidis. Xt1500.
Fic. 66. A poorly stained mney showing the division character-
istic of the prec cells of this
Fic. 67. An earlier stage of ticato
BIOLOGICAL RELATIONS OF DESERT SHRUBS.
II. ABSORPTION OF WATER BY LEAVES.
V. M. SPALDING.
DurInc a study of certain shrubs growing in the vicinity of the
Desert Botanical Laboratory near Tucson, Arizona, it has been
found that the leaves of some of them absorb water, while those of
others do not. Although leaf absorption is treated by leading physi-
ologists as a matter of indifference, or at any rate of secondary
importance, it has seemed worth while to inquire whether differences
of habit in this particular, on the part of these desert plants, may
not be correlated with other characteristic peculiarities; if so, even if
the fact should turn out to be of small importance physiologically,
it may be significant from a biological point of view.
Our knowledge of leaf absorption as yet is fragmentary and
uncertain. For the general subject it is quite unnecessary to cite
the voluminous and contradictory literature. DANDENO’ has given
a useful historical résumé, reference to which and to paragraphs
in BURGERSTEIN’S more recent work? is sufficient for the present
purpose. In regard to various highly modified plants, however,
the case is quite different. ScHurmper has made such detailed
observations of certain epiphytes as to leave no doubt that they -
normally absorb large quantities of water through their aerial parts,
and that this is a distinct physiological advantage, or even neces-
sity. In view also of investigations cited by BURGENSTEIN it becomes
necessary to accept the fact of leaf absorption in the case of various
other plants.
As for the plants of arid regions, the evidence has been less con-
clusive than could be wished. VotkENs, in his classical work,
describes various special structures by means of which, presumably,
many of the plants of the Egyptian-Arabian desert take up dew
* DANDENO, J. B. An investigation into the effects of water and aqueous solutions
of some of the common inorganic substances on foliage leaves. Trans. Can. Inst.
75230. 1908.
2 BURGERSTEIN, A., Die Transpiration der Pflanzen. 1904.
Botanical Gazette, vol. 41] [262
—S
1906] SPALDING—ABSORPTION OF WATER BY LEAVES 263
through their leaves, but the woody species growing in the arid
territory of the southwestern United States are so different in their
habits and in their environmental relations as to preclude the settle-
ment of the question for them, even within the bounds of probabil-
ity, in any other way than by direct observation and experiment;
in fact they are found, as regards-leaf absorption, to differ widely
among themselves. The object of the present paper, therefore,
is the presentation of such facts as have been determined for a lim-
ited number of species indigenous to southern Arizona.
In this region existing physical conditions give to the question
special interest. As is well known, precipitation is meager, except
at relatively high altitudes, and is distributed throughout the year,
with no distinctively rainy season. The rainfall, moreover, is
extremely uncertain, and for months at a time is often so slight
that it does not wet the soil for more than a few centimeters, an
amount of precipitation likely to be of very little positive advan-
tage as far as root absorption is concerned. Under such circum-
stances, in which delicate adjustment is the condition of survival,
it would seem that plants capable of leaf absorption might have a
distinct advantage in times of prolonged drouth, during which occa-
sional showers occur which are too light to penetrate the soil. As
will be seen, however, only a limited number of species appear to
enjoy this advantage to an appreciable extent.
Nearly all of the species selected for investigation grow in the
immediate vicinity of the Desert Laboratory. A single one, Hola-
_cantha Emoryi, which seems not to be indigenous here, was obtained
from the grounds of the University of Arizona. The following
classification of the plants employed into biological groups is pro-
Visional, but will serve to direct attention to the very diverse eco-
logical history of the species now growing together in this region.
BIOLOGICAL CLASSIFICATION OF PLANTS STUDIED.
I. Shrubs, with relatively slight modification of form and struc-
ture, their habits plainly indicating mesophytic origin. Celtis,
Covillea, Lycium.
II. Shrubs or small trees, more conspicuously modified, but
retaining manifest traces of mesophytic habits. Parkinsonia, Pro-
sopis, Acacia.
264 BOTANICAL GAZETTE [APRIL
III. Woody or partly herbaceous plants, exhibiting peculiar
modifications of distinctly xerophytic types. Fouquieria, Hola-
cantha, Koerberlinia, Zizyphus, Atriplex.
IV. Plants of the most pronounced xerophytic character. Opun-
tia, Cereus, and other cacti.
V. Plants adapted by habit, rather than structure, to desert
conditions. Sphaeralcea and many other half-shrubby or more or
less herbaceous forms.
Of the species employed in the experiments, Celtis pallida is a
shrub, growing commonly to the height of one to one and one-half
meters on the laboratory hill, where it is rather abundant. It holds
its foliage so well that it might be ranked as an evergreen, though
it suffers to some extent from the effects of frost. Its leaves are
rough-hairy, thin but firm in texture, and conforming in general
to the generic type. Covillea tridentata, the well-known creosote
bush, is the most abundant woody species of this region. Its small
coriaceous leaves, presented more or less edgewise to the sun and
covered with waxy varnish, are well protected against excessive
transpiration. Lycium Berlandieri is a small shrub, more than a
meter in height, of frequent occurrence on rocky exposures. These
species, of the three genera named, while well adapted to their
habitat, exhibit characters far less conspicuously xerophytic than
those of many of the plants with which they are associated.
Coming to the second group, Parkinsonia Torreyana attains
the dimensions of a small tree, and is conspicuous by reason of its
green bark, from which it has the common name of palo verde.
Though a denizen of the desert, it is not a dry ground form, but
frequents low places, where more water is available than on the
mesa or even on the adobe soil of the hills, where Parkinsonia micro-
phylla, a related species, does well. Prosopis velutina, the mes-
quite, grows chiefly in low ground, within reach of abundant water,
but it also occurs, though scattering and undersized, on the adobe
soil of rocky hills. Like the palo verde and many other legumi-
nous plants, the leaves of the mesquite exhibit in their structure
and position excellent adaptations for the prevention of excessive
transpiration. Acacia constricta, of similar distribution, occurs
on the mesa and also on rocky upland. It is a vigorous shrub,
1906] SPALDING—ABSORPTION OF WATER BY LEAVES 265
one or two meters in height. These several species of Parkinsonia,
Prosopis, and Acacia thrive well under the rather severe conditions
to which they have become accustomed; they all retain, however,
manifest traces of mesophytic habits, particularly in their choice of
habitat.
The species assigned to the third group, among which are Fou-
quieria splendens, Holacantha Emoryi, Koerberlinia spinosa, and
Zizyphus lycioides, present more striking modifications of form
and structure than do any of the members of the preceding groups,
and, though differing greatly among themselves, agree in possessing
such conspicuous adaptations to xerophytic conditions as easily to
rank next to members of the following biological group.
The cacti are commonly taken to represent the extreme type
of xerophytes, but notwithstanding various striking features com-
mon to members of this order, there are essential differences of
habit and adaptation, even between closely related species, ren-
dering it quite impossible to generalize from the study of “typical
forms” in the investigation of biological problems presented by
them.
The half-shrubby and herbaceous plants are much like those
of other regions, exhibiting as a rule no structures that would be
thought of as distinctively xerophytic, but accommodating them-
selves to desert conditions by their habits, especially such as enable
them to take advantage of periods favorable for rapid development
and production of seeds.
By way of first ascertaining whether any of the plants of these
several groups absorb enough water through their leaves or inter-
nodes to be readily detected by weighing, the following method
was employed: A small branch with leaves functionally active,
though often showing plainly the effects of long drouth, was severed
and the cut end immediately covered with vaseline. In a few
instances, which are specified, branches without leaves were used.
The branch was then weighed and directly afterwards immersed
in water, except at the cut end, for a definite time, usually about
three hours. At the end of this period, after exposure to the open
air long enough to be certain that the surface was fully dry, the
branch was again weighed, and the increase of weight, if any, was
266 BOTANICAL GAZETTE [APRIL
taken to represent closely the amount of water absorbed, though,
owing to loss during the operation of drying the surface, the amount
absorbed must often have been rather greater than the increase
of weight indicated. In the first preliminary set of observations,
a pair of large balances, weighing satisfactorily to ten milligrams,
was employed; but in subsequent experiments quantitative bal-
ances were used, the weighing being made to a milligram in each
case. Changes during the process of weighing rendered it as useless
as it was unnecessary to attempt a higher degree of accuracy.
Inspection of Table I shows that leafy shoots of Celtis, Covil-
lea, and Lycium, by immersion in water for three hours, gained
1.9 to 5 per cent. of their original weight; Atriplex in a little longer
TABLE I.
PRELIMINARY TEST OF CAPACITY FOR ABSORPTION. November 1904.
Species Date 7 Time baton ig = esa
Covillea tridentata (1)......... Nov. 1 10:12 A, M. 18.540
; : 1:12 P.M, 19.250 | 3.8 gain
Covillea tridentata (2) ........ i 9 10:55 A.M. 8.235
: : 1:56 P.M. So6ca ) fossa 3s
RMN IME NOM ss 0g 9 ld san wc 5 I 10:57 A.M. 24.920
: <a 1357 P.M. 26.060 .| 4.9 *
Lycium Berlandieri........... ee I 10:45 A.M. 7.105
: ; 1:45 P.M. e240 | ig ~
Peaeia COnstACta. so s.5 of < 7 hs I 11:14 A.M. 6.260
; ‘ I:14 P.M., 6.463 | £25 *
pruayes Velen. oo ah ae bs ‘ I 10:30 A.M. 4.200
; a 1:30 P.M. £6 ot 8 7 F
Parkinsonia microphylla....... a I 10:22 A.M. 10.410
: E 1:22 P.M. 10.490 | 0.8 “
Parkinsonia Torreyana........ eae” 11:03 A.M. 4.070
: 2202 P.M. 4.085 eee
Atriplex canescens............ ne SES 10:21 A.M. 2.140
, re 1:44 P.M. g0ep.: [2 eis
Zizyphus lycioides............. v4 9 11:39 A.M. 19.270
ins 3:25 P.M. 19.370°]:.0.3: *
Fouquieria splendens.......... Ss 9 11:21 A.M. 21.776
2:24 P.M. 21.920 | 0.7 “
ea tet 3:32 P.M. 21.770
Koerberlinia spinosa........... Sade: >. 10:41 A.M. 8.650
’ ; 1:40 P. M. 8.670 | 0.2 “
Encelia farinosa..... 2.05556... < 9 12:00 M 5.010
3:00 P.M Go PaaS
3:18 P.M 5-000
Sphaeralcea pedata........... meer ae 12:07 P.M 1.120
3:12° P.M i.e besa
3-21 P.M 1.110
2 sinscmmoneas ini
1906] SPALDING—ABSORPTION OF WATER BY LEAVES 267
period gained 5.1 per cent.; Parkinsonia, Prosopis, and Acacia
gained 0.4 to 1.5 per cent.; iil Zizyphus, Fouquieria, and Koer-
berlinia, all without leaves, showed almost no appreciable gain.
Species of Sphaeralcea and Encelia gained in weight 5.4 and 6.2
per cent. respectively, but promptly lost all they had gained by a
few minutes drying.
It will be noticed that of the plants employed in this prelimi-
nary work those without leaves absorbed no water to speak of,
while those in leaf fell into two categories, those absorbing and
those not absorbing water in quantity. The experiment, there-
fore, pointed to leaves rather than internodes as agents of absorption,
and indicated, apart from Sphaeralcea, Encelia, and the peculiar
Atriplex, only the woody species belonging to the first group as
likely to prove capable of absorbing much water.
Starting with the suggestions derived from these facts, a more
careful and detailed study was undertaken. Cut shoots were still
employed for a time, though it was understood that confirmation
of results would necessitate the use of entire plants, and these were,
as a matter of fact, employed to a large extent in the later work.
Care was exercised in the selection of material, and in each case
its source and any conditions liable to affect results were noted.
GROUP I.
Celtis pallida.
Four specimens of this species were selected, all in good condi-
tion, though apparently not as active physiologically as they would
have been earlier in the year. Numbers 1 and 2 were fresh shoots,
while numbers 3 and 4 were small branches taken from older bushes.
Those numbered 1 and 3 were cut so as to include a large leaf sur-
face as compared with the other two. In every case the cut ends
were covered at once with vaseline, and the first weighing was made
as soon as practicable after bringing them to the laboratory. They
were then wet at frequent intervals for a little more than three hours,
and; after drying the surface, were weighed again, after which
they were left in the laboratory to dry until the next day, when the
same steps were repeated. Finally they were immersed in water
over night and again weighed.
268 BOTANICAL GAZETTE [APRIL
TABLE II.
CELTIS PALLIDA. December 1904.
No.| Date Time bo nvcmcig ges Period of treatment ~
1 | Dec. 19 | 10:44 A.M. | 1.834
£255 P.M. | 1.853 | 2 gain After wetting mt 3 iy Ir min.
20 | 9:48 A.M. | 1.516 [18.2 loss ng tg hrs. 53 m
2250 P.M: | 1.997 )| 5 - gain “ wetting nearly Ma hrs. 22 min.
ar || 0535 A.M. | ¥.895 (7 3 ca 25
2 Ig | 10:54 A.M. | 1.392
2:02 P.M. | 1.410 | 1.3 gain After wetting nearly 3 hrs. - min.
20 | 9:57 A.M. | r.192.|15.5 loss drying t9 hrs. 55 mi
2:25 P.M. | 1.238 | 3.9 gain oh wetting nearly 4 hrs. 28 min.
21 | 10:52 A.M. | 1.440 |16.3 gain : 27
3 ¥ | 12213 A.M. | 2.137
2:20 P.M. | 2.185 | 2.2 gain After wetting nearly 3 hrs. A min.
20 | 0:07 A.M. | 1.734 |20.6 loss 1g hrs. 47 m
2:38 P.M. | 1.870 | 7.8 gain * wetting seed 7 hrs. - ‘min.
at | 11:05 A.M. | 2.182 |16.7 “ rs 7
4 IQ | 11:25 A.M. | 1.505 -
2°30 -P.M..| ¥.5§21-| 4.1 gain After pata ad 4 ma 6 min.
20 | 10:16 A.M. | 1.351 |11.2 loss Ts. 45 m
2:46 P.M. | 1.387 | 2.7 gain 3 wetting ‘nearly = hrs. 30 min.
SE.) 067564 Me. | oeaS ese - 29
Inspection of Table IE shows:
1. That all four specimens absorbed water very slowly just
after they were freshly cut, and that the rate of absorption was
greatly increased after they had lost weight by remaining over night
in the dry air of the laboratory.
2. The rate of absorption showed a correlation with extent of
leaf surface, being considerably greater in the two specimens with
large extent of leaf surface than in the other two.
3. The weight lost by drying for a given period was nearly or
quite regained when the leaves were given a full supply of water
for a corresponding length of time. The capacity of this species
for leaf absorption, under the conditions described, is thus fully
demonstrated. Its deportment in the seedling stage, which offered
for experiment perfectly fresh and unmutilated material, will next
be considered.
Seedlings of Celtis pallida were grown from seeds sown Novem-
ber 14, 1904. When used for experiment in January and February,
1906] SPALDING—ABSORPTION OF WATER BY LEAVES 269
1905, they were all in healthy condition, and when taken up were
found to have fine vigorous roots. In addition to the cotyledons,
which were still capable of photosynthesis, each seedling had two
or three perfectly healthy green leaves that had attained the length
of about one centimeter. The seedlings were transplanted into
earth contained in glass vials of convenient size for accurate weigh-
ing, sheet rubber being used to prevent evaporation from the soil.
In the case of seedling number 1 the earth was very moist when
the rubber was adjusted, and it was found that this plant, which was
transpiring vigorously, showed almost no capacity for absorption.
The case was different with seedling number 2, which was left some
five days after transplanting with the soil open to the air, so that
it became relatively dry before the rubber was adjusted. The coty-
ledons of number 1 were removed, their place of attachment being
carefully covered with vaseline; number 2 had one large cotyledon
which remained in place during the experiment. These details are
necessary to an understanding of the different behavior of the two
seedlings as shown by Tables III and IV, which cover the period
from January 21 to February 1, at which latter date the experiment
_ Was concluded.
It is seen that both seedlings transpired regularly and largely,
but that number 1, in spite of the fact that its transpiring surface
had been lessened by the loss of its cotyledons, exhibited a decidedly
higher rate of transpiration than number 2, which was in drier
TABLE Iii.
CELTIS PALLIDA. SEEDLING No.1. January 1905.
Date Time Weight in! 7 o¢s or gain Conditions
grams
Jan. 21 | 1:19 P.M. | 26.256 seit ;
4:05 26.229 | 0.027loss_ | After standing in dry air
23 | 10:15 A.M. | 26. PoE 5, ed Ps fe yet, Mer,
2:50 P.M. | 26.061 | 0.033 “ ~ Pw ee
25 10:50 A. M. 25.936 0.125 ce “ce “ce “ec oe “cc
26 9.50 2 ;.870 0.066 “c ee &e tS te ee
27 9:28 25.806 0.064 “ “ “ (2 ied | eae
2:53 P.M. | 25.810 | 0.004 gain After wetting i
3:41 25.803 | 0.007 loss standing i in dry air
28 | 11:02 A.M. | 25.762 | 0.041 “ x gulp eae ba
13:33 P.M. | 25.764 | 0.002 gain ns ng f
2:58 25.759 | 0.005 loss ie) cia divaie
270 BOTANICAL GAZETTE [APRIL
TABLE IV.
CELTIS PALLIDA. SEEDLING No. 2. January and February 1905.
Date Time ging Loss or gain Conditions
Jan. 30 | 10:09 A 22.638
1:10 P.M. | 22.630 | 0.008 loss After standing in dry air
3:40 22.626 0.004 “< “ce ““ “ «@
30)" O245 A: Me |) 22659 | 0.027 gain “wetting
12:45 P.M. | 22.628 | 0.025 loss ‘** standing in dry air
gi 22.634 | 0.006 gain “wetting
Feb. 1 | 9:48 A.M. | 22.592 | 0.042 loss ast ng in dry air
9:50 0.027 Weight of plant above ground
2:08 P.M. | 0.045 | o.o28 gain | After wetting -
soil, an interesting result in harmony with earlier experiments,
showing the direct relation between available soil water and rate
of transpiration.3
On the other hand, while the quantity of water absorbed by:
number I was so meager as to be negligible, that absorbed by num-
ber 2 was much more, in one case almost exactly 100 per cent. of
its own weight, i. e., of the part above ground when it was after-
wards severed from the root. Number 2, although apparently
perfectly healthy while the work was in progress, seems neverthe-
less to have reached a condition in which the diminished supply
of water from the soil was followed by a marked acceleration of
leaf absorption, while in the case of number 1, growing as it was
in moist soil, no such compensation was made or required.
Of interest as bearing on the validity of determinations of absorp-
tion by the use of detached shoots is the fact that while seedling
number 2, after it had finally been cut off at the surface of the ground,
absorbed in a few hours its own weight of water, it had done pre-
cisely the same thing before mutilation, only in longer time. It
may well be that a detached shoot, cut off from its normal source
of water supply, will absorb more rapidly through its leaves than
the same shoot, which, while attached, is supplied, even inade-
quately, from the soil; but this difference plainly does not justify
the degree of discredit that has been thrown upon evidence derived
from experiments with separated parts of plants.
Three other seedlings of Celtis pallida were treated like the
preceding ones, except that the observations were not begun until
3 SPALDING, V. M., Soil water in relation to transpiration. Torreya 5:25. 1905-
1906] SPALDING—ABSORPTION OF WATER BY LEAVES 270
the plants had been some weeks in the vials to which they were
transplanted, and weighings were made during a longer period and
with more complete records as to soil conditions, health of seed-
ling, etc. Their records as to weight are given in Tables V, VI,
and VII. All of these seedlings were in a healthy condition and
apparently capable of entirely normal development. The small
extent of surface through which absorption and transpiration took
place renders the consistency of the results all the more striking.
In every case water was absorbed when it was presented to the
leaves and internodes, and transpiration was resumed as soon as
their surfaces were dried.
TABLE V.
CELTIS PALLIDA. SEEDLING No. 3. February 1905.
Date Time Weight in Loss or gain Conditions
grams
Feb. 14 | 11:20 A.M. | 23.084 Weight of vi a and outfit
2:25 P.M. | 23.091 | 0.007 gain After i immersion raped water
15 -M. | 23.044 | 0.047 loss standin aii dry ai
T1755 23-053 | 0.009 gain 5 frametsioa! in rain qa
16 | 11:42 A.M. | 23.007 | 0.046 loss “standing in dry air
17 | 2:11 P.M. | 23.046 | 0.039 gain immersion in rain water
20 | 10:23 A.M. | 22.895 | 0.151 loss ‘standing in dry air
2:51 P.M. | 22.916 | 0.021 gain n immersion in rain water
21 | 10:28 A.M. | 22.0932 s0x6., rs
12:00 M. 22.907 | 0.025 loss 8 standing in dry air
3:02: Pi M. | 22.887 | 0.020" © se ae
22 {| 12:08 22.845 | @.052 * - - :
T2:145 22.828 Weight oe — cotyledon
act 22.8 0.007 gain | After immersing in rain water
a? a O. oe Weight ar cut is pares of earth
2228 22.799 4 , rubber, and ea i
25 1:58 0.015 * of plant above ground, air-
dried in laboratory
When the first weighing was made, February 14, seedling no. 3
had two cotyledons, still attached, and three foliage leaves. The
cotyledons showed some indications of drying. The earth in the
glass vial in which the seedling was growing was becoming rather
dry, but still contained sufficient water to maintain a transpiration
current for a week and probably longer. On February 16, the
note was made “one of the cotyledons drying, curled, and getting
stiff; the other paler than the foliage leaves, but still flexible, other-
wise the seedling is in good condition.”
On February 22 the cotyledons were removed and the subse-
ae |
{
272 BOTANICAL GAZETTE [APRIL
quent deportment of the seedling indicates that their failing condi-
tion previous to removal may be disregarded as not materially
affecting the results. As the table shows, the weight of the whole
plant above ground, including cotyledons, was less than the weight F
of water transpired in 19 hours (Feb. 14-15) and also less than the
gain of weight by absorption in 26 hours (Feb. 16-17), a conclusive
proof of the relatively large quantities of water absorbed and trans-
pired by this seedling during the period of experimentation. The
facts regarding seedlings 4 and 5 are so fully set forth in Tables VI
and VII as to render further explanation unnecessary. |
TABLE VI. |
CELTIS PALLIDA. SEEDLING No.4. February and March 1905.
Date Time braces in| Loss or gain Conditions
Feb. 14 | 3:30 P.M. | 20.130 Weight of plant and outfit
I5 | 10:02 A.M. | 20.090 | 0.040 loss After spent in dry air
12:47 P.M. | 20.093 | 0.003 gain immersion in rain water
16 | 12:00 M. 20.031 | 0.062 loss by saul in n dry air
17 | 3:11 P.M. | 20.051 | 0.020 gain ‘* immersion in rain water
20 | 10:46 A.M. | 19.821 | 0.230 loss «standing in
3:19 P.M. | 19.825 | 0.004 gain eC Sats mersion in rain water
21 | 10:54 A.M. | 19.830 | 0.005 “ 4
12:11 P.M. | 19.813 | 0.017 loss = standing i in dry air
22° tasdo 19.696 | 0.117 “ os
3:08 19.707 | 0.011 gain a mersion in rain water
March2 | 10:04 A.M | 19.080 | 0.627 loss is saad in dry air
2:04 P.M. | 19.085 | 0.005 gain a a in rain water
2225 19.080 | 0.005 loss “* standing in
2:30 0.070 Weight rakes cut Biss at surface of
rth
TABLE VII.
CELTIS PALLIDA. SEEDLING No.5. February 1905.
Date Time igo Loss or gain Conditions
Feb. 20 | 11:16 A.M. | 20.207 Weight of plant and outfit
3:28 P.M. | 20.204 | 0.003loss | After wetting with rain water (not
immersin:
2I | If:og A.M. | 20.219 | 0.015 gain | After immersion in rain water
12:15 P.M. | 20.214 | 0.005 loss af standing in dry air ~
22 12:34 20.172 0.042 oe ce té- Gt ce
3:18 20.178 | 0.006 gain in rain wate
3*33 0.025 Weight mye inky ean off at mates of
2G.) @txe 0.012 Weight after air drying
1906] SPALDING—ABSORPTION OF WATER BY LEAVES 273
Covillea tridentata.
The specimens of creosote bush selected for experiment were
taken from four different sources, for the sake of securing material
as different as practicable in regard to the amount of water in the
tissues. Number 1 was from a bush growing near an irrigating
ditch, where it had been abundantly supplied with water. Its
leaves were large, dark green, and fresh, and numerous flower buds
had been formed. Number 2 was from a plant growing on the
mesa a few rods distant. Its leaves were smaller and lighter green,
and in comparison with number 1 it was plainly a dry ground form,
though it did not have the appearance of having suffered to any
great extent from lack of water. Numbers 3 and 4 were from plants
growing on the mesa, near the foot of the laboratory hill, where
in a dry time the Covillea, the only shrub that keeps alive there,
shows the effects of drouth very badly. Their leaves were still
smaller and paler in color, those of number 4 especially, indicating by
their minute size and other peculiarities a plant that had long lacked
a sufficient supply of water. The contrast between this and the
first member of the series was very striking. It should be stated,
however, that none of the specimens were in quite so dried-up a
condition as those employed early in November before the Decem-
ber rains, which though meager—o.82 inch (21™™) thus far—
had freshened vegetation to some extent. The dried-up leaves
that were dying in November had been shed, and the leaves remain-
ing on the bushes when the experiment was conducted, late in Decem-
ber, were apparently in a vitally active condition.
It will be noticed by reference to Table VIII that, precisely as
in the case of Celtis, all the specimens of Covillea gained very little
in weight as the result of wetting soon after they were cut. Num-
ber 1, from the irrigated bush, gained least, and number 4, from
the dry ground plant, gained most. After prolonged drying and
again wetting, the gain was much greater than before, the greater
gain in each case being made by number 4, which, as already stated,
was from the most distinctively dry ground form.
The deportment of number 1, from the robust, well-watered
bush, is instructive, especially as it may throw light on the question
as to whether leaf absorption is a normal process that takes place
274 BOTANICAL GAZETTE [APRIL
TABLE VIII.
COVILLEA TRIDENTATA. December 1904.
No.| Date Time bears ie eo Samy Period of treatment
1 | Dec. 26 | 11:05 A.M. | 2.529
2217 P:M. |2.5383 | 6,4 gain After wetting ay 3 na 12 min.
28 | 10:32 A.M. | 1.957 {22.9 loss drying 4
2:18 P.M. | 1.996 | 2.0 gai = wetting nearly 3 hrs. e min.
29|| IorSo A.M. | 2.388 | 9.6 “ es 3
2 26 | 11:12 A.M. | 2.434 :
2:35 P.M. | 2.453 | 0.8 gain | After wetting nearly 3 hrs. 23 min.
28 | 10:40 A.M. | 2.234 | 8.9 loss ** drying 44 hrs. 05 min. :
2733 PM. | 2.258.| 1.1 gain “wetting nearly 3 hrs. 53 min.
20: | FR203 Ai M. | 2-416-| F.0 “* if oo 2. ga *
3 26 | 11:21 A.M. | 2.261
2:58 P.M. | 2.281 | 0.8 gain After wetting nearly MA hrs. 37 min.
28 | 10:48 A.M. | 1.980 [13.2 loss drying 43 hrs min.
2:45 P.M. | 2.020 | 2.0 gain sa wetting nearly 3 “hrs. : min.
29 | i1:14 A.M. | 2.225 a es ©
4 26 | 11:28 A.M. | 2.646
3:03 P.M. | 2.688 | 1.6 gain After wetting nearly 3 hrs. 35 min.
28 | 10:55 A.M. | 2.357 |r2.3 loss drying 43 hrs. 52 min.
2:56 P.M. | 2.415 | 2.5 gain ‘“* wetting nearly 4 hrs. 1 min.
2Q | 11:23 A.M. | 2.766 |14.5 “ is ee = ae ae
under natural conditions. This shoot, with its large, fresh, turgid
leaves, lost water by drying approximately twice as rapidly as did _
those from dry ground, with their much smaller leaves and firmer
tissues, and on subsequent wetting absorbed far less than the
latter in proportion to previous loss. Unlike these, moreover, the
leaves of the first specimen, in the course of alternate drying and
wetting, lost their fresh look and became discolored. The impres-
sion was received that this specimen, taken from a perfectly fresh
plant and requiring no additional supply of water, suffered patho-
logical changes in the course of the treatment to which it was sub-
jected, while the others, coming from dry ground plants in need
of water, absorbed it as by a perfectly normal process. Even these,
however, were not in a condition for rapid leaf absorption when
first cut, their gain per cent. being decidedly less for a given period
than that exhibited by individuals of the same species during
observations made before the December rains. In brief, the experi-
ments of December 26-29, in connection with those of November
1-9, indicate on the part of the creosote bush marked capacity for
1906] SPALDING—ABSORPTION OF WATER BY LEAVES 275
subaerial absorption after protracted drouth, but more limited
capacity for such absorption, even if artificial drying is resorted
to, when it is receiving a better supply of water.
Lycium Berlandieri.
At the time when the observations on Lycium were made, late
in December, most of the summer leaves had fallen and fresh ones,
following recent light rains, were only beginning to appear. Con-
sequently it was difficult to secure entirely satisfactory material,
but a few specimens were finally obtained for experiment which
were in a normal and active condition. The leaves of this species
are small, only about one centimeter in length, but otherwise the
plant gives the impression, as already stated, of having retained
up to the. present time distinctively mesophytic tendencies. The
rapidity with which the leaves were transpiring was at once obvious
when weighing was undertaken, and, as in cases previously cited,
absorption was found to take place extremely slowly while the leaves
were still fresh. Even after drying, water was absorbed in no case
as rapidly as it had been lost. Thus number 2 lost 3 per cent. of
its weight by drying three and one-half hours, and gained after-
wards by wetting nearly four hours 1.9 per cent. Of the actual
capacity of this species for subaerial absorption the experiments
leave no room for doubt; but the.specimens employed deported
themselves much like the well-watered Covillea, except that no
suggestion of pathological change in the course of the treatment
to which they were subjected was noted. When gathered they
were simply in the condition of fresh, actively transpiring plants,
TABLE IX.
Lycrum BERLANDIERI. December 1904.
No.| Date Time Iyer ge sl i Period of treatment
1 | Dec. 28 | 10:15 A.M. | 3.065
1:47 P.M. | 3.076 | 0.4 gain After ee aes? nearly 3 3 32 min.
AQ: NOTE E Ac M+[ 32235. |-5 22” paris 20 hrs. 28 m
1:42 P.M. | 2.956 | 8.7loss | ‘“‘ drying 3 hrs ‘
3:25 3-002 | 1.6gain; “ wetting cue I ae 43 min.
2 28:).107 92 A: Mi. | si ¥a9
E°G0 POM. |-3.028 | 370 loss After drying 3 hrs. * min.
29: | 10:00 A,M. | 2.602 |. 7-5 °
1:53 P.M. | 2.855 | 1.9 gain “wetting nearly : re 53 min.
276 BOTANICAL GAZETTE [APRIL
which apparently could: derive no advantage from an additional
supply of water presented to their leaves.
The record of these three species of Celtis, Covillea, and Lycium
has been given at length, on account of the importance of estab-
lishing beyond doubt the fact that in these plants, which have been
taken to represent desert species that retain in structure and habits
obvious indications of mesophytic origin, leaf absorption certainly
takes place, and apparently as an entirely normal process. We
have next to deal with a group of species genetically related, which
deport themselves quite differently from members of the first bio-
logical group in regard to leaf absorption. As representatives of
this second group, species of Parkinsonia, Prosopis, and Acacia were
selected, all belonging to the Leguminosae. The record of experi-
ments and their results is such as to admit of statement in few words.
GROUP II.
Parkinsonia Torreyana.
The specimens of palo verde employed in this work were seed-
lings some two months old. One was cut about thirteen hours,
the other (number 2) an hour and a half before weighing. After
weighing an attempt was made to wet the leaves by repeatedly
immersing the seedlings in water. The experiment might fairly
have been dropped at this point, since, as it was found impossible
to wet them, leaf absorption could hardly be thought of; but as
there remained a possibility of some slight absorption where drops
of water collected on the surface of the youngest parts, the attempt
was continued with number 1, which was repeatedly immersed
during a period of something over three hours.
As seen from Table X this seedling, so far from gaining by absorp-
tion of water presented to it, actually lost 1.8 per cent. of its weight
in three hours and thirteen minutes, its surface having remained
almost entirely unwetted, so that loss of water was possible during
the whole, or nearly the whole, of this period. Seedling number 2
was allowed to dry, after an unsuccessful series of attempts to wet
its surface. Its loss of weight, as might be expected, was greater
than that of number 1.
If these results are compared with those of November 1 and 9,
derived from similar experiments with shoots of Parkinsonia micro-
1906] SPALDING—ABSORPTION OF WATER BY LEAVES 277
ABLE X.
PARKINSONIA TORREYANA. SEEDLINGS. December 1904.
. Weight in|Loss or gain Boa
No.| Date | Time | erin per cent. Conditions
t | Dee. 31 | -g232 AM. | 0.901
12:45 P.M. | 0.944 | 1.8 loss | After repeatedly immersing in water
2 9.37 A.M. | 0.886 gtd drying, aera! be gees at-
5 tempts to i the s
12°52 P.M. | Ovd55 | 3.5408
phylla and P. Torreyana, the conclusion must be drawn that the
species of Parkinsonia represented here either absorb no water, or at
most an exceedingly small quantity, through their leaves. Before
making the experimental test it was thought that the fresh, rapidly
transpiring leaves of seedlings might exhibit a capacity for absorption
not shown by those of older plants, but this has not proven to be the case.
Prosopis velutina.
Work on the mesquite was carried on at intervals for a number
of weeks in January, February, and March, the material first em-
ployed being obtained from mature specimens, while in the later
experiments seedlings were used. Of specimens taken from mature
plants only the leaves were immersed in water. In some cases
the upper surface resisted wetting, while in others both surfaces
were easily wetted. This was followed, as indicated by some increase
of weight, by absorption of water in limited quantities. The seed-
lings which were employed in subsequent experiments remained
unwetted in all cases when they were immersed in water, and in spite
of the fact that two of the specimens had been left to dry as much as
forty-two hours and showed the effects of this treatment before im-
mersion, there is no evidence that they absorbed any water whatever.
It is apparent, then, that as long as the leaves of the mesquite
are perfect and resist wetting they absorb no water, even after dry-
ing for some time, but that they may absorb more or less after they
have become old and can be wetted. It is very questionable, to
say the least, whether in the latter case this process has any physi-
ological significance. It would seem that in the mesquite, as in
the palo verde, adaptations to xerophytic conditions have been
carried so far in the direction of preventing excessive transpiration
that leaf absorption, as a normal process, does not take place.
278 BOTANICAL GAZETTE [APRIL
Acacia constricta.
A series of experiments with this species was carried out, but
it was found unfavorable for exact results, owing in part to the
fact that its leaflets become tightly closed after wetting, thus rend-
dering it difficult to secure perfect drying of the surface without
overexposure and consequent uncertainty as to the true weight.
Accordingly, the conviction that the data obtained were unreliable
led to their rejection. For this second group, therefore, we are
restricted to the positive results obtained from Parkinsonia and
Prosopis, which exhibit either no capacity or very slight capacity
for leaf absorption, so long as the leaves are in perfect condition
and normally active.
. GROUP III.
This third group includes representatives of a number of genera
much modified in form and structure, and differing among them-
selves in their methods of meeting desert conditions. Several of
these are more commonly seen without ‘than with leaves, photo-
synthesis then taking place in their green shoots; while others, more
dependent on leaf activity, are commonly in a leafless condition
during a large part of the year, pushing out new leaves promptly
when conditions are favorable, and dropping them again when they
become adverse, .as is seen particularly in the case of Fouquieria.
Holacantha Emoryi.
Of this peculiar shrub a small branch with leaves was. cut and
left several hours to dry. At the end of this time it was still fresh,
with no indication of wilting. After weighing it was wet for two
hours and thirty-nine minutes, after which it was weighed again, the
weight remaining unchanged. Leaving the shoot now to dry until
the next day, and then wetting it for four hours and twelve minutes,
' there was a gain in weight of only o.5 percent. Part of the same
shoot, destitute of leaves, was treated in the same way, and, after
wetting four hours and eleven minutes, also showed a gain of barely
©.5 per cent. of its former weight. These results indicate on the
part of this species capacity for leaf absorption so inconsiderable
that it may be neglected.
Koerberlinia spinosa, a closely related species, agrees with Hola-
‘cantha as far as observations have yet been made. Only leafless
1906] SPALDING—ABSORPTION OF WATER BY LEAVES 279
branches have been obtainable, but these, as in the preceding species,
are green, and for a large part of the year the plant has no other
organ of photosynthesis. So far, then, as present evidence goes,
absorption through leaves or internodes is not to be predicated of
either of these plants.
Zizyphus lycioides.
After the preliminary experiments already recorded, very little
satisfactory material for the study of this species was obtainable,
as the plant cast its leaves and remained bare until after the period
of study was concluded. From observations made early in the
year, however, it appears that leafless shoots of Zizyphus do not
absorb water in appreciable quantity, but that leafy shoots have
considerable absorptive capacity, indicating that it is the leaves
and not the internodes through which absorption takes place.
Fouquieria splendens.
Leafy shoots of the ocotillo, as shown by Table XI, absorb con-
siderable water when wet for some time after drying. As in various
other cases, the loss of weight on drying the shoots after wetting
is considerably more rapid than the preceding gain by absorption.
TABLE XI.
FOUQUIERIA SPLENDENS. January I905.
No.| Date Time Weight in/I per pl as Conditions
I} Jan. 26 | 12:00 M. 1.614 In each case loss followed drying and
gain followed wetting the sp-eci
mens during the periods indicated
in the time column.
27 | 10:00 A.M. | 1.582 | 1.9 loss
28 | 11:40 1.664 | 5.2 gain
3:20 P.M. | 1.598 | 4.0 loss
3° | zio2 Ta§I6. 500"
3:28 1.530 | 0.8 gain
4:09 1.523 | 0.5 loss
31 | 4:15 1.490
Reb. = £532 1.585
2 | Jan. 26 2:53 POM. 4s2t4
27 | to:1§ A.M. | 4.112 | 2.4 loss
28 | 11:45 4.059 | 1.3 loss
30 | 2:04 P.M. | 4.363 | 7.5 gain
3:04 4-255 2.5 loss
4:02 4.197 | 1.4
Biol tae ACM: | S461, rea: 77
2:45 P.M. | 3.840 | 2.1 gain
3245 3-798 | 1.1 loss
280 BOTANICAL GAZETTE [APRIL
It appears, then, that for this biological group, which includes
a number of plants for the most part unrelated systematically, no
general rule can be formulated regarding leaf absorption. The
experiments go: to show that Holacantha and Koerberlinia hardly
absorb at all, while Fouquieria is certainly capable of absorbing
considerable quantities of water.
GROUP IV.
The only representatives of the cacti that have been studied
thus far are two species of Opuntia, O. Engelmanni and O. ver-
sicolor, A number of specimens of each species were selected after
a prolonged drouth, the results of which were plainly seen in their
much shrunken condition, very favorable, it would seem, for the
demonstration of absorption if this ever takes place. As the mate-
rial was rather bulky the large balances were employed, a terminal
joint in each case being severed and weighed; but there is no reason
to doubt the substantial accuracy of the results.
As shown by Tables XII and XIII, Opuntia Engelmanni did
not in any case gain more than 0.6 per cent. of its original weight,
even when immersed in water upwards of 45 hours, and it is safe
to conclude from this result, drawn from experiments with a num-
ber of specimens, that the species in question does not normally
absorb any considerable quantity of water in this way. Opuntia
versicolor, on the other hand, treated in the same manner, showed
TABLE XII.
OPUNTIA ENGELMANNI. December 1904.
No.| Date Time Weight in| pe st ocieed Conditions
1 | Dec. 3 | 10:15 A.M. | 82.360 In each case gain followed wetting
ad loss followed drying rs, =
oan indicated a the
1:30 P.M. | 82.850] 0.6 gain
§.| 18:00 A.M. | 83: 360] 0.6
2 6 | 10:12 A.M. | 72.085
I:10 P.M. | 72.470] 0.5 gain
3:05 72.080] 0.5 loss
3 10:39 A.M. |131.750
1:33 P.M. |132.290) 0.4 gain
3:20 131.760] 0.4 loss
ra
. 1906] SPALDING—ABSORPTION OF WATER BY LEAVES 281
TABLE XIII.
OPUNTIA VERSICOLOR. December 1904.
. | Weight in| Loss or gain’ de
No.| Date Time Pisano per cent. Conditions
1 | Dec. 3 | 10:30 A.M. 116.390
1355 P.M. |16.770 | 2.3 gain | See Table XII
f | FE Ae. [16.580 7|o21 7?
2 6 | 9:55 A.M. |10.640
12:14 P.M. /10.850 | 2.0 gain
3:02 10:630 | 2.0 loss
3 10:30 A.M. | 8.115
12:35 P.M. | 8.360 | 3.0 gain
3:10 8.115 | 2.9 loss
a gain of 2 to 3 percent. The specimens most shrunken with drouth
were found to absorb water most rapidly.
The rapid loss of water and the curiously close correspondence
in each case between the percentages of gain and loss, suggest that
in this species it is merely the tubercles that act as organs of absorp-
tion, and notwithstanding the fact that the water absorbed is so
promptly given off in a dry atmosphere, it appears probable that
in a period of frequent light rains the continued absorption of water
by the tubercles is precisely the necessary preparation for the develop-
ment of the young shoot which presently follows. Meantime the
different deportment of these two species of Opuntia as regards
amounts of water absorbed, corresponding as it does with a marked
difference in size of their tubercles, suggests the desirability of a
more extended comparison of these structures in different cacti
with reference to their marcas for absorption and the physiological
value of the process.
GROUP V.
A discussion of the annuals and partly herbaceous perennials
that have been referred to a fifth biological group, many of which,
structurally at least, are not to be thought of as characteristic desert
plants, does not fall within the limits of the present study. As
already stated, as many of them as have been studied agree in
promptly absorbing water when it is presented to their leaves and
internodes, which, however, is given off so rapidly in dry air that
it hardly seems possible that its absorption is of any utility. Cy.
Table I, Encelia and Sphaeralcea.
282 BOTANICAL GAZETTE [APRIL
SUMMARY.
From the preceding observations and experiments, in which
woody plants were chiefly employed, it has been seen that certain
species of desert plants of southern Arizona absorb water presented
to their leaves and internodes, while others do not. The species
represented in the vicinity of the Desert Botanical Laboratory may
be divided into several biological groups, based primarily on the
water relation, of which leaf absorption is a phase. Thus, in the
first group, including shrubs, which retain well marked mesophytic
tendencies, leaf absorption is characteristic. Members of the
second group, more distinctively xerophytic in various structural
particulars, are incapable of leaf absorption during their period
of normal activity. The third group, decidedly xerophytic, but
including species of widely different structure and habits, exhibits
corresponding differences in regard to subaerial absorption, which
takes place in some of its representatives and not in others. The
fourth group, including cacti which are assumed to represent the
extreme type of xerophytes, also exhibits interesting differences
in size and structure of the tubercles by means of which water is
absorbed. Finally, members of a provisional fifth group, which
in habit and structure are nearer than any other others to the meso-
phytes of moist temperate regions, absorb water largely, but very
quickly give it up again.
It may be doubted, perhaps, whether this classification, based
on biological relations, has in itself any permanent value, but mean-
time it serves to express and emphasize what is apparently no mere
theoretical conception, but a simple historical fact, namely, that
differences of habit on the part of these desert plants, as well as
the structural adaptations with which they are correlated, have
become established step by step together, during the long period of
geographical changes through which the land they now occupy
has been passing. A discussion of the physiological significance
of the facts which have been brought out does not fall within the
province of this pay er.
DESERT BOTANICAL LABORATORY,
son, Arizona.
NEW SPECIES OF CALIFORNIAN PLANTS.
ALICE EASTWOOD.
(WITH TWO FIGURES)
’ Zygadenus exaltatus, n. sp.—Bulb large, pear-shaped, covered
with brownish, membranous coats, about 1o°™ long, and half as
wide: radical leaves forming a conspicuous bunch sheathing
the base of stem, 6°" or more long, 2°™ wide, veins prominent,
midrib conspicuous on lower part of leaf, less so above, glabrous
except for the short, rather thick cilia on the margin: stem tall
and stout, hollow, 7-8 high, 1°™ in diameter at base, leafy to
the inflorescence: upper leaves merging into the bracts, lower cau-
line with broad clasping base, 3-44" long and as broad as the
radical leaves: inflorescence paniculate, 2-3°" long, the upwardly
spreading branches varying in length at different stages of devel-
opment, the lower branches generally bearing only staminate flowers,
the perfect flowers principally borne on the main stem above the
branches; staminate racemes with peduncles shorter than the bracts;
bracts attenuate; bractlets white, membranous, longer or shorter than
the slender pedicels, ovate-attenuate: perianth 17™™ in diameter,
outer divisions sessile, elliptical, obtuse, the gland 2™™ from the base
with parallel veins below forming a margined claw, veins on the
upper part proceeding from the teeth of the gland, branching;
inner divisions of the perianth with claw 2™™ long, 1™™ wide, the
gland oblong, obtuse, veins asin the outer division; filaments broaden-
ing towards the base, 5™™ long, attached to base of perianth, anthers
oblong, becoming explanate in age: fruit becoming 2°™ long, includ-
ing the rostrate styles, tapering a little more at base than at summit.
Type collected by F. E. Blaisdell at Prindle’s ranch, above Mokelumne
Hill, Calaveras County, California, at an altitude of 425™, in April 1900
The other specimens in the Herbarium of the California Academy of Sciences
are Amador, California, May 1886, M. K. Curran ae with root); Soda
Creek, Tulare County, C. A. Purpus, June 1896, no. 1549; Hindeman’s Trai]
gis Coyote Pass, Tulare County, California, July 19, 1903, collected by myself.
I also saw it on the trail on the way to Little Kern. In habit of growth and
283] {Botanical Gazette, vol. 41
284 BOTANICAL GAZETTE [APRIL
size of bulb it is related to Z. paniculatus, but it has flowers much larger and all
the floral organs differently shaped. It is probably the largest species of Zyga-
denus known. It is probable that there is no Z. paniculatus on the western slope
of the Sierra Nevada.
“ Silene deflexa, n. sp.—Stems several from a creeping root-
stock, about 24" high, erect, glandular-puberulent especially above:
radical leaves spatulate, 1-2°" long including the margined
petioles, rather thick in texture, obtuse; cauline leaves 3-4 pairs,
the uppermost very small, not more than 3™™, the lowest oblan-
ceolate to oblong, obtuse, 2°" long, connate-clasping and nodose
at base: flowers solitary in the lower leaf-axils, pedicels erect and
close to stem, flowers curved-deflexed in anthesis, erect in fruit;
terminal flowers few, cymose, pedicels capillary, 7-12™™ long,
generally shorter than the flowers; calyx narrowly cylindrical in
flower, enlarging and breaking apart with the expanding capsule,
ro-ribbed, 9™™ long, divisions rounded at apex, oblong, some-
times uneven, membranously margined, 1.5™™ long; petals with
the claws united tc the stamens and the stipe of the ovary at base,
woolly, gradually enlarging to the blade, not auricled, blade 4-cleft
into linear lobes, the two middle 2™™ long, o.5™™ wide, the two
lateral narrower and shorter, appendages oblong, reaching the base
of divisions, retuse with one side pointed; stamens and styles appar-
ently not surpassing the petals, the latter three and the upper thick
part of the ovary splitting into three valves, lower part of ovary
thin cylindrical, all together 4™™ long; stipe 1™™ long and almost
as thick: seeds suborbicular, brown, strophiolate.
Type collected in the Hudsonian Zone above the nies Canyon Creek,
Trinity County, California, by Vernon Baily, August 25,
It is related to S. Lemmoni, but differs in the foliage, eS shorter filaments
and styles, the close inflorescence, and the differently shaped floral organs. It
really resembles that species only in having the flowers pendent and the
blades of the corolla with four divisions.
’ Silene lacustris, n. sp.—Cespitose from+ creeping rootstocks,
slender, erect, 1-1.5 high, glandular-puberulent throughout
especially the inflorescence, flowering from the lowest axils: radical
leaves narrowly oblanceolate, acute or obtuse, tapering to a long
margined petiole, all together 2°™ long; cauline leaves 2-3 pairs,
linear, obtuse, connate-clasping at base, 1-2°™ long, 1-2™™ wide:
ing,
ee ee
1906] EASTWOOD—CALIFORNIAN PLANTS 285
calyx broadly cylindrical, 1°™ long, thin, with ten purple nerves, the
divisions rounded, 2.5™™ long, 2™™ wide; petals with claws woolly
and cohering round the thick stipe, gradually broadening to the
membranous laciniate auricles, 4™™ at top, blades purple, 2-cleft
with rounded divisions, the lateral teeth short or none, the appen-
dages bifid and laciniate, 1™™ long; stamens and styles exserted;
ovary oblong, 4™™ long: fruit unknown.
Type collected by the author at Monarch Lake near Mineral King, Tulare
County, California, July 21, 1903.
This species belongs to the group of high mountain Silenes, including S.
Grayi, S. Watsoni, and S. Suksdorfii. In appearance and shape of leaves it
most closely resembles the first, but differs from this and the others in the broader
auricles, and the bifid, laciniate appendages of the corolla.
‘ Silene pacifica, n. sp.—Perennial, with thick woody rootstocks;
stems generally several, erect, 4.5°" high, viscid throughout, espe-
cially the inflorescence, nodes prominent: leaves rather thin, radical
and lower cauline oblanceolate to obovate or elliptical, tapering
at base and decurrent on the long petiole; blade 5-6°™ long, 2-3°™
wide, slightly ciliate, sparingly pubescent, obtuse or acute; peti-
oles margined, dilated and sheathing at base; cauline leaves con-
nate-clasping at base, 4-5°™ long, the uppermost leaves lanceolate,
sessile: flowers verticillate in the axils or cymose on short peduncles,
pedicels 0. 5-2°™ long, the longest equaling the longest floral leaves:
calyx truncate at base, tubular, becoming somewhat turbinate
with the enlarging capsule, very viscid, prominently green or pur-
plish veined, the divisions deltoid, obtuse or acute, 4™™ long, green
with white or purplish membranous margins, entire calyx 1.5°™
long; corolla claret color, the claws of the petals white or tinged
with claret, 1.5°™ long, attenuate at base and broadening at trun-
cate summit to 4™™, exserted 5™™, blade deeply cleft, each part
entire, laciniate, or bilobed, the prominent divaricate teeth on each
side almost as long as the divisions, more than 1™™, appendages
laciniate-dentate, 1. 5™™ long; stamens monadelphous at base,
encircling the stipe and pubescent, glabrous above, varying in length:
pod 11™™ long, the tips of the three valves stellately reflexed and
often splitting into five or six over the calyx when the seeds are
ripe; stipe stout: seeds light brown, slightly pitted, reniform, 2™™
wide.
286 BOTANICAL GAZETTE [APRIL
Type collected by the author along the south side of Rodeo Lagoon, not
far from the ocean, Marin County, California, July 4, 1905. The species seems
to be isolated, as the number of plants is small and it is not elsewhere to be found
in the region. It is also in danger of extermination on account of the improve-
ments that are now going on in the vicinity of the military post
This is most closely related to S. grandis Eastwood Sin Bodega Head,
likewise a maritime species. It differs in having claret-colored flowers, a differ-
ently shaped calyx, the simpler inflorescence, thinner and different leaves, and
entire lack of the velvety pubescence so noticeable on S. grandis. A smaller
and more slender plant.
“Horkelia mollis, n. sp—Stems several, ascending from the
sheathed caudex, red-purple, villous with fine silky spreading hairs,
about 24" in height: radical leaves 6-9°™ long, less than 1°™
wide, the petiole less than half the entire length, often with a few
scattered simple leaflets near the base; leaflets crowded towards
the top, pinnately divided but apparently pedate on account of
the lower divisions surpassing the upper, the divisions lincar-spatu-
late, 3-4™™ long, finely villous; stipules adnate for 8™™, the free
tips filiform-attenuate, about 4™™ long, villous; cauline leaves
similar but with petioles becoming shorter as they ascend, stipules
often incised and always broader than those on the radical leaves:
flowers corymbose-capitate, terminating the stems, a few solitary
ones or few-flowered clusters in the axils of the upper cauline leaves;
hypanthium campanulate, 5™™ long, the bractlets linear, about
as long as the subulate sepals; petals yellow, the blades broadly
spatulate, 1™™ wide, a little longer, slightly shorter than the linear
claw; stamens 15 in three rows; ovaries 5-20, glabrous, the slender
styles tuberculate at base.
The type is 4405 of Carl F. Baker’s distribution, collected by Culbertson
July 19, 1904, at Hockett’s Meadows, Tulare County, California. In the her-
barium of the California Academy of Sciences are specimens of the same, col-
lected by the author along Volcano Creek in the same region, July 17, 1903.
This species probably is most closely related to Horkelia campestris (Jones)
Rydberg. A comparison with a duplicate of the type of the latter shows H. mollis
to be a larger, more villous plant, the appendages of the hypanthium longer, the
divisions more pointed, the petals more exserted and with blades orbicular
and claws more pronounced. In general the flowers are larger.
v STYRAX CALIFORNICA fulvescens, n. var.—Shrub a meter or
so high, with stiff divaricate branches; older stems gray-black,
i
i
;
:
1906] EA STWOOD—CALIFORNIAN PLANTS 287
younger white or tawny with dense stellate tomentum: leaves orbic-
ular-cordate, the apex obtuse or abruptly acuminate, generally
slightly longer than broad, 3-6°™, both surfaces stellate-tomen-
tose, the upper less than the lower, the fulvous hairs often outlining
the veins on lower surface; petioles 5~1o™™ long: flowers 1-3,
cymose, pendent, the pedicels as long as the peduncles; calyx cam-
panulate, cuneate at base, the margin truncate but marked with
5-6 short obtuse scattered teeth, densely clothed with white or
rufous tomentum; stamens 12, almost equaling the petals, attached
almost the entire length of the corolla tube, filaments glabrous,
ribbonlike, anthers with cell divisions white, the connective yellow,
thick; style thick, broadening at base, lower half tomentose, stigma
2-lobed, surpassing the corolla.
The type of this variety was collected by the author May 17, 1904, near
the Painted Cave Ranch in the Santa Inez Mountains back of Santa Barbara,
California. Mr. T. S. Brandegee collected the same in the same mountains
probably near San Marcos Pass in 1888. There is a specimen also of what
seems the same collected by J. G. Lemmon near San Bernardino, May 1878.
Near the head of Mission Creek a second collection was made by the author.
This bush grew in the shade and was taller and less rufous than the others on
the open hills.
This differs from the typical S. californica in the broader, rounder leaves,
heart-shaped at base, the much denser stellate tomentum, and the general prev-
alence of rufous ha.rs especially on the calyx.
v Diplacus calycinus, n.sp.—Suffrutescent, viscid-arachnoid through-
out, the young stems light brown, branching diffusely: leaves elliptical
to oblong, narrowed at each end, apex obtuse, base cuneate, margin
revolute, entire or somewhat sinuate-denticulate, upper surface glab-
rous, often viscid, lower tomentose and viscid, 2-6°™ long, 1-2°™ wide;
petioles very short, revolutely margined, woolly at junction: flowers
axillary, the peduncles 5~7™™ long; lower part of fruiting calyx cylin-
drical, 2°™ long, 5™™ in diameter, 5-ribbed, upper half dilating ab-
ruptly to thrice the diameter of the lower, with 5 strongly keeled
almost equal divisions 7”™ long, 3™" wide at base when folded,
1™™ at the rounded apex, total length of calyx 3.5°™; corolla light
yellow, the tube curved, uniformly slender for 1.5°™, dilating
above, the divisions having a spread of 1.5-2°™, exserted from the
calyx.
288 BOTANICAL GAZETTE [APRIL
This was first collected by Mr. T. S. Brandegee in Kaweah Canyon, Tulare
County, California, July 26, 1892. The type is 4407 of C. K. Baker’s distri-
bution collected by Culbertson in the south fork of Kaweah River, 1800™ alti-
tude, July 22, 1904.
This species is distinguished from allied species by the peculiar foliaceous
calyx described above. The corolla in the dried specimens cannot be satis-
factorily described, as in both collections the specimens are a little old.
“ Orthocarpus Copelandi, n. sp.—Stems about 1% high, simple
or divaricately branched, minutely scabrous with short, curved
hairs: lowest leaves narrowly linear-lanceolate, obtuse, 3-4°™ long;
upper on main stem as long but twice as broad; uppermost on
branches falcate, alternate or opposite: spike short and dense;
lowest bracts green, the middle division like the broadest leaves,
the lateral divisions spreading and very slender, about one-third
as long as the middle; upper bracts shorter and broader, ellip-
tical, rose-tipped: calyx thin and membranous, becoming globular-
inflated, pink with green ribs, cleft half in front, deeper in the back,
villous. with short gland-tipped hairs, 7™™ long, 4™™ broad, with
divisions triangular attenuate; corolla minutely glandular, 13™™
long, galea straight, obtuse, rose-color, ciliate, 6™™ long, lower
lip yellow, the three sacs inflated somewhat, 5™™ long, middle
tooth much larger than the other two: capsule bright brown, 5™™
long, 3.5™™ wide, obovate with obcordate apex, with few (appar-
ently only two) seeds.—Fic. 1.
Collected on Mount Eddy, August 18, 1903 at an altitude of 2130" by Dr.
Edwin Bingham Copeland, in whose honor it is named. It is a beautiful species
related to O. imbricatus and that group which contains so many closely related
species. H.E. Brown’s number 449 from the north side of Mt, Shasta is the
same but very immature.
v Veronica Copelandi, n. sp.—Perennial from slender, running root-
stocks, about 1% high, simple, glandular-villous throughout: leaves
five or six pairs, crowded on the lower part, sessile, oblong-ellip-
tical, entire, acute, veinless, 1-1.5°™ long, 4-8™™ wide: racemes
sometimes becoming 8°™ long, 5-15-flowered, the highest leaves
often with one or two axillary flowers; bracts lanceolate, the lowest
opposite, others alternate, shorter than the pedicels; peduncles
1-2°™ long, sometimes scarcely apparent; pedicels filiform, 5™™ long,
a small bractlet immediately below the calyx appearing like another
PPT —
1906]
EASTWOOD—CALIFORNIAN PLANTS
Fic. 1.—Orthocar pus Copelandi Eastw.
289
290 BOTANICAL GAZETTE [APRIL
sepal: sepals 4, oblong-ovate, obtuse, 3™™ long; corolla purple,
glabrous, g™™ across, the three larger divisions orbicular, entire,
4™™ in diameter, the smallest ovate-obtuse, 3™™ wide; stamens
exserted, 4™™ long, filiform, anthers obtuse and obtusely sagit-
tate at base, 1.5™™ long; stig-
ma exserted from the opening
bud, obscurely bilobed, style
7mm long, filiform at base,
flattening and slightly broad-
ening towards the apex: cap-
sules becoming almost twice
as long as the calyx divi-
sions, broadly oblong, 5™™
long, 3.5™™ wide, emarginate,
the lobes and sinus obtuse;
style persistent.—FIGc. 2.
This was collected on Mount
Eddy at an elevation of 2500™ by
Dr. Edwin Bingham Copeland,
August 18, 1903, distribution of
C. E. Baker, 1903. no. 3931. It is
near to V. Cusickii Gray, differing
in pubescence, shape of leaves and
sepals, and a larger and more open-
spreading corolla.
v Erigeron decumbens, n. sp.
—Stems several, from slender
creeping rootstocks, decumb-
ent or ascending, 1-1.5™
high, scabrous and somewhat
canescent with short appressed hairs which are glandular at base
(under a lens): leaves oblanceolate to spatulate, sessile, obtuse,
apparently veinless, 5-15™™ long, 3-5™™ wide, with pubescence
similar to the stems: heads few, rayless, 7™™ high, terminating short
branchlets, which are leafy near the junction with the stem and
have a few scattered minute bracts on the upper part; scales of the
involucre in four series, glandular-puberulent, outer ones small,
reflexed-spreading, inner green-tipped, ribbed, membranous at base,
FIG. 2.—Veronica Copelandi Eastw.
|
|
|
1906] EASTWOOD—CALIFORNIAN PLANTS 291
linear-lanceolate, acute, 6™™ long: corolla yellow, tubular, abruptly
narrowed 1™™ above the base, the border consisting of five short,
obtuse, incurved teeth; style branches exserted, the hairy tips very
short: akenes slightly hairy at top; pappus simple, barbellulate, as
long as the corolla.
Collected by Dr. Edwin Bingham Copeland on Mount Eddy, Siskiyou
County, California, at an altitude of 1400™, August 17, 1903. It belongs to
the group which includes E. miser Gray, as well as many species described by
Dr. E. L. Greene in Flora Franciscana, p. 394; but it agrees with none.
Erigeron Copelandi, n. sp.—Cespitose from an underground,
branched caudex, covered with black, scale-like, imbricated bases
of old petioles: radical leaves spatulate, subcanescent, with closely
appressed very short pubescence; petioles equaling or longer than
the blades, together 1-3°™ long, 4-8™™ wide, the petioles dilated
and closely imbricated at the reddish-purple base: stems 1-flowered,
5-10" high, sparsely leaved with narrow linear or linear-oblan-
ceolate leaves 5-10™™ long, becoming minute and bract-like on
the glandular-puberulent upper part which is like a peduncle: heads
about 6™™ high exclusive of the numerous, very narrow, lilac to
violet rays, which are 5™™ long; scales of the involucre in three
series, glandular-puberulent, the outermost shorter, clothed with
some scattered hairs, innermost linear-attenuate, sparsely ciliate,
green-ribbed, membranously margined, about 5™™ long; disk
flowers numerous, yellow, 2.5™™ long, narrowed 1™™ above the
base, glandular on the lower part, the border of five short acute
incurved teeth: pappus upwardly barbellulate, simple, that of the
ray flowers shorter than that of the disk, none as long as the corolla;
akenes clothed with upwardly spreading hairs; stamens exserted
in some flowers, pistils in others; fertile and sterile flowers in the
same head, ray-flowers sterile.
Collected on Mt. Eddy, Siskiyou County, California, at an altitude of 1250™
_ by Dr. Edwin Bingham Copeland, in whose honor it is a pleasure to name this
pretty plant. It is related to E. pygmaeus Greene and others of that group,
but differs from all in caudex, pubescence, leaves, and heads.
Y Chrysopsis gracilis, n. sp—Stems slender, simple, 3°" high,
loosely and sparingly villous-arachnoid, terminated by 2-4 cymose
heads: leaves thin, linear-lanceolate, narrowly acuminate, 3-4°™
292 BOTANICAL GAZETTE [APRIL
long, 6™™ wide, sessile, the upper surface somewhat dotted, lower
surface arachnoid (under a lens): peduncles with pubescence like
the stem but also somewhat viscid, bracts few, narrowly linear:
involucral scales in about 5 ranks, the outer narrowly linear- atten-
uate, the others lanceolate, acute, tipped with a green and glandular
spot, below yellowish, chartaceous, keeled, the innermost some-
times tinged with purple and considerably surpassing the others;
heads with about 15 flowers, rayless: corolla straw color, about
as long as the pappus, trumpet-shaped, gradually narrowed to the
base, border with acute teeth, 1™™ long; style branches filiform,
exserted, twining around each other at base; pappus thick, with
an outer shorter row, barbellulate; akenes flat, villous, white.
Collected on Mount Eddy at an elevation of 2225" by Dr. Edwin Bingham
Copeland, August 17, 1903.
This comes very near C. Breweri Gray, of which it may prove to be only
a variety. It differs however, in the simple instead of much branched stems,
more finely arachnoid pubescence, and leaves of different outline. The invol-
ucral scales are the most distinctive; in C. Breweri they are attenuate and not
keeled; in C. gracilis they are broader, acute, keeled, conspicuously green-
tipped and glandular; the corollas are paler and the pappus not so rough.
- Psilocarphus tenuis, n. sp.—Sparingly clothed with long loose
white woolly hairs; stems filiform, erect or ascending, 3-5°™ high,
with few slender divaricate branches: leaves oblong to elliptical,
5-10™™ long, 3™™ wide, veiny and submembranous, mucronate
at apex, the base of the opposite leaves connate-clasping: heads
in the forks and at the ends of the branches, the involucral leaves
4, ovate-oblong, folding over and almost concealing the flowers
within, texture similar to the other leaves: fertile flowers few or
many, completely enclosed by the obliquely-cuneate bracts, these
gibbous, veiny, membranous, slightly woolly, 2.5™™ long, the apex
orbicularly truncate or concave, the exserted membranous tips brown-
ish, conspicuous, generally curved upwards: akenes shortly stipitate,
narrowly obovate, 1™™ long: sterile flowers few, the corolla attenuate
to the base, divisions reddish-brown.
Type collected at Monterey, California, by Mrs. Joseph Clemens, July 1905.
What seems to be the same, but too young for certainty, was collected by the
author at Bakersfield, Kern County, California, April 4, 1893, and at Kaweah,
Tulare County, California, April 27, 1895.
1900] EASTWOOD—CALIFORNIAN PLANTS 293
This seems most distinct from all the other species in having the involucral
leaves almost closing over the flowers, the peculiar concave or truncate top to
the bracts enclosing the fertile flowers, and in the more veiny and membranous
foliage, less woolly pubescence, and more slender habit.
~ Senecio Millikeni, n. sp.—Stems tall, glabrous, hollow, ribbed,
paniculately branched, the slender virgate branches leafless in
the lower part: leaves linear-lanceolate, narrowed at both ends,
with acute apex, sessile base, margin dentate with small uneven
obtuse teeth, the lower 12°™ long, 2.5°™ wide, diminishing up-
wards: panicle thrysiform, the peduncles and pedicels slender,
bracts and bractlets attenuate, equaling or longer than the slender
pedicels: heads 1°™ high, bracteate at base, the involucre 5™™
high, with glabrous scales tipped at apex with a tuft of tomentum;
rays 6, 3-toothed, 7™™ long, style exserted 3™™; disk flowers
7™™ long, the acute triangular teeth of the corolla slightly gran-
ular, stamens exserted but style branches surpassing them: akenes
glabrous; pappus soft and abundant, about as long as the corolla.
Type collected in Natural Bridge Meadows, Tulare County, California, by
Culbertson, Aug. 10, 1904, C. F. Baker’s distribution 4268. It is named in
honor of Mr. Culbertson’s assistant.
This belongs to the polymorphous group of which S. triangularis was the
first described. It differs from all in the narrowed bases of the leaves, the thyr-
siform inflorescence, and the smaller heads.
SAN FRANCISCO, CALIFORNIA.
Pricrerk ARTICLES.
NOTES ON NORTH AMERICAN GRASSES. VI.
SYNOPSIS OF TRIPSACUM.
Tripesacum L., Syst., Ed. 10, 2:1261. 1759.
A GENUS of grasses confined for the most part to North America. The
type species is 7. dactyloides L.
KEY TO SPECIES.
Staminate spikelets all sessile or nearly so, outer glume coriaceous;
spikes single or 2- to 3-digitate. Section DACTYLOIDEs.
Blades 4 to 5°™ wide, pubescent on upper surface. . . Jatifolium
Blades mostly less than 2°™ wide
Blades 1 to 3™™ wide, involute . . - . + « + floridanum
Blades 1 to 2°™ wide, flat
Sheaths glabrous, blades sian except sometimes
along the midribabove. . . . . dactyloides
Sheaths more or less hispid, or sometimes es glab-
rous, blades hispid on upper surface dactyloides hispidum
Staminate spikelets with one of the pair sessile, the other pedicelled,
outer glume membranaceous; pistillate spikes branched, form-
ing a fascicle. Section FascrcunaTa.
Sheaths hispid . . pilosum
Sheaths glabrous daceps ‘oe fowccstiosé: or « Miepid ‘ally at the
throat
Blades 3°™ or more in width, glabrous . . . fasciculatum
Blades 2°™ or less in width, pubescent on upper surface
1.5 to 2°™ wide, flat or folded, culms robust . .lanceolatum
5 to ro™™ wide, more or less involute, culms
Ct, . . « . Lemmoni
Tripsacum latifolium, n. sp. ge ose vaginis glabris vel
apice pubescentibus, laminis amplis, ad 4.5°™ latis, 70°" longis, planis,
supra pubescentibus subtus scabris vel glabrescentibus, spiculis steril-
ibus geminis sessilibus, 3-4™™ longis, oblongis, obtusis vel breviter acutis.
Culm robust, 1°™ in diameter, glabrous; sheaths glabrous or pubes-
cent towards apex; blades ample, as much as 7o°™ long and 4.5°™
Botanical Gazette, vol. 41] . [294
de ee
1906] BRIEFER ARTICLES 295
wide, pubescent above, minutely papillate-scabrous or glabrescent be-
neath, scabrous-ciliate on the margin; ligule very short, scarcely 3™™
long, fimbriate; spikes 1 to 3, similar to 7. dactyloides but more slender,
pistillate section 2 to 3™™ wide, staminate spikelets sessile or nearly so,
3 to 4™™ long, outer glume coriaceous, oblong, rounded at apex, scab-
rous, ciliate on marginal keels, rather minutely striate with about ten
nerves. ;
The type specimen was collected by H. von Tuerckheim at Cubilquitz,
Dept. Alta Verapaz, Guatemala, alt. 350", Jan. 1902, no. 8333. The
only other specimen I have seen was collected by C. Thieme at San Pedro
Sula, Dept. Santa Barbara, Honduras, alt. 500™, March 1887, no. 55958.
Both specimens are in the National Herbarium (Herb. John Donnell
Smith).
The species is well distinguished from the other species with sessile
staminate spikelets by its broad pubescent leaves.
Tripsacum DActTyLompEs (L.) L., Syst., Ed. 10, 2:1261. 1759.—
Coix dactyloides L., Sp. Pl. 2:972. 1753.—Usually glabrous through-
out except the upper surface of the blades along the midrib near the base.
This and sometimes a considerable portion of the upper surface of the
blades may be sparsely pilose. The specimens from Florida and along
the Gulf Coast are usually pilose in this way, or occasionally the pubes
cence may extend to the young sheaths of the branches. The more pubes-
cent forms connect the species with the following subspecies, which occurs
in Mexico. The terminal spikes are usually in digitate clusters of two
to three, while the axillary spikes may be single. Sometimes, especially
in Texas, the terminal spikes are also single (TJ. dactyloides monostachyum)
(Willd.) Gray, Man. 616. 1848. T. monostachyum Willd., Sp. Pl. 4:202.
1805. Type locality ‘Carolina meridionali.””
Southern New England to Florida and Texas, mostly near the coast;
but extending inland west to west Texas, and north to Nebraska, Iowa,
southern Illinois, and eastern Tennessee.
If the spike is single the pistillate portion is cylindrical; if the spikes
are two or three, the pistillate portions are flattened on the inner surfaces
so that all together they form a cylinder, and the lower are more or less
peduncled.
TRIPSACUM DACTYLOIDES hispidum, n. subsp. —Laminae supra his-
pidae; vaginae hispidae vel glabrescentes.
The staminate flowers are less chartaceous than is usual in 7. dacty-
loides.
Mexico and southward. San Luis Potosi, rocky hills, Las Canoas,
296 BOTANICAL GAZETTE [APRIL
Pringle 3811 (type); Jalisco, Rio Blanco, Palmer 509; City of Mexico,
Holway 8; Lower California, El Taste, Brandegee, Nov. 1, 1902; Trini-
dad, Botanical Garden Herbarium 3303; Central Paraguay, Morong 675.
This form connects T. dactyloides with T. lanceolatum. In some
specimens the upper spikelet of the staminate pair is somewhat pedicelled.
T. dactyloides and possibly some of the other species may occur widely
distributed in South America. Information on this point is desired.
TRIPSACUM FLORIDANUM Porter, Contr. Nat. Herb. 3:6. 1892.
PortTEr’s herbarium name was published by Dr. VAsry in his monograph
of the grasses of North America. Type locality ‘Florida (A. P. Garber)
and Texas (G. C. Nealley);’ duplicate type in National Herbarium.
T. dactyloides floridanum Beal, Grasses 2:19. 1896. There are no
_ specimens of this species from Texas in the National Herbarium, nor
are there any so labeled by Dr. VAsEy; consequently the Texas locality
given above is uncertain and is probably incorrect.
Our specimens are all from the vicinity of Miami, Florida, Garber 454,
June 1877 (type); Pollard & Collins 272, April 1898; Eaton 530, Dec.
1903; Hitchcock, March 1903.
Distinguished from T. dactyloides by its smaller size and much nar-
rower leaves.
TRIPSACUM FASCICULATUM Trin.; Ascherson, Bot. Zeit. 35:521-
1877.—Well distinguished by its ample glabrous leaves, which are as
much as 6.5°™ wide and 70% long, resembling leaves of Indian corn
(Zea mays L.). Plant glabrous throughout; spikes branched, forming a
fascicle; staminate portion slender and more or less flexuous, the spike-
lets 5 to 6 ™™ long and broadest near the top.
The name first appears in the second edition of SrEUDEL’s Nomen-
clator. 2:712, as Tripsacum “‘fasciculatum Trin. Mpt. Mexico. T.
dactyloides Schlecht. in Linnaea VI.”” The latter name is a nomen nudum,
as is also T. fasciculatum Trin. in Steud. Gram. 1:363, and in Ruprecht,
‘Bull. Acad. Brux. 9:243. The first description appears to be by ASCHER-
SON’ in 1877, Bot. Zeit. 35:525, where a specimen from ‘“‘Pr. Hacienda
de la Laguna (Schiede)”’ is designated as the type. Fournter, Mex.
Gram. 69. 1881, includes the name without description and cites the -
following specimens: Hacienda de la Laguna (Schiede 947); Orizaba
* ASCHERSON had previously mentioned the species and given a brief description
as follows: ‘Diese Art besitzt Blatter von der Breite der Maisblitter, und die zahl-
reichen, schlaffen, mannlichen Inflorescenzzweige, deren Aehrchen kleiner als bei
T. dactyloides sind, erinnern ebenfalls an Euchlaena.” (Verh. bot. Ver. Pr. Brandenb.
17:79. 1875, in a footnote to an article on Euchlaena mexicana.)
a cogs ie rc I ace
1906] BRIEFER ARTICLES 297
(Bourgeau 3138); Mirador (Liebmann 549); Zacuapan pr. Jalapa (Gal-
eoltt 5796); Arumbaro (Galeotti 5844). The Bourceau and LirBMANN
specimens are in the National Herbarium; also Brade 16174, from Costa
Rica.
_TRIPSACUM LANCEOLATUM Rupr.; Fournier, Mex. Gram. 68. 1881.—
Leaves mostly 1 to 2°™ broad, pubescent on the upper surface; stami-
nate flowers 7 to g™™ long, spindle-shaped, often rather abruptly nar-
rowed above the middle.
Mexico. Sonora, Guadaloupe Cafion, International Boundary Com-
mission, 2035; Durango, Palmer 537; Oajaca, Villa alta, Liebmann 547;
Lower California, Sierra de San Francisquito, Brandegee 6, Sept.
1899; Jalisco, between Huejuquilla and Mesquitec, Rose 3570. In
addition to these specimens in the National Herbarium, FourNIER gives
the following: Inter Victoria et Rio Blanco (Karwinsky); Borrego prope
Orizaba (Botteri 1213 in herb. VAN HeEuRcK); Mirador (Schaffner);
Tacubaya (Schaffner 41 in herb. FRANQUEVILLE); Secus Amnem in her-
bosis- pr. Pedregal (Bourgeau 444); Aguas Calientes (Hartweg 252).
Liebmann 547 is also cited by FourRNIER and it is upon this specimen
that I have based my identification of the species. FoURNIER’s description
does not apply in all respects to the plants which I have included under
this species. He states that the culms are pilose, which is not true of any
of the specimens I have seen. Neither are both staminate spikelets pedi-
celled, as he describes.
The name first appears in Plant. Haitw: Addenda, p. 347. In the
body of the work (p. 28) no. 252 is listed without description as T. dacty-
loides ‘‘in saxosis, Aguas Calientes.” In the addenda this is corrected
as follows: ‘“‘n. 252 est species a Tripsaco dactyloide distincta, T. lan-
ceolata, Ruppr. ex cl. Rupprecht in Litt.”” Fournter (/.c.) cites T.
lanceolatum Rupr. in Benth. Pl. Hartw. 247. Under the circumstances
I think Hartwec’s no. 252 from Aguas Calientes should be considered
as the type of 7. lanceolatum rather than coca gomgse s specimen, the
first cited by FouRNIER.
FourNIER cites as a synonym of this “7. acutiflorum Rupr. mss. in
herb. Petrop.”” Under the rules of the recent code 7. acutiflorum was
not published. FouRNIER (/. c. 69) also mentions without description,
var. 8B monostachyum from San Luis Potosi (Virta 1447). I have not
seen this specimen.
TRIPSACUM PILOSUM Scribn. & Merr., Div. Agrost. Bull. 24:6. rgor.
—Type locality Mexico. ‘Collected on the road between Colotlan and
298 BOTANICAL GAZETTE [APRIL
Bolafios, State of Jalisco, 2841 J. N. Rose, September 7, 1897.’’ Speci-
men in National Herbarium.
The preceding species, together with this and the following, form a
rather closely connected series. The type of T. pilosum is distinguished
by the strongly papillate-hirsute sheaths, and the blades pubescent upon
both surfaces, but these characters are much less marked in some of the
specimens which agree with the type in other particulars.
I have referred here the following specimens: Jalisco, Rio Blanco,
Palmer 508; Cafion near Guadalajara, Pringle 2623, and hills near Guada-
lajara, Pringle 2611; San Luis Potosi, limestone ledges, Tinamel, Pringle
3993; and San Jose Pass, Pringle 3447.
TriesacuM Lremmoni Vasey, Contr. Nat. Herb. 3:6. 1892. Type
locality, ‘‘Huachuca Mountains, Arizona (J. G. Lemmon).” Type
specimen in National Herbarium. T. dactyloides Lemmoni (Vasey) Beal
Grasses 2:19. 1896.
Plant glabrous throughout except the lowermost sheaths, which are
more or less hispid. The leaves are long and narrow, 5 to 10™™ -wide,
and in herbarium specimens inrolled at the margins.
In addition to the type specimen I have included two Mexican speci-
mens, Jaral, Gebirgsthaler, Schumann 1718, and Jalisco, Mountains
near Guadalajara, Pringle 2610. These two specimens have the spikes
digitate instead of fascicled as in Arizona specimen, but the latter has
the lateral spikes in ones or twos.—A.S. HirrcHcock, U. S. Dept. Agric.,
Washington, D. C.
BOOK REVIEWS.
Vegetable foods.
THis well-known work of MOELLER on this subject," first put out about
twenty years ago, has played an important part in connection with the increasing
use of the microscope as a practical instrument for recognizing vegetable sub
stances in a more or less finely divided state. Many changes have taken place
essary.
now of the Connecticut eco ral Experiment Station, a pupil of MoELLER,
has furnished very important aid in the form of excellent figures as well as text.
The scope of the work is in general indicated by the title and those articles
here treated are with few exceptions used as food for man or beast, the term
food being defined so as to include such articles as flavoring agents as well
as tea, coffee, and cacao. Under the appropriate headings those substances
are also described and figured which occur as impurities, substitutions, and
adulterations. Since it often happens that condiments are also official ,
many chapters have a strong pharmaceutical interest. A few articles are con-
sidered which have their chief significance as coe products, e. g., sandal wood,
guarana, cubebs, cola, salep, and ca
In the treatment of the individual has. the book is distinguished by a
concise and exact statement of the features, gross and microscopic, character-
izing the structures concerned, dimensions frequently cited giving definiteness
to terms of size.
As valuable as the excellent text, are the numerous drawings illustrative
of it. A large number are original, many being by Dr. Winton. A bibliog-
raphy of the most important articles written on each subject closes the consid-
ation. One novel feature among the illustrations is seen in the gross pictures
of the leaves discussed. Here a direct print is made on a sensitive surface,
using the leaf itself as an opaque object. This method has been successfully
used before by a number of authors with various objects and here the result
is in general successful. Frequently a very considerable amount of detail has
: eect nee — ——T vege Nahrungs- und Genussmittel aus dem
fi te und unter Mitwirkung A. L. Winton’s
vermehrte Auflage. ous pp- iit sons jigs. 599. Berlin: Tolles” Springer. 1g05.
M 18; geb. M 20.
299
300 BOTANICAL GAZETTE [APRIL
oe is in the half tone reproductions of these so-called ‘‘autophoto-
‘The revision of this important work has again brought it to the front and
promises to continue it as one ies the valuable literary aids to the investigator
of pure foods.—Ropney H.
ALMOsT simultaneously with the foregoing has appeared in this country a
similar compendium by the same team; this time the pupil leads and the master
is the collaborator.2 The general plan and purpose of Dr. WinTON’s weighty
volume are similar to those of Dr. MoELLER’s. The fact that it is in English
will give it a sale that the German book could not hope to attain among the food
commissioners and inspectors and the official chemists, to whom at present
such a work makes its chief appeal. By reason of the existing agitation in this
country on the subject of pure foods and drugs, the enforcement of existing
laws, and the imminence of new and more exacting legislation, this publication
is peculiarly timely. The botanical features are on the whole reasonably accu-
rate, especially the anatomy, which is most fundamental. The definitions in —
the glossary are not always above criticism, and accuracy would not have ren-
dered them less practical. The illustrations are numerous and good, particu-
cited in the bibliography. The arrangement of material, analytic keys, lists
of adulterants, and the suggestions as to diagnosis are sure to be of great practical
Service in the new paaoraes against sophistication by unscrupulous manu-
facturers and dealers—C. R. B.
MINOR NOTICES.
Cryptogamic flora of Brandenburg.s—This monumental work begins its
seventh volume with the first fascicle of the Ascomycetes. Its character and
scope are so well known that the announcement of its publication and contents
Nn suffice to secure the orders of all who concern themselves with this group.
miasci are treated by G. Liypau; Saccharomycetineae by P. LINDNER;
Pobsatiein by G. Linpav; Exoascaceae, Erysiphaceae, Perisporiaceae, Macro-
thyriaceae, and Aspersiaceac by F. Necrer; Onygenaceae, Elaphomycetaceae,
Terfeziaceae, and Tuberaceae by P. Hennincs.—C. R. B.
2 Winton, A. L., The microscopy of vegetable foods, with special reference to
the detection of adulteration and the diagnosis of mixtures. With the collaboration
of Dr. JoseF MOELLER. Imp. 8vo. pp. xvit+7or. figs. 589. New York: John
Wiley & Sons. 1906. $7.50.
3 Kryptogamenflora der Mark Brandenburg, apn 7, Heft x. Pilze. Von P.
HENNINGS, G. Linpav, P. LinpNErR, F. NEGER. 8vo. pp. 160. figs. 17. pls. 8. Leip-
zig: Gebriider Decanneace: 1905. M1.50. (Not sci hear)
1906] CURRENT LITERATURE 301
NOTES FOR STUDENTS.
What is a species?—The many discussions as to what is a species have
resulted in a general appreciation of the facts that species are not all of equal
rank, that they are distinguished by more or less arbitrary characters, and that
although many species are real natural groups of individuals, many others are
simply arbitrary groups, associated for the sake of convenience. After review-
ing the various methods of distinguishing species, KUpPFFER* concludes that
no method will apply in all cases, that all methods are of importance, and that
when the several methods are used conjointly, little difficulty is experienced.
KupFFER then turns to the methods of K6LREUTER, based upon the sterility
of hybrids, as a method which has not been used to the extent its merits warrant.
Sterility of the hybrids being presumably due to defective germ-cells, he depends
for his measure of sterility upon the condition of the pollen, basing his method
upon the fact pointed out a few years ago by JENcIc5 that viable pollen swells
immediately upon the introduction of water, while the sterile pollen remains
- Shrunken, and that this capacity of the normal pollen to swell is retained for
many years in herbarium materials (more than 50 years in Viola, fide KUPFFER).
Although the author eee that considerable sterility of the pollen has
been observed in many “good” species, he has himself never found a pure
species in which more than a few (ein Paar) per cent. of the pollen grains remained
shrunken, the implication being that the reported instances would bear further
consideration.
After examining a number of species and their hybrids, especially among the
Violaceae, he concludes that when a supposed hybrid shows much less fertility
of the pollen than its supposed parents, it is not a mecessary but a sufficient proof
(1) that the supposed hybrid is truly a hybrid, and (2) that its parents belong to
distinct species,
Application of this aehed is then made with interesting results to forms
of Potentilla, Viola, Thymus, etc., which have puzzled the systematist—GEORGE
H. Suu.
Propagation of grain rust—Further comments by Dr. JaKosp Ertxsson®
on the question of the origin and distribution of the rust-diseases of plants have
recently been presented to the botanical public through separata. The author
has not essayed so much to put forth new facts, as to bring together and review
those recently published, in so far as they bear upon his mycoplasm theory,
giving especial attention to adverse criticisms.
4Kuprrer, K. R., Kélreuters Methode der Art Abgrenzung nebst Beispielen
ihrer Anwendung und einigen allgemeinen Betrachtungen iiber legitime and hybride
Pflanzenformen. Acta Hort. Bot. Univ. Imp. Jurjevensis 6:1-19. 1905.
5 JENcIc, Untersuchungen des Pollens hybrider Pflanzen. Oesterr. Bot. Zeits.
$0:1, 41, 81. 1g00.
6 Errxsson, J. Zur Frage der Sulsictoaing und Verbreitung ie Rostkrank-
heiten der Pflanzen. Arkiv for Botanik 53:1-54. 1905.
302 BOTANICAL GAZETTE [APRIL
He maintains that after taking into consideration the studies and observa-
. tions of MArsHALL Warp and PLlowricut in England; McALpine and Coss
in Australia; Bottey, HircHcockx, and CaRrLeTon in North America; BaRr-
cLay in India; KLEBAHN, DreTEL, SCHROETER, and MAGNus in Germany;
LAGERHEIM in Sweden, and others, the wintering of the uredo-bearing mycelium,
or of the uredospores, so as to be a source of infection for the coming season,
has not been proven. The evidence, chiefly as brought forward by KLEBAHN,
to show that the first appearance of the rust in spring can often be accounted
for by uredospores being carried long distances by the wind, is reviewed, and
the conclusion reached that this is an assumption based on no direct evidence
and highly improbable.
The author then enters upon the vital part of the subject and discusses the
mycoplasm theory and its recent criticism, especially that which has been most
ably presented by KLEBAHN and MarsHALL Warp. After an extended examina-
tion of the works of these authors, he finds that his theory has not been affected.
He directs attention to a report by BIFFEN of recent experiments in hybrid-
izing wheat carried on at Cambridge, England, in which the appearance of
rust on the plants can best be explained by assuming that the mycoplasm of
certain varieties was transmitted through the pollen to the resulting hybrid.
—J. C. ARTHUR.
Gynodioecism.—CorrENS? “has presented a second® report on the gyno-
dioecism of Satureia hortensis and Silene inflata, giving full confirmation of his
earlier conclusion that the pistillate form produces only, or mostly, pistillate
offspring when fertilized, as it must be, by the bisporangiate form. If the pis-
tillate form is a mutant from the bisporangiate and differs from the latter by the
possession of a distinct hereditary unit, as suggested by Burck,® all the seeds
produced by a pistillate plant are of hybrid origin, and the observed facts would
be best explained as a case of dominance of the newly risen character over the
older. In Satureia this dominance (?) is complete, but in Silene the offspring
of the pistillate plants were pistillate in only 87-93 per cent., the rest being bi-
sporangiate. Although this behavior looks very much like Mendelian inheri-
tance, a number of cases are cited in which quite contradictory results have
been obtained, so that while the author states it as a law that each sex has a
tendency to transmit its own sex form, he does not look upon this as dominance
in the Mendelian sense—Grorcre H. SHULL
An ear of corn.—The origin of such economic plants as wheat and maize,
which have a wide distribution in cultivation but are unknown in the native
tis Bot. GAZETTE 39:304. Ap. 19
, C., Weitere cae iiber die Gynodioecie. Ber. Deutsch.
Bot. Gesell. aie 452-463. 1905.
, W., Die Mutation als Ursache der Kleistogamie. Recueil Trav.
bot. pee I-2:95 sqq. 1905.
EDS lg _ ee
1906] CURRENT LITERATURE 303
state will doubtless always be an interesting subject for speculation. The most
satisfactory hypothesis for the origin of maize, and that which has been until
this time rather generally accepted, derives it from the teosinte (Euchlaena).
It has been thought that the ear was formed by an abnormal coalescence of
the pistillate spikes of that plant. The ease with which maize and teosinte
may be crossed gives strong support to the theory that they are nearly related.
An altogether different view of the origin of the pistillate spike of maize is
presented by MonrGomERy’® and much evidence is given in its support. His
ypotheses are that the ear of corn is the homologue of the central spike
of the staminate inflorescence; and that the progenitor of maize was a much
branched plant, bearing only terminal branched inflorescences of bisporan-
giate flowers. The chief support of these hypotheses is derived from abnormal
development of pistillate and bisporangiate flowers in the staminate inflores-
cence, and vice versa. A number of photographs show these abnormalities
and jig. 74 represents a plant, denuded of its leaves, showing that the same num-
ber of internodes intervene between the central axis and the ear as are found
between the ear and the tassel. Nothing in this new interpretation of the pistil-
late spike of maize need lessen the conviction of its near relationship with Eu-
chlaena.—GerorcE H. SHULL.
The laws of inheritance —CorrENsS*' published a lecture on the laws
of inheritance which presents in a very satisfactory manner the recent advances
which have been made in this discipline. He would include in hybridization
every union between two germ-cells having one or more different character-
The laws of dominance and of the purity of the parental gametes are illus-
trated from his own experiments on Urtica, Mirabilis, and Zea, and emphasis
is given to the fact that these two laws are absolutely unrelated to each other,
and that reference to them jointly as Mendel’s Law is ing.
Latency is considered at some length, but the present state of knowledge
of this subject leaves much to be desired. He makes a proper distinction between
latency in the sense of invisibility, and frue latency in which there is actual inac-
tivity of a unit that may be changed at times from a passive to an active state.
Regarding the relation between MENDEL’s and GALTON’sS laws, he holds with
DARBISHIRE,"? that both are correct and the antagonism only apparent, due
to the different manipulation of the data.
CorRens still maintains that sex is fundamentally unlike the unit-characters
which behave in accord with MENDEL’s laws. Touching on xenia and tel-
10 Montcomery, E. G., What is an ear of corn? Popular Sci. Monthly 68:
55-62. jigs. - Jan. 1906.
11 CORRENS, C., Ueber basi et gn 8vo. pp. 43. figs. 4. Berlin: Gebr.
Borntraeger. 1905.
2 DARBISHIRE, A. D., On the supposed antagonism of Mendelian to biometric
ari of heredity. Mem. and Proc. Manchester Lit. and Philos. Soc. 49. no. 6.
1905- 19 pp-
304 BOTANICAL GAZETTE [APRIL
ing he. holds that neither exists in the strict sense, namely that ids may escape
m the ‘germ-cells to produce modification in the surrounding maternal tissues,
or to be transferred thence into subsequent germ-cells.—GEORGE H. SHULL.
Heterostyly in Primula——The inheritance of heterostylism in Primula has
been investigated by BATESON and GreGory,' who find that there is general
agreement with Mendelian expectation, the short style being dominant over
the long style. A second character, a yellow flush in the center of the flower,
which was found associated with an “‘equal-styled” condition, also proved to.
be Mendelian and capable of being transferred by crossing to the short-styled
form. The investigation showed that whenever the yellow flush occurs in a
combination in which the long style would be expected, the styles do not develop
beyond the level of the anthers, thus forming the “equal-styled” type. Several
aberrant results were observed, the most noteworthy being a case in which a
single plant indicated a different composition of its germ-cells, according as
it was used as the pollen-parent or pistil-parent.—GrorGE H. SHULL.
Asparagus rust —Smitu‘+ has published a final account of his investigation
of the asparagus rust in California. One of the most important results of his
work is the demonstration of the fact that the spores of this rust depend upon
dew for the moisture required for germination. The more detailed account
of the water relation of this rust was published in this journal.'s This discovery
suggested certain practical methods of controlling the rust, such as planting
the rows with the wind and preventing weeds and other plants or trees from
forming a windbreak close about the asparagus field. In other words, the field
should be well ventilated. The bulletin will long continue to be the standard
work of reference for information upon the subject—E. Mrap WILcox.
Potato scab.—HENDERSON* has recently published the results of his studies
of the methods of control of the potato scab. He found that rolling the potato
tubers in sulfur did not prevent the scab, and this is in accord with results secured
by other investigators. Formalin and corrosive sublimate gave equally good
results with the factor of safety in use in favor of the formalin. If treated pota-
toes were planted in soil in which “scabby’” potatoes had grown the previous
season, the scab appeared in spite of the treatment. This emphasizes the neces-
sity of preventing new ground from becoming infected with the disease by plant-
ing none but healthy tubers.—E. Mrap WItcox.
"3 Bateson, E., and Grecory, R. P., On the inheritance of heterostylism in
Primula. Proc. Roy. Soc. London B. 76:581-586. 1905.
14 SMITH, R. E.—Asparagus and asparagus rust in California. Bull. Calif.
Exp. Stat. 165:1-99. figs. 1-46. 1905.
1s Smith, R. E., Bot. GazetrEe 38:19-43. figs. I-21. 1904.
*6 HENDERSON, L. F., Potato scab. Bull. Idaho Exp. Stat. 52: 1-8. 1906.
ee se ee |
1906] CURRENT LITERATURE cha 305
Nuclear division in Ascomycetes.—GUILLIERMOND"’ has continued his
studies on nuclear division in the Ascomycetes, which support in all essentials
the conclusions of HARPER and contravene those of MAIRE (except as to Gal-
actinia), though they are perhaps not irreconcilable with them. However,
his descriptions are not so detailed as those of HARPER in his last paper on Phyl-
lactinia, especially as it relates to the centers of spindle formation. In this
r GUILLIERMOND discusses chiefly the mother-cells of the asci and secre-
tion. The species studied comprise Pustularia vesiculosa, Aleuria cerea, Peziza
rutilans, P. Catinus, and Galactinia succosa. —B. M. Davis
Soil waters.—CAMERON and BELL show" that as a rule the various mineral
constituents of the soil solutions exist in sufficient concentration for the growth
of crops, and that the magnitude of the concentrations is practically the same for
all soils, because, generally speaking, soils contain all the common rock forming
minerals, some of each species presenting its surfaces to the solvent action of
the soil water; and on account of hydrolysis of the solutes this solvent action
is continuous. The paper strongly supports the previous work of the Bureau
of Soils which has been so much criticised, often on a@ priori grounds.—C. R. B.
Non-infection by rusts.—Erysiphe graminis has a number of biologic forms
which are confined to special hosts. Thus conidia from the form on wheat
will not infect barley and that on oats will not infect wheat. SALMon’? has
recently shown that the reason of the non-infection is not due to inability on
the part of the conidia to germinate, but because the haustoria cannot establish
relations with the cells of the host plant.—B. M. Davis.
Endoparasitic adaptation —SALmoNn’? shows that Erysiphe haar adapts
itself readily to an endophytic life. When spores are sown on oats
or barley the mycelium ramifies in the intercellular spaces a haustoria are
abundantly produced. Conidiophores develop profusely and perfect conidia
where they arise on a free surface; and they even break through a weak barrier
when they develop in intercellular spaces.—C.
Greening of seeds.— Ernst?" finds that during the ripening of the fruit of
Eriobotrya japonica the seeds become green, quite independent of light, by reason
of the greening of the amyloplasts. The process begins at the plumule of the
17 GUILLIERMOND, A., Remarques sur la karyokinése des Ascomycttes. Ann.
Mycol. witli pls. IO-12. 1905.
18 CAMERON, F. K., and Bett, J. M., The mineral constituents of soils. U. S.
Dept. Agric., me Soils Bull 30. pp.
19 SALMON, E. S., On the stages = paige’ reached by certain biologic
5
forms of Erysiphe in cases of non-infection. New Phytol. 4:217. 1905. pl.
20 SALMON, E. S., On
under cultural conditi tions.
21 Ernst, A., Das mesa: der pei yon Eriohbotrya japonica.
a ce agrees ae by Erysiphe graminis DC.
hae y. Soc. London B. 198:87~-97. pl. 6. 1905.
Beihefte Bot.
a
Centralbl. r9™: 118-130. pl. 2. 1905.
306 : BOTANICAL GAZETTE [APRIL
embryo and progresses from this region to the inner and outer faces of the coty-
ledons. Complete greening, however, only follows illumination.—C. R. B
The nucleus and secretion.—In the nectar glands on the stipules of the Vicia
Faba, according to STOCKARD,?? the nucleus does not give out granular material
directly to the cytoplasm, but it transmits a substance which results in the forma-
tion of granules. Changes which occur in the cytoplasm during secretion seem to
be controlled by the nucleus—CHARLES J. CHAMBERLAIN.
Black rot of cabbage.—HarpDING, STEWART, and PrucHa’s find much of the
cabbage seed in the market contaminated with Pseudomonas campestris, which may
survive and become a source of infection to seedlings. They advise sterilizing
seed by soaking for fifteen minutes in HgCl. 1:1000, or in formalin 1:240.—
C.K, B.
Movement of diatoms, etc.—Jackson suggests? that the evolution of oxygen
is the true cause of movements of diatoms, desmids, oscillaria, nostoc, etc. He
has been able to imitate the movements by those compressed tablets and bits of -
aluminum of proper shapes which evolve gas—C. R. B
Anatomy of Claytonia—A study of this genus by THEo. Hot forms one of
the Memoirs of the National Academy,?5 where it may be overlooked by botanists.
It contains some of the accumulating details which a master hand must some day
correlate.— C. R. B.
Apothecia of lichens —Gertr. P. Wotrr?® through some studies on the
development of the apothecia in a number of lichens argues against LINDAU’s
terebrator theory of the function of the trichogynes in lichens——B. M. Davis.
Intercellular ducts.—The intercellular spaces in the cotyledons of Legumi-
nosae function at the beginning of germination as conducting canals for aleurone
which becomes dissolved and diffuses through them.?27—C. R. B.
Mustiness.—The peculiar musty odor acquired by damp straw or corn is
due, according to Roussev,?® to the oospora form of oe Dassonvillei
and not to other of the fungus flora found thereon.—C. R. B
22STOCKARD, CHAS. R., The structure and cytological aegis aaa RY:
secretion in the nectar ne of Vicia Faba. Science 21: 204-5. 19
23H ARDING, H. A., STEwart, F. C., PrucHa, M. J., Vitality of be cba black
rot germ on cabbage seed. N. Y. Agr. Exp. Sta. Bull. 251: 177-19 05.
24JacKsoN, D. D., Movements of diatoms and ee microscopic perine Jour.
Roy. Mic. Soc. 1905: 554-7.
2SHOLM, THEO., ee a morphological and anatomical study. Mem. Nat.
Acad. Sci. 10: 27-37. pl. 7 1905.
26WoLFF, GERTR. P. aa zur Entwicklungsgeschichte der ha awe
thecien. Flora 95:31.
27JOFFRIN, H., io BUNT des méats intercellulaires pees les cotylédons
des Légumineuses au début de la germination. Rev. Gén. Bot. 17 : 421-2. 1905-
28BrocQ-RovssEv, Contributions a l’étude des causes qui provoquent l’odeur de
moisi des grains et fourrages. Rev. Gén. Bot. 17: 417-420. 1905.
nm
Sincere crcanesrticicenns
——
~
f
NEWS.
PRoFEssor J. C. ARTHUR spent the greater part of January at the New
York Botanical Garden in a study of Uredineae.
Proressor B. M. Duccar has been spending the winter in research at the
Botanical Institute at Montpellier, directed by Professor CH. FLAHAULT.
THE Bulletin de l’Académie Internationale de Géographie Botanique an
nounces the limitation of leading articles to thirty-two pages. We hope the
movement will become general among journals. By proper condensation an
author can say all he is entitled to say on one subject in such a space.
Dr, Jesse M. GREENMAN spent some six weeks in Yucatan and adjacent
Mexico collecting plants for the Field Natural History Museum, of whose her-
barium he is assistant curator. He had a violent attack of malarial fever which
interfered seriously with his work, but he has returned in good health and with
fair collections.
LAST SUMMER after lousy the Vienna Congress, Professor GrorGE F.
ATKINSON spent some time in the vicinity of Nice, Paris, and especially in the
Jura mountains in the vicinity of Pontarlier, studying the fleshy fungi. He
collected over 300 species, made photographic studies, and preserved material
for morphological investigation.
THE VIENNA ConcREss nominated as presidents of the Committee of Organ-
ization for the Brussels Congress of 1910 Lfo ERRERA and URAND. On
account of the lamented death of Professor ERRERA the Association internation-
ale des botanistes has named Senator Count Osw. DE KERCHOVE DE DEUTER
GHEM <s his successor. M. Emite” DE, WILDEMAN has been made general sec-
retary.
Mk. J. B. Extis, whose taxonomic work on North American fungi is known
the world over through his numerous publications and the important sets of
exsiccati issued by him and Mr. EveERHARD?, died at his home in Newfield,
New Jersey, December 30, 1905. A biographical sketch of Mr. Exzis was
published in this journal in November 1890. His herbarium and library have
been for some years the property of the New York Botanical Garden.
PROFESSOR WILLIAM WHITMAN BAILEY will retire from the faculty of Brown
University at the close of the present academic year. He has been connected
with the University for nearly twenty-eight years, twenty-five of them as pro-
fessor of botan’. For some years he has been suffering from ill health and
feels it wise not to carry longer the burden of regular classroom work. Yet
he will retain close connection with the University, and advise in many of its
affairs.
397
308 BOTANICAL GAZETTE [APRIL
A NEW JoURNAL, Le Bambou, has been established by JEAN HovuzEAU DE
Lrenare, Ermitage, Mons, Belgium. The English part of the trilingual pros-
~ pectus contains some interesting information for our readers, and is at the same
time so amusing in its construction and spelling that we reprint part of it.
“Our aim is the facility for botanists and lovers of Bamboo of communi-
cating their studies and desiderata, and exchanging their observations. We
claim likewise as design to let better know the horticultural value of these plants
and, giving information on the process of culture and on the places from where
they kan be obtained, to spread as much as possible their use in parks and gar-
dens. Each number shall contain: 1™ a technical part, 2¢ a practical part.
“The technical part for wohm we can rely on the cooperation of distin-
ghuished botanists wohm names are know from a long time, shall comprise the
description of new or little known species, with plates, or cuts, the critical exami-
nation of the nomenclature and synonymy and the bibliography.
“These studies shall compose an ensemble which will become a vade mecum
mere to all lovers of Bambusaceae who whish to make serious study. .
“Briefly, the classification of Bamboo is still on many points scnareiihat
preienat and the principal -design of the technical part is to cooperate to its
perfecti
“We: sail attend with much care to the bibliography: it shall contain not
only a list as complete as possible and up to the day of all the works relating to
amboo in any way; but we ask from our readers to insert in our ‘letter box’
their demands of books. . . . .
“We hope that our correspondents will be so good as to communicate the
tittles of the books on this matter within their knolege.
“The practical part shall contain such advices of culture our essais, began
since Ee entitle us to
e will smarty review hee plants noting the peculiarities distin-
penta each of them. We will fix the culture, the value, the rational arbori-
cultural use, the edurance to the inclemency of the weather of each of them.
“We will offer in our pages the largest hospitality to the discussions our
subscribers and correspondents could wish to hold out, leaving to them the
whole responsability of their propositions.
“Our letter box will allow everyone to ask questions and to send responds to
offer or to sollicit plants in exchange or to points outs plants they desire to buy.
In short we will endeavor to become the mediator of all the loevers of Bamboo.
“We will print “notices not only in french, but in latin, english, german
italian and esperanto, with the faculty of joigning, as the case may be a trans-
lation or a summa
“We wanted that the first number should be entirely of our one penn; not so
much to definite the way we wish to adopt, but to sustain alone the responsa-
bility of the beginning and to free from all responsability = Ris coprapnis
who spontaneously offered their instant collaboration. .. . .
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CONTENTS
NEW AND NOTEWORTHY WESTERN PLANTS. III. A.D. E. Elmer - - - - 309
SOME LITTORAL SPERMATOPHYTES OF THE NAPLES REGION. J. Y. Bergen - 327
NEW AND NOTEWORTHY NORTH AMERICAN SPECIES OF TRIFOLIUM —_
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—~
VOLUME XLI NUMBER 5
DOTANICA“E GAZETTE
MAY, 1906
NEW AND NOTEWORTHY WESTERN PLANTS. III."
A. D. E. ELMER.
’ Phacelia acanthominthoides, n. sp.—An annual or biennial,
2 to 5°" high or higher: stems many, profusely branched from the
base, erect or reclining, cinereous: leaves at least in the mature
plants all cauline, alternate, usually subtending the branches, those
from near the base 1o™™ long, pinnately 3 to 5-lobed or toward the
apex only pinnatifid; the pubescent petiole almost equaling the
blade proper; lobes hispidly strigose on both sides, 1°™ long or less,
margins with few much-rounded teeth; upper leaves finer dissected
and without petioles: inflorescence ample, in widely branched scir-
poid racemes; flowers bluish, upon short pubescent pedicels: the 5
sepals pubescent, 4™™ long, 1.5™™ wide, linear-oblong, very obtuse,
united at base, much exceeded by the flower: corolla 8™™ long;
petals 5, very short, obovate, lateral nerves extending from the middle
basal primary ones: stamens 5, exserted, inserted near the corolla
base and alternating with the segments; anthers elliptic, 1™™ long,
versatile; filaments glabrous, slender, 8 to 10™™ long, with minute
hyaline appendages at base: ovary ellipsoid, pubescent; style per-
sistent, 7™™ long, cleft nearly to the base, the united portion sparsely
pubescent; stigmas minute, terminal: herbaceous sepals of the mature
capsule 8™™ long, 4™™ across the widest part, ovate, acute, sub-
coriaceous, with ciliate margins, strongly 1-nerved with prominent
reticulations: capsule 2-valved, sessile, 4-seeded: seeds dark brown
2.5™™ long, oblong, triangular, pitted.
* The first four new species have been in manuscript more than two years, and
the types are in the herbarium of Stanford University.
59
310 BOTANICAL GAZETTE [MAY
Type specimen collected in May 1903, by Miss Laura M. Lathrop at Her.
nandez, San Benito County, California.
This species can be distinguished readily by its reticulately nerved, broadly
ovate, and ciliate mature calyx lobes, not unlike the bracts of Acanthomintha
ilicijolia Gray.
Trichostema rubisepalum, n. sp.—Erect annual, 2 to 3° high:
stems chiefly branched from near the base, the branches usually in
pairs and ascending, soit pilose and glandular, the lower ones becom-
ing reddish: leaves cauline, opposite, entire, subsessile, linear-
lanceolate, pilose on both sides and finely glandular, acute or acumi-
nate, 2°™ long or longer, about 5™™ wide: inflorescence in axillary
short-pedunculate cymes; flowers blue, solitary, on 2 or 3 glandular
pubescent pedicels, subtended by linear bracts: calyx united below
the middle, bristly pilose and somewhat glandular, about 6™™ long; _
the 5 subequal segments straight, acute, ultimately turning to a pink
or light purple: corolla exceeding the calyx, 7™™ long, curved,
pilose, throat oblique; its segments also pilose, thin, obscurely
bilabiate; upper lip somewhat shorter and 2-segmented; lower one
with 3 segments: anthers much exceeding the corolla, in two pairs
of unequal lengths; filaments curved, equaling the tube, slender,
glabrous, apparently adnate to the entire length of the thin corolla
tube; anther cells united toward the apexonly, attached dorsally
to the filament, ovoid, the base ultimately much spreading: style
glabrous, filiform, recurved, equaling the shorter stamens and insert-
ed in the depression of the ovary lobes; ovary short pubescent,
deeply 4-lobed: seeds amphitropous.
pe specimen collected by Miss Laura M. Lathrop at Hernandez, San Ben-
ito County, California, August 1go2.
This is closely related to T. laxum Gray, but distinguished by its lang pilose
and glandular pubescence, sessile or subsessile leaves, and by its usually pilose
corolla. The tips of the sepals soon turn red.
Collinsia Hernandezii, n. sp.—Annual, 10 to 20°™ high: stems
branched from the base, central ones erect, the outer reclining, soft
yellowish pubescent, glandular: leaves cauline, opposite, oblong to
oblanceolate, the larger ones 4°™ long, 1.5°™ wide, apex obtusely
rounded, gradually tapering at the base to a 1°™ long pubescent
petiole, margins entire, short and dirty glandular pubescent on both
sides, rather thick, the 3 to 5 obscure nerves parallel; upper leaves
ES os eee
1906] ELMER—NEW WESTERN PLANTS 311
becoming bract-like: flowers large, widely scattered along the spicate
racemes, half nodding upon short glandular pubescent peduncles,
subtended by leaf-like bracts: the 5 distinct sepals glandular pubes-
cent, 5™™ long, 1.5™™ wide at base, acuminate: corolla bluish, 2°™
long, strongly bilabiate, saccate at base, with gibbose throat; lower
lip obscurely 3-lobed, the middle lobe longer and prominently con-
duplicate; upper lip ascending, shorter, and broadly bilobed: fertile
stamens 4, equally inserted upon the tube near thé base, longer pair
1.5°™ long, shorter pair 2™™ less, jointed and papillate at base;
filaments winged, downwardly recurved, glabrous or the longer ones
glandular above the middle; fifth stamen represented by an oblong
membranous pouch on the lower portion of the corolla tube; anthers
2-celled, round or reniform, cells united at apex: ovary soft pubes-
cent and finely glandular; style usually straight, thick, about equal-
a the stamens, sparsely glandular toward the base.
specimen collected by Miss Laura M. Lathrop at Hernandez, San Ben-
ito ce California, June 1903.
Its habit and pubescence is that of C. Greenei Gray, but the leaves and flowers
are different.
Fritillaria succulenta, n. sp.—Stems glabrous, erect, simple, 2 to
3°" high, from a bulb of fleshy scales: basal leaves ascending, 5 to
10°™ long, 2 to 3°™ wide, in pairs or in whorls of three, elliptic-oblong,
obtuse, succulent and covered with a bloom; cauline ones few, erect, ©
alternate or the lower in pairs, lanceolate, also fleshy and glaucous:
flowers solitary or on the larger plants in racemes of three, nodding;
peduncle subtended by a leaf-like bract, glabrous, 1 to 2°™ long:
perianth campanulate, 2 to 3°™ long, wider than that when spreading;
the 6 segments 2 to 3°™ long, obtuse, oblanceolate to obovate, glab-
rous beneath, purple, entire, margins at apex yellowish, numerously
striate with darker purple and the upper surface pulverulent or
obscurely crested: stamens 6, inserted upon the base of the segments,
included; anthers 3 to 4™™ long, elliptic-oblong, versatile, extrorse;
filaments 8™™ long, glabrous, more or less expanded toward the
base: style 1°™ long, glabrous, cleft into three segments half way
down, the recurved segments subcompressed and bearing terminal
stigmas; ovary smooth, truncate at apex.
Type specimen collected in April 1903 by Miss Laura M. Lathrop, at Her-
nandez, San Benito County, California.
.
312 BOTANICAL GAZETTE [MAY
Its leaves are quite thick and fleshy, and are usually glaucous on both
sides.
Sanicula serpentina, n. sp.—Low spreading biennial or perennial
herb, from slender rootstocks, 2 high or less, wholly glabrous and
frequently somewhat glaucous, a rich brown color when cured: stems
chiefly branched from the base, the central one erect, the marginal
ones ascending: leaves mostly from the base, subtending the branches,
the radical ones upon membranously flattened 3-nerved petioles 2°™
long; blade proper 2°™ long or longer, ovate in outline, 3°™ across
the base, pinnately divided into laciniate lobes which are again
divided into slender acuminate usually somewhat recurved and
sharply pointed segments: inflorescence branched from near the
base, long-pedunculate; involucre of sessile leaf-like bracts; heads
3 to 5, the peduncles of the lateral heads usually much shorter at
least when in flower, densely flowered, about 4™™ in diameter;
involucels of entire lanceolate bracts slightly shorter than the yellow
flowers; marginal flowers sterile, pedicelled, the fewer inner ones
sessile and fertile: calyx 5-toothed: petals 1-nerved, quite broad
across the middle, the setaceously acuminate apex strongly inflexed
and emarginate on its bend: stamens incurved near the apex;
anthers broadly elliptic, o.5™™ long: ovary with uncinate prickles;
styles 2, slender, recurved, each persistent from the inner face of the
stylopodium: fruit not observed.
ype specimen no. 4498, collected in April 1903 near Portola, San Mateo
County, California.
This form is nearest related to S. laciniata H. and A., but the latter is a much
more rigid herb, with coarser, broader, spinosely toothed leaf divisions; and
with the bracts of the involucels often 3-parted or at least 3-nerved from near
the middle. It was discovered on serpentine rocks near Searsville Lake, of the
chaparral formation.
Trifolium bicephalum, n. sp.—More or less tufted, from an
annual root: stems slender, 8 to 18°™ long, decumbent or the outer
ones prostrate, rather numerous from the base, rarely branched,
sparsely pubescent: leaves both radical and cauline, the basal ones
somewhat smaller and more numerous, with slender flexuous pubes-
cent petioles 2°™ in length; stipules adnate, 6™™ long, membranous,
strongly nerved, subglabrous or finely ciliate along the edges, termi-
nated by two setae 2™™ long; leaflets 7™™ long, 4™™ wide, soft pubes-
1906] ELMER—NEW WESTERN PLANTS 313
cent with brownish hairs, obovate or truncate and usually emarginate,
entire or obscurely dentate above the middle, with prominent ascend-
ing nerves beneath: peduncles pubescent, equaling half the length
of the stem or branch, subflexuose above the middle; bicephalous
heads terminal, sessile, unequal in size, each subtended by a sub-
sessile trifoliate leaf with broad ovate stipules; involucre none: calyx
densely pubescent, 4™™ long including the 2™™ sharply acuminate
teeth: corolla exceeding the calyx teeth by 1 or 2™™, hyaline and
united with a stamineal tube below the middle; upper lip whitish,
obovately rounded and surrounding the lateral lobes or wings; lateral
lobes oblongish, slightly shorter than the banner, obtuse or acute
apical portion nearly white, the middle portion purple, the basal
portion hyaline and with an auricle; keel obtuse, shorter than the
wings, purplish: anthers very small: ovary glabrous, 2-ovulate;
style glabrous, equaling the stamens, terminally recurved, bearing
a capitate stigma.
Type specimen no. 4812, collected at San Pedro, San Mateo County, Cali-
fornia, May 1903.
his species comes nearest to the so-called Californian T. Macruei H. and
A., but is a much smaller and more prostrate clover, with leaves distinctly obo-
vate and emarginate. It forms dense prostrate mats on a high promontory near
the sea.
Eriophyllum Greenei, n. sp.—A cespitose perennial, from a
woody base: stems many from the crown, lanate, branched above
the middle: leaves numerous on the sterile stem, the lower ones
opposite, the upper ones alternate, petiolate, triangularly ovate in
outline, 2 to 3-pinnately divided, lanose on both sides; the segments
short, blunt, very narrow, with incurved margins inclosing a dense
matrix of woolly hairs; petiole about equaling the blade, as broad
as the segments with edges incurved: heads heterogamous, solitary,
terminating the leafy branches, ovoid, 1°™ broad; peduncle white
tomentose, without bracts; involucre quite rigid, cup-shaped, densely
lanate, united at base; bracts in one series, acute, 1o™™ in length:
ray flowers light yellow, 15™™ long including the achene, pistillate;
tube 2™™ long, pubescent, ligule 8"™ long, 3™™ wide, obovate or
oblanceolate, many-nerved, apex obscurely 3-toothed: style arms
1™™ long, narrowly flattened, obtuse: receptacle obscurely pitted,
somewhat raised and subconic: disk flowers perfect, very numerous,
314 BOTANICAL GAZETTE [MAY
6°™ long with the achenes: tubular corolla yellow, sparsely pubescent,
terminated by 5 obtuse segments: anthers 1.5™™ long, with apex
triangularly ovate, bases obscurely auriculate; filaments barely as long,
inserted upon the middle of the tube: style arms flattened, bearing
small capitate stigmas: achene brownish black, 3™™ long, usually
curved and attenuate from the base, subcompressed or 4-angled, edges
ciliate: pappus persistent, less than 1™™ in length, of unequal paleae.
Type specimen no. 4335, collected in the Mocho Creek Canyon, Alameda
County, California, May 1903.
Tt is intermediate between E. arachnoideum Greene and E. caespitosum Dougl.,
but sufficiently distinct from either. Named for Professor E. L. GREENE
Navarretia Abramsi, n. sp.—Densely lanose herbs, about 6°”
high: stems solitary or several from the base, rigidly erect, chiefly
branched from the middle; branches rather stout and straight,
ascending, terminated by solitary comparatively large heads: leaves
mostly subtending the heads, the larger ones 2°™ long, 1 or 2-lacini-
ately divided, soon becoming dry and brittle; the lobes very narrow,
becoming glabrous, usually recurved and terminated by a fine sharp
point: heads turbinate, the larger ones 1°™. across at top, densely
surrounded by a matrix of lanate hairs, 6 to 10-flowered, the sub-
tending bracts similar to the leaves but smaller: corolla easily sepa-
rating from the base, 8™™ long including the 3™™ long segments,
bluish or nearly white, glabrous, hyaline, conspicuously nerved; seg-
ments 5, subequally divided, linear spatulate, entire or finely dentate _
at apex: stamens 5, barely exceeding the throat of the corolla, sub-
equal in length, filaments threadlike, subequally inserted upon the
tube 2™™ below the throat; anthers ovate or elliptic, 1™™ long, apex
obtuse, base sagittate: style persistent, glabrous, minutely lobed at
apex: calyx of the mature capsule 6™™ long, divided nearly to the
base; the sepals straight and erect, linear, hyaline below the middle,
held intact by the hairy matrix; the upper part of the sepals glabrous,.
foliaceous, and acuminately pointed: capsule triangular, when
mature easily falling out from the persistent calyx, straw-colored,
smooth and shining, 4™™ long, 1.5™™ in diameter, apex pointed,
3-celled, loculicidally dehiscent: seeds solitary in each cell, subterete,
3™™ long, brown and very hard, with a gelatinous cover which
* adily dissolves in water.
{
ol.
1906] ELMER—NEW WESTERN PLANTS 315
Type specimen no. 4586, collected on Black Mountain, Santa Clara County,
California, July 1905.
It is a very late summer-flowering annual, chiefly confined to dry gravelly
soil immediately bordering thickets of the Californian chamiso (Adenostoma jasci-
culatum H. and A.). Named for Mr. L. R. ABRAMS, a former student of botany
and classmate at Stanford University.
Ribes Stanfordii, n. sp.—A rigidly branched shrub, 1 to 1.5™ high,
nearly as broad: bark on the younger branches light brown, becom-
ing grayish white with age, thin, separating into shreds; branchlets
subtended and protected by 3 spines, very short and rigid; spines
about 1°™ long, straight, shining brown, divaricate, distinct, the
middle ones usually longer, exceeding the axillary leafy branchlets;
branchlets terminated by 1 to 3 small tufts of leaves, subtended by
diminutive spines, each tuft provided with a subwhorl of 3 to 5 leaves
and terminated by a small inflorescence of 1 to 3 flowers: leaves
orbicular, 8™™ long including the finely glandular pubescent 2 to 3"™
long petiole, deeply cleft into 3 segments, soft pubescent on both
sides, rather thick, the segments usually terminated by subequal
obtusely rounded teeth or lobes, obscurely 3 to 5-nerved; petiole
gradually expanded at base into the adnate stipules: flowers 3,
upon a short and pubescent peduncle, each separately inserted and
sessile, subtended by conspicuously broad pubescent bracts: calyx
about the ovary densely pubescent, 3™™ in diameter, its tube 2"™ in
diameter, less pubescent, about 2™™ long, the 5 segments exceeding
the corolla by 1™™, triangularly obtuse, puberulous on the outer
surface, 2™™ long, yellow, rotate or much reflexed: corolla deeper
yellow, inserted upon the calyx throat and alternating with the seg-
ments, straight, obovate, 2™™ long: anthers 5, inserted upon the
calyx throat and opposite the segments, equaling the corolla; fila-
ments glabrous, flattened, 1.5™™ long: style erect, subterete, slightly
exceeding the stamens: anthers ovate, obtuse at apex, light yellow,
truncate or only obscurely lobed at base, 1.5™™ long, 1™™ wide at
the base: berry yellow and pubescent at least when young.
Type specimen no. 3958, collected on Mt. Pinos near Griffin’s Postoffice,
Ventura County, California, July 1902.
It was discovered in open pine regions in the vicinity of cliffs and rocky out-
croppings at the summit. Not common. Distributed as R. nubigenum Mc-
Clatch
316 BOTANICAL GAZETTE [MAY
Pedicularis Dudleyi, n. sp.—Perennial herbs, 2 to 3°" high,
usually from a branched caudex: stems solitary from each of the
scaly crowned caudices, the central ones erect, the outer ones ascend-
ing, simple or sometimes branched, not exceeding the basal leaves,
lanose especially toward the base, more or less curved; basal bracts
brown, lanceolate, entire, glabrous, marcescent: leaves chiefly from
the base of the stem, alternate, the uppermost at about the middle
of the stem but not exceeding it, the lowest ones longest and some-
what decumbent, lanceolate in outline, the larger 25°™ long, 6°™
wide; leaf segments about to pairs, subglabrous or short pubescent
on the nerves, membranous; lower pairs distinctly petiolate, the
upper pairs not only sessile but broadly united, those along the
middle largest; each lobe ovate or oblong in outline, 3°™ long,
2°™ wide, cleft into irregular lobes or merely dentate, the margins
unequally serrate or dentate, its teeth sharply pointed: inflorescence
spicate, densely flowered, at most 5°™ long and 3°™ in diameter,
upon peduncles equaling half the length of the stem, usually erect
but frequently somewhat curved; bracts subtending the flowers,
foliaceous, serrately toothed, the upper ones equaling the flowers,
the lower ones much exceeding them: calyx 1°™ long, unequally
5-cleft, the segments acute and obscurely toothed toward the apex,
densely lanose on the exterior: corolla 2°™ long, the narrow tubular
part half that in length, glabrous; upper lip conduplicate, slightly
notched at apex, pink or whitish, much protruding and arched; lower
lip subequally 3-toothed: stamens equal; filaments glabrous; anthers
broadly elliptic, attached to the basal dorsal side, the cells connate
and rounded at apex, the base not united, acute: style filiform, sub-
persistent, thicker and more or less flattened toward the apex, con-
spicuously recurved and protruding from the upper lip; ovary glab-
rous, dark brown, 2-celled, flattened, acuminately pointed: capsule
coriaceous, 12™™ long, 7™™ wide, acuminately terminating in an
upwardly curved point: seeds about 4, black when mature, pitted,
subterete or obscurely angular. 3
Good flower and fruit of this type specimen, no. 4289, was collected in May
and June 1903, at Iverson’s Ranch on the Pescadero Creek, San Mateo County,
ornia.
Only known from this locality, where it is rare and confined to the deep shade
of Sequoia sempervirens. Its proximity to a camping ground endangers its exist-
1906] ELMER—NEW WESTERN PLANTS I
317
ence. This denizen of the Santa Cruz Mountain redwoods is named in honor
of Professor W. R. Duprey of Stanford University, who first discovered it.
Orthocarpus longispicatus, n. sp.—A profusely branched decum-
bent biennial or perennial, forming rather dense mats: stems slender,
elongated and distantly branched, often 1™ in length, usually pubes-
cent with soft glistening white hairs: leaves alternate, evenly scat-
tered, sessile, membranous, puberulent on both sides, or with glisten-
ing hairs on the margins and along the 3 obscure nerves, cleft into
2 pairs of strap-like segments, the middle one longest: inflorescence
spicate, very long and usually curved; bracts not exceeding the
flowers, 5 to 7-laciniately cleft, the obtuse apices light red: calyx
4-cleft, soft pubescent, equaling the corolla, with colored tips: tube
of the corolla 2°™ long, externally pubescent, gradually expanded
from the constriction above the ovary; upper lip 1™™ longer than
the lower one, rather straight, apex obtuse, finely pubescent on the
lower surface, margins soft and hyaline; lower lip with 3 obtuse finely
pubescent teeth which bear moderate sized sacs: stamens 4, inserted
upon the middle of the corolla tube, the lateral pair shorter, the upper
pair nearly equaling the galea and enclosed by it; filaments hyaline,
linear-flattened, glabrous; the upper anther cell usually somewhat
longer and shedding its pollen before the lower one: style persistent,
glabrous, much exserted, thickened or expanded toward the base of
the capitate flattened or obscurely lobed stigma; ovary oblong,
truncate at the apex: capsule 10o™™ long, smooth, loculicidal:
seeds numerous, lenticular, with broad reticulate wings.
ype specimen no. 4938, collected in July 1903, at Point Reyes, Marin
County, California.
It was quite abundant among the pickle weed (Salicornia ambigua Michx.),
along edges of brackish water. Distinguished by its long decumbent fragile stems
and branches, numerous leaves, and elongated densely flowered spikes.
Godetia lanata, n. sp.—Erect annual, 3 to 6% high, single or
branched from near the base, quite rigid; mature stems shining,
straw-colored, scaling at base into membranous shreds, with ascend-
ing branches from or above the middle; the younger branches yellow-
ish tomentose: leaves cauline, lower ones soon falling, alternate and
clustered, sessile, very unequal, cinereous on both sides, semicoriace-
ous, lanceolate or linear-oblong, equally tapering at both ends,
acute, the larger ones 5°™ long, 15™™ wide, midnerve quite promi-
318 BOTANICAL GAZETTE [MAY
nent beneath, lateral ones obscure: inflorescence short, spicate or
subcapitate, terminating the branches, usually densely flowered, 3 or
4°™ in diameter; buds erect, from the terminal central axis; flowers
easily separating from the ovary, subtended by strigose lanceolate
acuminate bracts, subsessile: calyx tube obconic, 3 to 4™™ long,
lanose pubescent; its 5 equally pubescent segments 12™™ long,
acuminate and ultimately reflexed: corolla and stamens inserted
upon the rim of the calyx throat; petals straight, thin, pink, broadly
obovate in outline, 6™™ long and wide, irregularly or obscurely 3-
toothed, the middle tooth acute, usually the larger: stamens 8, in
2 series, those alternating with the petals nearly equaling them,
those opposite the petals barely more than 1™™ long; filaments glab-
rous, compressed, broadest at the base; anthers introrse, basifixed,
those of the upper series twice as long as those of the lower: style
glabrous. or with a few long hairs, barely equaling the stamens,
bearing an obscurely 4-lobed stigma; ovary densely and persistently
lanose pubescent, upon short thick pedicels: capsules subtended by
leaf-like bracts longer than themselves, loculicidally dehiscent from
the apex, straight, erect, lanose, subsessile, apex truncate, nearly of
the same thickness throughout, subterete or only slightly 4-sided,
8-costate, 4-celled, 4-valved: seeds numerous, in single rows, dark
brown, subterete or cubical, a little pointed at one end.
ype specimen no. 4376, collected in June 1903 at Bardins railroad switch,
Monterey County, California.
This characteristic species was found quite plentiful on the sandy plain between
Monterey and Castroville, and is quite variable in the density of its pubescence
branching habit, and size of leaves.
Pentachaeta laxa, n. sp.—A lax very much branched annual, 1 to
2 or 3™ high: stems branched from the base, softly but sparsely
pilose: leaves in pairs, subtending the branches, sessile, linear, gradu-
ally tapering from the base, the larger ones 3°™ long, 2™™ wide, very
thin, sparsely pilose on both surfaces: heads terminal, heterogamous,
turbinate, 6™™ long, about 9-flowered; the peduncle ascending, 1 to
4°™ long, pilose, somewhat thickened toward the apex; involucral
bracts 3 to 5, persistent, flat, acute, oblong, scantily pilose on the
exterior, the reticulate nerves quite prominent, equaling the flowers,
more or less membranous: receptacle pitted: each of the ray flowers
|
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1906] ELMER—NEW WESTERN PLANTS 319
subtended by an involucral bract, yellow, caducous, pistillate, tube
1™™ long, bearing a broad 1™™ long notched ligule: style arms
exceeding the ligule, recurved, acute: disk flowers perfect, tubular,
light yellow and caducous, 2™™ long, the upper half inflated, bearing
4 obtuse teeth: anthers well included, o.5™™ long, rather broad,
bases truncate, each with a very prominent apical appendage; fila-
ments thread-like, scarcely longer than the anther: style barely
exceeding the corolla, its arms subcompressed, recurved,
short, obtuse or truncate at apex: achenes subterete, 5™™ long, black
when mature, finely rugose, dotted with sessile yellowish brown
glands subtended by short setae; pappus of 2 or 3 paleaceous scab-
rous awns, usually persistent.
ype specimen no. 4437, collected in May 1903, on Cedar Mountain of the
Mount Hamilton Range, Alameda County, California.
This distinct species inhabits a steep shaded ravine of fertile soil, and forms
a tangled mass with its numerous decumbent branches. Not observed elsewhere,
and evidently very rare.
~»Nemophila Fremontii, n. sp.—Delicate annual: stems procum-
bent or prostrate, branched, subglabrous or sparsely retrorsely pubes-
cent, 10 to 30°" in length: radical leaves similar to the lower cauline
ones, frequently forming a rosette, 3°™ long including the 1.5°™ long
strigose petiole; blade membranous, ovate or oblong in outline,
usually oddly pinnate with two pairs of nearly divided lobes or
the uppermost merely sinuate, sparsely pubescent on both sides,
paler beneath; the lobes nearly as broad as long, rounded, finely
mucronate: flowers solitary, upon slender flexuose usually recurved
2°™ Jong peduncles which are clothed with retrorse bristles: calyx
persistent, campanulate, 3™™ long, the basal one-third united, equal-
ing or exceeding the corolla, pubescent with fine bristle-like hairs;
sepals oblong, obtuse or acutish, foliaceous, with a short blunt
recurved appendage from each sinus: corolla white, urn-shaped,
its lobes becoming reflexed, at most 3™™ long, 5-cleft, the basal
appendages quite obsolete; petals hyaline, ovate to oblong or obovate,
obtuse, sparsely ciliate on the edges above the middle: anthers 5,
alternate with the petals, erect, quite a little shorter than the corolla;
filaments inserted half way down on the corolla tube, glabrous, 1™™
in length; anthers 0.3™™ long, comparatively broad, apex obtuse,
320 BOTANICAL GAZETTE [MAY
base subcordate: ovary sessile, densely pubescent: style 1™™ long,
cleft into 2 recurved arms, terete, glabrous, persistent; stigma ter-
minal, capitate: capsule 4™™ in diameter, globular, sparsely ciliate:
seeds compressed, carunculate.
Type specimen no. 4991, collected in May 1903, on Fremont’s Peak of the
Gabilan mountains, San Benito County, California.
It was observed only at the very summit of the peak, among the moss-covered
rocks,
Monardella franciscana, n. sp.—A sprawling suffrutescent peren-
nial: lower stems reclining on the ground or in dense herbaceous
thickets, woody, one or more meters long; the leaf-bearing upper
stems usually clustered, herbaceous, woolly pubescent, 2 to 3° long,
erect or decumbent near the base: leaves opposite, mostly fascicled,
very variable in size, densely woolly pubescent on both sides especially
beneath; the larger upper ones 2 to 3°™ long including the 5™™ long
petiole, 2°™ wide near the base, broadly ovate, entire or with a few
obscure teeth, the edges recurved; the lower or axillary ones sessile,
lanceolate to elliptic-obovate: inflorescence capitate; heads densely
flowered, terminal, rarely more than one, 3 to 4°™ in diameter, sub-
tended by a whorl of pubescent leaf-like bracts equaling or exceed-
ing the flowers; flowers upon stout short pedicels: calyx about 8™™
long, the marginal ones usually curved upward, conspicuously 11 to
15-nerved, silky pubescent except near the base, tubular, equally
5-toothed; the teeth acute, 1.5™™ in length: corolla blue or light
pink, funnel-shaped, the longest 2°™, strigose about the middle,
glabrous toward the base, bilabiate; upper lip 5™™ long, erect or
straight, apex 2-lobed; lower lip divided into 3 linear segments,
equaling the upper lip, usually deflexed: stamens 4, fertile, moder-
ately unequal, exserted and spreading; filaments slender, glabrous,
inserted at the corolla throat or a trifle beneath it; anthers attached
to the base, the cells somewhat recurved: style equaling the stamens,
glabrous; stigma minute, terminal; ovary glabrous, distinctly
4-lobed.
Type specimen no. 4766, collected at San Pedro, San Mateo County, Cali-
fornia, July 1903.
It was repeatedly observed in dense herbaceous growths in the ravines on
the coast from San Francisco to Santa Cruz, and is a distinct seacoast species.
a
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1906] ELMER—NEW WESTERN PLANTS 321
HorKELIA BOLANDERI marinensis, n. var.—Tufted perennial,
r to 2% high, villous pubescent: stems deflexed and more or less
decumbent: leaves pinnately divided into 5 to 8 pairs of leaflets,
subradical though often the stem bears as many as 3 to 5 leaves with-
out subtending branches; average leaflets 10o™™ long, unequally 4 to
1o-toothed; basal stipules narrowly linear, 1o™™ long, those on the
stem broader and usually 1 or more-toothed: inflorescence branched
from near the middle, corymbosely paniculate; branchlets loosely
flowered: hypanthium longer than wide, saucer-shaped; bracts
broad, 3-toothed: stamens 10, opposite the outer and inner series of
sepals which must exceed them: outer sepals shorter, spatulate or
acute at apex, pubescent on both sides; inner ones acuminately
triangular, glabrous on the inner surface: filaments glabrous, flat-
tened, slightly unequal in length; anthers introrse, short, elliptic:
petals white, narrowly spatulate or oblanceolate, about equaling the
sepals, early falling: styles many, glabrous, erect: seeds bean-shaped,
smooth. |
Type specimen no. 5039, collected in sandy soil at Point Reyes, Marin
County, California, July 1903.
This variety has been distributed as H. Parry: (Wats.) Ryd. It has the
pubescence of typical H. Bolanderi Gray, but the stems are more or less decum-
bent, basal stipules strictly linear and almost twice as long as in the species,
leaflets larger and more toothed, and inflorescence more loosely corymbose and
with larger hypanthia.
Chrysopsis arenaria, n. sp.—A wiry perennial herb, from creep-
ing and much branched roots: stems 10°" long or very much
shortened and bearing a dense rosette of leaves, erect or more often
subreclining, hirsute: basal leaves soon withering; cauline ones
many, ascending, obovate to oblanceolate, the lower ones slenderly
attenuate from the base, alternate, entire, hispidly hirsute on both
sides, ascending: heads heterogamous, terminating the stems, 1 to
3, barely exceeding the leaves, upon densely hirsute often glandular
and few-bracteolate peduncles, 15™™ in diameter, nearly that in
length; jinvolucre of several series, campanulate; bracts linear,
imbricate, acuminate, pubescent, the longer ones 107™ in lengt
and usually with pink scarious margins: receptacle flat, favose: ray
flowers in one series, pistillate, showy; ligule yellow, 4-nerved, narrow-
ly oblanceolate, entire, 6™™ long; its tube slender, 5"™ long, glab-
322 BOTANICAL GAZETTE [May
rous, expanded at base: style barely exceeding the throat, with short
appendages: disk flowers many, all tubular, equaling the pappus,
perfect, terminated by 5 acute teeth, 6™™ in length, expanded at the
base, yellow: anthers included, 2™™ long, bases obscurely auricled,
apex triangular, acute; filaménts glabrous, inserted upon the tube
one-third from the base: style much exceeding the tube, its arms
flattened, short and truncate: achenes compressed, silky pubescent:
pappus bristle-like, chiefly of two series, rusty or yellowish white,
the longer series 5™™ in length, finely scabrous, the basal one very
short, lighter colored, smooth. . ,
Type specimen no. 4556, collected at Point Reyes, Marin County, California,
July 1903. | :
It forms dense prostrate mats on the windward side of the drifting sand dunes.
Castilleia Wightii, n. sp.—A tufted perennial herb, 3°" to 1™ tall:
stems several from the crown of the root, much branched from the base
to the middle, glandular pubescent with dirty yellowish hairs: leaves
alternate, sessile, quite membranous, pulverulent on both surfaces,
or sparsely hairy along the 3 nerves, broadly linear to oblong, the
larger ones 6°™ long and 15™™ wide, mostly with one pair of linear
lateral segments from the middle of the leaf, the terminal lobe much
longer and wider: inflorescence spicate, 10 to 20°™ long, terminating
the erect corymbosely disposed branches; subtending bracts barely
equaling the flower, densely covered with glistening subglandular
hairs, 3-nerved and 3-cleft, the upper ones with colored tips: calyx
pubescent, lateraily compressed and equally cleft nearly to the middle,
the halves 2-toothed, subequaling the corolla: tube of corolla proper
1o™™ long, glabrous, saccate, its nerves prominent; galea or upper
lip longer than the tube, quite broad and membranous at base,
straight, its blunt apex rather thickened and retrorsely pubescent
on the outer surface; lower lip 3-toothed, teeth 1™™ long, obtuse,
all alike: stamens equally inserted upon the middle of the corolla
tube, the lateral pair a trifle shorter, the upper pair nearly equaling
the galea and enclosed by it; , filaments glabrous, flattened, with
hyaline margins; anther cells broadest at base, subequally attached:
style little protruding, flattened, glabrous, persistent, often recurved
near the apex, beating a capitate or obscurely flattened stigma;
ovary smooth, somewhat compressed, acute toward the apex: cap-
*%
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1906] ELMER—NEW WESTERN PLANTS 323
sule 12™™ long, ovoid, sharply pointed, coriaceous, loculicidal:
seeds many, orbicularly compressed or somewhat elongated,. with
reticulate wings. .
Type specimen no. 4761, collected on the trail between Spring Valley Lake
and San Pedro, San Mateo County, California, July 1903.
It is a uniquely distinct subglandular species easily recognized by its abun-
dant foliage, densely flowered elongated spikes, and broadly linear unequally
3-segmented leaves. Named for W. F. Wicurt, a former student of botany and
classmate at Stanford University.
Phacelia flaccida, n. sp.—Delicate annual, 3 to 6% high: stems
sparingly but widely branched from the middle, rather weak
and more or less reclining, rarely. erect, beset with stinging white
hairs: leaves few, alternate, usually subtending the branches, the
larger ones 5°™ wide and 7°™ long, ovate to elliptic in outline, very
thin and flaccid, only sparsely setigerous on both surfaces, 3 to 5-
pinnately nerved, obscurely lobulate, the margins with irregular
low roundly obtuse teeth, base rounded or obscurely and unequally
subcordate; longer petioles 4°™ in length, setose, the younger ones
cinereously pubescent: inflorescence usually dichotomously branched;
flowers secund, not crowded, upon short ciliate pedicels: calyx of +5
persistent herbaceous distinct segments, the 4 smaller ones oblanceo-
late and in the mature state 7™™ long, bristly ciliate: corolla barely
exceeding the calyx, 3™™ long, blue or pale white; the 5 segments
short-ovate and comparatively very broad: stamens well included
within the tube, alternating with the segments and inserted near the
base of the tube; anthers short- elliptic, versatile; ‘filaments nearly
equal, glabrous, usually curved above the middle, subtended by
small hyaline entire membranous folds: ovary persistent, pubescent;
style persistent, equaling the anthers, cleft almost to the base, each
division bearing a minute terminal stigma: capsule ovoid, 1-celled,
divided into 2 valves, 1-seeded in each; seeds es flattened, 2™™
long, obscurely pitted.
cimen no. 4404, collected at Wright’s Station, Santa Clara County,
Type s
California, June 1
delicate eis + was discovered in a damp deeply shaded nook on the
banks of Los Gatos Creek. Otherwise not known.
Leptosyne Hamiltonii, n. sp.—Glabrous acaulescent annual:
leaves tufted, all radical, mostly erect, rather thick, 2 to 3°™ long
324 BOTANICAL GAZETTE [MAY
including the 1 to 2°™ long flattened petiole; blade proper triangular-
ovate in outline, 1°™ long, fully as wide at base, 2 to 3-pinnately
divided; the segments not wider than the petiole and the rachis, 1™™
wide, ultimate ones short, with bluntly rounded apices: scapes
usually 1 to 3 from each root, glabrous, 10 to 18°™ long, slender,
usually somewhat curved, each bearing an erect solitary head;
heterogamous heads subcampanulate or cup-shaped, 1°™ in diameter,
a trifle less than 1°™ in height; involucre glabrous, truncate and
united at the base, in two series; bracts of lower series 5, thickish,
linear, obtuse, 47™ long, dark brown; inner series of 8 to Io striate
bracts, shining, yellowish, about 10™™ long, 4™™ wide, submem-
branous, oblong, acute: ray flowers about a dozen or more, pistillate
and fertile; tube 1.5™™ long, subglabrous; branches of stigma barely
exserted, recurved; ligule 3 to 4™™ wide, 6™™ long, elliptic to oblong
or obovate, apex round, usually with a short obtuse tooth: scales
equaling the flowers, subpersistent, linear, hyaline, obtuse, 3-nerved,
those of the central flowers narrower: central flowers crowded, per-
fect, sterile, tubular; tube 4™™ long, hyaline, cylindric, the upper
half subinflated, its 5 teeth obtuse and more or less thickened along
the margins: anthers linear, 2™™ long, truncate at base, apex trian-
gularly appendaged: style equaling the stamens, its 2 branches short,
obtuse, and flattened; achenes linear-obovoid, compressed, mar-
ginally winged, ciliate on the edges, pubescent on the sides, brown
and glabrous when mature: pappus of 2 caducous hyaline finely
ciliate membranes.
Type specimen no. 2328, collected on Mt. Hamilton, Santa Clara County,
California, April tgoo.
It was in fine flower and fruit, and grew in dry gravelly soil on a steep slope
a few hundred yards below the observatory. Since then I have failed to find
it either in this same place or elsewhere.
EUNANUS ANDROSACEUS Curran.
This southern species was originally discovered at Tehachapi, California.
From the middle western part of the state it is only known at Ben Lomond, Santa
Cruz County, where fruiting specimens were collected by Mrs. K. Brandegee in
April 1890. In July 1903 the writer found excellent flowering specimens in the
same locality, which were distributed under no. 4519. It is evidently rare and
prefers hot and dry gravelly soil of the chaparral formation. The plants were
from 1 to ro%™ high, with single or branched stems, glandularly pubescent or the
—_ ee a aE
1906] ELMER—NEW WESTERN PLANTS 325
older ones subglabrous: leaves sessile, obovate or linear-oblong, entire or ob-
scurely apiculate above the middle, apex obtuse, attenuate toward the base, 5 to
2o™™ long, the larger ones 5™™ wide: pedicels of solitary axillary flowers not
exceeding 5™™ in length: calyx 8™™ long, compressed, somewhat inflated below
the middle; the two upper parallel lobes a trifle longer than the lower and lateral
ones, with 5 short and obtuse teeth: crimson corolla little exceeding the calyx,
slenderly tubular, conspicuously constricted below the limb, pubescent on the
exterior; limb barely bilabiate, the broadly rounded segments rotately spreading:
Stamens 4, all anther-bearing; filaments slender, inserted upon the corolla at
the middle; anthers o.5™™ long, comparatively broad, widely spreading: style
glabrous, erect, bearing flattened lobed stigmas; ovary smooth, conically elon-
ated.
LUPINUS POLYPHYLLUS Lindl.
This magnificent seacoast lupine was found as far south as San Pedro, San
Mateo County, California, May 1903. At this station it grew in wet adobe soil,
reached a height of 2™, and is succulent and apparently an annual. The larger
leaves were of an enormous size, and some of the flowering spikes exceeded the
length of a man’s arm. Flowers large, dense, with blue wings and purplish
banner. Pods persistent, densely covered with soft long yellow ish-white hairs.
Distributed under no. 4444
CAMPANULA EXIGUA Rattan.
No. 4357 was collected in May 1903 along La Puerta Creek, Stanislaus County,
California. It was found on a dry mountain side composed chiefly of small
rocks, and on gravelly embankments of the creek. At first it reminded me of
a Gilia in growth and habitat. The flowers are large and showy: corolla bluish
white, at first appearing tubular, ultimately campanulate: styles wholly included,
aring 3 revolute stigmas; dilated bases of the filaments not ciliate; capsule
dehiscing regularly on the sides, 3-celled; seeds numerous, 0,75™™ long, half
as wide, compressed, shining light brown, inserted upon 3 conspicuously enlarged
placentae centrally attached to the main axis.
SALIX BREWERI Bebb.
The type of this willow was collected by W. H. BREwrr on San Carlos Moun-
tains of middle California, and described in Bot. Calif. 2:88. 1880. Since then
it has been reported from only a few localities. In May 1993, while making a
botanical trip from Livermore, California, through San Antonio and Adobe
valleys, I found it in fine fruit, and distributed it under no. 4648. It is quite
abundant along the Little Colorado, Sweet Water Creek, and on the headwaters
of the La Puerta, all of which rise on Red Mountain, Santa Clara County, Cali-
fornia. This species was at once distinguished from the other willows, and
bears a remarkable similarity to Salvia mellifera Greene
It is a shrub 1 to 2™ high, rather gracefully branched from near the base;
326 BOTANICAL GAZETTE [MAY
branchlets sparsely branched, erect, about as tough as the western leatherwood
(Dirca occidentalis Gray), cinereous pubescent: leaves terminally oe wi
sessile, linear to oblanceolate, acute, the larger ones 6°™ in length and 10
width, densely canescent beneath, upper surface green and becoming partie
except the sunken cinereous midnerve, margins entire or sometimes remotely
apiculate: fruiting spikes immediately beneath the foliage, the short peduncle
oe by 3 foliaceous bracts, curved and slenderly elongated, 5 to 7°™ long,
ro™™ in diameter, densely flowered: capsules short-canescent all over, 5™™
ovoid, acuminate; styles persistent, not longer than 1™™, 2-cleft, each segment
again divided into recurved lobes: bracts spatulate or obovate, long ciliate
pubescent; axillary gland persistent, quite prominent and cinereous.
CUPRESSUS GOVENIANA Gord.
About sixteen miles southeast of Livermore, Alameda County, California,
is a mountain commonly known as Cedar Mountain. While collecting on this
mountain during the latter part of May 1903, I did not find any true cedars, but
found near the round-topped summit quite an area densely covered with a species
of cypress. It was too late for staminate flowers, but an abundance of mature
cones was collected. The trees are mostly of a young generation, of all sizes
from 34" to 5™ high and cone-bearing, widely branched from near the ground,
the terminal portion of the stem much exceeding the shortest uppermost branches;
leaf branchlets slender, more or less spreading: leaves without conspicuous dorsal
pits, acute, shining green: cones globose, 13™™ in diameter; the 8 unequally
sized scales grayish-white on the outside, almost smooth or only with low umbos:
angular, prominently margined or winged along the lateral edges above
the middle, reddish-brown and frequently somewhat glaucous, with a resiniferous
or.
This species was distributed under no. 4487. There is some doubt as to
this determination, and its relationship to the other closely allied species is not
as yet clearly understood. These flourishing trees might have been planted at
some very early date.
MUHLENBERGIA DEBILIS Trin.
The distribution of this handsomely tufted grass is given from Texas to
southern California. In May 1902 the writer found it in abundance on a dry
gravelly hillside near Santa Barbara, California. In June 1903 a few dwarfed
specimens were discovered on a cliff at Carmel Bay. The latter locality is its most
northern known limit along the coast. '
GOVERNMENT LABORATORIES,
Manila, P. J.
a a =
a a i
SOME LITTORAL SPERMATOPHYTES OF THE
NAPLES REGION.
Jj. ¥. BERGEN.
THE strand flora about the Bay of Naples differs so much from
most of those which have been studied with reference to the toxic
effect of sodium chlorid solutions that it seemed to the writer worth
while to investigate the conditions of existence of a characteristic
association.
Along a strip of beach sand not quite two meters above the average
sea level and less than ten meters from the water line, on the margin
of the Bay of Baiae, a well defined association of somewhat more
than fourteen members was found. All of these occurred within a
distance of a hundred meters, measured along the shore. The
species determined were:
Euphorbia Paralias, E. terracina, Polygonum maritimum, Matthiola sin-
uata, *Alyssum maritimum, Plantago Coronopus, Medicago marina, M. litora-
lis, *Lotus ornithopodioides, Eryngium maritimum, Echinophora spinosa,
*Senecio vulgaris, *Artemisia variabilis, *Inula viscosa.
I shall refer to this group as Association A. Other species occur
as members of the association, but none could be identified with cer-
tainty at the time when these studies were made (January, February,
and March). The five species designated by asterisks are much
more abundant in inland stations than they are as strand plants.
Conspicuous members of other strand associations neighboring the
one above given are: Narcissus Tazzetta var., Thymelaea hirsuta,
Glaucium flavum, and Verbascum sinuatum. Out of the fourteen
listed above, Euphorbia Paralias, Polygonum maritimum, Eryngium
maritimum, and Echinophora spinosa are the most notably psam-
mophilous species. All four of these are capable of growing out of
drifting sand, emerging again and again as they are partially buried.
Only a few of the structural peculiarities of halophytes, as enu-
merated by WARMING,' were to be noted in the association under dis-
cussion. Studies carried through a considerable part of the year
* Oekologische Pflanzengeographie. Zweite Auflage. Berlin. 1902. pp. 305-308 .
327] (Botanical Gazette, vol. 41
328 BOTANICAL GAZETTE [MAY
would be necessary in order to enable one to make a detailed com-
parison between plants of the same species found growing as members
of this strand flora and further inland. The points that could be
established during the months of late winter and early spring when
these observations were made were as follows:
Alyssum maritimum (littoral form): leaves with more, longer, and
stouter hairs on both surfaces than occur on the ordinary form; leaves
much smaller and darker green than in the ordinary form; leaves
from 1.5 to 3.5 times as thick as in the ordinary form.
Senecio vulgaris (littoral form): plants very low, often with the
leaves in rosette form and heads in full bloom only 2°™ high. The
maximum height observed was 8°™, while the average height of
plants growing in fairly good soil further inland was (ten specimens
taken at random) 35.7°™. The stems of the littoral form were pro-
portionately stouter and more hairy, and: the leaves were darker
green, more pubescent, thicker, more sessile (sometimes clasping
and almost decurrent). The root system of plants of the littoral
form is much more developed than in the ordinary form, but there
are no aerial roots, such as are commonly found along the lower part
of the stems of this Senecio when growing in ordinary situations.
Artemisia variabilis (littoral form): leaves with more slender
divisions, the tips more strongly mucronate and the surface more scaly
than the ordinary form; the young leaves of littoral plants were
often densely pubescent, while I have never seen them so on plants
growing in ordinary stations.
No decided differences were noted between the specimens of
Inula found growing in the beach sands and those occurring else-
where. The Lotus plants were too young to be compared with
inland specimens.
- The beach sand in which Association A was growing consisted of
grains for the most part ranging from o.80™™ to o.30™™ in diameter.
It was evidently mainly derived from comminuted trachyte and
scoriaceous lava, probably from the little extinct volcano of Monte
Nuovo close by. A sample of the sand was taken February 1 from
among the roots of the Euphorbia Paralias and Matthiola sinuaia
and analyzed gravimetrically for chlorin. It contained 7.3 pet
cent. of moisture, which contained a trifle more than 0.04™8 chlorin
1906] BERGEN—LITTORAL SPERMATOPHYTES 329
per gram. In round numbers, 0.2 per cent. of the moisture was sea
water. This extremely low per centage of chlorin accords with the
fact that waves never in ordinary storms wash up as far as the level
on which the plants were growing. They must, however, some-
times break into spray which reaches the station of the association
studied. For the purpose of getting data for comparison of the
salinity of the sand above described with other marine sands and
soils and with ordinary garden soil, I made some further analyses,
which are summarized in the table below.
Sea water
Per cent. | Per cent. Per cent.
a tok on water per cent. of
1. Beach sand, Bay of i eraemoncene A)| 0.0003 0.015 7.3 .19
2. Beach h sand, Mare" Motto iis acs 0.0163 0.807 21.4 2 76
3. Beach mud, Lake Lucrinus.......... 0.0125 0.619 52.5 2.23
4. Loam from vineyard near ( Poeeale -.+| 0,00025 25.0
The second column of the table shows what proportion of sea water
present in the sand or mud would account for the amount of chlorin
actually found. The third column gives the total moisture present,
and the fourth column shows what proportion of total water in the
soil was sea water. The calculations are based on the assumption
that the water of this portion of the Mediterranean contains in 1,000
parts about as follows:? NaCl, 28.76; KCl, 0.66; MgCl, 3.25.
In the beach sand from Mare Morto (no. 2) was found an asso-
ciation consisting of Polygonum maritimum, Salsola Kali Tragus,
and a species of grass undeterminable at this season. In the imme-
diate neighborhood occurred occasional specimens of Aster Tripo-
lium and much Plantago Corono pus.
In the beach mud from Lake Lucrinus (no. 3) I found no member
of Association A except Inula viscosa. This was extremely luxuriant,
with branches attaining a diameter of 4°™ instead of the diameter
of o.8 to 1°™ usual in ordinary localities. The only associates
identified in the very limited area examined were Statice Limonium,
Ficus Carica, and Inula crithmoides (?).
It is evident from the analyses given that none of the soils examined
were saline in any such degree as the familiar salt marshes of the
2 Roru, Justus, Allgemeine und Chemische Geologie 1:524. 1879.
330 BOTANICAL GAZETTE [MAY
New England coast and the maritime provinces of Canada.3 I
have not indeed been able to find any typical salt marshes in the
Naples region, since the beaches are often for long distances walled
or protected by riprap against the encroachment of the sea, and
nearly land-locked bodies of salt water such as Lake Lucrinus and
the Mare Morto are surrounded by vertical stone walls, to admit of
the utilization of the adjacent land to the water’s edge.
It would a priori seem probable that the plants of Association A
would belong to the category of littoral species rather than of genuine
halophytes, and the results of my cultures confirm this supposition.
It seemed to the writer that the questions most worthy of investi-
gation in connection with the association studied were: (1) the rela-
tive sensitiveness of the species to the effect of sea water or sodium
chlorid solutions;+ (2) the relative sensitiveness of strand-grown
and inland-grown specimens of the same species to the effect of such
solutions.
Thrifty self-sown seedlings of the species of Euphorbia, Matthiola,
and Senecio in the list of members of Association A were found in
great abundance, and therefore special attention was given to these
species, though larger plants of all the others which could be obtained
in good condition were also studied. Sea water and pure sodium
chlorid, in solutions ranging from 1 to 6 per cent. of the salt were
used for the cultures. The roots of the plants were partially freed
from adhering sand or earth by careful immersion in a solution of the
same strength as that employed in the culture, and then the plants.
were transferred to small tumblers containing the culture solution in
which the roots were immersed. Each tumbler was covered by a.
disk of waterproofed pasteboard, fitted around the protruding stems.
of the seedlings, sufficiently close to prevent much evaporation, but
not to hinder aeration of the solution. The room temperature
during the two and a half months devoted to the investigation usually
ranged between 12° or 15°C. by night and 20° or 22° by day, and.
the plants were about 1.5™ in front of a south window 2.5™ wide.
3 See GANONG, W. F., The vegetation of the Bay of F undy salt and diked mani a
Bot. Gaz. 36:286, 292.
4 The writer took a eucatS the accuracy of H. Couprn’s statement that the
fatal effect of sea water upon vegetation is due to its sodium chlorid. See his article:
Sur la toxicité de chlorure de sodium, etc. Rev. Gén. Botanique 10:177. 1898.
1906] BERGEN—LITTORAL SPERMATOPHYTES 331
In order to avoid the possible presence of traces of salts of copper or
other injurious metals sometimes found in distilled water the solu-
tions were made up with very pure cistern water.
Much difficulty was experienced in getting perfectly comparable
results from the fact that slight individual differences in the plants
(such as relative development of the root system) made decided
differences in their tolerance of the saline solutions employed. Many
values were discarded, from the fact that they were evidently errone-
ous from variations of this kind. In general, as was to be expected,
the duration of life in sea water was considerably greater than in a
pure sodium chlorid solution containing the same per cent. of this
salt (2.88) that is found in Mediterranean sea water. Some of the
principal results obtained are summarized in the following table:
ORDER OF RESISTANCE TO SALINE SOLUTIONS.
2.88 percent.| In 100 per cent.
pet NaC], sea water,
lived days lived days
Euphorbia — ee eee eee 17 20
PAOGES GEIS 6 305 os peewee oes eS 19
Euphorbia Actinic? ee ree eee ere 2-13 9
Echinophora eee Gi bcd, RRR OE eV Pe 7
Polygonum maritimum........5.....+:+- g-I0
Alyssum maritimum (ioral Peay ee 7
cx Plan cantar 7
Senecio vulgaris (itoral). EF re . 5
Sage eae oe .
It is noteworthy that the littoral and the inland specimens of
Alyssum and of Senecio were equal in their resisting power, as might
have been expected from the comparatively equal amounts of chlorid
in the beach sand and the vineyard soil.
Some experiments were made to ascertain the maximum per cent.
of seawater which could be tolerated by the species of Association A
without speedy appearance of symptoms of injury, such as partial
drying up or death of shoots. Difficulty was found in discriminating
between retardation of growth and weakening caused by culture
under artificial conditions, and that due solely to the excessive salinity
of the culture solutions. It was considered that a plant was unharmed
if it showed no decided ill effects from the solution (except retarded
332 BOTANICAL GAZETTE [MAY
growth) after the culture had continued for a month. The results
appended are only approximate.
PER CENT. OF SEA WATER TOLERATED WITHOUT SPEEDY INJURY.
BOC i a eR BG
eerea Ps oN a Oe ke oe
chinopnore Ses. a Se OOF More
Polygonum maritimum . ode 50
Bapnorbriterraciia . . 5. 4 sw tl ee Ce” GO Orless
For the sake of comparison a few cultures were made of species
growing in full sunlight in the vineyard the soil of which was analyzed
for chlorin as above given. All soon succumbed to the effect of 50
per cent. sea water (and in general still more readily to 1.5 per cent.
sodium chloride solution). Classed according to the readiness with
which they wilted and then died, their sensitiveness to the salts in
solution was about in the following order.:
1. Lamium amplexicaule. 6. Geranium molle.
2. Fumaria Gussonii. 7. Rumex bucephalophorus.
3. Papaver Rhoeas 8. Euphorbia Peplus.
4. Polygonum aviculare aks ian g. Anthemis arvensis var.
5. Veronica Tournefortii
Roughly speaking, the order above given is that in which these
annuals wither and die at the onset of the summer drought, except
as regards no. 3, which persists well into the dry season.
In order to compare the behavior of the plants of Association A
with that of true halophytes, cultivated in saline solutions, some
seedlings of Salsola Kali Tragus were procured from the sand adja-
cent to Mare Morto, above described. The plants were 2-2.5°™
high and cultures were made of these in clean sand, rinsed with the
saline solution to be used, and then flooded once a day with another
portion of the solution, which was afterwards poured off. The con-
_ centrations employed were respectively of 4, 5, 5.5, 6, and 7 per cent.
of sodium chlorid, and other specimens were cultivated in solutions
without sand.
In sand the plants appeared normal up to and including 6 per
cent. sodium chlorid solution, though growth was slow for all con-
centrations above 4 per cent. Without sand prompt loss of turgor
was noticed in 6 per cent. solution, corresponding pretty nearly to
that observed in the solution of 7 per cent. with sand. This dimin-
et a
sane
t
}
}
1906] BERGEN—LITTORAL SPERMATOPHYTES 333
ished effect of the salt in the presence of sand is in accordance with
the conclusions of TRUE and OGLEVEES in regard to the lessened
effect of toxic substances in aqueous solution in presence of inert and
insoluble solids.
None of the work done was shaped with special reference to deter-
mining whether the injurious action of saline solutions on the plants
examined was due to dehydrating or other physical effects, or whether
it was purely toxic. A notable desiccation and shrinkage was often
observed throughout the stem while the upper leaves and especially
the terminal bud remained vigorous. This fact would tend to con-
firm the hypothesis that the lethal action of dissolved salts is of a
physical character. On the other hand, the fact that a pure sodium
chlorid solution is usually more quickly fatal than sea water con-
taining the same amount of sodium chlorid plus other salts tends to
confirm the toxic hypothesis. Probably both factors may cooperate
to produce death. Apparently the relative ease with which root
hairs of the species studied can be plasmolyzed does not bear any
definite relation to the susceptibility of the species to the action of
sodium chlorid solutions.
The principal conclusions reached may be summarized as follows:
1. Association A consists of members very unequally resistant to
the action of pure sodium chlorid solutions and of sea water.
2. The tolerance of sodium chlorid on the part of some members
of this association is considerably greater than that of ordinary non-
littoral plants; in other words, they are facultative halophytes.
3- Many of the species of this association are typical psammo-
phytes, none are typical halophytes.
4. Growth of a non-halophytic species for many generations in
an atmosphere at times highly charged with saline spray does not
bring about greater tolerance of saline solutions when absorbed by
the roots.
5- The Salsola seedlings studied showed a tolerance of sodium
chlorid solutions up to almost 6 per cent. as good as that of any
member of Association A for the 2.88 per cent. solution.
NApLEs, ITALY.
5 TrRuE R. H., and OctevEE, C. S., The effect of the presence of insoluble sub-
‘stances on the toxic action of poisons. Bot. GAZETTE 39:1-21. 1905
NEW AND NOTEWORTHY NORTH AMERICAN SPECIES
OF TRIFOLIUM.
HoMER DOLIVER HOUSE,
(WITH TWELVE FIGURES)
I. NEW OR NOTEWORTHY SPECIES OF THE UNITED STATES.
THE following notes upon the genus Trifolium are based upon
material in the National Herbarium. All figures x 1}.
Trifolium Greenei, nom. nov.—(f7g. 1).
—T. bifidum decipiens Greene, Fl. Fran.
24. 1891; not TJ. decipiens Hornem.,
Hort. Hafn. 2:715. 1815.— Of much
broader distribution and apparently not
merging into T. bifidum Gray, though
closely related to that species. It has
more of the general appearance of T.
gracilentum T. & G., but distinguished
from it by its villous-pubescent peduncles
and cuneate-oblong, subglaucous, and re-
tuse leaflets.
Low moist places or natural grassy meadows,
from San Diego to Mendocino and Butte Counties,
California. The type, collected at Berkeley by
Greene, is in the herbarium of Professor Greene.
TRIFOLIUN BIFIDUM Gray (fig. 2) seems
to be a species peculiar to the bay region
of California only; the type, collected by
Brewer (no. 1184, 1862), “near Marsh’s
Ranch, between Monte Diablo and the
San Joaquin (Contra Costa Co.), among
[ grass in a ravine near the water, May ai
Fic. 1.—Trijolium Greenei is in the National Herbarium.
House: @, portion of type. Trrrotium BRrEwERI S. Wats., Proc.
Sea naa es Am. Acad. 11:131. 1876.—T. amabile Loja.
leaves; d, banner. Nuovo Giorn. Bot. 15:142. 1883, ex descr.
Botanical Gazette, vol. 41] [334
a
ere ee,
1906] HOUSE—NEW SPECIES OF TRIFOLIUM 335
A careful reading of Loyacono’s paper on North American species of Tri -
folium makes very apparent the fact that he had a very scanty representation
of the forms and distribution of our western species. While his descriptions are
excellent, he has failed in many instances to
Trifolium Douglasii, nom. nov.—T.
altissimum Dougl., Hook. Fl. Bor.-Am.
1:130. pl. 48., 1830; not T. altissimum
Lois. 1807.
An abundant clover in moist or natural wet
meadows of eastern Washington, Oregon, and
adjacent Idaho. Flowering in June and July,
fruiting from July to the end of August.
TRIFOLIUM HARNEYENSE Howell, FI.
Northwest Am. 134. 1898.—T. arcuatum
Cusickii Piper, Bull.
Torr. Bot. Club
29:642. 1902.
An examination of
the floral parts of the
available herbarium
material of T. Har-
neyense and T. arcua-
tum Cusickii shows
them to be identical.
Trifolium villi-
ferum, sp. nov.—
V
af
‘1G. 2.—Trifolium bifidum
A. Gray: a, portion of type speci- .
men; b and ¢, leaflets from lower F 1g. 3.—Related to
leaves; d and ¢, leaflets from JT, eriocephalum
upper leaves; /, flower; g, calyx Nutt.Stems slender,
expanded; /, banner; 7, legume.
erect, from peren-
nial, ascending, and branching roots, 25 to 35°™ ;
high, densely villous-pubescent with long hairs, —_ Fic. 3.—Trifolium vil-
stem nearly glabrous at base only: leaflets Ji/erwm —— : fila
mm 0,calyxexpanded; c, ban-
oblong-lanceolate, a5 to gh long, 610 5a oe wing, and Keel.
wide, obtuse at base, usually acute at apex,
irregularly and inconspicuously repand-denticulate, pale green, appear-
ing almost glaucous by the dense, whitish indument, scarcely less
336 BOTANICAL GAZETTE [MAY
pubescent above than below; petioles 3 to 7°™ long, uppermost
shortest; stipules lanceolate or the upper ovate-lanceolate, subfoli-
aceous, 2 to 3.5°™ long, acuminate, entire or sparingly toothed:
inflorescence pseudo-terminal; peduncles 6 to 13°™ long; heads
densely many-flowered, ovoid when young, flowers all becoming
strongly reflexed: calyx densely villous with shaggy hairs without
especially toward the apex, tube about 2™™ long, the 5 subulate
nearly equal teeth plumose, 3 to 4™™ long, somewhat bent in age:
corolla pinkish-purple, 12 to 14™™ long; banner oblong, rounded or
obscurely retuse at apex, broadest (about 6™™) below the middle;
wings shorter, tapering to a blunt apex, blade with a strong basal
auricle; keel still shorter and more acute: legume ovoid, sessile,
densely plumose-pubescent, pubescence extending nearly to tip of
style, 2-seeded.
Flowering in June and July, fruiting in July and August. Southern Utah,
Palmer (no. 91), 1877 (type in the U. S. National Herbarium); Burrville, Sevier
Co., Jones {(no. 5642a), July 17, 1894, 2100™ alt.;
Deep Creek, Jones, June 6, 1891
Trifolium atrorubens (Greene), comb. nov.
—T. Rusbyi atrorubens Greene, Erythea 4:66.
1896.
iType, collected by Parish (no. 3745), Buff Lake,
San Bernardino Co., California, June 21-27, 1895, in
the herbarium of Professor Greene.
\Examination of the type and several other sheets of
pedunculatum Rydb., and should properly be given
specific ran
Trifolium shastense, sp. nov.—Fig. 4.—
Related to T. longipes and T. oreganum. Stems
numerous from matted, branching, and slender
rootstocks (forming a sod), 10 to 15°™ high;
Fic. 4.—Trijolium shas- silky pubescent above and beneath on the
ype re leaves; stems purplish below: stipules lanceo-
late, green, aristate-acuminate, entire or few-toothed, 12 to eee
long; leaflets lanceolate, acute at base, often broadest above the
middle, apex aristate-acuminate, margins prominently spinulose-
1906] HOUSE—NEW SPECIES OF TRIFOLIUM |
denticulate, glabrous above, silky-pubescent beneath, 15 to 25™™
long, 4 to 7”™ wide; leaflets of the lower leaves relatively broader
and shorter, nearly obovate-cuneate, acute or rounded; petioles
mostly shorter than leaflets, but lower ones longer: inflorescence
usually solitary; peduncles 5 to 8°™ long, somewhat woolly-pubes-
cent above with whitish hairs, densely many-flowered in a globose
head; flowers sessile, the outermost spreading or becoming reflexed,
ro to 13™™ long: calyx silky-pubescent or becoming glabrate, tube
1.5 to 2.5™™ long, the five filiform-subulate teeth straight, 8 to
1o™™ long, upper ones shortest and scarcely more than 8™™ long,
sometimes shorter: banner sublanceolate, broadest (about 5™™) below
the middle, acuminate-pointed at the apex; wings and keel shorter,
wings conspicuously attenuate-pointed, keel acute: legume stipitate,
2-seeded.
North side of Mt. Shasta, Siskiyou Co., California, 1500-2700™ alt. Col-
lected by H. E. Brown (no. 362), type in the U. S. National Herbarium, June
11-16, 1897. No. 365, of the same collection is identieal.
Remarkable for its sharply serrated and pointed leaflets, acuminate-pointed
banner, and attenuate-pointed wings.
Trifolium Covillei, sp. nov.—Fig. 5.—Related to T. latijolium.
Stems very short and leafy, several from a solitary, perpendicular
thickened perennial root and appearing as a
dense green mat at its apex, the root 10 to
20° deep; stems 1 to 3° long: stipules
small, ovate, 5 to 7™™ long, blunt and often
rounded at apex, entire, adnate to the petiole
for two-thirds their length; leaflets obovate-
| subcuneate, rounded or retuse at apex, rarely
the upper acute, margins finely but not sharp-
ly or conspicuouly toothed, 6 to 12 ™™ long,
silky pubescent beneath, glabrous above;
petioles mostly shorter or but little longer
than the leaflets: peduncles 8 to 10°™ long,
much exceeding the leafy part of the plant,
silky-pubescent above; heads globose, 40 to
60-flowered, 2 to 2.5°™ in diameter, some of
Fic. 5.—Trifolium Covillei the flowers spreading or becoming reflexed,
House.
338 BOTANICAL GAZETTE © [way
all sub-sessile: calyx membranaceous, the tube 1.5 to 2™™ long,
sparingly hairy above, the 5 filiform-subulate teeth subequal, 2.5 to
3.5™™ long: banner yellowish, 12 to 14™™ long, inflated and includ-
ing the wings and keel, broadest (6 to 7™™) below the middle, acute,
wings subacuminate, keel acute: legume short-stipitate, 2-seeded.
Bog-lands in the Wenatchee Mountains, Kittitas Co., Washington, Coville
(no. 1180), Sept. 4, 1901 (type in the U. S. National Herbarium).
The group of small species related to T. monanthum Gray has
been not a little confused by various authors, and the location of
the type in the U. S. National Herbarium makes it possible to define
definitely its critical parts. The accompanying description and
drawing are from the type.
Q Wy
AW)
All Vv ayy
= AWA Wa
= <a
——
, BNA
Ay 2
Fic. 6.—Trifolium monanthum A.
Gray: a
a, entire plant (from type specimen);
6, involucre;
c, details of flower.
i
1906] HOUSE—NEW SPECIES OF TRIFOLIUM 339
TRIFOLIUM MONANTHUM A. Gray, Proc. Am. Acad. 6:523.
1865.—Fig. 6.—Perennial from numerous, slender, branching
roots; stems branching from near the base, ascending, 2 to 4°™
high, only sparingly villous: stipules ovate-lanceolate, green, 3 to
5-nerved, cuspidate-acuminate, 2 to 4™™ long, entire or rarely with
a few minute rigid teeth near the apex; leaflets obovate-cuneate,
retuse or truncate at apex, margins spinulose-denticulate, 4 to 6™™
long, sessile, pale green with a few villous hairs beneath, darker
green above, the ascending, branching whitish veins ending in the
subcallous mucronate teeth of the margin; petioles filiform, longer
than the leaflets: peduncles filiform, mostly shorter than the leaves,
one-flowered (rarely 2), involucre of two entire or sparingly toothed,
ovate-lanceolate, cuspidate-acuminate bracts, 2 to 3™™ long: calyx
cylindrical, twice as long as involucre, about 4™™ long, sparingly
villous, 10-nerved, 5 of the nerves more prominent than the others, the
triangular-lanceolate, spinulose-acuminate teeth about one-third the
length of the tube: corolla g to 12™™ long, purplish-white, the
slender tube elongated and somewhat glandular, not scarious or
inflated after flowering; banner lanceolate-obovate, retuse; wings
shorter and rounded; keel sub-acute: legume stipitate, 2-seeded.
Flowering from the middle of June to September; fruiting from July to October’
Mountain meadows, banks, and grassy places, Sierra Nevada in California
from Alpine to Tulare County, and in Ormsby County, Nevada; 2100 to 3000™
alt.
CALIFORNIA: Sierra Nevada, Lemmon 1875; Manachi Meadows, 2500™
, Rothrock (no. 307), Sept. 1875; Tuolumne Co.: ‘ Moist bank by Soda
swe alt. 2650™,”’ Brewer, June 26, 1863 (no. 1704), type in U. S. National
Herbarium; vicinity of Tuolumne Meadows, 2600-2900™ alt., Hall and Bab-
cock, July 1902 (no. 3625); Alpine Co.: Caple’s Lakes, 2600™ alt., Hansen,
June 21, 1892 (no. 351); Fresno Co.: meadows near Block mountain, 30007
alt., Hall and Chandler, July 1900 (no. 613); Tulare Co. near Mineral King,
a7so™ alt., Coville and Funston, Aug. 4, 1891 (no. 1473).
eiak. Ormsby Co.: Snow Valley, 2460-2615™ alt., C. F. Baker, July 8,
1902 (no. 1282).
TRIFOLIUM TENERUM Eastw., Bull. Torr. Bot. Club 29:81. 1go2.
“Higher meadows on the trail to the South Fork of King’s River
(Fresno Co.), California. It was collected by the writer at Summit
and Bearskin Meadows, July 1-13, 1899. The first named speci-
340 BOTANICAL GAZETTE [May
mens are considered the type samara Type in the Herbarium
of the California Academy of Sciences.”’
Characterized by the canescent and softly villous foliage, ate
nerved and setosely serrulate leaflets; heads 1 to 6-flowered; invo-
lucre glabrous, of 2-5 separate, laciniate-aristate bracts, 2 to 4™™
long; banner of the corolla with three rounded teeth at the trun-
cate apex; wings slender, as long as the banner, auriculate at base
of blade; keel two-thirds as long, tipped with an obtuse erect beak,
the keel itself purple, auricled at base; ovary obovate, pilose at
summit, one-ovuled.
TRIFOLIUM GRANTIANUM Heller, Muhlenbergia 1:136.
Undoubtedly distinct from T. tenerum Eastw., as HELLER indi-
cates, but it is unfortunate that the floral characters were not better
described [Calyx cylindrical, or somewhat campanulate, 4™™ long,
the tube 2™™ long, more or less membranous, veins prominent;
the narrowly lanceolate teeth aristate, green: corollas 1° long,
slender, 2™™ across, whitish, the hood of the keel purple”) when
it is considered that in this group of small species the floral char-
acters are of the utmost importance.
Based on Grant’s Number 6343, July 23, 1904, from San Bernardino Co.,
Calif.
TRIFOLIUM PARVUM (Kellogg) Heller, Muhlenbergia 1:114.
1905.—Fig. 7.—T. pauciflorum (?) var. parvum Kellogg, Proc.
Cal. Acad. 5:54. 1873; T. multicaule Jones, Bull. Torr. Bot. Club
Q:31. 1882.—The prostrate or slightly ascending stems:10 to 20°"
long, often many from a thickened root; softly silky-pubescent,
sometimes densely so: leaflets obovate, retuse or obtuse or some-
times those of upper leaves short acute at apex, subcuneate at base,
minutely spinulose-serrulate; stipules broadly ovate, sharply toothed
and acute; petioles filiform, the lower 1 to 3 times the length of
the leaflets, the upper scarcely longer than the leaflets: peduncles
exceeding the leaves, 1 to 3°™ long, 1 to 7 (usually 2 to 5)-flowered ;
involucre 5 to 7-divided into triangular-lanceolate acute and entire
segments, 1 to 2™™ long, these spreading in age, rarely somewhat
toothed: calyx 3 to 4™™ long, villous-pubescent, the 5 triangular
lanceolate teeth about equaling the tube in length, the lower one
7
OTT TTT, cy ta ae
—— It
.
i od
——
times the length of the leaflets: peduncles
' the length of the involucre, teeth subulate-
1906] HOUSE—NEW SPECIES OF TRIFOLIUM 341
a little shorter and broader than the others, all spinulose-acuminate:
corolla a little more than twice the length of the calyx, white with a
purple-tipped keel; banner narrowly oblong, about 10™™ long
and 3.5 to 4.5™™ wide, deeply retuse at apex; wings and keel much
shorter, the keel acute: legume 2-seeded. :
Around springs and in natural moist meadows of the Sierra Nevada, Cali-
fornia, from Nevada Co. to Acador Co., Yosemite Park, and Fresno Co., at 1500
to 2100” alt.
Nevada Co.: Soda Springs, Jones (no. 2592), July 30, 1881 (type of T. mul-
ticaule, in U. S. National Herbarium); low ground on south side of Donner Lake,
Heller (no. 6942), July 16, 1903.
Amador Co.: Bear River, Hansen (no. 1968), Aug. 30, 1896.
Yosemite Park: Hetch-Hetchy Trail, Hail and Babcock (no. 3385), July
1902.
Fresno Co.: Pine Ridge. Hall and Chandler (no. 256), June 15-25, 1900.
Trifolium simulans, sp. nov.—Fig. 8.—Perennial from slender,
branching roots; stems numerous, prostrate or ascending, slender,
5 to 20° long, glabrous: stipules ovate-lanceolate, 6 to 10™™ long,
green or the lower-ones subscarious, spinulose-acuminate, and with
a few aristate tecth, 5 to 7-nerved; leaflets oblong-lanceolate, the
middle one and often the others cuneate,
irregularly spinulose-denticulate, mucronate-
tipped at the acute apex, g to 14™™ long,
2.5 to 5™™ wide; petioles filiform, 2 to 4
as slender and-shorter than the petioles; in-
volucre divided into 2 or 3 parts, these cleft
again to make 5 to 7 narrow, spinulose-acu-
minate, conspicuously nerved, simple or few-
toothed segments to the entire involucre:
calyx short-pedicelled, glabrous, about twice
acuminate, 3 to 4™™ long, the lower slightly
shorter: corolla very large for the size of the
plant, white with a purple-tipped keel; ban-
ner narrowly oblong, about 15™™ long, 5"™ Fic. 8—Trifolium simu-
wide, retuse and mucronate at apex; wings Jans House.
342 BOTANICAL GAZETTE [MAY
and keel much shorter, blade of wings about 9™™ long, that of the
acute keel only about 4™™ long: legume 2-seeded, sessile.
San Jacinto Mountains, California, 1800 to 2700™ alt., H. M. Hall (no.
710), July 22, 1897, type in the U. S. National Herbarium); Strawberry Creek
(San Jacinto Mts.) 1600™ alt., H. M. Hall (no. 2200), June 20, 1gor.
Resembling T. parvum in size, but very distinct from it in the remarkably
large flowers for the size of the plant, the leaf, calyx, and corolla characters also
showing well-marked differences. It appears to be as distinct from T. parvum
as T. tenerum is from T. monanthum, and to show these differences descriptions
of all three species are given.
II. MEXICAN SPECIES.
TRIFOLIUM AMABILE HBK., Nov. Sp. & Gen. 6:503. pl. 593.
1823; T. Humboldtii Spreng., Syst. 3:313. 1826 (T. pauciflorum
Willd. herb.); 7. Hemsleyi Loja., Nuovo Giorn. Bot. 15:143. pl. 4.
jig. I. 1883.
One of the commonest — of Mexico and distributed from northern
Mexico to Central Americ
TRIFOLIUM GRACILENTUM T. & G., Fl. N. Am. 1:316. 1838;
I. denudatum Nutt., Proc. Acad. Phila. II. 1:152. 1848.
Lower California, San Quentin Bay, Palmer (no. 613), Jan. 1889.
Trifolium longifolium (Hemsley), comb. nov.—T. amabile var.
longifolium Hemsley, Biol. Cent. Am. Bot. 1:232.1879; T. gonio-
carpum Loja., Nuovo Giorn. Bot. 15:145. pl. 4. fig. 2. 1883.
HeMsLEy’s description is based upon Parry and Palmer’s no. 134, although
other specimens are mentioned. Loyacono’s description is also based upon
a plant collected by Parry and Palmer, but no collection number is given. His
description, however, agrees well with a duplicate of HemstEy’s type in the
National Herbarium, and the conclusion was forced upon me that they are iden-
tical.
San Luis Potosi, Parry and Palmer (no. 134), 1878: Chihuahua, Pringle
(no. 1208), 1887; Townsend and Barber (no. 177), 1899; Palmer (no. 385),
1885: Durango, Palmer (no. 237), 1896: Tepic, Rose, Aug. 9, 1897: Jalisco,
Palmer (no. 236), 1886: Aguascalientes, Rose and Painter (no. 7795), 1993:
Federal Dist., Pringle (no. 7492), 1897: Vera Cruz, Orizaba, Bourgeau, 1865-66;
Seaton (no. 93), 1891: Oaxaca, Rose and Hough (no. 4644), 1
Trifolium Lozani, sp. nov.—Fig. 9.—Related to T. mexicanum.
Stems numerous, spreading and ascending from a perennial root
densely silky-pubescent, 10 to 20°™ long; the internodes relatively
1906] HOUSE—NEW SPECIES OF TRIFOLIUM 343
short: stipules ovate, lower scarious and entire, upper green and
sparingly toothed, all acuminate, 6 to 1o™™ long, 4 to 6™™ wide;
leaflets cuneate-obovate or cuneate-oblong, sessile, 8 to 15™™ long,
| 5 to 10o™™ wide, rounded at apex, glabrous above, sparingly
pubescent beneath, subentire, the minute teeth
very inconspicuous; petioles mostly shorter than
the leaflets, or the lower slightly longer: pedun-
cles scarcely exceeding the leaves, 1 to 3°™ |
long; heads globose, ebracteate, 25 to 50-flow- .
ered; flowers short-pedicelled, all becoming \
+ — ei
reflexed: calyx tube subcampanulate, pubes-
| cent, slightly more than 1™™ long; the subu-
late teeth twice as long, four of them ascending,
lower one straight: corolla yellowish; banner
| broadly oblong, 5™™ long or less, retuse; wings
and keel much shorter, subacute: legume ob- Fic. 9.—Trijolium Lo-
long in outline, very blunt at apex, 3™™ long or 72”? House.
less, 4-seeded: seeds nearly as thick and broad as long, smooth and
brownish, less than 1™™ long.
a,
Eslava, Federal District, 2300™ alt.,
Pringle (no. 9512), June 15, 1901 (type
sheet no. 396298 in U.S. National Her-
barium). Distributed as T. amabile,
which it inno way resembles. It dif-
fers from T. mexicanum by its more
densely pubescent stems and pedun-
cles, smaller flowers, and relatively
shorter calyx teeth. Named in honor
of Sefior Filemon Lozano, assistant to
Mr. PRINGLE.
€
TRIFOLIUM MEXICANUM Hems-
ley, Biol. Centr. Am. Bot. 1:233.
1879.— Fig. 10.—T. potosanum
Loja., Nuovo. Giorn. Bot. 15:
144. pl. 2. 1883.
The type of T. mexicanum is from
San Luis Potosi (Parry and Palmer no.
Fic. 10.—T rijolium mexicanum Hemsley. 137; 1878), and upon the same number
344 BOTANICAL GAZETTE [MAY
is based T. potosanum Loja. A duplicate type is in the U. S. National
Herbarium.
The following specimens from central Mexico differ from typical T. mexi-
canum in having larger flowers, more pubescent stems, and blunter leaflets, and
are more spreading in habit. They may represent a variety, but scarcely more.
Mexico (state): hills near Ozumba, 2400™ alt., Pringle (no. 9775), Nov. 8,
1902; Flor de Maria, Pringle (no. 3238), Sept. 4, 1890; Rose and Painter (no.
7816), Oct. 13, 1903.
Trifolium Nelsoni, sp. nov.—Fig. 11.—Related somewhat to
T. mexicanum but scarcely resembling it. Stems spreading and
ascending from a perennial root, minutely
pubescent, about 50°™ long or less: sti-
pules ovate-lanceolate, green, rigid and
the lower scarious, aristate-acuminate,
entire, 15 to 20™™ long; leaflets ovate-
lanceolate to elliptic-oblong, sparingly
pubescent and pale beneath, green and
glabrous above, usually with a whitish
V-shaped blotch on the upper surface,
apex acute or blunt, callous-tipped, mar-
gins inconspicuously repand-denticulate ;
petioles of the lower leaves 2 to 4 times
the length of the leaflets: peduncles dense-
ly woolly-pubescent toward the summit,
: often 15 to 20°™ long; heads depressed-
Fic. 11.—Trijolium Nelsoni globose, 20 to 40-flowered, flowers sub-
i tended by subulate filiform bracts, 3 to
5™™ long; pedicels none or very short: calyx-tube 1o-nerved, about
1.5™™ long, the subulate green teeth 4.5 to 5™™ long, subequal:
corolla yellowish; banner 8 to 10™™ long, orbicular-obovate, slightly
retuse at the apex or rounded; wings and keel shorter but relatively
broad. .
Vicinity of La Parada, Oaxaca, E. W. Nelson (no. 1016), Aug. 19, 1894
(type in the U. S. National Herbarium). Remarkable for the extremely broad
petals.
Trirotium Patmerr S. Wats., Proc. Am. Acad. 11:132. 1876.
Guadalupe Island: Palmer (no. 26, 1875), type in Gray Herbarium; a dupli-
cate type in herbaria of Professor Greene and Columbia University; Greene
Apr. 21, 1885; Palmer (no. 859), 1889; Dr. F. Franceschi, 1893.
eee.
ae |
SSN Mee
1906] HOUSE—NEW SPECIES OF TRIFOLIUM 345
Trifolium cognatum, sp. nov.—Fig. 12.—Related to T. longi-
jolium. Minutely pubescent or glabrate; stems spreading or ascend-
ing from a thickened per- b
ennial root, 10 to 30°™ high, ;
often somewhat tinged with
dull purple below, striate:
stipules ovate or the lower
ovate-lanceolate, green, acu-
minate, entire and scarious
margined, 12 to 16™™ long;
leaflets _ elliptical- oblong,
subcuneate at base, glab-
rous, rounded and minutely
mucronate at apex, 8 to
20™™ long, 4 to 10™™ wide,
margins minutely and _ir-
regularly denticulate; peti-
oles 2 to 4 times as long
as the leaflets: peduncles
exceeding the leaves; heads
subglobose, 15 to 40-flow-
ered; flowers erect when Fic. 12.—Trifolium cognatum House: 4, stipule
young, soon becoming en- and leaflets; 5, flower; ¢, calyx expanded; d, ban-
: ner, wings, and keel.
tirely reflexed, their pedicels
2 to 4™™ long: calyx with a few scattered hairs, the tube 10-nerved,
about 1™™ Jong, the subequal, subulate-acuminate teeth 3 to 3.5™™
long: corolla 7 to 9™™ long; banner violet-purple, ascending, retuse
at apex, nearly s™™ broad; wings and keel shorter and yellowish:
legume 3 or 4-seeded.
Barren hills above Pachuca, Hidalgo, 2600-2g00™ alt., Pringle (no. 6933),
in U. S. National Herbarium.
Although fully matured legumes are not present on the type, the species is
purple and yellow flowers.
346 BOTANICAL GAZETTE [ May
TRIFOLIUM REPENS L., Sp. Pl. 767. 1753
Reported by authors from the valley of Mexico and specimens from Central
America have been examined. Probably introduced and sp about many
of the larger cities and seaports.
TRIFOLIUM MICROCEPHALUM Pursh, Fl. Am. Sept. 2:478. 1814.
Guadalupe Island, Palmer (no. 831), Apr. 1, 1889. Probably also occurring
in northern Lower California.
TRIFOLIUM RHOMBEUM S. Schauer, Linnaea 20:740. 1847.
As I have not been able to establish fully the identity of this species, further
than to advance the suggestion that it is perhaps the T. mexicanum of Hemsley,
the original description is given here in full.
“Trifolium (Trijoliastrum) rhombeum S. Schauer: caulibus ad-
scendentibus striatis cum petiolis pedunculis calycibusque villoso-
lanuginosis, foliolis rhombeo-ellipticis obtusis mucronatis arguta den-
ticulatis striato-venosis supra glabris subtus pilosiusculis glabres-
centibus, stipulis membranaceis lato-ovatis mucronulatis pilosulis,
capitulis axillaribus longe pedunculatis multifloris densis exinvolu-
cratis, floribus pedicellatis demumque deflexis, calycis laciniis subae-
qualibus setaceis erectis tubo brevi longioribus corolla tertia brevi-
oribus glabris.
“In montosis Mexici. Auahenks n. 164. (perennial).
“Ex typo Tr. hybridi nostratis; inter mexicana forte affine Tr. amabili
HBK., ceterum pubescentia, foliorum figura et serratura floribusque magnis.
insignis. Petioli 6-9 lin. longi, stipulis a longiores; foliola subpollicaria.
Pedunculi folium longe excedentes. Flores magni, yexillo 4 ve aequante
Corolla alba ‘vel pallide rosea ex sicco rae vexillum emarginat
TRIFOLIUM SCHIEDEANUM S. Wats., Proc. Am. Acad. oe
1883.—T. reflexum Schlecht., Linnaea 5:576. 1830; not L.
“Jalapa (Schiede), and at Lerios, 45 miles east of Saltillo, [E. Palmer] (201).””
The locality for Schiede’s plant as given by SCHLECHTENDAL is “Prope Jala-
pan ad latera montis Macultepec, San Andres inque graminosis.”
A duplicate of Palmer’s plant mentioned above is in U. S. Nat. Herbarium.
TRIFOLIUM TRIDENTATUM Lindl., Bot. Reg. sub pl. 1070. 1827-
Lower California, Todos Santos Island, Anthony (no. 194), 1897; San Quen-
tin Bay, Palmer (no. 697), 1889.
Trirotium WILLDENova Spreng., Syst. 3:208. 1826.—T. involu-
cratum Orteg., Hort. Matr. Dec. 33. 1797; Willd., Sp. Pl. 3:1372-
TR AE ETT: ALR, OES A nee
ee ae
a eT TTT ag,
f
{
1906] HOUSE—NEW SPECIES OF TRIFOLIUM 347
1801; not 7. involucratum Lam. 1778; T. sii ak Greene, Pittonia
a 186. 1897.
Both WILLDENOW and SPRENGEL seem uncertain regarding the native country
of this species and the identification of Willdenow’s name with an American
species is perhaps first made by . (Nov. Gen. & Sp. 6:502. 1823):
“Crescit prope Valladolid Mexicanorum alt. 1000 hex.”
is species, not rare throughout northern Mexico, is not at all closely related
to the species of California which for so long has passed as T. involucratum and .
which Professor GREENE has shown to be T°. Wormskioldii Lehm.
The important characters of T. Willdenovii are the linear-lan-
ceolate stipules, subulate-acuminate and _lacerate-toothed; the
involucre divided nearly to the base into 6 to 8 nearly simple, subu-
late-aristate segments, resembling in this respect the T. spinulosum
of northwestern United States, but not 7. Wormskjoldii of Cali-
fornia. The leaflets are all linear and apiculate, except those of
the lower leaves which are relatively broader; the purple flowers
are from 16 to 18™™ long, in large erect heads; the banner very
narrow and retuse at apex.
Chihuahua: Nelson (no. 6054), 1899; Townsend and Barber (no. 60), 1899,
Goldman (no. 430), 1899; Palmer (no. 309), 1885; ee (no. 1209), 1887.
Durango: Palmer (no. 238), 1896; Nelson (no. 4768
San Luis Potosi: Parry and Palmer (no. 135), 1878; : ae (no. 602),
187
Valley of Mexico: Bourgeau (no. 79), 1865-66, Pedregal, near Tlalpam,
Rose and Hough (no. 4518), 1899.
CLEMSON COLLEGE,
South Carolina.
*
BRIEGPER AK TICLES
THE BASIDIUM OF AMANITA BISPORIGERA.!
(WITH SEVENTEEN FIGURES)
THERE are among the Hymenomycetes certain species which have
basidia bearing only two spores instead of the usual four. Such a form
is sometimes found in a genus the other members of which have basidia
with the usual number of spores. Some time ago while studying the
structure of a white Amanita which resembles A. verna, Professor ATKIN-
SON discovered that certain of the plants had basidia with only two spores.
He also found that the two-spored plant could be distinguished from
the four-spored A. verna by other characters, and he has described it
as a new species, A. bisporigera.
AMANITA BISPORIGERA.
Amanita bisporigera iprincgd . sp.—Plants entirely ves usually occurring
singly, about 9—13°™ high; pileus 4-6.5°m broad; stem 5-8™m™ thick; 2-2.5¢m
thick. Pileus convex to expanded, ry often gibbous or somewhat broadly umbo-
nate, smooth, viscid when moist, thin. Gills subel eal coe tapering more
behind itis they are rounded and free but close, rather crowded, edge of gills floc-
cose. Basidia 2-spored. svn to subglobose or oboval, smooth, with a
minute pedicel where siinskectit to ae erigmata as in many species, 8-ro@. Stem
nearly even, or slightly tapering oa solid, when fresh finely floccose scaly both
above and below the annulus, in age tending to become smooth + below
e annulus. Annulus thin, membranous, fragile, sometimes entir metimes
torn, superior. Volva thick. with apical deiiectane iit with the free mb b sping
into two or heres lobes which are usually closely it ati against the s
e ground in woods. It has been found many times at nee Nu YX:
and mela and specimens under twelve or more sheet are in the Herbarium
of the Siecemre of Botany, Cornell Univers
The t bears a striking resemblance to | verna, but is distinguished by its
more Piers habit and the two-spored basidia.
As this plant so closely resembles a four-spored aa it seemed
desirable to study the nuclear phenomena in the basidium to determine
how the behavior of the nuclei compares with what has been observed
in the four-spored forms by WAGER (5, 6), JuEL (4), HARPER (2), and
others. The results of a number of investigators make it seem very prob-
able that in all Basidiomycetes the young basidium contains two small
primary nuclei which fuse to form the secondary nucleus ofthe basidium.
‘Contributions from the ci aac of Botany, —_ University, No. 108.
tanical Gazette, vol. 41] as ha
eee ET Aerie ANTE TEE Ie
1906] BRIEFER ARTICLES 349
WAGER, however, was of the opinion that in some cases more than two
nuclei move into the young basidium from the multinucleate cells of the
hyphae. After the fusion of the primary nuclei, the secondary nucleus
increases greatly in size. By two successive divisions of this nucleus,
four nuclei are produced for the spores. In basidia with more than four
spores, as in some Gastromycetes, there are probably more than two
divisions. In Dacryomyces, which has basidia with only two spores,
there has been some difference of opinion, but it seems probable that
the two nuclear divisions take place in the usual way. DANGEaRD (1)
observed only one division in D. deliquescens Bull. IstvANFFI (3)
found that in D. chrysocomus Bull. there are two successive divisions,
and the four nuclei arrange themselves in a row and move in pairs into
the branches of the basidium.
However, the young spore receives but one nucleus, as one remains
behind in each branch of the basidium. JurEt (4) worked with D. deliques-
cens Bull. and found that there are two successive divisions of the nucleus,
but each spore receives a single nucleus, the others remaining behind
in the basidium.
The material for this study was secured during the summer of 1904
from plants collected in the vicinity of Ithaca, N. Y. Small pieces were
cut from the pilei of young plants soon after their collection and were
placed in 1 per cent. chromacetic acid where they remained 12-24 hours.
The material was then washed 3-4 hours in running water, dehydrated
in grades of alcohol, and passed gradually into paraffin. Sections were
cut 3-5 » in thickness. In sections showing some mature spores several
stages in the development of the basidium are found. It is better, how-
ever, for the study of the young stages to take sections from younger plants.
In fixing material from plants so young that none of the basidia bear
sterigmata, there is a possibility of confusing these plants with A. verna.
To avoid this, small pieces were cut from the pilei of the young plants
and these plants were then placed in a moist chamber and allowed to
continue their growth until the spores were produced. Then by freehand
sections of the pileus the species could be exactly determined. A. bi-
Sporigera is so distinct in appearance, however, on account of its more
slender form, that after one becomes familiar with it there is no difficulty
in distinguishing it from A. verna, even before it is fully mature.
The sections were stained with safranin and gentian violet, which
gives very good results. The preparations were studied with Zeiss 2™™
apochromatic, 1.40 aperture, and oculars 8, 12, and 18. Drawings were
made with camera lucida and ocular 18.
oe
ae
350 BOTANICAL GAZETTE [May
Material of the two-spored Agaricus campestris was prepared for
study in the same way, but on account of the smaller size of the nuclei
and the dense contents of the basidium, this plant does not offer favorable
material. ;
Fic. 1. Young basidium which is densely filled with cytoplasm and contains two
primary nuclei—Fic. 2. The membranes of the two nuclei are in contact.—FIc. 3.
Two primary nuclei in the process of fusion, but the nucleoli indistinct—Fic. 4. Fusion
of the nuclei almost complete——Fic. 5. Basidium somewhat increased in size after
fusion of primary nuclei.—Fics. 6 and 7. Older basidia in which the secondary nucleus
occupies a position near the end of the basidium; structure of nucleus at this stage.
very distinct.—Fic. 8. Division of secondary nucleus.—Fic. 9. Chromosomes moving
to the poles of the spindle——Fic. 10. Chromosomes os iE of the spindle.—Fics. 11
and 12. Daughter nuclei occupying different positions in basidium; usually near the
end.—Fic. 13. Basidium showing four nuclei—Fic. 14. The four nuclei crowded
together at some distance from the end of the basidium and the sterigmata begin-
ning to form.—Fic. 15. The cytoplasm beginning to pass through the sterigmata to
form the spores.—Fic. 16. A basidium in which the spores are almost mature; two
nuclei still seen near the center of the basidium.—Fic. 17. Old basidium from which
spores have fallen; two nuclei near the center.
1906] BRIEFER ARTICLES . 351
The young basidium first appears as a club-shaped branch from the
sub-hymenial layer. It is more densely filled with protoplasm than in
older stages and contains two nuclei. The structure of these nuclei can
be made out very easily. Each nucleus has a rather large deeply staining
nucleolus, a network in which the chromatin granules are imbedded,
and a distinct nuclear membrane.
As the basidium increases in size the nuclei fuse together into one
large nucleus. At first the two nuclei lie side by side with their mem-
branes in contact and without apparent change. Then the membranes
disappear at the point of contact. The nucleoli remain distinct for a
short time, but finally fuse so that the large secondary nucleus produced
by the fusion contains but a single nucleolus. The manner of the fusion
of the threads bearing the chromatin could not be made out. After this
fusion, the nucleus increases in size and comes to occupy a position near
the upper expanded end of the basidium (jigs. 6 and 7).
At this stage the structure of the nucleus can best be determined.
It is so large that it fills more than two-thirds the diameter of the basidium.
The nuclear membrane is very distinct, so that the nucleus stands out
very clearly from the contents of the basidium, which at this stage have
become vacuolate. The nucleolus is large and stains deeply, taking a
reddish color with the triple stain. The nuclear network consists of one
or more coiled threads in which are imbedded the chromatin granules,
which stain blue or purple. Between the coils of the thread are the
colorless spaces which are filled by the nuclear sap in the living cell.
The nuclear division takes place in the manner described by WAGER
(5) for A. muscaria. The chromosomes are produced from the thread
bearing the chromatin granules. The number of chromosomes is small,
but I have been unable to determine the exact number. The chromo-
somes are small and stain deeply. The spindle consists of a small number
of fibers, but they do not show the structures of fibers very clearly because
they are crowded closely together. The spindle is arranged transversely
near the apex of the basidium and is long and narrow with a small deeply
staining body at each pole. Such spindles as the one shown in fig. 8
are found frequently in the preparations. After the chromosomes move
to the poles there are a few persisting spindle fibers which connect them.
The daughter nuclei are now formed and usually occupy a position near
the apex of the basidium (fig. rz). Each daughter nucleus has the same
form and structure as the parent nucleus.
The most important question now is whether these two nuclei divide
again to produce four nuclei, as is the case in plants with four spores on a
352 BOTANICAL GAZETTE ? [MAY
basidium. After careful search I have been unable to find the spindles
of such a division, but numerous basidia which contained four nuclei have
been observed. In some cases only three nuclei show in the section, but
it seems probable that in all such cases the fourth nucleus is in another
section. These nuclei are small and in most cases the structure is not
so distinct as in the earlier nuclei of the basidium (jg. 13). The four
nuclei move back from the apex of the basidium and become crowded
together in an irregular mass in which it is difficult to distinguish the indi
vidual nuclei (figs. 14 and 15).
Soon after the four nuclei are formed, the two sterigmata grow out
from the end of the basidium. The granular content of the basidium
moves up and becomes more dense near the apex. Then the proto-
plasm begins to pass out through the sterigmata to form the spores. The
question which now presents itself is as to the number of nuclei which
pass into the spores. It is difficult to follow the details of the passage
of the nucleus through the sterigma, and it seems to me that the best
evidence as to the number of nuclei which enter the spores is found in
the examination of old basidia in which the spores are fully mature or
may have fallen off. Such basidia which contain only a small amount
of cytoplasm show the presence of two nuclei (fig. 17). From the facts
that the basidium when the sterigmata are formed contains four nuclei
and that it contains only two when the spores are mature one may conclude
that two nuclear divisions take place as in those forms with basidia which
bear four spores, but that only two of the nuclei enter the spores —CHARLES
E. Lewis, Cornell University.
LITERATURE CITED.
I. DANGEARD, Mémoire sur la réproduction sexuelle des Basidiomycétes. Le
Botaniste IV, 1
2. Harper, R. A., Wimicleais cells in certain Hymenomycetes. Bot. GAZETTE
pee Igo2
3. IstvAnrr1, G., = die Rolle der Zellkerne bei der Entwickelung der
Pilze. Ber. Desech: Bot. Gesell. 13:——. 1895.
4. Juet, H. O., Die Kerntheilungen in den Basidien und die Phylogenie der
Rastdinaropreeen: Jahrb. Wiss. Bot. 32: 361. 1898.
5. Wacer, H., On nuclear division in the Hymenomycetes. Annals of Botany
7:489. 1893.
» ————.,, On the presence of centrospheres in fungi. Ibid. 8:321. 1894.
a
CURRENT LITERATURE.
MINOR NOTICES.
The algae —The second volume of OLTMANNs’ large work on the algae has
appeared.‘ This part treats of a variety of general topics, the first volume
having been devoted especially to the different groups. Among the subjects
are the algal cell, the development of reproductive organs, the nourishment of
algae, life conditions, response to stimuli, polymorphism, life histories, adapta-
tions, and a discussion of methods of collection, study, and culture. The two
volumes give an excellent digest of the large literature in phycology and will prove
very valuable as the starting point for many lines of further advance. Some
of OLTMANNS’ views, as for example that of the place of the tetraspore in the
life history of red algae, are not likely to be sustained, but the work is a very schol-
arly contribution to botanical science and will be welcomed as the only publica-
tion of its class in the field of phycology—B. M. Davis.
North American Flora.»—The general character and scope of this great
work were stated in this journal in connection with the appearance of the first —
part.3 Another part has now appeared, being a direct continuation of the former
one, and both belong to volume 22 in the general scheme
Saxifragaceae—Conimitella, Elmera, and Ceveorics: are established as new
genera, and 30 other genera are recognized; new species are descri under
Lithophragma (8), Tellima, Mitella (2), Pectiantia, Ozomelis, Heuchera (25),
Sullivantia, Therophon (3), Saxifraga, Muscaria (4), Micranthes (12), Spatu-
laria, Leptasea (3), and Heterisia.
Hydrangeaceae.—Neodeutzia is established as a new genus, and g other
genera are ca gee new species are described under Philadelphus (11) and
Edwinia (2).—J. M. C.
Philippine plants——Recent bulletins (nos. 29 and 35) from the Bureau of
Government Laboratories show commendable activity in the study of Philippine
plants. Ermer D. Merritt, botanist of the laboratory, is publishing a series
of papers on new or noteworthy plants, the third and fourth papers appearing
* OLTMANNS, F., Morphologie und Biologie der Algen. pki Band. Allge-
meiner Teil. 8vo. pp. vi+443. Jena: Gustav Fischer. 1905. M 1
North American Flora. Vol. 22. Part 2. Saxifragaceae, Hydrangea JoHN
KUNKEL SMALL, PER AXEL RyDBERG. Cunoniaceae, Iteaceae, Hamamelidaceae,
NATHANIEL Lorp Britton. Petrostemonaceae, PERCY Wnsox. Phyllonoma-
ceae, HENry Hurp Russy. 8vo. pp. 81-191. New York: The N w York Botanical
Garden, December 18, 1905. Subscription price $1.50 for ae part.
3 Bor. GAZETTE 40:74. 1905.
353
354 BOTANICAL GAZETTE [MAY
in the bulletins before us, and containing descriptions of nearly 150 new species.
There are also notes on the Gramineae by ACKEL, including descriptions
of 2 new species; an account of the Seikannucee by Henry N. RIDLEY, 8 new
species being characterized; and 10 new species of Acanthaceae by C. B
Crarke.—)J: M.-C
Aster —In 1902+ E.S. Burcess published a “History of Pre-Clusian Botany
in its relation to Aster;’’ and now a second paper on Aster has appeared,’ which
deals with the “Biotian Asters.’”” Under the head of variation, specific limits in
the genus are discussed; also normal characters and the comparative variability
of organs. There is no group of flowering plants in which such a discussion
would seem more difficult. A systematic treatment of the species is also begun,
84 species being presented with great fullness, 58 of which are published for the
first time; me 10 Sukepees and about 250 subordinate forms are character-
ized.—J. M
Festuca.—C. V. Piper® has published a monograph of the North American
species of Festuca, recognizing 34 species, and characterizing 3 of them as new.
A third subgenus is added to Vulpia and Eufestuca, to include F. conjinis Vasey,
and is called Hesperochloa. There are also notes on several Mexican species
including descriptions of 2 new species. A new word is added to the terminology
of grasses. The word “glume” is restricted to the “empty glumes;’’ while
the “lower palet” or “outer palet” or Se glume” of authors is the Jemma,
a Greek word meaning husk or scale-—J. M
Plants of the Bahamas.—Dr. C. F. Mirtspaucu, Field Columbian Museum,
has issued the first paper? of a series dealing with the flora of the Bahamas, Amar-
anthaceae, Euphorbiaceae, Rubiaceae, and Verbenaceae are presented, and a
new species of Solanum is described. New species are also described under
Iresine (2), Argythamnia (2), Euphorbia (3), Chiococca, Lantana, Valerianodes,
and Callicarpa; and two new genera (Nashia and Pseudocarpidium) of Ver-
benaceae are established.—J. M
Lichens of Santa Cruz—A. W. C. R. Herre® has published an account
of the foliaceous and fruticose lichens of the Santa Cruz peninsula, which is a
natural biological region lying west of San Francisco Bay and extending south-
4 Mem. Torr. Bot. Club, 10.
5 BurcEss, EpwaARD SANFORD, Species and variations of Biotian Asters, with
discussion of variability in Aster. Mem. Torr. Bot. Club 13: 419. figs. 108.
1906.
6 PrpeR, CHARLES V., North American species of Festuca. Contrib. U. S.
Nat. Herb. 10: 1-48. pls. 1-15. 1906.
7 MittspaucH, C. F., Praenunciae Bahamenses. I. Field Columb. Mus. Bot.
ieee 1906.
8 HerRE, ALBERT W. C. T., The foliaceous and fruticose lichens of the Santa
Cruz pcame Fy California. Proc. Wash. Acad. Sci. 7: 325-396. 1906.
1906] CURRENT LITERATURE 355
ward to Monterey Bay. Species are described under 22 genera, Parmelia being
the largest with 14 species; and new species are characterized under Cetraria,
Usnea, Parmelia, and Gyrophra.—J. M. C.
Die naturlichen Pflanzenfamilien —Part 223 continues the families of mosses
by BrotHERus, Hedwigiaceae being concluded; Fontinalaceae, Climaciaceae,
Cryphaeaceae, Leucodontaceae, and Prionodontaceae being completed; an
Spiridentaceae being begun
The first part of the second supplement has also appeared, including the litera-
ture of 1899-1904 in reference to gymnosperms and monocotyledons, with a
few pages beginning the dicotyledons.—J. M. C.
Index Filicum.—The sixth, seventh, and eighth fascicles of CHRISTENSEN’S
work® have appeared with great promptness, carrying the references from Glei-
chenia Cunninghamii to Polypodium basiattenuatum. It should be urged upon
colleges and libraries that so useful and thankless a task should be supported
by adequate subscriptions.—J. M. C.
Text-book of pharmacognosy.—A new textbook of pharmacognosy by G1Lc'?
is worthy an English edition. It is the best illustrated text for ordinary student
use that has appeared. The work would be still more valuable if a greater num-
ber of cuts showing the anatomical elements as they appear in powder had been
included.—Raymonp H. Ponp.
Plants of Bermuda.—A list of plants collected by the author in Bermuda
in 1905 has been published privately by A. H. Moore of Cambridge, Mass. The
pamphlet contains 22 pages, 3 plate reproductions of rs and descrip-
tions of new species of Rhynchospora and Elaeodendron.—. rg
Das Pflanzenreich.tt—Part 24, ee in amet of this year, contains the
Aponogetonaceae by KrAusE and ENGLE g d.—J.M.C.
NOTES FOR STUDENTS.
Items of taxonomic interest.—J. Carport continues (Bull. Herb. Boiss. IT.
6:1-17. 1906) his account of the mosses collected by the Swedish Antarctic Expe-
dition, describing nineteen new species from S. Georgia Island and 5 from the
Antarctic lands.—Pa.rprin adds (idem 18-22) 5 new species to the Chinese flora.
—H. Curist lists (idem 45-58) the ferns of Costa Rica, which is astonishingly rich,
and describes 8 as new.—I. Tuérr10r (Bull. Acad. Int. Geog. Bot. 16:40. 1906)
gives a 2-line diagnosis of two new Leptodontia from New Granada, with other
° CHRISTENSEN, C., Index Filicum, etc. Fasc. 6-8. Copenhagen: H. Hag-
erups Boghandel. 1905 and 1906. Each 3s. 6d :
= Gre, Ernest, Lehrbuch der Pharmacognosie. 8vo, pp. vii+368. Berlin:
™t ENGLER, A., Das Pflanzenreich. Heft. 24, Aponogetonaceae by K. KRAUSE
assisted by A. ENGLER. pp. 22, figs. 9 (718 M1.20. Leipzig: Wilhelm Engelmann.
1906.
356 BOTANICAL GAZETTE [way
new species from China.—E. L. GREENE characterizes (Ottawa Nat. 19:197.
1906) a new Antennaria from Athabasca.—F. von HOHNEL describes (Ann.
Mycologici 3: 404. 1905) a new genus, Unguicularia, near Pezizella, and 3 new
species of fungi—E. BRAINERD adds 2 new names to New England violets (Rho-
dora 7:245-7. 1905).—M. L. FERNALD (idem eae differentiates from Cyno-
glossum virginicum a new species, C. boreale—J. A. CUSHMAN concludes (idem
251-266) his enumeration of the desmids of New ee listing 253 species
and varieties against the 74 hitherto reported. He describes several new ones.—
M. L. FERNALD (idem 8:11 and 22. 1906) describes a new Geum from Vermont
and a new Salix from Maine; attempts to clear up (idem 31) the American forms
called Arenaria verna, including a new species; and characterizes (idem 69-71)
2 new species of Streptopus.—In a presentation of Astragalus and its segregates
as represented in Colorado, P. A. RypBERG (Bull. Torr. Bot. Club 32:657-668.
1g05) recognizes 17 genera, 7 of which (Alelophragma, Jonesiella, Phacopsts,
Ctenophyllum, Microphacos, Cnemidophacos, and Diholcos) are characterized
as new; and in his 16th paper on the Rocky Mt. flora (zdem 33:137-161. 1906)
he describes new species under Rumex, Sphaeralcea, and Senecio (6); establishes
as new genera Crunocallis, Naiocrene, Erocallis (all three near Claytonia), Cor-
nella (Cornaceae), Oreochrysum, Platyschkuhria, Chamaechaenactis, and Pren-
anthella (all four Compositae)—In his 6th paper on the Hepaticae of Puerto
Rico (idem 1-25), . Evans establishes Rectolejeunea and Cystolejeunea as
new genera.—In a sth paper describing new species of Uredineae (idem 27-34)
J. C. Arruur establishes the new genus Ceratelium.—J. K. SMALL (idem 51-57),
in a 2d paper on N. Am. Polygonaceae, describes new species under Eriogonum
(8) and Polygonum.—L. M. UnpERwoop and F. E. Lioyp (idem 101-124)
describe 17 new species of Lycopodium from the American tropics——L. M.
UNDERWOOD (idem 189-205) characterizes new species of pteridophytes from
the United States-under Asplenium (2), Stenochlaena, Tectaria, and Selaginella.
—R. ScHLECHTER (Engler’s Bot. Jahrb. 39: 1-100. 1906) in a study of the flora
of New Caledonia establishes 3 new genera (Coilochilus, Pachyplectron, Gon-
atostylis) of Orchidaceae and one (Trilocularia) of Balanopsidaceae.—U. Dam-
MER (tdem 20-22) describes Actinokentia and Nephrocar pus as new genera of New
Caledonian palms.—W. SuxsporF (Oesterr. Bot. Zeits. 12:5-7, 26, 27. 1906) has
described new species of Washington plants under Sanicula, Lomatium, Anten-
naria, Lasthenia, Pyrola, Navaretia, Orthocarpus, and Aphyllon (2).—A. A.
HELLER (Muhlenbergia 2:1-164. 1905-6) has published an account of his Cali-
fornian collections during 1905, including descriptions of new species under
Eriogonum (3), Montia, Delphinium, Ranunculus, Thysanocarpus (2), Litho-
phragma, Ribes, Amelanchier, Lupinus (14), Vicia, Acrolasia, Boisduvalia,
Glaux, Apocynum, Gilia (4), Solanum, Pentstemon, Castilleia, Orthocarpus,
Malacolepis (Compositae)—Max FLEIscHeR (Hedwigia 45:65-87. 1906), in
concluding his paper on new families, genera, and species of mosses, describes
Baldwiniella, Homaliodendron, Pinnatella, and Penzigiella as new genera of
1906] _ CURRENT LITERATURE 357
Neckeraceae.—U. DamMeErR (Notizblatt Kénig. Bot. Gart. 4:171-173. 1905
describes a new genus (Kinetostigma) of Guatemalan palms.—E. JANcZEWSKI
(Bull. Acad. Sci. Cracovie, pp. 13. Jan. 1906), in his second paper on Ribes,
presents the species of the subgenera Ribesia and Coreosma, including new Cali-
fornian and Mexican species.—A. Borzr (Notarisia 21: 14-16. 1906) describes a
new genus (Zoddaea) of Chlorophyceae (Chroolepidaceae) from a Mediterranean
island.
Heredity.—A lecture on heredity and the origin of species by MAcDoucaL?
not only presents the author’s views regarding the several more prominent
evolution hypotheses, but also makes the first public announcement of impor-
tant results of his own researches on the causes of mutatio
While not denying the possibility of other means of peodiicticns of species,
he holds that hybridization and mutation are the only demonstrated methods
by which new species have arisen. He attributes a popular belief in the Neo-
Lamarckian hypothesis to the supposed effects of garden practice, and these
supposed effects are supposed to be due to the prevalence of vicinism and the
vegetative propagation of bud-sports. Several ‘unsurmountable objections”
are opposed to the Neo-Darwinian hypothesis of natural selection of slight varia-
tions as a universal method. He would distinguish orthogenesis from deter-
minate variations, limiting the former to an internal perfecting force which
evolves rudimentary organs and develops them to ieangs structures without
any reference to selection; while the latter he would allow as a part of every
method of evolutionary procedure, in that no structure may vary to any other
Structure too much unlike itself. This is a very important discrimination theo-
retically, but it is clear that in most cases a practical distinction between ortho-
genesis and determinate variation as here defined would be an impossibility,
since the “morphological possibilities” may be estimated only by what does
appear.
The effects of isolation and of self- and cross-fertilization are held ix
be problematical.
€ greater part of the lecture is naturally devoted to the mutation cultures
of DE Vries and himself. Besides Oenothera Lamarckiana, the "following three
species have been shown to be in a state of mutation: O. grandiflora, O. bien-
nis, and O. cruciata. ‘Parallel mutations’’ are exemplified by two observed
origins of nanella-forms, i. e., forms with linear petals. A consideration of the
mutating and mutant species leads to the conclusion that plants are made up
of complex groups of unit characters, that some of these characters may exist
for an indefinite time in a latent state, that anew character that departs widely
from the parental condition is more variable than the homologous character
of the parent species, and that at the same time it is less closely correlated.
'? MacDoveat, D. T., Heredity and the origin of species. Lecture given before
the Barnard Botanical Club, Columbia University, ae 18, 1905. The Monist,
Jan. 1906. 32 pages. Printed and distributed in advan
358 BOTANICAL GAZETTE [May
The author substitutes for a period of mutation the conception of a nearly
constant frequency of mutation. Thus, one plant in twenty of O. Lamarck-
tana is a mutant, but only one in two hundred of O. biennis. In others there
may be one in ten thousand or one in a million.
Doubtless the most important fact presented is the result of investigations
to determine the cause or causes of mutation. The introduction of strong
osmotic and weak chemical solutions into the ovaries of Raimannia odorata
shortly before fertilization, appears to have produced a large number of individ-
uals of a hitherto unknown type. These new plants have a shorter life-cycle
than that of the parent and are profoundly different in many characters. ey
have already bloomed and fruited, and obviously constitute a potential species.
If this new species holds its characters in succeeding generations, this discovery
will be one of far-reaching importance, as the first real clue to the causes which
may effect mutative changes in plants—GrorGE H. SHULL.
Graft-hybrids——Noit has made a careful morphological, anatomical, and
cytological examination of the supposed graft-hybrids between Crataegus mono-
gyna (stock) and Mespilus germanica (scion) in the Dardar Garden at Bron-
vaux near Metz, Germany.'3 Three branches, starting from the callus where
stock and scion meet, present unmistakable evidence of their hybrid origin, each
branch showing a different combination of the parental characters.
A consideration as to the possibility of graft-hybrids, in the light of present
knowledge of the behavior of the hereditary substance, leads to the conclusion
that they must originate through nuclear fusions in the callus or not at all; and
moreover, that the studies of NEMEC upon asexual nuclear fusions gives a basis
of observed fact which warrants the affirmation that graft-hybrids are possible.
The cytological examination of the several hybrid branches showed that their
cells do not possess double the normal sporophyte number of chromosomes; there-
fore, if these hybrids originated from the fusion of two vegetative cells, this process
must have been followed by some method of chromosome reduction. This pre-
sents no insurmountable difficulty, since NEMEC found that after 78 hours no
nuclei were found which had more than the normal number of chromosomes,
though many such were observed soon after fusion.
The greater resemblance of one of the hybrids to Mespilus, and of the other
two to Crataegus, and the change of one of the latter from nearly typical Cra-
taegus to near one of the other hybrid forms, are explained by assuming that in
each fusion one nucleus remained in its accustomed cytoplasmic surroundings,
and that the other nucleus, moving into unaccustomed surroundings, w
weakened or injured that, when the degeneration took place which reduced the
romosomes to their normal number, the weakened or injured chromosomes
contributed the fewest determinants to the hybrid nucleus, thus giving the hybrid
13 Nott, T., Die Pfropf-Bastarde von Bronvaux. Sitzungsber. Niederrhezin.
Ges. f. Natur-u. Heilkunde Bonn, 1905. Separate, 34 pp.
|
i
1906] CURRENT LITERATURE 359
a greater resemblance to the species furnishing the stationary or ‘“mother-”’
nucleus.
The investigation shows that neither stock nor scion is itself of hybrid origin
and that there can be no reasonable doubt that these are true graft-hybrids.
The only other similar case that has attracted much attention is that of Lab-
urnum (Cytisus) Adami, and about this plant there has been so much contention
that, in the absence of other authentic graft-hybrids and with the disappearance
of the original tree, it seemed best to many botanists to consider the original obser-
vation and record to be in error. Nott prints the original account in full, and
decides, after considering the possible sources of error and misinterpretation,
that the internal evidence in favor of this statement compels belief in its truth.
The final demonstration must lie in a reconstruction of the same ora similar
hybrid, experimentally, and on this work Nott has been engaged for a number
of years, as yet with wholly negative results; but the rarity of the phenomenon
makes this quite to be expected, and the author still hopes by improving his
technic to succeed in re-creating Laburnum Adami.—GrorcE H. SHULL.
Experimental variation —K.rss' presents a paper which deserves special
attention because of the experimental data recorded, because of the author’s
effort to make a closer analysis of the problem of experimental variation, and
because a substitute for DE Vrres’s intracellular pangenesis is offered. Long
experience with the behavior of algae and fungi under artificial conditions, prob-
ably as much as the results stated in this paper, has convinced the author that in
the last analysis all variations must be referred to the influence which external fac-
tors exert upon the inherent potencies of the organism. From this point of view
the fundamental problem of experimental variation at once appears to be to deter-
mine the potential amplitude of variation for species. This problem is to be solved
by the application of as great a variety of conditions as possible. Some of those
used by the author are temperature, darkness, wounding, and artificial food.
The results obtained with Campanula trachelium and Sempervivum Funkii show
that the accepted taxonomic limits of a given species are easily transgressed when-
ever external conditions favor the expression of potencies inherent in the organism.
Trial clearly shows that the potency of external conditions is much greater before
the inception of organs than after. If, for example, nutrition is the determining
factor for a given variation, it makes little difference whether the necessary nutri-
tion status is established by one external condition or another
Over one hundred pages are used to expound the author’s view of the cor-
relation of variation and environment and to present a polemic criticism of intra-
cellular pangenesis. The results with Sempervivum Funkii show that those
characters which can appear as specific within the genus can by proper method
be induced to appear upon a single species. A species therefore is to be
4 KLEBS, GEorG, Ueber Variationen der Bliiten. Jahrb. Wiss. Bot. 42: 155-320.
pl. 1. figs. 27. 1906
360 BOTANICAL GAZETTE [MAY
characterized only by its constant relation to the outer world, and the author
believes any other definition is artificial and arbitrary. More explicitly, a
species is defined as comprising all those individuals which have arisen by
vegetative ee or by self fertilization, and which for many generations
r like conditions have shown identical characters. This definition is not
arbitrary to a reviewer, provided the assumption of a specific structure upon
which it rests is not arbitrary. If by definition potencies can never transgress
the limit prescribed by the specific structure and variation is merely the expression
of potencies, how have species arisen by variation? The potencies of the author
are merely inherent capacities to respond to certain combinations of external
conditions and are purely immaterial as compared with the pangens of DE VRIEs,
“which are material and carry the unit characters. To some investigators this
discrimination will probably appeal as being an interpretation closer to nature
and more logical from the strictly physiological standpoint. To others it may
seem as merely a restatement of the conception of DE Vries. The latter might
easily inquire what difference it makes whether a given variation has arisen by
an inactive pangen becoming active or by a hitherto impotent potency becoming
potent——Raymonp H. Ponp.
The lakes of Scotland and Denmark.—At the invitation of Sir JoHN MuRRAY
Dr. C. WESENBERG-LUND'S spent three or four weeks on the Scottish wie
in order to make a comparison with the lakes of Denmark. ile this was a
short time in which to make examination of a new country nevertheless we should
expect interesting results from one who has not only accomplished so much
thoroughly good work in the study of lakes, but has shown unusual skill and
originality in his interpretations.
The -general differences which the author finds between the Danish and
Scottish lakes are the differences which we should expect between shallow and
deep lakes. The Danish lakes have more plankton, more floating and sub-
merged vegetation, and more distinct littoral zones of vegetation. The greater
seasonal variations in the Danish lakes is noted; this, of course, would be
expected from the greater variations in temperature. There are more highly
colored crustacea in the Scottish than in the Danish lakes. The reviewer thinks
the author right in correlating this red color with low temperature, rather than
with elevation as has been done by some other authors.
Among the diatoms the author notes the absence of Melosira and Steph-
anodiscus in the Scottish lakes, with an abundance of Asterionella and Tab-
ellaria. These facts compare well with the differences in America between
the deep and shallow lakes. The Scottish lakes are remarkable for the large
number of desmids. These desmids are of forms that are common in the pools
of the hillsides. The occurrence of these desmids in the plankton, together
«5 WESENBERG-LuNp, C., A comparative study of the lakes of Scotland and
Denmark. Proc. Roy. Soc. Edinburgh 22: 401-488. pis. 2. 1905
1906] CURRENT LITERATURE 30i
with the occurrence of Entomostraca that are also common in pools, leads the
author to the generalization that the limnetic plankton of the Scottish lakes
is of littoral origin, and that the transportation of these forms to become a part
of the limnetic fauna and flora is favored by the steep hillsides surrounding the
lakes, and the extremely narrow littoral region.
The author enters upon a somewhat detailed discussion of the influence
of the organic life upon the lakes themselves, showing how in the Danish lakes
the algae and higher plants make deposits of lime which are partly thrown upon
the beach, and partly fall to the bottom in the limnetic region. In these bottom
deposits it is again worked over by worms and insect larvae, which devour the
remaining organic matter and leave the bottom sometimes composed almost
entirely of lime and clay. In the Scottish lakes the bottom in the deeper portions:
is composed of material largely derived from the littoral and shore regions, and
there is an absence of lime.
The general conclusion is that while the Danish lakes are filling up, the
Scottish lakes will remain with very slight alteration for ages.—C. DwicH
Mars.
Chlorosis.—One of the most notable papers recently published on the type
of diseases which may be classed as chlorosis is that of Baur on the infectious
chlorosis of the Malvaceae. The variegated mallows in cultivation were derive
from a form of Abutilon striatum known as A. Thomsoni, which appeared in a
collection of A. striatum imported into England from the West Indies in 1868.
This plant was found to be capable of transmitting its variegation by grafting.
Baur finds that if the leaves are removed from variegated plants, or if the shoots
are cut back so that no leaves remain and the plants kept in the dark, new shoots
form only two or three variegated leaves, and if those are removed the plants remain
permanently green in the light unless they are again infected from scions of varie-
gated plants. However, if latent axillary buds on the old parts are forced into
growth, these produce shoots with variegated leaves which in turn infect all newly
formed leaves on the plant. When all variegated leaves are removed from a
plant exposed to light, the plant becomes permanently green. Similarly
Scions of the green but susceptible A. arboreum are grafted on defoliated varie-
gated plants, the scions remain green, but here also if a variegated shoot is allowed
‘to develop from the stock it rapidly infects the whole plant. The author concludes
that the variegation in these plants is caused by a substance or virus which is
formed only in the light in the chlorotic parts of the plants; that this virus is
produced only in small excess so that it is rapidly used up if the variegated leaves
are continually removed. The substance is capable of infecting only the embry-
onic leaves and in those it is stored for months in an inactive form. By appro-
priate girdling and grafting experiments the approximate rate of movement and
the path followed was determined. Movement takes place in the cortex and
not with the transpiration stream. When scions of immune 4A. arboreum are
grafted on a variegated A. Thomsoni, they grow vigorously but are not infected;
362 BOTANICAL GAZETTE [MAY
but if scions of some susceptible species are grafted on the former these become
infected, showing that the virus can pass unchanged through the intermediate
piece of A. arboreum. These experiments seem to prove the existence in the
plant of a substance which in its behavior is analogous to the supposed shoot-
forming substance of SAcHs, or the growth enzymes of BEyERINCK.—H. Has-
SELBRING
Anatomy as a test of species—ALrrepD SaRrToN’® has made an elaborate
experimental study of the anatomy of related plants, to test the constancy of
anatomical characters under varying conditions of climate and of soil. .
work was done at the Botanical Laboratory of the Sorbonne and at the Labora-
tory of Plant Biology at Fontainebleau. He calls attention to the fact that there
are two kinds of species recognized in taxonomic writings: one he calls the “ Lin-
naean species,’’ which often bring together under a single name a large number
of different forms; the other he calls the “ Jordanian species,”’ which often consist
of dismembered Linnaean species. These two kinds stand side by side as of
equal rank, all of them based upon varying judgments as to the value of exter-
nal morphological characters.
ARTON Set out to discover whether real species could be detected by their ana-
tomical characters. He reasons that nearly allied species whose anatomical
differences may be exchanged under experiment are not separate species, however
unlike they may appear externally; and that those whose anatomical differences.
are constant under experiment are true species, however similar they may appear
externally. To test this dictum involved a large amount of laborious experi-
mentation and anatomical investigation. The result was to pronounce some
Jordanian species good and others not; and the Linnaean species shared the same
fate. This anatomical method, therefore, furnishes no basis for judgment between
the two types of species; and if it is used, it seems to the reviewer that it will
result in readjusting specific lines without settling anything.
The fundamental weakness in this whole point of view is the idea that there
can be any rigid test for that elusive conception known as a “species” which will
carry it beyond the reach of fallible and hence diverse human judgment. It.
is of great interest to know what anatomical characters will vary under given
conditions, and herein lies the chief value of this investigation; but even here
the conditions are not analyzed so as to be convincing. To regard these char-
acters as outweighing all others is to stir afresh the seething mess of taxonomy.
What we need is not more ‘‘specifics’’ but more hygiene.—J. M. C
Transpiration of evergreens.—Puc.isi"’ has published a paper on the trans-
piration of seven species of Chinese and Japanese evergreen trees and shrubs.
16 SARTON, ALFRED, Recherches expérimentales sur l’anatomie des plantes
affines. Ann. Sci. Nat. Bot. IX. 2:1-115. pls. I-4. 1905.
*7PuGLIsI, M., Sulla transpirazione di alcune piante a foglie sempreverdi. Annali
di Botanica 2:435-468. pi. 2. 1905.
- EE
a
1906] CURRENT LITERATURE 363
His objects were (1) to compare the winter transpiration of these species in Rome
with that already observed by Kusano in Tokio during the winter; and (2) to
compare the winter and summer transpiration in Rome. One set of experiments
was made with single leaves exposed 24 hours in GARREAU’S apparatus, and
another series with small twigs (in one case a leaf only) attached to MoLt poto-
meters. The potometer measurements were usually continued for about 8 days
and readings made at 9 A. M. and 5 P. M.
The author found the winter rate of transpiration at Rome decidedly greater
than that at Tokio. He obtained for an average value of the ratio of winter to
summer transpiration at Rome, for all the species examined, 1:3.10. The aver-
age ratio at Tokio of Kusano’s “typical plants” (species not given) was 1:20,
PUGLISI reports that transpiration continued at night during the minimum
temperature of his experiments, 2.6°. The rate of transpiration continued to
increase with the rise of temperature up to and including the hottest days of
July in which experiments were made
There is a notable difference among the plants experimented upon in the
sensitiveness to change of temperature. Measured by GARREAU’s method,
Ficus erecta showed an increase of 1.5 times in the amount of transpiration when
the temperature rose from 6.4° to 21.6°, and Raphiolepis japonica showed an
increase of 7 times for about a degree more of rise than that of the Ficus.
The paper contains many interesting data, but the author has not sufficiently
REAU method. All of the winter measurements were made by this method,
which eliminates the effect of changes in relative humidity at a season when the
actual range of this factor was from 58 to 95 per cent.—J. Y. BERGEN.
Plant breeding in the tropics —Locx" gives a further account of his studies
in plant breeding at Peradeniya, Ceylon. His general conclusions were given
in an earlier paper, and the present contribution describes in detail the experi-
ments with the genus Pisum. Records of climatic conditions are given, and
the changes which were induced in various European varieties on introduction
into Ceylon. There was no gradual adjustment or acclimatization, the change
of stature, habit, etc., being immediate and permanent during the several years
of the investigations.
In all of the experiments wherein the characters are clearly alternate, the
agreement with theoretical ratios is as close as the numbers used would warrant
one to expect, on the hypothesis that the union of gametes bearing the several
characters is purely a matter of chance. The author greatly weakens his paper,
however, by pointing out Mendelian ratios where they are wholly unwarranted by
his data, as for instance in width of pod (p. 371), where a variation curve with
"8 Lock, R. H., Studies in plant breeding in the tropics. Ann. Roy. Bot. Gardens
Peradeniya 2: 357-414. 1905.
364 BOTANICAL GAZETTE [MAY
millimeter classes showing the frequencies 1, 6, 5, 8, 4, 5, 2, I, is grouped in a
ratio of 7 wide :17 medium :8 narrow or nearly the expected 1:2:1. Every
variation curve of purely chance variates can be arranged in this way by counting
one-fourth of all the variates from each extreme, leaving the group between
the quartiles as the 50 per cent. intermediates expecte
The author reaffirms in a general statement the ealuseen offered in his
first paper,'? to account for the appearance of certain nova; but makes an inter-
esting observation in disagreement with that explanation, apparently without
noting the discrepancy—the new character of the pigmented parent which
was changed to the active state by crossing. He now states that he could
occasionally observe the mottled pattern like a faint water mark in the white
arent, and its occurrence there makes this an excellent new evidence that the
mottled character is not latent in the usual sense of being inactive, and that
it is not present in the pigmented parent, but being possessed by the white
parent is simply invisible owing to the lack of pigment.—GEORGE SHULL.
Spraying potatoes——Strewart, Eustace, and Srrrinr?° have published
the extensive results secured by them during 1904 in their seties of experiments
in the prevention of potato diseases by spraying. The results secured during
previous years should ‘be read in this connection.2? During 1904 a total of
of the Experiment Station at Geneva, while the remainder were conducted as
“farmers’ business experiments” in various parts of the state. The experi-
ments at Geneva form a part of a ten-year series of experiments designed to give
average results for various seasons. The other experiments should yield valuable
data year after year as to the actual net gains to be expected from the spraying
against potato diseases under actual farm conditions. At Geneva five sprayings
increased the yield 233 bushels per acre, while a gain of rg1 bushels was secured
from three sprayings. This gain was mostly due to the longer growth of the
plants made possible by the prevention of the late blight and the rot which follows
it. In the business tabu covering a total of 180 acres, the net gain per
acre due to spraying was $24.86. The average loss from blight in New York
State during 1904 was not less than 60 bushels per acre. The sugyestion is made
that the community hire some person to do all their spraying, thus effecting a
saving of time and labor—E. Mrap Witcox.
Alternation of generations in Phaeophyceae.—Strasburger?? agrees with
OLTMANNs that there is no alternation of generations in the Phaeosporeae. He
19 See Bor. GAZETTE 39: 303-304. 1905.
20 Stewart, F. C., Eustace, H. J., and Srrrine, F. A., Potato spraying experi-
ments in 1904. Bull. N. Y. Geneva Exp. Stat. 264:93-204. pls. I-16. I map. 1995.
2t Potato es experiments in tg02. Bull. N. Y. Geneva Exp. Stat. 221:
235-2063. 1
Sendi apeaion experiments in 1903. Bull. N. Y. Geneva Exp. Stat. 241. 1903.
22 STRASBURGER, oes Zur Frage eines Generationswechsels bei Phaeo-
phyceen. Bot. Zeit. 64:1~7. 1906.
1906] CURRENT LITERATURE 365
believes that here, and also in the Sco mare the germination of the zygote
will be found to be accompanied by a reduction of chromosomes, and that conse-
quently a diploid, or 2x generation cannot be present. ‘The absence of a diploid
generation explains why parthenogenesis occurs so readily. Forms like the Dicty-
otaceae, which have a diploid generation, must be widely separated phylogeneti-
cally from the Phaeosporeae. The thallus of the Fucaceae is diploid, while from
the initials of oogonia and antheridia to the mature eggs and sperms the condi-
tion is haploid, or gametophytic. The rather surprising view is expressed that
the antheridia and oogonia of the Fucaceae are not homologous with those of
the Dictyotaceae, but that they correspond rather to the tetraspore condition
of the latter group. SrRAsBURGER makes the statement that the gametophytic
generation begins with the complete separation of the 2x (doppelzahligen)
chromosomes, because this separation furnishes the condition for the formation
of sexual products. He does not indicate any more definitely that he would
regard the spore rather than the spore mother-cell as the first term of the
gametophyte.—CHARLES J, CHAMBERLAIN.
Diseases of sugar cane.—LEwToN-BRAIN finds that the root-disease of the
sugar cane in Hawaii?s is probably due to the fungus Marasmius sacchari, known
to cause a similar disease in other countries. In Hawaii the fruiting body of the
fungus has not yet been found. The Yellow Caledonia variety seems to be resist-
ant to the disease while the Lahaina and Rose Bamboo are most severely injured.
Ratoons are more injured than plant canes. Since this fungus is a soil-infesting
fungus it may be controlled by liming the soil and through cultivation.
Cops?: has recently published suggestions as to the inspection and disin-
fection of sugar cane cuttings to prevent the spread of sugar cane diseases. **
The cuttings should be made with care to prevent the shattering of the ends,
which permits the entrance of fungus parasites, and they should then be care-
fully inspected to get rid of any diseased ones that may be present.. “Pick-
ling” the cuttings in Bordeaux mixture is recommended, and a large part of the
paper is devoted to methods of doing this work on the large scale required on
a sugar plantation. The cuttings may also be sprayed with Bordeaux mixture
in the ditch just before being covered—E. Mrap Wi1cox.
Asparagus rust.—Smiru, as a result of his further studies of the asparagus
rust problem on the Pacific coast, finds that the oe may be effectively controlled
a duet spray of flowers of sulfur.?5 25 «
or entirely prevented by the proper app
23 LEWTON-BRAIN, L., Preliminary notes on root disease of sugar cane in Hawaii.
Div. Path. and Phys. Has Stat. Hawaiian Sugar Planters’ Association Bull. 2: 1-39.
1905.
ae Eosi. N. A., The inspection and disinfection of cane cuttings. Div. Path.
and Phys. Exp. Stat. of the Hawaiian Sugar Planters’ Association Bull. 1:1-35,
pls. 8. 1905.
2s SmiTH, R. E., Further experience in asparagus rust control.
Stat. Bull. 172:1-21. figs. 7. 1906.
Calif. Exp.
366 BOTANICAL GAZETTE [MAY
The important thing is to get the tops of the plants covered with a perfect coating
of the sulfur dust before the rust appears; the first application should be made
as soon as the tops have made some little growth, say about three weeks after
cutting stops, and a second and perhaps a third application should follow at inter-
vals of about one month each. In some cases it may prove advisable to spray
the plants first with some soap spray, to be followed by the sulfur dust to insure
the sulfur adhering to the plants. If the best grade of flowers of sulfur be employed,
it may be possible to cover an acre with about fifty cents worth. SmirH insists
also upon the supreme importance of destroying all wild asparagus plants near
the regular fields, since on these the rust first appears, and from them the field
soon becomes infested.—E. MEAD WItcox.
Germination in myxomycetes.—In a preliminary paper on the germination
of the gia of myxomycetes, JAHN recognizes two distinct types of germination.?°
The first type is represented by a single species of Ceratiomyxa, whose spore
contains four nuclei, the spore content escaping as an amoeba which immediately
ivides into eight swarmspores. In the second type, embracing all other myxo-
mycetes, the spores are uninucleate and produce a single swarmspore. Regarding
the conditions of germination, the following observations are made. The amoeba
escapes by rupturing the spore-membrane by osmotic pressure, and if this is
increased germination is prevented. The spores of Stemonitis do not germinate
when placed in water, but if after being soaked they are allowed to dry they will
germinate readily in water, an observation formerly made by LisTER. JAHN
concludes that such spores contain a latent enzyme which is made active by the
process of moistening and drying. Since maltose but not other sugars causes
germination, this assumption is strengthened, as maltose is the decomposition
product produced from glycogen stored in the spores—H. HASSELBRING.
Blight canker.—WueETzEL has. published the results of his study of a canker
of apple trees in New York state?® due to the same organism, Bacillus amylo-
vorus, that is responsible for the well-known fire blight of apples and pears. This
canker has been known in a general way for some years, but this seems to be the
first demonstration, by the usual inoculation experiments, of the bacterial nature
of the disease. Infection seems to take place only through wounds, and these
may be due to pruning, to accidental wounding or ‘“‘barking” of the tree, or to
the work of insects. The germ also enters at times through “water sprouts,”
since cankers are often seen to appear about the base of such blighted shoots.
Pear trees known to be affected with the blight should not be allowed to remain
in the neighborhood of an apple orchard, and great care chould be taken to prevent
the dissemination of the germs on the pruning instruments. Some variation
in resistance to the canker was noted—E. Mrap Witcox.
27 JAHN, E., Myxomycetenstudien. Ber. Deutsch. Bot. Gesells. 23:489-497- 1906,
28 WHETZEL, H. H., The scat canker of apple trees. Bull. Cornell Exp. Sta.
236:99-138. figs. 51-84. 1
eS”
1906] CURRENT LITERATURE 367
Excretion of acids by roots—Kunzr”® has extended the work of Mo.iscu,
PRIANISCHNIKOW, and CzAPEK on the general chemistry of the excretion of acids
by roots, including a study of similar activity as exhibited by mycorhizal fungi.
CzAPEK’s conclusion that the excretions do not contain free mineral acids is
confirmed, but the presence of acid salts of the mineral acids is denied and the
corrosive effect is attributed exclusively to the organic acids. Tests with about
two hundred different species widely separated in systematic standing shows
that many plants do not excrete enough acid from the roots to be detected by
litmus. Fungi excrete much more actively and it seems probable that they are
more potent as soil disintegrators than the roots of higher plants. The author’s
work tends to increase his belief in mycorhizal symbiosis. Whether intentional
or not it seems a serious deficiency to omit the date from 23 of the total number
of 35 citations—RAyMoND H. Ponp
Algae of northern seas—An interesting paper by Smmmons%° discusses the
history and relation of the algal floras of the North Atlantic and North Pacific
to one another and to that of the Polar Sea. The Atlantic and Pacific Oceans
are believed to have received a large number of species from the Polar Sea of
Tertiary times, especially just before the ice age, when the algae were driven
southward. Some of these became established and never returned to their old
situations, but settled and became variously modified in their new life ee
Others re-entered the Polar Sea with the retreat of the ice. This is a paper w
will bear careful study by those familiar with the algal floras of the North Meies
and Pacific, especially in comparison with BORGESEN’s Algal vegetation of the
Faeréese coasts noticed in this journal 41:71. 1906—B. M. Davis.
Grape diseases.— BUTLER?" has published some observations on three grape
diseases: red-leaf, shrivel, and root-rot. The red-leaf seems to be a disease closely
related to diseases known as folletage, rougeot, and California disease. Like the
other diseases named, the red-leaf is probably due to disturbances in the balance
between absorption of water by the roots and transpiration. It is possible that
the disease may be checked to a considerable extent by increasing the fertility
of the soil so as to render the plants more resistant. The shrivel is found mostly
among the white grapes and is also due to disturbed nutrition or deficiency of
water supply. The root-rot is similar to if not identical with the disease described
in French literature as pourridie. It is caused by one or perhaps several fungi
not yet fully determined. It often kills the vines in a single season but it may
only kill the vines after three or more years.—E. MEAD WILcox.
9 Kunze, Gustav., Ueber Siaureausscheidung bei Pokecage und Pilzhyphen
at tte ss Jahrb. Wiss. Bot. 42: 357-303-
3° Smumons, H. G., Remarks about the relations at di floras of the Northern
Atlantic, the Polar Sea, and the Northern Pacific. Beih. Bot. Centralbl. 19: 149-
194. 1905.
31 BUTLER, O., Observations on some vine diseases in Sonoma county, Cali-
fornia. Bull. Calif. Exp. Stat. 168:1-29. 1 pl. figs. 1-5. 1905
368 BOTANICAL GAZETTE [MAY
Preserving plants.—PoLvacct, speaking before the Italian Botanical Society,3?
commends his method, proposed in 1900, for preserving plants in a watery solu-
tion of sulfur dioxid. Specimens so preserved in 1900 have retained perfectly
their form and are in as good condition for sectioning as when fresh. He has
- improved the method of making the solution and has devised a means of retain-
ing perfectly the green color. To make the solution, place sodium bisulfid in a
large flask, add sulfuric acid drop by drop, and conduct the gaseous SO,
through water, which quickly becomes saturated and may be preserved for use
as needed. To retain green color immerse the material in a 1 per cent. watery
solution of copper sulfate, leaving it 24 to 48 hours ee to the consistence
of the tissues; then transfer to the preservative solution.—C. R. B.
Teratology in Salix.ss—Morr records various cases of teratology in the
flowers of two Californian willows, S. /asiandra Benth. and a hybrid of S. lasiandra
Benth. and S. babylonica L. In making the statement that no mention has been
made for Salix of an intimate association of microsporangial and megasporangial
tissue he overlooks an earlier account by the reviewer,3+ who described and figured
equally intimate associations. According to Mort, the abnormalities indicate
that the ancestral Salix flower consisted of a pistil and two stamens with a four-
parted perianth, the present unisexual condition having been reached by the
suppression of the organs of one sex. Hybridization seems to offer the most
‘likely explanation of the abnormalities—CHARLES J. CHAMBERLAIN.
Nectaries of Cruciferae——Vi1LANr has made an exhaustive comparative
study of the nectaries of Cruciferae35 and concludes that on the basis of their
number the Cruciferae can be divided into four types, and on the basis of their
position into generic groups. The diverse forms as to nectaries are referable
to one primitive type, having four nectaries, two of which are at the base and
external to each stamen, constituting an external dimerous cycle, and two at the
base and between each member of each pair of long stamens, constituting an
internal dimerous cycle. The tetramery of the corolla is only apparent, the
whole flower being purely dimerous. The nectaries function both for securing
Cross pollination and protection. —C. R. B.
Morphology , of Chloranthus.—Miss Herren M. Armour3® has published
the results of her study of Chloranthus, especially interesting as extending our
3? Pottacct, G., Nuovo metodo per la conservazione di organi vegetali. Bull.
Soc. Bot. Ital. 1905: 242.
33 Morr, Witt1am WarNER, Teratology in the flowers of two Californian wil-
lows. Univ. Cal. Publ. Bot. 2:181-226. pls. 19-20. 1905.
34 Bot. GAZ. 23:147-179. pls. 12-17. 1897.
35 VILLANI, A., Dei nettarii delle Ce e del loro valore morfologico nella
simmetria florale. Malpighia 19: 399-439. 1906.
36 ArMouR,’ HELEN M., On the morphol of Chloranthus. New gone
5:49-55- pls. +4. 1906.
.
i
1906] CURRENT LITERATURE 369
knowledge among the more primitive Archichlamydeae. The development
of the floral parts and both lines of sporogenesis. were studied, the general con-
clusion being reached that the characters agree with those of the majority of
the primitive Archichlamydeae. In the ovule the sporogenous tissue develops
as a mass of cells, from which usually a single mother-cell is selected, a late differ-
entiation of that cell which is quite characteristic of gymnosperms. The linear
tetrad is formed, ae there is the greatest irregularity in the selection of the func-
tioning spore.—J. \
Embryo of Symplocarpus.—C. O. RosENDAHL‘? has made a preliminary
announcement of a remarkable feature of the embryogeny of Symplocarpus-
The endosperm destroys both integuments and pushes into the basal tissue of
the ovule. There is a short, thick suspensor, and at this end of the ‘“ protocorm”
both hypocotyl and plumule are differentiated. The developing embryo destroys
the endosperm and all of the ovular tissue except at the very base, and thus comes
to lie free in the ovary cavity! This means that the ‘“‘seeds” of Symplocarpus
are naked embryos. The full paper, with illustrations, will be looked for with
interest.—J. M. C
Nucleoli in mitosis—The nucleoli in the vegetative cells of Equisetum
arvense, according to G. BARGAGLI-PETRUCCI, play an important part in mitosis.3*
In the resting nucleus there is a single centrally placed nucleolus. At the begin-
ning of mitosis, the nucleolus divides, one of the daughter nucleoli passing to each
pole of the nucleus, where it perforates the nuclear membrane and passes to the
apex of the achromatic figure.
While the figures are doubtless accurate, much more evidence will be required
to establish the contention that the nucleoli play such a rdle in mitosis. —CHARLES
Resistant potatoes.—The selection of races resistant to disease is one of the
most promising methods of meeting many kinds of plant diseases. The resist-
ance of Irish potatoes to blight, tuber rot, and scab has been worked out in an
admirable way by StEwarT.3° More than one hundred and fifty varieties were
tested and their difference in susceptibility is graphically represented. In general,
varieties having strong, woody, moderately branched, upright stems, and medium
sized, rather thick, firm, hairy leaves were more resistant than varieties possessing
weak, much branched, decumbent stems with large, thin, smooth leaves.—F. L.
STEVEN NS.
37 ROSENDAHL, C. Oro, Preliminary note on the embryogeny of Sym plocar-
pus foetidus Salisb. Science N. S. 23:590. 1906.
8 BARGAGLI-Perrucct, G., I nucleoli durante la cariocinesi nelle cellule mer-
istematiche di Equisetum arvense. Nuovo Giorn. Bot. Ital. 12: 699-708. pl.
39 Stewart, W., Disease resistance of potatoes. Vermont Agr. Exp. Sta. Bull.
122. 1906.
37° BOTANICAL GAZETTE [MAY
Glycogen and paraglycogen.—A posthumous paper on this subject by Prof.
Lfo Errera‘? has been edited from his notes by Dr. J. Massart. It contains
observations on the wide-spread occurrence of one or other of these bodies among
the fungi, and their sparse distribution, perhaps only less observed, among the
lower plants and animals, and possibly even among sea weeds and seed plants.
ErrerA had interested himself in this reserve food for many years and had accu-
mulated a great mass of bibliographical notes on it, which have been reduced
to order and herein published.
Epidermal gaps —Years ago MitpE and Kny and THomae described the
occurrence of interstitial gaps in the winged bases of the leaves of three Osmundas ~
and a Todea, and similar gaps have been found in the epidermis of floral leaves.
Now Lersurincer,‘" incidentally, in the course of some cytological studies, finds
such gaps in the epidermis of the scales of Alluim Cepa, which seem probably
connected with the secretion of mucilage —C. R. B
Germination of moss spores——Tresoux contributes testimony upon the
controverted question of the necessity of light for the germination of moss spores.*?
He finds twenty mosses of the most diverse families and three liverworts, a much
larger number than has ever been tested before, able to germinate without light
and (contrary to HEALD’s results) without cane sugar to replace its stimulating
action.—C. R. B.
Haustoria of Osyris——Pizzoni has published} an extended account of the
haustoria of Osyris alba, supplementing the note of FRAySsSsE*+ which unex-
pectedly forestalled Pizzont’s paper after all his observations had been ennieid
He treats of ‘the ee relations to host, contents, duration, and dimensions
of the haustoria—cC. R. B.
Nitrogen for maize.—Soave+s finds that nitrogen supplied to maize in ammo-
nium nitrate does not need to undergo nitrification in order to be available, so
that, other things being equal, this compound of nitrogen is to be preferred to
sodium nitrate, there being no delay in assimilation as affirmed by GERLACH
and Vocet.—cC. R. B.
L., Glycogéne et “ Bia sak chez les “rr Recueil
de og ae Brust I: 343-379.
4" LEIBLINGER, G., Ueber inters s tien ene ee, in der pflanzlichen
Epidermis. 0 Dekteck. Bot. Gesells. 23: 387-396. pl. 17.
#* TREBOUX, O., Die Keimung der Moossporen in ihrer Sate zum Lichte.
Ber. aoc oe Casal: 23:397-401. 1905.
43 Pizzoni, P., oars alla conoscenza degli austori dell’ Osyris alba.
Annali di Bot. 4: ol pl 3.
Frayssg, A., Surla SEE etl? anatomie des sugoirs del’ Osyris alba. Compt.
Rend, Acad. Sci. Paris 140: 270-1. 1905.
45 Soave, M., L’azoto ammoniacale e-l’azoto nitrico nello sviluppo del mais,
Annali di Bot. 4:99-114. 1906.
NEWS.
Dr. A. F. BLAKESLEE has been appointed recently upon the botanical staff
of the Philippine Commission.
PROFESSOR JoHN M. Coutrer has been elected an honorary member of the
Royal Botanical Society of Edinburgh.
Dr. A. B. RENDLE has been appointed Keeper of the Department of Botany
in the British Museum (Natural History).
Dr. C. F. Mittspaucu, Field Columbian Museum, is spending three months
in Europe, chiefly in the investigation of museums. .
Proressor Dr. Frrprano Cavara has been appointed director of the Botan-
ical Garden of Naples in succession to the late Professor DELPINO.
Mr. W. R. Maxon, U. S. National Herbarium, has just spent two months in
Costa Rica collecting plants for the New York Botanical Garden.
Dr. F. Rosen has been advanced to the position of professor of botany and
director of the Institute for Plant Physiology at the University of Breslau.
THE INFORMATION is just published that during 1903 there were 1,352,548
visitors at Kew Gardens; and during 1904 the number increased to 1,579,666.
Tue AcapeMy oF Scrences in Munich has made a grant of M2500 to Dr.
Rosz, curator of the Botanical Museum, for zoological and botanical investiga-
tion in Central America.
NG HIS PRESENT VISIT to the United States, Professor Huco Dr VRIES
will deliver lectures at the summer session of the University of California and also
at the University of Chicago.
AT THE RECENT Franklin Bicentenary at Philadelphia, the University of
Pennsylvania conferred the degree of doctor of laws upon Professor Huco
EVRIES, who was present to deliver an address.
Dr. Lester F. Warp, for twenty-five years the paleobotanist of the United
States Geological Survey, has left the ranks of professional botanists by accepting
the position of professor of sociology in Brown University.
IN THE RECENT DISASTER at San Francisco the building of the California
Academy of Sciences was destroyed, containing the very valuable collection of
Californian plants. It is reported that the types were saved by the heroic efforts
of Miss Atice Eastwoop.
Dr. E. N. Transeau, Alma College, Michigan, has been appointed a mem-
ber of the staff of the Station for Experimental Evolution at Cold Spring Harbor.
He will work at evolutionary problems from the ecological side.
if 371
372 BOTANICAL GAZETTE [MAY
Miss FreDA DetMeRs, formerly botanist to the Ohio Experiment Station,
lately teacher of botany in the Columbus North High School, has been appointed
instructor in botany in the Ohio State University and has already assumed the
duties of that position.
A BRIEF biography of the late Professor Lto ErrerA has been distributed,
containing, in addition to an appreciative notice of his life and work, a fine like-
ness in photogravure, and a bibliography numbering 168 titles—a marvelous
output, considering its high quality, for a man only 47 years old.
Dr. D. S. Jonson, Johns Hopkins University, is spending April and May
in Jamaica at the Cinchona station of the New York Botanical Garden. Dr.
ForrEST SHREVE, of the same university, is spending a year at the same station
in work on the physiology and ecology of the forest of the Blue Mountains.
ER CRONE, assistant in the Botanical Institute of the Royal Agri-
Salvnad College at Poppelsdorf, near Bonn, returned from a journey in Spain
ill with typhus, from which he died on the 23d of February last. He had already
published some recent studies on a cause of chlorosis and was prosecuting further
work in chemical physiology.
THE EDITOR of Flora and Sylva, having issued three volumes in serial form
at less than cost, has become convinced that it does not appeal to a sufficient
number of readers to justify its continuance as a monthly magazine. Hereafter
it is to appear as a yearly volume, but otherwise precisely as heretofore. The
next volume will appear in the autumn.
THe Marine Brotocicat Station of the University of Washington, which
is located at Friday Harbor, Washington, will open for its next season June 25
and will close August 5. The station is supplied with a steamer for transporta-
tion and deep dredging and offers good advantages for biological work, together
with the pleasures of camping and tramping. @Professor Bruce Fink, Iowa
College, will have charge of the botany.
Proressor Huco De Vries opened the co of spring lectures at the New
York Botanical Garden on April 21st by an ress upon “The correlation
of characters in plants.’’ Other lectures in thi§/course will be given by W. A.
Morrit_, ArtHuR Hotticx, L. M. UNDERWO6oD, C. S. GAGER, MARSHALL
A. Hows, G. V. Nasu, H. H. Russy, D. T. Doveat, and N. L. Britton.
INsTRUCTORs in botany at the Biological Laboratory of the Brooklyn Insti-
tute located at Cold Spring Harbor, Long Island, for the summer of 1906 are
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in charge of plant ecology, and Mr. H. H. York, associate in botany. The lab-
oratory will be open during July a August, the courses beginning July 5, and
continuing six weeks. |
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The Botanical Gazette
A Montbly Fournal cere all Departments of Botanical Science
Edited by — M. Coulter and CH S R. BARNES, with the assistance of other members of the
botanical Mall of the Guivenicy of Chicago.
Vol. XLL No. 6 e Issued June 30, 1906
CONTENTS
SOME STUDIES REGARDING THE BIOLOGY OF BUDS AND TWIGS IN WINTER
(WITH EIGHT FIGURES). Karl M. Wiegand eee
THE LIFE HISTORY OF POLYSIPHONIA VIOLACEA. CONTRIBUTIONS FROM THE HULL
BoranicaL Laporatory LXXXIII. Shigeo Yamanouchi - 425
THE STRUCTURE AND DEVELOPMENT OF THE BARK IN THE SASSAFRAS
(WITH NINE FIGURES). Howard Frederick Weiss - 434
BRIEFER ARTICLES.
THE DIstRIBUTION AND Hairs OF SOME Common Oaks. E. J. Hill - - - = us
CURRENT LITERATURE.
en REVIEWS - - - - - = = a Es my ae a = 448
ANICAL DICTIONARY.
MINOR NOTICES ee eters ent nec a ee pereat iron aires anernnepfimmmee ree orton orl
NOTES FOR STUDENTS 4. ee tg eS gee Rs Ber ee ey ek tS ey
NEWS mm > - - = -~ - - ~ ow. = e -~ - - - - 45
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Entered August 21, 1896, at th es d-cl; tter, under Act of Congress March 3, 1879.
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An Indispensable Book for Students of Botany
Methods in Plant Histology
SECOND EDITION—ILLUSTRATED
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VOLUME XLI NUMBER 6
DOTANICAL. GAZETTE
JUNE, 1906
SOME STUDIES REGARDING THE BIOLOGY OF BUDS
AND TWIGS IN WINTER.’
Karu M. WIEGAND.
(WITH EIGHT FIGURES)
Durinc the winter months in temperate and arctic climates,
the meristematic tissues of shrubs and trees assume a more or less
completely dormant or resting condition, and become separated
from the surrounding atmosphere by tissues of varying thickness
and varying degrees of resistance to the passage of water vapor.
A detailed study of these structures during the cold period has brought
out many interesting facts ordinarily escaping casual observation.
In the twigs the cells of the cambium lie close together without
intercellular spaces, but the cortical cells usually do not touch at
the corners, and consequently in the cortex there is a more or less
elaborate system of intercellular spaces. The main structural pro-
tective measure seems to be the firm epidermal layer with heavily
cutinized outer wall, which is always present at this period.
There were no stomates on the twigs in any of the species I exam-
ined. Gas diffusion takes place mainly through the lenticels; but
perhaps to a slight extent also through the cuticle its:lf. All the
living cells contain a large amount of water, 51-55% in most fruit
trees, 63% in Forsythia, and the quantity in each species is remark-
ably constant, rarely varying more than four to five per cent., and
usually even much less.
Regarding the time during the ‘ital summer when the bud
fundament is first distinguishable, ALBERT? found that, out of
r aay from the Department of Botany of Cornell University. No. 105.
2 ALBERT, P., Beitrage zur ee der Knospen einiger Laub-
hélzer. Forstlich naturw. Zeitschr. -3:345, 393- 1894-
Shoe
374 BOTANICAL GAZETTE [JUNE
15 species of trees bearing scaly buds, the first leaf fundament
in one (Betula alba) was present as early as May, in three at the
beginning of June, in eight at the beginning of July, in two August 1,
and in one not until September. The flowers were always formed
later than the leaves. Some of the naked buds he found to start
early in the previous season (Elaeagnus, Cornus); others, as for
instance Robinia Pseudacacia, did not start until the spring of the
year in which they were to unfold. He found that in general the
buds were further progressed at the beginning of the winter the
farther north the plants were native.
BEHRENS? found that in fruit trees the Sewers are first distin-
guishable at a later date, as for example, in the cherry during July,
and in the pear about August 11,
My own observations lead me to believe that in many cases the
fundaments are present quite early. The buds of the peach were
well formed July 15, and small buds were evident in the leaf-axils
of forest trees as early as June 1. This suggests that, in some cases
at least, the bud fundament may be present as early as the unfolding
of the previous winter’s buds.
Those that start quite early have usually reached an advanced stage
in development by the time cold weather overtakes them in the
fall. The rudimentary flower or shoot for the next season, together
with all its organs, is present in the buds of some species, as for
instance in the horsechestnut; while in others a varying number
of nodes and internodes are thus stored. To inclose so elaborate
a structure a certain number of leaves have been modified into
scales which closely overlap, or are firmly cemented together at
their edges around the young shoot. Such buds are found espe-
cially upon treés and shrubs with definite annual growth. The
scales are usually composed of several layers of parenchymatous
cells with intercellular spaces, moderately firm and slightly cuti-
nized epidermis, on the inner side, and a very strong heavily cuti-
nized outer epidermis, usually supported by mechanical tissue of
varying amount beneath. The parenchymatous cells of all or
3 BEHRENS, J., Entwickelung und Bau der Bliitenknospen unserer Obstbaume
und Obststraucher. Gartenflora 4'7:269. 1898.
4 For published descriptions of bud structure see: BEHRENS, J., /. ¢.; FEIST,
A., Ueber die Schutzeinrichtungen der Laubknospen dicotyler Laubbaume wahrend
1906] WIEGAND—BUDS AND TWIGS IN WINTER 375
all but the outermost scales are living and contain a large amount
of water throughout the winter. The inner scales are frequently
almost destitute of epidermal thickenings and are quite green and
fresh. Because of the much larger size of the cells in the scales,
and much larger vacuoles, there is much more water present in these
structures than in the young shoot whose cells are small and nearly
filled with protoplasm. This it will be seen is an important con-
sideration when the buds freeze during the winter. The abso-
lute amount of water in the whole bud is however very nearly the
same as that in the young bark, being about 51 to 55% for the
fruit buds examined; and, as in the bark, this amount is remarkably
constant for the species.
The proportion of space occupied by the young shoot varies with
the species and nature of the bud. In flower buds this proportion
is usually greater than in leaf buds. In many cases only a very
small fraction of the total volume is shoot-tissue, all the rest being
composed of scales; but in other cases, as for instance the flower
buds of pine, almost the whole volume is made up of cones, leaves,
and stem; while the scales are very thin, dry, and firmly ‘cemented
together. In this case of course nearly all the water is located
within the young shoot. The spaces between the various organs
and scales usually contain air alone; but in some cases, as for example
in apple and horsechestnut, there is also a large amount of wool
present in which the organs are seemingly imbedded. In Populus
and some other trees the spaces are more or less completely filled
with resin.
Buds of most indefinite growers differ from those of the majority
of definite growing trees in two essential ways: in the slight develop-
ment of the fundament, and in the usual absence of scales. The
young shoot is most frequently represented merely by a growing
ihrer Entwickelung. Nova Act. Leop. Carol. Ak. Naturf. 2: 303-344, 1887, Ref. Bot.
Centralb. 36:43. 1888; ScHUMANN, C. R. G., Anatomische Studien iiber die Knos-
Penschuppen von Coniferen und dicotylen Holzgewachsen. Biblioth. Botan. 15:32.
Cassel, 1889, Ref. Bot. Centralb. 42:275. 1890; Griss, J., saat zur Biologie
der Knospen. Jahrb. Wiss. Bot. 23:637. 1892; Lussock, J., On buds and stipules.
Jour. Linn. Soc. 30: 463-532. 1895; 33:202-269. 1897; CapuRA, R., Physiologische
Anatomie der Knospendecken dicotyler Laubbaiume. Breslau, 1887, pp. 42; Mrkosca,
K., Beitrige zur Anatomie und Morphologie der Knospendecken dicotyler Holzge-
Wichse, Sitz. Konig. Akad. Wiss. Wien Math. Wiss. Kl. 74':723-755- 1877-
376 BOTANICAL GAZETTE [JUNE
point without well-developed lateral organs, and can therefore be
protected more economically by being sunk in a pit produced by
a ring-like growth of cortex and cork, as is commonly the case.
This pit is then closed at the mouth by an ingrowth of the cork
itself, as in Gleditschia, or by a feltlike mass of hair, as in Robinia
and other species. .
In the case of the large buds with the shoot considerably advanced
in growth, the bud-scale method seems the only feasible way of
' covering them. Another advantage in this method lies in the tele-
scopic expansion of which scaly buds are capable early in the season
while unfolding. Growth is thus permitted, but at the same time
the protective qualities are not lost. In the maples and_horse-
chestnut the tube formed by the enlarged scales often reaches the
length of 2 to 8°". By this means buds may open early in the spring
and still be protected from excessive transpiration. Scaleless buds
usually remain nearly dormant until later in the spring when the
weather conditions are not so severe.
PHYSICAL PHENOMENA OF BUDS AND TWIGS WHEN NOT FROZEN.
The scaleless buds of the indefinite growing trees and shrubs
grow very little before or during the winter. In the autumn the
very limited growth is soon stopped by the advent of cold weather,
and from this time until late spring scarcely any change can be
detected. With the scaly buds however it is otherwise. From
the inception of the fundament in July or June until cold weather
there is a very considerable growth resulting in the buds of various
sizes found upon the different species of trees during the winter.
Little accurate work has been done towards determining the char-
acteristics of this growth, but the results obtained in our laboratory
by W. M. Morcan during the fall of 1901 seem to show that in
the case of fruit trees the growth is very uniform and gradual up
to about November 15. In some cases slight fluctuations occurred
which could not be accounted for, and in one or two instances these
seemed periodic; but on the whole there appeared neither accel-
eration nor retardation until the time mentioned, when the increase
in size ceased quite abruptly. From the middle of November until
March 1 there was no growth in peach buds, the curve remaining
almost exactly horizontal and fluctuating very little. On March 23,
1906] WIEGAND—BUDS AND TWIGS IN WINTER 377
several days of warm weather occurring, the peach buds began
to grow rapidly and uniformly until April 23, one month later,
when they came into flower. With the apple and apricot the results
were very much the same. Growth almost ceased November 15,
and from this time until March 1 the increase was apparent but
exceedingly slight, amounting to only $ to 1%. Renewal of activ-
ity began March 1, and from this time until April 23, seven weeks
later, when the apricots flowered, and eight weeks later, when apple
buds opened, the growth was very rapid. The curve after growth
began was not so gradual as in the peach, but became much accel-
erated just before the flowers appeared.
Mr. Morcan’s observations were as a matter of fact quite exten-
sive, but only the above summary can be given here. At intervals
of one week through the fall, winter, and spring, buds were taken
from the same tree and as nearly as possible from shoots of the
same vigor, a large number were measured, and the average taken
as representing the size at that time. It was found impracticable
to measure the same bud at different times, owing to the difficulty
of manipulating the micrometer out of doors on very cold days,
as well as to the fact that the measurements were liable to be taken
at different temperatures cach time. A Zeiss cover-glass measurer
was found the most convenient instrument for the work. From
the tables thus made a great many curves were plotted representing
the changes in various fruit trees. The results, however, agreed
very well, and in the peach, apricot, and apple were as stated above.
From these careful observations, therefore, contrary to the general
belief, it seems that fruit buds at least do not grow to any extent
in winter. Their swelling period is confined in the north to a few
weeks just previous to the opening of the bud. Regarding our
forest trees and shrubs no accurate work seems to have been done
toward the determination of their curve of growth. From casual
observations, ‘I am inclined to believe that a majority will be found
to agree with the fruit buds. This seems to be truce of the sugar
maple, whose buds are practically as large in November as in early
March, also of the ash, oak, etc. On the other hand, the buds of a
few plants, as, for instance, Salix discolor, Ulmus fulva, and Ulmus
scabra, seem to increase in size early in February. However, actual
measurements are necessary to determine these. points.
378 BOTANICAL GAZETTE [JUNE
Kuster’ found that during a specially mild winter the buds
of maple did show a very slight growth both in the lateral organs
and in the young axis. No new organs were started either in maple
or other species examined, except rarely in Alnus cordijolia.
ALBERT® found that practically all buds became dormant soon
after leaf-fall until spring again. The first change in spring was
a stretching of the tissues, further development of the parts taking
place only later.
It is a known fact in physics that the amount of heat absorption
varies, among other factors, with the color of the body investigated.
In other words, the same body if colored differently will absorb
a varying amount of heat from a constant source, depending upon
the color. Winter buds and branches are in many cases highly
colored, and the question naturally arises as to how this affects
the heat absorption of the bud during the winter and spring months.
Regarding the extent to which color will affect heat absorp-
tion, in addition to the records in works on physics, the experiments
of WHITTEN’ are interesting. He found that thermometer bulbs
wrapped in muslin of different colors, green, purple, black, and
white, or with pieces of muslin of these various colors spread over
them, or with the bulbs coated with a wash of similar colors, showed
a marked difference in reading when exposed to bright sunlight.
The average difference between the black- and white-washed bulbs
was 16°, between the white and purple 15°, and between the white
and green 13°. At one time a difference of as much as 21° between
the white and purple bulbs was found.
However, the actual experiments with buds have been rather
few and the results are not so definite as one might wish. The
most elaborate were those of WHITTEN described in the above-cited
report. He selected a row of peach trees containing several vari-
eties, and whitewashed them during the winter. During warm
days of the unusually changeable winter the unwhitened buds
swelled considerably, and during subsequent cold spells most of
5 KisTEr, E., Ueber das Wachsthum der Knospen wihrend des Winters. Beitr.
Wiss. Bot. (FUNFSTUCK) 2:401. 1898.
6 ALBERT, P., Beitrage zur Entwickelungsgeschichte der Knospen einiger Laub-
hdlzer. Forstl.-naturw. Zeitschr. 3: 345-376, 393-419. 1894.
7 WHITTEN, J. C., Winter protection of the peach. Bull. 38. Missouri Agric.
Exp. Station, April 1897.
&
|
|
:
|
1906] WIEGAND—BUDS AND TWIGS IN WINTER 379
them were killed. The unwhitened buds swelled and grew percep-
tibly before any swelling could be detected in those that were whitened.
The difference in size March 20 was plainly shown in drawings
of the sections of the two classes of buds. Whitened trees came
into bloom about one day later than unwhitened trees of the same
variety. In 1896-97, owing to a more moderate spring, the differ-
ence in time of flowering was still greater. The whitened buds
of each variety opened two to six days later than those that were
not whitewashed.
The differences in the actual time of flowering, however, does not
express the difference in time of the swelling of the buds. The
whitewashed buds did not begin to swell until almost time for the
flowers of normal trees to appear, while the unwhitened ones began
to swell three or four weeks earlier, as shown by the drawings above
mentioned.
These experiments of WHITTEN seem to show that in the peach,
at least, the dark-purple color of the buds tends to cause earlier
activity in the spring, accompanied by earlier swelling and flower-
ing. The only doubt, it seems to me, lies in the effect of the white-
wash upon the growing tissue. As mentioned below, some non-
porous substances seem to retard respiration perhaps to such an
extent that growth also is retarded, but the whitewash would seem
porous enough to escape this criticism. In a more recent paper
Wuirren® has shown that the temperature within whitened and
unwhitened twigs differs by several degrees. In bright sunlight
the difference was as much as 15° C., the unwhitened being the
warmer. The whitened twigs were nearly of the same tempera-
ture as the atmosphere.
WuitTENn® has also shown that purple peach twigs transpire
considerably more than green ones. This was probably due to
the greater temperature and is probably an additional factor in
the winter-killing of the peach.
8 WarrreN, J. C., Preventing frost injuries by whitening. Pacific Rural Press
60:276. 1900.
9 WHITTEN, J. C., Das Verhiltnis der Farbe zur Tétung von Pfirsichknospen
durch Winterfrost. Inaug. Diss. Halle. 1902. p. 35. See also in this connection
Macoun, W. T., Some results of experiments in spraying, etc. (whitewashing ”
Tetard bud development.) Ontario Fruit Growers Ass. Rep. 1899: 100, and “ Experi-
mental Farms,” Canada, 1899:92.
380 BOTANICAL GAZETTE [JUNE
Wishing to determine the effect of natural color and surface
of buds upon the absorption of heat, I carried through several experi-
ments with horsechestnut buds, which gave some interesting results
as follows.
On February 8, in bright sunshine, a large horsechestnut bud
was obtained and the scales dissected away, care being taken that
they were not unnecessarily injured. Two thermometers previously
tested as to accuracy were obtained, and over the bulb of one the
bud scales were carefully imbricated and firmly held in place by
a few turns of black thread. There was enough resin present to
cement the scales firmly together and thus form an artificial horse- °
chestnut bud with the thermometer bulb in place of the normal
shoot. The instruments were then placed on a table out of doors
’ and in the shade where they were allowed to lie. As soon as the
readings were nearly the same,'® the table and instruments were
carried to a place in full sunshine, care being taken that the two
bulbs projected about 6.5°™ beyond the edge of the table so as
not to be affected by direct radiation from the surface of the latter.
The readings were taken as follows:
‘TABLE I.
Horsechestnut bulb and naked bulb, from shade out of doors to sunlight. (See jig. I.)
Maked bein. | Momoeneh to ee ° Difference
ge" F. pire 0:00 oO PS (axe €)
33 32 0:10 I mo:
34 33 0:30 I ae
34.5 34 1:10 0.5 0.2
35 35:7 bee fe) 0.7 0.4
36 a5 2:00 I 0.5
36 38 2:30 2 L-6
or 39 3:00 2 EO
37 40 3:30 3 to"
37 42 4:00 5 2.8
38 44 4:30 6 ae
38 45 5:10 7 3.9
38 48 6:00 10 5.5
38 49 7:00 II 6.1
38 50 10:00 rz 6.6
30 5L 14:00 12 6.6
39 52 19:00 13 7.2
3) 53 24:00 14 7.7
39 54 29 :00 Ig 8.3
‘© They could not be made to read the same because the overcooling point in the
bud had been reached.
1906] WIEGAND—BUDS AND TWIGS IN WINTER 381
60
SS Ee
50 ——
ri
40 L
a a
a
380
me oe» « & & &® & & Sk & 8S & SK EF B
———horsechestnut bulb; .<.s.07:- naked bulb. Abscissas represent 5° F.;
BIG. he
ordinates, 2 minutes. See Table I.
No further rise was noted, although the instruments remained
in place more than an hour. The experiment was repeated several
times during the spring, both with the same bud and with a fresh
one, in every case with practically the same result, namely a much
faster rise in the bud-bulb, amounting finally to an excess cf 5 to
12° C. over the other bulb.
That the above differences were due mainly to the dark brown
color of the bud seems probable from another series of experiments
in which the naked thermometer-bulb was coated with brown
drawing ink. Readings were taken under the same conditions
as before. In these cases the difference was but slightly in favor
of the horsechestnut bulb, probably because of the less highly pol-
ished surface of the ink bulb. It would seem, therefcre, that the
point is fairly well demonstrated that in the case of the horsechestnut,
at least, the color does make considerable difference in the absorb-
ing power of the bud as regards heat. Although no experiments
were performed, there seems no reason why the same shculd not
be true also of other dark-colored buds.
With these results in mind I made a few observations one spring
to see if there was any relation between the color of the buds and
the time of swelling and opening. It seemed reascnable to expect
that the darker the bud, and consequently the more heat absorbed,
the earlier the bud would swell and open in the spring; and all the
more because this was the exact conclusion reached by WHITTEN
with his peach experiments. Unfortunately, accurate record of
the time of swelling and of opening were not kept; but still I believe
382 BOTANICAL GAZETTE [JUNE
some important general tendencies can be made out from the notes,
and therefore the following table is inserted:
Name Color of buds Ga
Magnolia acummaitay .. fe. osc. use ee yellow-gray or olive late
Ailanthus eli having Chg eR ea. bias yellow-gray late
Boer Negu nde n.6 naGkbs.- ecards hire a: pale, whitish medium
Ju _acsant senha hi pao ta ats be eens se pale, grayish very late
Quereus alla. o 53.16 Hees oad wae oes gray very late
Salix dite Lc c inte a oraitas. Stara tak ops yellow early
Pops Prandidentatd. 455 esses: pale very early
Ringe Wulearis. howl asis cows eso pale, yellowish early
Betula Bee: ee Or see ee olive late
Fagus SnGAiCaUe << os dress se cd euxs olive medium
Paes “dilatata ne Cre eae ee yellow-brown medium
Salix) COrdatan: 4 tactea erwiss ste itosnattcaaesivs, Ae olive or purple ly
Sie TOCA GATIO 6, 5 4.50 Cesena ar ah te een brown medium
Sank SeriGed ve coe okie sle, a2 worsiawesae eo). brown-purple early
MNES. POUR. se uals oa gs BOF eed wc brown-purple early
Acer obec eg wala PRR ata hee bark olive-brown medium
its ROIOONA ns oie be Se ee red medium
Aesculus Teecectaaet a Us ach otra ah dark brown » early
Aber pactariate 6 25 oo ns 8 cae Ys red very early
Prunus persica .......-00,.-00ssese> purple very early
Other species ee Sohgan and Pyrus.. red, brown, or gray early
CYRISORUE COCKER. 0 ooo ce occ deus a red o house olive medium
Dilnriis: Hikwase: tcc geseec cure oe heres dark he early
NENTS Sen Dita. ch cca c tat ele ees black very early
Stee WRN iy vinu ds 64 ex ba eadaes Hae black very early
It is not to be expected that the time of opening or even the time
of swelling will in all cases be proportional to the color of the buds
alone. The protoplasmic characteristics of the particular species
or genus undoubtedly play a very important part; but bearing
this in mind the following suggestions appear in the above table.
Nearly all of the light-colored buds are also late to swell and open.
None of the dark red or especially the black buds open late. On
the other hand, a few light buds, as for example those of Syringa,
Populus, and Salix alba, open quite early. This may be due to
more easily aroused protoplasm than is present in most buds. Pos-
sibly if these buds were black they would open still earlier and
therefore suffer injury from the frost; thus the present lighter color
may serve as a means of protection.
During the spring of 1901, about March 1, some experiments
were started to determine if possible whether other buds might
be influenced by color in a way similar to the peach buds white-
1906] WIEGAND—BUDS AND TWIGS IN WINTER 383
_ washed by WauirtTeN. Instead of whitewash, black paint was
used to see if they might be made to open earlier. Two kinds of
paint were prepared, one made of lamp black mixed with linseed
oil, the other of lamp black and xylol. Buds and twigs of Syringa
vulgaris, Ailanthus glandulosus, Populus dilatata, and apple were
treated to a coat of oil paint; while some others of Syringa, Ailanthus,
and apple were coated with xylol paint. The results were as follows:
Syringa.—Xylol-painted buds much behind the normal during vernation;
they looked unhealthy, one or two being entirely dead. Oil-painted buds never
began to swell, all dead.
Atlanthus.—The xylol paint made no difference with the killing back of
the branches nor with the development of the buds. Oil paint prevented the
swelling of the buds; they never opened.
A pple.—Xylol-painted buds much behind the others; one completely dead.
Oil-painted ones all dead.
Populus dilatata.—Oil-painted buds showed much more rapid swelling than
normal. When just opening the blackened buds were 6 to 8™™ longer.
These results are evidently in the main exactly opposite what
we were led to expect. I suspect that the explanation lies in this,
that the coating of the surface of the bud with paint prevented res-
piration, and thereby inhibited growth just as did varnish used
on naked buds as described later in this paper, although it is possible
that some toxic property of these substances might have had some-
thing to do with the matter. The xylol furnished a much more
porous layer than did the oil, and the inhibition was therefore much
less. The buds of Populus dilatata are normally almost completely
infiltrated with resin in the spaces between the organs and on the
surface, and consequently may have some other means of obtaining
oxygen for respiration. The coating even with oil paint, therefore,
did not injure them. On the contrary, the black color seemed
to cause an acceleration in growth.
In concluding this part of the subject we may say that in the
climate of New York, buds during the winter seem to remain in
an almost dormant condition until a short time previous to their
opening in the spring. In Missouri swelling of peach buds began
much earlier than in New York.
Color through its power of absorbing heat seems to have some
effect upon the growth of buds in the spring. Early buds are in
384 Pinna ‘BOTANICAL GAZETTE [JUNE
most cases dark, and artificial darkening, when unaccompanied
by deleterious factors, seems to accelerate the opening.
PHYSICAL CONDITIONS IN FROZEN BUDS AND TWIGS.
From many inquiries it would seem that very few people are
really sure. whether free ice is actually present in buds in winter.
Nevertheless, this is one of the most common phenomena connected
with the winter condition of trees and shrubs. To put the matter
on a firm basis of observation I undertook, during the winter of
tgo1, to section buds of various trees during cold pericds and to
determine under the microscope the amount of ice present. The
method employed was as follows. Early in the morning, at about
sunrise, after a fall of temperature to —18° C. or below, a table,
microscope, razor, needles, slides, and cover glasses were placed
in a shady situation in the open air, where they were allowed to
become thoroughly cooled. Free-hand cross sections of the various
buds were then made, and mounted on the slide. For a mounting
medium cedar-wood oil was found best. A small quantity of this
in a vial was allowed to cool with the instruments. One important
advantage in the use cf cedar-wood oil over those of a denser nature
lay in the fact that it did not congeal at the low temperatures of
the experiment. The ice remained unmelted in the preparation
and could be observed at leisure; or if the thawing process was
under study the slide could be carried to a warm room and placed
under another microscope. The melting ice was unable to evapo-
rate from the section, and therefore it was easy to determine whether
the water was all reabsorbed, and the approximate rate of absorp-
tion.
The ice was found to occur always in broad prismatic crystals
arranged perpendicular to the excreting surface; and usually formed
a single continuous layer throughout the mesophyll of the scale or
leaf, to accommodate which the cells were often separated to a con-
siderable distance (figs. 2, 3, 4). This ice sheet was composed of
either one or two layers of the prismatic crystals, depending on the
water content of the adjacent surfaces, and was often as thick as the
whole normal scale. The cells surrounding the ice, having lost their
water content, were in a more or less complete state of collapse,
1906] WIEGAND—BUDS AND TWIGS IN WINTER 385
depending upon the resistance of the walls, and often occupied
a space smaller than the ice itself. These cells were uninjured,
however, and would resume their normal condition on thawing.
In all cases more ice was found in the scale than in the young shoot;
never between the scales but always in the mesophyll. The cells
of the embryonic shoot were so much smaller and their water content
so much less, that frequently it was difficult to detect any ice forma-
Fic. 2.—Populus dilatata: cross-section of bud, showing ice in bud-scales and
foliage leaves.
tion whatever; but ordinarily very minute and numerous masses,
at least, were scattered between the cells, and sometimes there were
large masses such as appear in the outer organs. In young anthers
the ice often filled almost the entire anther cavity, and in it the pollen
grains were imbedded in a completely collapsed state.
The results of the observations regarding the occurrence of ice
in buds may be summarized briefly as follows. The temperature
Was ~33;5° C.to = 28°.
386 BOTANICAL GAZETTE [JUNE
1. Tissue packed full of ice in shoot and in mesophyll of scales forming
sheets parallel with the surface; rapidly and completely reabsorbed when the
sections were thawed in oil. Sponging out of sections very marked.—Populus
dilatata (fig. 2) and P. candicans, Prunus serotina and P. virginiana, Betula
enta, Acer Negundo, Pyrus Malus and P. communis, Aesculus Hippocasta-
num.
2. Containing a large amount of ice, but the water tardily reabsorbed on
thawing in oil.—Acer Saccharym, Tilia americana, Ulmus scabra, Crataegus
punctata (jig. 3).
Fic. 3.—Crataegus punctata: cross-section of bud, showing ice in bud-scales and
floral parts.
3. No ice could be found at o°C. Tissue dense, of small cells.—Castanea
dentata, Hamamelis virginiana, Fagus americana, Fraxinus americana, Jug-
lans cinerea, Corylus rostrata, Quercus alba, Hicoria ovata.
4. Other cases—In Pinus Strobus and P. sylvestris there was a moderate
amount of ice in the shoot and in the anther as well as in the inner scales. In
Syringa vulgaris there was a very large quantity of ice in the scales and young
shoot, especially in the anthers (fig. 4). In Viburnum dentatum and Prunus
persica the amount of ice was small, but water was quickly reabsorbed.
1906] WIEGAND—BUDS AND TWIGS IN WINTER 387
Of the twenty-seven plants examined there were only eight that
showed no ice in the buds at —18°C. These eight were sectioned
later at —26.5° C., with the result that in Castanea, Hicoria, Fraxi-
nus, and Juglans numerous minute ice crystals were found between
the cells. It would seem, therefore, that ice may be found in all
buds if the temperature is sufficiently low.
The accompanying illustrations are reproductions of photo-
micrographs taken by the writer during periods of low temperature.
Fic. 4.—Syringa vulgaris: cross-section through flower bud while frozen; the
light spaces filled with ice.
When the mercury registered at zero Fahrenheit or below, freehand
sections mounted in oil as already described were photographed,
the apparatus being set up in the open. The conditions for sec-
tioning were so strenuous that very thin sections could not be obtained,
and hence the rather poor quality of some of the photographs. The
palisade-like ice prisms fill the light areas through the mesophyll of
the scales and young leaves.
388 BOTANICAL GAZETTE — [JUNE
The question naturally arose as to the cause of the difference
in ice content and why ice was absent in the eight species mentioned.
Since lowering the temperature from—18° C. to—23.5° C. caused
the appearance of ice in some, it would seem therefore to be simply
a matter of temperature. But the degree of cold necessary to
cause the separation of ice is proportional to the force which holds
the water in the tissue. This in turn depends upon the relative
proportion of water to cell-wall and protoplasm. We should expect,
therefore, to find in those buds which are difficult to freeze a smaller
amount of water than in other buds; also smaller cell-structures,
since by this latter means the proportion of cell-wall and proto-
plasm is increased. When cells become smaller it is usually the
water content that most rapidly diminishes, the protoplasm follow-
ing at a much lower rate. I have made the following measurements
of the cells and water content in seven of the species in which there
was much ice, and in seven in which ice did not appear at —18° C.
Max. aver. | Min. aver. | Text. of wall | % of water
mm. mm.
A. Ice abundant in. bud-scales, leaves,
nd growing point—
Crntecwtis: pronttata. << 670.23 625 oss 0.040 0.012 thin 49-4
RT ENNOS Ste aie ots or een eae Be O51 0.015 : 46.6
Byrn Vileatigg 6.565. ..55 204 254 0.0045 | 0.009 ot 53-2
Pee MN id a deh 0 ware 0.021 0.009 : 45-9
SerNAl, cues ee a 0.021 0.015 ie 47.6
Populus racine LEP Dee wit 8 = sain to 0.025 0.018 2 39-3
meuttile lites es 5 vic 0 Pa Sh 0.018 0.006 se 37-5
B. Ice not sie at — 715" Cc 7 elles
Querc Pee PN Sa. 3s th elements 0.015 0.006 thick 22-9
orylus:rostratac: < .0csce 8 oe see: 0.018 0.006 a 29.7
Castanea dertata. .. 28 igs cds ees 0.018 0.015 = 25.4
F Westen. co ree ces | 0.008 0.003 26.8
a aphedia can foe ee ee 0.048 0.015 very thick 31-4
JUG CUMTOR, 055 65 eo eg oe | o12 0.003 thi ck 25:9
Fraxinus americana... ..........-;. 0.021 0.003 | 29.8
Our supposition regarding the smaller size of the cells and
smaller water-content in the second group, therefore, seems to be
upheld by these results.
In the twigs ice is also present in very cold weather, where it
may be found in three different localities. TThe largest quantity
occurs in the cortex, where the ice crystallizes in prisms arranged
1906] WIEGAND—BUDS AND TWIGS IN WINTER 389
in single or double series according to the law of freezing tissues.
The ice is more frequently in the form of a continuous ring, or really
a cylinder, extending entirely around the twig, prying apart the
cells of the cortex in which it lies. The outer cylinder of cortex
in such twigs is completely separated from the inner layers when
frozen. In a few species instead of the continuous layer, lens-
shaped ice masses are interpolated irregularly throughout the cortex.
The cortical cells after the withdrawal of the water are as com-
pletely collapsed as were those in the bud scales, but they also
usually regain their normal condition on thawing. In the wood
ice rarely forms in large quantities. It is usually confined to small
masses in the vessels themselves, or, according to some authors,"
sometimes extends in radial plates in the pith rays. In sectioning
twigs, I myself have never seen ice in the wood elsewhere than in
the vessels or wood-cells. In the pith the ice, so far as I have been
able to observe, always occurs within the cells and therefore in very
small masses.
At the time when the buds were sectioned, cross-sections of the
twigs were made and mounted in the same manner. Ice was
found in the cortex of all those in which it was present in the bud,
but usually in proportionately larger quantities. It was also found
in the following species which showed no ice in the buds: Corylus
rostrata, a small amount in large clefts in the cortex; Castanea
dentata, some ice in ordinary small spaces of the cortex but not
aggregated; Hamamelis virginiana, a ring of ice completely around
the stem in young twigs. In Fraxinus, Fagus, and Juglans none
could be found, and Quercus was not investigated.
Since water on freezing increases in volume, one would at first
thought expect the frozen twigs to be larger in diameter than normal.
Such, however, is not the case. In every instance a distinct con-
traction occurred, which in some cases was very marked.*?
_ ™ MULLER-Tuurcau found ice present in the large vessels of Syringa, Cornus,
and in pears, almost completely filling them; and several times he could also dem
onstrate it in the wood-cells. The ice was the most distinct, however, in the vessels
of the grape. Ueber das Gefrieren und Erfrieren der Pflanzen. II. Landw. Jahrb.
™2 Both Sacas and MU@LLER-TuuRGAU have shown that a similar contraction
occurs quite generally when herbaceous tissues freeze. SACHS, J., Krystallbildung
390 BOTANICAL GAZETTE [JUNE
To determine the exact amount of contraction the Zeiss cover-
glass measurer was used. . Pieces about 10°™ long of one or two
year old twigs taken at—18° C. were inserted in the clamps of the
machine, a record taken, and then the whole carried to the warm
laboratory. The increase in size on thawing could be followed
by watching the movement of the indicator as the ice melted, and
when at last stationary another reading was taken. Some results
are given in the following table:
: Frozen Thawed Difference Exp. or contr.
Cormis etolodifera < 00.25% 62. 4.3 2.58mm/ 2, 60mm o.o2mm| expanded
he oF i oP se acs Eire ares 3.38 3-43 0.05 i
"EWR QUGIYDNGUOS Soa ee 2.03 2.10 0.07 si
- Feces ok cath gare. tovalvee sid 3.18 3-39 O.12 i
Pennies Mlstates 66. Peony os 5: 3-17 3.28 O.11
" ee We is © pe ee gk 4.72 4.80 0.08 i
Acer piataniless yee kat 2.82 2.86 0.04 ig
Oe es gare hgh as 3.44 3.48 0.04 =
Pyrus Malus......... 0.2.0.6... 2.97 3.03 0.06 #
WANES Set. yas a arate 3.89 ze 2 0.06
ale MORO Gare nce diy es & 94 5.4 0.10 is
pee ne) Sit a ee eS: 5-9 0.14 .
Many twigs at — 18° C. or below appear very much wrinkled on the
surface as though dried and dead. This is especially true of the
polished shoots of Salix cordata. On very cold mornings shoots
of this species appear as though dead and dry, the bark being com-
pletely covered with fine longitudinal wrinkles. Some of these
shoots were brought to the laboratory and allowed to warm, during
which process the disappearance of the wrinkles could be watched
with ease. In about ten minutes the twigs were entirely smooth
and normal. It was from such twigs that the above readings were
taken. To show more graphically the expansion during thawing,
some twig-sections about 10°™ long were taken from the same willow
and the ends while still frozen dipped in melted paraffin. The
caps thus produced at the ends of the twig were in every case rup-
tured down the side on thawing, leaving in most cases a cleft of
considerable size between the two edges. Twigs of plum were
bei dem Gefrieren u. Veranderung der Zellhiute bei dem Aufthauen saftiger Pflan-
zentheile. Bericht Verhand. Konig. Sachs. Gesell. Wiss. Leipzig, Math.-Phys. Klasse
12:1-50. 1860. Mirrer-Taurcav, H., Ueber das Gefrieren und Erfrieren der
Pflanzen. Landw. Talnb. 9:187. 1880.
.
————ooo
a ee ee
1906] WIEGAND—BUDS AND TWIGS IN WINTER 391
also much wrinkled, those of Negundo and apple showed a slight
furrowing, those of black cherry and pear scarcely any at all.
The photographs of wrinkled twigs of Salix cordata, reproduced
in the accompanying illustrations (fig. 5), were made in the open
at a temperature of —20° C. The same twigs were then placed
in the laboratory, and after about one hour were photographed
again. The slightly wrinkled «appearance in the upper shoot in
D
Fic. 5.—Salix cordata: A and B, twigs ears in the open at—20° C.,
showing etic condition due to contraction; C and D, the same twigs thawed in
the laboratory; the furrows have disappeared except the minute normal striae on
the lower twig.
the second photograph was normal for that shoot when thawed,
during both winter and summer.
It seemed desirable to determine whether this contraction was
mainly in the bark or in the wood or in both. At a temperature
of —18° C. much wrinkled twigs of Salix cordata were collected,
and the following measurements made:
392 BOTANICAL GAZETTE [JUNE
With bark on twig the diameter, 7.80™™ expanded to 8.04™™ on thawing;
difference o.24™™. With bark removed from a small spot for the clamps of the
measuring instrument, the diameter, 5.05™™, expanded to 5.15™™; difference,
Lon
Therefore more than half of the total expansion was in the bark. Thickness
of the bark was 0.5 ™™ on each side; thickness of the wood and pith, 2.05™™
on each side; expansion of the bark, therefore, was 13.5 per cent.; of the wood,
only 2.5 per cent.
With thicker twigs, containing more hard wood, the expansion
would have been still less.
Where the bark was whittled away entirely around the end of
the twig and for some distance back, the expansion of the wood
was not detected; probably because water had passed to the bark
to freeze and being removed there was none to cause swelling again
when the twig thawed.
The explanation of the contraction of twigs on freezing probably
lies in the following considerations. When the water is extracted
from the walls of the wood-cells, the latter contract to a slight extent
just as they do when wood seasons. This accounts for a part of
the shrinkage. The rest and greater part occurs in the cortex.
Here the intercellular spaces are quite large and numerous, and
are normally filled with air. When freezing occurs the ice forms
in the spaces and the cells collapse, while the air is mostly driven
completely out of the twig. The contraction in the cortex will
be approximately equal to the volume of air expelled plus that of
the air compressed minus the expansion of the ice while freezing.
This: is for contraction in all directions; only a portion of this will
be radial, depending upon the structure of the particular species;
much the greater part, however, is radial in all twigs.
With buds the study is not quite so easy. The record of buds
measured at —18° C. and then again after thawing is shown in the
adjoining table.
From this table it will seem that in all cases, except in Populus
and Acer, there was a decided increase in size on freezing and a
consequent decrease when thawing out. In the two named cases
there was a slight contraction as in the twigs. It is not quite clear
why the buds should behave so differently from the twigs. The
only explanation I can offer at present is that the contraction of
1906] WIEGAND—BUDS AND TWIGS IN WINTER 393
| Frozen Thawed Difference oO +e ed
| eae) er een
Cornus stolonifera............ | 2.22mm 2,20mm 9.02mm | contraction
a“ SO veg fal NS Chote l= 2860 2.61 0.05 a
Tilia platyphyllos.......... 3.63 3.54 0.09 i
Pim. Tres dais okie *.i5 3.07 0.08 S
Populus dilatata.... «20.66 ..04. | 2.97 ey | ©.00 °
PS eet eT roe fae 2.03 3.04 0,01 expansion
Acer platanoides............. 5.06 5.08 0.02 -
iy cody emai S ee 4-34 4-34 0.00 s
“s pa rete Pe 4.16 4-17 0.01 =
ie a | ea ae as 4-92 4.88 0.04 contraction
“ee se ae
is Bo eRe Pee het ag eee 4.04 4-03 0.01 -
eee ee 4.92 4.68 0.04 :
Prunus persica............... 2.62 2.59 0.03 '
* Sige Te Te 2.50 2.46 0.04 .
‘a POD alte Sete ies 2.72 2. 66 0.06 .
Prunus americana............ 2.17 2.09 0.08
s Meee ree ee 1.78 Ei71 0.07 ie
ve fe he ec or ae 2.10 2.06 0.04 “
the wood is eliminated, of course, and that the formation of ice tends
to bow out the scales so that they stand less closely together. If
the bud scales curve like leaves in freezing this result is to be expected.
When the temperature rises sufficiently, the buds and twigs thaw
out and regain their normal condition. In the sections under the
microscope the reabsorption was so rapid as in most cases to be
entirely completed when the ice itself had finished thawing. This
results in an active sponging-out movement in the sections as
the cells recover from their collapsed condition (fig. 6). On account
of the rapidity it is frequently difficult to keep the point of observa-
tion in the field of the microscope. Thawing seems not to harm
these tissues in the least, no matter how frequently or how abruptly
it is done. I have often tried the experiment of transporting twigs
abruptly from —18° C. to the warm laboratory at 21° C. and back
again several times, thus alternately thawing and freezing them.
No matter how many times this was repeated no injury could be
detected in the buds, even when subsequently placed in the green-
house to grow.
Buds and twigs do not thaw at o° C. if the rise in the surrounding
temperature is gradual, as it is in atmospheric changes, but at a
much lower degree. The thawing like the freezing is proportional
to the temperature, and is almost if not quite completed when the
394 BOTANICAL GAZETTE [JUNE
freezing point of the tissue is reached. This, in the case of buds;
lies at about —3.5° C. to —2.3°C. Hence, if tissues which have been
subjected to—18° temperature in the open are to be observed with
the maximum ice content, it must be while the temperature is still
low. If in the morning the temperature has risen to —7° C. before
observations are made, very little more ice will be found than if the
cooling to —7° C. had just taken place.'3
6.—Syringa vulgaris: same section as in fig. 3 thawed in the laboratory;
note sponging out of tissue and closing of spaces occupied by the ice.
13 GOEPPERT gives a similar experiment. “Twigs with buds of Cornus mas-
cula, Prunus Cerasus, and Aesculus Hippocastanum were on January 2, 1871, placed
ten hours at a temperature of —16 to —20° C. Then while frozen stiff they were
eri into the warm tube of an oven at 25° C. and placed in water for further obser-
tion. They waht a later just as others that had not been subjected to this
riment.” Some other experiments with herbaceous plants led GOEPPERT to
iain that in most cases alternate thawing and freezing, when taking place many
times, gradually weakened the tissue. Ueber das tseFeleie Erfrieren der Pflan-
zen und Schutzmittel dagegen. Stuttgart, 1883, pe 33.
Warttrn believes that rapid thawing gat freezing is very detrimental to the
a eT
1906] WIEGAND—BUDS AND TWIGS IN WINTER 305
WINTER FUNCTION OF BUD SCALES.
Bud scales are obviously for the purpose of protecting the tender
inner shoot from detrimental external influences; but how is this
protection accomplished? This is a subject regarding which
opinion has varied widely and does still at the present time. I
believe we shall find that the most widely accepted views, strangely
enough, are not the correct ones, even though the subject appears
so simple. We can conceive of such protection taking place along
four lines: (1) by keeping out external moisture; (2) by preventing
the penetration of cold or sudden changes of temperature; (3)
by preventing the escape of internal moisture; (4) by warding off
external mechanical injury. It seems best to discuss each of these
in turn, and in this way determine the extent to which cach one
is operative.
External moisture.
There is a widespread belief that bud-scales function by keeping
out the wet. The subject, however, is a difficult one to determine
experimentally, and I can find no reference in literature to such
experiments having been performed. Let us first determine the
possible ways in which water might be supposed to injure the embry-
onic tissues. First, the cells might absorb too much water and
thus become more sensitive to frost. It seems quite reasonable to
believe that a cold spell following such an event might «nd the life of
the bud completely. Again, through gradual absorption of the air
by the water the latter might replace the air in the intercellular spaces,
thus preventng free respiration. Or again, if a thawing bud were
surrounded by water, the latter, instead of.air, would be drawn in to
fill the vacant intercellular spaces, the final result being the same
as in the last case. Lastly, one might expect that the freezing of free
water between the embryonic foliar and floral parts might cause
mechanical injury.
purple buds and twigs of peach. Green twigs and especially whitened ones warm
up less each day and this color would therefore be protective. I believe it may quite
likely be true that delicate buds might suffer by such violent treatment either from
stimulated activity or increased transpiration, even though hardy trees are apparently
indifferent. Das Verhiltniss der Farbe zur Tétung von Pfirsichknospen durch
Winterfrost. Inaug. Diss. Halle. 1902.
396 BOTANICAL GAZETTE [JUNE
Taking up these in turn, if the cells were so unprotected as to
be capable of absorbing water in this way, they would be expected
to lose a large part again when dry conditions returned, and thus
quickly following frosts alone could do harm. ‘There is also con-
siderable doubt whether sufficient water would be absorbed by
the cells to cause any perceptible difference in sensitiveness. Water
at winter temperatures absorbs air very little, and especially after
having fallen in the form of raindrops it may be considered as nearly
saturated. The air in the leaves would probably be absorbed very
little, if at all, although compression of the air due to capillarity
might allow some water to enter. If the thawing tissue has its
spaces filled with water instead of air, this will not necessarily cause
harm. In experiments on leaves it was found that only the ivy
leaf was unable to recover when the spaces were filled with water.
Many leaves allow the water to evaporate and then become normal.
Mechanical injury is not probable since the air spaces of the tissue
would be elastic enough to overcome the compression of the expand-
ing ice between the organs, and after the tissue froze slight pressure
from the outside on the compressed cells would do no more harm
than the pressure of the ice masses within ordinary tissuc.
However, the greatest objection to this theory, it seems to me,
lies in the fact that protection against moisture might be obtained
in a much simpler manner. The embryonic tissue might be densely
clothed with strigose hairs, or densely glaucous, either of which
would cause the rain drops to roll off without wetting, at the same
time allowing gas-interchange to continue; or a coating of resin
would effectually prevent all danger of water absorption. All
of these devices are more simple than the elaborate system of bud-
scales found on many trees. On the other hand, the wool produced
on many buds would tend directly to retard the drying of the bud
surface.
The result of an experiment may here be given. During the
winter of 1902, about January 24, several buds of Acer platanoides,
deprived of their scales but still remaining on the tree, were each
inserted in a rubber pipette-bulb previously filled with water. The
neck of the bulb was then fastened firmly around the twig by means
of twine. The experiment was allowed to continue about one
ee ee Oe et ee ee se hee eae a Serge A)
1906] WIEGAND—BUDS AND TWIGS IN WINTER 397
week, during which time temperatures of —23.5° C. had alternated
with those of 4.3° C., so that the buds were alternately frozen and
thawed. After removal of the rubber, the tissue appeared as fresh
and sound as ever; the twigs were then cut and placed with their
ends in water in the greenhouse, where the treated bud remained
fresh as long as did others whose scales were freshly removed as
check experiments.'4
There exists, it seems to me, insufficient evidence to sustain the
theory that the exclusion of external moisture has played an impcr-
tant part in the evolution of scaly buds.
Heat conduction.
The popular belief is widespread that bud-scales serve to keep
out the cold, and indeed such an explanation appears in some of
our leading textbooks and in various other works. A moment’s
consideration will convince us that this cannot be true. No plant
tissue yet known is a perfect non-conductor of heat, or, indeed, less
than a fairly poor conductor, and scale tissue is no exception; while
the very thin nature of the scaly covering on some buds, as those of
Salix, would absolutely preclude their offering more than a moderate
amount of resistance to the escape of heat. To keep out the cold
during an entire cold spell in winter would require, even in much
thicker tissue, an almost absolute non-conductivity, and that is
possessed by few if any substances in nature, much less by the bud-
scales. This erroneous impression has arisen probably through
comparing the action of bud-scales with that of clothing upon the
human body, forgetting the fact that in the body there is a constant
source of heat without which clothing could not keep it warm for
more than a few minutes.'S
™ Kny found that with the bud-scales and cortex intact average twigs will
not take up as much water through these organs as they give out in dry air pie ae
a similar time. He neglected, however, to experiment with naked buds. Ue
die Aufnahme tropfbar-fliissigen Wassers durch winterlichentlaubte Zweige von
Holzgewachsen. Ber. Deutsch. Bot. Gesell. 13:361. 1895.
8 It may be suggested that such a constant source of heat does actually exist
in a tree, at least so far as the buds are concerned, and that this is provided by the
heat accompanying respiration. However, reference to any textbook in plant physi-
ology will show that the amount of heat evolved in this way is but slight in the very
best examples, which are all herbs, and is mainly evident during the period of most
398 BOTANICAL GAZETTE [JUNE
Such substances can only retard, not prevent, the escape of heat.
As a final argument we may return to the fact that observation
shows that buds are always filled with ice during cold periods, which
of course could not occur if they were kept warm.
It is a more difficult matter to demonstrate whether the non-
conductivity of the bud-scales is of importance to the bud in any
other way. Recently Griiss'® has quite firmly upheld the theory
that one of their chief functions is to modify the temperatures reach-
ing the interior of the bud. We can conceive of several ways in
which such protective service might occur. First, poor conduc-
tivity might prevent injury from too rapid thawing. Second, bud-
scales might prevent extreme fall of temperature by preventing
excessive radiation. Third, they might prevent too frequent rapid
thawing and freezing due to fluctuating sunlight, and thus prevent
excessive water evaporation.
Before answering any of these questions let us try to understand
a little more fully the actual relation of bud-scales to heat. This
problem resolves itself into two parts, namely, a consideration of
the conductivity simply, and a consideration of the relation to normal
atmospheric heat changes in the open.
On the question of conductivity the following experiments seem
to throw some light: Two thermometers, previously tested as to
their readings, were selected, and the bulb of one was covered with
the imbricated scales of a fresh horsechestnut bud, as in the previous
experiment to determine the effect of color, thus forming an arti-
ficial bud with the thermometer bulb in place of the young shoot.
The other bulb was left naked. The experiments were all conducted
within the building where the conditions were more constant and
presented fewer uncontrollable factors than outside. The room
rapid growth. During the dormant winter period it must be very slight in all trees.
An ordinary thermometer probably could not measure it. It may also be suggested
that since the large size and mass of the trunk would retard heat changes, being
warmer than the air meee the temperature is falling, and cooler when the latter is
this, by conduction along the gp hae modify the temperatures in the
shoots and buds. UIRES has shown (M Bot. Stud. 1:453) that the average
empertr in a box elder tree was in ace 1.3°C. higher than the air, in Fe
ry the same as the air, and in March 1° lower. The differences between internal
‘esd i deena temperatures during the day was in all cases only a few degrees. he
idea that the branches can conduct such slight modifications so long a distance
without loss is so evidently unreasonable as to require no more discussion here.
16 Griiss, J., Beitrage zur Biologie der Knospen. Jahrb. Wiss. Bot. 23:651. 1892.
AL see ee
1906] WIEGAND—BUDS AND TWIGS IN WINTER 399
selected had a temperature ranging from 3.7° C. to 4.3° C. during
the several days on which readings were taken. The two ther-
mometers were brought to the same reading in a warmer place,
either in another room or over a water bath, then quickly taken
out and the readings recorded for every few seconds until they again
registered at the same degree in the cold atmosphere of the room.
Two classes of readings were taken, one from a temperature only
a few degrees above that of the cold room and the other from one
far higher. The readings in each class, taken with the same
bud, corresponded remarkably. A specimen reading from each
set is here given.
TABLE II.
Horsechestnut bulb and naked bulb transferred abruptly from a temperature of
19.5° C. to one of 3.5° C. (See fig. 7.)
Naked bulb a ane | ° Difference | Time difference
67°F. 67°F | osec.| 0 F.(0°C.) © sec.
66 67 § + eg 20
65 67 10 o: (ae) 20
64 67 18 gts 37 ;
63 66 | 25 3 rs) 45 (3 min.)
62 65 | 32 cee ke, _
6x 65 | 38 4 ap a 72
60 65 45 5 23) 60
Weegee gees
5 63. . 3.0 :
§7 Ps : 70 3 ? ba) 100 (1% min.)
62 80 6 3.3) 105
55 62 go 7 i) 125
54 62 100 9 30) 145
52 61 110 9 Ke) a
9° 130 .O °
3 ine i .o) 200 (34 min.)
47 6 185 9 so.) 255
46 6 200 10 ee) 285
45 5 215 10 a na
4 2 10 a 7 E
pe : Be 9 .0 ) 420 (7 min.)
42 I 325 9 re) ed
41 4.4 r
40 8 pe Blea) 765 (12 min.)
49 47 442 z.: 43-0 4} 765
40 46 485 Go t3.4,) 765
3) 45 555 6 (3-3) 960
3) 44 620 5 oe a, 960 ;
38 43 705 et) 1235 (204 min.)
38 42 845 S48 3
38 4? T160 2 1.0 )
38 3? 1515 t (es)
38 33 1942 ° 0.0 )
400 BOTANICAL GAZETTE [JUNE
aN
\ sie oe
a Se
ean —
See eeeeetee tebe sess
horsechestnut bulb; ........... naked bulb. Abscissas represent
FIG. 7;
5° F.; ordinates, 100 seconds. See Table II.
The first column of figures represents the readings in degrees
from the thermometer with the naked bulb; the second column
the same from the bud-covered instrument; the third column shows
the time in seconds from the beginning of the experiment; the
fourth column the difference in degrees at each reading; and the
fifth column is the ‘“time-difference,” so-called, which represents
the number of seconds elapsing after a reading on the naked bulb
before the same temperature was reached on the horsechestnut
bulb, in other words, the number of seconds by which the bud-
scales retarded the fall of temperature in the enclosed bulb.
While not attempting to deduce the physical laws governing
the fall of temperature in each case, we may note from the tables
and curves several points which bear upon our problem. It will be
seen that theoretically the time required for the temperature to fall
in either case is infinitely long, the curve becoming nearly horizontal
towards the end of each experiment. But for all practical pur-
poses, and as closely as my instruments would measure, the fall
was completed in about thirty minutes in each case. The greater
part of it, in fact, was completed in ten minutes. As regards time,
in Table II the very much more rapid radiation of heat more than
balanced the effect of the greater quantity of heat to be radiated.
As we should expect, the retarding effect of the bud-scales in
degrees, shown in the fourth column, was much greater in case
of the greater extremes of temperature, and was greatest when the
papas enh yer pn m2)
1906] WIEGAND—BUDS AND TWIGS IN WINTER 4o1
/ TABLE III.
The same thermometers and bud preparation transferred abruptly from a temperature
f 51° C. (over a water bath) to one of 2.7° C. (See fig. 8
Naked bulb | Horsechestnut Time °Difference Time difference
124° F 124°F. o sec riot © sec.
sai 123 5 a ks 15
116 122 10 6 (4.2 *
hee 120 15 1o (5.5 27
106 118 20 re ne 28
100 116 25 16 (8.8 42
97 114 30 my 184 45 (2 min.)
94 113 35 19 6 (10.5 50
oI 122 40 eo” (15,3 Be
88 109 4s ar (11.6 60
84 106 48 22 12.2 97.7
82 105 sy ag (12.7 80
80 103 65 230 «(12.7) 80
T7 Iol 65 24 144 go (14 min.)
74 99 69 25 (13.9 III
73 97 75 25 (13-9 115
7° 96 80 20> (1320 120
94 85 25 (14.4 125
67 93 go 26 (13.9 135
65 gr 95 26 (14.4 145
63 88 105 25 330 155
j 61 87 115 26 14.4 160
' 60 86 120 26 14.4) 165
58 83 130 #e-\- 08.6 180
58 81 140 23. {a2.9) 180
54 76 160 22 12.2 200 (3} min.)
52 75 170 23 42.4) 230
50 74 180 24 13.3 250
49 72 Igo 23 12.7) 265
47 69 210 22 12.2) 280
46 68 220 22 12.2) 330
45 65 240 20 ae A 335
44 63 260 19 10.5 240 .
42 61 275 19 (10.5) 435 (74 min.)
41 59 305 18 10.0) 465
41 57 320 16 8.8) 465
= 56 335 15 8.3). 465
ae 55 355 15 (8.3 545 (9 min.)
4o 80 13 7.2 545
39 aa 12 (6.6 580
39 49 455 10 5-5 580 :
38 47 4 g (5.0 ggo (164 min.)
38 46 55° 8 (4.4) 5
37 44 600 7 O20) 1300 (214 min.)
37 43 O..4s3
37 42 710 ee 9
37 41 770 aes fae |
37 40 Mee ee 6.
37 39 I Pais Ge A
37 38 1480 Beg ae
37 37 1900 yore toe ©,
Ra al a AO eh ore Rr ane
80
BOTANICAL GAZEFTE [JUNE
fall was most rapid. Of much more importance to our
problem is the retarding effect in point of time, shown
in the fifth column. This increased very rapidly towards
the close of the readings, but was for our purpose
practically the same in both cases. It was greater in
proportion to the slowness of heat penctration, and was
also somewhat greater at first in Table I than in Table
II. The greatest retardation capable of measurement
with the thermometers used was about twenty minutes,
while for most of the experiment it was only from one
to nine minutes. It was found that decreasing the
thickness of the scaly covering decreased this time
difference very markedly; while the presence of air
Oe i
between the scales tended to make it greater.
The mass of the thermometer bulb, or of a
shoot in a normal bud, and the extent of the
ai radiating surface, are important factors in deter-
a mining the length of time required for such a
structure to cool. While the mass of the mercury
N in this case is much greater than that of
ae
\ a _-
NJ
a2
c. 3
S688 © Sa 2s 8 2 SS s ce =
horsechestnut bulb; _........... naked bulb. Abscissas represent
Fic. 8.
5° F.; ordinates, 100 seconds. See Table III.
the shoot, its specific heat being only one-thirtieth that of water
would render the two not very dissimilar, so far as the present
problem is concerned. In apparent volume they do not differ
greatly, so that the radiation surfaces of the two would be nearly
the same.
I believe we are justified in saying that a normal horsechestnut
bud would not behave in any essential way differently from the
artificial one here used; and that the time for it to cool off would
—
1906] WIEGAND—BUDS AND TWIGS IN WINTER fe)
403
be for all practical purposes not over about thirty minutes, no mat-
ter whether it was cooled very much or only a little, providing it
was plunged directly into the cooler temperature.
We may also say, I belicve, that smaller buds with thinner scales
and smaller shoots will show a time period correspondingly less
than thirty minutes, and a time difference which will approach
more nearly zero. In the case of the willow buds with only one
thin bud-scale, the time period and time difference must be very
small indeed.
A number of readings were taken in which the thermometers
were warmed up instead of cooled, and it was found, as expected,
that the above generalizations applied in this case also. Providing
that atmospheric changes out of doors are abrupt, I fail to see how
the temperature at the center of a bud of medium size can be retarded
more than five or ten minutes over practically all of the range of
fall. A small bud would probably be retarded only about one to
five minutes. Of course the retarding would be greater than this
through the last degree and fraction of a degree, but this slight
change, it seems to me, would be of little moment to the present
question.
Buds in nature, however, are under slightly different conditions.
Instead of being transported from one temperature to another,
the temperature itself changes. We should therefore conduct
some experiments in which the air itself is varied. This change
is either very gradual, as when a thaw approaches, or more abrupt,
as when the sun shines from behind a cloud upon the bulb, which
is the only way in which abrupt changes are produced in nature.
In either case they are much less violent than were our laboratory
experiments. During warming by the sun, radiation from surrounding
objects may play an important part and introduce still another
factor. We should thcrefore conduct some experiments in which
the air itself is warmed. The experiments with the horsechestnut
bud already described in the discussion of the function of color
are to the point here. They show in a surprising way that instead
of retarding the rise in temperature within the bud, under these
very natural conditions the bud-scales actually scem to hasten it.
These experiments were with direct sunlight. It seemed possible
404 BOTANICAL GAZETTE [JUNE
that the readings might be different if radiated instead of direct
heat was employed, especially since there is a considerable difference
in the nature of such heat, as shown by the well-known fact that
direct heat from the sun passes easily through glass into the green-
house, but when radiated passes out with much greater difficulty,
thereby warming the house.
A number of readings taken with naked and_ horsechestnut
bulbs transferred from the shade to the surface of a black book in
direct sunlight with the bulb raised 3-4™™, or with the bulbs pro-
jecting several inches over the edge of the book which itself was
raised several feet from the ground, or with the bulbs raised 7.5—10°™
above the surface of the book, showed no appreciable difference
that could be referred to a difference in kind of radiated heat. There
was some difference in the readings, of course, but this could ke
traced directly to the fact that there was more intense heat where
the heat of radiation was also present. In case of more intense
heat the extra absorbing power of the bud-scales was at first more
obscured by the slightly greater retardation of heat-penetration due
to the greater difference in outside and inside temperatures, as we
should expect from the deductions from Tables III and IV. This
was partially shown by the difference in locality of the crossing
of the two curves plotted from each reading.
Looking at the matter from still another standpoint, we may
consider how much time is required for a bud to thaw. As shown
by the cover glass measurer, the wrinkled willow twigs thawed and
became perfectly normal in thirty minutes at the temperature of
the laboratory. Undoubtedly the ice had disappeared in about
half the time. Large buds of horsechestnut will lose all their ice
in about twenty-five minutes under similar conditions, and buds
of Negundo in about fifteen minutes. The small buds of the black
cherry require only about ten minutes for thawing. The time
required in the laboratory for the various buds, therefore, is ten
to thirty minutes. The question is whether when the temperature
changes are slow the buds thaw proportionately more slowly. The
answer must be that they will, slightly, just as a cake of ice will
thaw more slowly when the temperature rises gradually than when
the rise is abrupt. This difference is proportional to the size of the
1906] WIEGAND—BUDS AND TWIGS IN WINTER 405
ice cake, since it depends largely upon the non-conductivity of the
ice and the greater quantity of heat required to convert ice into
water. This heat is more slowly available when the change is
gradual. Although no experiments were made under these condi-
tions, it is to be expected, I think, that with long slow rise in atmos-
pheric temperature, the retarding effect would almost if not quite
disappear. Frozen peach buds, placed in the air at—5.5° C., which
gradually rose in 2 to 2.5 hours to a temperature of —1.0° C., were
completely thawed, apparently as soon at the temperature reached
about —2.3° C., thus following the general rule for frozen tissue.
We are now in position to consider the questions outlined on a
previous page regarding the various ways in which the bud-scales
May be supposed to act beneficially by modifying the temperature.
It was first suggested that they might retard the thawing out
and thereby be of benefit to the bud. From the tables already
given and the observations regarding them, it becomes at once
apparent that the temperature modification which scales are capable
of producing are, in the cases of moderate sized buds, of very little
Moment—not more than two or three minutes during most of the
time, and then only when the change from one temperature to
another is abrupt. When the transition is gradual, the retarding
effect will be very slight indeed, and is frequently wholly offset by
the absorbing power of the darker color. I cannot see how under
any atmospheric condition the modifications can become great
enough to be noticeable unless careful measurements are taken.
The idea that a slow thawing is beneficial to plants has come about
from analogy with the frosting of human tissue and from the con-
sideration of the treatment which gardeners successfully give frosted
plants. But the gardener’s treatment consists in keeping the
plant cool and dark for hours or even days after the freezing; while
recent investigators have shown that slow or rapid thawing (7. e.
conversion of the ice into water) in themselves bear no relation
Whatever to the extent of the injury. The gardener’s treatment
is essentially an after-treatment—while the plants are recovering
from the shock. I have already cited the fact that buds of many
trees, at least, may be thawed in an oven and then frozen alter-
nately many times and still come out in the greenhouse apparently
406 BOTANICAL GAZETTE [JUNE
as fresh as others not so treated.'7. The answer to this first question
then, is, that bud-scales do not function by preventing rapid thawing
of winter buds; neither does bark so function towards the twigs.
It has been suggested that bud-scales protect the bud by pre-
venting rapid radiation from the delicate tissue during the cold
nights, and thereby preventing a harmfully low fall of temperature.
MU ier-TuHurGAu,'® by placing one thermometer on some cotton
under a o.5°!" cloth screen fastened 1°" above the ground, and
another thermometer outside, was able on a clear night to get 4° C.
difference due to radiation. Griiss'® states that differences in
temperature due to radiation may be one or two degrees on cool
nights just before sunrise, and as great a difference as 6° C. has been
observed by other investigators. A difference of 4-6° C. would
frequently be of importance to tender exotic buds in winter, but
it is scarcely to be supposed that so slight a difference would be
of much moment to the great majority of perfectly hardy species
which withstand all of the fluctuations of our vigorous American
climate without injury. Indeed these species seem capable of
existing below any atmospheric temperature that has yet occurred
in this country, as freezing mixture experiments have shown. Besides,
the structure of buds does not lead one to expect a radiation screen
as efficient as those specially constructed. Strictly speaking, the ques-
tion here is not one of radiation of heat, since the scales are all more
or less in contact, but of conduction, and as such has already been
treated.
HENsLow’? has shown that it scems desirable for plants in tem-
perate regions to protect their delicate bud-structures from loss
of water when the bud is opening. Such loss he says is favored by
radiation and heat absorption. The above objection will apply here
also for the first part of this last statement, and the latter part is
treated elsewhere in this paper.
17 Motiscu, H., Untersuchungen iiber das Erfrieren der Pflanzen. Jena. 1897
18 MULLER-THuRGAU, H., Ueber das Gefrieren und Erfrieren der Pflanzen-
Landw. Jahrb. 15:563. 1886.
9 Gruss, Beitrige zur Biologie der Knospe. Pringsh. Jahrb. 23:651. 1891-92.
20 HENSLOW, G., On vernation and the method of meee of foliage as
protective against radiation. Jour. Linn. Soc. Bot. 21:624. 1886.
lt
1906] WIEGAND—BUDS AND TWIGS IN WINTER 407
In December 1901 some experiments were conducted to show
whether twigs and buds while continuing frozen lost as much water
by evaporation as when alternately thawed and frozen several times
during the same period. It was found that they did not quite,
and hence the question whether bud-scales may function by pre-
venting too frequent thawing and freezing. Several buds of Pinus
Laricio and horsechestnut, also several twigs 15°™ long of Syringa
vulgaris and apple, were sealed at the cut end with Venice turpen-
tine, weighed, and quickly placed on a tray in the open air. They
were divided into two equal lots, and one of these was brought
into the warmer laboratory for a few moments ten times, thus insur-
ing ten alternate thawings and freezings. During the experiment,
which lasted three days, the temperature ranged from —18° C. to
—7° C. in the open. The results were as follows:
Horsechestnut buds continued frozen lost 0.4% of their water.
«alternately thawed and frozen lost 0.6% of their water.
i he oe — continued frozen lost 3.4% of their water.
alternately thawed and frozen lost 5.0% of their water.
ea twigs continued frozen lost 1.3% of their water.
*¢ alternately thawed and frozen lost 2.4% of their water.
i a — continued frozen lost 1.6% of their water.
alternately thawed and frozen lost 2.4% of their water.
In every case there was a greater loss of water from the buds
which were alternately thawed and frozen. The difference was
very marked, and in each case amounted to about one-third of the
total loss. Considering the total quantity of water present, this
was really a very slight increase in loss, however, being 0.25% in
horsechestnut, 1.1% in lilac twigs, 0.8% in apple twigs, and 1.2%
in pine buds; and with me it is a serious question whether, in all
of these cases, so slight a difference would not be quickly equal-
ized during spells of thawing by conduction from the older wood,
if the twigs and buds were connected with the trunk in the normal
manner. Again, the thawings in nature would probably be fewer,
and it has not been shown that bud-scales prevent such thawings.
It seems to me that here again a beneficial functioning of the bud-
scales is very doubtful.
But the most vital argument against all these cases lies in the
fact that experiments have shown that dark buds tend actually to
408 BOTANICAL GAZETTE [JUNE
increase the heat absorption. Therefore, these considerations could
scarcely have been instrumental in bringing about the existence of
such structures.
The idea that bud-scales may protect the bud by warding off
the hot rays of the sun applies mainly to the tropics. It seems
to have been first advanced by TrEvuB,?" who cites several cases,
where in plants exposed to the hot tropical sun delicate young
tissues were inclosed in enlarged stipular organs or else well-shaded
by overlapping leaves or by other special structural provisions.
On the same subject, in 1891 another paper was published by
Porrer.?? According to this investigator many trees in the tropics
protect their young leaves and shoots from direct sunlight by means
of stipules. These organs were removed from a number of buds
and in every case the leaves from these when mature were deformed
and abnormal. The sunlight seemed to produce injury by causing
more water to be evaporated than could be replaced. For this
reason Artocarpus, the most pronounced type of this class, unlike
most trees, produced leaves throughout the dry season, probably
because of the stipular protection. Instead of by stipules some
tropical plants obtain similar protection by various methods of
leaf-folding, shading by older leaves, and coating with gum. Is
there not inaccuracy here in his interpretation? Rather than by
actually preventing the entrance of heat from the sun, which it seems
such structures could do only to a slight extent, is it not more proba-
ble that they function simply by preventing the escape of extra
moisture vaporized by the intense heat ?
The relation of bud-scales to the young shoot when the bud is
opening is discussed under internal moisture relations. Suffice it to
say that the results reached seem to indicate that even in this case
the scales do not function beneficially by modifying the heat.
It has sometimes been thought that the layers of hair and wool
found in many buds, as for example in the horsechestnut, are for
the purpose of modifying the heat conditions inside. To obtain
21 TrEuB, M., Iets over knopbedekking in de tropen. Hand. van. het eerste
Nederlandsch Natuur- en Geneeskundig Congres. Amsterdam. 1887, p. 139- Ref.
Bot. Centralb. 35:328. 1888.
22 PoTTER, M. C., Observations on the protection of buds in the tropics. Jour.
Linn. Soc. Bot. 28: 343. 1891.
Oa eal
1906] WIEGAND—BUDS ‘AND TWIGS IN WINTER 429
evidence upon this point I performed the following experiment.
The two thermometers used in the previous experiments were
selected, and the bulb of one was coated with black cloth; that of
the other was wrapped in a layer of cotton about twice the thickness
of the wool in the horsechestnut bud, and was then coated with
black cloth. The surface of both bulbs was therefore black.
TABLE IV.
Bulb covered with black cloth, and bulb covered with cotton and black cloth; trans-
erred from a temperature of 56° C. to a room of 9° C.
Cloth bulb Cotton bulb Time ° Difference Time difference
134-8. 134° F. o sec. o° F. (0° C.) © sec.
130 131 5 I (0.5) 7
128 131 to r Seda SS 10
125 128 15 3 (1.: 15
123 128 20 5 (2:7) 15
222 127 25 5 (2: 15
119 125 30 6 3.3) 20
i17 123 35 6 2-3) 20
114 T21 45 7 3-9) 20
ach a T19 50 8 4-4 25
109 117 55 8 4-4 26
106 Hie 9 (5.0) 30
104 112 70 8 4.4 30
100 110 80 Io oy) 37
98 107 85 9 -0 40
96 105 9 9 iO 40
93 103 105 Io 5 40
QI IOI 113 Io us 42
99 120 to fe 45
87 97 130 10 5 45
86 96 135 Io as 43
oy 93 145 9 (5-0) 59
82 ot 155 9 (5.0 55
80 89 165 9 (5.0) 55
78 88 173 10 8 _ 62
77 i 86 178 9 0 7?
76 85 185 9 -O 75
74 84 195 be) “5 75
73 83 205 Io 5 80
72 82 210 10 “5 85
71 81 215 10 5 95
7° 79 225 9 te) 95
69 78 235 9 .O 100
68 77 250 9 re) 105
67 76 60 9 .0 110
66 74 270 ee) 110
65 73 285 8 (4.4) 120
64 72 295 8 4-4 130
63 71 310 8 4-4 135
63 70 320 7 (3-9 135
62 69 335 7. (9 140
410 BOTANICAL GAZETTE [JUNE
TABLE IV.—Continued.
Cloth bulb Cotton bulb Time ° Difference Time difference
61°F. 68°F. 355 sec. or (3,90) 155 Bec.
61 68 370 6 aS 155
61 66 380 5 Ce 155
60 65 405 , aia 5 145
9 64 425 5 2a7 160
58 63 445 5 2.7 185
7 65.5 475 5-5 (2.7 210
56 61 510 5 2.7) 270
6 60 550 4 2.1 310
5 59 585 4 2.1) 545
4 8 630 4 2.1) 545
4 57 685 3 1.5) 545
3 7 745 4 2.1) 1083
3 6 780 a £25 1083
3 5 895 2 eae. 1083
2 55 IOI5 a (i.5 1300
2 54 1175 2 (1.0 1300
I aie 2155 a a | 480
49 -! 2635 2 (126 1440
49 50 3355 EG s§
48 49 40°75 Se Ne
As expected, the retarding effect was apparent in this rather
violent experiment, but it was not great. The maximum degree
difference of 10° was less than one-half that produced by the bud-
scales in Table III, while the time difference through the greater
part of the experiment ranged from o to 4 minutes. I think it may
be inferred that the wool in the horsechestnut bud retards the pene-
tration of heat, when the changes are at all great, by 0.5—3 minutes.
At any rate it seems evident to me that the retarding power of the
wool in such buds as horsechestnut is insufficient to explain the
presence of such a structure. This appears not only from experi-
ment but from a general consideration of the thinness of such struc-
tures compared with the relatively great temperature differences
which they are supposed to offer protection against, and must in
order to be effective. Their true function, it seems to me, lies in
an entirely different direction, as we shall see somewhat later.
In concluding this study of the relation of bud-scales to temper-
ature the following summary may be made: :
Bud-scales or bark cannot “keep out cold” during the cold
spells of winter. ;
They seem not to modify the temperature sufficiently to be of
1 hive apy age
1906] WIEGAND—BUDS AND TWIGS IN WINTER 4II
beneficial importance in preventing rapid changes, even if such
changes are detrimental.
Rapid thawing in itself is probably not detrimental to buds and
bark.
Bud-scales seem of no benefit in keeping out the heat from sudden
bursts of sunshine. They do not appreciably prevent the loss of
water by preventing alternate thawing and freezing. They do not
retard radiation to any important degree.
Dark-colored bud-scales indeed, instead of preventing tempera-
ture changes actually seem to absorb more heat than if they were
lighter colored.
“Wool” in buds does not function by modifying temperature
changes.
Bud-scales do not seem to function in modifying temperature
changes when the bud is opening.
Bud-scales may protect the delicate tissues in the tropics from
heat, but it would seem rather from excessive transpiration due
to great heat than from the heat itself.
Finally, we may conclude that as a factor in the evolution of
buds and bark in cold climates temperature considerations have
probably played a very minor part.
Internal moisture.
Of all the more important factors concerning the function of
bud-scales, perhaps that relating to their inhibiting effect upon the
loss of internal moisture is the least recognized by people in general.
In scientific literature, however, it has received considerable atten-
tion. Most authors now consider this one of the principal functions.
of the bud-scales and also of the bark. The subject has been dis-
cussed briefly by CapuRrA*3 and Groom,’* but also more fully by
Gritiss,?5 who performed a number of experiments to demonstrate
the point. His results may be summarized as follows. The first
function of the scales consists in protecting the inner meristematic
23 CaDURA, R., Physiologische Anatomie der Knospendecken dicotyler Laub-
baume. Breslau. pp. 42. 1887.
Pe Groom, P., Bud protection in dicotyledons. Trans. Linn. Soc. II. 3:255.
25 Grtss, J., Beitrige zur Biologie der Knospen. Jahrb. Wiss. Bot. 23: 649. 1892.
412 BOTANICAL GAZETTE [JUNE
tissue from loss of water. Even in summer and especially in fall,
when the sap flow decreases, the tender embryonic interior of the
bud must be protected from too great transpiration. Also in winter
this function is not interrupted, for then the cold wind can bring
into play its desiccating action. To prevent loss of water, cork
layers are formed, or in place of these felty hairs may be produced.
A third method. consists in the excretion of resin. If, under con-
stant temperature, the scales were removed from an oak bud, it
soon died, even though there was a moderate amount of moisture
present. The inner bud-scales dried out and perished, as well
as the embryonic tissues. The young leaves of a beech bud so
deprived of scales persisted much longer than did those of the
European oak; which he thinks was because the former were hairy
while the latter were not. Buds of horsechestnut proceeded to
develop in spite of the removal of the scales, probably, he thinks,
because of the thick wool among the young parts. Buds of Abies
pinsapo, whose pitch had been removed by carbon bisulfid, dried
out in a very short time. These experiments were all performed
on twigs cut from the trees and placed in water.
In 1895 Kny?° published a paper dealing with the transpira-
tion and absorption of water by buds and twigs in winter. He
cites WIESNER and PACHER as having shown that horsechestnut
loses water from twigs in winter, and also Hartic as having shown
that many trees do the same. Experiments are given to show that
in general not so much water is absorbed by these parts in saturated
atmosphere as may be given off at an ordinary degree of saturation.
In 1895 some determinations of the amount of water lost by
twigs with buds attached were made by the Cornell Experiment
Station.?”7_ The experiments lasted three days, beginning April 7.
The twigs were sealed at the cut end and kept in an open shed.
The percentage of loss ranged from 2 to 10%, with an average of
5-4%-
In 1875 WIESNER and PacHER?® found that twigs of horsechest-
26 Kny, L., Ueber die Aufnahme tropfbarfliissigen Wassers durch winterlich-
entlaubte Cutis von Holzgewachsen. Ber. Deutsch. Bot. Gesell. 13:361- 1895-
27 BaILEy, L. H., Cornell University Experiment Station Rep. 1896: 4.
28 WIESNER u. PacHER, Ueber die Transpiration entlaubter Zweige und des
Stammes der ete ge Bot. Zeitschr. No. 5. p. 9. 1875.
RE i ee Se ee are ot ae al ye
1906] WIEGAND—BUDS AND TWIGS IN WINTER 413
nut transpired an appreciable amount in winter at a temperature
of 13-17° C. and a slight amount also at —10° C. This was true
in older twigs. The leaf scars transpired more than the periderm.
The winter buds also lost some water.
That there is actual loss of water in winter probably every one
knows. My experiments given below show this very definitely,
but perhaps few understand that there can be a loss when the tissue
is frozen as well as when thawed, though less in extent. Water
may evaporate to a large extent from ice crystals themselves, as
is shown by the drying of frozen soil, damp clothing, and the frequent
disappearance of small quantities of snow at temperature below
the freezing point. In buds not all of the water becomes ice, and
the remainder is free to evaporate as at a higher temperature.
The fall of temperature on the approach of winter is always
accompanied by a decrease in the power of root absorption, and
it has been shown that, to a certain extent absorption is propor-
tional to the temperature. In the case of our native plants, the
decrease must be very considerable when the zero air temperatures
have chilled the soil to a depth of many feet. A compensating
decrease in transpiration must occur or otherwise the cells will
suffer from too small water content. This is mainly accomplished
by the fall of the leaves, but is greatly aided also by the coverings
of the bud and the waterproof bark. But so far bud-scales would
not be a necessity, because very little root absorption would probably
be sufficient to supply the slight amount of water that could evapo-
rate from unprotected buds, compared with that necessary to supply
the leaves. Besides, it has been found that considerable water is
present in the wood at all times, and in some trees even a larger
amount than in the summer. The necessary factor, I suspect,
lies in the decreased osmotic activity and vigor of the young tissue
itself, During the summer the tendency to transpire is probably,
a large share of the time, not so great as in the winter and spring
because of the greater saturation of the air; but there are times
during the summer when the transpiration is very great indeed.
The young tissues do not then dry up very readily, so that little
harm usually results. At this time I imagine the growing cells
are osmotically very active and more easily draw to themselves
414 BOTANICAL GAZETTE [JUNE
a supply of water sufficient to offset that lost in transpiration. In
winter, however, the cells are inactive, and on account of the cold
the osmotic force is much decreased, so that the cells find it impos-
sible to resupply quickly the transpiration water when this function
is very great.
That loss of water beyond a certain point is detrimental to the
cell needs no further demonstration. It has been shown that each
cell demands a certain percentage of water, depending mainly upon
its activity and water content, in order to maintain its life-prop-
erties. If transpiration even for a short time reduces the water
in the cells of the bud below the critical percentage, the cells will
cease to remain alive.
During January 1gor, I cut some twigs of horsechestnut, stripped
off the bud-scales from some of the buds, and exposed the whole
to an outside temperature of —18° C. to —12.3° C. for 24 hours,
after which the twigs were placed with their cut ends in water in
the greenhouse for further development. The buds all lived, although
those without bud-scales were the first to commence growth. Sub-
sequent experiments show that the reason why none died was because
the exposure to the dry air was not long enough. On March 1
of the same year, buds of the black cherry, Crataegus punctata,
horsechestnut, lilac, apple, and Pinus Laricio while still on the tree
were deprived of their scales and each divided into two lots. One
lot was left naked, the other was varnished completely with Venice
turpentine to prevent loss of water. When the normal buds were
opening May 8-10 it was found that both varnished and naked
buds were all dead except on Pinus Laricio. On this plant the
naked buds were all dead, but the varnished ones were alive, and
later all developed into normal shoots.
The varnished buds in all cases seemed to be all sound and turgid
until warm weather and time for swelling came, when they seemed
to decay rapidly, and in no case except the pine did any swelling
occur. I suspect that death here was due to the retardation of res-
piration owing to the lack of oxygen. The pine is normally closely
surrounded by resin without a space inside as in horsechestnut,
and possibly some other way is here provided for obtaining Oxy-
gen. The pine, therefore, is the only one of the series in whic
1906] WIEGAND—BUDS AND TWIGS IN WINTER 415
the results of varnishing are important to us here. The naked
buds of pine in every case began to dry and shrivel up after only a
few days’ exposure, and were quite dead long before the time for them
toopen. There seems no doubt whatever that the varnish preserved
the pine buds by preventing loss of water. Without the varnish
the pine was one of the first to succumb. That this thin layer of
varnish replaced effectively the thick layer of scales is also good evi-
dence toward the idea that the scales do not function by causing
temperature modifications.
To determine just how much more water is lost from buds without
scales the following experiment was performed. Several buds
of Pinus Laricio and horsechestnut were separated from the trees
by an incision at the base of the bud and the scales were removed
from all. One-half were quickly varnished, weighed, and placed
in the open air at —18° C., while the other half without varnish were
weighed and exposed at once. Care was taken to seal up the cut end
in both sets so that no water could escape that would not if the
buds had remained on the trees. After three days at a temper-
ature of —18° C. to —7° C. the results were as follows:
Orig. aa Final weight| Dry weight | Per cent. HO
Pinus Laricio—continued frozen:
Wath Trid-ecales. oo... caus eck 2.658m | 2.6158™)| 1.358™ 2.7
Without bud-scales.....:......... 1.28 0.96 0.58 45-0
Aesculus hippocastanum—cont. frozen:
Ser Btieh ain bet Go oe es Fk 4.03 4.02 1.80 pia
Peel aap EAicice alias 18 ic oso oases 2.21 1.94 1.04 33.0
uring experiment by bring-
ing into the laboratory:
With tead-deales. 35 c5 sis val es 5.30 5.28 2.26 0.65
_ Without bud-scales..............- 2.01 1.66 0.93 32.0
Syringa vulgaris—continued frozen
With Dich -atated 5 i405 awe cas i a 1.133 0.52 2.8
Without Sekxeabes Dell Sew ike elars ©.41 0.32 0.18 39.0
I think that nothing could show better than these experiments
the very great difference in amount of water transpired by buds
protected by scales and those having none. No wonder that the
loss of water oversteps the critical point and causes the destruc-
tion of the tissues.
416 BOTANICAL GAZETTE (JUNE
The buds in the above experiment were separated from the
tree upon which they grew, and therefore could not receive water
from it to replace that transpired. It would be interesting to know
how much water moves into the bud to replace the quantity lost,
thus giving a better idea of the actual decrease in percentage within
the cells. This has not been done for twigs at temperatures above
freezing, but the following figures are available for the frozen buds
of pine. I selected six vigorous buds of Pinus Laricio, all on the
west side of the tree, deprived them of their bud-scales and allowed
them to remain exposed three days. The temperature during
this time ranged from —18° C. to —6.7°C., so that the twigs as well
as the buds were constantly frozen. Three of the buds were cut
off, the cut surface sealed, and placed in a tray at the base of the
tree, while the other three remained attached. The results were
as follows:
baie’ pala Dry weight | Difference | Per cent. ™ wet at end
Cut buds—
i ©.1058™| 0.0618™| 0.044 41.9 |
" 2c Sk aati eg aR PE Tae PP 0.150 0.090 0.060 pies Maas average
Ree eed ree os 0.230 0.134 0.096 41.8
Sra Bas coe
No Ae Sy eo 255 65139 0.116 45-5
an 0.240 0.121 0.119 49-5 747-5 average
Des aie cP ees a es 0.210 0.110 0.100 47.6
It seems, therefore, that there was a rise of about 5% of water
into the bud while the tissues were frozen. This is quite possible,
since only a portion of the water was converted into ice, the remainder
remaining fluid in the walls and protoplasm and still capable of
movement. The figures given above for the loss of water from
desquamated buds are therefore slightly too large in every case.
It may be noticed by computing corresponding figures that the loss
of water during this last experiment is slightly less than in the two
previous experiments in which desquamated buds of Pinus Laricio
were used. This was due to the fact that the last experiment was
conducted in a different place, on the other side of the building.
‘I have found that exposure makes a very considerable difference
in the loss of water, and readings which are to be compared must
ve SE <a
1906] WIEGAND—BUDS AND TWIGS IN WINTER 417
be taken in the same place under the same weather conditions.
The rise of 5 per cent. is only a small part of the whole water lost.
How much will rise into the bud when the tissues remain thawed
was not determined.
The question arises whether the damage to the bud is done
while the latter is frozen or thawed. I believe that injury is done
both while thawed and while frozen, for the reason that the loss
of water from unprotected buds is sufficiently great to cause death
at either time. However, probably more damage is done above
freezing point, because here the loss must increase with the tem-
perature much faster than does the conduction of water in the bud.
The evidence therefore seems to be sufficient to warrant the
conclusion that the loss of water during the winter is a danger against
which the bud-scales and bark serve as protective organs. Next
to the warding off of mechanical injury this is probably their most
important function.
Protection oj the young shoot.
In many trees, notably in maples, horsechestnuts, oaks, etc.,
the growth of the young shoot is accompanied by a growth in length
of the bud-scales, especially of the inner ones, so that a sort of
telescopic tube is formed in which the young shoot remains concealed,
frequently until a growth of 2—8°™ has taken place. The function
performed by the bud-scales at this time has long been a subject of
interest to investigators, among whom Gruss’? has given us the most
comprehensive account.
According to Griiss, the protection lies in the ability of the
Scales to prevent the penetration of extreme cold on freezing, or
great heat when thawing out. The greatest danger to buds from
cold, he says, is in the spring after activity has commenced, when
a few degrees of frost will often kill the tissue. It is on just such
occasions that the frost is likely to be of short duration, and to occur
for a few hours only, just before sunrise. It is quite conceivable
that the non-conductivity of the scales would be sufficiently great
to prevent an extreme fall of temperature within the bud during that
short time. Again, if the cold was severe enough actually to freeze
0 Gruss, J., Beitrage zur Biologie der Knospen. Jahrb. Wiss. Bot. 23: 649.
1892. See also HENSLOW, I. c.
418 BOTANICAL GAZETTE [JUNE
the tissues, then when the sunlight fell upon the buds in the morn-
ing the scales would prevent injury from too rapid thawing. To
support this view he found that shoots of Picea Engelmanni deprived
of scale-caps perished, while normal ones did not. Shoots of Betula
alba projecting slightly beyond the sheath were overtaken by a
slight frost. The portion beyond the sheath was killed while the
protected portion was uninjured. At a temperature of —3° R. the
portion of the shoot outside of the sheath in Larix and Pinus Cembra
was completely killed. A horsechestnut tree at —5° R. had the
portion of shoot projecting beyond the scales killed, the rest was
uninjured. At —5° R. shoots of this species were killed, while at
—4° R. they were all right. Buds of Acer platanoides still in the
scales were killed at —4° R. Populus cannot stand a cold of —5° R.
after breaking out of the buds. Shoots of birch not inclosed were
killed at —5° R., but not at —4° R; those still in the bud-scales
were uninjured at —5° R.
I am unable to agree with Griiss that these cases of protection
are due mainly to the modification of the temperature. During
the spring of 1902 I removed the scales from a large number of oak
buds, also from birch and from maple. This was done about the
time that the buds were swelling rapidly; but no frosts occurred
afterward until the leaves were quite far developed. Many of
the oak buds so treated died, and the rest were retarded or deformed
in various degrees. The appearance in all cases was that of drying
out—as though the tissue had simply shriveled up from lack of water.
The birch and maple showed the same effect though in a less degree.
Any one seeing these experiments could scarcely believe otherwise
than that the loss of water caused the injury. When the shoot is
young the epidermal cells are but slightly cutinized, and are there-
fore much more pervious to water vapor than after having become
more mature. At this period also the cells have probably not yet
reached their full osmotic activity, and are still unable to replace
rapidly the lost water. It is not surprising, therefore, that shoots
should be injured at this period. Even in tropical regions, young
tissue is protected against transpiration by being folded within the
leaves, or in other ways. Groom’? has brought this point out nicely
3° Groom, P., Bud protection in dicotyledons. Trans. Linn. Soc. II. 3:255- 1893-
(RR eee a
1906] WIEGAND—BUDS AND TWIGS IN WINTER 419
for temperate regions. He points out that old parts often cover the
new. The blade of the young leaf is often covered by stipules. The
most critical time is when first exposed, the walls then being thin
and feebly cutinized, the chlorophyll dilute and easily decomposed.
The blades after coming out are folded and covered with hair in such
a way as to diminish transpiration and radiation as well as to reflect
light. According to Groom the function of mucilage and tannin in
buds is to help hold the water in the young shoots.
In this light I think we can see the function of the air and wool
which gave the negative results in the temperature experiments.
Water vapor diffuses through air quite slowly unless the air itself
is in motion. If a layer of substance containing air passages such
as wool, through which there is almost no circulation, is placed
around a damp object, the evaporation from the object is very much
retarded because the air in contact with the water is almost satu-
rated under certain conditions.
The horsechestnut wool, therefore, although not functioning
in the bud would become a most efficient protection against loss
of water from young shoots after leaving the bud-scales. I think
this is the purpose of nearly all hairy coverings of young flowers
and branches, which view is strengthened by the fact that in most
cases the hair disappears before maturity. The putting up of the
hair already in the bud insures its presence at the very earliest
moment when it shall be required. These and other similar obser-
vations have quite firmly convinced me that the growing out of
bud-scales and the presence of hair on the young organs is mainly
for the purpose of retarding transpiration. In some cases they may
be important also for mechanical support.
The observations of Griiss in regard to death at temperatures
slightly below the freezing point I think can be explained in this
way. When tissues freeze the water enters the intercellular spaces
and can from there escape more easily to the outer air. If, however,
hair or scales were present, not so much water would escape while
the tissue was frozen, and a lower temperature might be necessary
to cause fatal loss of water. In the case where —5° R. caused death
While —4° R. did not, it is possible that freezing just began at that
point which is near the over-cooling point of such tissue. It may
420 BOTANICAL GAZETTE [JUNE
be, however, that there are times when the temperature barely
falls to the death point, and then only for a very short time just
before sunrise, when the bud-scales may save the life of the bud
by preventing a temperature fall of from 0.5° to 1° for a very short
time, and this little we must admit they are capable of doing. But
we must consider that the period when the young shoot is covered
by the extending scales, in America at least, lasts only about three
or four days at the most, while the probability of a fall just to the
critical temperature during this short period is indeed very slight.
There may be no frost at all or there may be a very severe one.
Only in the case of such a slight frost just reaching the critical tem-
perature could the scales be of benefit, and even this injury would
not be fatal to the tree, since another crop of accessory buds can
grow out in a short time. The chance to function is therefore very
slight, and the effect could not compare in importance to the plant
with the benefits obtained according to the above theory, because
in the absence of scales nearly all buds of whatever crop would
run great danger of being killed by loss of water. It is not reason-
able to suppose, therefore, that the benefit derived by modifying
temperature is sufficient to have played any great part in caus-
ing natural selection to evolve such an elaborate structure for this
purpose.
Relation oj bud-scales to mechanical injury of the bud.
The idea that the bud-scales serve to protect the delicate young
tissues within from mechanical injury is of course not new.*' In
fact, probably everyone feels that this must be, if not the most impor-
tant function, at least a prominent factor in the work of protection.
Nevertheless the subject seems to have received little attention in
physiological works.
The young shoots of our definite-growing trees as they exist
through the winter are very tender organs, composed mainly of
thin-walled parenchyma. In most cases the epidermis is still thin,
no fibrous or other supporting tissue has been developed, and the
vascular bundles contain only spiral vessels, the result being that
they are very brittle and capable of resisting only the slightest con-
cussions. The effect of these abrasions is besides very much inten-
3« Groom, P., Bud protection in dicotyledons. Trans. Linn. Soc. II. 3:255- 1893-
ent -
1906] WIEGAND—BUDS AND TWIGS IN WINTER 21
4
sified by the comparative rigidity of the twig to which the buds
are attached. Danger to buds from mechanical causes during
the winter may be classified under three heads: danger from birds,
from passing objects, and from wind.
Birds—Young and tender plant tissue is a favorite food for
some birds, as for instance the purple finch.s? It is quite possible
that if the bud-scales were absent from our native trees, many species
of birds would take advantage of this easy method of obtaining
food, at a time when food is scarce. More birds might remain
in the North than now, so that altogether it seems probable that
trees would suffer severely, if they were not actually killed, by the
depredations. A firm hard armor is therefore desirable.
Passing objects—During my experiments with buds from which
the scales had been removed to determine the effect of evaporation,
€etc., many buds were prepared in a thicket of lilac bushes about
six feet high. I found great difficulty in passing through to inspect
the buds without breaking off some of them. A moment’s*absent-
mindedness while taking notes would frequently result in the de-
struction of several buds, a very slight touch only being necessary
to dislodge the tender shoot, and the brittleness was of course very
much increased when the tissues were frozen. If the above results
occurred when care was observed in passing, how much greater
would be the damage caused by animals both large and small run-
ning thorugh the dense copses in winter. Protective armor seems
here again to be a necessity.
Wind—In our American climate, at least, this is much stronger
in winter than in summer—in other words, at exactly the time when
buds are frozen and therefore most brittle. The beating together
of branches during a heavy wind storm could scarcely fail to do
incalculable damage to a tree with unprotected buds. ScCHUMANN*$
believed that one of the most, if not the most, important functions
of bud-scales is to resist injury from heavy winds. I myself was
32 Forsusn, E. fe Birds and woodlands. Mass. State Board of Agric. Rep.
1900:300. Brat, F. E. L., How birds affect the orchard. Year Book Dept. of
Agric. 1900: 2 291. Attu, Zerstérung von Baum- besonders Fichten- und Kiefern-
knospen durch Végel. Zeitschr. Forst. u. Jagdu. 29: 224-230. 1897.
33 SCHUMANN, C. R. G., Anatomische Studien iiber die Knospenschuppen von
3
Coniferen und dicotylen Holzgewichsen. Biblioth. Botan. 15:23. Cassel. 1889.
422 BOTANICAL GAZETTE [JUNE
much impressed by the abrasive power of objects beaten about
by the wind in winter in the case of firm paper tags attached by
strings two inches long to twigs of apricot. During the winter
they succeeded not only in marring the bark, but also in completely
disintegrating all the buds within reach by simple contact while
being blown about. It would seem that the beating together of.
larger and harder objects like the branches themsclves would do
even more damage. Therefore, in this casc also a firm outer coat
is demanded.
I believe that we are justified in concluding from these consid-
erations that mechanical protection is one of the most important
functions of the bud-scales—indeed the most important of all.
Some other suggested functions of bud-scales.
Gruss? included the storing of food material as one of the func-
tions of bud-scales. Bud-scales undoubtedly do store considerable
- @ . . 1
food in some cases, but this is a secondary rather than a primary |
function. We can scarcely consider this as having been an impor-
tant factor in the evolution of the scales.
GRooM®S suggests injury from excess of light as one of the things
from which bud-scales protect the delicate young tissue; that
when about to unfold the cell walls are thin and the chlorophyll
is dilute and easily decomposed. If we conclude that the red color
in young plants is for the purpose of modifying the light, then per-
haps there is some danger to the young tissues of the bud from too
strong light, since these usually become red. No direct evidence
however is at hand to warrant this conclusion, and since such pro-
tection could be obtained with the expenditure of less energy by
the use of hairs or bloom, there seems to be little reason for con-
sidering this a determining function of the bud-scales.
One of the most interesting suggestions is that of Capura*®
to the effect that in addition to protecting the delicate parts from
loss of water, great radiation, cold, and too great gas interchange,
34 Griiss, J., Beitrage zur Biologie der Knospen. Jahrb. Wiss. Bot. 23: 648. 189?-
35 Groom, P., Bud protection in dicotyledons. Trans. Linn. Soc. Il. 3:255- 1893-
36 CapuRA, R., Physiol gische Anatomie der Knospendecken dicotyler Laubbaiume.
Breslau, pp. 42. 1887.
i Es
1906] WIEGAND—BUDS AND TWIGS IN WINTER 423
bud-scales function beneficially by mechanically preventing too
early opening of the buds. That buds, as for instance the horse-
chestnut and lilac, do open several days earlier when deprived of
their scales I have frequently noted in connection with the experi-
ments conducted on desquamated buds in the greenhouse. Still
the evidence is not sufficiently strong to warrant the assumption
that this is an important and determining function of the scales.
Scalcless buds in nature instead of opening very early open quite
late, and, as ScHUMANNS’ has insisted, many buds during warm
wet autumns open in spite of the scales. My own observations
would tend to show that at best they can retard the opening not
more than three or four days.
SUMMARY.
Buds containing a considerable number of well differentiated or-
gans are usually protected by scales. Those sunk in the bark usu-
ally contain little besides the growing point or rudimentary leaves.
Bud-scales are not only the most feasible structures for covering
a large bud, but they also allow the bud to swell, and protect the
young shoot when unfolding.
The bud fundament in most trees is in dad down early in the sum-
mer, grows gradually till late autumn, remains dormant until early
spring, then passes through a period of swelling preparatory to
unfolding.
Dark-colored buds are usually warmer within than light-colored
ones; but the question whether there is any relation between dark
color and the early opening of the buds was not decided.
Ice may be found in most buds when the temperature has fallen
as low as— 18° C. and usually in large quantities. Its absence in other
cases is due to small-celled tissues and meager water content.
Frozen twigs are smaller than normal ones. Their contraction
occurs mainly in the bark.
Frozen buds do not show this contraction so plainly, probably
because of change of form in the bud-scales.
The thawing of fie if sufficiently slow, is per degree in inverse
37 SCHUMANN, C. , Anatomische Studien iiber die Knospenschuppen von
Coniferen und ak =r ieee Biblioth. Botan. 15:27. 1889.
424 BOTANICAL GAZETTE [JUNE
ratio to the freezing, so that at the freezing point of the tissue all
the ice will have disappeared.
Regarding the function of bud-scales, there is little evidence
that they function by keeping the water out; neither are they impor-
tant to the plant as modifiers of temperature.
Bud-scales have probably been evolved to prevent excessive
transpiration and to protect the delicate tissue from mechanical
injury.
When the bud opens the scales often grow out, forming a tube-
like structure which protects the young shoot from too great loss
of water. .
The wool in such buds as horsechestnut is not to modify the
temperature, but to protect the young shoot from too great transpi-
ration.
CORNELL UNIVERSITY,
Ithaca, New York.
ee
THE LIFE HISTORY OF POLYSIPHONIA VIOLACEA.
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY.
LXXXIII.
SHIGEO YAMANOUCHI.
Tuts preliminary paper will give a brief sketch of my cytological
studies on Polysiphonia violacea Grev., which were begun last sum-
mer at the Marine Biological Laboratory, Woods Hole, Mass., where
I occupied a Carnegie Research Table, and were continued at the
Hull Botanical Laboratory as a Fellow in the University of Chicago.
The problem was suggested by Dr. BRADLEY M. Davis, to whom I
wish to acknowledge my great indebtedness for his assistance and
criticism during the progress of the investigation.
Many points, which for the sake of brevity are omitted in this
paper, together with a discussion of literature will be presented in a
detailed account with plates to be published later.
METHODS.
The material was killed and fixed in Flemming’s fluids, in several
modifications which contained the osmic acid in various proportion,
Hermann’s fluid, I per cent. picric acid, and others; among which
the weaker formulae of Flemming proved most effective. The best
fixation for the study of spermatogenesis and the germination of car-
Pospores and tetraspores was obtained in material killed in weak
chrom-acetic acid (Flemming’s formula), without any osmic, as fol-
lows: 1 per cent. chromic acid, 25°°; 1 per cent. glacial acetic acid,
Io“; sea water, 65°.
Material was left in the fixing fluid five to forty minutes, and
then washed in a gentle stream of sea water. If material remains for
a longer time in chrom-acetic acid it becomes very soft and breaks
apart. The washed material was passed very gradually through a
series of alcohols beginning with 30 per cent., and imbedded in 52°
paraffin. The sections were cut 3-5 # in thickness and stained with
safranin-gentian-violet or with iron-alum-haematoxylin, sometimes
followed by some plasma stains as crange G, Bordeaux red, Congo
425] [Botanical Gazette, vol. 41
426 BOTANICAL GAZETTE [JUNE
red, or safranin. Preparations were studied with a Zeiss apochro-
matic immersion 1.5™™, N. A. 1.30, and compensating oculars.
GERMINATION OF THE CARPOSPORE AND TETRASPORE.
It is very easy to obtain the early stages in the germination of car--
pospores and tetraspores. Fruiting plants, placed in a dish of sea
water over night, will discharge great quantities of spores. These
fall to the bottom of the dish and germinate at once. The germi-
nating spores may be readily gathered from the bottom at the proper
hours to obtain critical stages.
The first division of the carpospores and tetraspores takes place
within 10-15 hours after their escape from the parent plants.
The cytoplasm before the first division shows a coarse network or
very irregular alveolar structure on the periphery, which becomes
much finer around the nucleus. The nucleus has a very delicate
membrane, within which lies the linin network, much finer in struc-
ture than that of the cytoplasm. The delicate transverse walls cf the
alveoli of the cytoplasm seem to end on the nuclear membrane where
the linin thread starts, which leads the writer to believe that there is
some relation between the positions of the walls of the cytoplasmic
alveoli and the linin of the nuclear network. The nucleus contains
one or two nucleoli homogeneous in structure.
Approaching the prophase of mitosis the linin threads become
more and more conspicuous and chromatin granules appear in rows;
but without constructing a uniform continuous spirem the threads
segment into a number of chromosomes. The nucleus becomes sur-
rounded by dense kinoplasm consisting of very minute closely crowded
granules, and the outer margin of this kinoplasmic mass assumes a
fibrillar structure which finally ends in the alveoli of the cytoplasm.
The distinct concentration of the kinoplasmic masses at the poles to
become the centers of the dynamic activities of the mitosis dces not
occur until the chromosomes are arranged in an equatorial plate.
The nuclear membrane persists through the prophase, which makes
it evident that the spindle is entirely intranuclear in origin.
The chromosomes at the equatorial plate split longitudinally, and
the two groups of daughter chromosomes pass to the opposite poles
of the spindle, where they become closely crowded in a mass near the
center of the accumulation of kinoplasm.
1906] YAMANOUCHI—POLYSIPHONIA VIOLACEA 427
The nucleoli sometimes fragment into two cr three small globules,
or decrease in size without fragmentation, finally vanishing during the
late prophase. New nucleoli appear after the formation cf the
daughter nuclei. No such close relation seems to exist between the
linin thread and the nucleolus as to warrant a belief that in Poly-
siphonia the nucleolar substance passes directly into the linin thread
to form the chromosomes, as is reported in Nemalicn.
No centrcsomes could be found during this process of mitcsis,
although the kinoplasm surrounding the nuclear membrane becomes
denser during later prophase and finally accumulates at the poles of
the spindle at the time of the metaphase. The masses of kinoplasm
present no radiations, yet it seems probable that they function as
centers of dynamic activity during mitosis, persisting until the daughter
nuclei are organized. |
The mitoses within the germinating carpospore and tetraspore cor-
respond in all essentials, except that it became at once apparent in
the investigation that the nucleus of the carpospore contained about
twice as many chromosomes as that of the tetraspore. Counts of the
chromosomes made during the later prophase and metaphase of the
mitosis made it clear that the nucleus of the carpospore contains
about 40 chromosomes and that of the tetraspore 20. :
SPERMATOGENESIS.
The mitoses in the vegetative cell of male or antheridial plants will
be described before those of spermatogenesis. During the prophase
the chromatin granules increase in size and become grouped as a num-
ber of short rod-shaped bodies upon the linin thread, without
developing a regular and uniform chromatin spirem. The linin
thread then segments into 20 chromosomes.
Polysiphonia, as a rule, is dioecious; however, cystocarpic plants
sometimes produce antheridia, a condition which will be described
later with other abnormalities. The antheridia develop as lateral
branches near the tips of the main filaments. Each branch consists
of a central axis from which clusters of sperm mother-cells or sper-
matocysts arise at the side on short stalk cells.
The cytoplasm of the spermatocyst has a delicate granular struc-
ture and contains a large vacuole, generally in the center of the cell.
Its nucleus in the resting state is similar to that of the vegetative cell,
428 BOTANICAL GAZETTE [JUNE
and during prophase the linin network becomes more conspicuous and
finally segments into 20 chromosomes. The kinoplasm around this
nucleus is rather scanty, even after the spindle fibers are formed.
The spindles are intranuclear. No centrosome could be found, but
there is a concentration of kinoplasm at the two poles.
After anaphase, the nuclear membrane dissolves and the vacuole
intrudes into the nuclear cavity between the two sets of daughter
chromcsomes, one set passing to the apex and the other remaining at
the base of the sperm mother-cell. The upper part of the cell, includ-
ing the vacuole, is then cut off as the sperm from the lower portion by
a cleavage furrow, which crosses the cell somewhat obliquely.
The nucleus which remains at the bottom of the sperm mother-
cell now repeats this mitosis, forming a second sperm, and perhaps
two or three more are developed before the antheridium ends its
fertility.
OOGENESIS AND FERTILIZATION.
The mitoses in the vegetative cells of the female or cystocarpic
plant are similar to those of the male. The number of chromosomes
is invariably 20.
The female organ or procarp develops from a central axial cell,
next to the apical cell of a short lateral branch. -The central axial
cell gives rise to a pericentral cell from which by successive mitoses a
four-celled carpogonial branch is formed. The terminal cell of this
series becomes the carpogonium, situated as a rule above the peri-
central cell, owing to the curved growth of the carpogonial branch.
The carpogonium is at first round and the nucleus lies in its center.
While this nucleus undergoes a typical mitosis to form two nuclei, the
carpogonium puts forth the process which is to become the trichogyne.
After mitosis one of the nuclei proceeds into the developing tricho-
gyne to become a trichogyne nucleus, and the other remains below
in the carpogonium as the gamete nucleus.
Ccincident with the development of the trichogyne, the pericen-
tral cell gives rise to the two sets of auxiliary cells, one of which is a
series of three or four, formed above, so that they lie just beneath
the carpogonium, and the other series consists of two cells formed
below.
When the sperm comes in contact with the trichogyne, the walls
c
_,
1906] YAMANOUCHI—POLYSIPHONIA VIOLACEA 429
between dissolve, the contents of the sperm cell enter the cytoplasm
of the trichogyne, and the sperm nucleus passes down into the car-
pogonium where the fusion of the male and female nuclei takes place.
The trichogyne nucleus may be found even after the sperm nucleus
has passed into the carpogonium. But later, when the trichogyne
becomes separated from the carpogonium, its nucleus can scarcely
be distinguished. The trichogyne then shrivels and dies.
The carpogonium after fertilization unites with one of the auxiliary
branch cells which lies beneath, and the latter also fuses with the peri-
central cell, thus providing a passage into the pericentral cell for the
fusion nucleus of the fertilized carpogonium. Then the remain-
der of the auxiliary branch cells fuse with one another and with
the pericentral cell, which results in a large fusion cell, the central
cell, that naturally contains a number of nuclei.
The nuclei in the central cell are of two sorts with respect to origin:
first, there is the fusion nucleus from the carpogonium (sporophytic) ;
and second, there are a number of nuclei derived from the auxiliary
cells, which are of course gametophytic. The fusion nucleus gives
rise to a series of nuclei by typical mitoses which present 40 chromo-
somes as a sporophyte number. The central cell now develops
several lobes into which these sporophytic nuclei pass. Further
mitoses increase this number, and each lobe then cuts off a carpospore
terminally, which is attached to the central cell by a short stalk.
After the carpospores are formed, the central cell increases in size
greatly, absorbing the stalk cells, and even the central axial cell also
becomes involved -in_ this general cell fusion. These very exten-
sive cell unions are probably concerned with the nourishment of
the carpospores.
Some of the gametophytic nuclei derived from the auxiliary cells
remain in the central cell, increasing in size and finally breaking down
after a number of peculiar changes. Others of the gametophytic
nuclei divide amitotically to form the paranematal filaments which
lie under the wall of the cystocarp.
TETRASPORE FORMATION.
The tetrasporic plant normally never produces antheridia or pro-
carps, and the cytological studies on the vegetative cells give proof
that it differs in an important respect from the sexual plants. The
430 BOTANICAL. GAZETTE [JUNE
mitoses in growing regions of the tetrasporic plant show that the
nuclei have 40 chromosomes (the sporophyte number), while it will
be remembered that the nuclei of the sexual plants have 20.
I shall not enter at this time into a detailed description of the events
which take place during the formation of the tetraspore mother-cell;
the only thing to be remembered is that the number of the chromo-
somes appearing during this mitosis is 40,so that it follows that the
nucleus of the tetraspore mother-cell contains 40 chromosomes.
The nucleus of the tetraspore mother-cell increases somewhat in
size, accompanied by the growth of the cell itself; yet the latter is
relatively slow until just before the first mitosis of the nucleus, but
very rapid after that.
The resting nucleus of the tetraspore mother-cell contains a fine
network of linin in which the chromatin is distributed irregularly in
larger and smaller granules. The nucleolus has no visible connection
with the linin thread. With the further growth of the nucleus the
linin thread increases in thickness; in such an irregular way, however,
that in some parts the threads are uniform in thickness and in the
others they appear to have knots. The chromatin thread now forms
a fairly well-developed spirem.
This condition presently passes into the so-called stage of synapsis,
when the spirem consists visibly of two parallel threads close together,
while in the other parts the two are in contact side by side or fused
into a single thread. The two threads may represent, according to
recent interpretations of synapsis, chromatin of maternal and paternal
origin.
After synapsis, the tangled thread becomes distributed throughout
the cavity of the nucleus. The spirem now shows the longitudinal
fission which precedes the separation of chromatin granules into two
sets, and then the spirem segments into 20 chromosomes, each show-
ing clearly its bivalent nature. :
While this process of chromosome formation is going on in the in-
terior of the nucleus, the kinoplasmic material surrounding the nucleus
becomes concentrated at two poles of a spindle, and when the chrc mall
somes are arranged in the equatorial plate a minute body cccuples
the center of each pole. The body might be called a centrcsome,
but it has not been possible to recognize its presence during prophase
or to follow it after anaphase.
1906] YAMANOUCHI—POLYSIPHONIA VIOLACEA 431
It is interesting to note that the two poles are not in a straight line
perpendicular to the center of the equatorial plate, but are asym-
metrical and less than 180° apart. Each of the 4o sporophytic
chromosomes composing the 20 pairs (bivalent chromosomes) ar-
ranged in the equatorial plate now splits longitudinally, so that a
large number of univalent chromosomes results, probably 8o in all,
although I was unable to count the exact number.
The group of 80 grand-daughter chromosomes separates into
two sets, but resting neuclei are not formed, and their further distri-
bution by the second mitosis begins at once. The axes of the two
spindles of the second mitosis lie perpendicular to each other, and
their complex relation to that of the first will be described in the
final paper. Kinoplasmic masses occupy the poles of the spindles
in the second mitosis, and each contains a centrosome-like granule.
Now, each group of 40 chromosomes, following this first mitosis,
separates into two sets of grand-daughter chromosomes, 20 in each
set, which are attracted toward the respective poles of the two spindles,
where the kinoplasmic material becomes more pronounced than be-
fore. These chromosomes, after reaching the four poles, become
massed together, lose their individual outlines, and larger and smaller
granules appear shortly after on linin threads which later become
contained in the four daughter nulcei.
It is a remarkable fact that the membrane of the original nucleus
in the tetraspore mother-cell persists through the two mitoses which
have just been described. The area included by this membrane
increases in size with the growth of the cell. The cytoplasm around
it shows larger alveoli, which become smaller in the vicinity of the
original nuclear membrane, and at last passes into the kinoplasmic
fibrils immediately surrounding it.
At this time constrictions appear simultaneously in the area marked
by the original nuclear membrane. The kinoplasm intrudes into this
area in a very interesting manner, which cannot easily be described
without figures, but results in the organization of the four daughter
nuclei that are to be contained in the tetraspores.
The division of the tetraspore mother-cell does not take place
simultaneously with the events described above. Cleavage furrows
start along four lines on the periphery of the cell, a little before the
end of the nuclear division, and prcceed more rapidly after its com-
432 BOTANICAL GAZETTE [JUNE
pletion. During the entire process of the tetraspore formation, the
mother-cell is connected by a strand of protoplasm with the stalk-
cell, and probably gets much nourishment through this strand, for the
tetraspore mother-cell increases greatly in size. Finally, the cleavage
furrows meet in the center between the four daughter nuclei, thus
dividing the protoplasm into four tetraspores.
ABNORMALITIES.
Normally, the male and female organs and the tetraspore are never
developed in the same plant, but it often happens that the male and
female organs are produced on the same individual, and cccasionally
antheridia are formed on the same branch with procarps and cysto-
carps. The sexual cells in these cases are developed normally, the
number of the chromosomes being always 20. Cystocarpic plants
have also been found producing cells whose lineage is identical with
that of the tetraspore mother-cell. However, I have never seen evi-
dence of nuclear division in such a cell; cleavage furrows appear
and cut deeply into the protoplasm, which nevertheless remains un-
divided, so that tetraspores are not formed. Whether this cell may
escape and germinate as a monospore has not yet been determined.
CONCLUSION.
The nuclear conditions in the life history of Polysiphonia may be
summarised as follows:
1. The germinating carpospore contains 40 chromosomes, and
the tetrasporic plant the same number; so it may be inferred that the
tetrasporic plants. come from carpospores.
2. The germinating tetraspore contains 20 chromosomes, and the
sexual plants (gametophytes) the same number; so it may be inferred
that the sexual plants come from tetraspores.
3- The nuclei of the gametes (sperm and carpogonium) contain
each 20 chromosomes. The fusion nucleus (sporophytic) in the fer-
tilized carpogonium presents 40 chromosomes, and gives rise to a series
of nuclei. Some of these enter the carpcspores, which are conse-
quently a part of the sporophytic phase to be continued in the tetras-
poric plant. The gametophytic nuclei in the central cell of the
cystocarp (with 20 chromosomes) either break down or form the
paranematal filaments.
1906] YAMANOUCHI—POLYSIPHONIA VIOLACEA 433
4. Tetraspore formation terminates the sporophytic phase with
typical reduction phenomena, so that the tetraspores are prepared to
develop the gametophytic generation.
5. There is thus an alternation of sexual plants (gametophytes)
with tetrasporic plants (sporophytes) in the life history of Polysiphonia,
and the cystocarp forms a part of the sporophytic phase.
THE UNIVERSITY OF CHICAGO.
THE STRUCTURE AND DEVELOPMENT OF THE
BARK IN THE SASSAFRAS.
HowarRD FREDERICK WEISS.
(WITH NINE FIGURES)
THE common sassafras occupies a somewhat isolated position
among northern trees. It is not only the single living representative
of the genus Sassafras, but it belongs to the Lauraceae, a family with
many arboreal genera in tropical and subtropical regions, but with very
few in the cooler parts of the earth. The tree is further remarkable
because its young branches remain green for a considerable period,
differing in this respect from the majority of the trees among which it
grows. For these various reasons it was hoped that a study of the
bark might reveal features of interest.
Métter has already studied the bark in several genera of the
Lauraceae and has included in his published account a short descrip-
tion of what he found in the sassafras.t According to his researches
the family as a whole is characterized by the following peculiarities in
the bark: a late appearance of cork; an epidermal origin of the phel-
logen; a slight development of collenchyma in the outer cortex, most
of the cells remaining thin-walled and parenchymatous; the occur-
rence of stone-cells in the medullary rays between the strands of
primary sclerenchyma; the presence of ethereal oil and slime in some
of the parenchyma cells; the scattered bast fibers in the inner or second-
ary bark. With regard to the sassafras in particular he notes that
the cork is homogeneous and composed of thin-walled cells and that
the inner bark is destitute of stone cells. It should be remarked that
most of MOLLER’s material in this family consisted of dried bark,
much of which was fragmentary and in poor condition.
In his more general account of the Lauraceae SOLEREDER accepts
the majority of M6LLER’s statements with regard to the bark.” Quot-
ing from J. E. Wetss, however, he notes the fact that the phellogen is
not invariably epidermal in origin, but that it is sometimes deriv ed
t Anat. der Baumrinden 103-110. 1882.
2 Syst. Anat. der Dicot. 795. 1899.
Botanical Gazette, vol. 4r] [434
——
>», . — Con a> =
= et
1906] WEISS—BARK IN SASSAFRAS 435
from the layer of parenchyma just within the epidermis. He also
remarks that the secondary bast fibers, although usually scattered,
‘form distinct strands in certain genera, and that the individual fibers
are normally four-sided in cross section with narrow lumina.
The present investigation is based on material collected near New
Haven, Connecticut, and is confined to the stem and its branches, no
reference being made to the bark of the root. The tissues described
may be classified as follows: .
PRIMARY TISSUES SECONDARY TISSUES
Epidermis Tissues derived from the cambium ring
Outer cortex The phellogen and its derivatives
Primary medullary rays
Primary bast
PRIMARY TISSUES.
Epidermts.
The epidermal cells are characterized by a strongly thickened
cuticle. Close to the growing point they are isodiametric and thin-
walled, but the cuticle begins to make its appearance very early and
OADOOSD
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Fic. 1.—Cross-section through bark one year old. X70. cam, cambium ring;
¢, epidermis; m, medullary ray; p, parenchyma; phx, primary phloem; s¢s, primar)
sclerenchyma; sc,, secondary sclerenchyma; st, stone cells; x, xylem.
Practically completes its development during the first year’s growth.
At the close of this period it occupies about half the thickness of the
epidermis (jig. 1). During the elongation of the stem the epidermal
cells retain the power of growth and division. Since their growth is
largely in a longitudinal direction, the cell-division is mainly brought
about by transverse walls, division by longitudinal walls being much
436 BOTANICAL GAZETTE [JUNE
more infrequent. In an epidermis a year old, seen from the surface,
the boundaries of the original epidermal cells can usually be distin- »
guished. They are somewhat thicker than the secondary transverse
oad) < walls, which in turn are thicker than the secondary
longitudinal walls (jig. 2). With the formation of
es cork the epidermis is of course split longitudinally
and soon begins to undergo disorganization. No
trace of it is left in a tree 8°™ in diameter.
Or The number of stomata produced varies greatly,
L000 SEE
but seems to be largely dependent upon external
conditions. A rapidly growing tree, for example, in
4 a moist locality has many stomata, while a slow-
e growing tree in dry soil develops very few. The
.
YO
stomata are depressed and the epidermal cells bound-
ing the guard cells are somewhat modified, being
longer and narrower than their neighbors (fig. 3)-
Fic. 2.—Surface Most of the stomata are transverse to the axis upon
view of epidermal which they are borne, a few are oblique, but appar-
ae nay Bag ently none of them occupy a longitudinal position.
ain ie miei’ ary Lhis is doubtless to be explained by the fact that
of the original cell, the stomata are formed late in the development of
which has under- the epidermis, the wall separating the guard cells
liad i representing one of the secondary transverse divisions
of an epidermal cell. In the majority of
cases the cells surrounding a stoma contain nie ‘ik
anthocyan, so that to the naked eye the
stomatal region looks like a minute red
speck in the epidermis. This peculiarity _}{ \@/¢ $#
affords a ready means for detecting the SS LE
stomata. LY aa
Epidermal hairs are developed on very ae
young twigs before the primary tissues are Fic. 3.—Stoma_ with sur-
fully differentiated. They are simple and TU"dins ie nana crcaals
unicellular, with thickened walls, and scarcely cage
extend below the cuticle (fig. 4). These hairs never persist through
the first vegetative period, but dry up and fall away as soon as the
cuticle begins to thicken. Their former position is often marked by
iG
ay 4,
x,
eS |
1906] WEISS—BARK IN SASSAFRAS 437
small concave depressions in the cuticle. The number of the hairs
"varies, and in a general way is inversely proportional to the number
of stomata. Thus, in a moist locality few hairs are formed, while
in a dry region they are very abundant. In a meso-
phytic area some trees bear few hairs, while others
under the same conditions bear very many. It would
appear from this that the production of hairs was
primarily due to individual peculiarities of the tree in
question and secondarily to the external conditions
under which the tree developed.
Outer cortex.
The outer cortex comprises everything external to yc, 4-—Epi-
the primary sclerenchyma except the epidermis. It is dermal hairs on
composed of a ground mass of parenchyma with ia iis
scattered stone-cells. No crystal cells occur. With ”
the formation of cork the outer cortex gradually becomes disorgan-
ized and eventually disappears. .
In cross section the parenchyma cells vary from elliptical to rec-
tangular in outline, the long diameter running in a tangential direction
(jig. 1, p). They vary considerably in size and some of the larger
cells have their walls slightly lignified. Most of the cells, however,
have thin walls, which may or may not be provided with simple pits.
Many of the smaller cells contain starch and this is especially likely
to be true of those which border the strands of sclerenchyma. The
presence of ethereal oil in the parenchyma can be demonstrated by
appropriate tests, but it does not seem to be localized in special cells.
In all probability the oil represents an excretory product of the proto-
plasm of the parenchyma cells, and this fact would account for its
general distribution.
The stone cells form a continuous or interrupted layer extending
entirely around the stem (fig. 1, st). They sometimes lie next to the
epidermis and are sometimes separated from it by one or two layers of
parenchyma cells. The stone cells are at first circular in cross section
but afterwards become flattened and assume an elliptical outline.
In radial section they appear rectangular, being about three times as
long as broad. Their walls are strongly thickened by deposits of
438 BOTANICAL GAZETTE [JUNE
ligno-cellulose in distinct layers, and these are pierced by numerous
simple and branched pits.
Primary medullary rays.
The primary medullary rays extend from the cambium to the
outer cortex, the ray cells merging into the cortical cells without a
distinct line of demarcation. The outer portion of the ray is of
course directly differentiated from the meristem at the growing point,
while the inner portions owe their existence to the activity of the
cambium. Some of the cells in the outer portion retain their power
of growth and division for several years, the majority of the dividing
i any.
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Fic. 5.—Cross section through old bast. X25. Fic. 6.—Radial section through
old bast. X75. c, cork; cam, cambium ring; e, epidermis; m, medullary ray,
p, parenchyma; ph, secondary phloem; phel, phelloderm; sc2, secondary sclerenchyma;
St, stone cells; x, xyiem.
walls being radial. Thus, in a stem one year old, the strands of
primary sclerenchyma are separated by from two to five layers of cells,
in a stem two years old by as many as fifteen layers, while in a stem four
years old the number may be increased to thirty or more. Since the
portions of the rays derived from the cambium do not undergo further
divisions, they remain permanently from one to three cells in width.
In consequence of these facts the rays gradually assume a T-shape
a
1906] WEISS—BARK IN SASSAFRAS 439
in cross section. This form is retained until the outer cortex has
become disorganized, after which they appear like narrow bands (ig.
5,m). In radial section the rays are from four to fourteen cells across
(fig. 6, m).
In most of the ray cells the walls are slightly thickened and pro-
vided with numerous simple pits. They usually contain starch and
sometimes ethereal oil as well. When the cells are cut off by cork
the starch disappears, showing that it is completely utilized; the oil,
on the other hand, persists. Some of the ray cells between the strands
of primary sclerenchyma become strongly sclerotic, and in some cases
cells of this character completely bridge the space from one strand
to another (fig. 1). They can be easily distinguished from the scle-
renchyma cells, even in cross section, by their larger size and distinct
lamination. In longitudinal section they appear short and resemble
the stone cells of the outer cortex.
Primary bast.
The primary sclerenchyma occurs in well-defined bundles, averag-
ing about fifty fibers apiece (figs. 1, 8, sc,). Most of these bundles,
in a radial direction, measure from three to eight cells across. In
most of the fibers the wall is so strongly thickened that the cavity is
reduced to a mere slit; in some cases, however, the thickening is less
and this is especially likely to be true in the middle of a bundle.
Apparently the deposition of ligno-cellulose upon the cell walls is not
completed until the second vegetative period.
The primary phloem lies just within the primary sclerenchyma,
between the latter and the secondary sclerenchyma, and forms a band.
from three to five cells across in a radial direction (fig. 1, ph,). The
Sieve tubes are more or less completely separated from the scleren-
chyma by a layer of phloem parenchyma. The cells of this layer
tend to be rectangular in cross section, and their slightly thickened
walls have numerous simple pits. The sieve plates separating the
segments of the sieve tubes are nearly always somewhat oblique 3
they are supplemented by numerous lateral sieve plates, especially
in the radial walls of the tubes. All of the sieve plates in the primary
phloem soon become covered by deposits of callus. The companion
cells conform to the usual type.
440 BOTANICAL GAZETTE [JUNE
SECONDARY TISSUES.
Tissues derived from the cambium ring.
The tissues of the bark, regularly derived from the cambium ring,
include the secondary sclerenchyma and the secondary phloem. In
addition to these, scattered groups of stone cells, which should prob-
ably be considered a part of the phloem, also make their appearance.
Of course the cambium also adds new elements to the primary medul-
lary rays and brings about the development of the secondary rays
(figs. 1, 5). The development of these various secondary tissues
begins during the first vegetative period.
The fibers of the secondary bast do not form bundles. Some of
them form interrupted layers arranged concentrically in the stem,
others are scattered through the secondary phloem. The layers ar?
usually but a single cell across and are separated from one another
by several layers of phloem. The individual fibers are rectangular
in cross section and about thirty times as long as broad; their walls
are very strongly thickened (jigs. 1, 5, 6, sc,). When the bast fibers
are cut off by cork all regularity in their arrangement disappears.
The sieve tubes of the secondary phloem, except those earliest
formed, are arranged in interrupted, concentric layers, one or two
cells across (fig. 5, ph,). Many of the sieve tubes are in direct con-
tact with the medullary rays, but very few of them adjoin the scleren-
chyma fibers. The tubes exhibit essentially the same structure ‘as
those in the primary phloem. On account of their delicate walls they
become practically indistinguishable when cut off by cork.
The bulk of the secondary phloem is composed of parenchyma.
When first differentiated from the cambium the cells of this tissue
are closely packed together, rectangular in outline, and destitute of
intercellular spaces. As they become pushed outward, their outlines
become more rounded and minute intercellular spaces appear. Their
walls are fairly thin but are provided with simple pits. Until they are
cut off by cork the parenchyma cells are arranged in layers, which lie
among the layers of sclerenchyma and sieve tubes.
The groups of stone cells are irregularly scattered in the secondary
bast but always abut against a medullary ray (fig. 5, sé). Such a
group in cross section is often larger than a bundle of primary scle-
renchyma and is composed of larger elements. The stone cells are the
1906] WEISS—BARK IN SASSAFRAS 441
most conspicuous structures found in the inner bark, and are even
more striking in appearance than those found in the outer cortex.
In longitudinal section (fig. 6, st) they show the same outlines as in
cross section (jig. 7) and are therefore isodiametric. Their strongly
thickened walls show a very distinct
lamination and their contracted cavi-
ties are connected by numerous
simple and branched pits. Prob-
ably on account of poor material,
these stone cells were not seen by
MOLLER.
The phellogen and its derivatives.
The derivatives cf the phellogen
are the lenticels, the cork, and the
phelloderm. The lenticel phellogen
is the first to make its appearance;
the primary cork phellogen is, at
least in part, a direct extension of Fis. 7.—Stone cells from inner bark,
the lenticel phellogen; and the suc- ‘S S*HO™ *45°
ceeding phellogens arise more or less independently from the deeper
layers of the bark. The primary cork phellogen first appears on
the south side of an erect stem and normally on the upper surface
of a horizontal branch. From these regions it gradually extends
laterally and usually forms a complete layer in the course of three or
four years. The development, however, follows no definite rule.
For example, in one eight-year old stem there was no cork on the
north side except in the immediate vicinity of the lenticels, while in
another stem of the same age there were five layers of cork on the
South side and three on the north. These observations show that
a phellogen layer may be active in one part although it has ceased
to be functional in another. They also show that there is no definite
relationship between the age of the stem or branch and the number
of layers of phellogen. The early appearance of cork in the regions
exposed to the sun is probably due to the fact that the sassafras is an
intolerant species and that the cork protects the deeper tissues from
Sun scalding.
442 BOTANICAL GAZETTE [JUNE
The primary lenticels are always formed directly beneath the
stomata, following in this respect the general rule first enunciated by
TRECUL. Some of the lenticels never break through the epidermis
but remain in an undeveloped condition. The lenticel phellogen
arises from the layer of cells just within the epidermis. The thin-
_ walled complementary cells are at first closely packed together.
After about twelve layers of these cells are formed the epidermis is
ruptured, and the complementary cells as they become exposed sepa-
rate from each other and present very irregular and distorted outlines.
The mature lenticel agrees with the second of the types described by
Devaux‘ and shows no distinct layers of cork among the. comple-
plementary cells (jig. &). In some
cases, however, a lenticel contains a
few scattered stone cells (fig. 8, st).
Secondary lenticels are developed from
secondary phellogens and make their
appearance in the splits of the bark.
These lenticels break through the
gt
Pig.ic:
*i
[|
i
OS
é OY
ERS
we
(f
ef
A
O
eo <a eVox Ss :
Sie Ogely es
FIG. 9. ere ee
Fic. 8.—Section through a primary lenticel. X55. Fic. g.—Section through @
secondary lenticel. X60. c, cork; com, complementary cells; e, epidermis; ~, paren-
chyma; phel, phelloderm; sc,, primary sclerenchyma; st, stone cells.
layers of cork and parenchyma cells which enclose them and eventu-
ally exhibit the same structure as the primary lenticels (fig. 9).
Since the primary cork phellogen is a direct extension of the lenti-
cel phellogen, it is never epidermal in origin but is always derived
3 Compt. Rend. 73:15. 1871. 4 Ann. Sci. Nat. Bot. VIII. 12:61. 1900-
EE
1906] WEISS—BARK IN SASSAFRAS 443
from the subepidermal parenchyma. In the majority of cases it
arises from the layer of cells just inside the epidermis. Sometimes,
however, it is derived from the second, third, or fourth layer, and this
is always the case when stone cells are present next the epidermis.
It thus frequently happens that the different parts of the phellogen
do not all arise from the same layer of cells. The secondary layers
of phellogen are largely derived from the parenchyma cells in the
secondary phloem. When stone cells are present in the parenchyma
the phellogen often bounds them on the inside. The phellogen forms
concentric layers in the stem, but these layers are not altogether
independent. In certain regions two layers will coalesce, in other
regions they will be separated from each other by several layers of
cells. Even the outermost of the secondary phellogens is more or
less united with the primary phellogen.
The cork, as already noted by MOLLER, is of the ordinary type.
It consists of empty cells arranged in radial rows, and the walls are
thin and suberised ( jigs. 5, 6, 9, c). In most cases from ten to twelve
layers are formed by each phellogen. The structure of the cork is not
uniform throughout the Lauraceae; in certain genera it consists of
two kinds of cells arranged in more or less definite layers; namely,
thin-walled cells and cells in which the inner tangential walls are
thickened.5
- The phelloderm in the sassafras forms a most characteristic feat-
ure of the bark. When derived from secondary phellogens it con-
sists almost entirely of strongly flattened cells with thick lignified
walls, provided with simple and branched pits. The flattening is in
a radial direction, and the cells show the same rectangular outlines
in both radial and transverse sections (figs. 5, 6, phel). The phel-
loderm is arranged in layers from one to three cells thick. The layer
derived from the primary phellogen differs from the others in being
composed of both thin-walled and thick-walled cells. In the case of
lenticels the thick-walled phelloderm cells are few and scattered and
are sometimes absent altogether. Lignified phelloderm does not
seem to be of very frequent occurrence. According to J. E. Wetss®
it is to be found in species of Cytisus and Philadelphus; KUHLA’
5 See MOLLER, Anat. der Baumrinden 103
. 1882.
© Beitrage zur Kenntniss der Korkbildung. Seiten. K®6nigl. Bayer. Bot.Gesells.
6:61. 1890.
7 Bot. Centralbl. '71:196. 1897.
444 BOTANICAL GAZETTE [JUNE
describes it for Ptelea trifoliata, and SOLEREDER® notes its appearance
in several genera of the Saxifragaceae other than Philadelphus. It
therefore occurs in widely scattered families and probably has but
little taxonomic significance.
SUMMARY.
Among the more interesting points brought out by this study are
the following: the early thickening of the cuticle; the variation in
the number of epidermal hairs and stomata; the early formation of
cork in regions exposed to the sun; the stone cells in the outer bark,
between the strands of primary sclerenchyma, and in the inner bark;
the regular layers of thick-walled phelloderm derived from the second-
ary phellogens.
The writer is indebted to Professor ALEXANDER W. Evans for
criticism and advice.
SHEFFIELD SCIENTIFIC SCHOOL,
YALE UNIVERSITY.
8 Syst. Anat. der Dicot. 360. 1899.
|
BRIEPER ARTICLES.
THE DISTRIBUTION AND HABITS OF SOME COMMON OAKS.
WHEN doing some work in Wisconsin last year for the Arnold Arbore-
tum, I found that Quercus ellipsoidalis E. J. Hill was well represented in
the woods of the southeastern part of that state. It was originally described
from trees growing in the vicinity of Chicago. It had been identified
by those studying the flora near Milwaukee, and is quite abundant on the
hills of the Kettle Range. It had also been recognized as distinct by those
unfamiliar with botanical works, as disclosed by the common name “pin
oaks.’”’ I had not before heard this name applied to any except Q. palustris
Moench. The original description mentioned the usual but not universal
drooping of the lower branches, as is quite common in the pin oaks.
When finding it in some new locality I have sometimes been at a loss to
decide which of the two species it was till the acorns were in hand,
As the branches often come low down, they are apt to die as the trees
grow older, and, breaking off a short distance above their base, leave stubs
along the trunk, so characteristic of the pin oaks. This was freely the
case in most of the trees seen in Wisconsin, and doubtless explains the
local name.
Quercus palustris was not seen in any of the localities visited, nor did I
learn of its presence from those familiar with the flora. In 1846 Dr. Lap-
HAM mentions its occurrence at Milwaukee in a book containing “A list of
plants which have not before been noticed as indigenous to Wisconsin.””!
It was mentioned again by him in a paper on the ‘Plants of Wisconsin.”’
Though no locality is specified, it is understood from a prefatory statement
to have been “ within thirty miles of Milwaukee.” If rightly identified (and
Dr. LapHam was a careful and competent observer), it would seem to have
disappeared. Yet there is the possibility that the tree with drooping
lower branches with stubs along the trunk, and finely divided leaves, going
by the common name of pin oak, was the one he alluded to, since the
common name is added to the botanical in both of the above citations.
That botanists have been bothered by some form ascribed to Q. palustris
or Q. coccinea is apparent from a statement of Dr. GrorGE Vasey in an
* Wisconsin: its geography and topography, history, geology, mineralogy, etc.
Milwaukee, 1846, p. 73.
2 Proceedings of American Association for the Advancement of Science 1849:19.
445] [Botanical Gazette, vol. 41
a
446 BOTANICAL GAZETTE [JUNE
article on “Our native oaks.” ‘It (Q. palustris) is found in low and
swampy ground, and in general appearance much resembles the scarlet
oak (Q. coccinea), and perhaps may yet have to be considered a variety of
that polymorphous species.”3 Dr. VAsEy resided for several years in
northern Illinois, and could hardly have failed to see such forms of Q.
ellipsoidalis as have led to its being confounded, by the common people at
least, with the pin oak. Buta typical scarlet oak is a tree of quite a different
aspect from Q. palustris, and from its habitat would be more easily con-
founded with Q. ellipsoidalis. It is true that Q. palustris is commonly con-
fined to low ground, though not always swampy, as along the margins of
streams which have cut their beds deep down into the drift, leaving a high
bank. Here the pin oak holds its place on ground that trends away from
the stream and is comparatively dry. I have seen it along the Kankakee
River move out of a swampy area to a bordering locality where the lime-
stone was but a few inches below the surface. And although Q. ellip-
soidalis commonly grows on dry or upland ground, it also occurs in lower,
even wettish, localities, as by the borders of ponds and sloughs in low
woods, becoming a near neighbor of the swamp white oak (Q. platinoides).
Those seen in Wisconsin were on hills of till, or by the borders of lakes
in the Kettle Range, or in soil of glacial drift. The least frequent of the
biennial-fruited oaks associated with it seemed to be Q. velutina. Q.
coccinea was quite common; Q. rubra the most abundant of all. In
Illinois I have most frequently met with it in woods adjacent to streams
not subject to overflow, the morainal hills being taken, when wooded, more
by Q. coccinea, Q. rubra, Q. velutina, and Q. imbricaria, in prevalence some-
what in the order given.
It is therefore a matter of some doubt whether Q. palustris now occurs
in Wisconsin. In Minnesota it is mentioned in UpHam’s “Catalogue of
the plants of Minnesota” on the authority of Dr. Lapuam, the locality
not being given; and on the authority of another collector as found in the
region of the Upper Mississippi. I have not been able to get these state-
ments verified. The pin oak of Minnesota may also be Q. ellipsoidalis.
Professor SARGENT identifies this in specimens collected at the Falls of
Minnehaha in 1878, and states that he himself first saw the species in 1882
at Brainard on the Red River of the North and at St. Paul. In his report
on the forest trees of North America, tenth census, volume 9, Q. palustris
is given for Wisconsin; but in his account of the tree in the eighth volume
of the Silva, this state is omitted from its range, as well as in his more
recently published Manual of the trees of North America. Both Wisconsin
3 The American Entomologist and Botanist 2: 376. 1870.
4Silva of North America 14:50. 1902.
item cttemant Deana
1906] BRIEFER ARTICLES 447
and Minnesota are rather far north for its range. The farthest north I
have found it in Illinois is in the town of Niles, just north of Chicago.
Nor can I find any record of its occurrence in the more northerly counties
of the state, where, if occurring at all, it is evidently very scarce. Dr.
FRIEDRICH BRENDEL of Peoria, in an article on ‘‘The trees and shrubs of
Illinois,” says ‘‘The pin oak (Q. palustris Du Roi) I have never seen
around Peoria, nor did, as I learn by letter, Mr. HALL in Menard County;
it occurs in St. Clair and Marion Counties; in Wisconsin and Cook County
(fide Mr. JacKson).’”’> The credit to Wisconsin is doubtless due to Dr.
Lapua, already cited. South of Chicago this oak appears in the southern
part of Cook County in the town of Thornton, extending sparingly up
Thorn Creek for a short distance, where it grows in company with Q.
ellipsoidalis. It is most abundant east of the village of Thornton, making
a good part of a wood growing in a soil of sandy peat, patches of sphagnum
being common under the trees. Eastward it is found in occasional spots
and in similar soils, and in the clayey soils of swamps in Lake and Porter
Counties, Indiana. It comes into the dune region of Lake Michigan north
of the village of Porter, in a sandy humus soil similar to that near Thornton.
Southward from here in Indiana it increases in frequency and abund-
ance. In eastern Illinois it reappears south of the Thorn Creek localities
after one crosses the range of hills here forming the water-shed of Lake
Michigan basin (the Valparasio moraine), and is oe along: the
Kankakee River at Momence.
Whether Q. ellipsoidalis occurs south of the most northern counties of
Indiana there is no evidence at hand to show. Some time spent in examin-
ing the flora in the vicinity of North Judson and English Lake in Stark
County did not reveal its presence, though the pin or Spanish oak was
common along the Kankakee River there. Specimens of oaks sent from
Bluffton in the eastern part of the state, a short distance south of Fort
Wayne, lacked this: species, but contained Q. palustris and Q. texana
Buckley. - '
It is evident from this survey that Q. ellipsoidalis replaces to a large
extent in the north of the Middle West the more southerly Q. palustris.
But it is usually with a different and drier habitat, and an adaptability
to a wider range of conditions. The boundaries of the two overlap in
southern Michigan, northern Indiana, northern ‘Illinois, eastern-central
Towa, possibly in southern Wisconsin. It may also be of interest to add
that the northern bounds of another biennial fruited oak, the shingle oak
(Q. imbricaria) correspond quite generally with those of Q. palustris.—
-E. J. Hitz, Chicago.
5 Illinois Agricultural Report 1859:596.
CURRENT LITEKATURE.
BOOK REVIEWS.
Botanical dictionary.
IN 1900 JACKSON published the first edition of his Glossary of botanic terms,
and last fall the second edition appeared.‘ We welcomed the first edition?
as being a marked improvement upon any existing dictionary, and criticized
but lightly the most obvious shortcomings. The compiler, most competent
in many respects, had certain limitations by reason of his unfamiliarity with
the content and consequently the terminology of morphology and physiology,
and our general criticisms lay along these lines.
In judging the second edition one looks to see whether this weakness of the
first has been removed, either by the author’s own efforts, or by his associating
with himself those who could supply the lacking knowledge. We find that the
“revised and enlarged” of the title page means only that typographical and
minor errors have been corrected in the plates of the first edition, and that a
supplement of 68 pages has replaced the former “Additions during printing.’”
ne can overlook much in a first edition that cannot be forgiven in a second.
Perhaps there will be a third with a resetting that will allow the necessary improve-
ment. In that hope we may point out certain objectionable features that should
receive attention.
In the first place it would be desirable to relegate to a separate list the many
terms which have become obsolete, most of which are adopted from LINDLEY’S
Glossary and were antiquated in his day. Technical language changes rapidly
and such terms should be put into a museum and labeled as exhibits, if shown
at all. We should then escape reading (except we were on antiquarian research
bent) that an ovule-tube is “‘a thread-like extension of the amnios, rising beyond
the foramen;” and, when we turn in wonder to see what the amnios in plants
could have been, learning that it is ‘‘a viscous fluid which surrounds certain
ovules at an early stage.”” We do not need often to know that prosphyses were
“abortive pistillidia of the muscal alliance,” and the youngster who has occasion
to look for the word should learn that both it and its definition are mere sur-
vivals from a past century.
Second, space could be gained by omitting to define common words which
have no technical meaning, such as congeries, enlargement, entangled, evapora-
« JACKSON, BENJAMIN Daypon, A glossary of botanic terms, with their deriva-
tion and accent. Second edition, revised and enlarged. 12mo. pp. 37!- London:
Duckwith & Co. (Philadelphia: J. B. Lippincott Co.) 1905.
2 Bot. GAZETTE 31: 68. 1901.
448
See
1906] CURRENT LITERATURE 449
tion, minute, parallel, sex, tall, wound. All of these and many others are now
include
Third, more care should be taken to make definitions sufficiently general
to include the various uses of the word, rather than so special as to refer only
to particular uses. Thus, conjugate appears as an adjective, but not as a verb;
conjugating tubes are defined in a special and unusual sense for the Rhodophyceae
and not at all for the Conjugatae; for pistil is given (after a wholly erroneous
definition in reference to spermatophytes) an obsolete sense which is restricted
to the genus Andreaea, when in the same sense it was formerly applied to the
archegonia of all mosses; retardation is not mentioned as other than the “‘influ-
ence of light on growth in certain structures;” and a fat enzyme is defined merely
as an enzyme ‘‘converting olein into oleic acid and glycerin.”
Fourth, greater aia is sadly ar A few examples will illustrate
this: Galvanotro pic, A istics, etc.;” geotropism, “‘the force of gravity as
shown By curvature;” geotaxis, Secveinias in plants caused by gravity;”
stamen, “‘a male sporophyll;” pistil, “the female organ of the flower;” stamz-
nate, ‘‘applied to flowers which are wholly male;” oogenesis, “the formation
of the oosphere, the early stage of the ovule”’ (but oosphere is correctly defined
later in the same paragraph!); sap-pressure, “the force exerted on passing
upwards through the tissues;” spermatogenesis, ‘the development of the male
elements, antherozoids, pollen-grains, and analogous bodies;”’ and so o
Fifth (a matter for the publisher), the use of a more flexible paper ee looser
binding would contribute much to the handiness of the volume.—C. R. B.
MINOR NOTICES.
The dynamics of living matter.s—In the spring of 1902 Professor JACQUES
LoEB was invited to deliver a series of lectures at Columbia University. In
these lectures, eight in number, he presented the gist of his researches upon
the dynamics of living matter. This book, forming the eighth volume of the
Columbia University Biological Series, is a somewhat more complete survey
of the field of experimental biology, says the author, than was possible in the
lectures. In ten “‘lectures’’ he discusses the general chemistry and physical
constitution of living matter, certain physical manifestations of life, the réle
of electrolytes, effects of radiant energy, heliotropism and other tropisms, fer-
tilization, heredity, and regeneration.
Through the publication of his collected papers in English in the Decennial
Publications of the University of Chicago+ Professor Lors’s point of view and
the general results of his experimentation have become even more generally
3 Logs, J., The dynamics of living matter. Columbia University Biological
Serizs VIII. 8vo. pp- xiit+233. figs. 64. New York: The Columbia University
Press. 1906. $3
4 Logs, J., Studies in general physiology, 1905.
450 BOTANICAL GAZETTE [JUNE
known than from the originals. The topics named above are naturally those with
which the author has chiefly concerned himself, and it cannot be said that the
present volume contributes to general physiological literature anything new.
The book is rather a new setting of the brilliant work and suggestive ideas of
the author, that have previously enriched physiology, and with them is related
the results of others in such wise as to round out the presentation. The lectures
are readable and instructive, and they are especially commended to the attention
of plant physiologists, who are too apt to pass over literature not strictly per-
taining to plants.—C. R. B
The problems of life——The third part of this book’ was issued last winter,
and extends the author’s fundamental hypothesis to the phemonena of fertili-
zation and heredity. To him, if one admits the premises, the difficulties of
these phenomena fade away like morning mists. The work does not cite defi-
nite observations, nor show, except in the most general way, how the known
facts can be correlated by this theory; but it presents a clearly reasoned, logical
series of deductions, which impresses the reader at once as too simple to be true.
Moreover, one is naturally shy of a theory, which, beginning with an assump-
tion regarding the molecular structure of protoplasm and the nature of assimi-
lation, makes reproduction a necessary and inevitable consequence of these
assumptions, while heredity likewise follows as a matter of course from the
phenomena of fecundation. We were inclined to welcome the molecular con-
ceptions of the first part,° as possibly embodying a fruitful theory, but we can-
not follow the author as he widens and heightens his construction upon the
acute fundamental assumption. Such inverted pyramids of logic can have no
stability.—C. R. B
Pfeffer’s Physiology —The third and last volume of this work was pub-
lished about the middle of March.? It treats at length of the movements of
plants, including the mechanical responses to various stimuli; and briefly of
the production of heat, light, and electric tensions, and of the sources and trans-
formations of energy. The translation, or rather the interpretation of the
original, is of the same satisfactory character as in earlier volumes. As before,
the editor has introduced supplementary and critical matter in footnotes; and
in an appendix of eight pages he has supplied some important facts not men-
tioned in the first two volumes, and a summary of the more recent literature,
especially that connected with the present volume. Throughout, his critical
5 GIGLIo-Tos, ERMANNO, Les problémes de la vie. IIJ¢ partie: La fécondation
et hérédité. 8vo. pp. vili+189. Cagliari: The author, at the University. 1905. /r. 8-
© Cf. Bot. GAZETTE 31:275. 1901.
7 PFEFFER, W., The physiology of plants, a treatise upon the metabolism and
and sources of energy in plants. Second fully revised edition; translated and edited
by ALFRED J. Ewart. Volume III. Imp. 8vo. pp. viii+4sr. figs. 70. Oxford: The
Clarendon Press. 1906. ats.
1906] CURRENT LITERATURE 451
care and acumen have enriched the already valuable work of the author, so
that English readers are indebted to him for far more than a translation of pecul-
iarly difficult German. To recommend the English form to all libraries and
laboratories as a standard work of reference is, at this date, really quite super-
fluous.—C. R. B
British flowering plants.—Under this title Lord Avepury,® better known as
Sir Jonn Lussock, has brought together a mass of desultory notes on various
things connected with a great many plants. The author says that this work
is “to describe points of interest in the life-history of our British plants; to
explain, as far as possible, the reasons for the structure, form, and color; ane
to suggest some of the innumerable problems which still remain for solution.’
A glossary and an introductory chapter indicate that the book may be used by
those with no botanical training; and perhaps it will be chiefly so used. Each
species is taken as the occasion for the presentation of all sorts of facts and fancies
and questions in reference to it, as though the author had emptied his note book
under that head. There is no distinct organization and no pam ae index;
so that the botanist will simply have to ‘‘run on” to things.—J.
Spring flora of Ohio.—Under the title ‘Spring Flora,” the botanical staff
of Ohio State University has issued a manual for beginners and amateurs.° It
is a revised edition of KELLERMAN’S “Spring Flora of Ohio,” and its range
as been extended so as to include Ohio and Indiana and the adjacent states.
The time range extends from the opening of the season into the first part of
June; and such difficult groups as grasses and sedges are not included. There
is also a key to the trees and shrubs based on leaf and twig characters.—J. M. C.
Flora of Norway.—AxeEL Buiyrtt’s completed Handbook of the Norwegian
Flora, including the vascular plants, has been issued under the peak of
AHL.'° In reality it has been in preparation since 1861, having
begun by the father, continued by the son, whose name is on the title page, <P
now finally edited by a third botanist. It is a model of compact and clear print-
ing, excellent arrangement, and good text figures. The sequence is that of Engler
and Prantl.—J. M. C
8 AveBURY, THE Ricut Hon. Lorp, Notes on the life history of British flowering
Plants. 8vo. pp. xxiiit+4so. figs. 352. New York: The Macmillan Company.
1905. $5.00.
9 Ketrerman, W. A., GLEASON, H. A., and ScHAFFNER, J. H., Spring flora
for beginners and amateurs. pp. xiii+188. Columbus, Ohio: Geo. W. Tooill. 1906.
75 cents.
*° Biytr, Axet, Haandbog i ee Flora. Efter forfatterens dod afsluttet og
udgivet ved ones AHL. pp. xi+78o. figs. 661. Kristiania: Alb. Cammermeyers
Forlag. 1906.
452 BOTANICAL GAZETTE [JUNE
Portraits of botanists —In 1903 WiTrRock published a set of photographs
of botanists selected from the collection at the botanical garden at Stockholm.
A second series has now been issued,'' containing full-page portraits of too bot-
_ anists arranged chronologically from Aristotle to Goebel; and 51 additional
plates, each containing 6 portraits. The biographic notes contain a large amount
of information which must have been brought together with great labor.—J. M. C.
British Desmidiaceae.—In 1904 the first volume of this work was issued
as a publication of the Ray Society. The second volume has now appeared,*?
containing the genera Euastrum (46), Micrasterias (18), and Cosmarium (50).
NOTES FOR STUDENTS.
Regeneration.—The number of recent papers dealing with regeneration
indicates a marked activity in this field of investigation. The work of [RMISCH
and others has made us familiar with the fact that the hypocotyls of a number
of plants can produce adventitious buds. “In some cases these occur normally,
but in others only in the presence of more unusual conditions of growth. BURNS
and HEDDEN"S have investigated these conditions, using seedlings of Linaria
bipartita splendida, Antirrhinum majus, and Linum usitatissimum. They
confirm KisTER’s results that when the cotyledon or the main vegetative tip
is cut away the tendency toward the development of adventitious buds is greatly
increased. On uninjured seedlings of Antirrhinum which do not stand erect
but are horizontal, buds arise only on the upper side, and when these plants
are fastened so that they must remain erect they produce no buds. The effect
of a moist atmosphere is to increase the number of buds and the rapidity of
their development. The same is true of higher temperature. The older parts
of the hypocotyl have a much greater capacity to produce buds than the younger
parts, and there is no tendency at all to bud production on the part of the hypo-
cotyl still elongating. Gravity seems to have no influence. Light, on the other
hand, is a necessary condition, for in one-sided illumination buds appear only
on the illuminated side, on a klinostat in the light on all sides equally, and in
the dark not at all. Experiments are mentioned which indicate that wounding
is not a cause of the regeneration here. The explanation of the phenomena
mentioned as given by the authors is that “‘when the cotyledons are removed
11 WITTROCK, VEIT BRecHER, Catalogus illustratus iconothecae botanicae horti
Bergiani ee ee notulis biographicis adjectis. Acta Hort. Berg. 3: No.3.
p. xcili+245. pls. 151.
*
2 West, W., and 2 ee A monograph of the British Desmidiaceae. Vol. I.
pp. X+206. pls. 32. London: Ray Society. 1
‘3 BurNS, GEORGE P., and HEDDE Pia ., Conditions influencing regenera-
tion of the hypocotyl. Beih. Bot. Coed 19: Suse 1906
1906] CURRENT LITERATURE 453
or cease to function, their work is taken up by the epidermis. The cells of this
develop a vast amount of chlorophyll and all movement is to and from them.”’
“Only those cells exposed to light function as cotyledons, and hence all flow
of material is to and from the lightest side. Light is then an indirect cause
of the location of the buds, while the principal factor is determining the loca-
tion in relation to movements of food materials in plants.” This would make
it entirely a question of nutrition, a rather unusual condition, for in most
cases of regeneration in plants, and in animals too, regeneration will occur
while the parts concerned are but poorly nourishe
Ficpor" cut off the apices of young fern fronds piepenciglir to the median
axis and very close to the tip, removing only a fraction of a millimeter of the tip.
Replacement occurs slowly, but the new tips become forked, two apical cells
forming, one on each side of the midrib. The two sides extend outwards, leaving
the midrib sunken in the center. By cutting very young fronds with a median
longitudinal cut about 5™™ deep, regeneration of each half occurs, and a sub-
sequent branching of the frond is obtained. The fern used was Scolopendrium
Scolopendrium, a variety of which (daedalea) occasionally occurs in nature with
forked fronds, and Frcpor considers this probably due to wounding of the tips
by insects and subsequent regeneration.
HILDEBRAND'S has continued his studies on regeneration in Cyclamen,
and presents further interesting observations. Two forms are mentioned,
Cyclamen Miliarakisii and C. creticum. On the former, when the leaf blade
of the cotyledon is removed, leaving the petiole, there arise a little below the
place of removal, froma point on one side of the petiole, four small leaves, each
having the form of the cotyledon, and the four together aggregating the size
of the blade removed. Each is borne on a distinct petiole of sufficient length
to bring the blades out far enough to prevent shading each other. In this the
author sees an exceptional example of the principle of utility in the development
of plant structures. In the other species, C. creticum, HILDEBRAND observe
a plant having no cotyledons, but upon which, arising from the center of the
tuber, were three leaves with long petioles. Each blade was almost one- -third
the size of the round cotyledon-blade, and in form intermediate between the
cotyledon -and the foliage leaves. Investigation showed that the cotyledon
had been destroyed to the base, and these three leaves arose together from
the axis of the plant just below the point of attachment of the cotyledon, The
originated as entirely new structures, replacing cotyledons, and were intermediate
in form between these and the later leaves.
4 Ficpor, W., Ueber bcaswageres der Blattspreite bei Scolopendrium Scol-
opendrium, Bes Deutsch. sell. 24:13-16. 1906.
ts HILDEBRAND, FRIEDRICH, Ueber eine eigentiimliche Ersatzbildung an einem
Keimling yon Cyclamen Miliarakis:i und einem anderen von — creticum
Ber. Deutsch. Bot. Gesell. 24: 39-43- 1906.
454 BOTANICAL GAZETTE [JUNE
SETCHELL’® gives an account of regeneration among kelps. He distin-
guishes between physiological and restorative regeneration, applying the terms
in the same sense as used by Morcan. In physiological regeneration he notes
two kinds, continuous and periodic. In the former the continuous growth
of the meristematic tissue at the base of the blade keeps pace with the constant
breaking off at the tip due to wave action, and so the blade retains a constant
length. In other species this growth is periodic, occurring in the spring and
inthe autumn. The growth of a new blade lifts the old one from the top of the
stipe and it is rapidly eroded, the new one thus taking its place. Restorative
regeneration involves the development of new branches and occurs as a result
of wounding. If the stipe is broken off a new blade is formed at its apex. Wounds
along the surface of the stipe result in new blades arising at the points. A ver-
tical wound at the tip results in a splitting of the blade and the appearance of
forking. The observations are followed by a discussion in which the author
contends that the phenomena of regeneration are to be explained best by the
assumption of a flow of materials toward the parts concerned. He does not
consider it necessary to assume a special organ-forming material, the impor-
tant thing being the control of the flow of already organized food materials.
This control of the food substances is due to certain cells being able to exert
a stronger ‘‘pull” upon them than others.
As this idea is so commonly used in explanation of regeneration, the reviewer
cannot forbear remarking that it removes one difficulty only to incur a greater
one. Soluble food materials, in common with all other diffusible solutions
in plants, move toward the region of least concentration, and if there is a more
rap:d flow of substances toward any region, it indicates that these are being
taken out of solution there, either by being used or otherwise transformed. The
more active the use, the lower will be the concentration, and the more active the
flow will tend to be toward that point. The increased activity of the cells,
either in using up by growth or otherwise transforming the food substances,
must precede any special flow (that is, apart from a general diffusion in all direc-
tions) of these substances into any particular region. The movement, or, if pre-
ferred, the ‘‘flow” of soluble substances (other than a general diffusion) toward
special cells is necessarily a result and not the cause of their activity. .
MirHe”’ has used an interesting method of studying the behavior of isolated
cells, especially in their relation to polarity. When a tissue is plasmolyzed,
the continuity of the protoplasm is broken and the protoplasts become separated
from one another. In this way a plant may be divided into its individual cells,
and the behavior of these, each acting independently, can be studied. MreHE
used this method on a marine Cladophora. The plants were plasmolyzed in
16 SETCHELL, WILLIAM le Regeneration among kelps. Univ. Calif. Publ.
Bot. 2:139-168. pls. 15-17.
17 MIEHE, Huco, Nia. Regeneron und Polaritit isolierter Zellen. Ber.
Deutsch. hon Gesell. 23:257-264. pl. 4. 1905.
*
1906] CURRENT LITERATURE 455
a strong salt solution (16.2%), and then transferred gradually to normal sé
water. In nearly all cases the protoplasts regained almost entirely their orleicisl
size, a few remaining in the plasmolyzed form. An active growth promptly
set in, by which the form of the alga was entirely changed. First the proto-
plasm of the last-mentioned cells, by means of rounded or tube-like outgrowth,
finally filled up the original space within the cell walls. Then all the cells grew
in this way: the basal end of the cell pushed into the cell below in the form of
tubes, often growing between the protoplasm and the cell wall; or occasionally
the whole cell bulged into its neighbor. hen one cell is dead, the next above
grows in and fills it completely. Often from the lower angles of the cells tubes
grow downward into the cells below. Many of these tubes assume the char-
acter of rhizoids. All of these outgrowths occurred at the basal end of the cell,
not a single one from the apical end. Later, unless the upper cells begin to
produce branches, they do so rs aes from the apical end. A. very Te
polarity of the cell is thus se
Some interesting results on eee and organ- Se ectuein on Caulerpa ‘ie
lifera have been contributed by JANSE.’® This plant he shows to possess a well-
marked polarity in the formation of “leaves” and rhizoids, and also in the
streaming of the protoplasm, which is always from the apex toward the base.
Following a wound there appears to be a division in the protoplasm, the chlo-
tophyll-bearing portion separating from a colorless turbid portion. It is the
latter, according to JANsE, that occasions the formation of new organs. The
polar phenomena he considers dependent upon a flow of energy in which
the force acts always in the direction of the base. This stream of energy he
calls the “‘basipetal impulse.’”? The opposite, an acropetal impulse, was not
to be detected, and JANSE concludes that ‘the lack of an ‘acropetal impulse,
implies the lack of a second pole at the organic tip.”? Thus we have a polarity
with only one pole. The author applies this conception to polarity as seen
in the higher plants. The point of view is more interesting than convincing.
OBLER’® uses some observations on Polysiphonia and Ceramium as the
basis of a lengthy and rather elusive discussion on regeneration and polarity.
He sees a difference in the lower and higher plants in respect to polarity, which
he considers rests on the differentiation of tissues, and accompanies the division
of labor in the plant—W. B. MacCatium.
Roots of Monocotyledons.—LINDLINGER?° has reopened the question of
the place of origin of the secondary growth shown by the roots of some mono-
18 JANSE, M. J., Polaritat und Organbildung bei Cawlerpa prolijera. Jahrb.
Wiss. Bot. 42: 394-460. pls. g-II. 1906.
‘9 TOBLER, FR., Ueber Regeneration und Polaritat sowie verwandte Wachstums-
vorgange bei { Polysiphona und andern Algen. Jahrb. Wiss. Bot. 42: 461-502. pis.
12-14. 1906
20 LINDLINGER, L., Zur Anatomie und Biologie der Monocotylenwurzeln. Beih.
Bot. Cent. 19: 321-358. 1905
456 BOTANICAL GAZETTE [JUNE
cotyledons, such as Dracaena. Contrary to the usually accepted view, the
author finds that cambial activity is present not in the pericycle but in the inner
layers of the cortex. Cell divisions in the pericycle are confined to the points
where lateral roots make their appearance, and this growth may have been
confused by other investigators with the true secondary growth in the cortex.
The second part of the paper is devoted to an account of the so-called ‘‘ Aussen-
scheide,’’ a zone of more or less Haaiies cells found in the inner cortex of
many monocotyledonous roots. This zone is not a real secondary tissue, 7. ¢.,
produced by the division of eek ia though it may assume the power
of secondary growth. This zone is considered to be equivalent to the secondary
tissues discussed in the first part of the paper. Several suggestions are offered
as to the function of the “Aussenscheide,”” varying with the habits of the plant
in which it occurs. Naturally the mechanical function seems to be the most
common one.—M. A. CHRYSLER.
. Morphology of Cucumis sativus.—TILLMAN?! has investigated sporogenesis
and embryogeny in the cucumber. The most interesting items reported are
as follows: the presence of two integuments that elongate greatly and invest
the remarkable beak-like prolongation of the nucellus; the somewhat irregular
development of the embryo; and an haustorial enlargement of the pollen tube
on its passage through the long nucellar beak. e fusion of the unequal polar
nuclei was seen, but no case of double fertilization was observed —J. M. C
Enzymes of Polyporus.—Butirr’? finds in the juice of P. squamosus the
following: laccase, tyrosinase, amylase, emulsin, a protease, lipase, rennetase»
and coagulase; but negative results were obtained by tests for pectase, maltase’
invertase, trehalase, and cytase. Yet the fact that it destroys the wood of Acer
pseudoplatanus indicates the presence of cytase and possibly hadromalase.—
Limiting factors——An illuminating paper on Optima and limiting factors
has been published by Dr. F. F. BLackMan,?3 which it behooves every physiol-
ogist toread. The argument shows, and it is sustained by the results of research,
that much physiological experimentation has been falsely interpreted —C. R. B.
Photosynthesis extra vitam.—Maccuiatr replies?4 to BERNARD,?5 criticiz-
ing his methods, maintaining that photosynthesis does occur in vitro, and stating
certain modifications of the process. He makes a weak case.—C. R. B
21 TILLMAN, Opa I., The embryo sac and embryo of Cucumis sativus. Ohio
Nat. 6: 423-430. pls. 29-30. 1906
22 BULLER, A. H. R., The enzymes of Polyporus squamosus Huds. Annals of
Bot. 20:50-59. 1906.
23 Annals of Botany 19: 281-295. 1905.
24 Maccaiati, L., Altri fatti e nuovi argomenti sull’ assimilazione fotosintetica
fuori dell’ organismo ees le richerche del dig. Dr. Ch. Bernard. Nuovo Giorn.
Ital. 12: 461-4 1905.
25 Bot. GAZETTE 41:157. 1906.
f
| i ee aa es i ia aa la
NEWS.
Dr. Jutius Wiesner, the well-known plant physiologist of the University
of Vienna, has been made a life member of the upper house of the Austrian
parliament.
Dr. Brapitey M. Davis has been spending the spring in Cambridge com-
pleting a textbook of botany in co-authorship with Mr. JosepH Y. BERGEN.
His connection with the University of Chicago will end July 1. He will be at
Woods Hole through the summer.
IN vIEw of the great service rendered by Dr. J. Brrquet at the Vienna Con-
ess, an international demonstration in his honor was arranged. The fund
obtained from fourteen different countries have been used in the purchase ae a
gold watch with congratulatory inscription, a check for 2000 francs for the further-
ance of Dr. BRIQUET’s scientific work, and an illuminated address. The details
of this movement have been published and also Dr. Briquet’s letter of thanks.
THE THIRD annual meeting of the Botanical Symposium will be held from
July 2 to 9, 1906, at Mountain Lodge, Little Moose Lake, Old Forge, N. Y.
Through the courtesy of the members of the Adirondack League Club the privi-
lege of occupying the Club House for one week is extended to the members of
the Symposium. Botanists are requested to notify Mr. Joseph Crawford, Secre-
tary, 2824 Frankford Avenue, Philadelphia, Pa., if they intend to attend the
Symposium.
A NEW JOURNAL bearing the title Annales de Biologie Lacustre is to be pub-
lished under the editorship of Dr. Ernest Rousseau, with the collaboration of a
very large board of editors. ‘The first fascicle as announced contains 192 ps ges
with figures and maps. Publication is to be in German, English, French, and
Italian. Each volume will contain 400 to 500 pages, and the subscription price
will be 20 to 30 francs. The address of the editor is Musée royal d’ Histoire
Naturelle, rue Vautier, 31, 4 Bruxelles.
A MOVEMENT is on foot to erect in Jena a statue as a memorial to Professor
ERNst ABBE, who died last year. The American Microscopical Society has
issued a circular letter appealing to its members to aid in this movement. The
Bausch & Lomb Optical Company of Rochester, N. Y., which has long had
business relations with the Zeiss works, has also sent out letters asking for con-
tributions from those who are not members of the society. Contributions in any
amount will be welcomed and will be acknowledged.
A FASCICLE oF Kew Buttetins has appeared recently, bearing various dates
from 1920 to 1906. This is an attempt to revive a dormant publication suffi-
457
458 BOTANICAL GAZETTE [JUNE .
ciently to permit the annual volumes to be bound. Heretofore these volumes
have been represented by the annual appendices, which led to the current gibe
that the Bulletin had. succumbed to appendicitis. The most curious illustration
of “closing up ranks” is the volume for 1900, the body of which consists of 32
pages, now issued as nos. 157-168, and which were necessary as a preface to
the four appendices.
Proressor W. A. KELLERMAN recently returned from his second collecting
trip to Guatemala. On account of quarantine regulations (because of yellow
fever) he was obliged to return three weeks before the time set. The part
traversed the entire country from east to west and went up as far as Quetzal-
tenango (alt. 2500™). Collections were made about Lake Amatitlan and also at
the still more beautiful Lake Atitlan, and on the ascent of three volcanoes. Per-
haps ten times as many species of parasitic fungi were gathered as in the same
time last year, and the collections seem to contain many new species.
Dr. F. Cavara reports as reasonably successful the attempts to establish
an alpine garden on the slopes of Mt. Etna. It is located behind the Casa Can-
toniera at an altitude of 1880™, the first cultures at 1440™ having failed on account
of the heat and drought. About 150 species are now thoroughly established,
and nearly 4oo more are more or less sucessfully grown. The garden is sur-
rounded by a stone wall which mitigates the violence of the winds. _Cisterns and
snow magazines (there are no streams) eke out the scanty supply of rain in the
growing season, which in 1904 was 56™™ in May, June, July, and August. The
director is to be congratulated on overcoming the many difficulties and solving
so many of the problems which confront him in this undertaking. The garden
has been christened Gussonea, in honor of “un valoroso studioso della flora
sicula,”
GENERAL INDEX.
The most important classified entries will be found under Contributors, Personals,
and Reviews. New names and nam
in bold face type; synonyms in he.
A
Abbe, Ernst, statue to 457
Absorptio on of mi by leaves 262
Abutilon, “cig ee m 361; striatum 361;
Thomso
Acacia saieeice absorption of water
278
Ac as, cers by roots 367
Acrolasia 356
Aether 356
Adaptation, anne 305
Aerotropism
Agaricus campestris 350
Agro opyron caninum, nodes of 4
m 150
wn pigment of 79; iron 225;
of northern seas 367
Allionia 151
‘teers, Cepa
Alternation of Sscimaasienee 222; in Phaeo-
eae 364
:
oo as of specie of
Andromeda politi 19; of Clayton
306; ecological, of bog plan 17
et argenteum, aia | in 11;
oe — asal ee sarge 10; sco-
pari mphivasal bundles
haste de Biologie Lacustre, a new jour-
nal 457
Antennaria 356; neodioica 149
pocynum 356
Apothecia of lichens 306
Aphyllon 356
Apple, pe canker 366; rot 223; and
rot
arineae 221
6
He eve ae on Chloranthus 368
Artemisia 150; variabilis 327
Arthur, J: C., 155, 217, 301, 356; per
sonal 160, 307; on AOE ie a 157
of new genera, species, and varieties are printed
Arundo Donax, cambium in 12; nodes
co)
Ascomycetes, nuclear reongs in 305
-Ascophanus, streaming in 217
pelea aha niger go
enium 356
ie ag internationale des botanistes
Ast 150
Astragalus 356
telophragma 35
pee George F., personal 307
Avebury, Lord, “Lite history of British
flowering plants’
Avena, barbata 24; ee 4; sterilis, 12
Bacterial diseases 214
3ailey, W. Whitman, personal 307
3alanopsidaceae 356
56
u, Le, a new journal 308
awe -Petrucci, G., on nucleoli in
mitosis
a. Sassafras 434
Barnes, C. R., 147, 148, 149, 153, 157,
ae 220, 221, 222, 224, 225, 300, 305,
306, 368, 370, 448, 449, 450, 456
Basidium of Amanita bisporigera 348
Bast in io fras 439
Bateson, E., on heterostylism 304
Baur, on chlorosis 61
Beauvar.l, o
Bell, J. M., on soil waters 305
Bennettites 79
Berberis 149
Bergen, J Y5 409,302
Bernard, C., on photosynthesis 158
Bessey, Ernst A., perso = 80
Betula, my: corhiza in
Biological _preceetas “Cota Spring Har-
bor 372; Ohio State nigrcaengg 27:
University of Minnesota i
or Ae of Washington 372; Woods Hole
Send teed Sad Sod eed
Ww
i]
227
Biondia 150
459
460
Black rot of cabbage 306
Blakeslee, A. F., personal 80, 371
“ ckman F. F., on Optima and limit-
ing factors 456; on photosynthesis 215
Blight canker 366
Blinn, P. K., on rust-resistant cantaloup
., “‘Haandbog i Norges Flora”
Bogs and bog flora 17
Bolscuralis 356
oletus 150
Borgen, Shi vegetation of Faeréese
oasts”’
Bor rzi, A., on Zoddaea 357
leria 1
Botanical S apeceiita. Sites annual meet-
in
Botrytis vulgaris 88
Boudier, Emile, personal 159
Boveri on Euglena 230
Brainerd, E., on violets
Brandenburg, rpms: ane of 300
Breazeale, J. F
pe Oskar, personal 80; work of 81
Briquet, J nal 457
a ue
373
eae of Poly-
456
Bulletins of Kew Pae! dens 457
euies ss, E. = ., “ Biotian pores 354
anni
ee we Hedden, on regeneration of
14
Biisgen Re on ® chaiucheuien 82
Supler, G. on grape diseases 367
C
Cabbage, og rot of 306
Calamagrostis Ca apenig 2 amphivasal
bundles py 9; cambium in 12
California, Academy of Seuns, destruc-
tion Sid building 371; new species of
plants 283
C Ramsey in grasses I
Canton F. kK. on soil waters 305
nula, exigua 325; trachelium 359
Canker blight 366
g's on measuring transpira-
tion I 158
Can a Tust-resista’
Cardy work of 149, ey ee
Carduus
Carey, Bae B., personal 227
INDEX TO VOLUME XLI
[JUNE
pi 356; Wight
2 a eons: Sans on n 455
ena, » per » 458
ee 151
Cell division in Empusa 229, 243
Celtis pallida, absorption of water 267
Ceramium, Tobler on 455
6
tis 356
Hh calyculata, ecological
chee teatiain cae Jo 148, 221, 223, 225,
226, 306, 364, 3 8, 369; on alteration
of genera ations oy ee 3 Methods i in plant
histology”
Chemotaxis of spermatozoids 76, 226
asada aes fungi 81
ee hel ey ecological anatomy
CHierwnshuns, morphology of 368
Chloroform, a stimulant 158
Chlorophyceae 357
—o are of sun and shade plants
Cigoresia
eee bse on ne erns of Costa Rica 355
Christ , C., “Index Filicum” 148,
355
gree ra nature of 220; in
Zygne
Chr viene ee 158; aberrant
225; in Zygnema
Chrysler, M. A., 1, fee 222, 455
a a avenaceus, amphivasal bun-
dles
Chhrysopss, ee 312; Breweri 292;
gracilis
Cladocephalus 150
Clark, J. F., on chemotropism 84
st on Philippine bea 353
ia Pag ke 5
Ciavarie 3
Claytonia, ater of 306
Cleomella 150
Cnemidophacos 356
Cobb, a ge - sugar cane 365
Coilochi
Coix dactpielis 2953 yore amphi-
vasal bundles 9, 10; in 12:
Cold Spring Harbor Piclogical laboratory
372
Collinsia Hernandezii 310
oa rg of Sargassum 167
Conimitella 35
Ganvans tion, yeasts 157
Connecticut, fungi of 215
Contributors: Arthur, J. C., 155, 217+
301; Barnes, C.
157, 215, 220, 223, 222, 224, 225, 3
305, 300, 368, 370, 448, 449; 45° 456%
1905]
“aes J. Y., 327, 362; Breazeale,
M., 71, 76, 791,740 aa 353,
367; Eastwood, A AL ay.
E., 309; Farmer, . i e pies Hi.
: nong, W. F. i
eer W. B., 73, 452} Marah,
Cy Merriman, M. L., 43;
seta A. vg os Newcombe, F. C., 76;
Olive, E. W102, 2295 Pond, R. jaar
¥50, 917.-219, ae 221, 226, 359, 367;
Schaffner, a H., 183; Shear, C. L.,
160; Shull, G. H. » 301, 302, 303; gee
358, 363; cae E. B., 161; Spal-
ding, V. M., 262; Steve ens, F. a 216,
369; Thiessen, R., 154; acon. E.
jae
INDEX TO VOLUME XLI 461
ea ag Goveniana 326
Cus ns fe on desmids 356
Gone I
Cyclamen, Hildslicnndlon regeneration
Cyclopedia, botanical 76
Cynoglossum, boreale 365; Virginicum
357.
Cystolejeunea 356
Dacryomyces, chrysocomus 349; deli-
quescens 348
D — I51
Dammer, U., work of 356, 3
Danish Arctic Lagan at 28
Darbishire, A. D., on Mendelian law 303
Darwin, F., pace 159
Davis, B. M., 71, 76, 79, 146, 157, 305,
300, 353 307
ary, A., work of 81
Deisiasin 150, 356
Denmark, lakes 360; shore formations 78
aren sia 150
Desert shrubs, a. relations 262
Desmids 350, 3
tmers, F eg personal 372
ai Osw. de eaties de, per-
1 307
; W E. M., 223 sona
365, 306, 3075 Yamanou chi, S., Diastase 158
Conzatti, C., géneros cette: Diatoms 360; movement 306
sicanos! 14 Dictyota, periodicity of sex organs 79
Cortana, E. B., “Polypodiaceae and D oo 150
edible fungi of the Philippines” 147 Diels, L., on Chinese flora 150
Coreosma 35 Dietel, P, on Japanese Uredineae 149
orn, an ear of 301 D holcos 356
Cornella 356 Dimorphella 150
Correns, C., on gynodioecism 302; on Di — calycinus 287 -
laws of i hueritencs € 303 Disease, apple rot 223; asparagus 365;
Cortex in Sassafras 437 bacterial 214; blight canker of apple
Cortinarius 150 s 366; chlorosis 361; Freeman on
Coulter, J. M., 353, 354, 355, 362, 368, aoe grape 367; pear rot 223; potato
451, 456; personal 371 364; sugar cane 365
Covillea tridentata, absorption of water Domin, work of 149
473 Ducts, ‘intercellular a
Cowles, H. C., Duggar, rsonal 307
Durand, Th., pe ersonal 307
77, 78
Crataegus 151; ere 358
Crone, ee der, death of 372 E
Croomi nam ora, “pagina bundles 8
C Earle, F. S., personal 1
Piatiierns idioblasts of 221; nectaries Eastw twood, Alic ce, 283; ioe 371
° Eatonia 150
Crunocallis Smee SERRA spinosa 327
Cryptogamic flora of a age — 00 Ecological survey 222
Gevitesa = of Sargas oat ae orgy and photdeynthesis 225
Ctenophyllum 356 Ellis, J. B., death of 307
Cucumis sativus, Tillman on morphology Elmer, A. D. E., 309
456 Elmera :
53 ‘
Cupressi Elymus 151; Americanus, nodes of 4
Embryology of Riccia 109
T§2
Simian 151
462
Embryo of Symplocarpus 369
Empusa, Aphidis 196; Culicis 203; mor-
pectoey and oe opment of 19
re
€ 149; “Natiir-
. lichen Pflanzenfamilien” 355; ‘‘Pflan-
zenreich”’ 1 ay
Entomophtho
woe
Delpinia
nophthoreae, cytaligionl ahs of
al 7°
rmis oF fax in Sassafras 435
uisetum arvense 369; chemotaxis of
sperms 226
anthus Ra avennae, cambium i
Etpeens she rogen 291; deokriines
290; miser 291; pygmaeus 291
Eriksson, tok on grain rust 155, 301
rlogonum 150, 356
Eriopho orum Vint ecological anat-
18
Eriophytium Greenei
Ernst, A., on greening of seeds 305
Erocallis -
Errera, Léo, personal 307; biography ie
372; on glycogen 379; on inhibito
acti ion 221; work o
mM 327
n 230; Keuten on 230
s Androsaceus 324
Speers Paisiias 327; terracina 327
oo Pag Ae on Sy mbes potatoes 364
geo n Hepaticae of Puert
ari
Pasa transpiration of 362
Ewart, A. J+ Pfeffer’s “Physiology” 450
F
Falck, R., on zygote formation 85
Farlow, W. G., “Index of fungi” 75;
rson 160
Fernald, M. L., 149, 356
Ferns 35
Fertilization in ae oa 428
150, arundinacea, amphi-
- 9
, transpiration. of 363
Figdor on regeneration {53
ink, Bruce, personal 160, is
Fischer, Walter, es
Fleischer 150, 356
Fliche, P., (and Zeiller) 0 a fossil gymno-
Sperm:
INDEX TO VOLUME XLI
[JUNE
sae: and Sylva 37?
oods, a
Ford, Sib on Araucarineae
Ly cuir pa seein absorption of wail
FE ee A., on haustoria of Osyris 370
i)
Freeman » personal 160; ‘‘ Minne-
sota plant diseases” 72
Freer, Paul C., personal 22
Freezing of buds and twigs 384
Fritillaria succulenta 311
Frith and Schré si ““Swiss moors” 144
Fulton, H. R..,
Fungi, heiotronstun of 81; of Connec-
ticut 215; edible 147; index of 75; pa-
rasitic 77
Fusarium Solani 77
G
Gager, C. S., personal 372
Gaidukov, N., on iron algae 225
Gallaud on ipeorhia 153
bat
-on Kine vines I
ae
Geu
Gibson, R. J. H., on scales of aquatic
monocotyledons, 156
Giglio-Tos, E., “Les problémes de la
vie” 450
Gilg, E., ‘‘Pharmacognosie” 355
Gilia 150, 356
Giraldiella 15
Glaucium flavum 327
Glaux 356
Gleason, H. A., (Kellerman and Schaff-
er) “Spring flora” 451
n es
Gloeosporium nervisequum 78
oe and Parag govern 370
a Veneta 78
~
.
( sadeti tia ‘edn 3
ae 356
Gothan, W., on fossil gymnosperms 151
Graftchybrids ape
G
G
367
7Tasses, of pa 215; nodes of 1; North
Ti 4
Greene, E.'L., 356
Greenm esse M., personal 307
Gregory, R. P., on heterostylism 304
Growth of scaly buds 37
Guilliermond, M. A., on a of
yeasts 157, on nuclear division 30°
sab ied ag . R. von, on array nse
organs 22
Gymnosperms, fossil 151
Gynodiorcism 3092
906]
H
Hackel, i ,on Aenea ee Sirti 354
Hardin ack rot 306;
on past tubers i
Harper, ot Pye ee roauction in mil-
dew:
Pe John W., personal 80, 227
Hartigiella 150
or nae Bh, 94; FS. 77) BGA THO, 57,
361, 36 ]
Haustoria of Osyris 370
Heller, A. “9 150, 356
Henderson, L. F., on Sood scab 304
Hennings, pe pe Tso 1 300
Hepaticae 356; of Fra rance 148
Heredity
Herpetineuron 149
oe . W. C. R., “Lichens of Santa
Cru
nnd 14
tl selene 356
Hesperochloa
Hecarostyiy in Primula 304
Hieronymus 150
Hildebrand on regeneration 453
bp E..J.,.4
Histology, ‘methods i in Leg 74
Hitch fa Ss 64,
Hoé von, on ‘hon gi 356
Holacantha Emoryi, See ek of water
Ho “4 k, A., personal 228
Hollick, ecnge oe al 372
n anatomy of Claytonia
3
nderi marinensis 321;
campestris 286; mollis 286
House, H. D., 334
owe, M. A, 150; personal 372
Huron river valley, bogs of 17
35
, Burns and Hedden on re-
generation of 452
I
Ice in buds 384
Idioblasts, of Cruciferae 221
Index Filicum 148, 355
Leonia n,
J
, “Glossary of botanic
terms E .
INDEX TO VOLUME XLI
463
Jackson, D. D., on movement of diatoms
ahn, E., on myxomycetes 366
Janczewski, E.,
Janse uler erpa Pore 455
Japanese vegetatio aah
Jetirey, E. Cs. 132;
Jentic, on po ollen Nn 301
Joff aa on ponies niie ye ducts 306
Johnson, D. S., 372
Tonesiella 356
ournals: Annales de Biologie oer
457; Flora and Sylva 372; Le Bambou
308; Philippine Journal of Snes 228
Jum melle, H., on tuberization 77
ee Z A. oie ase he
and Schaffner) ‘“Spri
Kelps, Setchel on regeneration of. oy
Keuten on Euglena 230
Kew Gardens, visitor 371; bulletins 457
Kidston, Robert, n Sigillaria 155; 0
ris 21
Kihlman, O., on chemotropism 81
eee 5
Klebahn, H., on parasitic fungi 77
Klebs, G., work of 85; on variation 359
Kleeman, "A, on diastase 158
f 85
ny, L., wo
Koeleria
Korni a M., on ne cilanoweaie reduction
158; on germination and radium ema-
nations 2
Kunze, Cuniae: on excretion of acids by
roots 367
Kupffer, K. R., on species 301
L
Laburnum Adami 359
Lacouture, C., ‘“‘Hepaticae of France”
148
oe tari
150
Lakes ‘a Scotland and ema 360
Land, W. J. G., 74, 79, 2
Larix laricina, scslogical nhaiouds 20;
Se Sper in 32
Lasthenia 356
Latham, uM E., on chloroform stimula-
tion 1
Leaves, I of water 262
Ledum ise sie Tog hag ecological anat--
omy 20;
agers onyzoides, » eset a bundles 9;
Leiblin nger, cong ~ oe apes gaps 370
Lenticels in Sassafras
Se 355
Leptosyne Hamiltonii 323
Lewis, C. E., 109, 348
464
Lewis, F. J., on weiss moors 22
Lewton- Brai ain, L., on disease at sugar
cane 36
3
as 149; apothecia of 306; of Santa
Cru
idforss, B., on chemotaxis of sperms 226
Lights tale: at high altitudes 156
Lignier, O., on Bennettites 79
ae aael tigrinum, chromosome reduction
183
Lisanthas 150
Lindau, G., ee 1 300
nger,
ae L., on eer of monocotyle-
dons 455
Pence P., personal 300
Lithophragma 150, 156
Litto ae spermaiophytes of Naples re-
gion
Heats Be. bs.230: —*. 159
ese F. = » 3363 personal 159
ock, R n plant sibiaa 363
Loeb r 4 tas of living matter”
449
— irc ne Avagiiarh in oi Sans
mbium
ne, c 12; nodes
Lotion 356
Longyear, B. O., on apple rot 223
ry; [2-2 per 2
Lotus baleinge e a 32
Lubimenko, M. W., on chloroplasts 219
eerie . or Pale -, on vascular system of
Mat
Lana 3 oe polyphyllus 325
Lycopodium
Lycium Berlandieri, absorption of water
2
Lyon, Florence, 156
Lyon, H. L., on alternation of generations
22
M
MacAlpine, on Uredineae 150
MacCallum, W. B.,.73, 452
MacCaskey, H. D |, pers onal 2
oa — ati, L.,- on amathed extra
i Doweat D. T., personal 159, 372;
on ig
MacMillan, C., personal 227
Macrae, Tilian J., personal 227
inum 226
Marine Biological Laboratory, Cold
Spring Harbor 372; Woods Hole 227;
University of Minnesota 227; Univer-
sity of Washington 37
Marsh, C. D., 360
Massee, G., ae of 84
INDEX TO VOLUME XLI
[JUNE
Matonia pectinata, vascular system of
21
Matthaei, G. L. C., on photosynthesis 215
Matthiola sinuata 32
Max ., personal
aise litora hi marina 327
Medullary in Sassafras 438
Mereschkovsky, C., on chromatophores
20
Meri 150
Merrill, ns D., on Mig leas plants 353
1L
Merrim n, Mabe
Miers germanica 358
emg ieee of 147
Me pe 149
Racrooha s 356
Micro ca aeaisa of eae 219
ee of Lilium
Miehe, H., on polarity of ciated cells 454
Mildews, reproduction 0 14
augh, C personal 371; “ Prat
”
Miscanthus sinensis, cambium i
— Botanical Garten, ae pte
Mitosis, nucleoli in 369
Miyos eas on Japanese vegetation 76;
wor 3
wich J ., ‘Vegetable foods’
Molisch, H., on brown Poor 0 work
of
Monardella franciscana 320
Mon er kg aquatic 156;
Lindlinger :
Montemartin, t on proteid-formation
roots of,
Montia 150, 356
Montgomery, E. G., aa ee oe
Montgomery, T. H., on aberrant chro-
Pisce ag a
., 69; on sporogenesis 67
“Plants of Bermuda” 355
Moore, G. T., personal 2
Moaes Scotch 224; Swiss 144 |
Mosses 140, 355, 3565 —— of
spores 370; sporo te of 15
M oe Ww. sr - on teratology in Salix 368
Movement of diatom
= ae 88;
pt asm 217
rian ve H,
S 3°
streaming of proto-
robryum 150
Mublenbergia debilis 326
Murrill, W. A., personal 372
BS a a 30
357
Myeoplasn and grain rust 155
Myc za 32; endotrophic 153
Meccaphentia a Ulmi 7
Myxomycetes, germination in 366
,
t
’
1906] INDEX TO VOLUME XLI 465
N
Naiocrene 356
Naples: eee cee spermatophytes 327
a i on Nevada plants 150
Ném regeneration 73
Nemoyhila Wremont i 319
ate eutzia 353
raperets -avis, brown pigment 79
Nephrocerpus 356
Newcombe, F. C., 7
Niklewsk, B., on the reserve food of trees
See for maize 370
Noll, F., on re faabrtat: 358
aus — sen 84
Nuclear division, naive $3055 Bia
Nucleus, division of in basidium 348; and
secretion 306
Oaks, distribution and habits of some
ommon 445
Ocrearia 353
pore: State University Lake Laboratory
Olive E. W., 192, 229
Caton Seffer , P., personal 227
Oltmanns, F. , “Mo Leia und Biologie
rt Algen”
Oocyst of Sargassum 175,
Oogenesis in Polysiphonia violacea 428
Optima and limiting factors, Blackman
on 456
Siteerrimeaes 356
ochrysum 356
O arpus 356; Copelandi 288; im-
veneresaa _ acai 317
Osmotropism
Osterhout, Ww. x V., on Colorado plants
Gore ser ed of 370
Oven, E. von, on tomato rot 156
Oxycoccu cus ‘macrocarpus, ecological anat-
omy 18; mycorhiza in 32
‘
Pachyplectron 356
Paddock, W., on apple rot 223
hinese flora 355
Palladin, W., on respiration 223
Pallavicinia, sporogenesis in 67
oe S357,
mel, , ‘Grasses of Iowa
Pani Americ ana, ets asal ee
m 12, nodes 3; nervata,
eee bundles
Panicum 64; - amaclonche m4 Crus-
galli, gprs th 12; demissum 64;
Enslini 64; lacuna 65; mls
65; lancearium oi aa 66;
sr og am undles 9,
mbium in 12; Pancha 66
Pantanelli, aur Led we 159; on me-
chanics of s
eee ‘imitation pe eal 307
ak dens and glycogen 370
Parkinsonia Torreyana, absorption of
r er
Pascher, A., on sexual reproduction of
Stigeoclonium 154
er my stoloniferum, amphivasal bun-
Paullinia 4
longistylum, cambium in 12
Ase laxa 318
Pentstemon 35
Personal: ‘Abbe E., Bek Arthur, Jac
9 3
ee, A. F., 80, 371; Boudier, E., 159;
Brefeld, ; ; rigquet, .J.,..4573
Britton, N. L., 372; Carey, H. B., 227;
Cavara, F., 371, 458; Coulter, J. M.,
71; ;
Léo, 3073 Farlow, W ee, 1605 an
; Fre
man, E. M,, a Free
Greenman, J. M., 307; Gager, C. S.,
372; MHarshberger, J. W. a27;
Hen JP, geo: Holeky: Ay .228:
Hollick, A., 372; Howe, M. A., 372;
Istvanffi, G. de, 80; Johnson, D. S.,
372; Kellerman, W. A., ; Lindner,
, 300; Lindau, G., 300 ingston, B
Lots
a Mil
C., 227; Macrae, L. J., 227; Maxon,
W. es 371; “Milispaigh, C. C. F., 371;
Moore, G. T., 227; W. A.,
R72; ek, G. V., 3723 Meare Es 300;
466
Olsson-Seffer, P., 227; Pantanelli, E.,
]
159; Porsi d, M. P.,, 228; Prain, ;
159; Rendle, A. B. 371; Richter. A.,
159; Rolfs, P. H., 80, 160; Rose, 5
1593 i
: Stickney, M. E.
rd ce E. , 80; Strong, R. P., at
Thiselton-Dyer, W., 1 159; Tracy, S.
M., 160; Transeau, E. N., 372; Un-
dé; 307;
s Wolte, J. J.,227
het is Se 2 es on ecological survey
Penaigela 356
, W., work of 81; “Physiology”
Phacelia acanthominthoides 309; flac-
23
Phacopsis 356
Phaeophyceae, alternation of generations
3
Phalaris, arundinacea, amphivasal bun-
dies ‘Soa = of 5; nervata, nodes of 4
Phellode n Sassafras 443
Phaliogen i in Sassafras 441
Philippine
228; plants
353
Phleospora Ulmi 77
Phieum ‘Ragtenaal re amphivasal bundles to
Photic sense organs 220
Pholiota 150
Photosynthesis: — Sapper Pollacci
a2: exits , Bernar do on 157,
Macchiati on n 456
Photosynthometer 2
Phycomyces te, eee in 217
dong a geen for
a Mari ecological aia 20;
pcbatitan { in
Piceoxylon 1
Pinus, fossil 151; Strobus, ecological
anatomy 20; mycorhiza 32
nusoxylon 15
Pityoxylon 152
i C. V., “North American species
per,
of Festuca” 354
Pizzo zoni, P., on haustoria of Osyris 370
ad
US 327
reeding in pris 363
Platyschkuhria a 356
Poa oailet
Podoc 152
Polarity ivy Tinlatod cells, Miche on 454
Polemonium 150
INDEX TO VOLUME XLI
[JUNE
Pollacci, G., on photosynthesis and elec-
; on Leet plants 368
; maritimum 327
Polypodiaceae 147
Ar atcanrvaceing Tobler on 455; violacea,
life history of 425
Pond, sis O; ae 21g, 220, 221, 226,
359, 397 ; ae es
Populus oe mycorhiza in 32
Porsild, M. P., personal 22
— resistant 369; scab 304; spray-
ing 36
r, R. L., on ecological survey 222
Prain, D., person nal 159
Prenanthella 356
rving plants
nie heterostyly in 304
Prosopis pe ape 4 bits of water 277
Proteids, formation o
Protoplasm, streaming in Mucors 217
Prucha, root ree ‘cmtees
216
Psilocarphus tenuis 292
Pterobry 150
teeter 150
Puglisi, M., on transpiration of ever-
greens 362
Pinnatella 356
Pyrenoids in Zygnema 44
Pyrola 356
Q
Quercus spp., distribution and habits of
445
R
Radium, and germination 217
Radlkofer, work of 149
Ranunculus 150, 151, 356
Raphiolepis japonica, transpiration of 363
i Sv! Sengaon :
Reduc on of chromosomes 158; 1n
Reduction division
Regeneration 73; recent papers
Reinhardt, M. O., on chemotropism A
Renauld, oe on Musci exotici 150
Rendle, A. B., personal 371 ~
’“ Aaand ges Flora”’451; Bor-
gesen’s “Algal vegetation of Faerdese
coasts” 71; ss’s ‘ Biotia:
354; Chamberlain’s “ Methods in plant
histology” 74; Christe ’s “In
Filicum” 148, 355; Con tti’s ‘‘ Los
géner nos’”’ 1473
os vegetales mexican }
Copeland’s “ Polypodiaceae and edible
fungi” 147; Engler’s “ Natiirlichen
1906]
sp agginag seein 355, . Pflanzenreich”
Ewart’s Pfeffer’s “ Physiol-
ogy” 450; Fatlow's "Bibliographical
i A. 75; Freema
144; Giglio-Tos’s “Les proble s de
la vie” 450; Gilg’s ‘Pha cuaoneace”
355; Gleason’s «Sprin g flora” 451;
Harper's “Sexual alegre in Sone
rre’s_ “‘ of
ry
n rm
“Studien iiber die Regenera ary
Oltmann’ s (““Morpho olo. xg ier Biolo ogie
der Pam
iper’s Peace "eg 545
sae aia eee flora” 451; Simons’s
e’s “Germ of mind in plants”
i “Bacteria a relation .
plant disease” 214; Wes s “Mon
graph of British Des sada aceae iis
White’s “Fungi of Connecticut” 215;
inton’s “Veget ble foods” 300;
Wittrock’s “ oe ee icono-
thecae botanicae”’ 452; ’s “Ob-
servations faites au Epuinpedeen” 146
Rham 150
Ribes 150 356, 357; Stanfordii 315
Ribesi
Bischofhii 117;
Say, biology of r10;
a sa 177; crystallina 116; develop-
109; embryology 117; fluitans
re glauca 116; : hirta 116; lutescens
155; perry: 117; minima
117; sexual organs 118; sperm
esis joe 9 ora 22%; ne sata
120; velutina 117
Ricciocarpus natans 117
Pets r, A., personal 159
Ri "HL N., on iw eee plants 354
Rolfs. P. H., personal I
oots, excretion of ony 367; of mono-
ns, Lin ranged n 455; rela-
wth to tops a wheat 139;
Rosén, F., personal 371
Rosendahl, C. O., on embryo of Sym-
plocarpus 369
Rosz, personal 371
Rot, tomato 156
INDEX TO VOLUME XII
l’s “Grass Ses -
467
Rumex 356
Rusby, H. H., personal 372
Rust, — agus 304, 365; tae on
155; ga-
Rydberg ., work of 1
Salix 3565 Rape apo 368; Breweri 323 $i
las sericea, ecological an
ne nny trioiogy 3
Ba i E. S., on endoparasitic adapta-
305; 0 n non-infection by rusts 305
Salts, a keer of 2
Sanicula 356; laciilats 312; serpentina
12
—— ea gu morphological
tudy of 1
ie Cc. . n Crataegus 151
ser apes pocaniok, eal anatomy
I
Sarton, A., on anatomy and cree, 224;
ex erimental anatomy
Sassafras bark 434
cab, potato
Scales, winter fracded of bud 395
ee ops! ee oe 1033 on Teduction divi-
on 22 33 ‘Spring flor 451
Schaudinn on Vaan 23
Schlechter, R., on flora of Now Caledonia
ao
Schmidle, work of 150
Sieket A., 149; personal 227
Schneider, C. K., “‘ Botanical cyclopedia”
6
repens A., on pears: in Mucor 217
Sch T, J. H n idioblasts 221
Save 152
ao etal Figdor on regeneration
Gecaceelie 3.
cotc
g
Settled ey ‘ak 360
Secretion, mechanics of 222; and nucleus
06
Se ae — of 54
Sonnen 356
Sem vum Funkii 3
Senecio 151, sa ele 293; tri-
angula ; gars 327
Sequoia, ‘Rails
rjania 149
Setchell, W. A., on regeneration i in kelps
Sopand. A. C., on Araucarineae 221
Shear, C. L., 160 ; personal 1
Shibata, ae on . motaxis of sperms 76
Shreve, F., 372
Shull, & Ht. “aon ae 303, 357) 3
Sigillarian stems 1 55
358, 363
468 INDEX TO
Silene deflexa 284; Grails — Grayi
5; lacustris 284; Lemmoni 284
pacifica ei Suksdorfii 285; : Wale
5
ae H. G., on algae of northern
S 397
Seas, A. M., France’s ‘‘ Germs of mind
ts” 148
ifs On gone potatoes 364
nail, oe K., work of 3
smi ith, , ““Bacte = diseases”? 214
smith, R £, ‘on asparagus rust 304, 365
smoke, injury by 1 152
e, M., on nitrogen for maize 370
soil, w ries
TS 305
Solanum 356; Gntiteneni 77; tubero-
m
oraucr, P., on injury by smoke
ere halepense, be aaron james 9
Spalding, V. M
Spaulding, P., y eacial 169
Species, definition of is I
Spermatocrst of Sargassum 174
Spermatogenesis in Folpsichobs violacea
Spermatozoids, Speer of 76, 226
Etinetalocs 356
eae a malorum 86
ce a stigma 150
Sphen cpt eris, microsporangia of 219
Spitzberge n, obse a in 146
oa of pa aed
Spor ation of S 370
Sporobolus \ Wright, amphivasal bundles
mbium
: Sporogenesi, in Pallavicinia 67; in Poly-
iphonia 429
S id ophyte, mosses 158
Stenochlaen a 356
Stephani, F., work of 149
Sterigmatocystis aa 86
seeks - dot 369
Stewart, F. C., on spra ying potatoes 364
Stew t, W., on r sistant potatocs 369
S rk 0 of 85
e¢
personal 227
al feprodliction 154
St , C. R., on nucleus and secre-
10 S$ 306
Stomate I in Sassafras 436
shea M. C., on cycadean integument
Stran a poe! ge of Naples 327
Strange, work of 8
ent a iss personal 80; on alterna-
of generations i in igs Heol Fy sei 364
str eptons 356
ong, R. P., 228
roa rae pero ay rapists 286
VOLUME XLI
JUNE
Sugar cane, diseases of 365
Suksdorf, W,, on Washington plants 356
Swingle, W. T., wo
Seusnioed pak ohare a a
Tansley, A. G., on vascular system of
Matonia 218
Tectaria 356
Telangium Scott
Teratology, Polysiphonia, 4325 Salix 368
pers formation in Polysiphonia 429
Thériot, I., on mosses 355
Thiselton Ae W., personal 159
Thiesse
on Araucarineae 221
327
Thysanocarpus, 150, 356
Tillman on morphology of Cucumis
_Sativus 4 456
d Ceramium 455
Tomato r
[rac
aluminum shells for ex-
i, measuring 158;
PA lag 362
réboux, O.,
spores 370
Trees, reserve food of 1
Trichostema rubisepalum 3
Trifo pen renga oF
on germination of moss
, abesgiaa
3353 bile 334; amabile 342; ama-
bile ‘ie wernt: 342; arcuatum Cusickit
bifidum 334; bifidum
Breweri 334; cognatum 345; Covillei
337; denudatum 342; Douglasii 335;
eriocephalum 335; & arpum 342;
gracilentum 334, 342; Grantlanum
340; Greenei 334; Harneyense 335;
Hemsleyi 342; Humboldtit 342; ™m-
volucratu 46, 347; latifolium 337;
longifolium 342; longipes 336; Lozant
342; 433 microcephalum
ie
species 334; oreganum 336; egae
; Palmeri 344; p 340; pau-
iflorum 342; pedunculatum 336; p9-
tosanum 343; reflexum 346; repens
346; rhombeum 346; Rusbyi afroru-
bens 336; Schiedeanum 346;
villiferum — 335;
Wormskjoldii 347
1906]
Trilocularia 356
Tripsacum acutiflorum 297; dactyloides
294, cambium in 12; dactyloides his-
se m 295; fasciculatum 296; Flori-
um 296; lanceola Spee 296,
iatiolium 294; Lemmoni 298;
2973 synopsis of 204
Triticum sativum ee bundles 9;
cambium in 12; nodes
sues R. H., 299; on sporophyte of moss
Tubercle, Toot 216
Twigs in winter 373
Ulearum 149
Underwood, L. M., 150, 356; personal
37
Unguicularia 356
ba eat is 356; amphispores in 157;
Japan
Dineces eek gins 86
Uromycladium 150
V
bie regi corymbosum, ecological anat-
my 295 ecarnber ot ge n 32
Var ariation, experi tal 350
Masculat syste em, of pion 1; of Matonia
218
= eratrum
ihaacuits sinuatum 327
Veronica ae di 288; Cusickii 290
Viola
ras 153; Hungarian Institute 80
Vri go de, — 150; 371, 372
Vuilleoia® a Saari 150
Ww
L. F., personal 371
Ward, H. M., work of 82
Warming, E., on shore formations 78
Water, absorption by leaves 262; soil 305
Weiss, H. F., 434
Villens A, on nectaries in Cruciferae 368
356
INDEX TO VOLUME XLI
469
Welwitschia 226
biarag gg § Lund, C., on lakes 360
Wes Bis, “Mono graph of British Des-
miiaceae” 452
, growth of root 139; growth of
i 54
Reet H. H., on blight canker 366
White, E. A., “Fungi of Connecticut” 215
Wiegand, K. nde
iesn 457; “— light rela-
a. . igh oa 15
Kien 223,226, nen 364, 365,
6, 367
Wilderan Emile de,, personal 307
n of salts 224
. aceae
Williams, R. S., personal 159
Winton, A. L., ‘‘ Microscopy of vegetable
foods”’ 300
Wittrock, V. B., Biers illustratus
iconothecae botanicae”
Wolfe, J. J., personal 227
Wolff, G., on apothecia of lichens 306
i
Xerophily, causes of 22
Yamanouchi, Shig2o 425
Yeasts, esneeton of 157
York, H. H., personal 372
Sonera Pd oie of 14)
ca Mays, saps bundles ro, nodes
Zeller, R., on fossil gymnosperm
Zizania aquatica, amphivasal Seestleg 3 10;
Barents iy tioides, absorption of water
279
daea 357
Zygadenus exaltatus 283
Zygnema, nuclear division 43
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have ever done before ?
Then here is a brief outline of
best two weeks you ever lived.
Leave Chicago (for example) any day after June 1 on
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ou may travel via Omaha, Pacific Junction, St. Joseph or
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From Denver take a side trip to Colorado Springs (no
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Returning to Denver, spend from one to three days in
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Leave Denver on the Burlington’s Yellowstone Park
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Big Horn Mountains; past Custer Battlefield, the most
tragic upon which
Yellowstone Valley to Gardiner, the official entrance to
the Park.
Don’t you think you would like to make this tour?
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The cost of a railway ticket for the entire tour (exclusive of side ae
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Too expensive? No!
five and a half
City to Omaha, inclusive.
P130
A Colorado-
Yellowstone Tour
Have you two vacation weeks at your disposal ?
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a tour that will bring you the
This tour provides for a stay of five and a half days in
ark — coaching
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scenery on the globe and being entertained at t
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after seven days $3.50 and up per day.
apolis and St. Pa
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finest river se
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The Mecca of the Leisure-loving
Lon sland
ALWAYS COOL IN SUMMER
Oe more attractions than ati atk = gone soon New
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descriptive literature.
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SOME RAILWAY PROBLEMS
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BUFFALO
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only a aie solvent, but has the additional virtue of containing substantial quantities of
the Alkaline Lithates. Patients should be encouraged to take two quarts per day, i
they can, and the relief they will obtain will be all the argument necessary after the
first day or so.
“The Results Satisfy Me of Its Extraordinary Value.”
Dr. tas, AGT of New Orleans, Ex- aur of the State Board of Health of
Louisiana, n affections of the kidneys and
‘“T have prescribed BUFFALO LITHIA WATER ... nary passages, particularly in
Gouty subjects, in Albuminuria, and in irritable euuatied of the Bladder and
Urethra in females. The results satisfy me of its extraordinary value in a large class
of cases usually pit difficult to treat.’
“I Have Witnessed Decided Beneficial Results from Its Use.”
Wm. B. Towles, M. D., formerly Professor of Anatomy “ge Materia Medica of
the University of Vir- are marked in causing a disap-
ginia: “ The effects of BUFFALO pearance of Albina from the
urine, and in cae nt of Bright’s Disease | have witmessed decided beneficial
results from its us
“Results, to Say the Least, Very Favorable.”
igs Griswold Comstock, A. M., M.D., Sz. Louis, Mo., says: ‘‘I have
mad in gynecological practice, in women suffering
use of LITHIA from acute Uremic conditions, with results,
to say the least, very favorable.”
Additional medical testimony on request.
For sale by the general drug and mineral water trade.
PROPRIETOR BUFFALO LITHIA SPRINCS, VIRCINIA-
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Be Fair to Your Skin, and It Will |
Be Fair to You--and to Others |
A Beautiful Skin can only be secured through Nature’s work. Ghastly, horrid
imitat‘ons of Beauty are made by cosmetics, balms, powders, and other injurious
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the skin that all incentive to the use of cosmetics is lacking.
HAND SAPOLIO is
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that it can be a part of the invalid’s supply with beneficial results.
SO SIMPLE
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