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BOTANICAL GAZETTE

EDITORS:

JOHN MERLE COULTER anp CHARLES REID BARNES

= VOLUME XLVII
a

JANUARY-JUNE, 1909

WITH TWENTY-FIVE PLATES AND ONE HUNDRED AND FIFTEEN FIGURES

CHICAGO
THE UNIVERSITY OF CHICAGO PRESS

TABLE OF CONTENTS

PAGE

On triple hybrids - ~~ Sennen ene. See ee ae Hugo de Vries I

Periodicity in Spirogyra - - - - W. F. Copeland 9
On the pollen of M sola te ene et plates

Tand II) - - Robert Boyd Thomson 26

A vegetative mutant, and the principle of homoeosis
in plants. Contributions from the Ames Botani-
cal Laboratory No. 9 (with nineteen figures) -

Robert Greenleaf Leavitt

Relation of soil and vegetation on es sea shores

(with twelve figures) - - - Pehr Olsson-Seffer
Some aspects of amitosis in Synchytrium (with fad
Ill andIV) - - - - - - - Robert F. Griggs

Vascular anatomy of the seedling of Microcycas calo-

c Contributions from the Hull Botanical

Laboratory 122 (with plates Vand VI) - Helen Angela Dorety
Mitosis in Fucus. Contributions from the Hull Bo-

tanical Laboratory 124 (with plates VIII-XI) - Shigéo Yamanouchi
The reduction division in the prin ae of

Agave virginica (with plates XII-XIV) - John H. Schaffner
Spermatogenesis in Dioon edule. Contributions te :

the Hull Botanical Laboratory 125° (wi

plates XV-XVIII and three figures) - - ene J. Chamberlain
Undescribed plants from Guatemala and other Cen-

tral American Republics. XXXI (with one figure) John Donnell Smith
Comparative histology of fruits and seeds of certain

species of Cucurbitaceae (with fifty-three figures) Kate G. Barber
The anatomy of Isoetes. Contributions from the Hull -

Botanical Pecunia 126 es seo XIX-

XXT) Alma G. Stokey
The megasporophyll of Be Sis and Micro-

cachrys (with plates XXI-XXV) - Robert Boyd Thomson
Studies on the oxidizing powers of roots

Oswald Saves and Howard S. Reed

Bog toxins and their effect £8 soils (with two

figures) - - Aljred Dachnowski

vi CONTENTS [VOLUME XLVI

Contributions from the Rocky Mountain Herbarium. VIII Aven Nelson
The leaves of Podophyllum -— - - + J. Arthur Harris
A botanical survey of the Huron River fie. VII.

Position of the greatest i sli in local bogs

’ (with five figures) George Plumer Burns
Pollination in Linaria with special reference to cleis-

togamy (with four figures) - E. J. Hill
BRIEFER ARTICLES—

Longevity ofseeds - - -~ one . William Crocker

Respiration calorimeter - - - ~ - George J. Peirce
Crataegus in Colorado — - - - - Francis Ramaley
The nature of balanced sefations - - - W.J.V. Osterhout
The extrafascicular cambium of Ceratoza amia.

Contributions from the Hull Botanical Labo-

ratory 123 (with plate VII) - - - Helen A, Dorety
The mounting ofalgae - = - - -- J. A, Nieuwland
Paul Hennings (with pose $e - - J. Perkins
Pure cultures of fun .
Parthenogenesis in Pine Pinaster bwith seven

gures) - :

Johanna Westerdijk

W. 7. Saxton
Carnation sienadi (with eight es Eng Se Siena: and J. G. Hall
CurRENT LITERATURE - -
For titles of books reviewed see cae indies au-
thor’s name and reviews
Papers noticed in “Notes for Students” are in-
dexed under author’s name and subjects

DATES OF PUBLICATION

No. 1, January 29; ae 2, saa 20; No. 3, March 23; }
No. 5, May 21; No. 6, Jun

PAGE
425
438

72, 153, 242, 336, 414, 467

vo. 4, April 17;

es ed ee = e Seas
ee i RN es eS Ss
ee, Bete Sy eine oot Pau alee

eer

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fone vl

ERRATA
VoLuME XLVI
302, line 7 from bottom, for KCI read CaCl.

. 326, line 5 from top, for distant read distinct.

400, line 6 from top, after division insert reduction; and in line 7, delete a
and reduction.

VoLtuME XLVII
69, footnote 1, for Ewart, A. L. read Ewart, A. J.
144, last line, for Goveniana read Goweniana.
170, footnote 24, for Leavitt, R. S., read Leavitt, R. G.
347, line 6 from bottom, for cones read, comes.
423, no. 9, for Abisdia read Absidia.

On Triple Hybrids _

Bos
-]

ee,

gles

sere

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)
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Che Botanical Gazette

A Montbly Journal Embracing all Departments of Botanical Science

Edited by JoHN M. COULTER and CHARLES R. BARNES, with the assistance of other members of the
botanical staff of the University of Chicago.

Issued January 29, 1909

Vol. XLVII CONTENTS FOR JANUARY 1909 No. 1
ON TRIPLE HYBRIDS. Augode Vries - - - : . - - - - : : I
PERIODICITY IN SPIROGYRA. W. F. Copeland ~ - - : - - “ i 9
ON THE POLLEN OF M/CROCACHRYS TETRAGONA CATE? PLATES 1 AND II). odert
Boyd Thomson - -
A VEGETATIVE sate a AND THE PRINCIPLE OF HOMOEOSIS IN PLANTS,
CONTRIBUTIONS FRO E AMES BOTANICAL LABORATORY No. 9 (WITH NINETEEN FIG-
RES). Robert awe Cee - - - 4 < : ae
BRIEFER ARTICLES
ONGEVIRN On Genie . Wile Grok <a ee ee
RESPIRATION CALORIMETER. George J. Peirce - - : a es 3 ‘ - ae
CRATAEGUS IN COLORADO. Francis Ramaley - - - - ‘ . * - we.
CURRENT LITERATURE
BOOK REVIEWS CO I rea ag ee nO Eee ea
E VEGETATION OF CHILE. THE PENDULATION THEORY
MINOR NOTICES mi Oe ana ah a a Break nc a i ate onl come nites ne
Wetns FOR STUDENTS OOO ee eee

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Entered August 21, 1896, at the Post-Office at Chicago, as second-class matter, under Act of Congress March 3, 1879.

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The Life History of Polysiphonia Violacea

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The English Rejormation and Puritanism, vith Other Lectures and
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VOLUME XLVII NUMBER 1

MeXTANICAL GAZEETG
JANUARY 1909

ON TRIPLE HYBRIDS
HvuGo DE VRIES

Twin hybrids are produced when the pollen of Oenothera Lamarck-
tana or of one of its derivatives is crossed with the European sub-
species of O. biennis or of O. muricata. These twins appear in about
equal numbers and are constant in succeeding generations. One
of them, O. laeta, is broad and smooth leaved; the other, O. velutina,
is more hairy and has furrow-shaped leaves.?

Triple hybrids may be produced by combining this phenomenon
with the hereditary qualities of O. lata and O. scintillans. Both of
them originated in my garden from O. Lamarckiana. OO. lata is
female, its anthers are barren. O. scintillans is an inconstant species
which repeats its type only in one-third or two-thirds of its offspring.
Both of them, when pollinated by O. Lamarckiana, give a mixed
Progeny, only part of which is like the mother.

Nn order to get triplets, therefore, we have to cross O. lata or O.
sciniillans with some species which will split them into Jaeta and
velutina (as it would do OQ. Lamarckiana itself or some others of its
mutant species), and which, moreover, does not prevent them from
Tepeating their own type in their progeny. The triplets will then be
O. laeta, O, velutina, and O. lata. The first two will drop the special
character of the mutant-parent (Bor. Gaz. I. c. 403), but all three will

intermediate hybrids between O. Lamarckiana or O. lata and the
species used as the other parent.

I found that O. strigosa Rydberg, O. Hookeri T. and G., and the
“emmon American subspecies of O. biennis L. comply with these
t Bor, GAZETTE 44:401-407. 1907.
* Die M. ulations-Theorie. Vols. I and II.

2 BOTANICAL GAZETTE [JANUARY

requirements. The experiments to be described in this article are _
limited to the effects of their pollen on O. lata and O. scintillans.

O. strigosa Rydb. has very small flowers, of the size of those of
_O. muricata, but in less dense spikes. They open their anthers
two or more days before unfolding the corolla, and the pollen comes
in contact with the whole outer surface of the stigma and causes
fertilization. The leaves are bluish green and furrow-shaped, and
the tip is bent sideways. The tops of young shoots, when seen from
above, therefore, present the aspect of a wheel with spokes all curved
to the left or to the right. This striking feature is repeated, though
somewhat reduced, in all of its hybrids and causes them to be easily _
recognized. I collected seed in the Yellowstone Park in 1904.
Another lot was kindly sent to me by Mr. T. D. A. CocKERELL from
Boulder, Colorado. Of both strains I have since cultivated two
generations.

In crossing O. strigosa with O. Lamarckiana, twins arise which
combine the characters of laeta and velutina with those of strigosa.
They differ more widely from one another than do the twins of any
other cross I have studied up to this time. The velutina is almost like
the velutina of O. muricata < Lamarckiana, but has the bluish tinge,
the more pointed leaves with bent tips, and the wheel-shaped tops
of the young shoots described above for O. strigosa. The laeta from
the strigosa cross is a very stout plant with very broad leaves (6°™),
blunt at the apex and indented at the base, with smooth surface, and
bright green. The flower buds are more narrowly conical than those
of velutina, the flowers open more widely, the fruits are conical with
four sharp and protruding teeth, whereas those of velutina are blunt
and short. By this mark and by the somewhat narrowed neck of the
fruits of velutina, the two forms are most easily distinguished when
flowering is over. The Jaeéa is usually poor, but the velutina is rich
in pollen.

In contrast to the species previously dealt with, O. strigosa pro-
duces twins from O. Lamarckiana when used as male parent as well
as when used as female parent. In these reciprocal crosses both of
the twins are identical.

The twins produced by O. strigosa from O. scintillans and from
O. lata comply with the description given. They cannot be distin-

1909] DE VRIES—TRIPLE HYBRIDS 3

guished from them externally, even when cultivated in large and pure
lots.

O. Hooker T. and G. is a Californian species and is also found in
Texas. I collected seeds in the vicinity of Berkeley, California, and _
another lot was sent to me from Riverside by Mr. Frep, M. REED.
It is a striking species, even more beautiful than O. Lamarckiana.
The flowers are of the same size, protruding their pistils high above
the anthers; but the petals are more deeply emarginate at the tip,
nearly obcordate, and a paler yellow. The plant is hairy and reddish,
and the leaves (especially those of the rosette) are long and narrow.

With Lamarckiana it produces twins identical in both reciprocal
crosses and both of them as large-flowered as the parent species.
The /aeta is bright green, with smooth leaves and slender flower buds.
The velutina is hairy and reddish, with furrow-shaped leaves and thick
buds. It flowers more profusely and resembles O. Hookeri almost
exactly, but is stouter, with dense and richly flowered spikes.

The twins derived by O. Hookeri from O. lata cannot be distin-
Suished externally from those derived from O. Lamarckiana. :

“The American subspecies of O. biennis used for my experiments
was collected by me in 1904 at Chicago, near Jackson Park. It seems
to be the same form as that which is most common in the eastern
states. Other subspecies I have collected in Pennsylvania, Kansas,

Missouri, and elsewhere. All of them are quite different from the
. European form, which is the one used in my Muations-Theorie and
m my article on twin hybrids (Bor. Gazette /.c.). A systematic
treatment of these numerous forms is still wanting, and therefore I
will provisionally designate my strain by the name of its source and
call it Chicago. Its most curious quality is that of producing twins
when combined as a male parent with O. Lamarckiana and not when
crossed with the pollen of this species. In this respect it is exactly
°pposite to the European O. biennis, and therefore very useful.

The O. biennis from Chicago is a taller plant, more richly branched,
and darker green than the European form. Its flowers are much
smaller, sometimes reaching the size of those of O. muricata, but

3OQen. Hookeri T. and G.=Onagra Hookeri Small=O. biennis hirsutissima

vast Bot. Calif. See H. M. Hatt, A botanical survey of San Jacinto Mountain.
niv. Calif. Publ. Bot. 1:98. 1902.

4 BOTANICAL GAZETTE [JANUARY

deeper yellow. The stigma is surrounded by the anthers which
open early in the bud. The velutina and laeta which it produces
from Lamarckiana can hardly be distinguished from those produced by
the European O. biennis, although the leaves are narrower and darker
green. I cultivated this strain during three succeeding generations.

O. scintillans and O. lata are the mutant species described in my
Mutations-Theorie. The scintillans used originated from Lamarck-
tana seed of 1889 which was sown in 1895. In 1906 I had the fourth
generation from continued pure self-fertilizations and used it for my
cross. From this same strain the lata used in the experiments of this
article arose as a mutant in 1901. It was artificially fertilized. My
crosses were made with specimens of the second and third generation
of its progeny.

I will now describe the crosses from which the triplets resulted.

Oenothera scintillans X strigosa-—This cross was made in July,
1907, between two biennial specimens. It yielded a small lot of seed
which was sown the next spring. From it arose 36 laeta, 21 velutina,
15 scintillans, and one lata, altogether 73 plants, most of which flowered
during thesummer. The Jaeta and velutina agreed with the descrip-
tion given above; the Jafa was a mutant. The scintillans were inter-
mediate hybrids, which had the habit and leaves of the mother
parent, or nearly so, but small flowers with the anthers surrounding
and touching the stigma like the father. I fertilized all three forms
with their own pollen and intend to sow the seed next year.

Ocnothera lata X strigosa.—This cross was effected in 190s, both
parents being annual specimens. I used different plants of Jata and
sowed the seed of one cross in 1906, of another cross in 1907, and a
third lot in 1908. I got the following results:

Year i Percentage laeta P pair Percentage Jata
Cae sears he os was 41 36 59 5
POP Sn eyes eo ewes sae 49 40 II
QUO einige urs. css 270 27 34 38
Fetal. scabs ee pecen 358 40 33 27

The laeta and velutina were the same as those from the cross with
Lamarckiana. 1 sowed self-fertilized seed from plants of 1906 in
1907; the mothers were velutina and gave 113 children, most of which

aT nS se eh

a as sr ee

SO oe eS eee a a

OE ea a ee ee,

4
|

1909] DE VRIES—TRIPLE HYBRIDS 5

flowered. All of them were velutina. In 1907 I self-pollinated some
laeta and some velutina and got 80 and 45 seedlings respectively.
The first were all Jaeta, the second lot was uniformly velutina. Of
each lot 25 specimens were preserved after the sorting period and
observed during the time of flowering and ripening of their fruits.
The lata were intermediate between O. lata and O. strigosa. In
1908 I cultivated one half of them in the open and the other under
an open glass-covering, both of them with cultures of ordinary lata
under the same conditions for comparison. The /ata from the
strigosa cross had narrower leaves but with the rounded tips; the
bracts were broad but less bent downward. The flowers were much
smaller than those of the mother, but somewhat larger than those

of the father. Their shape resembled that of O. strigosa, but the

stigma showed, although in a lessened degree, the peculiar hand-
shaped form of the ordinary O. lata. The anthers touched the
stigma, but only with their lower halves. The flower buds were
broad, and the tube was relatively short. In all these points and in
the other characters the lata hybrids were strikingly intermediate
between the two parents. Even the bent tips of the young leaves
were to be seen, and at once revealed the father. But the most
interesting feature was that of the pollen. O. Jata is purely female,
while O. strigosa has the ordinary supply of pollen. The hybrids
showed some pollen and a relatively small number of fertile grains.
These were, during ordinary weather, insufficient for fertilization,
even if the pollen of many flowers was brought upon one stigma.
But there were individual fluctuations, and so I succeeded in getting
self-fertilized fruits on at least one specimen.

Oenothera lata X O. biennis Chicago.—This cross was made in
1905, both parents being annual. Two specimens of O. lata were
used. I sowed their seed in different lots in the three succeeding
years and had the following results:

Year Number of [Percentage laeta alee a
a eee eS:
me) oo
td ohh an te Si 33 27 7 :
ae 78 34 ot | a

ee eee 167 36 a8 -
— 278 35 34 -
>: See OE as oe We

6 BOTANICAL GAZETTE [JANUARY

The Jaeta and the velutina were of the description already given.
I sowed the self-fertilized seed of 1906 in 1907 and 1908 and had
four lots of the second generation. The lots of the first included
69 and 139 individuals, all of which were Jaeta; the sowings of the
other strain extended over 38 and 158 samples, all of which were
velutina. Both of the twins thus complied with the rule of constancy
in the second generation. From the lots of 1907 I cultivated in 1908
a third generation comprising 70 children of laeta and go of velutina;
both lots were once more uniform and similar to their parents.

The /ata individuals of this cross were almost similar to those of
O. lata itself. However, they had the smaller flowers of O. biennis,
with the self-fertilizing position of the stigma, and the long lobes of
the father. Their flower buds were intermediate between those of
the two parents. In 1906 and 1907 the anthers seemed wholly
barren; but in 1908, during the very warm last days of July, they
yielded some pollen, which was used partly for self-fertilization,
partly for crosses.

O. lata X Hookeri.—This cross is the most interesting of all the
crosses with O. lata, since its lata hybrids are almost as rich in pollen
as any other evening primrose. This enabled me to study the second
generation of the Jata, which, in contrast to that of the Jaeta and the
velutina, repeats the splitting. The yield of the pollen was small in
the unfavorable summer of 1907, but large during the hot days of
July and August, 1908.

I have made this cross twice, in r905 and 1907; the parents were
annual specimens. I had the following results in the first generation:

|
Number of | Percent P t P
Year Seed of specimens J Lite baauar r ele
BOGE Ue ec re ey iy 1905 51 53 30 8
TO ee 1905 229 45 47 8
IQR rei i, 1907 72 49 28 22
i: RRR JE ieee 352 47 42 II

Here also the Jaeta and velutina were the same as those from the
cross with Lamarckiana. I sowed some self-fertilized seed of the
velutina of 1906, partly in 1907 and partly in 1908, and cultivated

1909] DE VRIES—TRIPLE HYBRIDS 3

from each lot 60 specimens, all of which repeated the velutina char-
acters.

The lata specimens also were cultivated from seed of 1906 in both
succeeding years. Their flowers were as large as those of both
parents and intermediate in the tinge of the yellow, the Hookeri
being paler yellow than the Lamarckiana and its lata. The anthers
did not reach the stigma, which was often hand-shaped. The spike
was much denser than that of Hookeri but thinner than that of the
mother. Stem, veins, and calyx were reddish; and leaves were
narrower than in O. lata. In all these and in other characters the
plants were strikingly /ata, but with the addition of the differentiating
marks of the Hookeri.

The self-fertilized seed of the Ja¢a plants of the first generation
gave a mixed progeny, consisting of velutina and lata, both resembling
the types of the first year. No Jaeéa specimens were produced. The
numbers were the following:

Waar Number of Percentage - Percentage

: seedlings velutina lata

= caf gare eee 53 85 15

EQOD Soh. ceo 134 81 19
fice ee 187 82 18

From the experiments described the following table may be
compiled :

TRIPLE HYBRIDS AND THEIR SUCCEEDING GENERATIONS

M First ti Second Third
other Father go iainssecrnyg generation generation
eens i Docume
©. scintillans X O. strigosa art Oe a uit 49 laeta
: 29 velutina
21 scintillans
O. lata xO. BUI OGG 6s cc nae ts 40 laeta 8o laeta
“ 33 velutina 158 velutina
: 27 lata
O. lata XO. bien. Chicago..... 35 laeta 208 laeta 7° :
34 velutina 196 velutina | 90 velutina
1 lata
O. lata *%O, Hookeri.. 5.0.3... 47 laeta
42 velutina 120 velutina
82 per cent,
velutina
11 lata 18 per cent
lata
ee

8 BOTANICAL GAZETTE [JANUARY

QUADRUPLE HYBRIDS.—The discovery of twin and triple hybrids
naturally suggests the idea of the possibility of a hybrid quartette.
As a matter of fact the experiments did the same. In the case of O.
lata X O. biennis from Chicago a fourth type appeared in one speci-
men, and in that of O. lata x O. Hookeri in two specimens. These
three plants belonged evidently to the lata type, but combined with
this the smooth, broad, and bright-green leaves of the Jaeta, whereas
all the other datas of these crosses had gray and furrow-shaped leaves
like the velutina. The lata-laeta of the first cross was sufficiently
fertile, but of two of the Hookeri cross one had barren anthers and
the other only some spare pollen in comparison with the rich supply
of pollen in the other /Jata plants of this cross. In this respect, there-
fore, they also showed the character of the Jaeta type.

It would seem probable that the lata-laeta should be produced in
the same number as the other Jata, or lata-velutina as we could now
call it. But then the Jaeta itself often appears in too small numbers.
The cause of this phenomenon has still to be investigated.

Summary

1. Triple hybrids are produced in crosses of Oenothera scintillans
and O. lata by such species as produce twins from O. Lamarckiana.

2. The species investigated are O. strigosa Rydb., O. Hookeri T.
and G., and one of the American subspecies of O. biennis.

3. Of the triple hybrids, two are the same as the twins from the
corresponding Lamarckiana crosses and bear the characters of O. laeta
and O. velutina combined with those of the other parent.

4. The third type resembles the mother (0. ata or O. scintillans) ,
but in its special marks is also intermediate between its parents.

5. The laeta and velutina are constant and uniform in their succeed-
ing generations, so far as experience goes. In this respect they follow
the rule for the twin hybrids of O. Lamarckiana.

6. The lata, however, in the only case tried, repeated the splitting
after self-fertilization, producing, however, only lata and velutina.

7- It seems probable that the whole progeny of the crosses named
should split up into two equal parts, Jaeta and velutina, and that each
of them should produce a certain percentage of Jata. In this way
quadruple hybrids would arise.

Botanic GARDEN
TERDA

PERIODICITY IN SPIROGYRA
W. F. CoPELAND

The object in beginning these studies with Spirogyra was to get
notes on the rearing of species and to see how far they could be
changed by conditions. With this end in view 1oo aquaria were
started and soon contained puzzling results. To interpret these and
to get on with the experiments, it soon seemed desirable, if not neces-
sary, to know how Spirogyra lives under natural environment. For
this reason the major part of my time was given to a consideration of
this plant out of doors. An attempt has been made to repeat all
observations and experiments under as great a variety of conditions
as possible in an ordinary biological laboratory. When field-work
was not practicable or necessary, laboratory work was resumed. In
this way scores of experiments have been in progress from the begin-
ning. -

After this work had continued for some months, considerations
were given to conditions appearing with some regularity whether
Spirogyra was being studied under natural or under artificial environ-
ment. As a possible constant condition, it was finally decided to give
Special attention to the subject of periodicity. In this paper only
those experiments will be mentioned which seem to have some bear-
ing on this subject. Although laboratory studies are of minor
Importance in the subject of periodicity, in the present discussion
they will appear first.

I take this opportunity to express my obligation to those who have
mm any way aided me in the present study. It is a pleasure to thank
Dr. C.F. Hopcx, who suggested the problem, for his help and criti-
“ism; HELEN REINHERR CopELAND for help in identification of
Species and preparation of material for microphotographs; O.

Mitts, Millbury, Mass., for collecting much valuable material in
at locality; and Dr. O. P. DELLINGER, Worcester, Mass., for help-
ful Suggestions and criticisms. The following persons have sent me
©onjugating specimens from more distant localities: W. D. Hoyt
and I. F. Lewis of Johns Hopkins University; B. A. PLace, Iowa
9 [Botanical Gazette, vol. 47

se) BOTANICAL GAZETTE [TANUARY

Wesleyan University; S. G. Winter, Illinois Wesleyan - University;
and W. A. MATHENY, Sardis, Ohio.

Review of literature

In 1803 VAUCHER (6) tells of his attempt to work out the life-
history of the Conjugatae. For this purpose the genus Spirogyra
was used and especially the species which MULLER had previously
described as Conjerva jugalis. VAUCHER’S observations were made
in the spring and it was found that this species would fruit and dis-
appear as early as February 15 and as late as March 20. The zygo-
spores which he had collected germinated July 15, and all seemed to
germinate on the same day.

A history oj the British freshwater algae by Hassa.u (3) appeared
in 1857. This well-known work contains descriptions of 42 species
of Spirogyra under the name Zygnema. A single hint is given as to
the time of year when conjugation takes place; namely, ‘‘The
species of this group of confervae may be found in a state of conjuga-
tion during the entire spring, summer, and autumnal months, but
more abundant in the spring.”

In 1874 The freshwater algae of the United States by Woop (9)
was published in the Smithsonian Report. Speaking of the Zygne-
maceae he says: “The family under consideration is among the
commonest and most widely diffused of all the freshwater algae.
They may be found in greater or less abundance at all seasons, but
the spores appear to be formed only in the spring and early summer,
at least these are the only times at which I have found fertile filaments.
In the neighborhood of Philadelphia I have collected them in excel-
lent condition as early as the beginning of April and as late as the
latter part of June. Further south conjugation of course commences
earlier, and fine specimens received by myself from Mr. CANBY
were collected by him in Florida in February.”

The work of Paut Perrr (5) entitled Spirogyra des environs de
Paris appeared in 1880. No general statement is made regarding
the fruiting season of this genus, but, with one exception, dates are
given with the description of the different species. These dates show
that 20 of the 36 species described matured during March, April, and
May; and all except two had matured by July. These were found

1909] COPELAND—PERIODICITY IN SPIROGYRA II

in fruit from Jul until October. One species fruited in April and
May and pa October. | -

Since 1880 a number of papers relating to Spirogyra have appeared,
but only a few of them give any facts that can be used in a study of
periodicity. BENNETT and Murray (1) write: “Germination some-
times takes place while still in the mother cell; but most commonly
both filaments perish after conjugation, with the exception of numer-
ous zygospores which fall to the bottom. These remain dormant
through the winter . . . . germinating in the spring . . . . instances
are recorded of filaments persisting through the winter. HOFMEISTER
states that the growth of Spirogyra is intermittent.”

The following case is reported by W. and G. S. West (7): “We
have melted out of the ice from Micham Common, Surrey, excellent
examples of Spirogyra catenaeformis in conjugation, the vitality of
which was in nowise impaired.’’

The few scattered remarks given include all the notes the writer
has been able to gather from papers published before 1905. In this
year a few papers appeared which had a more direct bearing on the
subject at hand. These seem to be the pioneer publications on the
subject of algal periodicity. In indicating some of the ‘problems of
aquatic biology” Frrrscu (2) calls attention to the fact that different
bodies of water, often far apart, will have the same dominant and
often the same subordinate vegetation; also that a plant may be very
abundant one month and almost, if not entirely, absent the next
month. An intimate relation was found to exist between the relative
abundance of an alga and its period of reproduction; the formation
of the sex organs frequently preceding the disappearance or dimin-
ished abundance of a given species. FrirscH writes: “In some
cases no doubt these features (maximum abundance and period of
Teproduction) are influenced by the periodically recurring factors
such as the rise of temperature and the increase in the intensity of
light in the spring; so that the maximum abundance and period of
Teproduction are definite phenomena.”

Another paper on the subject of algal periodicity appeared in
1905. So far the publications mentioned have had to do only with
freshwater algae; for that reason it is well to call attention to studies
made on marine forms. While at work on Dictyotaceae, WILLIAMS

¥2 BOTANICAL GAZETTE [JANUARY

(8) discovered that Dictyota had definite periods for the development
of its fruit. This is well explained in his introductory paragraph: |
“While studying the development of the sexual cells in Dictyota,
the interesting discovery was made that the process was practically
simultaneous, not only for a given plant but for all the plants of the
locality.”

This work by Wittams was done at Bangor, Wales, and at Ply-
mouth, England. During the summer of 1906, observations were
made on the same plant at Beaufort, North Carolina, by W. D.
Hoyt (4): “It was found at Beaufort, also, that this alga produced
its sexual cells in regular, periodic crops... . . One observation is
interesting in this connection. A female plant kept in the laboratory
for nearly two months and then examined showed sori only slightly
less advanced than those on plants growing in the harbor.....
This result is similar to the one observed by Wixtiams but goes a
step farther in showing that periodicity is transmitted to new struc-
tures formed from the original plant, even where these are not sub-
jected to alternating conditions... . . It still remains to be seen
whether the periodic habit can be transmitted through the proto-
plasm of the egg.”

Spirogyra under artificial environment

In the laboratories in which these experiments were conducted,
windows were used which faced toward the northwest, northeast,
and southeast. Whenever a new line of investigation was begun,
three sets of aquaria of different sizes were prepared. One set was
placed in a northwest, one in a northeast, and one in a southeast
window. The conditions made use of in the three sets of experiments
were the same with the exception of a few degrees in temperature
and a difference in light intensity. Other experiments were con-
ducted which had nothing to do with conditions of temperature and
light.

In the first experiments Spirogyra was brought into the laboratory
and small amounts placed in aquaria filled with water from the
pond from which the plant was taken. In some cases the plant was
washed before being transferred to the aquaria, in others it was not.

A second group of experiments contained the same conditions

1909] COPELAND—PERIODICITY IN SPIROGYRA 13

with the exception of the culture medium. In this group attempts
were made to repeat the experiments while using tap instead of pond
water. Asa result, it was found that the aquaria with tap-water were
as satisfactory, and, in most cases, much more so than those with
pond water. Hence tap-water aquaria were used as controls.

It is well to note concerning this point that the tap-water was from
reservoirs which were in turn supplied by small brooks and surface
drainage. In many cases the pond water was taken from ponds
supplied by brooks which drained the reservoirs just mentioned.

I next tried a group of experiments for the purpose of fixing some
value to the use of plant food solutions for the investigation at hand.
Knop’s solution was most frequently used; however, Sachs’s solution
was employed in a great many experiments and in most cases simul-
taneously with Knop’s. In comparing aquaria with and without food
solutions, no general rule could be established. On the whole, how-
ever, the writer is inclined to favor at least a weak food solution.
Knop’s solution as low as 0.04 per cent. gave favorable results.

A long list of experiments was prepared in which rain and distilled
water were used; and in others melted snow. Some of these were not
disturbed after being placed in their respective windows. Others
were left in these media for periods varying from one to ten days
and then changed to a weak food solution (usually 0.05 per cent.),
and this was raised by degrees to 0.2 per cent. where it was allowed
to remain. In this group care was taken to select Spirogyra from
different localities. Specimens were obtained from swift water, from
slow-running brooks, and from small quiet pools and ponds where very
little disturbance was possible. Notes on these experiments show
that no particular advantage was gained by using these media.

From the first the aquaria located in the southeast windows were
shorter lived than either of the other sets of aquaria. It will be
remembered that the only difference in conditions was a slight differ-
€nce in temperature plus a difference in light intensity. Experiments
were now conducted in order to try the effect of shading the cultures.
Tissue paper of different colors was used for this purpose. The best
Tesults were obtained when the aquaria were not shaded above the
Surface of the water. After scores of experiments the best results
_ Were obtained in the following way: when an aquarium was started,

I4 BOTANICAL GAZETTE [JANUARY

it was shaded with black paper; after two weeks, this was changed
to yellow; and after another two weeks the shading was not used at
all. This was a decided advantage to the aquaria in the southeast
windows. While the effect was not so noticeable in the other labora-
tories, the results were sufficient to justify this method in all the win-
dows. It is not to be understood that tap-water aquaria were the
only ones used with a shade. On the other hand, snow, rain, distilled
water, and food solutions were used in every case. The same applies
to the following series of experiments. ox

Up to this time no attention has been given to the substratum,
nor to other plants with which Spirogyra is most frequently asso-
ciated in its natural habitat. For this group of aquaria soil, débris,
and plants, for the most part, were used from the immediate locality
from which the specimens were taken. In some cases the soil was
first sterilized.

Among the plants which seemed beneficial to Spirogyra, none
proved more helpful than Oedogonium. In fact in every aquaria
in which Spirogyra lived for several months or a year, Oedogonium
was invariably present and often the predominating alga. The
Oedogonium could usually be found attached to the sides of the jar
or floating at the surface, thus shielding the Spirogyra from too much
sunlight. Chara supplied to the aquaria was helpful so long as it
was not too abundant. In some cases where aquaria were thus
started, the Chara died out after two or three months and was followed
by an abundant growth of Oedogonium, and later by Spirogyra. In
the field, masses of Spirogyra were often collected among water cress,
but in no case was a good growth of Spirogyra obtained in an aquarium
with this plant. A number of jars were supplied with oak leaves
taken directly from the trees where they had remained all winter.
To other aquaria pieces of charcoal were added. Both of these
factors seemed beneficial and in most cases excellent cultures of
Spirogyra were obtained which lasted until conjugation had taken
place.

Spirogyra under natural environment

Field-work was begun early in April. At this time 400 aquaria
of all sizes were in use and 100 of them contained excellent cultures
of Spirogyra which had been under cultivation for two months. On

ee ee ee ee See

1909] COPELAND—PERIODICITY IN SPIROGYRA 15

April 27 S. quadrata was found conjugating abundantly. This one
was followed by others until twelve different species had matured and
disappeared. At first all material was brought to the laboratory in
small jars and then examined, but this method was by no means
satisfactory. A microscope together with a crude temporary stand
was added to my collecting outfit, and an examination of all material
was made on the spot. There are so many chances of collecting
worthless material that no other method was found which would give
definite results.
The first task was to locate Spirogyra in a number of different
localities with different conditions of environment. Accordingly
a few localities were selected varying from a few rods to twelve
miles apart. Within these regions 40 different places were selected
which could be visited frequently and studied with care. Some
were in open places, while others were in deep shadow; some were
in pure running water, others in filthy stagnant pools; some with
numerous other algae, others almost alone; and under many other
conditions.
_ For identification of species some of the material was usually
killed in the field as soon as collected. From 2 to 4 per cent. solu-
tion of formalin was used for this purpose when larger masses were
killed, but in most cases chromacetic solution was used according
to the formula given in CHAMBERLAIN’s Methods in plant histol-
ogy, p. 139 (chromic acid 1, glacial acetic acid 4°°, water 400°°).
Several stains were used, but Haidenhain’s iron-hematoxylin was
most satisfactory. Some of the specimens stained in this way were
afterward mounted in glycerin, others in balsam, and others in Vene-
ian turpentine. I found the balsam more satisfactory than the

turpentine, but excellent preparations were obtained by using the
Venetian turpentine method recommended by CHAMBERLAIN (stain
with magdala red and anilin blue and mount in Venetian turpentine).
Stained preparations in glycerin were most quickly obtained by tak-
Ing the material as soon as washed from the chromacetic solution,
transferring it to a 10 per cent. solution of glycerin to which was
added 2 or 3 drops of Ehrlich’s blood stain, and allowing it to con-
Centrate. In some cases the stain obtained by this method has been
Permanent, in others it has faded after a few months. This method

16 BOTANICAL GAZETTE [JANUARY

of preparation has been used with much success by F. N. DuNcAN
and O. P. DELLINGER, of this laboratory, for animal tissue.

Periodicity of species studied

For the purpose of identifying the different species, the following
authors were consulted: VaucHER (6), Woop (9), Perit (5), and
Wo te (10). A brief account of the fruiting season of the different
species will be given below.

Spirogyra inflata (Vaucher) Rab.—VAucuHeER reports this species
in fruit from February 20 to March 19. Prrir gives April-May.
Unfortunately I was not able to find it except in a single locality.
This was in a small pond covering an area of perhaps rooo ft. The
surface was well covered with algae and floating débris of small sticks
and grass. This species was found floating on the water together
with S. catenaeformis and S. calospora. On May 2 all three species
were found in a state of conjugation interwined with each other. No
indication of S. inflata was found after May 12, although this pond
was visited frequently during the summer and fall.

Spirogyra quadrata (Hassall) P. Petit.—This is the Zygnema
quadratum of HassaLt. Perit gives the fruiting season as April-
May. The writer studied this species in eight different localities,
varying from a few rods to twelve miles apart, but under no great
variety of conditions. In every case it was found in quiet surface-
water pools or streams which were entirely dry by the middle of July.
All the eight places were in low pastures or meadows and specimens
were collected from among dead grass and leaves. S. quadrata was
found fruiting abundantly April 27, and within a week the eight
places mentioned above were visited and found in about the same
state of conjugation. By May 5 the Spirogyra at two localities had —
fruited abundantly and had passed the maximum of conjugation.
By May 12 this was true in every case and no fruiting material was
found after May 19. The temperature of the water recorded at each
visit to the different ponds Save a range of 10-25°C. It was very
difficult to collect any vegetative filaments of this species after the
period of maximum conjugation had passed.

: Spirogyra calospora (Cleve).—Woop found this species fruiting
in a ditch in a meadow late in April, and as late as May 25 in the

1909] COPELAND—PERIODICITY IN SPIROGYRA 17

“neck”? below Philadelphia. The season given by Petit is April—
May, and the habitat “ditches in low ground and swamps.” For
this species the present paper includes notes from observations of
five different ponds and one brook in different localities, and includ-
ing a greater variety of conditions than those described for S. quad-
rata. Early in April the plant in question was growing abundantly
and continued in increasing abundance until about the second week
in May. In two of the ponds the disappearance was by no means
rapid. A few scattering filaments were found May 6 and the last
June 4. The average temperature was 7-14°C. In the pond in
which most fruiting material was found the temperature ranged from
7 to 11° C at the time of maximum conjugation.

Spirogyra Hantzschii Rab.—In 1906 fruiting material was col-
lected first on April 28, and, with one exception, no fruiting material
was gathered after May 23. In the fall of 1905 this species was found
in active vegetative condition in three localities. Examinations were
made frequently during the fall and, when the ponds were free from
ice, during the winter. This species lived beneath the ice and in two
of the three localities the plant did not come to the surface at the time
of conjugation, but was collected from the débris at the bottom of the
ponds. Temperature of the water varied from 12-28°C. There
was no great range of habitat; generally in shallow ponds near the
mouth of a small stream, and more or less protected from the mid-
day sun.

Spirogyra mirabilis (Hassall)—This species is described by
Hassatt and also by Petit as producing spores without conjugation
and by no other method. Woe states that lateral conjugation is
more frequent than scalariform. I was not able to find this species
in fruit in the pond from which this material was taken. It was
brought into the laboratory in the vegetative condition April 2 and
within a week was fruiting in several aquaria. On May 7 a few
filaments with scalariform conjugation were found; all other spores
were formed in cells where no conjugation had taken place. Com-
paratively few cells formed spores and very few cells conjugated.

TIT gives March-July as the fruiting season. In the material
Mentioned all fruiting specimens had disappeared in all the aquaria
by the second week in June.

18 BOTANICAL GAZETTE [JANUARY

_ Spirogyra Jurgensit (Ktz.).—This species was found in fruit in
the neck of a small pond May 4, 1906. In the same locality were
found vegetative filaments of S. Hantzschii and Oedogonium, besides
sticks, dead grass, rusted wire, and tin cans. The algae mentioned
were found here in small patches January 8, and were visited at least
once each week until June, at which time no algae of any kind were
present. The fruiting specimens were most abundant when the
temperature averaged about 12°C., and lasted about a week. No

fruiting specimens were found after May 25. Prrir gives the con- |

jugating season as April-May.

Spirogyra catenaeformis (Hassall) Ktz.—This species, so far as I
know, has not been reported for this country, and is so placed because
it corresponds so closely to the descriptions in the works of HASSALL
and Petit. The latter author gives the fruiting season as April-May,
and the habitat “very humid ditches, wooded ponds, etc.”” The first
fruiting material was collected May 4 and the last May 22. It was
found in two localities about two miles apart. One was a shallow
ditch and the other was in cow tracks near a small stream. The
period of conjugation was the same for both localities. The tempera-
ture of the water ranged from 14 to 17°C. This species was found
with S. dubia in one pond and with S. inflata and S. calospora in
another pond.

Spirogyra varians (Hassall).—This plant was studied in five
localities from one to eight miles apart. In one case the water was
rather swift and the Spirogyra was fruiting wherever it happened to
lodge against a support. The other localities were shallow ditches
and ponds. The first fruiting material was collected May 2, and
the last May 23. In all localities except that in swift water this
species fruited abundantly. The average temperature of the water
was 8-12° C.

Spirogyra longata (Vaucher)—For this species my observations —

were confined to two ponds about one mile apart. In one S. longata

covered an area of perhaps 200°4‘t and in the other fully 100 sq. —

rods of surface. In both ponds the plant in question was well
exposed to bright sunlight, receiving its main protection from numer-
ous algae which were present. The temperature of the water ranged
from 20 to 25° C. and the depth of the water from a few inches to

EL OC

ee ee ee a ae

;
:
:
]
:
.
:
|
3

1909] COPELAND—PERIODICITY IN SPIROGYRA 19

several feet. VAUCHER says that this species conjugates from Febru-
ary 20 to April 12; Petir gives April-May. In the ponds mentioned
this species appeared in July and with increasing abundance until
early in August. The first fruiting specimens were collected August
2, and the last August 21, and neither vegetative nor conjugating
filaments as late as September 1.

Spirogyra Lutetiana (P. Petit)—I was not able to identify this
species except in a single locality. Its habitat was a small open ditch
in a meadow unprotected from the sun. PrtIT gives the fruiting sea-
son as March-May. The only date at which I found it in a state
of conjugation was October 14, and when collected it had all the
appearance of being old material which had fruited earlier. This
may have been a week and it may have been a month.

Spirogyra dubia Ktz.—Concerning this species Woop writes: “I
have found this species fruiting abundantly in May. When in this
condition it forms a mass of dirty yellowish-green.”” WoLLE agrees
as follows: “Fruiting abundantly in May; forming dirty yellowish-
green masses.’’ J have found this species in fruit in seven different
localities. The first date for each was as follows: May 1, May 2,
May 21, July 31, July 31, August 10, and August 10. Those fruiting
in May were found in rather sluggish streams and in water varying
in depth from one to four feet; those fruiting in July and August were
in cattle tracks and in a small ditch. |The conjugating period in the
spring lasted about four weeks; in the summer the period was about
two weeks. The temperature of the water in the former ponds varied
from 12 to 18° C.; in the latter places from 22 to 28°C. All seven
localities were situated where the plant was protected from direct
sunlight for a part of the day.

Spirogyra orthospira (Nageli)—I had an excellent opportunity
to watch the life-history of this species in a small pond of about
ie. Mh ates, two.to four feet deep, and well shaded with low
The surface of the pond was entirely covered with Spirogyra
to a depth of several inches. On May 22 it was found to be fruiting
abundantly both at and beneath the surface. It reached its maximum
about the second week of June, and by June 30 either vegetative or

Conjugating filaments could be found only by examining the débris at
the bottom of the pond.

alders,

20 BOTANICAL GAZETTE [JANUARY

General conclusions

In forming conclusions concerning whether any given plant passes
through a series of regular recurring phenomena, it is well to with: —
hold positive conclusions until the plant in question has been carefully
observed for a number of successive seasons. ‘This method of ap- ©
proach becomes more complicated, but at the same time safer, if the |
plant can be studied from a number of varying conditions of environ- —
ment. However valuable external conditions may be in effecting —
any given phenomenon, it seems well not to overlook whatever influ- —
ence may be due to internal conditions. There may be conditions
that cannot be seen, felt, weighed, or measured, that initiate and —
support even the simplest plants through a series of complicated
phenomena. Notes on Spirogyra for this region seem to credit much
influence to internal conditions. Here were forty different places
in which this plant appeared, matured, and disappeared, in a single
season. Without a single exception the disappearance was imme-
diately preceded by a period of conjugation. Special importance is
given to the fact that in no case did all of the filaments enter into —
spore formation either with or without conjugation. It is to be —
further noted that both the vegetative and conjugating filaments dis- ©
associated almost at the same time and disappeared, even from —
places where this plant was abundant over comparatively large areas
of surface. ; ‘oO
Of the twelve species studied, ten were in fruit in May, and —
in every case the maximum abundance of conjugation occurred within —
this month. In a few places more than one species was in fruit at the —
same place and time, but usually if several forms grew together the! a
conjugated at different times, their period of maximum conjugation -
succeeding one another at intervals ranging from one to two weeks. — :
For example, early in March, within a distance of forty rods, fout |
species of Spirogyra were found growing together at three different
Places at the edge of a slow brook. In two of the three places Spiro- - |
Syta was very abundant and covered an area in each of about 200%
P robably all had had the same source earlier: in the year during
high water, but remained distinct until all had disappeared in June. |
|

)

No fruiting material was found in either of the larger patches; tw0 t
Of the four fruited in the smaller one, but not at the same time —

ae a ee RE eee eT ee ee Or EN

Sg a

a OI 8 at a ta |

PH Se Ne EN PR NOOR PRM) ne = tg ee

1909] COPELAND—PERIODICITY IN SPIROGYRA 21

S. Jurgensit was found in fruit May 1-May 15; S. dubia was found
fruiting May 21—June 4.

Under the description of S. inflata mention was made of the fact
that on May 2 this species together with S. calospora and S. catenae-
jormis were found in fruit at the same time and place. _ It ought to
be noted, however, that S. inflata had passed its period of maximum
conjugation and that there were comparatively few filaments of
either of the other two species. No S. inflata was found after May
4, but from this until May 22, S. catenaeformis was abundant. In
giving a description of the latter species it was noted that this form
was found fruiting abundantly at the same time and quite near S.
varians. I did not find the two species intermingling, although grow-
ing abundantly almost in the same mass and having the same fruiting
season. :

I am inclined to think that not enough emphasis is given to the
fact that the lower plant forms do not have a continuous period of
growth, either vegetative or reproductive. In other words, the normal
period for either activity is continuous for only a few weeks or months
at most; it remains a problem of research to show how inclusively this
generalization applies to the freshwater algae. For Spirogyra the
observations reported in this paper seem to offer overwhelming evi-
dence in support of the view, for this region at least, that it is not a
‘perennial plant. Ihave only a single exception to this conclusion and
will speak of it later. A few of the species studied appeared more or
less abundantly in the fall; in this connection S. Hantzschii furnishes
4 specific example. In a majority of the forty localities kept under
observation during 1905-6, several forms of algae succeeded one
another in maximum abundance in the course of a year. In only
four of the forty places was any trace of Spirogyra found in the fall
where it had been in a state of conjugation in the spring. The man-
ner in which different species appeared, matured, and conjugated was
for most of the forms quite distinct and constant for the several locali-
ties. Following the fruiting season the disappearance of the plant
took place sometimes suddenly and sometimes gradually.

__ It is a well-known fact that vegetative material is very difficult to
identify; the same might as well be said of conjugating material.
For this reason it is difficult to follow any given species with certainty ;

22 BOTANICAL GAZETTE [JANUARY

where several closely allied species are associated, this is practically
impossible. Fortunately this did not occur except in a few places.
However uncertain and unreliable may be one’s observations and

conclusions relative to a given species during its period of vegetative _

growth, it is possible to determine with some certainty whether or not
the different species of Spirogyra have any definite or fixed periods
at which they conjugate, or whether this phenomenon takes place only
under certain conditions of environment. In all cases notes were
taken at each visit to a pond, and the determination of the species
left until the fruiting season; then by going back to my notebook, the
life-history could be worked out with some certainty. Leaving out
of account uncertain conditions such as the vegetative activities, and
placing special importance on the period at which conjugation takes
place, it is possible to concludé that one of the two important phe-
nomena (vegetative and reproductive activities) in the life-history of
this alga appears periodically, not due exclusively to seasonal condi-
tions nor to environment.

No one can say with absolute certainty whether the natural forces

that produce conjugation are internal or external. It is a fact,

however, that whenever a condition was present, in any part of a

mass, which was able to initiate and control reproductive activity, —

there was at the same time and place either the same or some other
condition which brought about the destruction of all vegetative
filaments. In this study no exception was found to this rule, whether
under laboratory or under natural environment.

Up to the time when conjugation was abundant in ponds, brooks,
and ditches, I still had over 300 aquaria in the laboratory. A small
percentage of them contained fruiting Spirogyra. The time of fruiting
for any given species began about one week earlier and lasted about
one week longer in the laboratory than that of the field. The disap-

pearance of the vegetative filaments was also more gradual. It was :
not difficult to keep aquaria early in the year, but after the middle of |

May they began to decline and were kept only with increasing care
until mid-summer, when out of 700 only one remained.

As to the different sizes of aquaria, it ought to be said that some

were ordinary test-tubes; some contained 2 5-30 liters; and the

majority ranged from one to ten liters. The results were not constant

a Pare

1909] COPELAND—PERIODICITY IN SPIROGYRA 23

for either the large or small sizes, although all sizes were usually
arranged at the same time. In dozens of test-tubes two-thirds full
of water or culture media Spirogyra kept well two to three months.
On the average, however, aquaria of four to ten liters were most satis-
factory. In three large aquaria of about forty liters each no successful
cultures were made, although frequently attempted.

An attempt was repeatedly made to grow Spirogyra in some of the
larger aquaria after a species had died in it and without renewing the
culture media. In every instance where this was tried the plant
died down in a few days, showing that the solution contained a toxin
injurious to Spirogyra. It is left as an open question whether a mass
of Spirogyra in nature dies from the accumulation of this same destruc-
tive element. ;

Aside from the twelve species herein described, there was still
another which has been kept under observation since early in the fall
of 1905. It is a large form and was found growing in rather large
bright-green patches in the running water of three different brooks
one to three miles apart. When first found, a few aquaria were
started, one of which has remained to the present time without any
indication of decline. This is an ordinary aquarium of about 10 liters;
it is kept well covered and for that reason very little evaporation takes
place. The main algal associate is Oedogonium, and a few brown
and green hydra have lived in the aquarium from the beginning.
This jar is the one exception to the 700 aquaria that died down during
the season of 1906. In concluding, I wish to call attention to the
fact that no trace of a conjugating filament has been found in this
aquarium; and furthermore, that none was found at any of the
three brooks mentioned, although in one case the plant was very
abundant. It is also remarkable that there has not been a time
from the first when vegetative specimens of this species could not be
obtained,

Tn addition it ought to be said that this study was continued

during the following year (1906-7), in the same locality, under about

the same conditions and methods of study. This second year’s

observations and experiments did not vary enough from those of

ze Previous year to warrant any change in the conclusions above
ven.

24 BOTANICAL GAZETTE [JANUARY

Summary

1. Few of those who have published papers on the genus Spirogyra
have given any notes on the subject of periodicity.

2. Out of 7oo aquaria it was found that the best way to arrange
a culture was to place some sterilized earth in the bottom, add some
dead leaves or dead grass, allow to settle well, then add.a small amount
of Spirogyra, and place at first in a window not exposed to direct
sunlight.

3- Of 300 aquaria in the laboratory at the time when Spirogyra
was fruiting most abundantly under natural conditions, there were
about.5 per cent. which contained conjugating material. The time
corresponded with that of the field.

4. Leaving out of account the number of species which may have
been collected in the vegetating condition and for that reason doubt-
ful of identification, there were at least thirteen different species
collected, of which twelve fruited more or less abundantly.

5. Ten of the thirteen species passed their period of maximum
abundance in May, one in August, and one in October. A few doubt-
ful fruiting filaments of S. Hanteschii were found a second time in
August. The only reliable example of a second fruiting period found
was S. dubia, which fruited in May, and again in small patches, but
relatively abundant, during the latter part of July.

6. When field-work was first begun, material was collected and

then brought to the laboratory for examination, This method was by

cop:

iat

Peust: eres!

Soe

1909] COPELAND—PERIODICITY IN SPIROGYRA 25

dition, both in the laboratory and in the field, since October, 1905,
but no trace of fruiting material has been found.

11. Notes taken in the field and supplemented by those in the
laboratory offer overwhelming evidence in support of the view that
the phenomenon of conjugation results not so much from external as
from internal conditions.

12. The writer therefore concludes that Spirogyra has definite
periods of growth and activity.

CLARK. UNIVERSITY
WORCESTER, Mass.

LITERATURE CITED

I. BENNETT AND Murray, A handbook of cryptogamic botany 266. London.

1889.
. Fritscu, F. E., Problems in aquatic biology, with special reference to the
study of algal periodicity. New Phytol. 5:149-169. 1906
Hassat1, A. H., A history of the British freshwater algae. London. 1857.
Hoyt, W. D., Periodicity in the production of the sexual cells of Dictyota
dichotoma. Ror. GAZETTE 43: 383-392. 1907.
. Petit, P., Spirogyra des environs de Paris. 1880.
VAUCHER, J. P., Histoire des conferves d’eau douce. 180
West, G. S. anp W., Observations on the Conjugatae. “Annals of Botany
12:29. 1898.
Wituams, J. Liovn, Periodicity in the sexual cells in Dictyotaceae. Annals
of Botany 19:533. 190

9. Woon, H. C., Coabaues to the history of the freshwater algae of North

America, Sinithsoinian Report. 1874.

10. Wort, F., Freshwater algae of the United States.- Bethlehem, Penn. 1887.

Ss]

: ae x

o won

ON THE POLLEN OF MICROCACHRYS TETRAGONA
ROBERT Boyp THOMSON
(WITH PLATES I AND It)

“Microcachrys tetragona occurs only on the highest summits of the
Western Range and Mount Lapeyrouse in Tasmania. It was intro-
duced to the Royal Gardens at Kew about the year 1857 by Mr.
Wit1Am ARCHER, on whose property it grew. Although of great
interest in a botanical sense, its only value as a garden plant is for
conservatory decoration, for which the elegant habit it can be made
to assume under pot culture, its neat foliage, and bright-red fruits
render it highly suitable.”* The specimens for this work were ob-
tained through the kindness of the present director of Kew Gardens,
Colonel PRAIN, whose courtesy and that of the staff is much appre-
ciated by the writer.

The general features of the fruiting branches of microsporangiate
and megasporangiate plants are indicated in pl. I. The cones ar
borne terminally and their sporophylls are spirally arranged, in
contrast to the opposite and slightly concrescent vegetative leaves
(pl. I, and $l. IT, fig. 1, the branch to the left).

The microsporophylls bear two somewhat spherical, pendant spo-
rangia (fig. 2), whose form and structure, after the discharge of
the pollen, is indicated in figs. 3 and 4. The terminal scale of the
sporophyll is broadly triangular in outline ( jig. 2), very much extended
dorsally (fig. 3), but not, or scarcely at all, in the ventral direction.
The inner layers of the wall are much collapsed at the stage indicated,
but the epidermis retains a very definite structure on account of its
peculiar thickened bands. These are shown in transverse and in
longitudinal section in jigs. 5a and 5b.

In fig. 4 some linear and branched structures are apparent in and
around the sporangia. These are the hyphae of a fungus, and to
their presence the retention of many of the pollen grains in the
dehisced sporangia of my material is due. ‘The hyphae are often

* Kent, Apotpuus H., Veitch’s manual
to the discovery of this form and its descriptio
Botanical Gazette, vol. 47]

of Coniferae 161. 1900. The references
n are given here.

[ 26

1g09] THOMSON—POLLEN OF MICROCACHRYS 27

in intimate association with the pollen, they frequently branch, and
the branches form contact with the grains. This pollen is no different
from the less abundant found in uninfected sporangia.

The pollen of Microcachrys (fig. 8) is small as compared with
that of Saxegothaea (jig. 9), Podocarpus (fig. ro), or Pinus (fig. 12),
all these figures being the same magnification for the purpose of
comparison. The grains vary considerably in size, a feature which
my material of Saxegothaea also shows. They are winged, though
in some instances I have thought that no wings were present, but
more material was needed to determine this point than was at my
disposal. The wings arise in the ordinary way by a separation of
the exine from the intine. In longitudinal section the grains usually
show two rather poorly developed wings (see various grains in fig. 6),
and in following the series these two are often all that one can be
certain of. In gross material, however, three Wings are readily
apparent (jig. 8).2_ In transverse sections, also, which pass through
the ventral part of the grain, the three-winged condition is clearly
seen (see middle of jig. 6). In some cases one of the wings is very
small (jig. 6, grain in upper part of the field). The lowest grain in
jig. 6 shows the presence of four wings (the central one very dark).
Some four-winged grains have one pair of wings much smaller than
the other. Exceptionally, five and six-winged grains are found. A
student, Mr. W. P. THompson, kindly made a careful determination
of the number of wings in gross material from several cones, rolling
the grains in fluid under the cover glass. He found that of sixty-four
grains, fifty were three-winged, three of these having one very small
Wing; nine were four-winged; two had five, and three had six wings.
The pollen studied was in the mature condition and the wings must
have been fully developed. Fig. 7 shows the greatest expansion of
the Wings that I have observed. They usually do not extend laterally
beyond the body of the grain (jig. 8, also some grains in fig. 6).
They project, however, beyond the body of the grain ventrally,
though not to such a degree in either direction as do those of Podo-
carpus (jig. 10) or Pinus (fig. rr). A small amount of material from
the apex of one of the cones showed another feature. When the

ot re I studied the grains in section first, and this led me to misinterpret the number
© wings, as stated in Bor. GAzETTE 46:465, 466. 1908.

28 BOTANICAL GAZETTE [JANUARY

pollen grain has a five-celled gametophyte, the wings are very small,
not much larger than they are in Pinus when its pollen is still in the
tetrad condition, The wings of Microcachrys arising thus late onto-
genetically give indication of their recent acquirement.

With regard to the microgametophyte, I find that in the mature
condition of the pollen four prothallial cells are often present, though
three are more usual. When three only are present, it is the second
prothallial cell that has divided (fig. 7). The first and second
prothallial cells have walls which turn blue with chlor-iodide of zinc.
No further cellulose walls were demonstrated. Fig. 7, in addition
to the prothallial cells, shows a lateral derivative of the body cell and
the tube nucleus. The gametophytic structure thus conforms to
that which recent investigators have shown characterizes the related
forms. ' Perhaps it is not out of place here to give credit to THrBoUT,?
the first person, so far as the writer knows, to describe and figure a
multicellular gametophyte in the Podocarpeae.

NoreNn in his recent work on Saxegothaea* has emphasized the
relationship of this form to the Araucarieae. He finds that, in addi-
tion to the supernumerary prothallial cells, there is a curious remi-
niscence of this group in the mode of pollination, the pollen sometimes
being deposited in the cavity around the ovule and growing over the
tissue into the micropyle, a condition which so far as can be judged
is essentially similar to that which I have found in Agathis. He also
calls attention to the wingless condition of the pollen in both Saxego-
thaea and the Araucarieae.

With the present contribution our knowledge of the occurrence
of excess prothallial tissue in all genera of the Podocarpeae is com-
plete. The winging of the grain in Microcachrys is of varied and
indefinite character, late in development, and undoubtedly of primi-
tive type, the form occupying in this respect an intermediate position
between Saxegothaea on the one hand and Dacrydium and Podo-
carpus on the other. This is the more evident from the presence of
three wings in some species of Podocarpus. Turpout describes. the

$ THIBOUT, E., Recherches sur Vappareil m4le des Symnospermes. pp. 265. is.
16. Lille. 1896. See pi. 14, figs. 8 and 9, of Podocar pus polystachya.

4 Norén, C. O., Zur Kenntnis der Entwicklung von Saxegothaea cons picua Lindl.
Svensk. Bot. Tidskr. 2: ror—r29. pls. 7-9. 1908.

BOTANICAL GAZETTE, XLVII PLATE I

THOMSON on MICROCACHRYS

BOTANICAL GAZETTE, XLVII PLATE II

THOMSON on MICROCACHRYS

ee ee Ne Ee TON

PF SE a ay ae Fee RR aT as ne nee ea

iin

PO ge oP aa ee eT,

1909] THOMSON—POLLEN OF MICROCACHRYS 29

pollen of P. dacrydioides as having normally three wings (op. cit., pl.
14, figs. 10,11). I have found three wings present exceptionally
in the usually two-winged P. ferruginea (fig. 12). Unfortunately, no
data nor material of the more probable genus Dacrydium were
available. ;

The bi-winged condition of the pollen on the pines and podocarps
has often been referred to as an indication of affinity between the
groups. This view has no longer support, since the winging of the
grain has arisen in the Podocarpeae within the group itself, and so
is distinct from that of the pines. In a further contribution on the
character of the megasporophyll of Saxegothaea and Microcachrys,
the writer hopes to show the essential difference of this structure in
the two great phyla of the conifers.

UNIVERSITY OF TORONTO

EXPLANATION OF PLATES I AND I
PLATE I
Twigs from staminate and ovulate plants. X2.25.

PLATE II

Fic. 1.—Branch with microsporangiate cones. X4.

Fic. 2.—One of the same enlarged; the sporangia can be seen in pairs
beneath the terminal scale of the sporophyll. Xo.

Fic. 3.—A microsporophyll in longitudinal section, to one side of the axis.

Fic. 4.—Same in transverse section; the linear structures in the sporangium
are the hyphae of a fungus shown in contact with pollen grains in fig. 6.

Fic. 5¢.—The wall of the microsporangium in tangential section.

Fic. 55.—The same in transverse section.

Fic. 6—A field of sectioned pollen grains; below is one with four wings;
above it one with three wings cut transversely and a small part of the body
uniting these; various other aspects of the wings.

Fic. 7—The microgametophyte.

Fics. 8-11.—Pollen grains at same magnification: fig. 8, Microcachrys,
from below; fig. 9, Saxegothaea; fig. 10, Podocarpus ferruginea (a and 6 two latera
views at right ‘angles; c, dorsal view); fig. 11, Pinus resinosa.

Fic. 12.—A three-winged grain of Podocarpus ferruginea.

A VEGETATIVE MUTANT, AND THE PRINCIPLE OF
HOMOEOSIS IN PLANTS
CONTRIBUTIONS FROM THE AMES BOTANICAL LABORATORY, NO. 9
ROBERT GREENLEAF LEAVITT
(WITH NINETEEN FIGURES)

When viewed in their relations to morphogenetic and_ broader
evolutional problems (as distinguished from narrower phylogenetic,
or genealogic, problems) numerous facts of teratology—a descriptive
cult without unifying principles, heretofore pursued chiefly by the
vaguely curious, and lending itself discreditably with equal readiness
to either side of many a morphological discussion in the past—mani-
fest a special and high value. The production of form from form-
lessness in the egg-derived individual, the multiplication of parts and
the orderly creation of diversity among them, is an actual evolu-

It is this relation, to the central rather th
lems of evolution, which has le
upon abnormalities in plants.
the bearing which some of the

an the peripheral prob-
d me to carry out certain observations
Yet I shall try to make clear, also,
observed individual variations have

1909] LEAVITT—HOMOEOSIS IN PLANTS 31

The present paper is sequel to a casual observation made several
years ago, when as I was passing near a horsechestnut tree in the autumn
I noticed, upon a depending branch, a leaf which had lost several of
its leaflets. I found that the palmately compounded leaves of this
species fall to pieces as they are cast off by the tree, or even before
this event. The leaflets are removed as if by a clean cut; and some-
times the petiole, quite devoid of blades, is left standing alone upon
the stem.

This complete dismemberment seemed to me a curious thing.
Abscission at the base of the petiole is not a simple decay, but is, as
everyone knows, a somewhat complicated process, wherein there is
formed across the leaf-stalk, through suitable cell divisions, a plate
or layer of cells, by the disintegration of which the leaf is ultimately
allowed to fall away. The cell walls of the scar-surface become
suberized, so that the wound is sealed against the loss of water.
Leaf-abjection is thus a complex adaptive process.

The question arose, Why should the abscission proper to the
petiole-foot be repeated in all of its details at the bases of the petio-
lules ?—for I ascertained that the process is carried out in full at
these points in this species. Of what use to the tree is it that the
frame of the leaf, now emptied of its valuable contents which have
been withdrawn into the stem, should be carefully disarticulated? I
have never been able to imagine any utility nor have those to
whom I have propounded the riddle suggested any. I believe that
in fact the habit is neutral in the tree’s economy.

If this is so, the evolution of absciss-layers in petiolules cannot
be referred to natural selection. For in the plan of evolution by
the accumulation of variations under the guidance of natural selec-
ion, developing organs must pay their way as they go.

An explanation of the presence of the useless structures occurred
to me. It seemed probable that the absciss-layer perfected in the
natural course of evolution at the foot of the leaf-stalk had, sub-
sequently to the compounding of the blade in this species, been
transferred, So to speak, to the bases of the petiolules. The disartic-
ulations of the leaflets seemed to be a series of ecologically meaning-
less echoes of the primal, useful disarticulation at the junction of stem
and petiole.

32 BOTANICAL GAZETTE [JANUARY

The horsechestnut is not peculiar in this respect, and further con-
siderations, with regard to compound leaves generally, have confirmed
in my own mind my first understanding of the matter. Disarticula-
tion of the leaflets seems to be universal in deciduous-leaved species.
It occurs in many families separated in a natural system by entire-
leaved groups, as in Juglandaceae, Berberidaceae, Anonaceae,
Rosaceae, Rutaceae, Sapindaceae, Vitaceae, Oleaceae, Bignoniaceae,
Caprifoliaceae. That is, it must have arisen in evolution many times
independently. There must, then, be some wide underlying prin-
ciple capable of bringing the disarticulation in question to light,
wherever compounding of the blade establishes the conditions for its
operation. This principle seems, as has been argued above, not to
be natural selection. There is, however, in plants a recognizable
principle of morphic translocation, mobility of characters, or homoeosis

.(BaTEsoNn), to which the phenomenon may very well be referred."
The conditions favorable to its operation in this case are very obvious,
since the structural relations at the junction of the petiole and stem
are imitated at the junction of petiolule and petiole (or rhachis).

The Pierson fern

Recently evidence corroborating the above interpretation has
come to hand in a case where advance in complexity has been all
but observed. In the title I have alluded to it as a case Of mutation;
but perhaps I am using the word somewhat loosely, if the DeVriesian
sense is to be insisted upon. The plant is the Pierson fern, now well
known to horticulturalists and to the public generally. The meta-
morphosis of the Boston fern, to which the Pierson owes its origin,
seems not to have been hitherto described as a homoeotic trans-
formation; yet such it is.

The first plant of the variety was found while it was still small,
among vegetatively propagating Boston ferns in the greenhouses of
Mr. F. R. Pierson, the well-known horticulturalist of Tarrytown-
on-Hudson, N. Y., in 1900.2 While it was not a “pedigreed” plant,
* My first paper on this topic, entitled “On translocation of characters in plants,”

Ora 7314-19, 21-31. 190s. en it was written I did not know
i ed the same subject in relation to variation of ani
and had devised the name homoeosis. I find his term applicable to the present pur-
pose and more convenient to use than my own expression.
? Mr. F. R. Prerson, ina letter to the writer.

1909] | LEAVITT—HOMOEOSIS IN PLANTS 33

its derivation is patent, for when its descendants are grown under
poor conditions, they generally revert temporarily, in a part of their
fronds at least, into the primitive state, which is seen to be that of
the Boston fern. This constitutes highly desirable proof of parentage.
Whether the sudden alteration of character took place in a spore, or
in a bud from a runner, is not surely known. Nevertheless the cir-
cumstances leave little doubt on this head; for the Boston fern, while
richly stoloniferous, is almost universally sterile. Mr. Pierson
believes the original individual to have been a bud-sport. The
young plant caught the eye of a gardener, who brought it to the
notice of Mr. Prerson, and it was set aside so that its development
might be followed. All Pierson ferns and numerous varieties have
descended from this individual.

In this case we may say that the transformation, which affects
the leaves of the fern and consists not in the introduction of alto-
gether novel outlines or
proportions, but rather in
the relocation of forms
already in existence, has
taken place under observa-
tion. A single individual 7
has been noted to emerge
from the common mass—
under cultivation as an
isolated colony—with very
markedly dltered thasactets. Fic. 1.—Pinna of Boston fern.

New features have not been slowly developed through several
Senerations, but have appeared suddenly.

The Boston fern has simply pinnate fronds. The pinnae are
fntire, serrulate-margined, oblong, and usually bluntly toothed or
auriculate on the upper margin at base (fig. 1). In the new plant,
the pinnae have become divided,3 and at the same time elongated—
= much elongated. Their divisions, the pinnules, are oblong and,

€n fully developed, toothed or auriculate on the upper margin at

(fig. 2). The pinnules, in fact, are very good copies of the

3 .
sei: it the new form is fully expressed. The division may affect only part of

34 BOTANICAL GAZETTE [JANUARY
original pinnae; and if we lay a well-developed pinna of the Pierson

fern by the side of a small frond of the Boston fern, we shall be
struck with the fact that they are almost identical. In short, the

f\
:

Fic. 2.—Pinna of Pierson fern.

pinnae of the new variety are transformed from the original condi-
tion in such a way that the plan of the whole frond of the Boston
fern is now seen in the primary segments of the Pierson frond.

The completeness of the
imitation may be indicated
still further:

1. The pinnae are elon-
gated, the growth being less
strictly determinate than that
of the original pinnae, and
more like that of the frond.

2. The apical growth of the
pinnae is now circinnate, as
: shown in fig. 3, after the
fashion of the original frond. The pinnae of the Boston fern on
the other hand, while lapped together in the young state at the
summit of the developing frond, are not in the least circinnate.

3. The blades of the pinnae are divided.

4. The outline of the pinnules is like that of the original pinnae.

5. Finally, the peculiarity of the original form, that its pinnae in
age become disarticulated by the development of an absciss-layet;

—Growing apex of pinna of Pierson
a (ea and Boston fern (right).

|
,

1909] LEAVITT—HOMOEOSIS IN PLANTS 35

and ultimately fall off after several layers of brown tissue have been
developed on the scar surface-to-be, is now shown by the pinnules
of the Pierson fern. The pinnules of the latter are discovered to be
deciduous, and a minute investigation shows that deciduity is secured
by the development of an absciss-layer at the base of the pinnule,
with the formation of brown scar-tissue. Thus the translocation of
the structures concerned in disarticulation of leaflets is in this case an
ascertained occurrence: these structures have passed at once and
unaltered from the bases of the pinnae to the bases of the pinnules.

The beautifully soft and luxuriant aspect of such ferns as the
Whitman‘ is due to the fact that each frond is in effect made up of

Fic. 4.—Pinna of Whitman fern.

Many small fronds. In this last-named variety, and in some others,
as Pierson’s elegantissima and superbissima, homoeosis has gone one
step farther than in the Pierson, so that we find a thrice-compounded
leaf (jig. 4), and the segments of the third order have the frondlike
character—even to the circinnate apical growth.

To return to the matter of normal casting of leaflets, with which
We began. The fact that in an instance now before us positive evi-
. the translocation of absciss-layers, etc., from one part of
a setae has been secured, strongly corroborates the infer-
ate er € on more speculative grounds, that leaflet-abjection
0 be understood as an imitative or repetitive phenomenon.

ihe . originated in the greenhouse of H. H. BARROWS & Son,
Sport from the “aa a gt stesessi dee upon a runner of the Barrows fern (itself a

. trom H, H, Barrows. & Son to the writer.

36 BOTANICAL GAZETTE [JANUARY

The principle of homoeosis

But whether or not the transference of absciss-layer from leaf to
leaflets in this fern verifies in any material degree the idea of a trans-
locational origin for leaflet-abjection, it alone, were there no other
available demonstration, would suffice to establish the important
truth, that a character perfected in the course of evolution under one
relation in the plant body may make its appearance suddenly under
another relation and in a region of the body to which it is not native.
Here is a principle of wide import in morphogenesis, and not without
bearing—as has just been shown—upon some of the minor problems
of plant evolution; concerning which principle some further obser-
vations may be offered.

The described involution of frond-plan belongs to one among
several categories of homoeosis all subsumed under the larger type,

but of this type it is a rather com-
plicated example. A simpler illus-
tration will make the nature of the
general principle clearer, and at the
same time exhibit its distinctness
from another aspect of morphogene-
sis with which it is frequently con-
fused—that of reversion.

In 1906 there was discovered in
the county of Dorset, in England, a
solitary plant of Platanthera viridis,
the flower of which was remarkable.
s, with three spurs. for the possession of three spurs
(fig. 5), one under the lip in the

normal position, and one on each of
the lateral sepals.’ The extraordinary spurs entirely resembled the
normal one. The general conformation of the flower was very
nearly that of the species, though slight modifications were observable
throughout.

The conditions of the occurrence make it highly probable that
we have here the offspring of normal parents, rather than one mem-
ber of a long series of gradually deviating forms; for in a botanically

5 HEMSLEy, W. B., Journ. Linn. Soc. 38:3. 1907.

Fic. 5.—Abnormal flower of Pla-

tanthera viridi
After Hemstey

1909] LEAVITT—HOMOEOSIS IN PLANTS 37

often- and well-explored country a race of deviants undergoing a
gradual evolution from the typical condition of the species could
hardly have escaped observation. It is necessary to suppose that
the change of floral structure, in the line of descent embodied in
this individual, supervened suddenly. The supernumerary spurs
came into existence fully formed. They are not, however, new struc-
tures, except when regarded as sepallary appendages, and their
origin morphogenetically and historically considered is easily dis-
covered. The spur elaborated and perfected by the usual evolu-
tionary processes—whatever the “usual” processes may be—as an
appendage of the lip, has in a moment been transferred in full char-
acter to parts which heretofore have not been concerned—namely, to
the sepals.

The principle is not new in biology. More than a century ago it
began to be observed that when a part of such an animal as a worm
is cut off, a new part different from that removed, but like some
other part, may grow from the cut surface.6 Bonner believed that
there are special germs for the development of the various organs;
as head-germs and tail-germs. From his experiments he made cer-
tain inductions concerning the distribution of these germs. LOEB?
has investigated the conditions under which substitutions for lost
Parts may be induced, this process being regarded by him as essen-
ually different from regeneration and deserving the distinct name
heteromorphosis. Logs also discovered that in some cases hetero-
Morphosis can be produced without any organ being cut off, or any
Wound being inflicted upon the animal. Other investigators have
dealt with the matter on experimental lines, especially with reference
‘0 the physiology of the process. WEISMANN has independently
argued the translocation of characters from segment to segment in
mia a ot some insect larvae. He uses the word hetero-
fee : Pepnection, Z BATESON, who appears to have discovered
es wg e€ ms an original way in his study of variation, gave it the

Sfosis.° He describes cases in several phyla; for example,

° Bonnet, Cy
287. i749, » ©H., Oeuvres d

: Lozs, YE Studies
ae

histoire naturelle et de philosophie 1:191, 215; 3:

: : in general physiology 115, rgr, 627. 1905.
hg theory (transl. by THOMPSON) 1:365, 367.
€rials for the Study of variation 85. 18094.

° Ma

38 BOTANICAL GAZETTE [JANUARY

in Arthropoda the development of the extremity of a sawfly’s antenna
in the form of a foot; and in Vertebrata, the not uncommon anoma-
lous presence upon the under side of turbots, etc., of the pigment and
the tubercles proper to the upper side.

An interesting case in animals, which has lately come to my
notice, is furnished by a crab, the claw being modified as shown in
jig. 6. Thesuperfluous claw, developed
upon the normal dactyl, repeats in detail
the style of the larger claw, as regards
the general form and even the teeth.’®
The formation is believed not to be
congenital, but to result from a wound
to the normal claw near the time of

Fic. 6.—Crab claw, showing ‘
homoeosis. moulting.
: The presence of the principle in
animal development greatly enhances the interest which the botanical
student of morphogenesis must entertain with respect to homoeosis.

In plants the transposition of organs has frequently been described
in teratological works; but the phenomena of this class have been
confused with those of a different nature, and the larger relations of
homoeotic formations have not been recognized by teratologists.
MASTERS devotes a short chapter to the subject, under the caption
heterotaxy**—an expression which, as used by him, seems only in
part to cover the ideas I am here endeavoring to formulate. MAs-
TERS’ chapter is a miscellany of teratological facts which will not go
conveniently into any other chapter of his compendium. His term
metamorphy"? is too general for the present purpose, being employed

ro Photograph in St. Nicholas Magazine 25:177. 1907. The specimen is in my
possession. A number of similar anomalies have been described by W. Faxon, Bull.
Mus. Comp. Zoél. Harv. Coll. 8:257. 1881; by F. H. Herrick in “The American
lobster,” Bull. U. S. Fish Com. 1895, p. 145, pl. 47; and by other writers of whose

work a summmary appears in BATESON’s Materials. ‘These authors do not specifically
recognize the homoeotic nature of the anomaly.
tt Vegetable teratology 156. 1869. |
ex It will be remembered that GorTHE employed the word metamorphosis in 4
different sense. Sacus, again, has a definition: “Metamorphosis is the varied develop-
ment of members of the same morphological significance resulting from their adapta-
25 definite functions.”—Sacus, Text-book of botany, tr. by Bennett & DYER
1875), 131.

1909] LEAVITT—HOMOEOSIS IN PLANTS 39

“to distinguish cases where the ordinary course of development has
been perverted or changed” (p. 241). This author’s Vegetable tera-
tology, while it is a rich storehouse of facts for the study of homoeosis,
is—as might be expected from the period of its publication—entirely
wanting in illuminating discussion and appreciation of relations to
the problem of organic evolution such as are to be found in BATESON.
The latter author recognizes the presence of homoeosis in plants.*3

Sacus distinguished clearly between atavistic formations and those

which are merely translocational. His explanation of the aberra-
tions now being considered would indeed give a degree of literality
to the expression mor phic translocation, since he attributes them to
changes in sap movement, with disordered nourishment and abnor-
mal distribution of the formative stuffs.t4 Prnzio’s perception of
the real nature of the anomalies now under discussion is clear.*s

Modes of homoeosis in plants

In the vegetable kingdom homoeosis has many interesting phases,
some of which I may indicate. The facts being superabundant, it
1s well to begin to categorize them, not for mere convenience, but in
order that new and possibly suggestive points of view may be won.

I. The translocation of characters may be acropetal.

Everyone recalls numerous instances where details of the foliage
leaves—toothing, texture, hairing, etc.—have appeared in the floral
leaves. A case of acropetal translocation is that of the corolla fringe,
Peculiar to Gentiana crinita, from its normal site to the summit of
the carpels.76 4 remarkable transposition has more than once been
noted, of the secretory hairs (tentacles) of Drosera foliage to the
Sepals, petals, and even interior of the carpels.'?

2. The transference may be basipetal.

A igs and sepaloid characters—color, texture, outline—not
Tequently appear in the involucre, and sometimes the effects reach
Even the foli

age. The most remarkable case that has come to my
- BartEson, Op. cit. 111, 570.
4

488, 9g “i Form der Pflanzenorgane, §3. Arbeiten Bot. Inst. Wiirzburg 2:452-
Me ra Gesam. Abhandl. Planzenphysiologie Bd. 2:1159-1231. 1893.

ae €n-Teratologie 23335, 489, etc. 1890-1894.

: ae Rhodora 7:14. 1905.

7
Odora, 2. ¢.; PLANCHON, Ann, Sci. Nat. Bot. III. 9:84, 86. pis. 5, 6. 1848.

40 BOTANICAL GAZETTE [JANUARY

notice occurs in a hybrid of the saffron (Crocus sativus). Stigmatic
characters in this plant have wandered so far that not only the anthers
and floral bracts, but also the foliage leaves are sometimes surmounted
by portions of clearly characterized stigma, furnished with papillae;
and even the sheaths below the leaves are tinged with saffron color."*

3: Translocation may be lateral.

Peloria, whether regular or irregular, illustrates this form of
homoeosis. In the well-known Uropedium Lindenii (Selenipedium
caudatum, Orchidaceae) the form of the remarkably long pendent
petals invades the labellum, entirely subduing the saccate character
of that member and converting the strongly zygomorphic into an
actinomorphic corolla.*® Contrariwise, the irregular member may
impose its form upon the remainder of the corolla, as in the well-
known peloric Linaria vulgaris.

4. The invasion of migrating characters may be partial in any
degree.

The curious modifications of the pistil and its contents in abnormal
Drosera intermedia illustrate this truth in several different ways.”°
As vegetative influences begin to evince their presence in the flower,
the first modification of the gynoecium is an elongation of the ovary,
especially its lower part. The carpels, while still concrescent, ovulif-
erous, and terminated above by the usual stigmatic apparatus, in a
slight degree respond to the-influx of foreign morphogenetic forces
by the lengthening of their basal portions, corresponding to the
petioles of foliage leaves: the carpels sensibly approach a petiolate
condition. In flowers yet more affected the carpels separate in.
various degrees, finally falling apart altogether; their bases become
elongated to form true petioles; the blades become more and more
complanate and spathulate; the ovules after suffering a series of
changes are finally replaced by tentacles; and the styles and stigmas,
still retained even when the carpel-foliage-leaves spread horizontally
from the center of the flower in rosette fashion, lose much of theif
rightful character. Thus is the gynoecium transformed step by
step into a whorl of leaves. The metamorphosis of each part of

*8 CHAPELLIER, P., Journ. Royal Hort. Soc. 243277. Igo0.

*9 LINDEN, L., Pescat. pl. 3; Retcus. r., Xen. Orch. 1: pl. 15.
20 PLANCHON, /. c. 86.

1909] LEAVITT—HOMOEOSIS IN PLANTS 41

the carpel is marked by the same flexibility and miscibility of charac-
ters which pertains to the whole gynoecium. In ovaries affected by
the anomaly, but not yet resolved into separate carpellary leaves,
ovules become stalked, the integuments may compose a small cup
from the edge of which a few tentacles arise, or may be converted
into minute modified discoid or concave leaf-blades bearing numerous
minute tentacles. Between these blades, representing many charac-
ters of the radical leaves of the plant, and apparently perfect ovules,
many stages of gradation have been observed. Similarly the stigmas
and styles, in the most aberrant carpels, combine with their own
characters the structure of marginal tentacles.

The accompanying jig. 7 represents advancing petaloid modifica-
tion of the calyx of Ranunculus bulbosus. The normal sepal A is
green and hairy on
the back, while the
petal is glabrous and
yellow. Sepals B,C,
and D are progres-
sively invaded by
yellowness and glab-
Tousness (and doubt-
less by other corolla
characters), until in

in Ranunculus

Fic. 7.—Progressive |
bulbosus. After O. AMEs.
D all that remains of sepal nature is hairiness and slight greenness
in the median line of the back.

The intermediate formations described here, which are of very
Common occurrence in plants, are the “anamorphoses”’ of CELA-
KOVsky.

5- Migrating characters may transgress boundaries of homology.

€ term “homologous,” applied to organs supposed to have had
é common evolutionary origin, is very often vague and not rarely is
Practically meaningless for the reason that, while degrees of homology
are Infinite, the specific degree of homology in many cases cannot be
ss even approximately, In spite of this vagueness in the word,
‘ici aa. ponies will probably agree that the ing is in =
Stamen, nor of ice bat a wg pepe rel Cee ‘Yet

42 BOTANICAL GAZETTE [JANUARY

the fundaments of ovules have been observed to develop into all of
these structures. LerrcEB reported?! a Marchantia polymorpha in
which cupules of various degrees of development (presumably with
gemmae) took exactly the places of archegonia. Yet cupules and
archegonia are not homologous.

6. When against homology, the invasion may be partial in any
degree.

Without entering into details, the literature of ovular abnormality
may be cited in support of this opinion.2?_ Even when the imitating
and imitated structures are known not to be related historically, the
presence in any organ of form-giving factors derived from a different
organ may merely be indicated by some faint resemblance, or may
be pronounced in any degree up to the total replacement of the local
member by the foreign.

A corollary of the above propositions (5 and 6) may be stated:
Homology of two organs cannot be inferred on the ground alone of
the replacement of the one by the other, or of the translocation of
features proper to the one into the other of the organs. The exist-
ence of even a so-called “complete” series of gradations between
two members does not necessarily imply the homology of the mem-
bers. I think that the anamorphoses of CELAKOvsKyY have not the
extraordinary phylogenetic significance, as compared with other
kinds of abnormality, which this author has attributed to them.

7. In homoeosis characters may pass from one to the other of the
alternating generations.

YAMANOUCHI has recently published? an account of the very
interesting conditions attending apogamy in Nephrodium molle. The
apogamous prothallia bear no functional archegonia, but they pro-
ceed to initiate the sporophyte directly. A superficial cell on or neat
the cushion divides perpendicularly to the surface, and underlying
cells divide in various directions. From the complex so originated,
the sporophyte develops. The cytological history being followed

t Bot. Zeit. 332747. 1875.

22 WIGAND, A, Grundlegung des Pflanzen-Teratologie 39. Marburg. 1859;
MASTERS, op. cit. 186, 262-271; WyDLER, Denkschr. Regensb. Bot. Gesells.
LA 1859; PLANcHoN, article cited; CELAKovsky, Bot. Zeit. 332129-177- 1875:

23 Bot. GAZETTE 44:142. 1907; 45:289. 1908,

1909] LEAVITT—HOMOEOSIS IN PLANTS 43

throughout the critical stages, no nuclear fusions were found; and as
a matter of fact the chromosomes of the sporophyte continue to be
of the reduced (or x) number. Here we have a sporophytic form
imposed homoeotically upon a gametophytic cell basis.

The contrary or aposporic case in ferns has been studied in detail
by several workers. In Athyrium Filix-foemina, BowEr?4 found
the sporangia arrested at various stages, the development being carried
on by prothalloid growths which organized typical wedge-shaped
cells at one or more points on their margins, while rhizoids were
formed at the same time by the outgrowth of individual cells.
Drurry’s had already observed sex-organs and even young plants
arising from these prothallia. Woraston?® found in Polystichum
angulare that the tips of pinnules were converted directly into pro-
thallia, bearing archegonia and antheridia. In these cases a game-
tophytic form is imposed homoeotically upon a sporophytic cell
basis (with little doubt; though the cytology, I believe, has yet to be
worked out).

Entirely parallel is the production of protonemata from the
setae of mosses, as observed by PRINGSHEIM?” in Hypnum and
Bryum, and by Srant’® in Ceratodon. LANG?9 has demonstrated the
power of Anthoceros capsules to produce thalli aposporically.

The adventive structures are, of course, gametophytic, yet they
arise by the prolification of sporophytic tissues.

8. When the boundary between alternative generations is so trans-
Sressed, the invasion of extraneous characters may be partial, and both
Senerations may be represented side by side in the same body.

GOEBEL° has recently induced the formation of prothalloid
stowths from cut fronds of very young plants in several species of
ferns. Some of the aposporous prothallia bore stomata, as well as
Sex organs (p. 132). LANG3" found sporangia in various stages of

*4 Journ. Linn, Soc. 21 2360. 1886,

*S Ibid. 213354. 1886,

oi Teported by BowER, /. c. 362.

a aoe ane. Wiss, Berlin, July 10, 1876,

34:689. 1876,
% Annals of Botany 153503. rgor.
“longetas gee Wiss. 3'7:119-138, 1907.
i + NOY. Soc. B. 1902194. 1898,

44 BOTANICAL GAZETTE [JANUARY

perfection, produced in groups having a superficial resemblance to
sori, upon prothallia of Scolopendrium vulgare ramulosissimum which
had been growing for a long time without fertilization. Lowe’s
cultures of Scolopendrium vulgare exhibited the combination of alter-
native generations in an interesting manner.3? “We have here
: consequently a posi-
tion as nearly as
possible intermediate
between _sporophore
and oophore, the
sporophoric character
distinctly existing in
the shape of a circle
of stalked fronds gen-
erated spirally from a
regular axis of growth,
while the oophoric
character as distinctly
appears in the fact
that these fronds are
practically stalked
prothalli bearing the
sexual apparatus
proper to them.”

Fic. 8.—Abnormal flower of Cattleya labiata. After a: Phe exact ne

a colored drawing by O. Ags. acter of a homoeotic
metamor phosis is often
determined by the nature of the nearest normal organs.

In many cases proximity appears to be a predominant factor.
The following instance conveys more definitely the meaning of this
general statement. A plant of Cattleya labiata, formerly grown in
the greenhouse annexed to this laboratory, produced. flowers uni-
formly exhibiting petaloid homoeosis of the calyx (fig. 8). The
three sepals—which in normal flowers are rather narrowly lanceolate
—were broadened, and their Margins were crisped in imitation of the

32 Journ. Linn. Soc. 322536. 18

: 96. The quotation above is from DRUERY’S
report in Mr. Lowe’s communication,

Pee eee

1909] LEAVITT—HOMOEOSIS IN PLANTS 45

ovate crispate petals. ‘The two lateral sepals, however, differed from
the dorsal in that the inner half of each became colored like the
labellum. The latter organ forms in this species, by the inflection of
its margins, a loose tube, embracing the column, open distally and
expanding to form the landing-stage for the pollinating insect. The
floor of the tube has a broad yellow stripe, widening toward the
mouth, and at the extremity replaced by a large purple or crimson-
magenta patch. These colors are bounded by the pale magenta
body-color of the flower. In the abnormal specimens the halves of
the lateral sepals next the labellum reproduced the color-pattern of
half the labellum. The halves away from the labellum were like
the rest of the flower in hue. This peculiar distribution of the colors
becomes interesting when an examination of the base of the perianth
shows that in Catéleya labiata the foot of each lateral sepal is close to
the foot of the labellum and overlaps it by just half—the half corre-
sponding to the homoeotized half-limb in the abnormal flowers;
while the other perianth segments stand somewhat apart from the
lip. In these particolored sepals the discrimination between homoe-
otic and non-homoeotic areas seems to be directly related with con-
tiguity to the “source”’ of the derived features.

PENz1G%3 has dwelt upon this phase of the phenomenon which we
sa now calling homoeosis, in his general considerations of the sig-
nificance of monstrosities and in his arguments for the theory of
DELPINO regarding the nature of the fertile scale in Abietineae. He
was impressed, as everyone must be, with the agreement between
the facts of this category and the formative-stuff theory of SACHS.

10. A compound member may be changed in such a way that some
or all of the parts exhibit the plan originally characteristic of the whole
member (entropic homoeosis).

The general plan of organization is transferred from the member
. a whole to the constituent parts. Compound.members may thus,
. entropism of form, suddenly become decompound. This form
Pees. has already been illustrated in the description of the
ie s ern above. Further examples in leaves of ferns and flower-
a P ants may be noticed here, since this variation has an interesting

ation to the question of the evolutionary origin of compounding;

*° Pflanzen-Ter. atologie, 1. c. (and elsewhere).

46 BOTANICAL GAZETTE [JANUARY

and besides is interesting from the purely morphogenetic point of
view. The same species of variation is also to be observed occa-
sionally in inflorescences.

A glance into two or three private and as many institutional
herbaria makes it evident that entropic homoeosis is prevalent in
ferns. I have found it in the ten species following, repeatedly in
several of them: Aspidium Thelypteris, A. spinulosum, Polystichum
acrostichoides, P. angulare, P. Braunii, P. munitum, Asplenium the-
lypteroides, Dicksonia punctilobula, Polypodium vulgare, Osmunda
cinnamomea,. :

Among these species I have found the phenomenon most fre-
quently in Polystichum (Aspidium) acrostichoides, in which species

its operation takes some
interesting turns. The
fronds of P. acrosti-
choides are somewhat
dimorphic, not only in
the sense of being

“A OH ae Y soriferous and non-

Se Oa aa aa soriferous, but also in
se € Y {/i/\}.| the fertile and sterile

Ex fronds having slightly
eae I different general out-
f Fic. 9.—Abnormal Pinna of Poly- lines. The sterile fronds

re stichum. acrostichoides. have their apical pinnae
: non-auriculate, obtuse
and oblong or elliptical to obovate. In the fertile fronds,
the pinnae, again, are dimorphic, the apical ones being
shorter, narrower, and abundantly soriferous. Homoeotized pinnae
which I have found on various specimens represent the entire range
of this dimorphism. A specimen in the herbarium of A. A. EATON;
ex-herb. E. J. WaNsLow, collected in Lowman, Chemung Co., N. Y:
has the two basal pinnae transformed into miniature fronds. Theif
length and breadth are increased, and the segmentation is quite
perfect. One of them very nearly represents a fertile frond, its tet-
minal portion being constricted and soriferous, and many of the
pinnules being auriculate or toothed (fig.9). Some of the pinnules,

saat

1909] LEAVITT—HOMOEOSIS IN PLANTS 47

however, are of the type found toward the extremity of the infertile
frond, as above described. The proximal fourth of this pinna is
occupied by pinnules much shorter than the rest, so that the total
width of the pinna here is about normal. This region is the part of
the pinna corresponding to the stipe of the frond, and would be bare
of pinnules were the frond-plan fully realized; as a matter of fact
two form-forces are
expressed, the pin-
nar, which gives a
WIRNINAK blade of proper
; width on either side
Fic. 10.—Normal pinna of Polystichum Braunii. of the mid-vein, and
the frondescent,
which divides the blade. In the opposite basal pinna of this specimen,
segmentation is complete, but the homoeosis refers to the terminal
pattern of the sterile frond.
A specimen of this species in the Gray Herbarium marked “ Herb.
A. Gray. Near Philadelphia, Bourcurn,” exhibits an interesting
Interplay of form-factors, wherein that which at first
Sight appears to be extreme irregularity gradually es
Tesolves itself into definite adherence to |
Pre-existing types, Frond-plan in the
pinnae is expressed by , A
Segmentation and by § we,
the character of . oe _
the segments, by ft PS ;
their increased => A
length, and by Re 3
Fic. 11.—Homoeotic pinna of Pol-ysti-
chum Braunii (specimen in herb. GRAY).

increased width
of the distal two-
ot eg moe Of the lower pinnae by the absence of pinnules
third or . Bie 0 the pinnar rhachis, throughout the basal one-
Petiolate oS shbn of its length—in an “effort” to realize the
ate those of “ee of a frond. The segments, more or less complete,
and the frond 9 as al region of the infertile frond of the species,
infertile + “OFM, In so far as it is realized in the pinnae, is of the
Ype. In these pinnae, the old fashion (pinnar character)

48 BOTANICAL GAZETTE [JANUARY

is able to express itself simultaneously in the morphogenesis, by
more or less successfully reducing the segmentation, by maintaining
normal width in the basal region of the pinna, and by accentuating
the basal segment on the upper side of the pinna, in correspond-
ence with the large tooth or auricle which stands there in normal
pinnae. The homoeotic character of the variation is unmistakable,
though partial.

In a specimen from the Torrey herbarium, at the New York
Botanical Garden, the frondescent pinnae of the lower section of the
aberrant frond
imitate an occa-
sional trick of the
species, by which
sorif may

Fic. 12.—Normal pinnule of Osmunda cinnamomea. run down the

margin of the sterile
region, on the tips of the pinnae. In the Torrey specimen, the lower
frondlets (pinnae) are soriferous and constricted in their apical
region, and the soriferousness runs down their margins on the tips of
the pinnules.

Dicksonia punctilobula in the herbarium of Mr. F. G. FLoyp
shows entropic homoeosis of an interesting type, inasmuch as the

4 : j

7\ 5 ( eee

Le Q

ae ONY a ET x. a ee en
——_

g — a, ; Q

Fic. 13.—Homoeotic pinna in Osmunda cinnamomea (in herb. FLOYD).

primary segments become frondescent in imitation, not of the adult
but of the infantile, or nepionic, leaves.

Asplenium thelypteroides collected by DAVENPORT and FLoyD at
Coon Hollow Brook, Milton, Mass. (hb. Froyp), has the second and
sixth pinnae frondescent (others in a less degree), the modification
proceeding upon the basis of the lobing already normally present in

‘

1909] LEAVITT—HOMOEOSIS IN PLANTS 49

the pinnae, and resulting in the formation of segments of a second-
ary rank imitating the normal primary segments; and further, in a
change in spacing of the divisions, which brings the modified pinnae
into conformity with the main frond-plan as regards the distance of
the segments. Increased spacing, correlated with perfected divisions,
in very obvious homoeosis, is also seen in Osmunda cinnamomea
(fig. 13) collected by F. G. FLoyp at Mt. Desert, Me., where, accord-
ing to the collector, the form is evidently hereditary, numerous modi-
fied plants covering a considerable area.34

Coming to flowering plants, the principle is seen in curious varia-
tions of the leaves of an Aster which Mr. C. Stuart GAGER has
kindly shown me. They were briefly noticed by him in Torreya for
January, 1908. Basal laciniae have become increasingly distinct,
some of them even petiolate (petiolulate); and when so, the new, or
secondary, blades imitate the main blade, even down to a peculiarity
of asymmetrical development.

I regard a leaf condition found by me in Gleditsia triacanthos
as homoeotic. At any rate the variation noted is very abrupt. The
character leaf of the species is simply paripinnate, the leaflets short-
stalked, the blade slightly unequal at base, elliptical, obscurely ser-
ulate or crenate, apiculate. In the leaves of vigorous running shoots
(not spur shoots) I find some of the leaflets replaced by segments
mutating the normal leaf in all the features specified above; the
segments are, in effect, character leaves of a second order. The
modified, or compound, leaflets, which are nearly twice as long as
the unmodified ones, occur irregularly among the others (fig. 14).
The condition here, essentially the same as that described for the
Pierson fern, has been noticed also by GaceER (I. c.), one of his
specimens—as also one of mine—carrying the homoeosis to the
are Nia a few pinnules becoming perfectly compounded (as
e = sees fern). There is no evidence, so far as I know, that
Wis i, condition looks back to an older, normal state of division.
bins. ae Is that segments of the leaf are transformed into the

something now existing—the normal, or character, leaf of

cat Britain, etc., Nature printed. London. 1857; and in Moore, Nature
London. 1859 (?).

NO, Bot. G
1909

ardea

50 BOTANICAL GAZETTE

[JANUARY

the species. The likeness is perfect. Whether the variation here is
admitted to be at the present time in each instance a fresh display

lJ
Uy

w), NW

—_

of homoeosis, or is sus-
pected of being rever-
sionary, the abruptness
of the change and the
manner of its occurrence
seem to me plain indi-
cations of the origin of
the decompounding. It
seems to me far more
likely that this is en-
tropic homoeosis, past —
or present, than that
the complex form of
the compound segments
was worked into perfect
likeness to the original
blade slowly by natural
selection.

BONNET recorded a
variation of the jasmine
(Jasminum officinale)

WK CE
ff Ning; VERS
GY We
KOT

ve | way. Fig. 15 shows the
normal and _ variant
forms side by side, as
| delineated by BONNET
(Oeuvres 2: 363. pl. 24)

Inflorescences of
certain marked types
tropic h ne sometimes undergo en-

P omoeosis; for example, umbels and pseudumbels. Of the
chica T have seen two cases, the first in Aralia nudicaulis (specimen
in hb. N.Y. Bot. Garden, coll. by Austin C, APGAR, near Lambertville,
N. J., 1887). This species normally bears simple umbels at the ends

Fic. 14.—Leaf of Gleditsia triacanthos with seven
homoeotic pinnae,

1909] LEAVITT—HOMOEOSIS IN PLANTS 51

of the three main branches of the scape. In Apcar’s specimen some
of the florets in each umbel are replaced by long-pedicelled umbellules.
That is, the form resident in the flower cluster as a whole has instantly
passed into several of the parts (florets). A similar variation I have
seen in Hydrocotyle umbellata (specimen in hb. N. Y. Bot. Garden,
Jamaica, Ferry, no. 6168):
and in Daucus Carota it was
noted by CRAMER,35 radial
florets being replaced by
umbels.

Two false umbels of
Pelargonium in my posses-
sion are abnormal in a
similar way. In each case
one of the florets, eccentri-
cally situated, has been Fic. 15.—Normal leaf of jasmine (left) and
transformed homoeotically, nea Se
with the result that it rises
aS a pseudumbel of a secondary order from amid the flowers of
the first. The subordinate flower cluster is like the chief cluster as
Tegards centrifugal development, involucre, etc., though the flowers
are fewer (8 instead of 18) and shorter-pedicelled. The secondary
Peduncle is articulated to the expanded summit of the primary, as
are the pedicels of the flowers, and apparently would have been cast
off, in event of failure to fertilize, by an absciss-layer, as with the
flowers among which it stands. :

Homoeosis and reversion

Thave said that homoeosis is often confused with reversion; indeed
3 oe the confusion may be said to have been habitual.
Teversio oes having applied to homoeotic phenomena the word
little ni bi ats peeent sense, some contemporary authors have with
Which schon retained the antiquated language in expressing ideas

are altorether modern,36

°S CRAMER, C

63. Zurich, 1864. : Bildungsabweichungen bei einigen wichtigeren Pflanzenfam., etc.

36] :
reversion = Odora 7:18, 19, 1905, I have discussed briefly the origin and use of the
"Idea as applied to plants,

52 BOTANICAL GAZETTE [JANUARY

In order that the real relations of the forms concerned shall be
more generally recognized, one needs but to direct attention to them;
argument is scarcely needed. For example, the appearance of two
supernumerary spurs in Platanthera viridis is palpably anything but
atavistic. No one at all conversant with Orchidaceae will for a
moment imagine that there ever was an ancestral race of three-
spurred Habenarias to which the curious Dorset plant harks back.
In the above-mentioned gentian with the fringed carpel, the relation
of things in evolutionary time is perfectly clear. While the carpel
is an ancient, the fringe is a very recent, structure; the former was
not derived from a petal, and the fringe has never until now been a
carpellary appendage. When the carpel puts on the fringe, therefore,
it adopts a character not to be found in its own phylogenetic line.
Simply the fringe is borrowed; there is no reversion. The non-
atavistic nature of the stigmatic papillae on bracts and foliage of
Crocus sativus is equally patent. The assumption of foliage charac-
ters by carpels of Drosera is no more reversionary. The carpel of
Drosera, it is safe to say, had its origin in common with that of other
angiosperms, and runs back through a series of forms, none of which
is a foliage leaf, to the megasporophyll of the earliest angiospermous
seed-plants. Likewise the foliage leaf of Drosera is a derived struc-
ture, with characters probably more recent than the family Drose-
raceae even. Its peculiarities of contour and its appendages arose
long subsequently to the establishment of the angiospermous, and even
the droseraceous, carpel. To reach a point whence these two lines
of derivation diverged, i. e., a point where the reproductive and the
vegetative organs were one and the same member of the plant body;
we must probably go back to the spore-bearing foliage leaf of the
fernlike ancestry, far antedating the first flowering plants. The
common original of the Drosera carpel and the Drosera foliage leaf
was probably a kind of fern frond. The aberrant carpels in question
bear no resemblance to fern fronds. They do not reproduce a form
from which they are descended. They have simply taken to them
selves properties of coordinate derivative members, the foliage leave _
This is so obvious that the statement would seem superflous were it
not still the custom of inconsiderate writers to speak of such meta
morphoses as reversions in the phylogenetic sense. The transforma

1909] LEAVITT—HOMOEOSIS IN PLANTS 53

tion is not retrograde in any except an imaginary sense; since the
normal unmodified carpel is to be regarded as already “retrograde,” or
conservative, as compared with the tentaculiferous leaf; and the ovule
as vastly more ancient than the tentacle into which it suffers change.
~ In peloria the morphogenetic relations are not so instantly obvious.
However, the phenomenon is usually, if not always, better viewed as
a homoeotic, than as an atavistic, occurrence. In notices of peloric
monstrosities one often meets with the statement that these forma-
tions have a historical value. Thus, with respect to a peloric
Laelio-Cattleya, MasTers3” suggests that here is a reversion to the
earlier and simpler conformation from which the peculiar orchid
structure, as we know it, has evolved. While this is true abstractly,
in a merely descriptive sense—since actinomorphy doubtless preceded
#ygomorphy in the monocotyledonous phylum—yet it is probably
untrue if we are to take it in any real phylogenetic sense, with the
understanding that actinomorphy has remained latent as a heredi-
tary character through the enormously long period of the evolution
of this family from an actinomorphic condition. Regular peloria is
to be considered in conjunction with the opposite change, which
frequently occurs in orchids. The antithesis of the two pelorias
makes it evident that we have here something besides atavism; since
if either form is atavistic the other cannot be. This outcome,
coupled with the fact that we have a less objectionable construction,
discredits the entire idea of reversion in peloric orchids. The argu-
ment extends to other families.
- et further expatiation, it will be evident—if the standpoint
a lg paper is correct—that the word reversion is a much-
~ ‘tm. True reversions, except those which occur periodi-
cepa eae are, I Suspect, rather rare. Atavism is never
Sie wie: hae in teratological cases, but it is to be admitted
es a5 we by aid of independent proof. It should be
often to be sy, . at antecedents of monstrous forms are much more
dies &ht in contemporary normal parts than in ancestral con-

The place of these facts in botanical theory

. to which I am referring from a special point of view
In botanical theory in at least three different relations:
hron, . 31235, 239. 1902.

The fact
have Values
37 Gard. C

54 BOTANICAL GAZETTE [January

frst, a study of the modes or phases of homoeosis helps us to estimate
at its true worth teratological evidence applied to the solution of
phylogenetic problems; secondly, the facts seem to throw light on
the method of evolution of some normal structures; thirdly and chiefly,
the facts of homoeosis constitute, as has been already pointed out in
the introductory passage, an important section of the data of morpho-
genetics. Let us examine these relations of the subject a little more
in detail.

1. The study of homoeosis must somewhat increase the caution
with which we use deviations from the normal as aids to morphological
interpretation.

In the past the commonest use of abnormalities has been to make
them the ground of phylogenetic inductions. The stereotyped
remark of writers describing monstrous specimens has been that such
aberrations “are very instructive” —an expression of faith either in
the phylogenetically reversionary nature of abnormalities, or in
the eternal inviolability of homology in morphogenetic sequences.
Surmises from monstrosity alone were naturally more common if
the days when the evolutionary story of plants was less complete
than it is now, and when morphologists were driven to indirect and
speculative methods. The history of this subject is voluminous;

truth; and it will not be entered into here. Some of the contentions
for which teratological formations were used have been abandoned,
and others have been settled by the discovery of direct evidence from
comparative morphological studies of living and extinct plants. AS
we are enabled by extension of knowledge and maturing of opinion
to understand better the relations of both normal and abnormal forms,

botanists in the absolute integrity of morphological categories; am
if the observations assembled in the present essay are accepted 1

1909] LEAVITT—HOMOEOSIS IN PLANTS 55

the sense in which they are presented, it is indisputable that stages
of development supposed to be concatennated in a fixed order are
subject to the most violent dislocation—that a stage belonging to one
morphological category may pass into another of a different category.

Abnormalities which are traceable to very ordinary features of
contemporaneous organization and may be brought under the rule
of homoeosis, will for the moment lose all historical force which they
may ever have been thought to possess. Historical significance can
be restored to them only by application of the laws (if we may so
speak of the operations) of homoeosis. If homoeosis much more
easily follows lines of homology, and only rarely transgresses them
under very special conditions, then frequent homoeotic metamor-
Phosis of a particular organ in a particular direction may be thought
to be indicative of the derivation of the organ. But I think that the
admission of homoeosis in any case is practically fatal. The method
of inference then becomes too roundabout to be acceptable to modern
taste. If in any case homoeotic formations agree with the results
of comparative morphology, of anatomy, and of organogeny, still
they add only a reflected light to the general illumination.

In writing these words I have in mind particularly the long debate
— the morphological nature, or origin, of the ovuliferous scale in
Conifers °r so much of it as relates to malformations. Perhaps no
Single organ of vascular plants has afforded morphologists more mat-
ter for disputation than this scale, which has a vast literature of its
own by the most eminent authorities—Linnarus, A. BRAUN, SCHLEI-
PEN, VON Mout, EICHLER, Sacus, BAILLON, STRASBURGER, VAN
Tiecuem, DELPINo, PENzIG, Masters, Nott, CELAKOVSKY, and
neg The theories of these authors have been various, and based
yeaa oo consideration, not least upon abnormalities.
ie ope as appealed very differently to different students—to

a cael: One doctrine, to others as favoring a different
having - aan as of great or even decisive weight, to others as
Re titan at all. CELAKOVSRY, who has studied the
cially eg conan has relied upon abnormalities, and espe-
betwee aR ss en series of intermediate formations, such as those
ance of a ch a € scale as it separates into parts (at the appear-

) and the leaves (or bud scales) of the shoot. - This

56 BOTANICAL GAZETTE [JANUARY

author assumes that such gradations can exist only between homolo-
gous organs. He regards the entire composite structures found in
the monstrous cones, made up of the fertile scales and the shoots —
which grow through them, as such intermediate formations (ana
morphoses), and looks upon them as sufficiently proving the shoot-
nature of the fertile scale.38

From the numerous clear and detailed drawings of the abnormal
cones which have been published,39 certain truths are apparent
enough. First, no new organ or form of any organ, not proper to
the species of today, except intermediate formations between present-
day vegetative and reproductive parts, is to be found in the cones.
No ancient structure, nor anything suggesting ancestral structures,
nor any organ of paleobotanical aspect, makes its reappearance out
of the past; there is no reversion. If the shoot in the axil of the
bract, replacing the scale, were truly atavistic, we should expect
that lost characters would appear in the axis and its appendages.
But the shoot turns out always to be an ordinary leafy branch with
the reduced foliage representing the xerophytic adaptation of the
group. This shoot is not reversionary in even a barren formal and
descriptive sense; for if the fertile scale represents a shoot, now
reduced to two ovules and their expanded integuments, the relics of
two sporophylls, and if the original development of axis and spore
phylls is to be restored, we ought to have in the restoration an axis
terminated by sporophylls or by ovules as representing them, since
megasporophylls when reduced to a pair are terminal appendages;
but this development is not realized in the abnormal cones, in which
we find that the shoot arises sometimes above or below the scale, and
when in the midst of it, then in the form, not of a stalk, but of @
proliferation, upon which the parts of the fertile scale become basal
appendages—if they are to be taken as appendages of this shoot at all.

The transformations of the cone are homoeotic. And therefore

3° CELAKOvSKY’s writings summarize the whole controversy (Abh. Kgl. Bohm.
Gesell. Wiss. VI. 11:1882; VII. 4:1892; ENGLER’s Bot. Jahrb. 24:202. t

= E. g., CELAKovsky, Abh. Kgl. Bohm. Gesell. VI. 11:1882; PARLATORE, =
Sci. Bot. IV. 16:1. 13. 1862; STENZEL, Nov. Act. Nat. Cur. 38: pls. 12-15. 1878
VELENovsky, Flora JI: pl. Tr. 188s.

1909] LEAVITT—HOMOEOSIS IN PLANTS 57

while it is in itself not unlikely that the fertile scale represents a pair
of sporophylls belonging to an axillary shoot, and while comparative
morphology and anatomy may make this view even probable, the
monstrous cones, it seems to me, add nothing material to the satis-
faction which one may take in this solution of the conundrum.

Nor is their value improved by the anamorphosis idea. To me
at least the “transitions” from fertile scale to axillary shoot do not
appeal with the same compulsion with which they appeal to CELA-
KOVSKY and some others. I can conceive the most perfect of these
transitions to represent combinations of historically unrelated form-
factors, reproductive on the one hand and vegetative on the other;
and in the conception I find nothing incongruous with other facts of
homoeosis. If there were originally no shoot in the axil of the fertile
leaf (the bract) but only an ovuliferous segment (SACHS-EICHLER) or
pair of lobes (DELPINO) —supposing for argument that these notions
were admissible on any but teratological evidence—and if by influx of
Vegetative forces the cone were converted gradatim into an ordinary
branch, with buds in the axils of the leaves; then it seems to me that
we might expect to discover morphogenetic impulses toward the
formation of ovuliferous segments and impulses toward bud-formation
coexistent and cooperative in the same body of axillary tissue, with
such a result as the monstrous cones exhibit. After reviewing the
combinations of diverse formative impulses in teratological occur-
tenes generally, I do not feel that CELAKOVvsKY’s case for the pre-
eminent value of anamorphoses is established. In comparison with
other Sources of suggestion the monstrous cones seem to have a
minimal value. The interest and worth of CELAKOvskKy’s thorough-

§0lng study of the whole subject does not lie in his treatment of
nionstrosities,

Yet a small de

gree of approval for the theory which makes the
Scale Tepresent

EO ag a shoot bearing sporophylls might possibly be derived

and a ey with which a shoot arises in the site of the scale;

a . M spite of the facts that there are irregularities and that

% an one plausible reason might be given for the frequent

Ppearance of the shoot. The latter has been found in the genera

ees Pinus, Tsuga, Cryptomeria, Cunninghamia, Glypto-
» Sequoia, Taxodium, and some other genera.

58 BOTANICAL GAZETTE _ [aNvary

I hope that it may be clear that the shoot-and-sporophyll theory
is not here called in question, but only the use of teratological forma-
tions as competent evidence. The ground is taken that these
formations do not remove all doubt as to the origin of the fertile
scale, but on the contrary only after all doubt has been removed as
to the nature of the scale, by legitimate argument from comparison
of normal structures, do the monstrous formations begin to have any
considerable historical significance.

2. Homoeosis has played a part—necessarily from its nature,

which is essentially anarchical, a small part—in the evolution of

plants. We can trace to a

homoeotic origin certain

established sequences in de-

velopment, of which specific
examples may be adduced.

(a) Habenaria quinqueseta
(or Michauxii) of the southern
states carries on vegetative
reproduction by certain of its
roots. At the apex of these —
roots, close to the punctwm —
vegetationis, pointing back-
ward in the embryonic tissues,
a stem apex is organiz
Fig. 16 shows its relations t0
the apical regions of the root. —
Two leaves and a bud in the

Fic. 16.—Median longitudinal section of axil of one of. them -_
root of Habenaria quinqueseta. already been differentiated.

Subsequently to the stage here
represented, the root-apex enlarges and forms at the base of the
new shoot a spherical tuberoid growth evidently with storage
functions.

As to the evolution of a shoot fundament in this curious position,
no one, I suppose, will imagine that the entire evolutionary history
of the stem, leaf, and bud in cormophytes has been repeated in the

1909] LEAVITT—HOMOEOSIS IN PLANTS 59

interior of a Habenaria root. It is perfectly clear that the form-
giving agencies which shape the leafy shoot in this species have been
set to work upon tissues which up to a certain point have been domi-
nated by other forces. There is nothing new except the extraordinary
morphogenetic sequence. In this instance, morphic translocation
has become habitual, or normal.

(b) In Phyllonoma ruscifolia of Mexico the flowers are regularly
produced from the upper surface of leaves, near the apex (fig. 17).
There is no adnation in this case, for the anatomy
of the blade below the inflorescence and of the
petiole shows no addition to the normal vascular
structure, indicating any concrescence. Here
again an abrogation of ordinary morphogenetic
Sequence has become fixed in the development.

(c) The specific form of the vegetatively
derived embryo in some plants must be con-
sidered homoeotic; as for example in Opuntia
vulgaris, described and figured by GANONG.‘
At the time when fertilization should take place,
the egg cell, according to GANONG’s observation,
has become disorganized. The place of the
normal embryo is taken by several embryos Fic. 17.—Leaf of
budding in from the nucellus. The noteworthy 7” spac can ie
feature of these apogamous individuals is their pe
adoption of the form of the abdicating embryo proper. Here is a
homoeosis which has become established and provides a regular
Means of propagation for the species. A considerable number of
such cases is known,

lt 18 natural to imagine that conditions in the embryo sac deter-
es the form of the buds and cause the homoeosis; yet it is not at
certain that such is the explanation of the assumption of form.
Sa _ aps by myself the embryos organize outside of the
fo - Teler to s piranthes cernua of the meadows. The upland
sis? of the Species has a normal development. In the variety
raise oy Where in rich meadows I have seen only polyembryonic

- Having followed the development with care, I find that the

* Bor, Gazerre 252221. 1808, oe :

60 BOTANICAL GAZETTE [JANUARY

embryo sac ceases to develop after a few nuclei have been formed in it,
and is pushed aside by the hypertrophic inner integument. This
envelope, normally composed of a few flattened cells overlying the
sac, in this variety of the species early takes on a very active growth,
forming a mass of cells which, as it comes to maturity, splits up into
rounded bodies simulating the embryo of the genus.4* The forma-
tion of embryos is often incomplete and growth then results in amor-
phous masses; but on the other hand, it is often successful, and
gives such seed-contents, consisting of several
well-made embryos, as that figured herewith
(fig. 18). It is true that the form is very simple,
for the normal embryo has neither suspensor nor
cotyledons. But that cells of the inner integu-
ment, ordinarily forming a saccate one-celled
layer, with not the remotest resemblance to an
embryo, should so much change their habits, and
in the transformation should select the fashion,
though simple, of embryos, is sufficiently singular.
IT look upon the process as homoeotic in its

* Wenn
x
ca

aS
ep,
ary,

F nature. as
IG. 18.—Seed of : ee
Spiranthes cernua The formation of embryos here is quite inde-
(lowland form). pendent of pollination, as I have proved by

carefully castrated and guarded flowers—an
ecologically important item for a species blooming so late in the
season. The biological significance of the process, indeed, is—if I
may digress for a moment—that the plant by this means combines
the certainty of issue which pertains to vegetative methods of repro-
duction, with the swiftness of dispersal and range of dissemination
secured by the seed apparatus. This js doubtless the combination
of favorable circumstances which has secured the wide adoption of
apogamy in Compositae also. In Spiranthes cernua, homoeosis
supervening in the integument has doubtless contributed to the
abundance of the species in Suitable soils from Massachusetts and

4 Rhodora 23227. 1900, SS

1909] LEAVITT—HOMOEOSIS IN PLANTS 61

ROSENBERG,** of a whole series of characters from the megaspore
and embryo sac to a somatic cell in Hieracium flagellare. The
embryo sac is sometimes formed normally, but in the greatest num-
ber of cases this structure aborts. A neighboring cell, which may
be situated in the epidermis of the nucellus or in the chalazal region.
or in the integument, takes upon itself the function of a megaspore,
though without reduction of the chromosome number. This cell
enlarges to form an embryo sac. Its nucleus divides as that of
the megaspore would do; egg apparatus and antipodals are formed;
and even polar nuclei fuse to form an endosperm nucleus. The
gametophyte is thus reproduced, after the failure of the proper organ
‘o construct it, by a cell of the sporophytic generation. And this
homoeotic process is habitual, hereditary, and established in the
species.
If HEcerMarer’s observations upon Lycopodium Selago are cor-
rect,*° we may attribute the formation of the gemmae of this species
and its near allies to a homoeotic origin. The gemmae—detachable
shoots—arise in exactly the position of leaves, from which in their
earliest stages they are indistinguishable.

: The form of homoeosis most often entering into the diversifica-
hon of species of plants is apparently that which has above been
called entropic. Abscission of leaflets has already been discussed
in this connection. Another seemingly homoeotic feature of com-
Pound leaves is found in their stipels. An examination of stipels
1 a considerable number of groups—as in Xanthoxylum, Staphylea,
: Phis, Turpinia, Robinia, Bradburya, Desmodium, Galactia,
Dolicholus, Vigna, Amorpha, Sambucus—strongly suggests that the
eae have arisen as echoes of the antecedent structures, stipules,
i the evolution of the leaf suitable conditions have been
little do - hey occur in plants possessing stipules, and with
in ubt already in possession of them before leaf-compound-
Mes 7 2 they are generally useless, as far as one can see; when
eis: ave ‘a visible use it is the same as that of the stipules, in a
weakened degree; they follow punctiliously the greatly vary-

* Bot. Tidsskrift 28:

2150. 1907.
43 Bot. Zeit. 302841. 1872,

62 ' BOTANICAL GAZETTE [JANUARY

ing character of the stipules—being large, stout, and thorny; slender,
terete, rigid, and sharp-pointed; long, weak, and membranaceous;
small or evanescent or flattened and glandular-capitate, etc.—in
the different groups; they occur in situations much resembling the
situations occupied by the stipules. All of these facts suggest that
the compounding of the leaf with organization of partial leaf-stalks,
in evolution, has been the occasion for the production of stipels
homoeotically.

Moreover, we may well suspect that in many cases decompounding
of the blade has followed upon compounding from the same general
cause; the partial blades borrowing the compounding tendency from
the parent blade, the whole leaf thus becoming by a single step
decompound. The circumstance that the ground-plan of the whole
leaf is repeated in miniature in the several parts, in a vast number
of decompound leaves of both phanerogams and ferns, and the
occurrence of the corresponding type of homoeosis as individual
variation in both divisions, lend a color of probability to this con-
ception. Decompounding, on this hypothesis, would have no initial
relation to utility, and would not be a product of natural selection.

To consider this matter a little further with respect to ferns: If
the variation which we find so frequently in ferns has been the basis
of evolutionary advance in complexity of the frond, we should be
able to discover in the decompound-leaved species certain relations
which would necessarily follow from such an evolution. We shoul
certainly find the ground-plan repeated in the segments. We should
expect to find, further, that the historically earlier and simpler condi-
tions would be retained in youthful leaves; and that these youth-
ful leaves would be matched by the segments of the fully developed
fronds of the mature plant. Both these expectations are met in many
species. Ferns on every hand illustrate the first. Of the second, a
single example will be enough to direct attention to the facts which are
easily observable. In fig. 19 are reproduced a youthful frond of
Polystichum angulare proliferum, and a pinna from an adult leaf.
In Asplenium Filix-foemina we have a depauperate variety, Val
exile D. C. Eaton, the mature frond of which is very precisely
matched by the pinna of the frond of the typical form.44

44 See Eaton’s Ferns of North America pl. 76.

1909] LEAVITT—HOMOEOSIS IN PLANTS 63

Further, if entropic homoeosis has given us decompounding widely
in ferns, we shall expect to find species so related to each other that
some represent the original types and others the homoeotic deriva-
tives. We shall look for, first, the simply pinnate types; secondly,
types in which the pinnae match the fronds of type z; and perhaps
thirdly, a type in which the fundamental plan is worked out in the

pinnules. If the simple type has two
modes, as above described for Polystichum

bo acrostichoides, we may expect to find
either or both represented in the more
complex patterns.

Anyone having access to a collection
of Polystichum may see this projected
scheme filled out by existent species.
First we have the simply pinnate forms,
such as P. acrostichoides, P. munitum,
P. lonchitis; in which we have two frond
patterns, za being the fashion with more
or less oblong, non-auriculate pinnae,
probably historically earlier, seen in a

iu pical regions of sterile fronds; and 2)

Fic. 19.—Youthful frond being the characteristic plan, the pinnae
Pin eg adult frond of falcate-auriculate. Then we find many
erum. cy Sa anc species with pinnae frondescent in fac-

; simile of pattern za or 1b; as P. cali-
uf ormicum (1a) and P. Lemmonii (1a), P. aculeatum (1b), and P.
Braunii (jig. 10, 16, sometimes za); and others, a considerable
weal of species. Finally, frondescence of the second degree

eu Tepresented in P, angulare tripinnatum (ta, in the ultimate

aA in P. scopulinum the (supposititious) homoeosis is

when as e, the Proximal region of the pinnae alone being seg-
hted (za), the distal merely lobed.

Caice find species so much alike (except that one of two is simply
. Ae decompound) that one appears to have been
Indian Saga from the other by entropic homoeosis ; as the West
pound) ae bala (decompound ta) from P. triangulum (com-

: urope P. aculeatum and P. lonchitis stand in a similar

64 BOTANICAL GAZETTE [JANUARY

relation to each other; the slight differences—apart from homoeosis—
are such as might be expected to arise in specific isolation.

In homoeosis a character or a system of organization which has
been evolved in one part of the body is transferred, ready made, to
another part. The great mass of instances are of the class called
teratological. By this designation we mean, substantially, that they
are suddenly appearing deviations from the customary structure,
“Monstrosities”’ in general have the special value that their chronol-
ogy is oftentimes ascertainable; we know that such and such identical
plants have arisen in the midst of normal relatives. They exhibit,
as has been said, discontinuous or saltatory variation. That homoe-

otic monstrosities typify homoeosis in general, as a saltation phe-

nomenon, may be inferred from the very nature of the process. When,
therefore, we discover homoeosis at work in normal evolution, diver-
sifying lines of descent, we are able to augment the steadily increasing
collection of evidences for discontinuity in the origin of specific
differences.

- Homoeotic changes may thus be classed with mutational phe-
nomena. But grouping them with the mutations exhibited by the
Oenotheras rests only upon some such negative property as that
homoeotic transformations fall outside of any law of the evolution of
characters by natural selection. It seems unlikely that the two classes
of alterations have anything further in common. Mutation proper

catory scheme of variations under mutation seems to rest on purely
formal and abstract descriptive resemblances.

3- The idea of homoeosis unites for descriptive purposes a great
number of facts of ontogenesis which, even though they may not

i OO _ _
Ste aR Skee ee Pa nn Re ae ee ERS Foe rer
Fe eg ee eae ere

1909] LEAVITT—HOMOEOSIS IN PLANTS 65

at the present juncture point a way to the correct mechanical ex-
planation of development, possess in this connection a considerable
prospective value. An adequate theory of ontogenesis must take
these facts into account.

Their value lies in their exceptionality. Homoeotic occurrences
are fundamentally antithetic to our usual conception of the method
of development. The most general and fundamental of our notions
of a mechanically autonomic evolution from egg to adult, is that of
a series of states (A, B,C, D ..... X, Y, Z), so related that each
one necessarily serves as the stage upon which the next state arises.
Every one of the several states seems to be the needful condition for
the appearance of the next following. This is the primary concep-
tion of ontogenesis derived from ordinary experience. It is contra-
dicted by homoeotic formations. We see the usual sequence violated
at some point, and a state (as X) arising upon a state (as D) from
which it does not normally arise, and from which we have supposed
that, in the very nature of the process, it cannot arise directly.

In homoeosis, then, we have a new ontogenetic phenomenon.
Herein lies its worth; for every datum of a new sort adds to the
Materials for a true theory of development, and increases the chances
of our finding a clew to the right construction and combination of
the materials

Although the times are doubtless not ripe for the appearance of
an adequate theory, and further attempts in this direction may be
Profitless for the present, yet almost universally there has been felt
= desirability of biomechanical hypotheses going behind the bare
i of development. We have had from Darwin, NAGELI,
sia eee Roux, Wetsmann, Driescu, and others a series
‘eica . ess elaborate attempts at an explanation. I think that

all familiar with this line of biologic endeavor will recog-
_ = — in substantial knowledge effected under the influ-
ch speculations—even though they deal with gemmules,
“Sos iad biophores, and less concrete “organizations,” far
some of th. Tange of the microscope. And it is entirely possible that
which _.. already offered may be germinal points from
ea able system will develop.
Persistent attempt at a solution has been made by WEIS-

66 BOTANICAL GAZETTE [JANUARY

MANN, who has erected a theory of the nucleocentric type, wherein
the control of development is made to reside within the several cells,
and specifically within the chromatic matters of the nucleus. Nuclear
divisions are conceived as being of two types; namely, (1) equating,
in which the germ-plasm is conserved in its integrity, all characters
of the mother nucleus being shared equally by the daughter nuclei;
and (2) differentiating division, in which the daughter nuclei receive
from the parent nucleus unlike assortments of character-giving bodies.
All the determinants, capable of giving form to the whole organism,
being present in the nucleus of the egg, differentiation is concomitant
with an orderly distribution of the governing bodies, effected in the
successive nuclear divisions, each different part of the body ultimately
receiving its proper kind of determinant.

In view of present limitations of our knowledge, the theory of
WEISMANN is a highly speculative system, and as such has been

the proposal of this hypothesis has prompted investigations leading
to interesting discoveries. And even as a means of throwing obsel-

vations into some sort of order, any such theoretic scheme has its —

use. It seems to me, therefore, well worth while to consider all
kinds of facts of ontogenesis in the light of this and other generaliza-
tions in morphogenetics. The facts of regeneration have thus been
arrayed with relation to the Roux-Weismann hypothesis. Regenera-

whole subject gains in spirit. A tentative relation is established
between regeneration and morphogenetical theory in general. .

The effect upon the Roux-Weismann theory has been the intro-
duction of a fundamental modification. For the power of reproducing
lost parts, so Widely possessed by animals, and in some degree by
plants, is not accounted for by the Roux-Weismann idea in its most

: ants sufficient for purposes of regeneration
This auxiliary equipment is, however, tightly stowed. Under ordi-

ck, eR aaa ra hs

Vee ON ey TEEN Ae ee ee Se ee Sen ee eee

1909] LEAVITT—HOMOEOSIS IN PLAN1'S ° 67

nary conditions only the differential determinants are free to influence
the development of the cell and its activities, the regenerative or
reserve determinants being held strictly under restraint until they are
called for, to reconstruct lost members.

Homoeosis has a similar influence with regeneration upon the
Roux-Weismann idea; and perhaps a more destructive influence.
For while the facts are quite as striking, they are very much more
varied and abundant. And if the vast array of data, the existence of
which has been suggested in the present article, is added to the evidence
from regeneration, the necessity for providing full reserve-funds of
determinants in differentiated parts is much increased. Many parts
of the body—this statement is made especially with reference to
plants—are able to produce almost any other part. We are prob-
ably warranted in postulating a pangenerative capacity for every
Vital member. We know that a single epidermal cell of the leaf in
Begonia may originate a complete plant. Indications are not wanting
that single living plant cells in general intrinsically possess the same
Power, ordinarily latent, its exercise inhibited by circumstances.
If each cell possesses a complete character-fund, the characters
Capable of being severally activated upon motion of factors external to
the cell, does not the necessity for imaging special individualized
determinants disappear? Can we not as well think that differences in
cells and forms of organs spring from the nature of the molecule of the
form-giving substance (probably chromatin), this substance being the
Seat of morphogenetic powers ready to be evoked and responding
variously to the diverse conditions in which, in the course of develop-
ment, the substance finds itself placed? Obviously, in proportion as
" it of an organism are made to appear equipotential in a morpho-
eenetic sense, does the need of assuming the existence of different

me of form-giving substance diminish.

a the basis assumed for an explanation of the micto-
be. ich we call ontogenesis—whether the existence of special
. ng bodies, or the general properties of the organic
wie ea fo ming stuffs capable of diffusion, or some
ormation of € abrupt diversion of formative currents and trans-
tea « members into others of usually dissimilar origin, the

Ppearance of forms in locations not expected in the ordi-

68 BOTANICAL GAZETTE [JANUARY
nary sequence of development, and the potentiality of all parts in
each part, indicated by the general phenomenon which we have been
calling homoeosis, will need to be provided for in our ultimate theory

of development.

STATE NORMAL SCHOOL
TRENTON, N., J.

SonipreR ARTIGEERS

LONGEVITY OF SEEDS 8

In a recent article on longevity of seeds, Ewart makes a number of
statements which merit comment. He assumes as correct the claim of
BERGTHEIL and Day? (working on Indigofera arrecta) that théy have
priority in discovering that the water-resisting power of the seeds of legumes
is due to the character of an outer layer of the coat. NosBeE,3 however,
pointed out the fact for Trifolium pratense 31 years before the publication
by Bercruer and Day. Both find that stains dissolved in water penetrate
in the hard seeds only through the very thin outer layer, called cuticle by
Nope; but that they do not pass through the palisade layer. This is
shown in fig. 1 of BERGTHEIL and Day’s article. Ewart finds the resist-
ance in Adansonia digitata to be due to the impermeable nature of all layers
of the integument, and I find the same to be true of the hard seed of Axyris
amaranthoides. It is generally assumed that in the Leguminosae the imper-
meability to water is never due to oily deposits. I find that the seed of
ee Mesquite (Prosopis juliflora) is an exception. By soaking these seeds
in ether for several days and then allowing the ether to evaporate, a large
Percentage is caused to germinate when germinati litions a pplied,
while a direct supply of germinative conditions brings only 5 to 10 per cent.
The evaporation of the ether in which they have been steeped always shows
an oily residue. . Absolute alcohol is less effective in this case.

Ewart says: “The seeds of the hawthorn are supposed not to germinate
Ra after a year in the ground. CrocKER obtained no definite confirma-
kane chalga of this fact, but here also it appears to be a case of the slow
ae ona the seed coats.” This is hardly consistent with the state-
T kee 5a In my paper on the rdle of seed coats.4 In this case, however,
pa rai the significance of the coats. I find that in Cra-
thes ae embryos taken from apples just ripe and entirely freed from

S and endosperm begin growth within a few days after being
* Ewart, A. L., O

N.S n the longevity of seeds. Reprint Proc. Roy. Soc. Victoria
ce ere Pp. 210. 1908.
? BE ve,
arrecta, rm HEIL, C., and Day, D. L., On cause of ‘“‘hardness” in seeds of Indigofera

nals of Botany 21: 57-60. 1907.
3
; — F., Handbuch der Samenkunde 117. 1876.
ROCKER, Wo. . . me ‘
42:265-2091, ; 906: » Role of seed coats in delayed germination. Bot. GAZETTE

[Botanical Gazette, vol. 47

70 BOTANICAL GAZETTE [JANUARY

put into the germinator, whether in light or dark. The removal of the
inner coat and endosperm is done after sterilization, and the whole process,
including germination, is conducted under aseptic conditions. One finds
this a very tedious task and the resulting germination is of a peculiar type.
In the light the cotyledons begin to expand and turn green. A small per-
centage of the roots begin growth within a week, but in a larger percentage
the roots begin growth only after several weeks and after the cotyledons have
expanded to several times their original size. Many of the radicles do not
grow even after two months in the germinator. In the dark the growth is
similar except that the cotyledons turn yellow and the radicles are even more
tardy in their development. The coats then seem to play an important
part in the delay, but the tardiness of the radicle in its development is of
especial interest and reminds one of the behavior of the fungus-free orchid
seeds,5 or the upper seeds of the cocklebur with coats intact and in 76% of
oxygen pressure.* A full investigation of the physiology. of the germina-
tion of these seeds is now in progress by Mr. W. E. Davis and myself.
Ewart again says: “CROCKER has, however, overlooked the fact that
both the early and late seeds of Xanthium echinatum will germinate at
20 to 25°C. if the temperature is maintained for fourteen to twenty-one oF
more days instead of for eight to nine days.” I assume that Ewart meals
with coats intact, for that is the connection in which I have made my
statement. On November 5, 1908, burs of this species were collected from
the plants, the seeds removed from the burs, and the upper seeds soaked
18 hours, so as to show up any defective coats. ’ Upper seeds with perfect

coats were placed between wet filters in baths; one maintained at 24-25" ne

and one at 27-28°. On December 5 none had germinated. Of course

with coats removed these seeds germinate within three days, even at 23°.
In collections of this species from the crop of 1906 a small percentage of

upper seed with coats intact germinated at 30°. In collections of the cfop —
of 1905, on which the work for my paper was done, the minimum te™ —

perature for the germination of these upper seeds lay between 32° and 33°
even when they were kept in the germinator for a month. EWwarT says
nothing about the time of gathering, precautions against defective coats,
or the percentage germinating at 20-25°. In the absence of all these relat

data his statement can mean little. In X . canadense high temperatures ar
far less effective in overcoming seed-coat effects, and here a temperatul’ —

fluctuating between 2 5° and 41° is most effective. A temperature of 4°
° .
43° for a few hours is often more effective than a lower temperature for ®

5 BERNARD, N., On the germination of orchids, Roy. Hort. Soc. Rep. 3rd Meas

nat. Congress on Genetics 292-296. 1906

Laid

1909] BRIEFER ARTICLES oe

much longer period, a fact that led to the discovery of the temperature
effects. In the light of this fact, Ewart’s statement, “If burs are heated
at 40° C. for a day or two, 50° for a few hours during soaking, a variable
percentage of the later seeds will germinate within ten days,” is not at all
new. My data have shown that the effectiveness of high temperatures
in overcoming seed-coat effects varies greatly with different species. Further,
my data indicate that there is in this respect a slight variation in different
crops of the same species gathered from the same locality, and it appears,
if Ewart’s data have been obtained with proper attention to sources of
error, that there is a decided variation in seeds gathered from different
regions of the globe.

Ewart’s assumption that the coats in the seeds of water plants secure
this delay by excluding oxygen does not seem to be true for some of these
species. I have mentioned evidence for this in the case of the water
hyacinth. One of our students, working with the effects of oxygen on
germination, tells me that seeds of Alisma Plantago germinate rather
readily in entire absence of oxygen, provided the coats are ruptured. In
the case of the upper seeds of X. canadense she finds that about 3° of
oxygen pressure (o.2 that of the ordinary atmosphere) with two weeks’
exposure is necessary to produce germination, even when the coats are
removed. These results still need to be thoroughly tested. ‘TAKAHASHI?
has shown that seeds of rice germinate in entire absence of oxygen. It is
Probable, therefore, that the coats of the seeds of water plants secure delay in
germination mainly by limiting the water supply, as I have pointed out.®

Through a discussion of minor and less significant details, however,
we Must not lose sight of the main conclusion, which is being more firmly
¢stablished as more data are accumulated,® that delayed germination in
—. eed , though not always, related to seed-coat characters rather
sicrti as _ < dormancy of protoplasm. The coat may limit the oxygen
ply ile € cocklebur; it may exclude or merely limit the water sup-

» 4S In the seeds of legumes on the one hand and of Iris and other water
any Sa ‘a or it is possible, though not proved, that in some cases

bat . ot * chemical compounds necessary for germination.
to tell from gs omg Ewart no injustice when I say that it is impossible

: paper in how far it is a contribution and in how far a
515-38 mae Wu, Germination of the seeds of water plants. Bot. GAZETTE 44:
Tide teat me = Saag an possible in absence of air? Bull. Coll. Agr,

®§ Kiucy
28:816, 1908. G. F., Some cases of delayed germination in seeds. Science N. S.

72 BOTANICAI. GAZETTE [aNvary

compilation. In this way credit due other investigators appears to belong
to Ewart, and no one has suffered more in this respect than the writer—
Witt1am Crocker, The University of Chicago.

RESPIRATION CALORIMETER

On p. 133 of the second edition of Professor W. F. GANoNc’s admirable
Laboratory course in plant physiology, which has just appeared, and 4
copy of which has come into my hands through his courtesy, I am inter-
ested to find a Dewar flask figured as a respiration calorimeter. Before
my recent paper was published (Bot. GAzETTE 46:193-202. 1908), !
wrote to Mr. Ganonc, knowing that he was preparing a second edition of
his book, asking him to put my calorimeter into it. He wrote that his
book had already gone to press. When my paper appeared he wrote —
again, saying that he “had been using the Dewar bulbs as a respiration
calorimeter some four years past. . .-. . Of course the point about the
prior use of the bulbs is of no consequence whatever, and I mention it
now because of the coincidence in your asking me to mention their use it
my book.”

So far as priority of use is concerned, it lies obviously with Mr. GANONG.
To acknowledge this, and to record another of the curious coincidences
which after all are not altogether rare in the history of science, is the pur _
pose of this note-—GrorcE J. Perce, Leland Stanford Junior Universit).

CRATAEGUS IN COLORADO

The attention of the writer has been called by Mr. W. W. EcciesTon t0
a misstatement in an article in this journal for November, 1908. On P:
382, line 4, I should have said that the species described resembles most
C. erythropoda, of the forms known to the writer in northern Colorado.—
Francis RaMALry, Boulder, Colorado.

CURRENT LITERATURE

BOOK REVIEWS
The vegetation of Chile ;

The eighth volume of ENGLER and DRupe’s Vegetation der Erde is a mono-
graph on the vegetation of Chile by Dr. Kart Reicue? of the National Museum
at Santiago. This is the first volume of the series to deal with American vegeta-
tion. RercHe’s long first-hand acquaintance with the Chilean flora makes this
contribution a masterpiece, and all the more since fourteen years of effort have
been spent with this volume in mind. Chile is to botanists the best-known part
of South America, partly by reason of its peculiar accessibility, and partly by
reason of the large number of foreign botanists who have made Chile their home,
for a time at least. Among those who have contributed largely in recent years, and
thus made REIcHeE’s work more readily possible, are Puitippi, JoHow, NEGER,
and DuséN. Of particular importance is the work of R. A. PHILIPPI, who was
active for over half a century, and who died in 1904 at the age of 96. A short
account of botanical investigation in Chile forms the introduction to the work,
and there is given a bibliography of Chilean botany comprising 550 titles, of which
R. A. Putuirpr and his son contributed nearly one hundred.

No country in the world presents distribution problems of greater interest than
those of Chile, as may be suspected by reason of climatological variation. The
wer tion ranges from that of the desert of Atacama in the north, perhaps the
driest of all deserts, to the rain forests of the south, where there is a rainfall of
25o™ per annum. In northern or tropical Chile (18°-30°) there is the region of
desert where there are no marked seasons, and where agriculture is confined to the
oases and river banks. In central or subtropical Chile (30°-38°) there are sharply
marked dry and wet seasons, and the vegetation varies from steppes northward to
sclerophyll forests southward. In southern or temperate Chile (38°-55°) there
's a sharp distinction between the very rainy coastal district, where seasonal
Ui are relatively slight, and the interior, where the climate is dry and where
. Winters are cold and the summers hot. This rainy coastal strip is characterized

Y evergreens (temperate rain forest), while there is a strip of deciduous forest
pricaneg the only such forest of consequence in the southern hemisphere) in the
Peisay Periodic climate to the east. It will be noticed that this distribution of

€parts from that given by Scummper in that the deciduous forest lies east
ér than south of the evergreens.
Ne arate one

tT
— a UnD Drunk, O., Die Vegetation der Erde. VIII. Rercue, Kart,
Mb nzenverbreitung in Chile. pp. xiv+374. maps 2. figs. 55. pls. 33.
Leipzig: Wilhelm Englemann, 1907. M30.

74 BOTANICAL GAZETTE | [JANUARY

The second part of the volume presents a detailed account of the most impor-
tant families of vascular plants and their representatives, the vegetation forms,
the formations, and the “biology” of the representative plants. Among the
more interesting of the forest trees are the beeches (Nothofagus), of which five
species are deciduous and three evergreen, and the conifers (Araucaria and Fitz-
roya); the latter forms swamp forests, perhaps comparable to our tamarack
swamps. These conifers and beeches sometimes form pure forests, but most of
the Chilean forests contain many tree species. Other important formations are
the bamboo (Chusquea) thickets, xerophytic acacia thickets, and steppes.

The most detailed portion of the volume is that presenting the floristic features
of the Chilean vegetation from north to south, and the delimitation of floral prov-
inces. Many endemic species and monotypic genera are found in the country.
The final chapters consider the relations of the Chilean flora to other floras
(notably those of California, New Zealand, and Argentine), the life-history of the
Chilean flora, and the modifications due to human influence. From the develop-
mental standpoint the flora is made up of (x) a tropical contingent, the oldest of
all, dating from the Mesozoic; (2) the Andine contingent, a xerophytic element
associated with the rise of the Cordilleras; (3) the Californian and Mexican con-
tingent; (4) the Antarctic contingent, mostly in southern Chile, and related to
the New Zealand flora; (5) the boreal contingent, perhaps the most interesting
of all, there being genera and even species in southern Chile that are common with
the far north; (6) ubiquists and littoral pantropists; and (7) adventives. Many
admirable plates add much to this important volume.—H. C. CowLES.

The pendulation theory

Now and then a geologist attempts to account for Permian glaciation within
the tropics by supposing that the poles have shifted their position during the
course of geologic history. Such theories are usually dismissed because they intro-
duce more difficulties than they dispel. A few years ago PauL REIBISCH, 40
engineer, laid before the Verein fiir Erdkunde at Dresden such a theory, known
as the pendulation theory. There is now presented by Professor SeroTH? of
Leipzig a detailed account of the theory, together with a new alignment of facts
of distribution. ‘The essence of the pendulation theory is that the earth swings
slowly to and fro upon an axis whose poles are in Ecuador and Sumatra. These
poles are supposed to remain fixed, but the axial extremities that we commonly call
the north and south poles are such for but a moment, speaking geologically.
It will be seen that Ecuador and Sumatra must have been in the equatorial
from the beginning, while for points now on the equator but go® distant from
these fixed poles (i.e. in the French Congo region and in the Pacific Ocean north
of Samoa), there may have been in times past any conditions between polar

IMROTH, Hetnricu, Die Pendulationstheorie. pp. xii+564. maps 27- Leip:
zig: Konrad Grethlein’s Verlag.] 1907. M12.

1909] CURRENT LITERATURE 75

and equatorial. According to this theory, past variations in climate in any given
place have been due to pendulation.

Glacial periods, such as the Permian and Pleistocene, have been developed
through a poleward swing of regions now temperate; while warm periods, such
as the Cretaceous and Eocene, have been developed by means of a swing toward
theequator. At the present time Europe and eastern North America are supposed
to be swinging southward and getting warmer, while western North America is
swinging northward. Pendulation causes constant redistribution in the oceanic
waters, by reason of the earth’s oblateness, thus accounting for the submergence
of coast lines.

The major portion of the volume is devoted to the presentation of the facts of
distribution in animals as related to the pendulation theory. It is claimed that
the various groups are more or less symmetrically distributed with reference to the
fixed poles, owing to the control exerted on migration by the swinging of the earth
on its axis of pendulation. One chapter only is given to plants, and in this chapter
chief attention is paid to the conifers and Campanulaceae. Three maps are
presented, showing the distribution of the conifers. In these and other maps
southern Europe figures largely as a center of origin of forms and a center of
migration, and the attempt is made to show that migration has taken place sym-
metrically from that region.

The volume as a whole has a strangely medieval atmosphere. Students of
geographic distribution in these days are so accustomed to look carefully for
facts that they have largely ceased to care for hypothetical disquisitions such as
that of SmmorH. One feels that the author regards the pendulation theory as a
panacea, and that he selects for consideration those facts of distribution which
fit it best. Certainly the problems of migration are vastly more than the sym-
metrical movement of organisms from a center under the control of the direction
of pendulation. And the idea of pendulation itself seems more like an iridescent
fancy than a reality. Biologists may well wait until there is some astronomic or
geologic basis for such a hypothesis before they attempt to readjust their facts to
the new theory.—H. C. Cowzxs.

MINOR NOTICES

Purple bacteria.—A monograph on Rhodobacteria’ is the natural outcome
- the results of shorter studies on the subject which have been presented from
lime to time by Moriscu. After a discussion, partly historical, of methods of
ire, the author describes eleven new species recognized by him and gives a
th cation, based upon those of WINoGRADSKY and Mrcuta, in which he divides
e onder Rhodobacteria, containing all known purple bacteria, into two families:
T Which do and those which do not show sulphur granules in the cell substance.
“ning to the biochemical side of the study, Moriscu examines the relation of

rapier x ‘

$ Motiscx, Hans, Die Purpurbacterien nach neuen Untersuchungen. pp. 95- plse
4. Jena: G. Fischer. 1907

76 BOTANICAL GAZETTE [JANUARY

these organisms to light, oxygen, and organic substances. With regard to light,
the purple bacteria do not ordinarily show positive phototaxis, but are incited
to motility which continues for some time after the light is removed. ‘They are
not able to obtain carbon from carbon dioxid in the presence of light. Some
forms are even anaerobic, and, unlike most pigment bacteria, can produce pigment
under this condition. As to the pigment itself, Moriscu distinguishes two kinds:
the red (bacteriopurpurin) and a green (bacteriochlorin). ‘The latter is distinct
from cholorophyll, which fact agrees with that of their inability to use CO,.
Mottscx# concludes that nutrition from organic substance is somewhat related
to light and the presence of pigment as shown by the increased energy caused
by light; and that thus these forms stand between the colorless bacteria and
the green algae——Mary HEFFERAN.

The typhoid-coli group of bacilli—Numerous methods have been proposed
for the ready separation and identification of the typhoid and the colon bacilli in
water. Such special media as LOrrLer’s malachite-green, MACCONKEY’S lac-
tose-bile, Enpo’s lactose-fuchsin, and Conrapi-Dricatsky’s crystal-violet, have
been more or less successful in the hands of various workers. These are based
upon substances which restrain the growth of one type of organism while allow-
ing a characteristic development of the other. Ducamp‘ proposes for this purpose
the use of an “‘antibacilliary” broth prepared by cultivating in a lactose-peptone
solution several strains of B. coli, for example, derived from different sources.
This broth, when finally filtered germ-free, will be exhausted as a medium for
B. coli, but will still allow the growth of B. typhosus. For the rapid detection of
the latter in water, the sample is first plated in phenol broth and inoculations
made from the colonies into lactose broth. If a race thus obtained grows in the
anticoli and not in the antityphoid broth, and is agglutinated 1:50 by typhoid
serum, it is undoubtedly B. typhosus.

Studies on the fermentative activities of the typhoid-coli-dysentery group
resulted in the confirmation of some facts already known, and brought out some
new aflinities. B. para-typhosus, B. enteritidis, B. psittacosis Danysz and
Thomassen, and hog cholera ferment the same sugars except for two races of hog
cholera, which are inactive on xylose, dulcite, and mannose. B. para-typhosus

in addition ferments saccharose and raffinose. B. para-typhosus A differs
with respect to xylose, mannose, and dulcite—Mary HEFFERAN.

NOTES FOR STUDENTS

Subterranean fungi.—Ep. Fiscuer has recently made a contributions to the
morphology of the fungi. The paper is based on the study of material collected

4 aides Louis, Contribution 4 étude de la différentiation du colibacille et du
bacille typhique. Action des bacilles du groupe ee sur les
sccuien de carbone. pp. 181. pi. r. Thesis. Lille. 9
- Pee FiscHEer, Ep., Zur Morphologie der ‘eiciea: Bot. Zeit. print

1908,

1909] CURRENT I.ITERATURE 77

by Dr. W. A. SETCHELL and Dr. N. L. GARDNER in the region of Berkeley, Califor-
nia, during the years 1903-1905. It is of especial interest and importance in
view of the fact that so little attention has been given heretofore to the collection
of subterranean fungi in North America, and because this new material, some
of it in young stages, has enabled the author to put some of the imperfectly
described genera of HARKNESS on a better footing, and to revise some of his
own opinions as to the systematic position of certain genera which have occupied
an unsatisfactory position.

Some of the more important results are as follows: Myrmecocystis cerebriformis
Harkness and M. candida Harkness are shown to be identical, the former being
an older and mature stage, while the latter is unripe material of the same species.
The former name has precedence. M. yrmecocystis Harkness (1899) is also shown
to be generically identical with Pseudogenea vallisumbrosae Bucholtz (1900), and
the latter becomes M. vallisumbrosae (Bucholtz) E. Fischer. Young material
of Piersonia, a genus imperfectly described by HARKNEsS from old material,
shows that this is a very interesting genus. ‘The small nests of asci in the interior
of the fruit body are arranged in separate, pouchlike segments of hymenia, with
the free ends of the asci facing open passages or chambers terminating the venae
exlernae, the point of junction being rather abruptly narrowed. Paraphyses are
irregularly distributed among the asci in groups or partly wanting, but line the
Surface of the venae externae. The latter in the deeper parts of the fruit body
are filled with a loose weft of hyaline hyphae developed from the ends of certain
of the paraphyses; while toward the external portion of the fruit body brown
hyphae are intermingled and become more abundant as the openings of the

externae are reached. In the arrangement and form of the asci Piersonia
tesembles Pachyphloeus, the absence of hyphae in the hymenia-lined passages
recalls Hydnotrya; but the most characteristic feature in which Piersonia differs
his: ae meee is the sharply localized condition of the hymenial parts, since
inna er But i th xternae are lined throughout by the hymenium.
“é this respect Piersonia represents a special type at one extreme of an arm in the
nes Eutuberineae, in which the ascus hymenium has disappeared from a large
ea ok the vende externae and is found only at the innermost terminations of
. olding of the same. FiscHEr suggests that Piersonia may give the clue
proper interpretation of the structure of Choiromyces which he has formerly

P red with the Plectascineae, where it certainly occupies a rather anomalous
Position with its distinct hymenium. His suggesti is that the large, irregular
Pouchlike hymenial portions in the fruit body of Choiromyces may, like the

: ones of Piersonia, stand at the terminations of the venae externae, which
'n Choiromyces h

aS a suggestion. It will require developmental studies of
decide the point. Should this prove to be the true interpreta-
Some support to MatTiroto’s view that the difference between
*s and Tuberales is not fundamentally sufficient to warrant their

78 BOTANICAL GAZETTE [JANUARY

separation into two groups, although FiscHER has contended, and still maintains,
that there is no intermediate type between the two groups; since if his suggestion
as to the interpretation of Choiromyces proves to be correct, this anomalous struc-
ture would indicate subsequent modification of the venae externae and not an
ontogenetic connection with the Plectascineae.

Pseudohydnotrya, founded by FISCHER in 1896 on material from California,
he now finds is not related to Hydnotrya Berk. and Broome, a member of the
Eutuberineae, but is generically identical with Geopora Harkness, which is closely
related to Hydnocystis Tul. The fruit body of Hydnocystis possesses a single large
hollow space which opens to the outside, though the opening is filled with hairs.
The wall of the hollow space is clothed with the hymenium. Geopora is a Hydno-
cystis in which the hymenial walls are deeply infolded in an irregular and com-
plicated manner, in some species closed from the outside, in others communicating
in some places with infoldings of the external walls. All recent students of this
group agree in placing Hydnocystis among the Pezizaceae, and FIscHER locates
Geopora here also, although he formerly placed these two genera among the Bal-
samiaceae, where they occupied an anomalous position.

e of the very interesting forms proved to be the type of a new genus, Pseudo-
balsamia, which resembles Balsamia in external appearance, but differs in the
presence of venae externae which open to the outside, thus agreeing with the
Eutuberineae. It also differs from Balsamia in the absence of distinct trama
plates or veins (venae internae), or rather in the masking of them by the irregular
distribution of the asci among the tissue elements. In this latter character it
resembles the Plectascineae. The venae externae, however, are lined with para
physes, and occasionally asci are found in this layer parallel with the paraphys¢s-
Pseudobal ia, then, is regarded as Cee ee , in which by secondary
modification the asci have withdrawn from their regular position in a hymenium
and have become intermingled with the elements of the trama, thus simulating oné
of the characters of the Plectascineae, without showing any ontogenetic connec
tion with that series. This leads FiscHER to regard Hydnobolites, formerly
placed by him in the Plectascineae, as one of the Eutuberineae, since the hymenium
has probably undergone a similar modification, and the venae externae open to the
outside. This view of the relationship of these two genera is strengthened by the
well-known fact that the asci are often distributed in the trama in certain species
of Tuber, as in T. brumale, and T. rufum; while BucHoxtz has shown that
the development of T. puberulum the tissue corresponding to the trama ares
(venae internae) become compressed and changed by the pressure of the develop”
ing asci.

The modification which FiscHer’s views on the systematic arrangement of
the ascomycetous Hypogeae have undergone as a result of this study are exP
ina résumé. Briefly this is as follows: :

I. The Plectascineae series, with asci scattered in the tissue of the interiot
iat fruit body, or in groups, not forming hymenia, includes the two families

aphomycetaceae and Terfeziaceae. In the latter family remain the gene™

1909] CURRENT LITERATURE 79

Eoterfezia (as a simple form), Terfezia, Tirmania, Terfeziopsis, Picoa (incl.
Phaeangium), Delastria, Delastreopsis (as higher differentiated forms). Genabea,
Choiromyces, and perhaps also Hydnobolites and Pseudobalsamia are excluded
from the Plectascineae, and probably go to the Eutuberineae series.
2, The Balsamiaceae, with asci in definite hymenia lining the walls of chambers
closed to the outside, includes the single genus Balsamia. Hydnocystis and
Geopora go to the Pezizaceae. (See Ep. FiscHER, Hedw. 30:56-60. 1898.)
3. The Eutuberineae series, with hymenia lining the walls of interior passages
which open to the outside and are either hollow or more or less filled with hyphal
welts (asci rarely withdrawn from hymenia into the trama), includes the gymno-
carpic forms and are probably derived from the simpler Helvellales like Rhizina
and Sphaerosoma.
Since the second series is represented by the single genus Balsamia, with a
hymenium lining interior passages of the fruit body, one is led to inquire if it
would not be a more satisfactory arrangement to recognize two series: (1) the
Plectascales as outlined above, and (2) the Tuberales, including the Eutuberineae
and Balsamia. May it not be possible that Balsamia has been derived from
= of the Eutuberineae by a secondary modification of such a nature that the
Interior passages have become closed from the outside; just as in Geopora, as
FISCHER points out, examples occur in which such a secondary modification has
oe taken place? The development of Balsamia should be studied with
view.

Among the basidiomycetous Hypogeae several collections of Hysterangium
furnish additional evidence of the existence of a Hysterangium-Clathraceae
De ing with Gautieria, and then passing from Hysterangium through

allogaster, Protubera, etc., to the Clathraceae. :

HP paper abounds in speculative discussion as to relationship and ontogeny,
hoe Is a characteristic of FiscHER’s contributions. In a number of instances
pita seem to be based on rather insufficient evidence, which is perhaps the
e verse criticism which may be made on this contribution. Some of them
ae to be well founded, and certainly his present views on the classification of
or iene Hypogeae are to me much more satisfactory than his arrange-

t in ENGLER & Pranti’s Pflanzenfamilien. It should be said, however,
all his Suggestions and speculations are stimulating to thought, and I trust
oe collectors and investigators to bring to the light the riches in
Gro ean fungi which are awaiting us in this large field of North America

TKINSON.

will
ce

acc, —— basis of Mendelism.—Grécorre’ has published a critical discus-
the interpre t cytological theories, with particular reference to their bearing on
tation of Mendelian phenomena. Certain fundamental hypotheses

can ge

6 2 £ £

hes ot. V., Les fondements cytologiques des théories courantes sur Vhérédité
nne. Ann. Soc. Roy. Zool. et Malacol. Belgique 42: 267-320. figs. 4 1907

80 BOTANICAL GAZETTE [JANUARY

will be mentioned first. His view of the individuality of the chromosomes is based
quite largely on his well-known observations, and those of his pupils, on the resting
stage of the nucleus compared with the late telophase and early anaphases of
mitosis. The attempts of Fick,’ TELLYEsNiczky,® and others to disprove indi-
viduality, come in for pointed criticism. GRfcorrE concludes that ‘II est certain
que les chromosomes persistent dans leur individualité, sous la forme de continus
structuraux, a travers toute l’ontogénése.” Regarding reduction, he considers
it certain that the heterotypic mitosis dissociates the » chromosomes, received
by the reproductive mother cells, into two groups of m/2; and probable that a
paternal chromosome always conjugates with a maternal of the same form. But
he finds nothing to prove that a pair of allelomorphic characters is fixed only in
one pair of chromosomes, nor that the chromosomes conjugated in the heterotypi¢
gemini are homologous maternal and paternal chromosomes.

The hypotheses necessary to explain Mendelism on a cytological basis are
given as follows: (1) The chromosomes play a preponderant réle in the trans-
mission and determination of hereditary characters. (2) The different chromo-
somes of a given cell are bearers of different properties. (3) In the chromosomes
of a hybrid egg a Mendelizing character is represented only by two chromosomes,
one maternal, one paternal. In one of these the character is represented in 4
recessive condition (modalité recessive), in the other in the dominant condition.

It must be said (and Grécorre would probably agree with this) that the con-
ception of representative particles in the dominant and recessive condition me
projects the phenomena of dominance back into the germ cell without attempting
an analysis of its meaning, or how it comes about, and hence explains nothing.
This appears to the reviewer to be a serious and probably fatal objection to the
last hypothesis.

On the basis of these hypotheses the germ cells would receive of each pair 4
single recessive chromosome (maternal or paternal) and a single dominant chromo
Some (maternal or paternal). In the prophase of the heterotypic mitosis the
chromosomes join in pairs, and observation favors the view that these are homolo-
gous maternal and paternal chromosomes. Then after reduction half the ger™
cells would receive a “dominant” chromosome, and half the corresponding “Te
cessive’ chromosome. We thus arrive at MENDEL’s conception, and the chances _
of meeting he described between germ cells are here conceived between chromo
Somes. Granting the three hypotheses then, Mendelian phenomena would be
expected to result.

In Pisum eleven or more pairs of allelomorphs have been observed and the
reduced number of chromosomes is only seven; which shows that in this case, #

* Fick, Betrachtungen iiber die Chromosomen, ihr Individualitat, Reduktio®
ve Vererbung. -Waldeyer’s Archiv. 1906; Vererbungsfragen, Reduktions- und
hromosomenhypothesen, Bastard-Regeln, Engeb. Anat. Ent. 1907.
* Tettvesniczxy, Zur Kritik der Kernstrukturen. Archiv. Mikr, Anat. 60%
681-706. 1902; Ruhekern und Mitose. Idem 66: 367-433. 1905.

1909] CURRENT LITERATURE 81

least, several characters must reside in one chromosome. The characters must
then be confined to separate particles or corpuscles of the chromosomes, and an
interchange of homologous particles according to chance during maturation would
give the Mendelian combinations. Many observers, including STRASBURGER,
ROSENBERG, ALLEN, and SCHREINER, have described such an interchange of
particles; but Gr&Gorre’s conclusion, which he has emphasized before, is that
nothing in the observations of the nuclear reticulum, the somatic spirem, or
the heterotypic spirem justifies the admission of representative particles, chromatic
or achromatic. The “‘chromomeres” observed particularly in the heterotypic
spirem, he considers not as autonomous granules imbedded in a substratum, but
merely as a substratum impregnated with chromatic material and rather regularly
alveolated, giving the appearance of a single or double row of ‘‘chromomeres.”’

Grfcorre further denies that there is an interchange of particles between
the parallel filaments in the double spirem stage, such as various cytologists have
described. These two facts, namely, the presence of autonomous particles and
their free interchange at some time during the reduction processes, would appear
to be essential to a cytological basis for Mendelian phenomena.—-R. R. GATES.

Liezic’

wh : :
€n present in excess. The classic study of these organs is DELPINO’s treatise

hes ie BELT proposed a similar theory for Acacia sphaerocephala, and from
necophiles, or ant
Darwin, Frrrz MULLER, TRELEASE, and ScHIMPER. BONNIER (1878)

incid
ity and Kerner (1878) regarded extra I g
mecophily > sage Beginning with the skeptical attitude taken toward myr-
Y VON THERING in 1894, there have been critical contributions by
Wee
9 Nie:
sheidungen NHUIS VON UEXKii1-GULpENBANDT, M., Extraflorale Zuckeraus-

und Ameisenschutz. Ann. Jard. Bot. Buit. II. 6:195-327- 1907-

82 BOTANICAL GAZETTE [JANUARY

Rertic, ULE,*° and others, all of which are out of harmony with the myrmecophile
hypothesis. The work of NrEUWENHUIS-UEXKULL confirms these more recent
views.

After a detailed account of extra-floral nectaries by plant families, the author
summarizes the data presented, and some of the chief conclusions follow.
structure and form of the nectaries do not favor the theory that they originated
as adaptations for ant protection; in many cases they specifically oppose such an
assumption, and their position on the plant (largely on the leaf undersurface)
is such as to be of no purposive significance. The secretions often begin late in
life, so that the plant is without protection in youth, when it is most needed. In
other cases the secretion begins in early youth and soon ceases, thus leaving the
plant for a long time without ant protection, if such exists. The nectaries usually

ar t ST Li ly during their period of activity, and are often
dry. The nectar of many species is avoided by ants and other animals.
view that the honey-seeking ants drive off crawling insects and other “unbidden
guests” that mutilate the flowers, robbing them of honey or pollen, is quite
untenable, there being no relation between mutilated flowers, ants, and extra-
floral nectaries. Floral mutilation depends on the structure and position of the
. flower or the weather; furthermore, most mutilated flowers produce as many seeds
as flowers that are not mutilated. The honey-seeking ants are not combative
and do not attack other insects on the plants they visit; indeed, these other insects
often attack and repel the ants. The nectaries, therefore, so far from being bene
ficial structures developed by natural selection, are harmful to the plants of which
they are a part, in that they attract insects of all kinds, which not only eat the
sugar but do harm in various ways. Observation showed that individual plants
which secreted little or no nectar are less harmed by insects than are those that
produce nectar.

This paper, in addition to other recent work, makes it clear that myrmecophily _
is a figment of the imagination, and the word should be dropped from botanical —
literature. Ants may “‘love” plants, but there is no evidence that plants “Jove”
ants. Plants inhabited by these insects, if it seems worth while to group them,
may be called myrmecophytes.—H. C. Cow es.

eecreat } ll
SCULCLE SU.

A Mendelian ratio and latency.—SuHuLi'' in a suggestive paper makes
further contributions to Mendelian theory. In certain bean hybrids three distin¢
units were shown in earlier papers‘? to be involved, namely, a pigment factor,
blackener, and a mottled pattern. In the last character a peculiar condition is
found, namely, the mottled pattern depends upon the presence of a mottling allelo-

*° See Bor. GAZETTE 44: 314. 1907.

*t SHULL, Geo. H., A new Mendelian ratio and several types of latency. AM
Nat. 42:433-451. 1908.

"4 ———, The significance of latent characters. Science 25:792. 1907; Some
latent characters of a white bean. Idem 25:828. 1907.

1909] CURRENT LITERATURE 83

morph in a heterozygous condition, the homozygous giving unmottled seeds.
This peculiarity results in a new ratio, 18:18:6:6:16, instead of the anticipated
27:9:9:3:16. Latency is held to mean invisibility and not inactivity or dormancy.

ATESON’s “presence and absence” hypothesis, in which the presence of any
character is said to be dominant to its absence, is believed to be of general validity;
and his's more recent terms “‘epistatic’”’ and ‘‘hypostatic,” as applied to the capa-
city of one unit to hide or be hidden by another, are accepted. Thus in MENDEL’s
original case, yellow in cotyledons is not to be considered *‘dominant”’ over green,
but dominant to the absence of yellow and “‘epistatic” to green, i.e., according
to SHULL, causing its “invisibility” but not its “inactivity.” This change of view
involyes some nice distinctions, but appears to obviate some of the difficulties
of the older view of dominance, especially in connection with ontogeny. Inciden-
tally all that remains of the Mendelism of MENDEL is his hypothesis of gametic
sh The superstructure erected upon this has grown in complexity with great

With latency thus clearly defined, four types of latency are discussed: (1)

y due to separation, in which an allelomorph when acting alone has no
extemal manifestation, and is only rendered patent by combining it with another
allelomorph.” This type of latency is not uncommon, and gives rise to such ratios

“

give such a to carry invisibly a light-yellow allelomorph. ‘This condition may
aracters mg as 12:3:1. (4) Latency due to fluctuation. Disappearance of
hilbeagas an 26 unfavorable conditions of nutrition, etc.; a very common phe-
Cases former! oad Sey discrepancies from the expected ratio. Some of the
classed here ee incomplete or partial dominance” would probably be
Morphic ¢¢ - ag may also rarely be modified by the failure of certain allelo-
combinations to form a zygote which will develop.—R. R. GATES.

Res; .
Y agiingy chromogens.—Patiapin"+ has devised a new, very simple, and
method to sh a deteciing the respiratory chromogens in plants. He uses
ow the wide distribution of these chromogens in the plant kingdom.

Ih 71 speci :
i sre Tanging from liverworts to dicotyledons, this method showed these

3 Bate
660, 1907, a Wiittam, Facts limiting the theory of heredity. Science 26:649-

4 Pattap :
agg IN, W., Die Verbreitung der Atmungschromogene bei den Pflanzen.

Ber. Bo
t. Gesells. 26a: 378-389. 1908.

84 BOTANICAL GAZETTE [JANUARY

chromogensin 67. Their existence in three of the other four species can be demon-
strated by other methods. He mentions various fungi that other investigators
have shown to contain chromogens, as well as various other higher forms. The
points in the literature of this very important subject are briefly and clearly stated.
The same investigator finds's that portions of leaves in a 20 to 25 per cent. j
h lution for seven days show a great increase in respiratory chromo :
over checks immediately taken from the plant, or those kept in distilled water for
the same length of time. Illumination during the treatment increases somewhat
the chromogen production. If this treatment is continued for 17 days in light,
the portions of leaves take on a bright-red color. The color he believes originates
from the oxidation of respiratory chromogens. He holds that the sugar greatly —
increases the respiration and therefore the respiratory chromogens. Whether
the chromogen shall become chromatic depends upon whether the oxidases
exceed the reductases in activity. In long-continued exposures this seems t0
occur, hence the red color. He believes that OVERTON’s explanation of spring and
autumn coloration of leaves is not complete with the consideration of low tempera-
ture (as lowering respiration) and abundant supply of sugar as the factors, and
considers the relative activity of oxidases and reductases on the chromogen prod-
ucts of respiration as very important—Wr1LtIaM CROCKER.

ear

Graft hybrids. —WinkLER™ has begun a series of experiments in the endeavor

to produce graft hybrids, such as the well-known Cytisus Adami is believed to be. —
€ uses for this purpose certain members of the Solanaceae and Capparidaceat. 4
The method is to graft one Species on another in the ordinary manner, and after _
the scion has “‘taken,” to sever the stem at a point where the tissues of bo ;
Species will be cut. Adventive shoots then grow out from this cut surface. These

shows simultaneously the characters of both parent species.—R. R. GATES.

5 PALLADIN, W., Ueber die Bildung der Atmungschromogene in den Pflanze?
Ber. Deutsch. Bot. Gesells. 26a: 389-394. 1908.

*© WINKLER, Hans, Ueber Propfbastarde und pflanzliche Chimiaren. es
Deutsch. Bot. Gesells, 25:568-576. figs. 3; 1907. |

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CONTENT

n of Ceratozamia :

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The Botanical Gazette
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Issued February 20, 1909

XLVII CONTENTS FOR FEBRUARY 1909 No. 2

RELATION OF SOIL AND VEGETATION ON SANDY SEA SHORES Sens TWELVE
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VOLUME XLVII NUMBER 2

BOTANICAL (AZETTE

FEBRUARY s909

RELATION OF SOIL AND VEGETATION ON SANDY
SEA SHORES

PEHR OLSSON-SEFFER

(WITH TWELVE FIGURES)

The studies on which the present paper is based were carried on
during a number of years on a variety of sandy sea shores on the Baltic
coasts, in Denmark, Holland, Scotland, and France, on the Mediter-
tanean shores, along the coasts of Australia and New Zealand, in
Hawaii, California, Mexico, and Central America.

tn this paper I propose to give some of the observations made by
myself regarding the conditions for plant life on coastal sand forma-
Hons, and also to compare these investigations with the accumulated
Tesults from studies on this subject obtained by others.

Although considerable attention has been given to some of the
Bey P portant points, I have as yet not been able to study in detail
n se “dually weighty ; and in trying to interpret several of the phe-

om na of the distribution of sand vegetation I have found myself
oe Nae preany problems, for the solution of which there is
ttle definite evidence at hand.
oe the factors that influence plant life, I have found it
hydrod = 40 Classify them into the following groups: atmospheric,
To = ape edaphic, topographic, and historical factors. ee
ine. heric factors all those are here referred, which directl
ture, the jj "5 Wrectation through the air. The atmospheric tempera-
Bite Aaah conditions, the variations in air moisture, the movements
kind, ead and the electricity are the principal factors of this
th Ydrodynamic factors I understand all those connected
* Water content of the substratum, and edaphic factors are

86 BOTANICAL GAZETTE [FEBRUARY

those relating to the chemical and physical quality of the soil. I call
topographic those factors which have reference to the external features
of the ground, and they affect indirectly more or less the factors belong-
ing to previous groups. Historical factors are those which in the
course of time exert an influence on the topography and thus indirectly
on the plant covering.

Another series of factors analogous to the physical factors influen-
cing plant life are the biotic. They are either zoobiotic or phytobiotic.
Of the former especially the influence of man has to be recognized in
connection with the study of vegetation on coastal sands. The
phytobiotic factors are those caused by the plants themselves, and the
mutual relationship of sand plants will be discussed in another paptt.

Atmospheric factors

It is impossible to determine the relative importance of different
factors influencing plant life, or to give one of these factors precedence
in rank before another, because this depends in different cases
different conditions. It may be said, however, that the whole group
of atmospheric factors is the most important, especially because of their
influence on the transpiration of the plants. Sand vegetation is
particularly affected by: (1) the large amount of heat absorbed by —
the sandy ground and reflected from its surface; (2) the intensity of
illumination, both direct on the open, unprotected formation,
reflected from the white sand; (3) the exposure to winds, which com —
stantly change the atmosphere. |

TEMPERATURE.—If we compare in a general way the temperalul® ‘
conditions in a few of the localities under consideration, we find :
there is no significant difference in results as regards development of
sand formations or their vegetation in cold and warm countries.
the dunes along the Gulf of Finland, where the vegetation is in 4 dor- :
mant state for at least three months of the year, the sand drifts bes
during winter, when the grains are covered with a thin surface of 10% ©
and aresmooth. The friction is less and they are able to move slowly
forward. The herbaceous vegetation is absent at this time,
deciduous shrubs are without leaves. There is consequently 0
to arrest the movement of the sand.

On the Queensland coast, with a semitropical climate, the vegeta: 2

1909] OLSSON-SEFFER—VEGETATION ON SEA SHORES 87

tion period extends over the whole year, but the sand drifts here much
better during the summer, because winter is the rainy season, and
when wet the coherence of the sand is greater. Plants like Ammo-
phila, Cakile, Salsola, and Atriplex, of which the parts above ground
die every winter on the Baltic coasts, grow all the year round on the
shores of Australia, and there is no difference, external or internal,
instructure. Convolvulus soldanella on the coast of Holland does not
differ from the same species on the shores of tropical countries, where
it is common (fig. r0). :

Dunes are formed in warm countries more generally than in cold
because of the longer periods of drought, which favor the drifting of
the sand. We refer to the enormous areas of sand formations in
Africa, both on the coasts and in the interior, in the deserts of Asia,
on the coasts of India and Australia, in the interior of the latter con-
tinent, on numerous tropical coasts as Java, Hawaii, etc. The dunes
which occur in really cold countries, as in certain parts of the United
States and northern Europe, are insignificant in comparison with the
former.

It would be of considerable interest to have correct data of tempera-
ite conditions from the various localities where the author has made
his studies of the coastal sands. The field-work on which this paper
's based, however, has been conducted for comparatively short periods
at each place, and the temperature observations made do not offer,
acs any reliable basis for comparisons. Official data, obtained
pes Sete stations on the coasts, are also unsatisfactory for —
buil aR purpose, because the instruments usually are kept close to
‘idk <2 onl vegetation, in more or less sheltered positions,
ay . of these facts the observations cannot serve for any defintte
ecccis, Sih i the real temperature conditions under which the
BE Soins .. eveloped. It is not necessary to burden these pages
sie a : lons of any general data of the kind referred to. The
differ mpared a great number of temperature statistics from

.__ Tent coasts, but he has entirely failed to find any apparent rules
applicable to the devel ; se teat
elopment in general of sand vegetation 1n er

his negative result is not due to absence of such laws,
oie : must exist. It merely shows that our knowledge is
€ present method of taking temperatures at meteoro-

ent climates. T
Which certain
defic

88 BOTANICAL GAZETTE (FEBRUARY

logical stations is entirely inapplicable to the subject under considera-
tion.

For ecological purposes temperature observations in the field have
to be made very carefully if we are to draw from them any conclusions
of value as to the influence of heat on the vegetation. And further, they
have to be extended over a long period of years before we are justified
in advancing any general laws of temperature influence on distribution
of plants.

Let us here draw attention to the opinion held on this question by
the greatest authority on ecology, Professor WARMING. In his re
nowned handbook on ecological phytogeography (14) he says on page
22, speaking of the many attempts to determine the sum_-totals of
temperature in relation to geographical distribution, that these investi-
gations need in a very high degree to be supported by really scientific
experimental determinations of the cardinal temperatures for the
phenomena of different species. And even the results of such observa
tions would hardly be sufficient for the solution of the very difficult
and complicated questions of the importance of heat conditions for
distribution of species and phenological phenomena, as other factors,
perhaps, to some extent can replace a higher temperature.

One feature of the heat conditions on coastal sands is apparent.
That is the great fluctuation of diurnal temperatures. On account
of the low specific heat of sandy soil, the surface layers are rapidly
heated by the sun in daytime and as quickly cooled by night. These
variations of temperature are conducted by radiation to the lowet
strata of the atmosphere, or those in direct touch with the plants
which consequently are greatly affected by such changes.

Of some occasional observations by the writer on the diurnal rang?
of temperature on’ sand dunes the following may be mentioned 4S
examples of the great divergence between temperature extremes in
such localities.

Observation 64.—Dunes at Hangé, Finland, September 10, 1897. Maximum
temperature in the shade (thermometer from unknown maker) 28°8 C., betwee®

6 A.M. and 6 p.m Minimum (thermometer from WALLMANN in Stockholm) o

2°6 between 6 p. M. and 6 A. M. Range 25°6. The instruments were pl :
on an open sand surface 25°™ above the ground, and were shaded by white
canvas, 2™ high. Ordinary thermometer readings in the shade were taken
every hour in the daytime, giving the following results:

1909] OLSSON-SEFFER—VEGETATION ON SEA SHORES 89

6am. 8°5C. 10 A.M. 14° C. 2P.M. 23°6C.

7 A.M. 9.2 tt A 37 3 P.M. 27.2

8Aa.M. 9.8 I2M. 19.4 4 P.M. 27.5

g A.M. I1.4 I P.M. 20.6 5 P.M. 22.4
OP tS 6

This place of observation was on an open spot, unprotected from the winds.
About 60™ inland between the pines, and sheltered from winds, the thermometer
gave 7°4C. at 6 A.M., 9°2 C. at 9 A.M., 16°6 C. at noon, 28° C. at 3 P.M,
16°2C. at 6 p.m. This shows that the temperature was slower to rise in the
morning and forenoon, but once high it was also slower to decrease, when the
energy of the solar rays was diminishing toward evening.

Observation 576.—Dunes at North Beach, near Perth, Western Australia,
September 12, 1902. Maximum in the shade 31°1 C. (8 A.M-6 P.M.). Mini-
mum 6°8C.(6P.mM-8a.M.). Range 24°3. The instruments, from NEGRETTI
& Zawpra in London, were elevated 30°™ above ground and shaded by a white
canvas tent with open sides. The hourly variation was the following:

8a.M. 7°9C. 12M. 23°6C. 4P.M. 19°5 C.
“QO A.M, 11.2 I P.M. 29.4 5 P.M. 16.3
Io A. M. 15.6 2 P.M. 30.7 6P 9.6
II A.M. 18.6

3-P. M. 20.3

These data show, as in the previous observation, that the temperature rose
steadily until about 2 P.m., although the rise is so much more rapid in these
latitudes on account of the greater energy of the sun. In this case, however,

Over 10° in a single hour, and nearly 10° more between 4 and 6 P. M.
si the coast of Western Australia a sea breeze always sets in about noon,
ard to Cooke (4), the temperature then begins to fall until evening, and
are generally cool the whole year round. :
The influence on vegetation of such a wide range of temperatures
Must necessarily be of considerable importance. Although these air
smperatures have been largely affected by radiation, the direct radi-
ant heat of the sun is still more important. Actinometric methods
oe Tegistering intensity of solar radiation are, as yet, very unsatisfac-
saree only instrument available for field observations is
are. lack-bulb thermometer in vacuo. ‘The rather casual
another = . this kind made by the writer will be referred to in
May ie ace In connection with some transpiration phenomena. It
Jeeta however, to say here that these occasionally taken
Writer 4 in spite of their discontinuity, have convinced the
lems of heat = i and most effective way of attacking the prob-
ines of Saeiee dogg hay hs scat a is to pursue investigations on the
air nor the — records. Neither the mean temperature of the
m-total of atmospheric temperature is of such importance

go BOTANICAL GAZETTE [FEBRUARY

to vegetation as the amount of direct solar radiation and radiation of

heat from the ground. The value of the former factor in climatology
has long been recognized by meteorologists.

We must never lose sight of the fact, however, that it is not one
factor alone that determines the development and distribution of
the vegetation, but a resultant of the many different conditions to
which the plants are subjected. This has been duly emphasized by
WARMING (14).

Licut.—In the closest relation to atmospheric temperature, and
especially to radiation of heat, is the factor of light. The intensity of
illumination is remarkably large on the open sand formations of the
coast, and this circumstance is noticeable in the many protective adap-
tations of plant structures against the influence of light. There are
as yet no reliable means of ascertaining the intensity of the light, and
we have consequently no basis for comparisons on this subject. The
strong insolation on the white surface of the sand favors a greater varia-
tion of temperatures than on other formations of the coast. The radi-
ation is, however, generally less on the coast than in localities far away
from the sea because the larger quantity of aqueous vapor in the
atmosphere in the former place tends to check this terrestrial radiation.

HYDROMETEORIC CONDITIONS.—This term has here been used 10
distinguish the factors of atmospheric humidity from those of the watet
content of the soil or the substratum from which the plants take their
supply. On coastal sands it is perhaps more apparent than on othet
formations that there is some difference in influence and effect 0?
vegetation of the moisture contained in the air and of the water in the
soil. It is, however, always extremely difficult to decide to what extent
certain adaptations are due to one factor more than to another,
especially when we do not possess detailed observations.

AIR MOISTURE AND EVAPORATION CAPAcity.—The supply of
atmospheric moisture is to a great extent obtained from the ocea?
and it follows that on the coast the amount of humidity must be much
greater than farther inland. This is in fact an everyday observation.

We do not know for certain to what extent the plant is able to con
dense water vapor and absorb atmospheric humidity, but we do know
that moisture in the air greatly lessens transpiration, and, other factors
being equal, transpiration ought therefore to be less on sea shores than

1909] OLSSON-SEFFER—VEGETATION ON SEA SHORES gi

inland. Now it is not the absolute humidity that determines the
amount of evaporation, but the saturation deficit or the amount of
water which the atmosphere at a certain temperature is able to absorb
(Warminc). In the continental interiors the evaporating power of
the climate is very great during summer, and on account of the cold
quite inconsiderable during winter (HANN 7). If we compare the
habits of certain plants which occur both in the interior deserts, and
on coastal sands we shall find them instructive. For instance Mesem-
bryanthemum aequilaterale has the same appearance when growing
on the beach or in the immediate neighborhood thereof, as when occur-
ring on the interior sands of Australia, many hundreds of miles from
the coast. But when found at a certain distance, 0.5*™ or so, from
the beach, outside the influence of the spray from the sea, and sheltered
from the full force of the ocean, it grows higher, its leaves are two or
three times as long, and its succulence is less marked than in specimens
from the beach or from the interior. In the case of the writer’s obser-
vation, the places of growth were as nearly similar as possible with
regard to exposure to the rays of the sun and moisture in the soil.
Some notable differences in anatomical characters observed in this
connection will be alluded to in another paper. These characters also
Proved that the devices for protection against excessive transpira-
tion were not so well developed in the specimens which were not
exposed to the strong salt-laden winds, although they had the full
benefit of the coastal moisture.
i allowed to draw any inferences from this fact, we
a ae e that, as a rule, ¢ranspiration is less on coastal than on
ie : There are other factors, however, ‘to be taken into
ri ee somewhat equalize the conditions, as the wind
Bester o. the atmospheric moisture. a ik
i ie 1, > warm coasts, especially in the neighborhoo of the
terized by s oy we can notice a peculiar sand flora. It is charac-
hand am which do not occur farther inland, and on the other
eas ot ag as a rule do not go down to the beach. The great-
ith sis. strand flora consists of halophytes or plants growing
: ations.
Sai assumed that the conditions in which halophytes
pendent on the amount of salt present in the soil. As

92 BOTANICAL GAZETTE [FEBRUARY

we shall show, when speaking of the edaphic factors, the quantity of
salt present in the soil in some of the coast formations is quite incon-
siderable, but still the plants exhibit the same characters typical of
those which grow where salt occurs in the soil.

We cannot here escape the conclusion that the influence which is
exerted on the plants in one case by the salt-water content in the soil,
in the other case is brought about by some other cause of similar kind.

Analysis of spray from the sea reveals the presence of a large amount
of salt, usually even more than in the sea water, because water is
evaporating from the drops of spray and the fine particles of moisture
near the ocean. This salinity of the spray is greater at a high tempera-
ture, as the evaporation is then more intense, and it is common to find
on hot days, with sea wind carrying moisture landward, that salt is
deposited in form of crystals on the plants and on other objects, as
well as on the ground.

A few analyses will be given to illustrate this large quantity of salt in the
atmosphere on the sea coast. In all instances when samples of spray were
secured, the method was as follows. Pieces of muslin, thoroughly examined
and found free from salts, were dried and kept in tight-closed jars until expo
at the place were the sample was to be taken. The muslin was then exposed
to the sea spray; the temperature and time of exposure were registered, the
muslin bottled, and later examined in the laboratory. At the same time sam-
ples of the sea water were secured for chemical analysis. The following results
were obtained from four observations:

I. Nagu, Hégsar, Finland, August 22, 1897. Atmospheric temperature
19°5 C. Sky clear. Velocity of wind about 7™ a second. Muslin expo for
15 15™, at a distance of g™ from the water. Salinity of spray sample 0.673
per cent., of sea water 0.662 by areometric measurement, and 0.632 per =
by chemical analysis. Temperature of water 15°0C. As is the case in this
observation the areometric value of salinity is always somewhat higher than
that obtained by analysis.

Il. Hangé Tulludd, Finland, September 9, 1897. Atmospheric temperature
21°2C. Sky partly overcast. Velocity of offshore wind about 1o™ a secon’
Muslin exposed on the beach, 5™ from water, for 2. Salinity of spray sample
0.625 per cent., of sea water 0.607. Water temperature 13°6 C.

Ill. North Beach, near Perth, Western Australia, September 16, 1902. Atmos:
pheric temperature 24°6C. Sky clear. Velocity of wind, measured with
anemometer (of Crova type, from Necrett1 & ZAMBRA, London) averaging
12.3™ a second during time of observation. Muslin exposed on the beach,
from the water, for 4 aS”. Salinity of spray 4-68 per cent., of sea water 3-24
per cent. Temperature at the surface 10°6 C.

1900] OLSSON-SEFFER—VEGETATION ON SEA SHORES 93

"* TV. Beach at Pialba, Queensland, on the eastern coast of Australia, June 17,
igor. Atmospheric temperature 16°C. Sky clear. Velocity of wind about
6" a second. Muslin exposed 5™ from the water. A heavy surf was rolling at
the time, but as the water is shallow far out from shore, and sheltered by clumps
of mangrove, the breakers did not strike the shore with any force. The sample
showed a salt content of 4.1 per cent. Salinity of ocean water 2.91 per cent.,

and temperature 9°4 C.

Although the presence of sodium and chlorine, as common salt,
can be shown in many inland plants, a larger percentage of these salts
is found in the ash of strand and marine plants than in that of the
former type.

Whether these salts are absolutely essential for any plant we do not
know for certain. If that is the case, the amount of salt needed is very
small, as has been shown by several investigators. Even for many
marine algae only the smallest quantities of salt are necessary, if at
all essential.

Inland plants are, however, unfavorably influenced by a percentage
of salt which strand plants bear without injury; on the other hand, it
has been proved by cultures, that the halophytes can grow without the
usual amount of salt contained in the soil or atmosphere of their
natural habitat. |
ot experimental study of strand and other plants with

He found that 1.5 per cent. of common salt in soil or in water is
Polsonous to plants which do not naturally grow on the sea shore.
.. — about 2.5 per cent. of common salt, and the soil
still ck € at as well as the atmosphere near the coast, contains
te than this proportion. We can thus readily understand the
ee i which separates the marine and strand floras from those of
ie: enor. Covuprn attributes the poisonous property of sea water
a. mainly to its content of common salt, for the ~~
chlorid, are iis a abundance, magnesium sulfate and magnesium
Proportions ores aaa which he considers below the toxic
Per cent. m agnesic sulfate is poisonous at a concentration of 1
Water me agnesic chlorid at 0.85 per cent., but they occur 1 sea
tively y to the extent of o. 75 per cent. and o.5 per cent. respec

Th : ‘
© question of the influence of salt on strand plants and of the

94 BOTANICAL GAZETTE [FEBRUARY

absorption by the plant of saline water has been discussed consider-
ably, and several theories have been advanced.

PRECIPITATION is a factor, which on naturally dry soil, such as pre-
sented by sand formations, is of considerable importance, not only on
account of the quantity of water which in this way is brought to the
plants, but also because of its influence in giving coherence to the sand,
thus preventing it from shifting, and because of its weathering action
on the soil particles.

That precipitation must to a great extent influence the develop-
ment of vegetation on marine coasts is evident, and it is easily seen
that the composition of the plant covering of sand formations varies
somewhat in rainy and rainless climates, although the atmospheric
humidity on the coast tends to minimize this difference. The latter
factor is especially important as the precipitation often is so variable.
The absolute amount of rain during the year does not in fact give any
correct basis for comparison of the conditions in different localities,
as it is far more important for the vegetation how this quantity is
distributed over the period in question. On the Baltic shores the
number of rainy days of the vegetative season is much greater than,
for instance, on the coast of Australia or California. The easterm
coast of Australia, at Brisbane, has an average yearly rainfall of
129.5°™ while the Aland Islands in the Baltic have only 52.9% but
the vegetation in the latter region has a much more even supply of
moisture during the vegetative season, because the precipitation is
distributed over'a greater number of days, about 70 of the 160 rainy
days of the year falling in the growing season.

Edaphic factors

: When Scuimper proposed this term (1898) he apparently regarded
it as covering all the peculiarities of the ground. It seems to the
present writer that it would be more convenient in this connection
distinguish between the soil as such, and the media, water and aif;
filling the interstitial spaces. This distinction has been made in this
paper by separating the factor of soil moisture under the heading
hydrodynamic, and the factors pertaining to the soil proper as edaphic:

The hydrodynamic factors are now generally admitted to be of the
very greatest importance for the vegetation and its distribution. J

1909] OLSSON-SEFFER—VEGETATION ON SEA SHORES 95

have treated this subject in another place (12). With regard to the
edaphic factors, sensu scriptoris, a wide difference of opinion has pre-
yailed. While some writers have maintained that the chemical influ-
ence of the soil is the most important, others have been in favor of the
theory which gives to the physical properties of the soil the largest
bearing. Since the physical conditions mainly determine the amount
of water, most recent authors upon the subject, among them WARM-
ING, hold that these are of greater consequence.

In discussing the physical conditions of sand formations we will
confine ourselves to the question of soil temperatures, and further
briefly refer to some measurements of the size of sand grains made by
the writer.

Som. TEMPERATURE.—The heat-absorbing power of sand is low in
comparison with other darker-colored soils, but because the radia-
tion is great the vegetation on a surface of sand is subjected to a com-
paratively high temperature. As the sand is always moist only a
little below the surface the heated layer of sand does not reach deep
before it meets lower temperature.

The roots always penetrate to this moist layer, and only the upper
part of the root is under the influence of the heat of the surface
stratum of sand. We find corresponding adaptational protections
on the roots of most plants growing in sandy soils.
es Seslage of heat within the soil is influenced by several factors
. : _ from one layer to another. It is impossible,
nt a a e€ natural state of the soil to eliminate these factors, the
‘Sa : of water and air in the soil, the evaporation of water from
hie dain : and condensation of water vapor in the colder strata.
Sell nds: ae supplied therefore represent the temperature of the

“ar tions such as it presents in the field.
irda es made by the author ordinary soil thermometers
eas = n : ae temperature was taken at following depths: 9, 5
in sae — . The bulb of the instrument can easily be placed
ia. at the desired depth. The number of complete series
of these a: made on different sand formations reaches 876. Some
will be of ere be referred to, and in other cases average values

given,

On i
the front beach the temperature of the soil is varying more

96 BOTANICAL GAZETTE [FEBRUARY

than on other formations of the sand strand, because of the frequent
inundations and subsequent changes in evaporation. It is generally
low as compared with that on the higher parts of the beach. Seven
measurements taken at Hégsar, Nagu, Finnish Archipelago, in June,
1894, averaged 16°4 at a depth of 2°™. The corresponding data,
obtained at varying depths, are shown by Table I.

TABLE I
Jind velo- i
2cm. | 5cm. | 10cm. Air | Sein Time any — age
see nee 18.3 | 16.3 | 17.4 | 19.6 | 18.4 | 12:30 P. M. 1.5 2
ee re 14-9 | 15.5 | 16.3 | 13.8 | 16.6 | 2:00 P.M 0.5 6
Baer cee 19.6 | 18.9 | 18.6 | 19.8 | 16.7 | 11:30 A.M 4.1 4
fe aera 16.5 | 15.8 | 15.9 | 16.5 | 16.9 | 2:00 P.M. 0.3 g
Somes es 14.2 | 14.6 | 15.2 | 14.4 | 16.5 1:30 P. M. 2.6 1.5
Weil ies 13-3 | 13.6 | 12.9 | 14.0| 15.6] 1:30 P.M 1.8 4
, ee 16.7 | 15.4 | 14.7 | 16.9 | 16.5 | 2:00 P.M 0.4
Boe

In all cases except 3 the sand was covered with vegetation,‘in 5 and 7 with
Glaux maritima, in 2 with Erythraea litoralis, in 1 and 4 with Argentina anserin4,
and in 6 with Triglochin maritimum and Eleocharis uniglumis.

In September, 1902, similar measurements were taken on the
front beach at Freemantle, Western Australia. No vegetation
occurred on the formation. The following results were obtained
(Table II):

TABLE II :
Be
. d i .
2cm. | scm. | rocm.| 20cm.| Air —. Time ay, m. per gee
‘ sec.
pee
I 32.6 | 25.4 | 22.9 | 20.2 | 26 I :
3-4 | 9:00 A.M 2
2 37-4 28.7 | 27.1 | 24.9 3052 | t4.6:) 1:00 Pi 2 pes
3 mo-8 | 40-2.) 47.4.) 15.0 1 97.5 | ra,.9 | 2:00 P. 1-5 3
4 25-4 | 20.4 | 18.5 | 15.8 | 24.4 12-5 | 11730 A.M 0.3 .
5 21.3 | 15.6 | 14.2 | 13.6 | 22.7 | 12.8 12.15 P.M 35 7
na

The average temperature of these five series at 2°™ depth is thus
28°5 C., that is, 12°: higher than the mean of the previous series:
The sea water in the latter case was much colder, while the atmo>
pheric temperature was considerably higher.

Of sixty measurements taken in day time on the front beach undet
conditions as similar as possible, the highest temperature obtained for
2°™ depth was 42°6 C. on the Queensland coast at Pialba, in Dece™

1909] OLSSON-SEFFER—VEGETATION ON SEA SHORES 97

ber, 1901, while the lowest was 2°1 near Mariehamn, Finland, in
September, 1896. The average of these sixty observations was
1894 C.

The soil temperature on the middle beach is already much higher,
as following data will show. The observations were made at the
same time and in the same place as those mentioned in Table I.
The distance from the water was 6.5™ and the sand pure quartz of
medium size and yellow color.

TABLE III

2cm, 5 cm. ro cm. 20 cm. 50 cm.
Deeeeeee esa, 22.6 re ee 17.8 14.1 12.6
19.1 16.7 14.9 Lai} 12.1
3 23-4 ar.3 18.9 15-3 13-3
4 20.1 18.5 17.0 13.9 12.7
: 18.4 16.9 15.6 12.7 12.2
© heerecwny actos 16.9 15.2 13 | 12.6 IIl.9
r 19.8 1753 5.5 13.8 12.9

The vegetation consisted in x and 3 of an open community of the following
stituents:

Factes: Leontodon autumnale, copious.
SECONDARY: Festuca rubra arenaria, subcopious, Agrostis vulgaris, sub-
“pious, Plantago maritima, gregarious, Erythraea litoralis, sparse.
lutj : Erythraea was scattered about in patches, between low shrubby Alnus
: sme solitary individuals of Elymus arenarius, and Rosa canina.
oe 4, 5, and 7a Juncus Gerardi community occurred on the middle beach,
fe i Erythraea litoralis and Plantago maritima. In 6 Elymus arenarius
fuca rubra arenaria formed an open community.

fe parallel table to Table II shows the temperature conditions on
middle beach at Fremantle, W. Australia. The sand was here

= Consisting of pure light-yellow quartz. Time and atmospheric
Conditions as in IT.

eae aes TABLE IV oe
at _— 5 cm. ro cm. 20 cm. 50 cm.
Sree ERS pe: ecteeeer, ER ee panera at
Be en 2 5 ee 23 4 22.4 20.7 18.4
KE oe 38.6 25.8 24.6 23.6 20.0
a 29.5 18.6 18.3 16.5 15-8
oe 27.2 15.3 19.6 19.4 16.3
ae pe

98 BOTANICAL GAZETTE [FEBRUARY

A comparison with the results in Tables I, II, and III reveals the
fact that, while the surface temperature (2°™) in all cases was higher
in the series given in Table IV than in Table II, the temperatures at

-5°™ and lower were higher in the former case, presumably on account
of a more intense evaporation, which caused a corresponding loss of
heat. No such difference existed between Tables I and III, where the
solar radiation was less, and both the atmospheric and sea-watet
temperature lower. .

We shall now proceed to a statement of the temperature conditions

on the upper beach. Table V gives the results of some observations

made on that formation at Jerwe on the Oesel Island in the Baltic, in
July, 1896. The beach has a low grade and is limited landward bya
littoral dune in the shape of a steep and high bank, on the top of which
small dunes are developed. The sand on the upper beach at the foot
of this bank is rather coarse, consisting of a reddish-yellow quartz.

TABLE V

Wind velo- :

2cm. | 5§cm. | Iocm.| 20cm. Air = Time ‘ty, m. per Cloudiness
er sec.

aes Ueaaeee aoa e Re cai

I eo 2.4 | 22.8 | 21.4 | 19.6 | 21.6 B71) 30730 A, M. I 3

2 +++) 27-3 | 26.2 | 24.5 | 22.7 | ac.2 16.4 | 1:00 P.M. 0.5 :

3 | 26. 24.3 | 22.8 | 21.1 | 23.6 | 16.1 3:15 P. M. 2 5
wk ial ee mle

The next table is a continuation of the measurements given it
Tables II and IV from Fremantle, W. Australia, and the genet
conditions supplied in regard to those tables refer also to these obse™
vations on the upper beach, except that the time in each case was about
15 minutes later. The sand was of medium-sized, white-yellow quat™
mixed with an abundance of shell fragments.

TABLE VI

2cm. 5 cm. ro cm. 20 cm. a
: eS ais 34.8 24.5 22.6 21.1 ee
aoe 38.9 24.9 23.4 22.0 ae
a 28.8 19.2 17.6 16.4 a4
er a 27.6 22.5 20.7 18.2 4

seeeh pues 25.3 ba 16.9 14-5 +

As will be seen, the temperature on this formation does not differ

|
4
|
q
|

~ 1909] OLSSON-SEFFER—VEGETATION ON SEA SHORES 99

essentially from that on the middle beach at the same locality as given
in Table IV. The differences existing may be accounted for by the
topography and by the fact that the upper beach here was covered
with a sparse vegetation consisting of various low herbs.

On the littoral dune the temperature conditions are somewhat vary-
ing, usually higher on the landward slope, and a rise in temperature
can also be noticed with an increase in the height of the dune. The
summit and the landward slope of the littoral dune are frequently
covered in large patches with vegetation, and the temperature differ-

“ences between the open spots and those where plants occur are
considerable.

Table VII shows a series of measurements made on the seaward
slope of the littoral dune at Fremantle, W. A., under conditions similar
to those given for previous observations from that locality. The
sand was fine white quartz.

TABLE VII
ae 2cm. 5cm. ro cm. oe one
: eet ea
i Cee 35.2 26 4 23.5 21.5 19.2
; es | 38.8 27.3 22.6 20.6 18.4
oe ies 29.4 20.5 18.1 16.2 15-4
; = 28.9 21.3 19.4 17-7 10:3
25.6 1537 16.6 te al —s
ee i es

' Table VIII gives the temperature on the summit of the same dune,
ss ? height of 6™ over the ocean level. The dune material consisted
o medium sand, somewhat yellowish in color.

ee TABLE VIII

TUL rere en ook 5 cm. ro cm. 20 cm. Gee name

T —————

iste ak ee oa |

2 | 35-5 27.4 24.2 21.7 19.8

3 ts 38-8 28.1 23.0 21.2 18.9

4 39-3 21.0 18.2 16.8 15-7

5 "| 30.1 21.8 18.9 18.3 16.3
e364 18.8 16.7 15.7 £4.90

: On the landward slope, some 3™ from the top, the following meas:
Fements were obtained:

100 BOTANICAL GAZETTE [FEBRUARY

TABLE IX
2cm. 5 cm. ro cm. 20 cm. 50 cm.
Bi vie seep tea aie 35-4 27.0 23.6 205 19.4
re PENS 39-2 27.6 22.8 20.4 18.2
3+: 29.8 20.8 18.1 16.5 15.6
qe. 29.5 21.7 19.0 17.8 16.1
Eos 25.9 18.9 16.8 15.6 14.3

In this last case the sand was rather fine quartz, of yellowish color.
The maximum soil temperature measured by the author on a littoral
dune formation was obtained in December, 1901, on the leeward slope
of a high dune at Southport, Queensland, where the thermometer
2°™ under the surface registered 58°4C. at 2 p.m. The formation
was devoid of vegetation.

The temperature of the dunes and the sand fields varies greatly.
Some averages will be here given. Of the 34 readings made under
general conditions as similar as possible a mean temperature of 26°2
C. was obtained from dunes in Finland for a depth of 2°" and 25*4
far ee. The average of 19 readings at Fremantle, W. A., was
28°7 C. and of 12 readings at Southport, Queensland, 29°8. On the
dunes of North Cape, New Zealand, the author measured on the same
day within one hour the following series: 26.1, 25.8, 24.2, 27:
25.4, 26.3, 26.1, 2799 C. The atmospheric temperature at the |
time was 25°4, cloudiness 3, time December 7, 1902, 11:30 A- Me
12:30 P.M.

The daily variation of temperature must naturally be of some —
importance to the vegetation. Only a few observations have been
made by the writer to this end. One series will be given as a sample
of the extent to which such variations take place. The readings wet
made at Southport, Queensland, in December, 1901.

In the light of measurements obtained the local distribution of
certain plants on the coastal sand formations seems to indicate that
the temperature factor is of the greatest importance for the mode of
association of plants into communities. On the coasts of the Baltte
the writer made frequent observations which tend to show

this.

At Ahus in Sweden there occurs on the upper beach an Ammo
phila-Elymus community, consisting of the following plants:

1909] OLSSON-SEFFER—VEGETATION ON SEA SHORES IOI

Factes: Ammophila arenaria, Elymus arenarius.
Seconpary: Triticum junceum, Carex arenaria, Festuca rubra arenaria,
F. ovina, Poa pratensis, Cakile maritima, H. alianthus peploides.

TABLE X
Hour or | AtmosPHERIc DEPTH UNDER: SURFACE Cromer
READING TEMPERATURE reas CS ae
2cm. 5 cm. Io cm. 20 cm. 5° cm.
Oa, Bi. .6c: 9°0'C. 4.9 449 Say Q.t 14.3 6
eee eee 10.8 6.5 5.6 5-9 9.4 14.4 2
Beker. 11.4 7.7 6.9 6.2 9.4 14.6 3
ae eae 15.6 9.0 8.9 6.9 9.6 14.6 rs
ny Ce 17.7 it 3 10.2 8.0 9-7 14.5 4
trees 19.5 14.6 1r.8 6 8 14.5 3
12 NOON .... 23.2 1742 13.0 ee 5 Bae TAL 3
oe 26.3 19.6 14.3 11.8 10.7 14.4 5
sh oS ae 27.2 19.4 14.8 12.6 Li4 14.5 4
DN ae 27.0 18.1 14.6 12.7 11.6 14.5 4
‘ 26.4 18.0 14.3 12.6 11.8 14.4 5
é Se 25.1 57.3 14.0 11.8 Tx0 14.5 4
22.3 15.1 1 Tr4a I 14.6 a5
i ethic 18.6 12.9 129.4 10.4 II.9 14.6 x
whey 16.2 Ir.0 | 10.7 9-5 11.6 14.7 o-5

The two species which constitute the facies of this community
usually occur in small separated patches, and measurements of the
temperature in the small sand elevations formed by these plants
ree the fact that in the Ammophila patches the soil temperature
mie heey was two-tenths to six-tenths of a degree higher that
exact diff eeeees which would explain the lower temperature, but the
: erence in moisture has not been ascertained. Many similar
eo of temperature differences have been noticed. Halianthus
sn — grows in colder places than Argentina anserina,
temperatur ag often form a community. This question of
however dee fferences influencing the formation of communities,
tak be < : S further investigation before any decisive statements
influenced a = Punt also be remembered that the temperature 1S
on the ph ae SEAT, which in its turn depends to a great extent

ysical conditions of the soil.
ia . ANALYSIS OF SAND.—A considerable number of such
given to show iy nade by the writer, and a few series will here be
the various oY the differences in size of sand grains on
rmations on different sand strands. In the table the

102 BOTANICAL GAZETTE [FEBRUARY

letter A refers to a series of samples from Kurische Nehrung on the
northern coast of Germany, D to a series from sand formations near
Amsterdam in Holland, E to sand from the west coast of France
south of Bordeaux, F to samples from the west coast of Australia, neat
Fremantle, G to sand from Port Fairy, Victoria, H to the sand at
Southport, Queensland, J to a series from North Cape of New Zea-
land, and J to a series of samples from the Pacific coast of North
America near San Francisco.

ne, OE ae apa ace ees Medium Medium | Fine Medium
ME Gee ye Finest Finest
eer
eee
F G H I J
ee a wee
Submerged beach............. Coarse Medium | Fi Coarse Medium
Pronk Gi 2 fo Medium | Fine Medium | Grits Fine
Middle beer Mediu Medium | Coarse | Coarse | Fine
Upoet bea Coarse | Medium} Coarse | Grits Medium
Aattotel dane: 2255 2g Medium | Fine Medium | Medium | Fine
ME. ee ee Fine Fine Medium | Fine Fine
sand Held oe ae Finest
Beare

In each of the above cases the result represents the average of 10
samples, secured at approximately corresponding places on |
formation. As these data show, the coarsest sand occurs on the upp
beach. The material that builds up the littoral as well as the ordinal)
dunes is usually of the same grade of coarseness. It is only when We
analyze the sand from various places on the dunes that difference
appear, which explain the formation of ripple marks and dunes 48 o
cussed on previous pages. It will be seen when we describe the vege
tation on the various formations that the coarseness of the sam '
some cases seems to determine the composition of the plant acts
munities. This is easily understood when we consider that the siz
of the sand particles determines the water-holding capacity of the

; CHEMICAL COMPOSITION OF SAND.—The nutritive value of sand
different according to the chemical character of the sand grains.

1909] OLSSON-SEFFER—VEGETATION ON SEA SHORES 103

a tule, sand is very deficient in plant food, and this is especially the
case with the commonest form of sand, that which consists mainly
of quartz. The quartz grains are insoluble, or only to a very small
degree soluble. Only in the case of lime or organic matter in the
form of humus entering into the composition of the sand is there
plant food in sufficient quantities to allow the development of a more
luxuriant vegetation. Generally the chemical composition of coastal
sands is very uniform, and this may to some extent account for the
evident similarity in vegetation on these formations.

A number of analyses of sand have been made by the writer, and
some typical results will be here related.

* °
No. 1—MiIppir BEACH, ECKERO StorBy, ALAND ISLANDS, BALTIC

Per cent.
Insoluble matter.............. So.25 . Alumina....c. 4. cee ees
onthe at aga 3.44 Water and organic matter..... a9
ORE 1.06 Nitrogen. .../.0.. 2.545233 0.16
Ph o.18 Other constitutents.........--- 1.02
sora acid 4 (oluble).. uae sits Total. 6... bee 100.01
Peroxid 0 Se ee ee
ie. ae DUNE, ENGELHOLMSHAMN, SKANE, S. W. SWEDEN
In Per cent.
Pre eee... ... 84.38 Alumina........ 63.2 oa
2G ot eae 4-53 Water and organic matter ....- 4-93
ican: ates 104. . Nitrogen... :. 2.2.5 5055 ses 0.12
Phoapthoei ae ‘a Sas eee 0.21 Other constitutents.........--- 2.00
¢ acid (soluble eee
Peroxid of iron ee oe os Total: 7 FAS eee 99-99
No. 3—Uprer BEAcH, NorTH BEACH, NEAR PERTH, W. AUSTR.
Insolu Per cent
Sie Woe. 86.38 - Alumina... 3.0. es 0.93
x cs ata 3-61 Water and organic matter... --- 2.89
site Peet revs 1.59 Nitrogen... «ec: rk ee Gee 0.03
Phosphoric a, < cit 5s OvgO . Other constitutents eee a 3-33
Petoxid of ir . coluble.. ae a Total. eee 100.02
No. 4, > onset Sovnonr, QUEENSLAND
Tnsoluble matter Per cent.
a hClUC CU or px Alamins...0. i.6s56 oe: ‘
Me te 3-18 Water and organic matter . 2.15
ee 6.86. Nitrogen. ...6205 5-0 0.11
Phosphoric 4. i ais id .. ©.21 Other constitutents.......----> 0.44
Petoxid of iro (0 uble).. Se aes valet. 2 Psat ee Os 100.00

104 BOTANICAL GAZETTE [FEBRUARY

No. 5.—Dvunes, GOLDEN GATE ParK, SAN FRANCISCO, CAL.

Per cent.

Insoluble matter.............. 88.2 Alumina..2... 06.522 6 1.08
Renee Be i ee a 4.42 Water and organic matter...... 2.15
EROS OO e Re og Sarena Rd NNTOSEN. 65.45 2 ss ee 0.05
ee ee a OuI Other constitutents.......... 4 1.80
Phosphoric acid (soluble)... ... 0.08 TO). oo. os oe 100.01

oS 6 in 0.05

These analyses show what a small amount of plant food is available
in the dunes in comparison with that in ordinary agricultural soil,
where the insoluble substances do not comprise more than 70 pet
cent. of the total volume. And it must be remarked that the analyses
here given represent soil from places more or less covered with vegeta-
tion, where the organic constituents are better preserved from decom
position and from being washed out by water than on open sand.
They therefore show a higher percentage of humus and soluble
material than the barren quartz unprotected from the influence of
sun, air, and water. Where sand has recently been deposited after
having been exposed for some time to sea water it is naturally very
deficient in plant food, and it has therefore to be considerably changed
before it is able to sustain a vegetation covering.

The amount of lime contained in the dunes varies to a great extetl.
On tropical coasts it is generally very large, especially where the s
is formed by disintegration of coral rocks. On such shores carbonate
of lime is dissolved by the rain water and the sand is at a low depth
under the surface consolidated into limestone. A similar process of
calcification can be observed also on many coasts where the amount of
lime is quite small, as on some coasts of Europe. BANG (1) has ob-
served that the dune sand near the sea contains up to sixteen times
more lime than farther inland. This is a natural result of the wash
Ing-out process and decomposition, which takes place on the ope
sand, and is more effective farther inland, because the supply dimin-
ishes with the distance from shore.

On the upper beach and on the seaward slope of the littoral dunt
are frequently found fragments of shells that have been Cat
ashore by the waves. In places where the littoral dune is broke
shells are often accumulated in the depressions, while more landw

the lime in the animal remains is disintegrated by the carbon dioxid

1909] OLSSON-SEFFER—VEGETATION ON SEA SHORES 105

The greater or smaller amount of peroxid of iron in dune sand
determines to some extent its color. The usually colorless grains of
pure quartz are covered with a thin coat of ferric hydroxid, which gives
the sand its yellow color, and in some places almost a red tint.

THE SOLUBLE SALT CONTENT IN COASTAL SAND.—Of the soluble
salts that saturate the coastal sands sodium chlorid is the most impor-
tant. Its presence, as common salt, in all plants is well known, and its
influence on the littoral flora is very apparent. Whether sodium chlorid
isessential to plant life is still an open question, and the investigations
hitherto conducted in order to ascertain this fact seem to indicate that
such isnot the case. In experiments it is difficult to eliminate salt
entirely, but it has been conclusively shown that the smallest quan-
tities only, if any at all, are needed for the development of plants, even
for those which apparently prefer salty situations, when growing under
natural conditions.

That common salt is injurious to plants, when present in excessive
quantities, is certain. It is commonly believed that this unfavorable
influence of salt is due to the amount of magnesium chlorid it contains.
It 'smore likely, however, that all the chlorids are injurious, and exper-
(ments by the writer have supported this view, previously maintained
by several authors, :

It 1S generally stated by various writers that the formations on the
Nees contain a considerable amount of common salt. Thus
ELE G (14) says that on the sandy beach the salty ground water Is
: ae 08 slight depth under the surface. CONTEJEAN (3), in
hae oe conditions in southwestern France, considers that his
bck i si sea-shore vegetation, that is our middle and upper
feet ea eae in a saline soil. Mascter (9) found the salt con-
150" from eae the sea to be 0.351 per cent., while at a distance
bins, as ore he found 0.17 per cent. of sodium chlorid, and

= aan ©.041 per cent. :
authors whether "9 some doubt, however, in the minds of certain
ic i. * coastal dunes are impregnated with common salt
Various coasts 8 . is MASSART (to). The present writer has on
analysis hem € tests for salts in the sand by means of chemical
circumstances . of these observations shows that under ordinary
mes do not contain sodium chlorids in perceptible

106 BOTANICAL GAZETTE [FEBRUARY

quantities. When salt is found it has been deposited as spray from the
sea, but this is rapidly washed out by rain water, and when no precipi-
tation has fallen, the sodium chlorid does not come into contact with the
ground water but is detained on the surface by the upward movement of
the water. Because of this the roots of the plants are not exposed to
sodium chlorid. On the littoral dune the uppermost half an inch
layer of sand usually contains some salt, but deeper in the soil no salt is
found before we reach the sea-level. ‘The upper beach has very
similar conditions, as a rule, except at times when inundated by high
water. Even on the middle beach we cannot find that the sand would
be impregnated with salt. On the contrary, for quite a considerable
depth there is fresh water, which, on account of its being lighter than
the salt water, flows on top of the latter. This fresh water is a pati
of the continuous stream of rain water, which slowly works its way 10
the sea. The roots of the plants do not, as a rule, penetrate deepet
than to the bottom of this fresh-water layer, and it is therefore wrong
to assume that the plants are growing in salt water on the beach.
Even on the front beach, the layer in which the roots of the plants
are situated has more of a brackish character, because the water from
the beating waves runs off before it has time to sink through the layer
of fresh water, which flows on the surface of the salty ground watel.
On a superficial investigation of the beach it appears that the
ground is thoroughly soaked with salt water, but careful sampli
from various depths and subsequent analysis has made it apparent
the writer that this is not the case. It is a well-known fact, howeveh
that the ash of strand and marine plants contains a much larger P®
centage of sodium chlorid than that of inland plants. This is due, at
course, to the presence of a greater amount of salt on the sea shot
than inland. But when it comes to a comparison between the com
ditions on sea shores and salt-impregnated formations in the interior,
the amount of salt in the latter is much greater. This fact brings
forward the question whether all sea-shore plants are halophiles *
not. Kearney (8) has investigated this question and comes t© as
result that they are not. The present writer made numerous exper
ments in this direction and the results confirm those of Kear’
as the following discussion will show.
Ithas long ago been proved by experiments that most inland plants

1909] OLSSON-SEFFER—VEGETATION ON SEA SHORES 107

are injured by the presence in the soil of sodium chlorid in certain
quantities, which the strand plants are able to tolerate without evident
injury. There appears to be a certain maximum amount of salt for
every species, to which it is very accurately adapted, and this maximum
cannot be overstepped without fatal results to the plant. In some
cases investigated by the writer this maximum has been found to be:

Per cent. Per cent.
Argenlina anserima...............1.9 Glaux maritima...............--. 2.7
Aster Trspolum................. 2.6 Juncus Gerards,.. 2.5 0c ee oe
Airipex hastata maritima..........3.1 | Matricaria inodora maritima...... 2.3
Cakile maritima................. 2.9 Plantago maritima...........5.-- 2.8
Crambe maritima................ 2.5 Sonchus arvensis maritima.......- 2.6
Elymus arenarius................ 2.6 Triglochin maritimum .....- Te
Erythraea vulgaris............... 1.9

These experiments were conducted in the summer of 1894 with
plants from the Baltic coasts. Sand cultures saturated with normal
solution of sodium chlorid were used. In these cultures young seed-
lings as well as older plants were grown, and the results given above
teler to seedlings, about two weeks old at the time of transplanting.
= were grown in fresh water for five days, after which time the salt
See rere gradually applied. It was found that plants which had

\ growing on strands with low salinity were considerably more
oe an increase in the amount of salt than those which were
pit In from the open shores with higher salt content in the water.
ni well-developed plants adapted themselves more readily than
would tag to the gradual transfer to stronger salinity. At
limi of considerable interest to ascertain whether this specific
the eo rntration could be raised much higher by growing

S through a succession of seasons. The ability of the sea-
— to endure salt in the soil without injury and by adapting
of “ahi these conditions has, no doubt, been the ultimate cause
Petition fro he many cases confined to the strand, precluding com-

We i am poeeng this power of resistance.
not need sodj wed from experimental cultures that strand plants do
then stises aR chlorid in order to develop normally. The question
lower than ene the common salt, even when present in quantities
SCHIPER : maximum, exercises a poisonous influence or not.

» Who paid much attention to this matter, came to the con-

108 | BOTANICAL GAZETTE [FEBRUARY

clusion that the chlorids produce abnormal conditions in the plants
and disorders in the nutritive processes. In this regard most writers
agree, but in explaining the means by which the plant neutralizes
this injurious effect of common salt there is a wide divergence of
opinion. While ScHmmPeR maintains that the structural adaptations
of halophile plants are caused by the necessity of keeping the rela-
tive amount of sodium chlorid in the cell-sap below the specific danger
point, Drets (6) considers that this is effected by chemical decompo-
sition of the salt. This process is not known, but DreLs assumes that
in respiration the succulent halophytes differ from other plants in that
the oxidation does not proceed so far in halophytes, but stops at malic
acid or some isomer, with which the cell-sap becomes saturated, while
only small quantities of carbonic acid are evolved. The malic acid
then combines with the hydrochloric acid and is excreted by the roots.
BENECKE (2) has severely criticized these conclusions of D1zts.

In regions having a hot climate the evaporation of water is very
great on the coastal sands and the salinity is naturally higher. The
concentration of salts is also increased in countries where the rain falls
only during a rainy period, leaving a long time in which no leaching
of the salts takes place in the soil. In places with frequent rains the
salts are rapidly washed out and carried deeper into the ground, until
the lateral flow of water toward the sea is encountered.

The observations on salinity of strand sand made by the author
are all based on chemical analysis. The electrical method of deter
mining the salinity as employed by the United States Bureau of Soils
was not familiar to the author at that time, but careful observations
and determinations of the salinity with that method ought to reveal the
causes of distribution of certain plants on the strand. The writer
has found that the small embryonic dunes formed by certain strand
plants contain a greater amount of salts than those occupied by others.
Thus, for instance, the small, embryonic, Elymus arenarius dunes
always contain ©.005-0.009 per cent. more sodium chlorid than the
Ammophila arenaria dunes. Likewise the Mesembryanthemum
dunes on the California coast have a higher salinity than the Abron™
dunes, while the elevations formed by Abronia latifolia contain mor
salt than A. umbellata hummocks. These are the only exam es
which have been verified by analyses, but more extended investig®-

i909] OLSSON-SEFFER—VEGETATION ON SEA SHORES _t09

tions will, no doubt, give an explanation of certain hitherto unex-
plained features of the local distribution of strand plants.

We often find on sandy sea shores a number of immigrants from
inland formations, and this occasional occurrence of plants which
do not naturally belong to such habitat shows that it cannot be the
chemical composition of the salt water that keeps so many island
plants from the sea shore, but other adverse conditions, which allow
only the peculiar sand-strand. flora to develop. Even on the front
beach, where the salinity is greatest, we cannot attribute the scarce-
ness of the plants to the salt content, but to the easily movable sand
soil

As we have already mentioned, the lateral current of fresh water
flowing on the surface of the salty ground water near the sea has to be
taken into consideration when we discuss the salinity of the strand
soil. Our assumption that the conditions of the strand are not such
as to characterize this formation as halophytic is borne out by the
analyses made of the salinity of the soil at different depths. Many
true halophytes, of course, occur on the sea shore, but the strand
flora as such must rather be classified as a halophile flora, while the
ae halophytes are those plants which are confined to saline situa-
= in the interior, or where we know that the hydrodynamic condi-
tons do not change to any marked degree the salinity, as is the case on
wea shore. If this holds good, the halophytes occurring on the

d must be regarded as immigrants from dry saline habitats.
Several Salsolaceous plants, widely spread in the interior of
ian sometimes occur on the sea shores of that continent as
. peor specimens, but reach their best development in the dry
ae cay of the interior. ScHmMPER (13) maintains that the cliffs
marsh Sea shore have a much less halophile flora than the sandy or
; le This is evident to everyone who has studied the
many a but we shall find that the plants even on the cliffs exhibit
acters of the halophytes, and are sufficiently differentiated
as bist, aay of inland situations to warrant a classification
imprep he physical nature of the substratum prevents Its
the _ aor aie ee salt. We have here to account for
8© much th eles adaptations so characteristic for halophytes, not
Tough the influence of salt in the soil as through the salt

IIo BOTANICAL GAZETTE [FEBRUARY

contained in the spray, to which the plants are constantly exposed.
On sandy soils the protectional adaptations are caused more through
the physical conditions of the sand, than through the salt content of
the soil. The characteristic vegetation developed on all sand forma-
tions, inland as well as on the coast, is so much alike, that there is no
reason to assume that the sodium chlorid content of the sea shore,
which in fact is not very large, would be responsible for the aspect of
the vegetation on marine sand strands. On coastal marshes the con-
ditions are different, and this is also evident in the vegetation on such
formations, which in no way differs from that on saline marshes in the
interior, and always is composed of true halophytes.

A series of samples of the soil was taken with earth-auger on the
beach and dunes at Fremantle, Western Australia, at various depths,
and subsequently examined for soluble salts. The results appear in
Table XI.

TABLE XI
3
a 5
aeio we Se ee
= rae gs) 8 a. eam
ormation of %s g 5) gig - _
bale =| ‘ -) 7) £ et %
aS $ wi ee ge ae z
di] § (2138/2 [St] &
3 a 3) q o or 5
A 6 Gl¢ieja | & |e
Dec. 18, ’02| Lower beach, 2 | Medium | 20 | 15 | 1 9 | 9-005
upper limit nd
Middle beach 5 | Medium | 35 | 21 | 20 | 6 | 0.004| Sparse,
vegetation
Border of mid- | 7 | Coarse | 30 | 23 | 20] 6 | 0.004
up- sand
per
Upper beach to | Medium | 30 | 26 | 22 | 4 | 9-009
Dake tanta . sand
une inside 28 | Fine ier
littoral dune san zeit) et . /
Littoral dune | 15 | Fine 4o | 30 | 21 | 4'| 0.007
Dune marsh Finest 00
inside littoral i aa or coal
dune
Dec. 21, 02) Middle beach 8 | Medium | 25 | 8/14] 4 0.006
5 sand :
ame place 8 | Medium | 50| 8| 12| 3 | 0-004
sand
Upper beach 14 | Medium | 25 | 15 | 16 3 | 0.011 Spars ‘on
sand | 50 15 | 521-23 8
oo ei oath

1909] OLSSON-SEFFER—VEGETATION ON SEA SHORES III

DEVELOPMENT OF HUMUS.—There is no other soil which so little
favors development of humus as the loose shifting sand. The organic
substances that happen to be deposited on the sand are very rapidly
decomposed by the admission of air, and the physical structure of the
sand allows-rain water to percolate and thus to carry the fine humus
particles deep into the soil and out of reach of the roots. The earth-
worms, which are the most active agents in mold formation in forests,
as DARWIN and MUELLER have shown, are entirely absent in sand
and the mycorhizal fungi seem not to thrive on the beach, where they
are likely to be exposed to occasional contact with sodium chlorid.

When a shrubby vegetation has got a foothold on the sand, the
humus is developed to the best advantage. In the shade of the
bushes remains of plants do not decompose so easily as on the open
ground, they are more sheltered from the rain, and an accumulation
of humus can take place, so that grasses and herbs are able to get a
footing. When this has happened the sand is usually made perma-
nently stable. The few animal remains that are thrown up on the
front or middle beach enrich the soil on these formations only tempo-

tatily,and do not play any important réle in the f tion of the humus
on the sand.

Topographic factors
foes ty as a factor influencing the development of vegeta-
in ety often overlooked by writers on plant geography. Its
Tea, however, is so considerable that it cannot be omitted in a
ie. of the agents which exert their influence on plant life.
bie dn’ €getation topogr aphy acts principally indirectly, by determin-
the tem i extent the moisture content of the soil, by influencing
We mar i oo exposure to winds, and also the light relations.
topograph €re to mention briefly only the principal features of
hon. y as far as they influence the conditions on coastal sand
Ons,
apnea, NDINGS.—From our previous discussion (11) of the devel-
Toundin € various sand formations it is apparent that the sur-
nee =e : the eetest consequence to the evolution of dunes.
have been : coasts investigated the topographical conditions
Sanh, a character as to prevent any greater development of
as the case, for instance, on the southern shore of the

I12 BOTANICAL GAZETTE [FEBRUARY

Gulf of Finland, where almost the whole shore line consists of a
steep wall of rock, leaving only a narrow strip of beach along the
water edge. In places where this rocky barrier was broken and the
winds are allowed free play over a wider stretch of land, dunes ap-
peared at once. The vegetation on the beach of the former type
presents a somewhat different aspect from that on open shores with
the background of a dune-complex. .The best evidence of the influ-
ence of surroundings on the composition of the vegetation can be seen
if we compare that on a sand field and on a dune-complex with its
diversified topography. Also on the slopes of an unbroken dune, the
vegetation is usually quite different from that on a train of dunes fre-
* quently cut through by furrows and valleys.

On beaches a similar difference can be noticed, and the cause
underlying this effect can only be attributed to the topography. Where
_ we have a long continuous beach the plants associate according t0
rules different from those which have determined the composition of
the communities on cuspate forelands. This was especially evident
on the shore stretches of sandy beaches that are so common on the
shores of the islands in the Baltic.

ELEVATION.—This factor is of minor importance in regard to the
sand-strand vegetation. The sand formations do not rise to aly
great height, but it seems that certain plants choose their place of
growth with reference to altitude, even on these formations. Without
taking into account the fact that humus naturally accumulates mot
rapidly in the depressions, we find that some plants prefer the foo! of
a dune, while others are found only on the middle of the front slope,
and others again do not thrive except on the top of the dune, where
they are constantly being covered with drifting sand.

On the beach a corresponding selection of habitat takes place.
Some plants never occur on a low beach although the conditions
otherwise seem to be favorable, but only a short distance away, where
the beach rises more abruptly, they appear again. We have ee
sumably two different causes for this. While on the dunes the selee
tion of a place of growth is determined apparently by the plants
Sreater or less power of resistance against the drifting sand, :
sft the dominant cause must be the sensitiveness of the plant Y
inundations of salt water. :

1909] OLSSON-SEFFER—VEGETATION ON SEA SHORES 113

Depressions between the dunes offer to many plants a refuge from
the sand-laden winds, and the richer soil in the troughs induces other
plants to settle.

GRADE OF SLOPE.—We have elsewhere (11) referred to this
factor as being of great moment in the growth of dunes and in the
development of sand formations generally. Its influence on the
moisture conditions is of no less importance. The higher up on the
slope the drier is the soil, the greater the evaporation, and the more
intense the influence of the wind.

The exposure of the slope is another matter of the greatest con-
sequence to the vegetation on the sand formations. The various
degrees of slope ought always to be considered when a description of
a habitat is given in order to arrive at a correct understanding of the
conditions that have determined the composition of the plant com-
munity. Southern slopes in the northern hemisphere and northerly
slopes in the southern are drier than those facing other directions, and
the vegetation has a corresponding aspect. On sea coasts the expo-
“sei ” the prevailing winds has to be noticed.

Light relations are further to a certain degree determined by the
Section of the slope, and this is of special importance in northern
latitudes, where light even during the period of growth is not too
abundant.

Historical factors

Paes ‘ his heading we include all those factors whose influence
. Plant life is determined by time. This must not be understood
3 time was not involved in the action of other factors, but that the
fg Miintluence and the time required for attaining any results is of

@ long duration, that it cannot be ascertained within a few
iss - te plants. Physiographical changes of land and sea,
considered in i spac) - within long geological periods, have to -
factors to be . ee One of the most important baie
sand strands { ag Been. mn explaining the present conditions 6
Shore by wa is the oscillation of the coast line. The erosion of the
equally im ves and the deposition of sand or other material are
rapidly ..... Sand deposits are in many places formed >
influence of a effect can be noticed within a very short time. The

mals, principally through grazing, and the interven-

114 BOTANICAL GAZETTE [FEBRUARY

tion of man belong strictly to the group of biotic factors, but have here
been considered in connection with the historical factors as a matter
of convenience.

OSCILLATION OF THE COAST LINE.—Many coasts are slowly rising,
while in other instances the coasts are sinking. We have excellent
examples of both kinds of movements on the Baltic. While the
whole coast of Sweden north of Stockholm, and the coasts of Finland
are in a state of elevation, the southern shore of the Baltic is in a cor-
responding state of depression. Besides having a great influence on
the development of dunes, this oscillation of the coast line has hada
marked bearing on the evolution of the flora on the coastal sands.

On the shores mentioned which are rising, one may see in some
instances how long stretches of land are slowly raised above water
and in a few years carry a cover of vegetation that gives an instructive
demonstration of the successive stages of development of the plant
associations. Again, on the sinking shores of the southern Baltic
may be frequently found examples of plant communities being
destroyed in the course of a few years through the submersion of the
shore.

On the coasts bordering upon the oceans oscillations also take
place, but they are usually neither so regular nor so rapid as these
changes on the Baltic.

In postglacial times considerable changes of the coast line of the
Scandinavian countries have taken place, and as we are able to follow
these changes with the assistance of fossil remains of plants, found in
old sea beaches now raised high above the present level of the sea, We
can to some extent interpret the various stages of development which
have been passed before the flora arrived at its present state.
question will be discussed in another paper.

EOLIAN DEPOsITs.—The influence of the wind on formation of
plant communities on the coastal sands is shown not only in the
peculiar arrangement of the plants in patches, but also in the aspect
of the aggregations of plants, which (especially in the case of te
and shrubs) give evidence of being continuously attacked by the
strong winds laden with spray or sand. Shrub associations 0M open
strands are usually lower toward the shore, gradually increasing
height inland under the shelter of the more exposed specimens.

1909] OLSSON-SEFFER—VEGETATION ON SEA SHORES

II5
Most of the sand-strand plants are not able to withstand partial
burying by the shifting sand; consequently it very often happens that

1G. 1.—Cupressus macrocar pa on Cypress Point, Monterey, Cal., showing influ-
ence of wind. (Photograph by author.)

Fic >

“=

—Rejuvenated Salix dune at San Francisco, Cal. (Photograph by author.)
who ‘7s
le Communities are su

Y plant: ddenly destroyed and their place taken up
mei, se to endure more or less complete covering by sand.
“Sit may be observed that trees, which are being buried by

116 BOTANICAL GAZETTE [FEBRUARY

an encroaching dune, are bent leeward (fig. 1). The cause of this is
to be found in the continuous pressure, or in sudden gusts of wind
which bend the trees while the onrushing sand prevents return to the
original position. As a rule the sand on the lee side of a dune is
moister, and the slope is consequently steeper. Often slides of sand
take place, and they also bend or even break the trunks of the trees.
Dunes which have been made stable by a cover of plants are
sometimes again broken up by the wind (jig. 2). Such formations
are often met with on the Baltic coasts. On these broken-up dunes
the usual series of development of vegetation begins anew and thus

San

Fic. 3.—Embryonic dunes inland from littoral dune, south of Cliff House,
Francisco, Cal. (Photograph by author.)

they have a peculiar character, remnants of the old communities
being mixed with the new immigrants. A new life-history of the
plant community is started, and during the course of development "
may take a direction entirely different from the former series.
Vegetation covering on the ground will greatly slacken the speed
of the air current which comes into immediate contact with the
ground, and if bushes or other obstructions are in the path of the
wind, dunes are often formed behind them (fig. 3). The plants that
= first struck by the whole force of the wind are md y
injured, not only by its mechanical action, but to a greater extent

1909] OLSSON-SEFFER—VEGETATION ON SEA SHORES Ii7

by the sand carried by the wind. While the wind in itself dries
the plant, the sand particles, which often have a high temperature,
are still more apt to increase the evaporation, and thus to hinder the
development of the plant (fig. 4). Further, stems of plants growing
on exposed places on coastal sands are often eroded by the sand, and
even the green leaves are sometimes cut into shreds during severe
storms by the sharp angular sand grains.

Fic. 4.— :
hihis, 6 Effect of wind on Leptospermum scoparium Forst. on dunes at New
anterbury, New Zealand. (Photograph by Dr. L. CocKAYNE.)
OTHER sEpryE
mout S, and
on the sand

‘DIMENTS.—As we have pointed out in another place
any likely to accumulate in the neighborhood of river.
in such places heavy floods often carry down and deposit
ee ble quantities of mud, which then enrich the
exclude a the appearance of a quite new flora, that soon will
On alm ee eand plants.

ashore by ae “oan coasts quantities of seaweeds are thrown
Posed and th lee) but in warm climates they are so rapidly decom-

| mains washed away, that no accumulations can be

118 BOTANICAL GAZETTE [FEBRUARY

effected. In more temperate regions, however, these seaweeds lie
in banks on the beach for some time, and add to the fertility of the

Fic. 5.—Bank of Macrocystis on the beach at New Brighton, Canterbury, ¢s
coast of South Island of New Zealand. Height of scale 41cm. (Photo tograph .
Dr. L. Cockayne.)

Fic. 6.—Kelp banks on West Australian coast. (Photograph by author-)

soil, Figs. 5 and 6 hase such kelp banks from the coasts of New
Zealand and West Australia.

Along the shores of the Baltic a considerable amount of seaweeds

1909] OLSSON-SEFFER—VEGETATION ON SEA SHORES IIg

principally Fucus vesiculosus, is deposited on the beach at high-water
mark, On open shores these banks mark the limit of vegetation
toward the sea, and are characterized by a vegetation quite different
from that on the rest of the beach.

Man’s INTERVENTION.—The principal influence of man on coastal
sand floras is a result of his endeavors to arrest the drifting of sand.
This is mainly done by planting so-called sand-binding plants, or by
covering the loose sand with refuse or other material. Either action
brings about a considerable change in the natural development of the
sand vegetation.

Near cities the sand dunes are sometimes used for supplying sand
for industrial purposes, and in such cases the removal of the sand
will naturally change the conditions for the original vegetation. Fires
are sometimes started through the carelessness of man, but as the
oe vegetation is very seldom dense, the influence of fires on sand
formations on the coast is not great, except in artificial plantations.

_ ~tazing animals do more injury to the sand vegetation by tramp

= and uprooting the plants than by actual feeding on them, and in

43 oo of many cities, where sandy beaches and dunes
» tuman agency is equally detrimental to the plant covering.

Summary

Summing up the physical conditions prevailing on the various sand
always — marine coasts we would say that the submerged beach is
)» Sovered with water, and therefore is the most salty of all the
and the : She soil is loose, the temperature that of the sea water,
Special ; ~ 48 continuously beating. The vegetation is therefore
ao to these conditions, and in most cases no plants
The fr ete a a footing on this formation.
mati Shs periodically washed by the waves, presenting
oo and aquatic conditions. It is almost con-
Under the surg gas aad and has a salty ground not very deep
Submerged a The soil ay Wary loose, still more so than on the
Constantly any Strong insolation, rapid evaporation, and a
Of these i temperature are characteristic. It is on account
ion of a conditions usually devoid of vegetation, with the
“W unicellular algae, often Cyanophyceae, but where

120 BOTANICAL GAZETTE [FEBRUARY

the formation has not been inundated for a few days only the spores
of the algae retain their vitality. On the Baltic coasts where the
salinity of the sea water is low, conditions approach more or less those
on fresh-water shores, and many green algae occur in the sand, The
width of the barren front strand varies not only with the slope, but
also with the force of the surf.

When a higher vegetation occurs on this formation it is open and

Cal. 02

Fic. 7.—Middle and upper beach south of Cliff House, San Francisco, goat

upper beach small dunes made by Cakile americana. In background littor@
with Ammophila. (Photograph by author.)

very poor. The loose soil, the wind, and the impact of the surf pa
a limit on the types that are able to develop. These are a
annuals or perennials with long creeping rhizomes and the flora *
always poor both in species and individuals.

The middle beach is characterized by its light color, its abundar

Moisture, its low salinity, its loose soil, and its comparatively low
temperature. Occasional inundations, spray, and wind are Be
nn

direct causes of the scattered vegetation, consisting mainly of @

1909] OLSSON-SEFFER—VEGETATION ON SEA SHORES I2I

or a few perennials. These plants also are more or less dwarfed,
because of the wind and the cold substratum. The tension line
between this and the following formation is very marked. Some-

left, se 8.—Ammophila dunes at San Francisco, Cal. Embryonic Salix dune to the
Nn winter, (Photograph by author.)

ere!
ne . autho ems littoral dune, north of Fremantle, Western Australia. (Photo-

122 BOTANICAL GAZETTE [FEBRUARY

times a transition belt can be distinguished, marked by a darker strip
of humus-mixed sand, and covered with a denser vegetation (fig. 7).

The upper beach has a higher percentage of humus, abundant
moisture, and a higher temperature than the middle beach. The
illumination is much greater, especially during certain times of the
day, when the formation is not shaded by the littoral dune. Evapora-
tion and radiation from the soil, however, are less intense, because the

Fic. 10.—Soldanella community on dune slope at New Brighton, —
New Zealand. On summit, Scirpus Jrondosus. (Photograph by Dr. L. CocKaAYN

ground is well covered with plants. The amount of spray is less the
farther w

very long intervals. The distance to the salty ground water 1s ©”
siderably greater than on the lower formations, and very few

which often is covered with small embryonic dunes (fig. 8). i .
wind favors transportation of seeds and shoots from the mid

Fic. r1.—Sand dunes at Studeli Mile in north Jutland, Denmark, covered by Elymus arenarius, Ammophila arenaria, Carex
arenaria, etc, In the foreground especially Salix repens. (Photograph by Dr. F. BORGESEN.)

[6061

1 OA A—AAAATS-NOSSTO

SHYOHS VAS NO NOILVI

124 BOTANICAL GAZETTE [FEBRUARY

beach, and in many cases the occurrence on the upper beach of plants
belonging to the middle beach can only be explained by their having
been blown landward from the original position.

The vegetation on the upper beach consists principally of perennial
herbs, shrubs, or low trees. The tension line toward the littoral
dune is not so marked as in the direction of the middle beach, but
where no dune formations are developed the upper beach usually
borders upon a forest. In the latter case it often happens that inland
plants have wandered out to the sea-shore formations, while it never
happens that sea-shore plants have been able to establish themselves
inland on the meadow or forest that usually follows the strand forma-
tions.

The littoral dune is much exposed to the wind, its moisture content
is low, constant oxidation of organic water goes on, and the tempera
ture is lower than on the upper beach, because of more intense
radiation. The soil is very loose, shifting, and sterile (fig. 9). The
vegetation shows the results of these conditions very plainly. It is
prostrate and dwarfed in habit, scattered and poor in variety of
orms.

The active dune (the white dune of Warmtnc) has all the character
istics of the littoral dune in excess, and its vegetation is generally i
more monotonous. Some difference can be observed in regard to the
plants on the slope and summit of the dunes. It is usually richer
species on the latter part of the formation.

The stationary dune (the gray dune of WARMING) is formed at #
greater distance from the sea, where the sand has to some extent col
solidated, and a heather vegetation has been established. Form
tion of humus goes on, the plants grow closer and closer, mosses
lichens occupy the ground between the higher plants, and finally
the soil is completely covered with a carpet of vegetation. This
heather association is the final stage in the series of sand-plant com
munities beginning on the small embryonic dunes to leeward from the
littoral dune (fig. rz).

The sandy field is richer in humus than any of the former sand for-
mations mentioned. It has a comparatively level surface, is better
to retain moisture, and has a higher temperature. The sand grain
are of such uniform size as to prevent ripple or dune formation, and

Fic. 12.—Sand field at Lodskovvad Mile in som ee Denmark. The place was formerly a shallow lake, which had
been filled in with sand three years previous to the time (August 18, 1898) when the picture was taken. e vegetation consists of
Salix repens, Juncus balticus, Agrostis alba maritima, Hochars Palas and Phragmites communis. In the background sand dunes
with Elymus and Ammophila,. (Photograph by Dr. F. BORcEs

[6061

SHAYOHS VAS NO NOILVLYDF A—aXddAATS-NOSSTO

Cer

126 BOTANICAL GAZETTE [FEBRUARY

the vegetation covering is consequently developed in quite a different
way from that on dunes (fig. 12).
Mexico, D. F.

LITERATURE CITED

1. Banc, J. P. F., Om de nord- og vestjydske Klitters Beplantning. Tidssk.
for Shovbeus, Kjobenhavn 12:—. 1880.

. Benecke, W., Ueber die Diels’sche Lehre von der Entchlorung der Halo-
phyten. ih. Wiss. Bot. 32:179-196. 1901.

3. ConTEJEAN, C., Géographie botanique. Influence du terrain sur la végéta-

tion. Paris. 1881.

4. Cooke, W. E., The climate of Western Australia from meteorological obser-
vations made derine the years 1876-1899. Perth. 1901.

. Courin, H., Sur la toxicité des composés du sodium, du potassium et de
Eatnenhic 4 légard des végétaux supérieurs. Rev. Gén. Bot. 12:17

1g00.

. Drets, L., Stoffwechsel und Struktur der Halophyten. Jahrb. Wiss. Bot.

32: — 1898.

7. Hann, J., Handbuch der Klimatologie. Stuttgart. 1897.

8. Keavnzy, 2. H., Are ae of sea-beaches and dunes true halophyte?
Bot. GAZETTE a 424-436. 1904.

9. Mascrer, A., Etudes sur la aul botanique du nord de la France.
Journ. de Bot. 2:177. 1888.

10. Massart, J., La biologie de la végétation sur le littoral Belge. Mém. Soc
Roy. Bot. Belgique 32:7-43. 1893.

II. OLSSON-SEFFER, PEHR, Relates a — to topography of coastal drat
sands. Journ. Geol. 16:549-564. 1

12. ———, Hydrodynamic factors ane plant life on sandy sea i“
New Phytologist (in press).

13. Scuimper, A. F. W., Pflanzengeographie auf physiologischer Grundlage
Jena. 1898.

14. Warne, E., Plantesamfund, Grundtraek af den oekologiske Plantes
grafi. Kjébenhavn. 1895.

a=,)

SOME ASPECTS OF AMITOSIS IN SYNCHYTRIUM:
ROBERT F. GRIGGS
(WITH PLATES III AND Iv)

Previous papers on the cytology of Synchytrium have announced
very striking peculiarities in the nuclear behavior of this interesting
fungus. The idiosyncrasies, only a portion of which have yet been
described, are so abundant at a certain period of the life-cycle of the
plant that it is very difficult to consider any one set of phenomena
without quickly becoming involved in all the rest, either because of
the occurrence of different types of structures in the same coenocytic
cyst, or because of transitional forms apparently connecting diverse
structures. While no final interpretation of any one series of nuclear
transformations can be made until it has been brought into relation
with the whole life-history, it-is apparent that it is out of the question
‘0 work out all of the peculiarities at once. The present paper is an
attempt to isolate and describe one of the most conspicuous groups of
=a Phenomena. Further correlation of this with other mani-
‘stations of nuclear activity will be undertaken in later papers.

= the Preparation of a former paper on Synchytrium (GRIGGS
a — is under very great obligations to his friend, Professor
i, for the information which aroused his interest in the
ty tic os ce criticism of the results. This obligation is increased
rs at Dr. StEvENs also supplied the material from which
Species, § made. The present paper deals entirely with one
taken hc: y ee decipiens Farlow. The drawings have all been
toxylin, a — stained with Heidenhain’s iron alum hema-

Tn the Sea i. has also been used. ae
the variation hy this plant there is no more striking feature than

uently found haw of the nuclei. In the same cyst nuclei are

iameter ss a | Tanging all the way from 8 or ro # down to I # in
the small a ‘a first reported by STEVENS (12, fig. 2). Very often
“el are bunched together, either in a close morula-like

ss i . .
No. XL, butions from the Botanical Laboratory of the Ohio State University,
127]

[Botanical Gazette, vol. 47

128 BOTANICAL GAZETTE [FEBRUARY

cluster (fig. 33) or in a looser group (fig. 18). The origin and fate of
these small nuclei is the subject of the present paper. Although such
variations in the size of the nuclei are sometimes found in almost any
stage of the period of nuclear division, they are most conspicuous
immediately after the division of the primary nucleus and continue
prominent until there are 200-300 nuclei in the cyst. It is at this
same stage that the other peculiarities in the cytology are most prv-
nounced. While this period of irregularities is not sharply marked
off from the succeeding phases of the life-history, yet as the nude
become more and more numerous there seems to be a tendency fot
them to settle down, so to speak, and to conform more nearly to the
usual habits of dividing nuclei in growing tissue.

The isolation of these groups suggests that their constituent nude
have a common origin. Because of the absence of any pairing, and
because of their great differences in size, one is inclined to suspect
that they have been derived by some process other than mitosis. Since
mitosis in this plant is always simultaneous, involving all the nuclei in
a cyst, the differences could not be due to the failure of some nuclei 0
divide, while their neighbors became smaller and smaller by repeated
division. They might of course be due to some process of mitosis 18
which the products were unequal, as in the reduction division of
animal egg. But all the mitoses observed gave rise to equal daughter
nuclei. Further, mitoses in cysts of this age are uncommon. This
led STEVENS (12) to suggest the possibility of an amitotic origin for the
nuclei of this stage.

: There are several processes of direct nuclear division in Synchyt
num. Two of these are quite different from the commonly observel
division by an amoeboid constriction of the parent nucleus. While
they may be considered under the general term amitosis, which =
come to include several forms of non-mitotic division, they require e
tinctive terms for their designation. Indeed, there is considerable
need for a classification of the different forms of direct divis®™
especially in view of the increased importance amitosis is likely
assume in future cytological discussion. The first process, W™
ee of a budding-out of a small nucleus from a larger; may be
designated nuclear gemmation. The second differs from ordinaty
amitosis in that the nucleus loses its membrane and vacuole of karyy

1909] GRIGGS—AMITOSIS IN SYNCHYTRIUM 129

lymph before the division, which is a multiple fragmentation. This
form of division I shall term heteroschizis (érepos, different, and
cy (Sev, to split).

NUCLEAR GEMMATION

In nuclear gemmation, as is usual in amitosis, the division of the
chromatin is not nearly so frequently observed as the separation of
the two nuclei. In the resting nuclei of Synchytrium the whole of the
chromatin content is usually concentrated in a single globular karyo-
some (nucleolus). At the beginning of nuclear gemmation the margin
of this karyosome becomes crenate, and rounded lobes develop, which
separate from it and become smaller independent karyosomes (figs.
4-8). Sometimes only one daughter karyosome migrates from the
parent at a time (jig. 6); sometimes the parent undergoes a process
of bipartition resulting in equal daughter karyosomes (jig. 7); OF
sometimes several form at once, in which case the whole karyosome
breaks up (jig. 4). Fig. 8 shows a very large nucleus where the
daughter karyosomes were unusually numerous. They were not free
in the nucleus, as appears from the drawing, but all of them were
lying against the nuclear membrane, only one hemisphere of which is
Tepresented.

After the separation of the small karyosomes is complete, they mi-
si oe the nuclear membrane. This process is probably rather
ee ae all stages are easy to observe: figs. 8, 19 show them
= met against the membrane; in figs. 9, 20 they are pressed
“bainst it; in figs. 8, 21 they have begun to pass through; ig. 27

* one lying almost exactly half-way through the membrane;
Ms . show karyosomes which have passed through, but still lie
gta the membrane.
cavity “a as the passage is completed, a vacuole, similar to the
is is ok nucleus, appears around the migrating karyosome.
of the a surrounded by a membrane, extending out from the
. eae ent nucleus into the cytoplasm next the vacuole (fig. 10).
Nucleus ig of ome be observed satisfactorily only when the daughter
branes of the considerable size, because of the delicacy of the mem-
nucleus moy smaller nuclei. When the membrane is complete the new
€s away from the parent and becomes an independent

figs.

wall

130 BOTANICAL GAZETTE [FEBRUARY

small nucleus free in the cytoplasm. The stages in this process are
also easy to follow: jigs. 3, 9, rz show cases where the karyosome is
still in contact with the membrane of the parent; jigs. 6, 12, 13, 18
cases where the karyosome has separated from the parent, but the
membranes remain in contact; jigs. 3, 5, 9, 14 cases where the two
nuclei have separated, but still lie close together.

Division by nuclear gemmation occurs also in the spirem stage
(figs. 15-17). In this case the division of the chromatin takes place
at the time of spirem formation and cannot be definitely connected
with nuclear division, but the manner of the separation of the daughter
nuclei is the same as that already described. Figs. 1, 2 show groups
of small nuclei from a cyst where all the large nuclei (fig. 16) are in
spirem. |

In cysts where the nuclei are numerous and evenly scattered
through the cytoplasm it can be seen that the peripheral nuclei
divide much earlier than the central ones. Groups of small nuclei
are always found at the periphery before the large nuclei in the center
are much divided. Thus a lateral section of a cyst (fig. 1) shows only
uniform groups of small nuclei, while the central sections show numer
ous large nuclei, of which jig. 16 is an example.

Nuclear gemmation may take place at very different rates in dif
ferent cysts. In the cyst from which jigs. rz, 12, 14 were taken, the
few small nuclei present are scattered singly through the cytoplasm.
In this case the appearances indicate a slow and orderly formation of
small nuclei. In other cysts the chromatin seems to be extrul
with almost explosive violence (figs. 24-26, cf. also GLASER 6). In
these cases a large proportion of the migrating chromatin never forms :
nuclei but degenerates in the cytoplasm. Some members of almost
every large group are imperfect and disintegrate, forming in their last
stages deeply staining spots in the cytoplasm. Such disintegration §
seldom seen in cysts where the small nuclei give evidence of mor
gradual formation. It is more pronounced in younger cysts where
there are only a few parent nuclei, than in later stages where they at
numerous,

The deeply staining granules on the nuclear membrane vary ae
karyosomes half the size of the mother karyosome to microsom” —
similar to those usually found in the nuclear membrane in both

1909] GRIGGS—AMITOSIS IN SYNCHYTRIUM 131

animal and plant cells (jigs. 7, 18, 21, 22). No optical distinction can
be drawn between these extremes. The very smallest granules, how-
ever, do not form small nuclei but may function in metabolism.
In mitosis and in the degeneration of the large nuclei (jig. 22) they are
cast aside with the old nuclear membrane and lost in the cytoplasm.
But no distinction can be drawn between these granules and those
which form small nuclei, for some of the latter are excessively minute.
Besides these, there are yet other granules on the nuclear membrane
from which conspicuous radiations proceed into the cytoplasm as
from centrosomes (fig. 39). The discussion of these bodies involves
other questions than those considered’ in the present paper and can-
not be undertaken here. Another complicating factor is the frequent
presence of asters near nuclei which are giving off gemmae. I have
avoided using such cases for the figures of the present paper, but in
many instances nuclei adjoining those drawn had conspicuous asters,
and it would be possible to duplicate most of the drawings herewith
given from nuclei showing asters. But though the centrosome prob-
lem, one aspect of which was touched in a former paper (GRIGGS 7),
s Very puzzling and far from solution, my belief is that it is inde-
pendent of the phenomena discussed in the present paper.

HETEROSCHIZIS

oe en nd process of amitosis is a multiple division or fragmenta-
nuclear . nucleus, which occurs for the most part at later stages than .
and ap but is sometimes found in young cysts (7g. 33)
area even in segmented cysts (fig. 34). Nuclei derived by
gemmati are at once distinguished from those due to nuclear
like clus on, because they form not a loose group but a close morula-
Matin ae sured by Stevens (12, fig. 3/). As in nuclear gem-
be ni = stages in their formation are easy to observe and may
at a eng a single cyst. But while the new nuclei are formed one
a that process, here they originate simultaneously by the

_ 40n of the mother nucleus. The first indication of division
Miclear : . loss of the nuclear membrane and the vacuole of
the UN allaas the karyosome lying naked in the cytoplasm, like
some then mes in the metaphase of mitosis (jig. 27). The karyo-
” *pparently enlarges to nearly double its former size (/ig-

132 BOTANICAL GAZETTE [FEBRUARY

28). This statement is based on the fact that the naked karyosomes
are, in the cases observed, larger than those of any other nuclei in the
same cysts, and that the resultant clusters of small nuclei are greater
in mass than any single nucleus in the cyst at this stage. The varia-
tion in the size of the nuclei, however, is so great in other stages that
it is not impossible that these may have been larger nuclei in the
beginning. Lobes (jig. 29) now appear on the margin of the karyo-
some, each of which rounds off and becomes the karyosome of a small
nucleus. When these karyosomes have separated, vacuoles of
nuclear sap appear around them; surrounding membranes are next
formed in the meshes of the cytoreticulum bounding the cavities, thus
completing the process. The membranes, however, do not appear
simultaneously around all the nuclei of a cluster. There is usually
sufficient difference to allow some observation of the process of mem-
brane formation. The vacuoles which become the nuclear cavities
are at first indistinguishable from those between the meshes of the
cytoreticulum which are filled with cell sap, but they are gradually
surrounded by membranes which are apparently precipitated from
the cytoplasm next the cavity. Neither in heteroschizis nor in nucleat
gemmation is there evidence of any connection of the centrosomes
with membrane formation such as occurs in the reconstruction of
the nucleus after mitosis (Kusano 8, GrRicGs 7).

Besides the two sorts of amitosis just described, a third method has
been observed a few times. In this process, which has been seen only
when the nuclei were in spirem, the nucleus becomes strongly lobed;
each lobe contains a portion of the original unchanged spirem; the
lobes become more pronounced and are cut apart by continued com
striction. There may be only two lobes, as in ordinary amitosis,
there may be several, as in heteroschizis. Although even a si
nucleus of this kind (fig. 40) would seem to indicate the general nature
of the process, there is much concerning it which is doubtful, and its
occurrence is rare in my slides. I refrain, therefore, from more that
mention of the matter at this time.

LATER HISTORY OF THE SMALL NUCLEI >

If amitosis leads to degeneration and death, as has been held
almost universally until recently, we should expect to find a large

190] GRIGGS—AMITOSIS IN SYNCHYTRIUM 133

percentage of degenerating small nuclei in every cyst where they
occur, either during the period of their formation or later. After
nuclear gemmation, however, degeneration of chromatin is relatively
small in amount and is almost altogether confined to masses which
never organize nuclei. It occurs not in the later portion of the period
of gemmation but only during the early portion, when there are few
latge nuclei in the cysts. In clusters due to heteroschizis, degenera-
tion may also occur at later stages, but is infrequent at any time. If,
on the other hand, these amitoses are due to pathological conditions
allecting the whole parasite, we should expect to find a large number
of dying cysts. Fully three-fourths of all the few-nucleate cysts give
evidence of amitosis. Of the remainder only a small number show
mitoses at this stage. This hypothesis would therefore require that
three-fourths of the cysts should degenerate sooner or later. But no
such thing occurs, Degenerating cysts are seldom found, and the
degeneration gives no indication of being connected with earlier
_ amitoses,

The clusters of small nuclei arising from heteroschizis tend to
remain close together, and when mitosis is resumed they may form a
cluster of small spindles. Fig. 35 shows such a cluster between pro-
phase and Metaphase, in which the remains of the nuclear membranes
ee sult evident. Below them is the solitary spindle of a large nucleus,
of which there are 40-50 in the cyst. Fig. 36 shows three objects
of nat cyst assembled in one drawing. At ais a similar cluster
ia ora at b is one of the solitary spindles of the larger
oi rs s which in this cyst are in a later phase than the clusters;
z eeply staining mass which has the appearance of a cluster
plisiee te Figs. 37, 38 show similar clusters in ana-
sive S though the spindle fibers in fig. 38 are distorted so as to

mewhat the appearance of a pathological multipolar spindle,

Som, ;
Pai the spindles are perfectly normal. The spindles of fig. 37
crample closely ‘the solitary spindles of the cyst and are typical

‘ of the peculiar anaphases of this genus. |
quickly eg - small nuclei arising by nuclear gemmation scatter
hich ea there is no means of connecting them with the mitoses
‘ytoplasm aa They have the usual relations, however, to the
appear normal in all microscopic characters. When

134 BOTANICAL GAZETTE [FEBRUARY

not too small they bud off other small nuclei in the same manner
(fig. 18). This process usually continues till all the nuclei in the cyst
are approximately equal in size (figs. z, 2). Sometimes all the
daughter nuclei given off are so much smaller than the parent that
the mother karyosome is never divided up among the daughters, but
remains behind full size, after giving up its chromatin, like the
nucleolus in the prophases of mitosis. In this case the large nude
degenerate and leave the small ones as the functional nuclei of the
cyst. Fig. 20 shows the first indication of this in the vacuolate karyo-
some of the parent nucleus. In fig. 27 all the chromatin has migrated
from the old karyosome but some of the small karyosomes stil
remain inside the nuclear membrane. The larger of these are about
the same size as the numerous small nuclei of the cyst. Fig. 22
shows another large nucleus from the same cyst, which is entirely
bereft of chromatin. Fig. 23 is the last stage of the process; here
the old nuclear membrane has disappeared and the faintly staining
old karyosome (nucleolus) lies naked in the cytoplasm. Beside it
is shown one of the functional nuclei. We are therefore led to the
conclusion that the nuclei derived by these processes of amitosis ae
normal, and that they with their descendants become the functional
nuclei of later stages, capable of perpetuating the species.

GENERAL CONSIDERATIONS 3

Although the processes by which these nuclei are derived at
novel, the formation of normal tissue by amitosis is by no meals
without parallel. C. M. Camp (1-5) has recently shown that
amitosis is a frequent occurrence in regenerating organs, embry%
and in some adult animals. He records instances from most of the
Sreat animal phyla, including coelenterates, flat worms, trematodes
cestodes, insects, amphioxus, fishes, amphibia, and birds. In thest
canes contrary to what would be expected, there seems to be
especial distinction between the soma and the germ plasm as to the
origin of the nuclei. In Moniezia, a tapeworm infesting sheep, whic
t most fully, the germ plasm is almost exclusively

—— and the spermatogonia may even undergo 4 *

of amitotic reduction by which sperms are formed without ever havité
pamed through mitosis, In general, amitosis is most common

1909] GRIGGS—AMITOSIS IN. SYNCHYTRIUM 135

regions of excessively rapid growth, where the nuclei are small and
have scant cytoplasm, while the larger nuclei, better supplied with
cytoplasm, divide by mitosis. This leads Cumtp to conclude (5, p.
292): “In short I am inclined to believe that amitosis is associated
with conditions where the demand for material or perhaps for some
particular substances exceeds the supply.’’ The behavior of the
nuclei of Synchytrium is distinctly opposed to the generalization of
this hypothesis, for in Synchytrium amitosis is most marked when the
nuclei are largest and the ratio of nuclei to cytoplasm is at a minimum.
While a condition of “hunger”? may very well be assumed to exist in
the cells of a rapidly r egenerating organ or in a growing embryo, it
ane be ascribed to a dividing cyst of Synchytrium, because growth
is very slight after the division of the primary nucleus, while the sup-
ply of nutriment from the host is presumably as great as before.

But in all of these cases the conditions of growth demand an
“xcessively rapid multiplication of nuclei, and indicate that the
process of nuclear reproduction is pressed on so rapidly as to give no
*pportunity for the rhythmic pause occasioned by mitosis. If the
Bs a growth and reproduction are independent, as many obser-
ae indicate, we may suppose that when the stimulus to division
Pe excessive the nucleus divides directly, without waiting for
but So pause necessary in mitosis. If the stimulus were

gnuly stronger than in mitosis, a slow and orderly division of the
thongs would result, and the daughter nuclei would be mostly
Oey if the stimulus became greater the process would be
Portion of ue finally the nucleus would explode and a large pro-
at all. r.: chromatin would never succeed in forming new nuclei
orthodrom; Is exactly in line with CHILp’s view that amitosis 1s an
always a Process which “.... pushed to the extreme mast
that “it ig ae the total destruction of the original substances,” So
ut there is Coens that degeneration frequently follows amitosis,
eee reason for supposing that it must always follow, and
Cover rie that it does not.” While this hypothesis may not
0 accord Cee again amitosis, which superficially would seem
iating the . hunger hypothesis, it would afford a basis for
"esehetating hon-mitotic divisions in Synchytrium with those in
Members, embryos, and other rapidly growing tissues.

136 BOTANICAL GAZETTE [FEBRUARY

Knowledge that in certain instances the reproductive cells of a
species are independent of mitosis for their origin must affect current
theories of heredity, which, since the renaissance of MENDEL’s law,
have leaned very heavily on the individuality of the chromosomes and
their separation in the reduction division. CHILD rejects the chromo-
some theory in any universal application. He believes (p. 290) that
“these processes appear to consist essentially in the production of new
nuclear material like that already present and without the periodical
recurrence of metamorphosis. The act of division is very probably
a mere incident of the increasing volume of substance.’’ Accordingly
he is inclined to doubt the constancy of the chromosome number it
the tapeworm, although he feels that the facts are too difficult of
determination to admit of certainty. In Synchytrium, likewise, the
minuteness of the nuclei makes determination of the chromosome
number so difficult that one hesitates to dogmatize. But in all the
many cases in which the chromosomes could be counted on the spindle
the number seemed to be constantly four (cf. fig. 36b). The same
number was given provisionally by SrEvENS (13) in his first paper
and is shown by the drawings of his second paper (12). This maiter
may, however, be left for consideration later, after the mitoses have

en worked out in detail. But whether the chromosome number is
found to be constant or variable, it is obvious that our theories of
heredity will require considerable revision.

SUMMARY OF RESULTS

The numerous peculiarities in the cytology of Synchytrium occu!
mostly in a somewhat definite period of irregularities immediately
following the division of the primary nucleus.

_In this stage direct division of the nucleus is more frequent than
mitosis. ‘This takes place by at least two processes:

a Nuclear gemmation.—The karyosome of the parent nucleus
Sives off a small karyosome which migrates through the nuclear mem
brane, forms a vacuole and a membrane about itself, and becomes 4”
independent small nucleus, the whole looking like a budding y -
Plant. This process is repeated until the parent nucleus is CO
verted into small nuclei, often forming a definite group.

2. Heteroschizis.—The membrane of the parent nucleus dissolves

1909] GRIGGS—AMITOSIS IN SYNCHYTRIUM 137

and the karyosome fragments into a number of pieces, each of which
becomes a new nucleus, thus giving rise to a morula-like cluster of
nuclei.

These nuclei at later stages undergo mitosis and their descendants
form spores and become the nuclei of succeeding generations.

No variation in the number of chromosomes in any of the nuclei of
the plant has been detected.

Onto STATE UNIVERSITY
US

LITERATURE CITED

1, Cap, C. M., Studies in the relation between amitosis and mitosis: I,
The development of ovaries and oogenesis in Moniezia. Biol. Bull. 12:
89-114. 1907.

2. , Idem: Il, The development of the testis and spermatogenesis in
Moniezia. Biol. Bull. 12:175-224. 1907.

3: ——,, Idem: III, Maturation, fertilization, and cleavage in Moniezia.
Biol. Bull. 13:138-160, 1907.

4. , Idem: IV, Nuclear division in the somatic structures of the pro-
glottids of Moniezia. V, General discussion and conclusions concerning
amitosis and mitosis in Moniezia. Biol. Bull. 13:165-185. 1907.

5 Sg, Amitosis as a factor in normal and regulatory growth. Anat.

‘ Anzeig, 30:271-297. 1907.

' Glaser, 0. G., Pathological amitosis in the food ova of Fasciolaria. Biol.
Bull. 13:1-4. 1907.

7. Grices, R. F., On the cytology of Synchytrium. IL, The rile of the cen-
‘somes in the reconstruction of the nuclear membrane. Ohio Nat. 8:
, 277-286. 1908,
: — S., On the nucleus of Synchytrium puerariae Miyabe. Bot. Mag.
. okyo BLSII8. 1907.
10, ___? On the cytology of Synchytrium. Centralbl. Bakt. 197:538. 1907:

os On

ue a disease caused by’ Synchytrium puerariae. Bot. Mag. Tokyo

22:1, 1 :
ai;
jue Karyodermatoplast, a nuclear-membrane-forming body {in
I. Seen Bot. Mag. Tokyo 22:205. 1908.

Ye. 5: Se Some remarkable nuclear structures in Synchytrium. Ann.
13. StE : * 4. I 7 :
decipiens Pre ano A, C., Mitosis in the primary nucleus of Synchyirium
- Bor. Gazerre 353405. 1903.
The EXPLANATION OF PLATES III AND IV
“ompensating Were all made with a Spencer 1.5™™ immersion objective and
cular 12, giving magnification of 2130, excepting fig. J, for

138 BOTANICAL GAZETTE [FEBRUARY

which ocular 2 (magnification 355) was used. They were reduced } in repro-
duction, canceling the enlargement due to the camera and rendering them the
same size as they were seen in the microscope.
E II

Fic. 1.—A lateral section of a cyst, showing numerous groups of small nuclei
due to nuclear gemmation.

Fic. 2.—One of the groups shown in fig. 1.

Fics. 3-6.—The breaking-up of the mother karyosome preparatory to the
migration of the chromatin.

Fic. 7.—Bipartition of the mother karyosome to form equal daughters.

Fic. 8.—Nucleus with a large number of daughter karyosomes lying on the
nuclear membrane, only one hemisphere of which is shown.

IG. 9.—A nucleus with one of the daughter karyosomes pressed against the

nuclear membrane; three small nuclei which have budded off from it near by.

Fic. 10.—Daughter karyosome constructing its nuclear cavity and membrane.

Fic. 11.—Daughter nucleus complete but still closely appressed to the mem-
brane of its parent.

Fics. 12, 13.—Karyosomes of daughter nuclei separated from the parents
but their membranes still in contact.

Fic. 14.—Daughter nucleus separated from its parent but lying close by.

Figs. 15~17.—Nuclear gemmation from the spirem stage. (Fig. 16 is one
of the large nuclei from the center of the cyst from which jigs. 1 and 2 were taken.)

Fics. 18-20.—Resultant groups of small nuclei.

PLATE IV

—Stages in the degeneration of the parent nuclei.
Fics. 24-26.—Nuclei from which chromatin has been thrown out in large
quantities and is mostly degenerating without forming new nuclei.

Fic, 27.—Beginning of heteroschizis; nuclear membrane di ving, karyo-
some slightly irregular.

Fic. 28.—Membrane and nuclear cavity lost, karyosome much enlarged.
Same cyst as Jig. 27,

a 29.—Karyosome lobed. Same cyst as figs. 27 and 28.
Bo, Ait 3°-32.—Karyosomes broken up, nuclear membranes appearing —

Pics oh karyosomes, (Fig. 32 is from the same cyst as figs. 27-29:)
— 33-—A very large cluster complete. 5 ided
by uaa Segment from a summer sorus whose nucleus has a?

Fic. 35.—A cluster of
Sig. 33, together with a sing

Figs. 21-23.

uster of spindles similar to Jig. 35; b, a solitary spindle from
2s ; bably a cluster of spindles disintegrating.
°S. 37, 38.—Similar clusters of spindles in anaphase.

Fic. 39-—A nucleus with dee;

* * m
i i . Ply staining granules on its membrane, {
which radiations are 8iven off into the sone
ting by constriction.

Fic. 40.—A nucleus fragmen

ATE TL

ray

GS on SYNGHYTRIUM

TE FF

PLA

GRIGGS on SYNCHYTRIUM

VASCULAR ANATOMY OF THE SEEDLING OF
MICROCYCAS CALOCOMA [j

CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY 122
HELEN ANGELA DORETY
(WITH PLATES V AND VI)
HISTORICAL

DeCanDoLLe (5) in the first account of the genus Microcycas
expresses the opinion that it holds a position among cycads inter-
mediate between Dioon and Zamia.

A historical sketch of the literature concerning M icrocycas calo-
“oma may be found prefacing the taxonomic account of the species by
CALDWELL and Baker (z), who succeeded in procuring material
from Cuba. The only part of their description which concerns us
here is the statement that the stem has a single vascular cylinder.

In a second paper, intended primarily to present the conditions
the Teproductive structures, CALDWELL (2) has described incidentally
—s the superficial characters of the embryo and seedling. Certain

» Which he naturally overlooked in so comprehensive a work,

pi = brought to light by a more detailed study having the seedling
alone as its object.

INVESTIGATION

ee first place, it is not the root which is seen to emerge from the
t! bend downward, as described and pictured by this author;
Pa of events in the process of germination is the same as
Cyeads which * Ceratozamia (7), the same which occurs in the other
Species of "ae have germinated, Dioon edule, D. spinulosum, and
elongating ¢ a. The root is not yet formed when the base of the
i : embryo Tuptures the seed coat in the small, less indurated,
When it emerges, it still bears the small brown
only remains of the coleorhiza, an organ which, in
*Potophyte Th He embryogeny, is by far the larger portion of the
the soil, Th united cotyledonary stalks bend downward toward
ty] ot May not pierce the coleorhizal disk until the latter
{Botanical Gazette, vol. 47

. region.
disk “onstituting the
‘arly stages of t

140 BOTANICAL GAZETTE [FEBRUARY

has touched the soil; indeed its appearance in some seeds is preceded
by the exit of the plumule.

A second detail in which this description must differ from the
one cited is that the usual number of cotyledons is only two. They
never wholly emerge from the seed. Their common stalk, which
forms a sheath about the plumule, is soon ruptured in many places
by the radial growth of the latter, and its decay causes the seed to drop
away.

The appearance of embryos of Microcycas, like those of Cerato-
zamia described by CHAMBERLAIN (3), examined at short intervals
during the period between fertilization and germination, indicates
that there is no resting stage; and if at any time the seeds are allowed
to dry, the embryos are killed. LANnp (10) relates that he has reason
to suspect the same condition in Ephedra.

CALDWELL has called attention to the adhesion of embryo to endo-
sperm. This adhesion is greater than in any of the cycads I have
studied, except Zamia. So intimate is the union, that by using ordi-
nary precautions, one does not separate them entirely, but always
carries some few layers of the endosperm through the microtome
and stains with sections of the embryo.

Coincident with the close adhesion of embryo to endosperm is the
remarkable fusion of the cotyledons. Although these always arise
as two distinct organs, the fusion in older embryos is so complete
that several were sectioned from the apex to within a distance of
°.5™™ above the tip of the plumule without discovering any trace of
the characteristic seam made by the meeting of the adaxial epiderm
layers (fig. 2). The epidermal cells themselves disappear in mally
cases slightly above the meeting of the inner faces over the plumule.
The seam was not seen to extend to the surface at any level in these
embryos, although there were conspicuous superficial sutures (fig- 1)
Leo them. The plumule is liberated by the splitting of the
sheath into strips, each of which simulates the petiole of an individu
cotyledon.
has not been Teported of a cycad.

Each cotyledon may have eight or ten vascular strands; all at
collateral, with normal orientation. Apparent exceptions may occur
immediately above a dichotomy or immediately below a fusion oF 4

To my knowledge so complete a fusion of cotyledons

1909] DORETY—MICROCYCAS CALOCOMA 141

approximation. In these situations, the xylem of one strand faces
that of the other, giving to the inner one an inverse orientation
(@, jigs. 1, 2). When the fusion of two bundles is complete, the
combination presents the appearance of a single concentric

All the cotyledonary traces are derived from the branching of
three, which join the central cylinder (fig. 3) in a manner similar to
that described by Marre (11) and TuressEN (12) for Dioon edule,
and by the present writer (7) for Ceratozamia. The wood remains
endarch as far out as the sheathing base of the cotyledons (jig. 13);
it becomes mesarch in this region; and in the upper portion of the
blade the wood in the greater number of the strands is exarch.
Transfusion tissue is abundant, and in close connection with the cen-
inpetal xylem. Mucilage ducts alternate with the cotyledonary
traces. Tannin cells are conspicuous in the peripheral region.

The hypocotyl has no vascular plate, no protostele; the passage
from stem to root is therefore easily studied. The four cotyledonary
strands remain distinct throughout this portion of the axis, only fusing
laterally with the few elements of the leaf traces still remaining to
sai a@ very imperfect siphonostele. Their elements finally unite
with those of the four root poles. The metaxylem and phloem
re as usual, and the resulting portions swing to right and left,
5 Pies half of the phloem of each joining with the left half of that

ne, With sometimes the lowermost extremities of leaf-trace
Se alee (fig. ga). There is thus produced the character-
fark cture, four groups of phloem alternating with four double-

zy xylem groups. Irregular proliferation of the medulla

- 6)

a tes. the phloem group again into its two constituents
oo of the hy pocotyl were discovered the remains of a
tissue could . * cambium. No traces, however, of any vascular
tWo expan ed | etected outside the central system In seedlings with
"The root .. and several others developing.
4, 6) With a red the seedlings under observation was tetrarch (figs.
Mucilage ja uction to triarch toward the tip, in some cases.
into the root <o numerous in the hypocotyl, but do not penetrate
Neither pericycle nor endodermis is distinct in these

142 BOTANICAL GAZETTE [FEBRUARY

young roots. ‘The root tip differs in no observable respect from that
of Ceratozamia.

In the stem four large groups of leaf traces alternate with the
cotyledonary strands, just above the cotyledonary node (fig. 3).
Higher up (fig. 7), they close in, and together constitute the central
cylinder of the axis. Taking a generally vertical course, these traces
branch and anastomose until they reach a position so near to the
growing points of leaf and stem that the vascular tissue is still pro
cambial. In this position, even before the procambial strands from
leaf and stem apex have united, the traces destined for the leaf margins
manifest the phenomenon of girdling.

At first the wood of the leaf traces is endarch. F ig. 8 represents
a portion of a leaf-trace girdle, and jig. 9 a cross-section of two vertical
strands from the middle of the same leaf at the same level. Figs. 10,
11 illustrate stages in the transition from the endarch to the mesarch
condition. The wood becomes exarch at a relatively low level in
some of the foliar strands, and there is considerable irregularity in
different traces in this respect. Fig. 12 is a drawing of one of the
traces (5) in jig. r4, still in the region of the stipules. The othet
traces of the same leaf retain, at this level, a few elements of centrifu-
gal xylem; this strand is entirely destitute of them. Further—and
I have attempted to represent it diagrammatically—the wood in al
the traces of this petiole is more nearly exarch than in those of the
older leaf (/2), although the section of that leaf which is here repre:
sented is higher up in the petiole than that of the younger leaf. Inspité
of such irregularities, however, the statement holds that the xylem
entirely centrifugal near the base of both cotyledons and leaves; :
that it gradually diminishes to the vanishing point in proportion 10 -
appearance and increase of centripetal xylem in the ascent of the
cotyledonary blade and leaf petiole. ;

The section represented by fig. 13 is very close to the leaf bas
It shows the increase in the number of bundles entering consecull¥®
leaves, and also the meriphyte’s gradual assumption of the 2 arrang
ment from the open arch of the early leaves. Transfusion tissu s
present in these traces, a

The strands are all collateral. Frequent branchings and appro
Mations occur, and real fusions are common, most noticeably of thos

1909] DORETY—MICROCYCAS CALOCOMA 143

strands which are brought to the center of the meriphyte and form the
flanks of the 2. When two of these strands are fusing, the centripetal
elements of both xylem groups are gradually eliminated and the
protoxylem groups are therefore brought together and finally united.
This common group of protoxylem, then, is surrounded by the united
metaxylem, and outside of this the phloem of both bundles may
almost encircle the xylem, completing the delusion of a concentric
bundle (figs. 15, 16).

ucilage ducts occur in stem and _ leaves. They sometimes
extend through the petiole as far as the bases of the pinnae.

The characteristic cycad ramentum is prominent, especially upon
the unfolded leaves. The hairs are one-celled. Fig. 18 shows the
ip of a young pinna bearing these epidermal outgrowths.

Tannin cells occur in the periphery of the petiole, and may extend
well into the pinnae. They are in close relation with the mechanical
tissue, as represented in jig. 17.

DISCUSSION

The suppression of one of the cotyledons of Ceratozamia illustrates
. « Process by which the monocotyledonous condition may be reached;
* fusion of these organs in Microcycas affords an illustration of
method, one in harmony with the well-known theory of
= saath Experimentation with some of the monocotyledonous
inat] ons listed by Courter and CHAMBERLAIN (5) shows that
ae of them the former process has taken place.
ing ss Sous of the cotyledons of Microcycas has a further mean-
ade - ea study of a series of juvenile gymnosperms.
'o me to re ng at the tips of the cotyledons in so many cycads seems
Present an ancestral condition of polycotyledony. Many
mia and Dioon spinulosum in my collection have four
= bes, extending, in some cases, three-fourths of the
cotyledon. A young embryo of Dioon spinulosum
edons, and one of Pinus edulis with twelve cotyledons
the pine aig groups have a remarkable similarity. Of course,
Osperm 4 ©ns soon escape from the small, comparatively dry
and thin seed coats, and develop exteriorly to their full

144 BOTANICAL GAZETTE [FEBRUARY

size, which is in foto much greater than that attained by the two
cotyledons of a cycad. But let the conditions bé such as to cause
in the pine seed the production of the massive endosperm and indurated
coats that characterize the cycad seed; and let the cotyledons be con-
fined within the moist endosperm until they attain their full size, -
with the pressure of this growth forcing them into such intimate con-
nection with it as to cause difficulty in distinguishing between them,
and bringing the tips of the cotyledonary vascular strands into intimate
contact with the endosperm, a condition reported by WorsDEtt for
Cycas revoluta (13) and by Tutessen for Dioon edule (12): under
such pressure, the inner faces of the pine cotyledons would be very
intimately united and the question naturally suggests itself, What
would become of the epidermis of these inner faces ?

The alternation of mucilage canals with the cotyledonary vascular
strands in cycads, and its ready relation to the peculiar condition
found in pine cotyledons, may be used as evidence for a theory 0
fusion as well as for one of splitting, as Hitz proposes (8).

The absence of the protostele in the hypocotyl of Microcycas in
contrast to the condition found in Dioon edule and Ceratozamia may
not have any significance in the light of recent investigation. That
the protostele is, in general, the most primitive condition of the vascular
axis may be true; but that this structure must occur in every prim
five vascular plant is, of course, not true; neither are we to regard as
primitive all plants in which it is found. CHRYSLER (4) has found
in members of the Araceae. There are other characters, howevtt,
which seem to indicate a greater advance than that made by Cycas%
Encephalartos, or even Ceratozamia. These are the single stele and
the degree of elimination of the cortical cambium, which, in the cyeaé
stems, produces this vascular tissue. However, the large proportion .
centripetal wood in the foliar traces is an offsetting primitive charactél,
which must be weighed in the same balance. :
ise polyspermy would seem, at first sight, to y

iiae, ght of evidence on this side ; but it is possible that
primitive feature is a recurrence rather than a direct inheritance
what JEFFREY calls a Coenogenetic, rather than a palingenetic, =
acter, JUEL (9) found as many as twenty sperms in the pollen "
of Cupressus Goveniana, and no one can believe that Cupressus -

1999] DORETY—MICROCYCAS CALOCOMA 145

tetained this primitive feature when all the closely related genera
have discarded it.

However that may be, this curious combination of characters, and
the absolutely unique archegonial development, are features to be
reckoned with by those who, in the future, when all the evidence is in,
will be in a position to decide upon the phylogenetic place of Micro-

cycas.

What is to be thought of the early appearance of girdling is scarcely
worth saying until we learn something definite concerning the cause
of girdling itself. The theories now in the field approach only
temotely to the causes lying at the foundation of the phenomenon.
It is probable that it may be relegated, like so many other problems,
to the domain of cytology. That cell division takes place much more
frequently in the horizontal than in the vertical direction in every
Portion of the axis is clearly evident.

SUMMARY

1. There is no resting stage in the development of the embryo of
Microcycas calocoma.

2. The germination is hypogean.

3: The root is a delayed organ, as in Dioon and Ceratozamia.

4 There are two cotyledons as in all cycads (except, perhaps,
Encephalartos),
5 The cotyledons are often fused to form one organic whole, the
Plumule escaping by bursting the sheath, |

» Mucilage ducts alternate with the 8-10 cotyledonary strands.

7. The cotyledonary node is similar to that of Dioon edule and

oo but the vascular cylinder of the hypocotyl is a siphono-

. tle © hypocotyl contains no cortical vascular tissue, although
= nants of a br oken-up cambial zone.

i vescular strands of cotyledons and leaves are endarch at

“xarch in the upper portions. The exarch condition obtains

out Most of the length of the petiole.

is catia ° of the marginal leaf traces takes place while the tissue

I, : .
The root is tetrarch, but may reduce to triarch toward the tip.

146 BOTANICAL GAZETTE [FEBRUARY

Grateful acknowledgments are due to Professor JoHN M. Courter,
and Dr. W. J. G. LAND, under whom the investigation was conducted,
and to Professors CHARLES J. CHAMBERLAIN and Otis W. Catp-
WELL for material.

THE UNIVERSITY OF CHICAGO

LITERATURE CITED

1. CALDWELL, O. W., AND BAKER, C. F., The identity of Microcycas calocoma.

AZETTE 433330. 1907.

2. CALDWELL, O. W., Microcycas calocoma. Bot. GAZETTE 44:118. 1907.

3. CHAMBERLAIN, C. J., Preliminary note on Ceratozamia. Bot. GaAzerTE
43:137. 1907.

4. Curyster, M. A., The development of the central cylinder in Araceae and
Liliaceae. Bor. GAZETTE 38:161. 1904.

5. Courter, J. M., anD CHAMBERLAIN, C. J., Morphology of angiosperms
206. 1903.

6. DeCanpoite, A., Microcycas calocoma. DC. Prodr. 16:558. 1868.

7. Dorety, Heten A., The seedling of Ceratozamia. Bor. GAZETTE 46:205.
908.

I
8. Hitt, T. G., anp DEFraine, E., On the seedling structure of gymnosperms.
Annals of Botany 20:47. 1906.
9. JuEt, H. O., Ueber den Pollenschlauch von Cupressus. Flora 93:56. 19°
10. Lanp, W. J. G., Fertilization and embryogeny in Ephedra trijurca. Bor
GAZETTE 44:273. 1907.
11. Matte, H., L’appareil libéro-ligneux des Cycadacées. Caen. 1904:
12. TutEssen, R., The vascular anatomy of the seedling of Dioon edule. Bot.
GAZETTE 46: 357. 1908.
13. Worspett, W. C., Comparative anatomy of the Cycadaceae. Jour. Lin’
Soc. Lond. 33:437. 1898.

EXPLANATION OF PLATES V AND VI —

The drawings were made with the aid of an Abbé camera lucida. The following
abbreviations have been employed: A, B, C, D, main cotyledonary traces; ©
cotyledonary sheath; ¢fx, centrifugal xylem; cpx, centripetal xylem; -_
¢, epidermis; /, group of leaf traces; J, leaf; m, medulla; md, mucilage ”
mx, metaxylem; ph, phloem; px, protoxylem; r, ramentum; #, tannin cells

PLATE V ‘

Fic. 1.—Transverse section near the middle of the cotyledons, showing tie?
complete fusion and the large number of cotyledonary strands. <8. le
=e 1G. 2.—Transverse section of cotyledons 0.6™™ above the tip of pl

Fic. 3.—Diagram of stele 40 # above the cotyledonary node.

®,
go

aon

— aN “7
a—ae¥

sheet ie
Cer
ZR
SON
HEY FHA
a =e=.»

Ie
id

f,
(
Sez

if
ey
ad
Yo
Bs

fh
Hy
ec)
ie
@.

an
ee. &.
Wet
UE:
pe oie
“dese,
<1 Pris
am

| Lis
ik

eke 4 Lore def

. PLATE. VI

199] DORETY—MICROCYCAS CALOCOMA 147
Fic. 3a.—Detail from fig. 3 showing one of the four cotyledonary traces near
its point of insertion. X60.
Fic. 4.—Base of the cotyledonary trace shown in fig. 3a. X60
Fic. 4a.—Diagram of stele of hypocotyl showing method of formation of root

arrangement.

Fic. 5.—Exit of lateral root. 60.

Fic. 6.—Diagram of root stele.

Fig. 6¢—Detail of one of the root poles. 60.

Fic. 7.—Transverse section of the central vascular system above the coty-
ledonary node. It is composed of four groups of leaf traces. Semidiagrammatic.
_ Fic. 8—Longitudinal section of portion of girdling leaf trace taken from
transverse section of seedling. X60.

PLATE VI
Fic. 9—Transverse section of median traces of same leaf from same section
X 380.

Fig. 10—Transverse section of leaf trace slightly above that represented in

Fic, 11.—Transverse section of same bundle 60 & above section represented
in fg. 10, X 380.

* 12—Exarch bundle from petiole 8™™ from base. X 380.
ine » 13.—Transverse section of embryo, showing the sheath formed by the

of the cotyledonary petioles, the increase in the number of leaf traces in

ee leaves, and their gradual assumption of the Q arrangement. X8.

‘ '4-—Transverse section of three leaves in their natural arrangement,
ree the relative amount of centrifugal wood at different levels of the petiole.

: % Le ae of two leaf traces. X 380.
_. Fusion of. some of the xylem elements of same to form a quasi-
ee strand. X 380, =

1. 1 . : ; : fe
ce se ag of peripheral region of tissue of petiole showing position of

Fig. 38.

—Tip of young pinna sho wing unicellular hai entum X 760.

BPerereR ARTICLES

THE NATURE OF BALANCED SOLUTIONS

In his recent “Note on balanced solutions”! Professor LoEw criticizes
some of my statements. The following reply is inspired solely by the
desire to obviate if possible any misunderstanding regarding the nature of
a balanced solution.

A balanced solution is defined by Lors as one in which the toxic
effects which each salt would have, were it alone present in solution, are
inhibited by one or more antagonistic salts in the solution.

Professor Lorw objects to the term toxic as applied to calcium and
potassium salts. His statement that I and a pupil claim to have dis
covered the poisonous action of potassium and calcium respectively is
evidently due to a misapprehension. On the contrary, we treated them as
fully accepted facts, and it was a surprise to us that he should call them
in question. The poisonous action of a salt must be determined by com-
paring its effects with those of pure distilled water, or, in the case of strong
solutions, with the effects of an isotonic balanced solution or an isotonic
solution of an indifferent substance, if such can be found. In the absence
of the facts needful for such a comparison, it is not possible to say whether
the effects observed by him are to be regarded as toxic or not. At te
concentration chiefly used in my experiments (.12 M) roots of wheat
reached a length in KCl of 63™™, in CaCl, of 84™™, in an isotonic balanced
solution of 360™™, and in distilled water of 740™™, I may add that for
certain forms of Vaucheria KCI and CaCl, at the dilution of .oor Mf (of
even less) may be toxic, inasmuch as they kill the algae in three oF four
days, while in distilled water or dilute sea water of a hundred times greatet

* Bor. Gazerre 46: 302. 1908. :
Botanical Gazette, yol, 47] 2

1909] BRIEFER ARTICLES 149

toxicity of the anions is negligible) by supposing that the sole poisonous
constituent is magnesium, whose toxic action is completely inhibited
by the calcium present. But on this view it is clear that the potassium
and iron are completely superfluous from the standpoint of a balanced
solution. Knop added them for nutrient, not for balancing, purposes,
nor is there reason to suppose that he was aware of antagonistic salt effects.
At the concentrations at which he worked these effects are not at all evi-
dent with such flowering plants as were used in his experiments. Under
these circumstances the discovery of antagonistic salt effects is very improb-
able. For most of the plants for which Knop’s solution is employed at
its ordinary concentration, it is not a balanced solution, because its indi-
vidual components are not sufficiently toxic to require balancing.

That to Professor Lorw is due the very great credit of investigating
the antagonistic action of magnesium and calcium, and of making clear
- ¢conomic importance, is acknowledged by all. These and other inves-
tigations made by him in the difficult and obscure field of the function of

mineral salts are of the highest value. Together with the experiments
of other investigators they have thrown much light on antagonistic action.

uch, however, as Professor LOEW apparently does not believe in
Seneralizing the principle of antagonistic action, as Professor Logs has
done in his theory of balanced solutions, but prefers to restrict it to the
Single case of Mg vs, Ca, I find myself quite unable to agree with him.
In the course of a series of experiments on wheat I have found antagonism
“tween each of the following pairs of salts:

NH, vs. Ca NH, vs. Na Mg vs. K Na vs. Sr
Ks. Ca NH, vs. K Na vs. Ba K_ vs. Sr
Na us. Ca Na vs. K K_ vs. Ba Mg vs. Sr
Mg vs. Ca

. One Who has to deal with such a series of facts can hardly be expected

adopt a view which accounts for only one of these cases and ignores the

Pas €xplains them as due to the formation of double salts, particularly

pin og explanation is wholly untenable in view of the facts of dissocia-
~~": J. V. Osrernour, University of California, Berkeley.

150 BOTANICAL GAZETTE [FEBRUARY

THE EXTRAFASCICULAR CAMBIUM OF CERATOZAMIA
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY 123
(WITH PLATE VII)

The anatomical features presented by the seedling of Ceratozamia
described in a previous paper (3) gave promise of such phylogenetic impor-
tance that the study has been continued upon older stems. The interest
centers mainly in the extrafascicular cambium—its origin, its distribution,
and its failure to differentiate xylem and phloem which could be clearly
recognized as such. In a careful examination of microtome sections of
over eighty seedlings, varying in age from a few months to two years,
only one small extrafascicular bundle was found.

The origin of the concentric vascular zones ‘of cycad stems puzzled the
early anatomists. BRoNnGNIART (1), failing to distinguish the phloem in
these zones, regarded them as the equivalent of the seasonal wood rings
of dicotyledons. Von Mont (8) said that they were an aggregation of
bundles which passed out from the central cylinder and, running down-
ward in the cortex, grouped themselves in a ring. LEsTIBoNDOIS (4) S4¥
individual bundles in the cortex, but thought they foreshadowed the
' breaking-up of the central stele, and so considered the cycads as a transi-
tion group from dicotyledons to monocotyledons. METTENIUS, to whom
we are indebted for much of our knowledge of cycad anatomy, made n0
attempt to explain the extra vascular zones. CoNsTANTIN and Morot (2)
thought that the tissue arose from the pericycle.

All the early research was confined to mature stems of Cycas and
Encephalartos. In 1890, Sotms-LauBacH (7) reported the absence of
such zones in Ceratozamia mexicana. In 1896 WorsDELL (9) added
Macrozamia to the list of cycads in which these thickenings occur, and in
1898 (10) recorded their absence from the stem of Stangeria; in 1900 (11)
he found them in the root of Bowenia spectabilis. His study of seedlings
of species of Cycas (10) demonstrated to WorspELL that the extralas
cicular zones in mature stems arose as independent cortical cylinders
arranged in distinct series, the innermost ones being composed of primary
Haale, By a later growth of the central cylinder they become 4pP
to - periphery and flattened radially. This study brought to him @
to his readers a conviction of the truth contained in his earlier suggest?”
that the cycads are closely related to the Medullosae, which are polystelic
_ ferns. Martr’s study of Cycas siamensis and Encephalarlos

(5) is an almost perfect demonstration of WorspEL1’s theory:

1909] BRIEFER ARTICLES 151

My work upon Ceratozamia will come as cumulative evidence. SOLMs’s
statement concerning the absence of extrafascicular zones is correct as
far as the seedling is concerned; but the presence of extrafascicular cam-
bium in great abundance led me to make a careful search to discover its
relation to the central cylinder. In the study of seedlings described in the
previous paper I was unable to do this on account of the disturbances
caused by mucilage ducts, which are large, abundant, and irregular in
distribution. In the study of older stems I have been more fortunate.
Four-year-old seedlings have in the hypocotyl clearly distinguishable
tings or cylinders of cambium. The cylinders are arranged in several
series. Those of the innermost series, though decidedly flattened, are
the most distinct.

Fig. 1 represents diagrammatically a section of the hypocotyl slightly
below the exit of the cotyledonary traces. The innermost cylinders
(a, b §) arise in the pericycle near the transition region and are of primary
mika They extend well up in the stem, though pushed outward by the
horizontal cotyledonary traces. The other rings appear later. It would
seem that the single bundle described in the previous paper (3, /ig. 30, 2)
Was differentiated from one of the outermost series of cylinders.

Fig. 2 is a detail of the inner portion of half the section represented in
Ag. 1. One large cambial ring (a) is represented, and ends of two others
5). Several small rings (e, r) suggest how concentric bundles might

ise cause of the flattening is manifestly the enlargement of the central
Ea ae consequent pressure upon the inner side of the cortical
sa (a, b, s). The final result is a central cylinder surrounded by
ha cs or less imperfect zones of cambium cells. The xylem and
i hich these cells might produce would be oriented differently ;
| — on the centripetal side of the zone would be differentiated toward
aca the phloem toward the center of the stem. But in
we aa, inner cambium of each zone would cease to function, and
with oo have successive zones of alternating xylem and phloem all
Would be i. Mentation. Occasionally a bit of the elongated cylinder
*ylem would ennected, the cambium would round out, the growth of
All these ¢ climinate the pith, and a concentric bundle would result.
like —. are found in the Medullosae, from the distinct fern-
- stellata Fg Medullosa Solmsii and M. anglica to the condition of
have oy Ich Closely resembles Cycas revoluta. :
PELL (9) ha yet examined mature stems of plants of this genus. Wors-
8 confirmed Sorms-LauBacn’s statement that the extrafas-

152 BOTANICAL GAZETTE [FEBRUARY

cicular zones are lacking; but MATTE (6) reports that he saw them in stems
which he examined. I am therefore in doubt whether it is an absolute
failure to function, or only a delay; in either case we have an indication
that the cortical vascular tissue is a disappearing character.

Another feature to which attention should be called is the constant
occurrence of centripetal xylem in the cylinder of the hypocotyl and in the
bases of the cotyledonary bundles. This is represented in fig. 28 of the
paper already cited and also in fig. 2 of the present one. It is often rela-
tively more abundant than in some stems of Lyginodendron. The stem
cylinder above the cotyledons is endarch, the leaf traces becoming mesarch
almost immediately after leaving the cylinder.

Acknowledgments are due to Professor Joun M. Courter and Dr.

. J. G. Lanp.—HELEN A. Dorety, The University of Chicago.

LITERATURE CITED

1908,

4. LEsTIBONDoIS, Mémoire sur la structure des Cycadées. Comptes Rend.
Acad. Sci. Paris 51:651. 1860.

5. Marre, H., Recherches sur Pappareil libéro-ligneux des Cycadacées. Caen.
1904.

6, » Note préliminaire sur des germinations des Cycadacées. Rennés:
1907

Miinch. 1832 Vermischte Schrifte
n 195. 1845. .

9. Worsvett, W. C., The anatomy of the stem of Macrozamia compared wit
that of the other genera of Cycadeae. Annals of Botany 10:601. 1896.

» Comparative anatomy of the Cycadaceae. Jour. Linn. Soc. Lon
don 33:437. 1898,

II. ———,, The anatomical Structure of Bowenia spectabilis Hook. Annals of

Botany 14:159. 1900,

10,

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i @ ZOO TN IES WL
| ON BS Pisens ant Os Pe ee
SRY Ph Osercsrcnne: o
AHO RUS Candas T? IY LOH a Me ay
WIN YOR (ees ee iperasit:
NY Reese eeratetteers
a Roe rei ASH SIE LOSS
a ANS OW, ace TE A Genes
AOS Wo eee | ER OY
A) XIV SN SGan! eee
HAL LYN SOAK \ fig ES
OG Oh caret:
OSA RT Neesan etree can
isa ACCU Es
line apace ieee cee:
ee piguasae
erely del. 2 q ey Le 4) a Ly e, ip 4
<i SE RE,
YOSEH e.

DORE
TY on CERATOZAMIA

CURRENT LITERATURE

BOOK REVIEWS

Gray’s new manual of botany

It has been generally known for some time that Grav’s Manual was under-
going a revision, so that the appearance of the seventh edition was no surprise:
in fact no book has been awaited by the botanical public, especially during the
past three years, with so much interest as this one.

The present edition, contrasted with the sixth edition, presents the following
more conspicuous changes: (1) a change in the geographical limits, namely by
excluding the region west of the 96th meridian instead of the rooth, and in extend-
ing northeastward to include the maritime provinces and a portion of Quebec and
Ontario; (2) the Sequence of the families, which for the most part is in accord-
ance with the system of EICHLER as elaborated by ENGLER and PRANTL; (3) the
removal of the keys leading to the species, in the case of most of the larger genera,
from the body of the text to a position immediately preceding the specific descrip-
te (4) the introduction of numerous text- or marginal figures; and finally
(5) in the use of a different system of nomenclature, namely the strict observance
of the Vienna Code, or the nomenclatorial rules adopted at the International
Congress held at Vienna in 1905.

The results of these innovations are that the flora treated is a somewhat
en neous one, the general arrangement and sequence of families is in
— with advances made during recent years in the classification of plants,

facility and certainty in the identification of species by the use of direct
= sis descriptions associated with accurately executed and reliable figures,

Ste “onsistency and uniformity in the use of plant names.

— *ppearance of the printed page is essentially the same as in the
tant specific She tions; the use of italics in emphasizing the more impor-
ba ae eg eristics is also retained. The treatment of genera and Species
ecroed with Conservative and rational; and generic limitations are in close
in Previons oe usage. ‘he number of species is considerably larger than
Our flora 7 ene Owing to the very active, careful, and exhaustive study of

The § recent years.
where they _ are judiciously distributed throughout the volume in groups
Salicaceae, 4 a useful, as, for example, in the Gramineae, ee
type of illustrat: Tuciferae; the Cyperaceae especially lend themselves to this
—__“"n, and here they are certainly at their best. In some cases,

1
beni %. L, AND FERNALD, M. L., Gray’s new manual of botany: a
States ang adj . flowering Plants and ferns of the central and northeastern United
Tevised, 9. Jacent Canada. Seventh edition, illustrated, rearranged, and extensively
* PP. 926, Jigs. 1036. New York: American Book Co. 1908. $2.50.

£53

154 BOTANICAL GAZETTE [FEBRUARY

for example in the Compositae, the illustrations have apparently suffered some-
what in the process of reproduction; thus for instance in the genus Crepis the
illustrations, as reproduced, add little or nothing to the text. In general, how-
ever, the illustrations are excellent, and one only regrets that they are not more
numerous

The presswork is exceptionally good; there are few typographical errors.
On page 500, however, ‘“‘Abizzia’’ occurs instead of Albizzia, which is evidently
a misprint.

The value of this work as a textbook should not be overlooked. The dis-
criminating text and complementary illustrations present the subject-matter in
a satisfactory way for teaching purposes. ‘The illustrations themselves are for
the most part insufficient for the hasty determination of the species by the student,
and they can be used to advantage only in connection with the brief but clear
descriptions. In this regard the book has no equal.

the whole this new edition of the well-known Gray’s Manual presents
a flora of the central and northwestern United States, and adjacent Canada in
a single volume of convenient size and moderate price, thoroughly revised to
date, incorporating the verified results of recent years of research, and fully
accords with the most advanced and universally accepted views of taxonomy—
J. M. Greenman.
Heredity

A recent volume on heredity, by Prorrssor J. ARTHUR THOMSON? of the
University of Aberdeen, “is intended,” as the preface states, “(as an introduction
to the study of heredity.” The writer has long been known as the joint author
with Geppes of the Evolution of sex, but is perhaps most widely recognized -
the translator of WrIsMANN’s works and the exponent of Weismannism. At
view of such a book in a botanical journal needs no apology, for much of the more
recent work in heredity has been done with plants, and moreover the book deals
with those general fields of biological research which must always be of equal
interest to botanists and zoologists. ‘These fields will ever be the meeting-ground
of botany and zoology, because in this class of problems the organism is treated
as such, and the fact that it is a plant or an animal is of minor importance.
The work is divided into fourteen chapters, and among the topics dealt ge
may be mentioned the physical basis of inheritance; heredity and vine
reversion; telegony; transmission of acquired characters; statistical and expel:
mental study of inheritance 3 theories of heredity and inheritance; heredity and
sex; and a final chapter i ‘ ‘ological results.

Heredity and in i

? THomson, J. ARTHUR, Heredi : York: GP:
, : 4 ty. pp. xvit+605. s. 49. New XOrs:
Putnam’s Sons, London: John Murray. Sk. te?

1909] : CURRENT LITERATURE 155

sive generations; and by ‘inheritance’ we mean ‘organic inheritance’—all that
the organism is or has to start with in virtue of its hereditary relation to parents
and ancestors.”

In the chapter on the physical basis of inheritance the author discusses the
phenomena and experiments connected with chromosome reduction and fertiliza-
tion, and concludes from the evidence that the chromosomes are the bearers of

reditary characters, but that ‘‘we should be chary in committing ourselves unre-
servedly to the conclusion that the heritable organization is exclusively resident
in the chromatin of the nuclei of the germ cells.” The chapter on heredity and
Variation contains a clear exposition of the facts and theories of mutation and
continuous variation. The author believes that both are important evolutionary
factors; that mutation, so far as present evidence goes, may have been a much
more important factor in plants than with animals; and that the distinction be-
tween “large fluctuations” and “small mutations” is merely a verbal one. Re-
garding the causes of variation he considers it “useful to say that variation is the
expression of a qualitative asymmetry beginning in gametogenesis.” ‘‘ Variation
18 a novel cell division.”

There is a lengthy treatment of the question of the transmission of acquired
characters or “somatic modifications;” with a critical analysis of the data usually
eg asevidence. The result may be stated in the author’s own words (p. 242):
- question resolves itself into a matter of fact. Have we any concrete evidence

oo believing that definite modifications are ever, as such or in any
oie. degree, transmitted ? It appears to us that we have not. Be to
BS rai that such transmission is impossible is unscientific. The
ee, of Gatton, PEARSON, and others are summarized, and under
Wes 5c study of inheritance an array of data from the work of MENDEL,
which hea . . CoRRENs, and many others of the recent school of sosebics
Sao to illuminate some of the obscure problems of hybridity, A
another part . discussed. These are largely the facts of Mendelism.
pietented € book blended, preponderant, and particulate inheritance are

. m

“nonstate are devoted to theories of heredity, which are largely theories
Which the a:

well-kn utho

sae muthor’s theory is presented, namely that the difference in the
Metabolism” ng ht difference in “protoplasmic gearing” or in the

Much-p ‘

too litle gea;cPetiments on the subject. The author also apparently attaches
accom sBnificance to the discovery that in many insects an extra chromosome
: Panies the female ax.

At
Very 2 am “ the work there is a representative bibliography of 48 pages,
Subject-index to the bibliography, and a general index to the volume.

156 BOTANICAL GAZETTE [FEBRUARY

The book is a broad and comprehensive treatment of the subject of heredity, a
veritable mine of valuable data concisely presented and clearly dis . The
worker in these fields will find it almost indispensable for reference, and the more
general reader will find it a very satisfactory and fascinating exposition of present-
day views on these problems.—R. R. GATES.

Laboulbeniales

In 18096 the first part of a monograph of this group was published by THAXTER.
The review in this journals gave a general account of its contents, which presented
the history, literature, and morphology of the group, in addition to the description
of genera and species known at that time. During the last decade material has
accumulated rapidly, and several preliminary papers describing it have been
published. There has now appeared a second part of the monograph,‘ which
brings together the material and illustrates it by a series of handsome plates. By
means of visits to European collections and to South America, and through numer-
ous correspondents, the new forms have multiplied remarkably. In the present
part nearly 350 forms are illustrated, which i th ber described to ab
500, included in more than so genera. In addition to these, more than 100 new
Species have been assembled since the completion of the present plates (in 195),
and these will be described and illustrated as soon as possible.

A brief review of the literature since 1895 is given, with comments on the
morphology, development, etc., of the group, based upon the new data available.
There is some modification in the conceptions of generic types, in the distinctions

vetween series, etc.; but the comparative morphology of the group remains esse
tally as it was presented in 1895. This will be somewhat disappointing to thos

Florideae; but the author wisely remarks that “‘it is foolish, on the basis of out
pee knowledge, to attempt an arbitrary settlement of the complex phenomens
of series among the fungi.” If he declines to be arbitrary about Laboulbeniales,
no one else can afford to be so. Nevertheless, he thinks the statement safe that
the group resembles the Florideae in some respects more closely than they 4
any other plants, while at the same time they’are more surely ascomycetes that
many forms included in this group.” He sees no reason why they should not wo
placed in the Pyrenomycetes, as a group coordinate with the Perisporiales, BYP”
creales, etc, :

This contribution, as the former one, is a model of painstaking and exact wot
Ee am carefully weighed statement; and it is also an illustration of the wealth
material available for those who can see.—J. M. C.

3 Bor. Gazette 23:216, 1897. i
a *THAXTER, ROLAND, Contribution toward a monograph of the Laboulbent
ceae. Part II. Mem. Amer. Acad. Sci. 13:219-469. pls. 28-61. 1908.

1909] CURRENT LITERATURE 157

African plants

Under the title Die Bliitenpjflanzen Afrikas,’ Dr. FRANZ THONNER, after list-
ing the chief botanical works and directing attention to some of the more
scattered literature pertaining to the flora of Africa and supplying a detailed
table of contents, gives a convenient key to the determination of the families of
African flowering plants; then he presents the main body of the work, namely a
key for the determination of the genera, which occupies about 540 pages. The
text is supplemented by 150 plates, and a map indicating the floral regions and
provinces of the continent.

; Each family is described as to the essential and most striking characters and
is Tepresented, moreover, by a full page illustration of a characteristic genus;
the number of genera and species of each family, so far as it occurs in Africa, is
- Furthermore, the number of species in each genus and their geo-
graphical distribution is indicated.
The author presents no bibliography in connection with the text, and there
8 comparatively little in the way of synonomy; it is, however, definitely stated
in the introduction that the limitations of families and genera are in accordance
with ENGLER and Pranrti’s Die natiirlichen Pflanzenfamilien and DALLA TORRE
and HarMs’s Genera Siphonogamarum, so that for critical identification it will
hecessary to use the book largely in connection with these and other important
Works of reference.
— The keys are ingeniously arranged, well contrasted, and lucid; the illustrations
clear and advantageously portray the general and detailed characters of the
Plant, and. well represent the different families. The volume also contains a
Carefully prepared comparative table giving the number of families, genera, and
rag their general geographical distribution; it also contains a glossary,
mc cal authors, and a useful catalogue of the common African plant-
ea. with the proper scientific name. oe
‘eleice oe : ; the work brings together in an epitomized form and in a single
“es in ormation concerning the flora of Africa that hitherto has been
Ted through many different volumes; hence it is a work which will be of great
Practical use both in the herbarium and in the field—J. M. GREENMAN.

ie... MINOR NOTICES
Nepenthaceae at 36 consists of a monographic treatment of the
Professor J ae the well-known writer and authority on insectivorous ey
following the os ape An excellent general account of the family,
————. ‘Sequence of this series, precedes the taxonomic consideration

‘Tuonner, FRanz,

5 Eng ;
Mocrariane” A., Das eich. Heft 36 (iv. rrr). Nepenthaceae von J. M.

. t 36 (
* PP. 92. figs. 19 (95). Leipzig: Wilhelm Engelmann. 1908. M 4.60.

158 BOTANICAL GAZETTE [FEBRUARY

of the group. A concise dichotomous key leads one direct to the species under
which are numerous references to literature, synonomy, a detailed description,
concise statement of geographical range, and a rather full citation of exsiccatae,
The author recognizes 58 species and several varieties for the one genus Nepen-
thes, of which 8 species and 4 varieties are here published for the first time.
The main body of the work is followed by an alphabetical list of artificial hybrids.
These are designated by the binomial under which is given, so far as known,
the names of parent species. The family is illustrated by 19 figures; a com-
plete index concludes the part. The production is quite in accord with previous
publications of this comprehensive and admirable series, and it is pleasing to
note the tendency toward international cooperation which is already manifest
in the Pflanzenreich.

Part 37, treating the Araceae? (begun in part 21 of this series), comprises:
(x) @ supplement to the Pothoideae in which a new genus (Epipremnopsis) is
Proposed with a single species, (2) an exhaustive treatise of the Monsteroideae,
which reach their highest development in equatorial Asia and America, and in
which group the authors recognize 12 genera and 190 species, 30 being new
science, and (3) an elaboration of the Calloideae with 4 monotypic genera. A
concise key to the species precedes the larger genera, the species are clearly defined,
and the numerous clean-cut illustrations happily combine general with essen
detail characters.—J. M. GREENMAN.

Flora montana Formosae.’—This work concerns the mountain flora of the
Island of Formosa, embracing the region lying at an elevation of 3000 to 13,0
feet. The total number of species recorded for this region is 392; the sebelong
to 79 families and 266 genera. The author enumerates the various composing
floral elements, such as the arctic, antarctic, alpine, tropical and North America”,
Malayan, Himalayan, southern, central, and northern Chinese, Japanese,
endemic. These upon summation show that ‘the flora is, in general, temperate,
having as many as 320 species of temperate character, or 81 per cent. of the w”
number of elements.” The flora of the island has its strongest affinity #1
central and southern China and Japan, particularly as to the ratio of comp”
nents, but as to their character “the flora of Formosa has as triking affinity |
that of Japan.” After a discussion of the general aspect of the vegetation and ® : 7
— of the Montane zone into four briefly characterized regions, the author

rs with a detailed enumeration of the plants. In this list 69 spect

9 varieties are published as new to science. The descriptive matter is supple
SS Spi

7 ENGLER, A., Das Pflanzenreich. Heft 37 (iv. 23 B). Addita c
Araceas-Pothoideas von A. ENGLER, Araceae-Monsteroideae von A. ENGLER ye
KRavse, Araceae-Colloideae von K. Krause. pp. 1-160. figs 60 (498) as
Leipzig: Wilhelm Englemann. 1908,
oS HAYATA, B.. Flora montana Formosae. Jour. Sci. Coll. Tokyo aie
pls. 1-41, 1908.

1909] CURRENT LITERATURE 159

mented by several text-figures and carefully reproduced full-page illustrations.
The work will serve as an excellent basis for future taxonomic investigation on
the interesting flora of this island—J. M. GREENMAN.

The United States as seen by de Vries.—Professor DE VrrEs has published in
he most attractive f account of his experi on his second American trip.?
The volume is written in popular style, and is amply illustrated with unusually
good half-tones depicting American scenery and universities. ‘There are chapters
on North Carolina with its cypress swamps and insectivorous plants; Arizona
aid the Grand Canyon; southern California with descriptions of San Diego,
the marine vegetation of Santa Catalina, Pomona College, and a camping trip
in the San Bernardino Mountains; the San Francisco earthquake, with special
ilustrations and descriptions of the disaster at Santa Rosa and Stanford Uni-
versity; the University of California, together with accounts of excursions to
Mill Valley, Monterey, Mt. Hamilton, etc.; Great Salt Lake and Salt Lake City;
‘gnculture in the central states, giving descriptions of the Kansas prairies, experi-
ment stations and agricultural colleges in Kansas and Iowa, and maize culture
in Illinois; and the dunes of Lake Michigan. One notices slight mistakes in
the legends of two illustrations, a cut of Drosera being called Dionaea, and a
“ene among the University of Chicago buildings being attributed to the Uni-
‘asity of California. One in perusing this book longs for facility in the Dutch
language, for the book contains the American impressions of one of the ablest
men of our day. Botanists in these days too rarely write such volumes as this,
za because they feel that most of us are now globe-trotters, and able to

own interpreters —H. C. Cow gs.

— and bryophytes of Connecticut.—The algae of the fresh waters of
Het wut bave been described by Professor CoNN and Mrs. Wessrer in a

_ lary report. The descriptions and analytical keys and numerous
ease (from nature) bring these forms within easy reach of collectors and
The bry

“oe i i EVANS
and Mr, Ria - Meeps have been described by Professor

1 Pihoorrotales 3, Sphagnales 31, Andreaeales 2, Bryales 247, @ total
9D : .

1907, i Huco, Naar Californié IT. Haarlem: H. D. Tjeenk Willink & Zoon.
10 Co

the vei AG AND WEBSTER, Lucta W., A preliminary report on the algae :

Nes c: of Connecticut. pp. 78. pls. s. 291). Hartford: State Geol. an
Hist, Survey, Bull. Io. 1908. p 44 (jg 9 )

Ey ANS
Hartford: ha W., ano Nicuots, G. E., The bryophytes of Connecticut. pp- 203-

Geol. and Nat. Hist. Survey, Bull. 11. 1908

160 BOTANICAL GAZETTE [FEBRUARY

of 387. Of these, 68 are peculiar to America, 244 are common to Europe and
Asia, 61 are common to Europe but not to Asia, and 14 are common to Asia
but not to Europe. The bibliography of “Connecticut bryology” contains &
titles.—J. M. C.

British Basidiomycetes.—In 1905 the trustees of the British Museum secured
the descriptions matle by Mr. W. G. Smrru when preparing the series of colored
drawings of British Fungi exhibited in the Department of Botany at South Ken-
sington. Now these descriptions, accompanied by many line drawings ils:
trating generic characters, have been published as a handbook,’? which te
hoped will be useful as an introduction to the field study of the fleshy fungi of
Great Britain. A short introduction (8 pp.) gives a description of the general
features and terminology of the group. The sequence followed is that of Fames
Hymenomycetes Europaei (1874), which is followed also in Great Britam by
‘BERKELEY, Cooke, and SrTEveEnson. Space has been saved by reducing the
descriptions of species to the salient distinctive characters, which must be supple
mented by the generic and sectional characters. The total number of species
Presented is about 2130, distributed among 128 genera and 11 families. y
Hymenomycetes include about 2050 of the species, 106 of the genera, and 6
the families. A full glossary and a complete index conclude the volume, which
should certainly stimulate the interest and activity hoped for.—J. M. C.

Tabulae Botanicae.—This excellent series of botanical charts, —
Gebriider Borntraeger (Berlin), has been appearing during the last two years,
has proved to be of unusual value. They are larger than the ordinary a
so that they can be seen well in a large lecture-room. Even more importan

the group presented, and executed by an artist under his supervision. ea
Baur has directed the illustrations of Myxobacteriaceae and Lichens, JAHN

-onn.) has been asked to act as the American agent, and he will give inform®
tion and transmit orders if desired.—J. M. C.

British Fungi
of British Fungi,
Cas History). The purpose of the models was to exhibit to the public s¥
a series of edible and ro} : A
takes often made from eati : : i of restoring
wee ee ee fating poisonous species. The wor

12 SMITH, Worrtnincton GrorceE, S ° ‘tish Basidiomycetes;

sae » Synopsis of the Britis :
pen So ee © Oe dinwings kad specimens in the Department of Bota?

the
tish Museum, 8yo, . 6 az, - The Trustees of
British Museum, 1908. a 531. pls. 5. figs. I45. London:

1909] CURRENT LITERATURE 161

fragile models was committed to Mr. WorTHINGTON G. SmiTH, and in connec-
tion with this the Guide was published. There has now appeared?3 a second
edition which has been carefully revised, and a glossary has been added.—J. M. C.

Natiirlichen Pflanzenfamilien.—Parts 231, 232, and 233 continue the pres-
entation of mosses by V. F. BRorHERUS, completing Thuidieae, and presenting -
Hypnaceae, Leucomiaceae, Sematophyllaceae, Rhegmatodontaceae, and Brachy-
theciaceae —J. M. C.

meet G&S FOR STUDENTS

Current taxonomic literature.—A. D. E. ELMER (Leaflets of Philippine Botany
1:272-359. 1908) describes 100 new species of flowering plants, belonging to
Various genera, and (idem 2:375-384) 9 new species of Lauraceae all indigenous
to the Philippine Islands—J. D. HooKER (Hook. Ic. Pl. pls. 2851-2875. 1908)
describes and illustrates 24 new species and one new variety of the genus Impatiens
from China. The types are deposited either in the Paris, Le Mans, or Kew
Herbarium.—v. L. Komarov (Acta Hort. Petrop. 29:1-176. 1908), under the
title of Prolegomena ad floras Chinae nec non M ongoliae, makes a valuable con-
tRbution to the literature concerning the flora of China; it includes, moreover, a
— revision of Clematoclethra Max., Codonopsis Wall., Epimedium and
Nitraria L., in which genera g species and one variety are proposed as new to
ee D. House (Muhlenbergia 4:49-56. 1908) gives a Synopsis of

California Species of Convolvulus. The author recognizes 26 species, two of
are new.—W. P. Hiern (Journ. Bot. 46:273-278. 1908) records the
cccurrence of a Sagittaria in the river Exe, near Exeter, England. The plant is
described a8 a new variety of a North American species. SPENCER LE M. Moore
(idem 290-208) describes 12 species of African plants as new to science, and pro-
ae new genus (Grossweilera) of Compositae; the same author (idem 305-313)
of the Ascle I new species of African plants and a new genus (Swvynnertonia)
Desi adaceae, and also a new genus (Eylesia) of the Scrophulariaceae.—

(Bull. Hb.

ha egy Specialists (idem 62 5-640), describes 32 new species of African
lip, Toone 2 NeW genus (Pseudotragia) of the Euphorbiaceae.—A. BRAND

A P- Journ. Sci, 2- : ‘ ae of
the Philip i - 3:I-10. 1908) gives a synopsis of the Symploca
€ Islands, in which 16 species are recognized, 6 of which, in addi-

a he TH, WORTHINGTON GrorcE, Guide to SowerBy’s models of British Fungi.

u : wy ised. pp. iv+85. figs. gt. London: The Trustees of the British

162 BOTANICAL GAZETTE [FEBRUARY

tion to 3 varieties, are described as new—T. Naxatr (Journ. Coll. Sci. Imp. Univ.
Tokyo 23:1-28. 1908) presents a careful synopsis of the Polygonaceae of Corea
and describes one new species and a new variety in the genus Polygonum.—
N. L. Britton (Bull. Torr. Bot. Club 35:337-345. 1908), under the title of Siud-
ies of West Indian plants, I, describes 5 species as new to science.—Ic. Ursay,
in collaboration with different specialists (Engl. Bot. Jahrb. 42:49-176. 1908),
under the title Plantae novae andinae imprimis W eberbauerianae, IV, has published
173 new species and 25 new varieties of South American plants, and also 4 new
genera: Fiebrigiella (Leguminosae), Centradeniastrum (Melastomaceae), Guran-
iopsis (Cucurbitaceae), and Huthia (Polemoniaceae).—HEnR1 LEcoMTE (Jour.
Botanique II. 21:101-109. 1908) describes 7 new species of Eriocaulon from
Indo-China.—AtFreD CHasert (Bull. Soc. Bot. France IV. 8: 305-310. pls.
12, 13. 1908) proposes 2 new varieties of Campanula rhomboidalis L. from France.
—G. Bonatt (idem 310-314) describes 4 new species of Pedicularis from China—
F. GaGNEPAIN (idem 322-325) describes 2 new species of the Capparidaceae
from China.—E. A. Finer (idem 333-343) has published 11. new species and1
variety of orchidaceous plants from South America and from different parts of the
Old World.—H. Lévernié (idem 407-409) recognizes 5 species of Mtcuna indi-
genous to China, 2 of which are described as new.—S. T. Dunn (Jour. Linn.

Henry Prrtier (Contrib. U.S. Nat. Herb. 12:1 59-169. 1908), following a general
discussion of the genus Sapium and an analytical key to the Mexican and Central
American species, describes and illustrates the 9 species recognized for this regio®;
of these 6 are new to science, and 4 are published in joint authorship with the si
Prof. Kart ScHUMANN.—Ep. Parra (Oesterr. Bot. Zeitschr. 58: 389-392 1908)
describes 3 new species of Cyperaceae from Mexico and Colombia.—CHARS
Brooxs (Bull. Torr. Bot. Club 35: 423-456. 1908) gives an account of The fri
Spot of apples and records a new species of the genus Cylindrosporium—P. 4
RyDBERG (idem 457-465) recognizes 6 species of Philotria for this country,
being described as new.—EvcEne P. BIcKNELI, (idem 471-498), under the Me
Ferns and flowering plants of Nantucket, III, has published 2 new species and
proposes 7 new combinations chiefly in the genus Carex.—F RANK D. KERN ar
499-511) in Studies in the genus Gymnosporangium publishes 3 new species ant
makes 3 new combinations,—F. Petrak (Fedde, Rep. Nov. Sp. 53329335
1908) describes 9 new hybrids and x new variety in the genus Cirsium fe
southern Europe.—E. Hacker (idem 333-335) records a new species of se!
from Australia—F. KrAnziern (idem 369, 370) has described a new species
Calceolaria from Bolivia.—k. RosENSTOCK (idem 370-376) has published o_

1909] CURRENT LITERATURE 163

species and 5 varieties of ferns from New Guinea.—J. BORNMULLER (idem 376,
377) records a new species of Reaumuria from Persia.—D. GrirriTus (Rep. Mo.
Bot. Gard. 19:259-272. pls. 21-28. 1908), under the title I/lustrated studies im
the genus Opuntia—I, has described 15 new species of Opuntia from Mexico
and the southwest—W. GucterR (Ann. Mus. Hung. 6:15-297. pl. rz. 1908),
under the title of Die Centaureen des Ungarischen National-Museums, records
in detail the species and subordinate categories of the genus Centaurea represented
in the Hungarian National Museum; the work is preliminary to a monograph of
this genus—F. SrepHANt (Bull. Herb. Boiss. II. 8:837-866. 1908) has described
45 new species of Hepaticae, of which several are American.—B. DE LESDAIN
(Bull. Soc. Bot. France IV. 8:420-424. 1908), under Notes lichénologiques,
describes 3 species and 2 varieties as new to science.—H. L&vEILié (idem 424-
427) enumerates 11 species of the genus Pueraria for China, of which 5 are pub-
lished in joint authorship with Vaut.—P. Dop (idem 427-430) describes 3new
species of the Malpighiaceae from Indo-China.—F. GAGNEPAIN (idem 430-436)

the
Zeal ee (idem 419-421) records a new species of the genus Bagnisia from New

n €conomic studies, has published 7 new species and 2 new varieties of

Ber Denna Plants of Africa, chiefly from the Congo region—B. ScHROEDER

and ne
aay varieties of Australian flowering plants.—G. E. MATTEI (Boll. R.
- Giard. Col. Palermo 7:85-112. 1908), under the title Contribuztont alla
Cs della Somalia italiana, has described 9 new species of flowering plants.—

69 new species of Crataegus from Missouri, and in another place
- State Mus, 122: 26-130. 1908) 83 from New York.—C. H. Peck
131-160) has published 8 new species of fungi from New York.—
(Philip. Journ. Sci, 3:269-276. 1908) has published 12 new species

164 BOTANICAL GAZETTE [FEBRUARY

and 2 new varieties of Philippine ferns.—E. B. CoprLAnp (idem 277-284) de-
scribes 8 new species and 3 new varieties of ferns from China; and the same author
(idem 285-300) presents A revision of the Philippine species of Athyrium in which
46 species are recognized, 5 species and 1 variety being described as new to science.
—E. D. MERRILL (idem 307-315), under the title of Philippine Freycineti
24 species of this genus from the Philippine Islands, 8 being described as new;
and the same author (idem 317-338) has published 6 new species of oaks and 6new
species and 1 new variety of the genus Radermachera from the Philippine Islands.
—R. Wacner (Oesterr. Bot. Zeits. 58: 435-439. 1908) has described a new
species of Tropaeolum from Columbia.—F. SENNEN (Bull. Acad. Intern. Geogr.
Bot. II. 17: 449-480. 1908), under the title of Plantes d’ Espagne, has published in
joint authorship with C. Pav 6 new species and several new varieties of flowering
plants from Spain—R. Fries (K. Sy. Vet. Akad. Handl. 42:1-67. pls. 1-7.
1908) has published 23 new species and 14 new varieties of Malvales, chiefly from
South America; the same author (idem 43:1-114. pls. 1-10. 1908)
37 species for the genus Wissadula, of which 11 species and 4 varieties are described
as new, and a new genus (Pseudoabutilon) of the Malvaceae is proposed, to which
are referred 9 species, 3 being new to science.—K. Jouansson (Arkiv fér Botanik
7*no. 12, pp. 48. pls. 1-5. 1908), under the title Hieracia vulgata Fr. fran Torne
Lappmark, enumerates 35 Species, 20 of which and 3 varieties are described as
new.—J. M. GREENMAN.

Effect of light on germination of seeds.—HEInRICHER" has recently added
two more papers to his series on the effect of light on germination, and KrnzEt's
publishes a second paper (preliminary statement) on his extensive researches 0
this subject. Data enough are now at hand to get at some general principles. is
respect to the effect of white light upon their germination, seeds may be divid!
into four groups: those requiring light for germination (Rhododendron javanicum,
R. hirsutum, R. ferrugineum, Drosera capensis, etc.); those germinating more
quickly and fully in light (Veronica peregrina, Allium suaveolens, etc.); those
germinating equally well in light and darkness yrmecodia echinata, etc.); and
those retarded in germination by light (Phacelia tanacetifolia, Pedicularis Scepir™™
Carolinium). It is agreed by both authors that the favorable effect of light isnot

Ha. records
7

oe HEINRICHER, E., (1) Beeinfliissung der Samenkeimung durch das as
Wiesner-Festschrift, Wien. 1908. (2) Die Samenkeimung und das Licht. :
Deutsch. Bot. Gesells. 26a: 298-301. 1908.
Ss Kinze, W., Die Wirkung des Lichtes auf die Keimung. Ber. Deutsch. Bot
Gesells. 26a: TOS-I15. figs. 4. 1908,

1909] CURRENT LITERATURE 165

to support this conclusion. I see no grounds, however, for concluding that the
effect is upon the digestive enzymes rather than upon some other mechanism of
the protoplasm.

Kinzer’s attention has been largely centered upon the effects of rays of different
refrangibility. HEINRICHER*® early found that the red end of the spectrum was
most effective in Veronica peregrina. KiNzEL’s results indicate that this is
generally the case. Among different species, however, there is a great variation in
the relative effectiveness of various rays. Green is by far the most effective with
Nicotiana, while with Veronica yellow gives the greatest stimulation. NZEL
finds the blue rays least effective; in fact they often cause marked retardation. In
many cases of seeds favored in germination by light, blue gave a much slower and
lower percentage of germination than darkness. K1INzEL comments upon the
general retarding effect of blue light, while HEINRICHER later points out that in
the seeds of Phacelia tanacetifolia which are retarded in germination by white

t, blue markedly stimulates germination.

Many of the “‘light-loving” seeds demand a considerable period of rest after
harvest, during which they become thoroughly dried out. In Veronica bellidioides
three and one-half months was the most effective period. In the short-lived seeds
. Drosera fifteen hours of drying in the laboratory best effected their “‘after-ripen-
mg. HEINRICHER says, “On the whole the experiments indicate that the
results in the germination of such seeds as are helped by light depends upon the
age of the seed, upon the quickness of drying after harvest, further, also, upon
Whether this takes place in light or darkness, and, if in the first way, whether in
one layer or several, F inally, even the moisture content of the air during storage
must be considered as a factor. is evident that tl liti traordinarily
complex and that conformity of results is to be expected only under the considera-
“on of all these factors,”
light ig aa con uapuRgers in behavior as ne sea

nnected with thei i ionshi an WI

of cg aise heir phyletic relationships
ota @ reviewer that in cases where evident coats appear, the investiga-
son of oxy ork with coat-free seeds to make sure that the coats by partial exclu-
light bas — Salts, or even water are not hindering germination. In such cases
tion. It is — of compensating some other limiting condition of germina-
problem will evident, from the variable results, that the real solution of the
i aroused come from learning the particular dormant process in each case that
made in o y light. This ought to be possible in view of the great advance being

ur knowledge of the catalytic nature of protoplasmic activity, but it will

Upon — attack on the problem from other points than the mere effects of light
living seeds Wat, Crocker
Dera ee

16 Her
Gesell, p>. CHER, E., Ein Fall beschleunigendes Wirkung. Ber. Deutsch. Bot.
7:308-311. 1899,

166 BOTANICAL GAZETTE [FEBRUARY

Observations on Welwitschia.—Prarson has communicated further studies

on this peculiar genus to the Royal Society, London, of which the following is an
abstract.
Macrospores and embryo sacs are frequently present in the pith region of
the female cone-axis. This confirms the view, already adopted by most authors,
that the ovule of Welwitschia is cauline. Sporogenous cells have not been
found in a similar position in the male cone.

It is suggested that the female cone and the male flower are derived by redue-
tion and specialization from an amphisporangiate strobilus of a type similar to
that of Bennettites.

At the end of the free nuclear division the embryo sac contains about 1024
nuclei which are equivalent in all visible characters, Cleavage of the cytoplasm
occurs, resulting in the septation of the whole sac into compartments. Those
near the micropylar end contain few nuclei which are functionally sexual; most
of those of the lower three-fourths inclose many potentially sexual nuclei. The
former send out embryo-sac tubes into the nucellar cone and into them pass the
cytoplasm and free nuclei ; all the nuclei in each of the latter fuse so that each com-
partment becomes a uninucleate cell. The compartments containing the fusion
nuclei form the primary endosperm, whose later growth is distributed over two
periods, one before and the second after fertilization. The endosperm of Gnetum
is probably formed in the same way. In respect of the morphological character
of the endosperm, Gnetum and Welwitschia are widely separated from Ephedra,
in which the endosperm is a prothallus of the normal gymnosperm type. It is sug-
gested that the endosperm of the primitive angiosperms was homologous with that
of Welwitschia,

The embryo-sac tubes meet the pollen tubes in the lower half of the nucellar
cone. Fertilization occurs within the generative cell, which enlarges after leaving
the pollen grain and its nucleus divides. The daughter nuclei are functional
gametes.

Several oospores are commonly formed in each nucellus. The cytoplasm a
the oospore is mainly, if not entirely, provided by the generative cell. A resting
nucleus is formed. The oospore elongates toward the top of the endosperm.
The first nuclear division within it is followed by the formation of a centripetally
cope wall which separates the upper “primary suspensor” from a lower
terminal cell. From the latter are developed: (a) 24 cells which, surrounding -
/ower part of the primary suspensor, form with it “the secondary suspens0r;
(0) a terminal group inclosing a presumed embryonic plate of eight cells. The
later stages of embryo development have not been seen; they possibly occur, 4 in
Gnetum, after the seed is detached from the plant. '

It is suggested that (r) the Gnetum-Welwitschia alliance has its origin ™
the same stock as the angiosperms, but separated from the angiosperm line
before the carpel became the pollen-receiver; (2) Welwitschia is the most i
ized living representative of the race to which it belongs.

1909] CURRENT LITERATURE 167

Mucilage ducts in Pipereae.—In considering the genera of the Piperaceae,
two tribes are recognized by VAN TIEGHEM,*7 namely Pipereae and Peperomieae;
while the Saururaceae are kept as a distinct family, as suggested a hundred years
ago by L. Ci. Ricnarp. By Casmurr DECA theS were replaced
in Piperaceae-as‘a tribe; and then separated by ENGLER. In speaking of Piper-
aceae, therefore, Van TIEGHEM does not include Saururus and itsallies. From an
anatomical point of view the Piperaceae have long attracted much attention,
especially on account of peculiarities in stem structure, which at the same time are
characteristic of the respective tribes. For instance, in the Pipereae the stem
exhibits a normal monostelic structure, with the broad stele surrounded by a
well-differentiated endodermis, and possesses at least two concentric bands of
mestome bundles. In the Peperomieae, on the other hand, the stem structure
is of the schizostelic type, the numerous meristeles being scattered, not arranged
in bands, and each being provided with a special endodermis. Common to
both tribes, however, is the presence of roundish oil cells with the cell wall sub-
rized or at times lignified; these oil cells are widely distributed through stem
and leaf. In certain Pipereae still another secreting system occurs, which is now
for the first time described. It consists of a single duct or several broad ducts
containing mucilage and extending through the full length of stem and leaf; these
ducts are lysigenous, since they arise from the destruction of a row of secreting

They occur in the pith of the stem, mostly a very broad one in the center
and several harrower ones in a band around this and alternating with the inner-
most mestome strands, They contain a colorless mucilage, and are surrounded

— f secretory ducts belonging to the stem stele is readily followed
in. a me internodes, but disappears completely in the nodes. In the leaf
hadro: = eccur in the petiole, in the parenchyma located on the ventral (the
be en face of the arch formed by the mestome strands; thence they may
the auth, a the midrib of the leaf blade, from the base to apex. Although
ee fxamined various representatives of the Peperomieae, he failed to
in P pe ducts in any of them. Among the Pipereae they occur in Piper (as
“mgrum, P. Cubeba, P. macrophyllum, etc.), in Chavica Blumei, C. sphaero-
ai and some species of Heckeria; while they are not developed in Macro-
? Nematanthera, and Zippelia—Tuxo. Horm.
wo in aquatic plants.—FRancors'® has offered a very interesting con-
nal as eo " knowledge of aquatic plants with notes on their structure, cu
drawn ay internal, and on their seedlings, the text containing many well-
Special attention is given to the vegetative reproduction of such

17 V , :

IX. TIECHEM, Pu., Sur les canaux & mucilage des Piperées. Ann. Sci. Nat.
Bot, » 72117, 1908,

18 Fp

IX, 7:25. 1908, » L., Recherches sur les plantes aquatiques. Ann, Sci. Nat. Bot.

168 BOTANICAL GAZETTE [FEBRUARY

species as occur on river banks, the stolons of which grow in the water, creeping
over the muddy bottom. Among these are Mentha aquatica, Lysimachia vulgaris,
Lycopus europaeus, Stachys palustris, partly also Potentilla reptans, Ranun-
culus repens, and Cynodon Dactylon. In these the vegetative reproduction is
amply secured by the ability of the fragments of the rhizomes and stolons to
root very easily, and at the same time the water currents help to disperse such
fragments over wide areas. The seedling stage of Butomus, various species of
Alisma, Sagittaria, Najas, and Potamogeton is described. The slow growth of
the primary root is characteristic, while the hypocotyl attains its final length
in a very short time, and before the root actually commences its increase in
length. The primary root stele in Butomus and in Alismaceae consists of a
single central vessel and of two strands of leptome diametrically opposite each
other. In the Najadaceae, on the other hand, several vessels are developed, the
largest of which is usually located in or near the center, and there are also several
strands of leptome, corresponding in number with the rays of hadrome. The
hypocotyl exhibits a bilateral structure in Butomus and Alisma and no pericycle
was observed inside the endodermis. In the Najadaceae the bilateral structure
is much less pronounced, and no stomata were observed in the epidermis of the
hypocotyl of any of these plants. A very simple structure characterizes the
cotyledon; the chlorenchyma is homogeneous and contains only one vein neat
the ventral face. The seedling stages are -very carefully described and figured,
adding several interesting points to the knowledge of the structure of aquatic
plants.—Turo. Hota.

Organic correlations.—East"? attempts a classification of correlations with
especial consideration of plant data. This is a little-known field at the present
time, but one of great promise for the future. The writer of course realizes that
this tentative classification awaits the accumulation of further data to place it 0
any satisfactory basis. Correlations are considered as “somatic” and “gametic.”
Under somatic correlations are classed: (1) correlated reactions to environment;
here are placed the experimental results of MacDoucat in Raimannia and
TowER in Leptinotarsa, although the indications are that these changes 3
germinal and not somatic; (2) growth correlations between (a) non-homologo™s

interdependent and exclusive development; here is cited the case of O

lata, in which the broad-leaved character is associated with the ‘inability to P™”
duce healthy pollen;” but this association is not constant, for P. 2
England having the O. lata characters have been successfully self-pollinated;

: + pp. 12
"9 East, Epwarp M., Organic correlations. Amer. Breeders’ Assoc. 4:PP: *

Sint . and
and relationships of the Oenotheras. Carnegie Inst. Publ. 81:pp- 92: ie
unpublished results of the reviewer.

* MacDoveat, D. T., Van, A. M., AnD SHutt, G. H., Mutations, variations
1907

1909] CURRENT LITERATURE 169

_(5) heterozygotes; in the numerous cases where heterozygotes differ from either
parent, “the ability to transmit certain characters is correlated with other apparent
characters.” Under gametic correlations are placed the phenomena of partial and
complete “coupling,” so called, developed chiefly by BATEsoN.—R. R. GATEs.

Tyloses in ferns.—It has been noted by various writers that in the stems and
petioles of ferns the protoxylem groups suffer disintegration, and into the cavities
so formed the wood-parenchyma grows, forming the “‘cavity-parenchyma” of
Russow. Proliferations from these cells frequently fill the cavities, and present
the appearance of tyloses. These growths have recently been studied in detail by
two independent workers, KrrscH?* and Miss McNicuot.??_ Both writers show
that the phenomenon is widespread, being found in nearly every family of the
true ferns, as well as in Marsilia and the Ophioglossaceae. In both papers the -
cells in question are carefully described and their origin as stated above is proven.
Kirsc has studied Pteris aquilina in most detail, and finds cavity-parenchyma
in the stipe and in all regions of the rhizome, where it occurs in the outer system
of bundles which he erroneously regards as cortical (p. 388). He offers the fol-
lowing as a theory of the cause of these growths: the cavity formed by disintegra-
tton of the protoxylem at first functions as a water duct; later the metaxylem
(Secondary xylem according to Krrsc#) makes its appearance and performs the
duty of water carrier. Hence the pressure in the cavity is reduced, and as a con-
Sequence tyloses grow into it—M. A. CHRYSLER.

Composition of a field of maize.—A brief paper by SHULL?S calls attention
to the view, already expressed by DEVries and others, that a field of corn, like
<. and other grains, is made up of a number of elementary species or biotypes.
rca the fact that inbreeding in corn results in deterioration, and points

that the old hypothesis that the deleterious effects of inbreeding result from
oa rama of disadvantageous individual variations to form an organism
ties se inharmonious or unbalanced constitution, is untenable, in view of the

“s of cleistogamy, self-pollination, and parthenogenesis in plants which have
tesa been successful in the struggle for existence. A cornfield is conceived to

“Series of hybrids between elementary species, and on the basis of the common
dither that hybrids between nearly related forms are more vigorous than
.__ Parent, he believes that over-selection, which eliminates down to a single
Pi Tesults in deterioration, not intrinsically from inbreeding, but because the

“igor which comes from the crossing of biotypes has been eliminated. The
a

; ;

sis esi Suwon, On the development and function of certain structures In the

. saa of Pteris aquilina and other Pteridophytes. Trans. —
= ~ 14°353-412. figs. 27+ 27. 1907. Anna

MeNicxor, M., On cavity parenchyma and tyloses in ferns. aes

22:401-413. pl. 25. 1908.
23
SHULL, Gro, H., The composition of a field of maize. Amer. nes
*4°PP. 6. 1908,

170 BOTANICAL GAZETTE [FEBRUARY

ideal of the corn-breeder should then be continuous hybridization between
biotypes, rather than the isolation of pure strains.—R. R. Gares.

Isolation and mutation.—While the final adjudication of the claims of the
various theories of evolution must be made on an experimental basis, such data
must be in harmony with the facts of plant and animal distribution, as is pointed
out in a suggestive paper by Leavirt.*4 It is of much interest to observe that
zoologists, as a rule, have been less inclined to believe in mutation than have
botanists. This is in part due, Leavitt thinks, to a less perfect grasp of the
theory by some of the zoologists, but in part due also to the fact that most students
of animal distribution believe that isolation of closely related species is a most
important principle in evolution. The author shows that there are innumerable

without geographic isolation, although cases suggesting the latter are not wanting.
Therefore, it is concluded, many facts of plant distribution favor the mutation
theory, though they do not show that this is the only valid theory of evolution —
H. C. Cowrgs.

Osmotic Properties of root hairs.—Hy.125 has investigated the osmotic prop-
erties of the root hairs of Glyceria maritima, Suaeda maritima, and Salicornis
herbacea, which grow in a salt marsh subject to great changes in the osmotic pres-
Sure of its soil water, due to Periodic flooding by the tides and to occasional drench-
ing rains. He finds that the hairs show marked and rather rapid variation in
Osmotic pressure Corresponding in variation to the osmotic pressure of the soil
water. This variation is not due to the entrance of the abundant chlorids of the

dify their osmotic properties readily in response to and
Tapidly varying external osmotic pressures.— WILL

Statolith theory ‘ —Bupr R? 6

comes to the support of the statolith theory with a
Set of well-chosen and critical
rena

experiments that seem to justify his conclusions
_ 4 Leavirr, R. S., The 8eographic distribution of nearly related species.
Nat. 41 *207-240. 1907,
. * Hu. bservations on the Osmotic properties of the root hairs of cer-
n salt marsh plants. New Phytologist 7:133-142. 1908. i
estschrift

BUDER, JOHANNES, Unte i
: gs , Tsuchungen zur Statolithenhypothese.
zur Feier des 25-jahrigen Bestehens der Deutsch. Bot. Gesells. Ber. 26:162-193- 19%

1999] CURRENT LITERATURE 171

which are briefly as follows: Contrary to Firrrnc’s conclusion, in a combination
of the rest position with various angles, the statolith starch takes the position that
would be expected by the statolith theory. Centrifugal acceleration causes the
movement of the starch that the hypothesis assumes, as shown by accelerations
from 0.13 gto gg. In these various accelerations the time of the movement of
the starch to the side of the cells coincides with the presentation time as deter-
mined by Bacu. In the intermittent exposures of opposite sides when these
exposures are of short duration the starch moves to the side of the cell of the
most effective exposure only after the process is long continued, corresponding to
the slow reaction in these cases. However well this paper may answer a number
of the arguments against the statolith theory, there are yet a number unanswered
and this whole matter of geotropic reaction seems too complex to be entirely
explained in such a simple way.—WM. CROCKER.

Ray-tracheids in Cunninghamia.—The complex structure of the medullary
rays of living Abietineae, consisting of parenchyma cells, ray-tracheids, and an
elaborate system of ligneous resin-canals, has been used as one of the evidences
of a highly specialized and relatively modern group. JEFFREY?? has studied the
harginal ray-tracheids that occur occasionally in Cunninghamia and has found
a Ly due to wounding, being most numerous in the region of the injured
annual rings opposits the wound-callus. They resemble in general those de-
ne * a genera of the Taxodineae and Cupressineae, and JEFFREY
the why “ this is additional evidence that these two tribes have been derived from

is vi ca the ray-tracheids being “vestigial or reversionary.” He emphasizes
aie if eiing attention to the fact that there is no evidence that the Taxo-
sions ia Cupressineae existed before the end of the Cretaceous. Such conclu-
oe the fact that apparent simplicity of structure may not indicate

Sater antiquity than greater complexity of structure.—J. M. C.
Rs 3 ystem of Ranales.—WorspEi”* maintains that the primitive angio-
ina scat large leaves, and that as a result the vascular bundles were disposed
there is in all Saad as is seen in the monocotyledons. He considers that
Split and the i a single terminal cotyledon in the embryo, but that it may
inal organs pe terrier through 180°. Like the cotyledon, all the leaves are
“ondition ocaaagee hence dominate the stem (‘‘grandifoliate”’). From this
n derived the one in which the stem is dominant and the leaves

Fa aeaaas although he claims to adopt the “recapitulation theory.
ete Paper he outlines the results of an extensive study of the leaves in

th | * . .
tnnals of Botan Epwarp C., Traumatic ray-tracheids in Cunninghamia sinensts.
Wo any 22°593-602. pl. 27, 1908.
Pa ie study of the vascular system in certain orders of the
of Botany 22:651-682. pls. 32, 33. 1908.

92° BOTANICAL GAZETTE [FEBRUARY

certain representatives of Ranales, and shows how the scattered bundles of a
petiole may be converted into a ring, and the bundles of one side of the ring
approximate the opposite side so as to produce a single arc.—M. A. CHRYSLER.

Solution of mitoses.—Experiments of Oxrs?9 with various root tips, embryo
sacs, and pollen mother cells show that cells capable of growth and division con-
tain a chromatin-dissolving enzyme (nuclease), which dissolves chromatin when
toluol, chloroform, carbolic acid, etc., are added. Metaphases, anaphases, and
telophase are most quickly attacked, the prophases being less susceptible, and the
resting nucleus still more resistant. In autolyzed objects the spindle is dissolved,
but the nucleolus and nucl f resting nuclei remain unaffected. The
effect of temperature, neutral salts, free acids, and alkalies was observed in various
objects. The writer believes that the diminution of chromatin in the telophase,
_ observed by SrRAsBuRGER and others, may be due to nuclease. If nuclease func-
tions in the normal, living plant, thus causing irregular fluctuations in the chro-
matin, the question arises whether chromatin is the exclusive bearer of hereditary
qualities—CHartes J. CHAMBERLAIN.

the relation of the mite to infection has not been completely worked out in
case.—F. L. Stevens

*° OES, ADOLF, Ueber die Autolysis der Mitosen. “Bot. Zeit. 66 389-120

1908

3° CHRYSLER, M. A., Tyloses in tracheids of Conifers. New Phytol. 7:7
pl. 5. 1908.

s Stewart, F. C., anp Hopcxiss, H. E., Tech. Bull. 7, N. ¥. Agric :
Sta. Oct. 19, r908.

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THE

March 1909

Editors: JOHN M. COULTER and CHARLES R. BARNES

CONTENTS
2 | Mitosis in Fucus Shigéo Yamanouchi

_ The Reduction Division in the Microsporocytes of

Agave virginica John H. Schaffner
Spermatogenesis in Dioon edule Charles J. Chamberlain

: Br iefer Articles :
Ze The Mounting of Algae J. A. Nieuwland
Paul Hennings J. Perkins
Pure Cultures of Fungi Johanna Westerdijk

: Current Literature

- The University of Chicago Press
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The Botanical Gazette
Montbly gournal Embracing all Departments of Botanical Science

Joun M. CouLTER and CHARLES R. BARNkEs, with the vii a of other members of the
botanical staff of the University of Chica

Issued March 23, 1909

CONTENTS FOR MARCH 1909 ee 4 ok,

DUCTION DIVISION IN THE MICROSPOROCYTES OF AGAVE VIRGINICA
(WITH PLATES xII-x1Vv). John H. Schaffne 198

TOGENESIS IN D/OON EDULE. CONTRIBUTIONS FROM THE HULL BOTANICAL
Laboratory 125 (WITH PLATES XV-XVIII AND THREE FIGURES). Charles J. Chamberlain 215

ER ARTICLES

MOUNTING of ALGAE. /. 4. Nieuwland - - ee A A ee
Pau. HENNINGs (WITH pereurry de Perkins 6 4 8 Pie ee ee
BE SUEtUERS OF FUNGI: Johonna Westerdjt - - 9 = = + +> + + 2M

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E XLVII NUMBER 3

MARCH 1909

MITOSIS IN FUCUS
ONS FROM THE HULL BOTANICAL LABORATORY 124
SHIGEO YAMANOUCHI
(WITH PLATES VIII-XI) -
es. INTRODUCTION
___ The first cytological study of Fucus is that of FARMER and
: Wataus (8), published about ten years ago as a preliminary note.
The next year SrraspurcER’s paper (24) on Kerntheilung und Be-
_ frchtung bet Fucus appeared. In 1898 the final paper of FARMER
and Wittaus (9) was published.
. FARMER and Wittrams’ material, Fucus platycarpus, F. serratus,
: ‘ ad F. vesiculosus, was collected on the coast of England. Their
3 noma deals with egg-formation in the oogonium, particular atten-
_ lion being devoted to the third division; also with fertilization and
‘arly segmentation divisions. Ascophyllum nodosum and Pelvetia
vom wlata were used in a supplementary way. STRASBURGER’S
i. m “th based chiefly on material from Heligoland, Germany,
hws ciel in detail the third division in the oogonium of
. Paryearpus and the fertilization processes in FP. serratus and
vesiculosus

T a
es 0 the brilliant results of these authors we owe most of our present
Wledge

W of the cytology of these forms. Since the work of FARMER,
ao and STRASBURGER, cytological conditions have been studied
28) a “Sea such as Dictyota (WitraMs 27), Nemalion (WOLFE
.. olysiphonia (YAMANOUCHI 29). The morphology of chro-
he. » M connection with the theoretical problem of alternation
: Problem, F » specially in the algae, is becoming a more important
; °r the solution of such a problem, there must be a
173

174 BOTANICAL GAZETTE [maRcH

thorough study of the life-cycle of chromosomes; unfortunately not
much of such work has been done in algae. The work of FARmer,
Wirttams, and STRASBURGER dealt only slightly with the first two
mitoses in the oogonium, where it was inferred, but not actually
observed, that the reduction of chromosomes takes place. Mitoses
in the antheridium, from the first division to the development of
mature sperms, were not studied. In the present investigation,
special attention is paid to the behavior of chromosomes in the first
and second mitoses in the oogonium, and to mitoses in the antheridium.

The results here presented are based upon a study of Fucus
vesiculosus L. Material was collected and fixed at Woods Hole,
Mass., during the latter part of March and early April, 1908. As
fixing reagents, Flemming’s weak solution containing osmic acid,
with various modifications, proved to be most satisfactory. There
are several points of interest and importance in regard to the relation
between the frequency of mitotic figures and environmental con-
ditions, both in the oogonium and antheridium and in the young
thallus. In general, the plants collected one or two hours after being
covered by the tide were full of figures. Material was imbedded in
paraffin with a melting point a little less than 52°C. Sections were
cut 3, sometimes 5 thick. Flemming’s saffranin and Heiden-
hain’s iron alum hematoxylin, with or without counterstains, We
mostly used in the slides from which the accompanying illustrations
were made.

To Professors Joun M. Coutrer and CHARLES J. CHAMBERIAIN
I am indebted for their kind suggestions and criticism during the
progress of this work.

MITOSES IN VEGETATIVE CELLS OF THE THALLUS

Any young part of the thallus, when well fixed, showed figu"®
available for study, especially in the apical region, in the adventitious
outgrowths which are not infrequent, and in the early stag®
conceptacles or cryptosomata. The nuclei in the thallus, except
in cases of young sporelings, are generally very small an ee
to study; but to make certain as to whether the number of sine
somes remains constant in the thallus grown under normal condition»
a thorough study was made of the typical mitoses at various stage

1909] | YAMANOUCHI—MITOSIS IN FUCUS 175

development. The following brief account of the essentials is
illustrated by figures from the apical portions of male and female
plants. Since the mitoses in male and female plants are precisely
alike, the following account may be understood as applicable in
either case. :

The nuclei in the growing apex of the plant are somewhat larger
than those in older regions of the thallus, sometimes filling almost
all of the lower half of a narrow elongated palisade cell, such as
constitutes the surface layer of the thallus. The cells are filled with
an abundance of contents, such as plastids, physodes, and other
granular substances of undetermined nature. In regions where such
contents are scarce, the cytoplasm shows a very fine alveolar structure,
which gradually becomes granular, even, and homogeneous toward
the nuclear membrane.

In early prophase the chromatin network of the resting nucleus
changes to a structure in which numerous chromatin knots become
more and more pronounced, until they become transformed into
well-developed chromosomes (figs. ra, 1b, 7). The chromosomes,
pisos at first appear very irregular in size and shape, now become
que similar in form, slightly bent, and in this condition they are
arranged in an equatorial plate (figs. 2, 8).

During the chromosome development within the nucleus, the
: has a tendency to become transformed into kinoplasm at
the two poles. As a rule in these kinoplasmic accumulations, centro-
Somes appear first in the late prophase, when the chromosomes are
‘manged in the equator (fig. 2). The nuclear membrane persists
ri ay peo late metaphase, being especially well marked toward
_ “uatorial region. In the polar region where the centrosomes
ded emorane will perhaps be very faint, so as to allow the
astral ied spindle fibers to intrude into the nuclear cavity while the
A ae are formed outside. Owing to the minuteness of the
tthe ‘i 1S rather difficult to make any positive statement in regard
treated a. of the centrosome and spindle. These features can be
Saale satisfactorily in connection with the divisions in the
: = the early segmentations of the fertilized egg. :
esi — T of chromosomes in the early prophase (jigs. eae )

an 60, but 64 chromosomes can be counted with certainty

ae BOTANICAL GAZETTE [ace

in the late prophase (jigs. 3, 7) and anaphase (jigs. 6, 12a, 12) in
the polar view of the equatorial plate. In metaphase, the chromo-
somes at the equator split and separate, and in anaphase two sets
of daughter chromosomes proceed toward the poles (jigs. 4, 10).
The sets of daughter chromosomes when they reach the poles no longer
remain in one plane, but become aggregated into more or less irregu-
lar spherical masses which then become vacuolate. Probably through
the interaction of the nuclear sap, derived perhaps from the vacuola-
tion, and of the surrounding cytoplasm, a new membrane is organized,
thus completing the process of typical mitosis.

The centrosomes that were always observed staining black at
the poles lose gradually their sharp identity, until they can no longer
be differentiated by stains.

MITOSES IN ANTHERIDIA

In Fucus antheridia develop from wall cells of the conceptacle.
A wall cell of the conceptacle puts forth a papilla which is cut of
by a transverse wall (fig. 13). The papilli” grows for a time and
divides, forming the antheridium and its stalk (fig. 14). A stalk
cell may produce again either several antheridia directly or a papilla
which gives rise to an antheridium and a stalk; the latter often
repeats papilla formation again and again, so that there are produc
conspicuous branching systems bearing numerous antheridia.

The young antheridium enlarges after its formation until 1s
length becomes 2-4 times greater than its breadth, the growth of the
cell being accompanied by that of its nucleus. The cytoplasm

contains deeply staining granules and is very dense, especially in the 5

neighborhood of the nucleus. The nucleus in the resting condition

contains a comparatively large amount of chromatin substance

arranged in a network evenly distributed throughout the nuclear
cavity. At this time neither kinoplasmic accumulations nor cei
trosomes are differentiated. The nuclear network, com of
tagged chromatin, now becomes transformed into a somewhat thicket
thread (fig. I 5).

This transformation of the chromatin from a fine ragged reticulum
to a thread accompanies the first manifestation of polarity '
nucleus; for it does not occur simultaneously throughout the cavity

1909] YAMANOUCHI—MITOSIS IN FUCUS 177

but is more active near the nuclear membrane. The fine ragged
material previously scattered throughout the central portion of the
nucleus moves toward the peripheral region and becomes trans-
formed into chromatin threads, leaving the central part of the cavity
comparatively free from chromatin (fig. 16). The chromatin threads
become thicker and an eccentric distribution of them takes place
(ig. 17), until finally they are grouped in synapsis at one side of the
cavity (fig. 18).

The chromatin threads, thus eccentrically grouped in synapsis,
have a certain regularity, i. e., they are not in a tangled mass or ball,
but are in groups of almost parallel loops, converging to a spot where
they are attached to the inside of the membrane. While this eccentric
synapsis of chromatin threads is going on within the nucleus, in close
association with the threads in synapsis, the cytoplasm directly out-
side the membrane becomes transformed into dense kinoplasm.
Not infrequently there are two synaptic groups (fig. 19) at two
*pposite poles within the nucleus; and naturally in such cases two

plasmic accumulations appear in association with the two
synaptic groups.
The loops grouped in synapsis thicken and shorten, the two arms
af each loop touching each other (figs. 20, 21). They now condense
Considerably, appearing therefore a little smaller, and become de-
from the spot where they lay during synapsis. Each of these
condensed loops becomes a pair of bivalent chromosomes (fig. 22).
Later, the two halves of such a bivalent chromosome become closely
applied to each other, so that the whole chromosome appears to be a
mall spherical mass. Such is the condition of prophase.
: Tn late Prophase a single centrosome appears in the kinoplasmic
enaulation at one pole (fig. 23), the centrosome at the other pole
making its “ppearance later. Radiations and achromatic spindles
‘ia ‘2 connection with the centrosomes. The spindles then
pai to the chromosomes and an equatorial plate is

ag (fig. 24).

OS oielaea to note the origin of the chromatin threads or
ines synapsis and the relation between the spirem and chromo-
the ees € chromatin threads in synapsis, which have arisen from

Srmation of a delicate ragged chromatin reticulum in the

178 BOTANICAL GAZETTE [MARCH

resting nucleus of the young antheridium, are not paired, but single,
As a consequence, the loops in synapsis are single in nature. An
examination of the loops cut transversely during synapsis showed
that the arms of the loops are altogether about 64 in number (fig. 21).
Finally, both ends of the loops being detached, 64 chromosomes are
formed, each pair of which, being derived from two arms of a loop,
becomes a pair of bivalent chromosomes. The number of these
bivalent chromosomes may be readily counted again in the polar
view of the equatorial plate (fig. 25). When the two halves of a
bivalent chromosome begin to separate, the figure (fig. 26) shows
the characteristic aspect of the heterotypic mitosis. The two sets of
the daughter chromosomes then separate and proceed to the poles of
the spindle (figs. 27, 28, 29, 30). The central spindle is of short
duration; when the daughter chromosomes aggregate in a mass al
organize a new nucleus, the spindle fibers entirely disappear. The
centrosomes disappear at the end of telophase.

The two daughter nuclei, after a short rest, commence the second
division, which is simultaneous, the antheridium remaining without
much increase in size. In prophase, 32 chromosomes are differ-
entiated from the chromatin reticulum, and in the later part of this
phase two centrosomes appear (jigs. 31, 31a) one after the other; the
achromatic spindle is developed in connection with the centrosomes
and an intranuclear mitotic figure is established (figs. 32, 320).
Metaphase (fig. 32), anaphase (fig. 33), and telophase in the two
nuclei proceed simultaneously and finally four nuclei are formed.

Soon after the telophase, the cytoplasm between the four newly
formed nuclei shows a fibrillar arrangement connecting the nuclei;
but the display is of short duration and the four nuclei remain either
in. @ §roup or scattered with no regularity along the longitudinal ax!
of the antheridium. The second division does not differ much from
ee al mitosis, except that the nuclear membrane dissolves at
earlier stage in prophase, and that no cell plate is laid down between
the daughter nuclei.

The four nuclei in the antheridium, after a short rest, begif =
in each of these four nuclei naturally resu">,
eight nuclei (figs. 34, 344, 35, 35a, 36, 36a). The eight nuclei giv

1909] YAMANOUCHI—MITOSIS IN FUCUS 179

rise by simultaneous division to sixteen nuclei (fig. 37). The fifth
division follows at once in each of these sixteen nuclei, resulting in
the formation of thirty-two nuclei (fig. 38). These simultaneous
mitoses take place with only short resting periods between, and
precisely the same as the second mitosis. The centrosome is most
brilliant at the first mitosis and gradually becomes fainter in the
successive divisions. The number of chromosomes in early meta-
phase (figs. 34a, 37a, 38a) and late anaphase (fig. 36a) of these
mitoses is 32.

The formation of partition walls in the antheridium begins to
take place at the 32-nucleate stage. Up to this stage, the nuclei of
the antheridium are free, but finally in telophase of the fifth mitosis,
with the disappearance of the central spindle in each mitotic figure,
there could be seen in the neutral region between any two nuclei the
faint manifestation of a protoplasmic plate formed by the transverse
walls of fine alveoli becoming perceptibly thicker and arranging
themselves in such a way as to appear as an uneven or somewhat
aigzag line in section. The unevenly continuous walls of the alveolar

ellae grow gradually thicker, and soon uniform plates are laid
down simultaneously, so that the antheridium is divided into 32 cells.

The nuclei in these cells of the antheridium undergo one more
mitosis, the sixth, which results in 64 nuclei (fig. 39). Thirty-two
catomosomes are present at this mitosis (fig. 39a). ‘This last division
Bako accompanied by the laying down of thin protoplasmic parti-
ton walls, so that the antheridium now contains 64 cells, which are
Spematids or sperm mother cells.
Hag nuclei in the spermatids undergo a peculiar modification, and
an accompanying change of cytoplasm surrounding the nucleus,
hag Produced a sperm with two cilia. The details of the events
a. in the antheridium following the 32-celled stage, a ep
a. evelopment of the sperm from the spermatid, will be treate
Parate paper which will be published later.

MITOSES IN OOGONIA
e : 'S well known that oogonia in Fucus develop from the wall cells
.. COnceptacle. The wall cell puts forth a papilla which divides

Int :
° two cells, an oogonium and its stalk. The oogomium enlarges

180 BOTANICAL GAZETTE [MaRcH

to a considerable size, and three mitoses occur within, naturally pro-
ducing 8 nuclei, each of which with its cytoplasm becomes an egg.

The mitoses in the oogonium of Fucus have already been studied
chiefly in Fucus platycarpus, and in a supplementary way in F.
serratus, by STRASBURGER, and mainly in F. vesiculosus, and in a
supplementary way in Ascophyllum nodosum, by FARMER and Wa-
L1AMs. The detailed accounts given by these authors are devoted
chiefly to the last one of the three mitoses, the first and second mitoses
being touched only slightly. The following is a description of the
first two divisions in the young oogonium in Fucus vesiculosus.

The resting nucleus in the oogonium contains a delicate chromatin
reticulum which is scattered irregularly throughout the cavity. The
amount of chromatin substance seems rather scanty in proportion
to the size of the nucleus (fig. 40). One or two very large nucleoli
generally lie isolated in the center. The cytoplasm in general has a
very delicate alveolar structure, which is very frequently interrupted
here and there by plastids, physodes, and _ black-staining spherical
bodies of undetermined nature. Toward the periphery of the nucleus,
the cytoplasm assumes a mixed structure of fine granules and fibrils
The nuclear membrane seems extremely delicate. No polarity is
manifested in this resting condition.

In very early prophase, a ragged chromatin reticulum gradually
passes into a thread, at first branched and then becoming simple.
As was described for the first mitosis in the antheridium, the trans
formation of the ragged chromatin into a thread is more active at the
periphery of the cavity, so that after a while the chromatin threads
are observed running irregularly and more abundantly along the
periphery than in the center, thus leaving the center nearly free from
chromatin (figs. 41a, 41b). When the partial distribution of the
chromatin thread proceeds farther, the most of the tangled mass .
threads is located at one side of the nuclear cavity (figs. 41), #%
42b), showing the beginning of a typical synapsis.

Coordinate with these internuclear changes, kinoplasm develops
and accumulates close to the nuclear membrane at a spot where
associates with the synaptic group of the threads within the nucleus.
The threads Sradually shorten and thicken. The irregularly tangled
threads now become Tegularly arranged into’ loops. These shunt

1909] YAMANOUCHI—MITOSIS IN FUCUS © 181

are evidently formed by the folding back repeatedly of long con-
tinuous threads, the blunt ends of which protrude toward the cavity,
while the opposite ends become closely attached to the nuclear
membrane (figs. 43a, b, c; 44a, b). The loops are therefore not
independent of one another, but are connected also at the base, which
is in contact with the membrane in such a way that an arm of one loop,
by turning back, passes directly to the arm of the next loop. These
connections of the loops at the base are detached and there results
a number of loops in synapsis. The number of the loops is not
easily counted from profile views; however, a section cut transversely
through the loops showed that there are 64 cut ends of arms of the
loops (fig. 45). Consequently, the number of loops is 32, each loop
consisting of two arms.

The loops differ at first in their thickness and length, but by
thickening and shortening they gradually become similar, a change
which is more rapid in the thinner and longer ones. The loops
now become more closely associated with one another during the
culmination of synapsis. The two arms of each of these loops in
contact with each other gradually become more compact and con-
sequently appear smaller. The two arms of each of these loops then
“parate at the bend—the point of connection—and form a pair of
bivalent chromosomes in prophase of the first division. The bivalent
aeevingg remain for a while at the spot where they were grouped

synapsis, and then become distributed in the nuclear cavity (fig.
4). Therefore in Fucus, pachynema, strepsinema, and diakinesis
stages are very much modified.

a attention was paid to the centrosome. When the kino-
9 ae accumulation is first visible at one pole of the nucleus in early
Sion ae small body is differentiated distinctly in the midst of
The era and it soon becomes surrounded by radiations.
rept: osome with its radiations is always in association with the
Scattered ag of loops. When the bivalent chromosomes are
Makes, its ughout the cavity (figs. 47a, b, c), a second centrosome
sometimes aaa generally at a distance from the first one,
The .. apart from it, but often not so far away (jigs. 49a, b).
but then sie — within the kinoplasm sometimes fragments into two,

y remain side by side without separating or establishing

182 BOTANICAL GAZETTE [MaRcH

a new sister centrosome with new radiations. The two centrosomes
in the prophase of the first division in the oogonium in Fucus seem
to be entirely independent, one appearing after the other. Often the
second centrosome has not yet appeared even at a late prophase, when
the chromosomes are well organized (figs. 48a, 6). The radiations
seem to increase in number and elongate, probably at the expense
of the cytoplasm, as the mitotic process proceeds from prophase to
metaphase.

The spindle fibers at first are clearly seen developing from the
area surrounding the centrosome, where the nuclear membrane
seems to be so thin as to allow the intrusion of the achromatic sub-
stance. The rest of the membrane holds its contour very sharply,
so that the mitotic figure is intranuclear. ‘Thus the intranuclear
spindle of Fucus seems extranuclear in origin.

In late prophase the bivalent chromosomes are arranged in the
equator. The nucleolus often remains as a vacuolate structure.
The axis of the figure of the first division is variable, either parallel
(fig. 50) or perpendicular (fig. 51) to that of the oogonium. The
nuclear membrane, as a rule, dissolves after metaphase, and yet the
outline of the figure remains even to late anaphase (fig. 53) without
the intrusion of the surrounding cytoplasm.

The number of chromosomes in prophase emerging from synapsis
is 32, each being bivalent (figs. 48a, b). The same number is counted
from the polar view of early metaphase (fig. 52b). In metaphase
the two halves of the bivalent chromosomes separate. These tw?
halves are not formed by the splitting of one chromosome, but aré
two independent chromosomes which were two arms of one loop:
Later metaphase (fig. 53) and anaphase (fig. 54) follow; in -
anaphase the chromosomes near the pole are straight rod-shape,
without any apparent indication of partition (fig. 55). Probably the
Initiation of the splitting which provides for the second division m4)
be very much delayed in this form. After telophase there are organ
ized two daughter nuclei and the centrosomes become unrecognizable.

The second division in the oogonium follows the first after omly
a short test. The differentiation of the chromosomes from the é
chromatin Teticulum and the appearance of centrosomes seem -
tially the same as in the typical mitosis. ‘The mitotic process #

1909] YAMANOUCHI—MITOSIS IN FUCUS 183

the two nuclei, from the beginning to the end, always proceed simul-
taneously (figs. 56-62). In the prophase generally a remnant of the
nucleolus is seen at the side of spindle fibers (jig. 56) which persists
to a late anaphase (fig. 59). The relation of the axes of the two
figures varies (fig. 57). In early metaphase the polar view of the
chromosomes in the equator shows the number to be 32 (fig. 580).
Anaphase and telophase follow as was described in typical mitosis.
The centrosomes always persist with a beautiful display of radiations.
When the daughter chromosomes reach the poles and become vacuo-
late, some of the central spindles seem to be replaced by fibrillar
cytoplasm. The cytoplasm between the newly formed nuclei of two
sister figures also changes to a fibrillar structure; thus the four nuclei
are connected with cytoplasmic fibers (jig. 63) that resemble the late
telophase of the second division of spore mother cells of some higher
plants. Soon after, the fibrillar cytoplasmic structures fade entirely
away, and the four daughter nuclei come into close association with
one another at the center of the oogonium, and rest for a considerable
petiod. There then follows a rapid growth in the oogonium, which
almost reaches its full size before the third division begins.

Detailed descriptions of the third division were given by FARMER,
Wittams, and STRASBURGER, and therefore need not be repeated
here, A point or two concerning chromosomes seems worth men-
tion, In the early metaphase of the third division, when the chromo-
oe of a slightly bent rod-shape are arranged in the equator, they

- Such a position that their long axes lie parallel to the equator
without overlapping one another. As a consequence, the profile
nti the figure in this stage (fig. 64) shows the end view of the
: ae and the polar view their whole length. It is very easy
then onstrate that there are 32 chromosomes. The chromosomes
nes split longitudinally in the equator (fig. 65), and after keeping
the a (fig. 66) for a short time, they become directed toward
ee 67), and then the usual anaphase and telophase follow.

€ daughter nuclei contain 32 chromosomes.

FERTILIZATION AND THE FIRST SEGMENTATION DIVISION

Ps €vents which take place during fertilization as well as during
Segmentation division have been described by FARMER, WILLIAMS,

184 BOTANICAL GAZETTE eanen

and STRASBURGER. Avoiding an unnecessary repetition, a few points
concerning the centrosome and chromosomes may be noted.

The resting nucleus of the discharged egg has shown no mani-
festation of polarity. Cytoplasmic alveolar structures as well as
plastids, and spherical globules of various sizes are arranged radially
about the nucleus. The cytoplasm surrounding the nuclear mem-
brane has a finely granular aspect. When the sperm has entered
the protoplast of the egg and is advancing toward the egg nucleus,
a change occurs in the latter. At a certain spot outside the nuclear
membrane, there is first observed a dense kinoplasmic accumulation,
in which there lies a single deeply staining body very close to the
membrane. Faint radiations are formed from the kinoplasm sur-
rounding this centrosome (jigs. 68a, 68b). The egg nucleus, there-
fore, is furnished with a single centrosome before the sperm reaches
it. The second centrosome has been found to appear in connection
with the sperm.

While the sperm is proceeding toward the nucleus, there appeat
numerous irregularly crowded granules, surrounding the nuclear
membrane. The size of these granules at first is not very different
from that of the centrosome of earlier occurrence, but they rapidly
grow larger and are either spherical or (sometimes) elongated. Theit
growth, thus, is different from that of the centrosome, so that small
granules of the same size as young centrosomes can be distinguished
from genuine centrosomes. Such is the condition of the region su!
rounding the nucleus just before the appearance of the second centro”
some. The sperm then reaches the egg nucleus, becomes closely
applied to it, and seems to slip in through the nuclear membrane
(fig. 696). At this very instant, there is first observed a new centro
some with radiations, appearing at the spot where the sperm ent

The second centrosome might have been brought in some ied
by the sperm, as was suggested by STRASBURGER (24): Or .
probable that one of the granules surrounding the nucleus might
have been brought to the spot mechanically by the streaming move
ment of kinoplasm caused by the progress of the sperm. At any
Tate two centrosomes do appear, one after the other, the first .
being visible before the entrance of the sperm, and the second oor
in connection with the entry. That the appearance of the seo”

1909] YAMANOUCHI—MITOSIS IN FUCUS 185

centrosome is always associated with the sperm is evidenced by cases
of polyspermy (jigs. 76-79).

Coalescence of the egg and sperm, the entry and progress of the
sperm in the egg cytoplasm, and the entry of the sperm into the
nucleus, all occur with rapidity. The chromatin of both the sperm
and egg nuclei forms the reticulum of the fusion nucleus. The
chromatin of both nuclei is mingled so as to become indistinguishable
(figs. 71a, b, c). The mitoses at the segmentation of normally
fertilized eggs (figs. 72a, b; 73; 74; 75a, b, c) take place as described
by STRASBURGER and by FARMER and Wrrtrams. The number
of the chromosomes in the prophase is 64 (fig. 752).

In cases of polyspermy, when two sperms enter the egg nucleus,
two centrosomes appear in the two spots where the sperms entered ;
when three sperms have entered, there are three centrosomes. In
case of bispermy there are developed three poles, and in case of
trispermy (fig. 76) four poles (fig. 77) are present; for one pole has
already appeared before the sperm enters. In the nucleus with three
poles, there are tripolar spindles, and 96 chromosomes become dis-
tributed upon the three spindles. The chromosomes split longitudi-
nally at the metaphase, and at telophase two sets of 32 chromosomes
meet at each of the three poles to form three daughter nuclei.

Ina quadripolar spindle (fig. 78) 128 chromosomes are dis-
2 six spindles, and each of the four poles receives three
to § aughter chromosomes, numbering 21, 21, and 22 (jig. 799);

orm daughter nuclei. In cases of polyspermy, the formation of
— nuclei occurs simultaneously. |
re 1S very interesting to note that in these cases of polyspermy,
a of the number of the chromosomes is maintained by
agen multipolar spindles. Whether or not polyspermy may

I natural conditions has not been determined.

DISCUSSION OF CYTOLOGICAL PHENOMENA
“os gaa of cilia-bearing structures and centrosomes and
Which bok relationship is treated best in such a form as Fucus, .
lem is quit epharoplasts and centrosomes are present. As the pro
At Sit important, it will be treated in detail in the next paper.
ent only a brief account of the chromosomes will be given-

186 BOTANICAL GAZETTE [MARCH

Origin of the bivalent chromosomes.—Although the actual seg-
mentation of the chromosomes in Fucus occurs just after the nucleus
has emerged from synapsis, their virtual preformation, as continuous
chromatin threads from which the chromosomes develop, begins very
early in prophase. As was described before, the ragged reticulum
of chromatin in the resting nucleus gradually becomes transformed
into a thread running in various directions, the transformation being
very much more active at the periphery than in the center of the
nuclear cavity. The threads in their beginning are uneven and
branched, then they become much evener and the transformation
continues, so that long continuous threads are formed, running mostly
in the peripheral region of the cavity. The threads thus formed
seem to have no ends (jig. 42), and apparently form one continuous
thread. Moreover, any part of the thread shows its single nature
from the early beginning of the transformation up to its completion
as a continuous structure. Entering into the synaptic condition, the
single thread then shortens and thickens, and becomes eccentrically
grouped as a loose tangled mass at one side of the nuclear cavity; 9
that eventually a number of loops are formed by the repeated folding
of the thread (figs. 18, 20, 43, 44). The loops so developed are there-
fore still continuous with one another at the bases where they come
in contact with the nuclear wall. The loops then become arranged
in a loose bunch, parallel and regular, with their bases attached !
the nuclear membrane, while the opposite folds protrude into the
cavity. Then the loops continue to shorten and thicken and become
more and more aggregated; each loop then folds at its bent end s0
that the bent arms are in contact with each other, when synapsis =
reached its culmination. As they emerge from Synapsis (7i8- #);
there are present 32 bivalent chromosomes, which become deta
from the nuclear membrane, moving toward the various regions of
the nuclear cavity.

__ The relationship of the chromatin thread in prophase, the mitt
In synapsis, and the bivalent chromosomes of postsynapsis, may
clearly followed. A pair of bivalent chromosomes corresponds ,
one of the loops in synapsis; the loops being formed by a folding
of the chromatin thread, so that a loop in synapsis should be
sidered as composed of two sporophytic chromosomes arranged ¢

-back |

PaaS POT ART RR eS ne ae eter ye a NI a ee ee ee na) ee ee ae ee eee

Tay ee gt he NO EEE Site patentee res

FRE er ee ee

1909] YAMANOUCHI—MITOSIS IN FUCUS 187

toend. If we apply a modern interpretation of synapsis to this case
of Fucus, the chromatin of paternal and maternal origin becomes
arranged in early prophase, not in parallel threads, but with the
chromosomes end to end, so as to form a single thread, which, passing
the so-called leptonema stage, enters into the synaptic condition,
during which there probably takes place a close association of the
chromatin of the two origins. In this case, the pachynema and
strepsinema stages (if they occur at all) must be of very short duration,
and consequently the chromatin thread of the zygonema condition in
synapsis passes directly into the diakinesis stage. The two elements
of the bivalent chromosomes then separate from each other, thus
effecting what may be regarded as a reduction. Generally in Fucus
the initiation of the longitudinal splitting which provides for the
second division does not occur even in late anaphase of the first
division, but probably may occur before the organization of the
daughter nuclei, as in the generally accepted account of sporogenesis.
Neglecting for a moment the many points which differ in particu-
lars, the results in Fucus, namely, that the chromosomes emerging
from synapsis show the reduced number, and that the reduction has
— Place by an end-to-end fusion of sporophytic chromosomes,
agree In essentials with the views published by FARMER and Moore
(6, 7), SCHAFFNER (21, 22), Morrrer (15, 16), and STRASBURGER
and by one group of zoologists, such as vom RATH (17),
“CKERT (20), and MONTGOMERY (13, I4)-

poem the origin of bivalent chromosomes, however, the
ei 1S fully convinced of the correctness of the interpretation that
tac majority of cases now investigated, two independent threads
ee in early prophase and become associated side by side in
Pion and that when the two threads emerge from synapsis they
ees 2 eaepai of the bivalent chromosome. Such cases were
, . established by Gritcorre (10, II, 12), BERGHS (3, 4), ALLEN
%s » RoseNBERG (18, 19), and some others, including the author
genesis - 31). The author, in a forthcoming paper on - gscet
as has the — cinnamomea has reached the same conclusion
ever, are tter group of investigators. The results in Fucus, how-
distinct not deniable. It is not inconceivable that there are two
types of arrangement of sporophytic I somes at synapsis-

Cilboiuy
J

188 BOTANICAL GAZETTE [MaRce

Constancy in the number of chromosomes.—After the appearance
of STRASBURGER’S classical paper (23) on ‘‘ Periodic reduction of the
number of chromosomes in the life-history of living organisms,”
investigators of many forms added to the evidence in favor of the
proposed theory. A plant is known to have a certain number of
chromosomes, without much variability, in one phase of its life-history
When the number is not too great, an accurate counting is not difficult.
The larger the number, however, the more difficult the counting
becomes, especially when the chromosomes are long and filamentous,
because the stages favorable for exact counting then become more
and more narrowly limited.

Unfortunately the rarity of the favorable stages has led some
investigators to the hasty conclusion that the counting is almost
impossible, while others, being unable to find the favorable stage,
have tried to make a rough estimate of the number from such stages
as they had. It is no wonder that such rough estimates, based upon
stages unfavorable for counting, should vary. It is curious to note
that even in Nephrodium molle, which contains 66 chromosomes in
the gametophyte and 132 in the sporophyte, the number was clearly
counted by the author both in apogamous and in normal forms,
while FARMER and Dicpy (5) claimed that the number of chromo
somes varied in the allied forms of Nephrodium molle which they
studied. The constancy in the number of chromosomes in norm
cases has been cited as one of the important proofs of the individuality
of the chromosome, and the importance of this theory in any discus
sion of heredity cannot be neglected.

metaphase (figs. 3, 9) and anaphase (figs. 6; 12a, b), 64 chromosome
were counted. Although the antheridium is very small, the polat
view of the mitotic figures in early metaphase showed clearly the sam?
number, as 32 bivalent chromosomes in the first division - j #
univalent chromosomes in the mitoses following the second division
In the first mitosis in the oogonium 32 bivalent chromosomes “°
Present (jigs. 47, 48, 52) and, as in the antheridium, 32 univalent o®

Se SHES ay ees eee eto ae et omar Thee ise

1909] YAMANOUCHI—MITOSIS IN FUCUS 189

appear in the second (fig. 58) and third (fig. 64) mitoses. In the
first division of the fertilized egg there are 64 chromosomes arranged
in the equator. Thus, the number is constantly 32 and 64 in Fucus
vesiculosus,

Farmer and Wittrams (g) state that in Ascophyllum nodosum
the approximate number estimated in the mitotic figure of the oogo-
nium mother cell is about 26-30, and later on they counted in the
third division in the oogonium 14-15 as the reduced number. STRAS-
BURGER (24) considers 30 to be the probable number in Fucus
platycarpus, in which he studied chiefly the division in the oogonium.
Such a difference in the number of chromosomes in different species
of the same genus or in allied forms which grow in normal conditions
has also been known in other cases; for instance, in Osmunda, where
Osmunda regalis has 12 and 24 chromosomes and O. cinnamomea
22 and 44.

Aliernation of generations.—It has been suggested by STRASBURGER
(26) that the antheridia and oogonia in young stages (Anlagen der
’ gonien und Antheridien) should be regarded as corresponding not
with antheridia and oogonia of Dictyota but with its tetrasporangia,
although the exact phenomena of reduction which occurs in the first
two divisions in these structures was not then known in detail. The
wei result may confirm the correctness of his suggestions.

Briefly summarizing the nuclear conditions of Fucus: The
utes cells of the plant contain 64 chromosomes, and the same
ein 1s present up to the formation of antheridium and oogonium
appear, a the first nuclear division in these initials 32 chromosomes
of the oe reduced number, but they are bivalent. At the telophase

ee second division there are 32 univalent chromosmes. Conse-
— ys the four nuclei resulting from the second division In both
nrg and antheridium initials are the first nuclei which contain
Within a8 chromosomes. Each of the four nuclei divides sae
anther; ‘am structures, once in the oogonium and four times in the

Wg tsa and after the division or divisions ‘there result 8
— oF 64 Same nuclei, each nucleus containing 32 chromo-
doubled i . enon of the sperm and egg nuclei, the number 1s
evelops the sporeling with the diploid number of chromosomes

Into a Fucus plant.

190 BOTANICAL GAZETTE [arcu

It would follow that the antheridium and oogonium initials up to
the second division may be well compared with spore mother cells
in the higher plants, and that the four nuclei in these structures thus
produced may be compared with microspores and megaspores, which
in Fucus germinate at once within the oogonia and antheridia,
and the gametophyte generations, thus initiated, undergo only one
mitosis in the oogonium and four mitoses in the antheridium. Thus
in Fucus the gametophyte generation with the haploid number
extends from the tetranucleate stage both in antheridium and oogo-
nium initials, up to the formation of the sperm and egg. With the
union of the gametes, the sporophyte generation with the diploid
number of chromosomes begins, and it terminates with the develop-
ment of the four nuclei in the antheridium and oogonium initials.

SUMMARY

The nuclear conditions during the life-history of Fucus vesiculosus
may be summarized as follows: .

1. The Fucus plant contains 64 chromosomes and the number Is
reduced at the end of the first two’nuclear divisions in the oogonium
and antheridium initials,

2. Each of the four nuclei produced at the end of the first to
divisions contains 32 univalent chromosomes, and this number
persists up to the formation of the sperm and egg; the phase com
taining 32 chromosomes may be regarded as the gametophyte s°™
eration. :

3- The union of the gametes doubles the number, and 64 chromo
Somes are present in every mitosis through the development oe
Fucus plant up to the formation of the first four nuclei in the oogomu™
and antheridium initials. The phase containing 64 chromosome
may be regarded as the sporophyte generation.

4. There is thus present in Fucus an alternation of the game
phyte generation containing 32 chromosomes, with the sporophy!®
Seneration containing 64 chromosomes.

THE Untyersiry or CHIcaco

1909] YAMANOUCHI—MITOSIS IN FUCUS Ig!

LITERATURE CITED
1. ALLEN, C. E., Chromosome reduction in Lilium canadense. Bot. GAZETTE
37:464-470. 1904.
—— , Nuclear division in the pollen mother cells of Lilium canadense.
Annals of Botany 19:189-258. pls. 6-9. 1905.
Bercus, J., Formation des chromosomes hétérotypiques dans la sporogénése
végétale. I. Depuis le spirtme jusqu’aux chromosomes mars dans la micro-
sporogénése d’ Allium fistulosum et de Lilium lancijolium (speciosum).
Cellule 21:173-189. pl. r. 1904.
, Idem. II. Depuis le sporogonie jusqu’au spiréme définitif dans la
microsporogéntse de |’ Allium jistulosum. Cellule 21:383-397- pl. 1. 1904.
ARMER, J. B., AND Dicpy, L., Studies in apospory and apogamy in ferns.
Annals of Botany 21: 161-199. pls. 16-20. 1907.
FARMER, J. B., anD Moorg, J. E. S., On the essential similarities existing
between the heterotypic nuclear division in animals and plants. Anat. Anz.
11:71-8o. figs. 1-29. 1895. ,
7- ——,, New investigations into the reduction phenomena of animals and
plants. Proc. Roy. Soc. London 72: 104-108. figs. I-6. 1903-
- Farmer, J. B., and Witttaus, J. L. On fertilization and the segmentation of
the spore in Fucus. Annals of Botany 10:479-489. 1896.
+ ———, Contributions to our knowledge of the Fucaceae; their life-history
vr Ses Phil. Trans. Roy. Soc. London B. 190:623-645- pls. 19-24:

“38

; Grécore, V., Les cinéses polliniques chez les Liliacées. Cellule 16:235-
297. pls. 1, 2. 1899. :
~_—> La réduction numérique des chromosomes et les cinéses de matura-
tion. Cellule 21:297-314. 1904.
ST, La formation des gemini hétérotypiques dans les végétaux. Cellule
13 aii pls. ap a 1907.
" *NTGOMERY, T. H., The spermatogenesis of Peripatus (Peripatopsis)
jouri up to the formation of the spermatid. Zool. Jahrb. 14:277-3°8-
14. bls, 19-25, 1900.
~_..? The heterotypic maturation mitosis in Amphibia and its general
1s, aga Biol. Bull. 4:259-269. figs. 1-8. 1903.
thet, Ter, D. M., The development of the heterotypic chromosomes 1n pollen
6. lee cells. Bor. Gazerre 40:171-177. 1905.
Wie | ditto (final paper). Annals of Botany 21:309-347- pls. 27, 28. 1907-
TH, O. vom, Zur Kenntniss der Spermatogenese von Gryllotalpa vulgaris
18. Rie Arch. Mikr. Anat. 40: 102-132. pl. 5. 1892.
H NBERG, O., Ueber Reduktionstheilung in Drosera. Meddel. Stock.
88. Bot. Inst. r904.
— Zur Kenntniss der Reduktionstheilung in Pflanzen. Bot. Notiser
905: 1-24, jigs. I-14.

12.

Ig.

192 BOTANICAL GAZETTE [MARCH

20. Rickert, J., Zur Eireifung bei Copepoden. Anat. Hefte 4:261-351. pls.
21-25. 1894.

- SCHAFFNER, J. H., The division of the macrospore nucleus (of Lilium phila-
delphicum). Bot. GAZETTE 23:430-452. pls. 37-39. 1897.

, Chromosome reduction in the microsporocytes of Lilium tigrinum.
Bor. Gazerre 41:183-191. pls. 12, I3. 1906.

23. STRASBURGER, E., Periodic reduction of the number of chromosomes in the
life-history of living organisms. Annals of Botany 8:281-316. 1894.

,» Kerntheilung und Befruchtung bei Fucus. Jahrb. Wiss. Bot.

30: 351-374. pls. 17, 18. 1897. ’
5: » Ueber Reduktionstheilung. Sitzungsb. Kénigl. Preuss. Akad. Wiss.
18:587-614. figs. 1-4. 1904.

6. » Zur Frage eines Generationswechsel bei Phaeophyceen. Bot. Zeit.
64:2-7. 1906.

27. WiLttams, J. L., Studies in the Dictyotaceae. I. The cytology of the tetra-
sporangium and the germinating tetraspore. Annals of Botany 18:141-160.
pls. 9, 10. 1903. II. The cytology of the gametophyte generation. Annals
of Botany 18:183~-204. pls. 12-14. 1904.

28. Wotrr, J. J., Cytological studies on Nemalion. Annals of Botany 18:609-
630. pls. 4o, 4I. 1904.

29. YAMANOUCHI, S., The life-history of Polysiphonia. Bor. GAZETTE 42:4°F
449. pls. 19-28. figs. 3. 1906.

30. ———, Sporogenesis in Nephrodium. Bor. GAzETTE 45:1-30. pls. 4
1908

31. ————,, Apogamy in Nephrodium. Bor. GAzETTEe 45: 289-318. figs. 3: 1908.

EXPLANATION OF PLATES VIII-XI
The figures were drawn with the aid of an Abbé camera lucida, under Ze
apochromatic obj. 1.5™™ N. A. x, 30, with compensating ocular 12; except fi

2 1) 29) STA, 324, 340, 35, 360, 374, 38a, 394, which were drawn Wi
pensating ocular 18:

PLATE VIII
Mitoses in the vegetative cells of the male plant
Fics. 14, 1b.—Two sections of the same nucleus in the cortical layer of the
thallus; no centrosome has appeared, the nucleus is in early prophase and :
ec number of chromosomes can be estimated from the two sections
ear O4, '

Fic. 2.—Late prophase: two poles; centrosomes in the center.
E 1G. 3.—Stage similar to fig. 2, viewed from pole: chromosomes ee
Fig. 4.—Metaphase: two sets of daughter chromosomes separate’.

1909] YAMANOUCHI—MITOSIS IN FUCUS 193

Fic. 5.—Anaphase: each of the two sets of daughter ss i i arranged
almost in a plane.
Fic. 6.—Stage similar to jig. 5, viewed from pole: chromosomes 64.

Mitoses in the vegetative cells of the female plant
Fic. 7.—Early prophase of the nucleus in vegetative cells of the thallus:
chromosomes (estimated) about 64.
Fic. 8.—Early metaphase: centrosomes with a few radiations.
Fic. 9.—Stage similar to fig. 8: chromosomes 64.
Fic. 10.—Anaphase.
Fic. 11.—Late anaphase.
Fics. 12a, 12b.—The same stage as fig. 11, viewed from pole: two sets of
64 diranncaomes,
Mitoses in antheridia
Fic. 13. Sage papilla, to become later an antheridium: nucleus
cae prop
IG, aie a of papilla: nucleus in anaphase; when this mitosis
is icin there will be formed a stalk cell and an antheridium.
Fic. 15.—Nucleus of the antheridium in resting condition, showing delicate
chromatin network: no centrosome.
Fis. 16.—Nucleus with chromatin network beginning to be transformed into
* more or less pronounced thread structure: nucleolus without any connection
with the network; no centrosome.
Fi. 17. pWhcleus with first indication of polarity: chromatin thread more
ckly tangled at one corner of the nuclear cavity; cytoplasm begins to show
plasmic nature,
5 a 18.—Nucleus in synapsis: parallel chromatin loops protrude from one
of nuclear membrane into the nuclear cavity.
Fic. 19, —Nucleus in synapsis: most of the chromatin loops aggregated at
oc few threads traverse the nuclear cavity, connecting the poles; this

thick!

Fic. 20
Fic, 21
— 22

-—Nucleus still in synapsis: the loops sndokennd. and shortened.

—The same stage, viewed at right angles.

-—Early prophase just after synapsis: chromosomes showing bivalent

eS 23-—Prophase: a centrosome at one pole; the two constituents of the
chromosome come in close contact, so its double nature cannot be recog-

at the oe = st hs ones bivalent chromosomes in the equatorial plate
®' Separation, revealing characteristic feature of heterotypicm mitosis.

194 BOTANICAL GAZETTE [MARCH

Fic. 27. anne: two sets of daughter chromosomes proceeding toward

the poles.
Fic. 28.—Late anaphase
Fic. 29.—Telophase of the first (heterotypic) division in the antheridium:

centrosomes faintly discernible.
1G. 30.—The antheridium after the first nuclear division: two daughter
nuclei in the resting condition; no centrosome.
G. 31.—Prophase of second mitosis in the antheridium: two daughter
nuclei in similar stage; centrosomes present. :
IG. 31¢.—One of the two nuclei shown in fig. 31, under higher magnification:
chromosomes 32.
IG. 32.—Metaphase: two figures in the same condition
Fic. 32a.—One of the two nuclei shown in fig. 32, etidey higher magnification.
Fic. 33.—Late anaphase: mitosis proceeding simultaneously in the two
nuclei. at
Fic. 34.—Late prophase of the third nuclear division in the antheridium:
four figures in similar condition.
Fic. 344.—One of the four nuclei shown in fig. 34, under higher magnification.
Fic. 35 .—Metaphase, viewed from the pole: each of the 32 chromosomes has
just split.
Fic. 36.—Anaphase: the four nuclei in the same condition.
Fic. 36a.—One set of daughter chromosomes from jig. 36, under higher
magnification: chromosomes a3. ‘
Fic. 37.—Late prophase of the fourth mitosis in the antheridium: eigh
figures in the same sta age.
Fic. 37a.—One nucleus in late prophase from fig. 37, under higher magnifica
tion: chromosomes 32. .
Fic. 38.—Late prophase of the fifth mitosis in the antheridium: sixtee
figures in similar condition. -
Fic. 38¢.—One beta from jig. 38, under higher magnification: chrom
somes 32.
Fic. 39.—Late prophase of the sixth nuclear division in the antheridium:
thirty-two figures in the same stage. aour
Fic. 39¢.—One nucleus from fig. 39, under higher magnification:
somes 32,
PLATE IX
Mitoses in oogonium ragged
Fic. 40.—Resting nucleus of the oogonium: chromatin showing
structure and nucleolus without apparent connection with it; no ee
Fics. 41a, 41b.—Two sections of the same nucleus in very early PP
ragged erp esa into a thread; a centrosome has made its appet
ance with a few radiat

not shown if
1G. 42.—Early iene a synapsis: centrosome with radiations
this figure.

1909] YAMANOUCHI—MITOSIS IN FUCUS — 195

Fic. 42a.—Nucleus from fig. 42, under higher magnification: chromatin
threads very much tangled; centrosome a shown —

Fics. 430, 436, 43c.—Th in synapsis: chromatin
threads in form of loops cane attached by their ends to a part of the nuclear
membrane, outside of which there lies a centrosome with radiations

1Gs. 442, 44b.—Two sections of the same nucleus in synapsis, similar stage to
above: there a black staining body associated with a nucleolus.

Fic. 45.—Section through the base of crowded loops, at contact with the
nuclear membrane, showing 60 or more isolated chromatin dots, some of them
connecting with one another; the dots are either the ends of the loops or their
optical sections.

Fic. 46.—Nucleus emerging from synapsis: chromatin loops moving from the
place of aggregation in synapsis; two arms of each of these loops are always in
close association, forming bivalent chromosomes; centrosome in next section.

Fics. 47a, 476, 47¢—Three sections of the same nucleus in prophase: 32
bivalent chromosomes; now two centrosomes lie at two poles, one of the centro-
somes being newly foemed, independent of the other that appeared at a previous
Stage; some of spindle fibers beginning to intrude into the nuclear cavity.

Fics. 4 , 48b.—Two sections of the same oogonium: 32 bivalent chromo-
a; these figures show the case where there is still only one centrosome.

Fics. 49a, 495—Two sections of the same nucleus in Hie two —

than 180° apart; intruding fibers attaching to chromosom

Fic. 50. — metaphase: intranuclear figure autiniale its axis parallel
to that of oogon
ice s-Tnranadea figure in prophase from fig. 50, under higher magni-

: nant of nucleolus still visible near the spindle. .

PLATE X
Fic. 51, —Metaphase a little later than the rit in fig. 50, with the axis of
. at right angles to that of the previous on
1G. 51¢.—Nucleus from jig. 51, under le magnification: the nuclear
pay has disappeared.
‘8S. 524, 52b, 52c.—Three sections of the same nucleus in metaphase: the
Section shows 32 bivalent chromosomes, although their bivalent nature
: y discernible,
ne ~Anaphase: the case where the contour of the nucleus still remains
oa d even after the dissolution of its membrane. ;
a. * 54-—Nucleus in anaphase, similar stage to fig. 53, under higher magnifica-

Fie ee -—Portion of one set of daughter chromosomes in late anaphase,
F &y IT rod- rete while seowanniia to the — —
o '—Prophase of the second onium: two figures similar.
Tome —Nucleus from fig. 56, under higher caguameaiid: the figure is
‘nuclear; nucleolus still remains.

196 BOTANICAL GAZETTE [MARCH

Fic. 57.—Oogonium in which two nuclei show early metaphase: two figures
perpetidiculas to each other.

Fic. 574.—One of two figures in jig. 57, under higher magnification.

Figs. 58a, 58b, 58c.—Three sections of the same nucleus in early metaphase:
the middle one shows 32 univalent chromosomes in the equatorial plate.

tox, 59.—Oogonium with two nuclei in early anaphase.

G. 59a.—Nucleus from fig. 59, under higher magnification: nucleolus still
remains; two centrosomes still showing conspicuous radiations.
. 60,—Oogonium with two nuclei in late anaphase.

Fic. 60a.—Nucleus from jig. 69; under higher magnificati

Fic. 61.—Telophase: tw with their sash stall recognizable;
central spindle about to disappear.

Fic. 62.—Late telophase: chromosomes aggregated at poles beginning to
vacuolize; meshes of cytoplasm arranged ssicacsbinous radially from two poles
toward the equator.

Fic. 63.—Section of oogonium cut transversely through its as, | aie sae
telophase of second mitosis: only three of four
every two of these three nuclei is an irregular fibrillar arrangement of ee

Fic. 64.—Late prophase of the third division, viewed from pole: chromosomes
~ in the equatorial plate before splitting.

G. 65.—Metaphase: nuclear membrane still present; most of the chromo-
somes they in the equator show their ends, the stage being just after splitting.

Fic. 66.—Late metaphase: nuclear membrane almost dissolved; daughter
ees beginning to separate.

7—Anaphase: nuclear membrane has disappeared, th
the stile teat nucleus undisturbed.

e contour of

PLATE XI
Fertilization and segmentation of fertilized egg

Fics. 68a, 68b.—Two sections of the same nucleus in resting condition, a
a discharged egg before fertilization, showing delicate ragged chromatin a
two nucleoli; a ag centrosome close to the nuclear membrane, without. ‘

radiations (fig. 68 a
rm
Fics. 690, "a be wo sections of the same nucleus of an egg when =

ae) te-
Fics. 71a, Ph = —Tives sections of the same fusion nucleus: the the dist
grating sperm nucleus has completely mingled with the contents of egg ®

Tg ag

Vigo. YD Ne
~~

a cate Sr

acs

nee

om,

PLATE X1

YAMANOUCHI on FUGUS

1909] YAMANOUCHI—MITOSIS IN FUCUS 197

so that there is now a homogeneous chromatin reticulum throughout the whole
cavity of the fusion nucleus.

Fics. 72a, 72b.—Early prophase of the first division in the fusion nucleus:
parts of chromatin threads begin to become pronounced, suggesting prochromo-
somes

Fic. 73.—Late prophase.

Fic. 74.—Early metaphase: nuclear membrane has disappeared; chromo-
somes not yet split.

Fics. 75a, 753, 75c—Same stage as fig. 74, cut perpendicular to the axis of
the spindle: fig. 75b shows polar view of 64 chromosomes arranged in the equator.

Cases oj polys permvy

Fic. 76.—Nucleus of fertilized egg with three sperms: a centrosome with
radiations started in connection with each of the spots where the sperms
entered

Fic. 77.—Early prophase: one of the four poles not shown.
Fic. 78.—Prophase, showing quadripolar spindle: one of the four poles not

Fics. 792, 79b.—Two sections of the nucleusinearly metaphase; six equatorial
shown from various views, one of them showing a polar view with 21
chromosomes,

THE REDUCTION DIVISION IN THE MICROSPORO-
CYTES OF AGAVE VIRGINICA'
Joun H. SCHAFFNER
(WITH PLATES XII-xIv)

This investigation, my fourth on the reduction karyokinesis, was
undertaken to test the correctness of my former conclusions on a subject
apparently beset with many difficulties, judging from the numerous
contradictory reports of various observers. Having obtained a
year’s leave of absence from the university for travel and study, |
prepared suitable material of A gave virginica L., which was found very
favorable for my purpose. The stamens were collected and killed at
various hours of the day during the last week in June and the first in
July, 1907, a number of vigorous plants being in bloom on the cam-
pus of the Ohio State University. The killing fluid used was a weak
chrom-acetic acid solution (0.3 per cent. chromic and 0.7 per cent.
acetic in water). After imbedding in paraffin, the sections were cut
to-20m thick and stained on the slide. After experimenting with various
stains and combinations, Heidenhain’s iron-hematoxylin was found
satisfactory, while Delafield’s hematoxylin and the various safranin
combinations gave very poor results. This was probably due to -
readiness with which the cytoplasm took up and retained these stains
In the whole investigation great care was taken to have the sections
correspond somewhat to the size of the nuclei, for sections too thick
or too thin may frequently give misleading figures. The nuclei ri
py microsporocytes of Agave are comparatively small, 15-20” :
diameter, so it was possible to obtain rather complete spirems am
spindles with rather thin sections. :

Tam greatly indebted to Professor Dr. Hans Scutnz, of the agp
versity of Ziirich, where the major part of the investigation was cart
on, for his kindly assistance and courtesy shown me during ™Y stay
in his laboratory

* Contributions from the Botanical Laboratory of Ohio State University,
Botanical Gazette, vol. 47] iy

1909] SCHAFFNER—REDUCTION DIVISION IN AGAVE 199

INVESTIGATION

Incipient stages of division.—The sporogenous tissue is differen-
tiated very early in the young stamens and all vegetative divisions
come to an end long before the earlier stages of the reduction division
are apparent in the microsporocytes. ‘There is therefore no danger
in Agave of mistaking belated prophases of vegetative divisions for
stages of reduction. The nuclei of the incipient sporocytes are quite
small (fig. r) and usually contain but one or two nucleoli and a rather
coarse chromatin net, in which are prominent dark-staining granules.
The cytoplasm is rather dense, with a spongy structure. As the
sporocytes grow in size the nucleus enlarges considerably, and at vari-
ous points in the enlarging net, masses of chromatin material appear
6. 2). Studied in detail the net reveals single chromatin granules
lying here and there in the linin meshes, and the clumps of chromatin
also show definite granules (fig. 2a). ‘These masses do not appear
to be of a definite number, but approximate the reduction number of
chromosomes. They continue to become more conspicuous as the
early stages of division progress, until they have the appearance of
tue protochromosomes. The meshes of the net at the same time
become larger, and the finer branches disappear, being probably
awn into the larger threads and masses, like the pseudopodia of
athizopod. The linin network appears to be the active agent, the
“Saag anerely carried apart or together as the linin is setts
chromoso ' ay is no question but that these masses are the “ pro-
apnroy: mes” of OverToN and STRASBURGER. As stated, they are
Pproximately of the same number as the reduction number of
.... The evidence is strong that they represent pairs of
to the bie, a nes which are orienting themselves panels
these ‘ag ion of the spirem. Since the massing and lengthening .
hot aay sual nae be synchronous, the apparent number 00s

definite pairi a with the reduction number. At this oon
be that ig of the individual chromosomes must occur, and it may
e eg a only definite pairing during the whole hee
Teduction ny Td rotochromosomes may also appear comets t gin
at two or i ", for the accumulation may at first be taking aor

the ee, points of the paired chromosomes. It appears
Omes, extended and spread out like a rhizopod or an

200 BOTANICAL GAZETTE [arc

amoeba in the chromatin net, mass themselves together as definite
individuals, probably in pairs; for thus alone are the later stages of
reduction intelligible.

During these early stages the tapetum is in the very beginning of its
development. It is therefore of value in helping to determine the
exact sequence in the development of the sporocytes. And it is of
course evident that if the successive stages cannot be determined
with absolute certainty, the whole investigation is vitiated.

After the development of the chromatin masses, they seem to
elongate, as is shown to some extent in fig. 3, and more perfectly in
fig. 4. Finally they are stretched out into a very long and delicate
continuous spirem, with rather uniformly distributed chromatin
granules (fig. 5). ‘The masses are probably all connected in series and
thus elongate into a continuous delicate strand.

If the individuality of the chromosome is admitted, we may conceive
the influence which causes paternal and maternal chromosomes t0
conjugate in pairs in reduction to be of much the same character as
that which induces cells to develop as male and female gametes with
subsequent union. This property may develop in the chromosomes
only at the reduction stage, and if this were the case, the paternal
and maternal units might be indifferent in regard to each other dur-
ing all the vegetative divisions. The evidence on this point must

1909] SCHAFFNER—REDUCTION DIVISION IN AGAVE 201

stage, even as the cells of a myxomycete lose their individuality in
the plasmodium.

First stages of synizesis——As soon as the extended and delicate
spirem is formed, the nuclei mostly appear in synizesis. There are
all types of contraction. The chromatin may stretch across the center
of the nuclear cavity (figs. 6, 8); it may be contracted at one side with
the nucleolus (fig. 7); or it may be balled up at one side of the cavity
with the nucleolus lying free (fig. 10). In some cases the mass is in
the center, and often the nuclear membrane is injured by the irregular
expansion of the nuclear cavity (fig. 9). The period of development
during which synizesis occurs is comparatively long, the anther
lengthening greatly in the meantime. The anthers of Agave thus
make a most favorable object on which to determine definitely the
stage when contraction must be looked for in the living material. A
thorough study was therefore made of unstained as well as of stained
sections, in order that I might become familiar with the appearance
of the cells in unfixed anthers.

Study of living cells.—Having ascertained the stage when synizesis
occurs in killed material, a study was made of living anthers during
the last two weeks in June, 1908, at Columbus. The anthers were
‘xamined immediately after removal from the plant. In some cases
“foss-sections were cut, in others the stamens were cut into short
Pieces and the sporocyte tissue squeezed out and mounted in water.
Both methods were satisfactory. In none of the numerous anthers
eehed during the two weeks was there the slightest evidence of
‘ynizesis. In the great majority of cases the nucleolus is in the center
Og nuclear cavity; occasionally it is somewhat to one side; and
We Tarely near the nuclear wall, as is almost universal in the synizesis
ag ae material. The nuclei look large, clear, and a.
fa. es and flaky material (no doubt the chromatin) agin
ce — the cavity. The synizetic knot would certainly be es i
Sia... ae In the killed material the synizetic mass shows
lad aed in unstained as in stained material. The aia
ape fo condition was found in all the cells in every ae
fea, € period before chromosome formation. The fact t : :
on eile nearly uniform position near the center of the nuclea

¥, while in synizesis they are usually near or against the nuclear

202 : BOTANICAL GAZETTE [MARCH

wall or flattened out in the “sickle stage,” is in itself sufficient proof
that synizesis is an artifact. But as stated, the chromatin can also
be faintly recognized in the living nuclei, and it should be still more
evident if in a contracted ball, since the cavity in typical synizesis is
entirely empty of threads, flakes, or granules. The granular material
in the nucleus often radiates outward from the nucleolus, and some-
times it is prominently distributed over the surface of the nuclear
membrane.

With salt solution and also with 95 per cent. alcohol, the cells
contracted considerably and soon became indistinct, so that it was
difficult to make out any details. The nuclei were displaced to some
extent. The weaker chrom-acetic acid solution, used for the paraffin
material, caused the whole mass of sporocytes to contract violently,
but not much displacement of the nuclear contents was noticeable.
This was probably because the cells were lying rather free and could
contract readily from all sides, or the fluid may not have acted long
enough. However, it is probable that the synizesis occurs rather
suddenly.

_ An attempt was made to stain the fresh material, both before and
after treatment with killing fluids; but this proved unsatisfactory,
the stained material showing no more detail then the living cells.

A study of the living microsporocytes of Agave virginica indicates
that synizesis, as seen in the usual paraffin sections, is an artifact
When the chromatin is comparatively free in the nuclear cavity and
is expanding, we find the most decided synizesis. Meanwhile, as
appear further on, synizesis is not confined to this stage, but occu!
to a greater or less extent until the chromosomes are fully developed.
It is'largely on account of the erroneous idea that synizesis occuls al
but one stage of division that a number of inaccurate interpretations
have been advanced, through which the whole subject of reduction
has been confused. :

Development of the chromatin loops.—The spirem begins to thicken
while the chromatin granules are still in a single row (figs. 17) sie
At this stage synizesis is still frequent, the spirem usually
crowded to one side, but occasionally lying entirely around the ee
wall (figs. 12, 13). The spirem now becomes very distinct, 8° -
is often possible to trace out great lengths of the thread by

1909] SCHAFFN ER—REDUCTION DIVISION IN AGAVE 203

properly. It begins to twist into loops and the chromatin granules
now appear double (figs. 14, 14a). Although the spirem is much
thicker at this stage than it was earlier, synizesis is occasionally present,
the spirem filling one-half of the nuclear cavity, as shown in fig. 15.
The double granules are at length prominent, although the spirem
does not split (figs.:16, 16a). Finally the whole spirem is thrown into
definite loops of various shapes and sizes. It is difficult to represent
the perspective of these loops in a drawing. One can trace out the
position and depth by focusing up and down, but in the camera
projection they appear nearly in a plane (figs. 17-20). There is no
question but that the spirem is continuous, since one can often follow
the thread through more than half of the loops without losing the
connection, and in uncut sections no free ends are present. In tan-
gential sections or half-sections one can frequently also follow through
three or four loops before coming to a free end (fig. 19). Practically
also, it seems impossible that such twists and loops could be formed
unless the spirem were continuous. In jig. 18a a number of twisted
loops are shown. Some of the loops are produced by a single twist,
which results in ring-shaped chromosomes (fig. 20). There are three
of these ring-chromosomes in the nucleus and they are developed
side by side. The three main types of loops are shown in figs. 204,
20b, and 20c. The loops are not formed, as in Lilium, with a central
knot, but more openly. In this stage synizesis was also present in
some of the material (fig. ary:
After the loops are developed, they are pressed and curved against
Oe nuclear wall, the whole central part of the cavity becoming very
- €. At the same time they break apart to form the individual
reduction or bivalent chromosomes (figs. 22, 23). It was exceedingly
“utcult to determine the number of chromosomes on account of the
a shapes of those bodies in some nuclei, as appears in figs.
é, 25; but it was finally determined that the number is twelve
bia » 37, 38). In jig. 26 only fragments of the twelve chromo-
= i shown, a large part of the nucleus being cut away.
none 2 where the number of chromosomes is said to —
: aes a greater or less number may not be of any significance, if the
, appears in vegetative division. Two or more chromosomes
might become united through a failure of transverse segmentation,

204 BOTANICAL GAZETTE [MARCH

but longitudinal division could proceed in the normal way and the
identity of the chromosomes not be lost. But in reduction the number
should be definite, if the karyokinesis is to furnish normal cells.

The nucleolus is still present when the chromosomes are fully
developed, but often shows signs of fragmentation, as in the examples
shown in fig. 31a. After the chromosomes are developed, the cyto-
plasm also shows a change in structure, having passed from a spongy or
reticulate arrangement to a more or less radiate structure (figs. 31, 32).

Individuality of the chromosomes.—The chromosomes continue to
become more indefinite in shape until they appear as irregular, dark-
staining, apparently structureless masses, very unequal in size. The
real character of the chromosomes can be studied to advantage only
during the formative period, although the larger ones can be recog:
nized even in the mother star. In the incipient chromosome loops
individuality is very marked. As stated, there are three small ring
chromosomes (figs. 20, 20a, 29, 63); four large long chromosomes,
two of which are very prominently coiled and always side by side
(figs. 20b, 22, 27, 29, 30, 32, 63, 64); and five smaller chromosomes
of various shapes and sizes. Since these are bivalent chromosome,
it is evident that, on the theory of the conjugation of maternal an
paternal chromosomes, the conjugating pairs must be quite similar
in shape and activity. In the microsporocytes the bivalent chromo-
somes have an individual shape and size easily distinguishable, and
the inference from this is evident, as also in the massing of the
matin in the early prophase, that these bodies are individualized and
retain their individuality from one division to another. Were the
chromosomes not individualized, they could not preserve such dete
forms and numbers from generation to generation. During 15
ontogeny, the chromosome passes through a series of forms, only t0
return, as any other organism, to a definite type at a definite stage :
From the present study and the investigations of others, it is evident
that the mechanics of chromosome reduction is simple, the ve
spirem orienting itself into folds, twists, or simple loops, giving
to all the various shapes, as rings, rods, coils, tetrads, and cro
The actual form observed in any individual case may be @ mere Pro”
jection, and great care should be taken to ascertain the actual shape
by observation from various points of view.

1909] ‘SCHAFFNER—REDUCTION DIVISION IN AGAVE 205

The spindle and late stages of synizesis.—The incept of the spindle
is laid down immediately over the surface of the nuclear membrane
while that structure is disappearing. At the same time, connecting
lines, which appear prominently in heavy-stained sections, are present,
forming a sort of network between the chromosomes (figs. 37, 38).
The incipient spindle appears as a dense wall of material that was at
first mistaken for the modified nuclear membrane, which, however,
lies on the inside. This double layer about the nucleus, together
with the connecting strands between the chromosomes, makes an ideal
arrangement for abnormal contractions, and at this stage there is
present a final prominent synizesis of the chromosomes, together with
the dissolving nuclear membrane inclosing them. The chromosomes
at this stage have not yet fused with the surrounding spindle. A few
examples of this appearance are shown in figs. 32-38, all about in the
same stage of division. Those which show the connecting fibers less
distinctly are from the lighter-stained preparations. In fact, without
a heavy stain, the connecting threads are barely visible. The con-
tracted nuclei are seen in slides side by side with cells having a normal
appearance. There is no doubt in the writer’s mind that the phenom-
enon is an artifact.

The spindle—The incipient spindle soon begins to show a fibrous
character, the fibers at first being few and indistinct, and running
more or less parallel toward opposite poles of the nucleus (fig. 39)-
- many cases two more or less pointed caps extend from opposite
Sides of the nucleus and become prominent before the longitudinal
cS me Nisible (figs. 4o-42). The points sometimes show delicate
ano fa my figs. 41 ete The spindle fibers develop rapidly, and soon
Ren slightly pointed structure is produced, in which the chromo-
Song one or more nucleoli lie scattered about (fig. 4 3): The
Pa fibers are also prominent, giving the spindle an irregular
and oe The spindle is bipolar from the beginning, originating

developing in the same way as in the vegetative divisions.

oa in 1899 showed that in Hemerocallis the spindle originates

nearly eg structure surrounded by a dark zone. This zone was

yy in Agave, but the difference may be due to sega’

inside of * owed, however, that the incipient spindle 1s entirely
e dark zone.

206 BOTANICAL GAZETTE lien

The spindle becomes narrower and more pointed and the connect-
ing fibers, apparently contracting while the spindle is lengthening,
gradually draw the chromosomes into a perfect circle in the equatorial
plane (figs. 44-60). In fig. 48 the spindle is distorted. This was
probably produced by the unequal contraction of the cytoplasm.
Figs. 39, 43, 47, 57, 60, 61 make a series, showing how the chromo-
somes are drawn from their scattered positions into the symmetrical
figure of the mother star. A large number of figures of this stage
have been included in order to show all the ordinary types of develop-
ing spindles to be seen in Agave. In some, the connecting fibers are
prominent; in others, especially as the chromosomes approach the
equator, one sees only a dark-staining central mass. It is important
to note that the spindle fibers appear thickest and densest in their
central parts, even in very young spindles. Apparently the chromo-
somes are attached to the spindle fibers from the beginning. The
crowding of the chromosomes against the nuclear wall, as shown
in jigs. 23, 39, brings the chromosomes into a position where their
fusion with the spindle fibers can be accomplished.

The mechanism for bringing the chromosomes from their scattered
position into the symmetrical wreath of the mother star is compat
tively simple, requiring only the shortening of the connecting fibers,
combined with a pull from the spindle threads exerted from the
poles. The action of the spindle as well as the attachments must be
looked upon as being accomplished by a viscid substance, perhaps
under the influence of attractive and repulsive forces. If the sub-
stance is contractile in the ordinary sense of the word, it must
acquire this property after development. his

Multipolar figures.—Multipolar figures were not numerous: T dé
may have been because of the comparatively small size of the nu :
and the thickness of the sections. A special study was made of tH?
multipolar figures found, and the conclusion was reached that they
were.all artifacts. The various types are shown in the series -
64-73. Fig. 6gisa diagonal section, included to show the charact
and position of the chromosomes in the mother star. Both poles ar
cut away, one end more than the other. F igs. 05; 66 i om
sections representing small parts of the nucleus and spindle. :
fibers are both spindle and connecting fibers and make an appest

TS eres eee RE eT a eee

FER ee et a OM Re

Suse eb iets = Re oa a

1909] SCHAFFNER—REDUCTION DIVISION IN AGAVE 207

ance very much like the figures usually given to represent multipolar
spindles. The writer believes that these connecting fibers have
caused much trouble in the interpretation of spindle sections. Fig.
67 might be taken for atripolar spindle. The few projecting fibers
were probably disturbed in the cutting. Fig. 68 is a spindle broken
and distorted by the knife. Fig. 69 is another torn spindle, the fibers
at one end being spread out by the knife. Fig. 70 has the fibers of one
pole cut diagonally. In fig. 71 one pole is perfect, with a well-
developed aster, while the other pole is cut away. In figs. 72, 73
both poles have been cut off. Such figures are common, as is neces-
sarily the case with cells in which the spindles lie in all directions.
The division of the bivalent chromosomes.—The chromosomes are
arranged symmetrically in the mother star (fig. 61), with the closed
end of the loop extending outward, at least in the long chromosomes
(fg. 67, a, b, c,d, e). The spindle fibers are attached very near or at
the free ends. In the following division the general appearance is
entirely different. The larger chromosomes are V-shaped and are
attached to the spindle fibers at the head of the V, the two free ends
Projecting outward (fig. 62). The individual character of the
“romosomes may occasionally be seen from the polar view, even
as late as the mother-star stage (fig. 63). The chromosomes are
pulled apart very rapidly and are considerably scattered before they
teach their new positions in the daughter stars (figs. 74-77) In some
cells one can see large nucleoli in the cytoplasm along with micro-
nucleoli (figs. 5/750; 70, 75).
re daughter chromosomes are arranged in a loose ring or plate,
then begin to contract, until they form a compact dark-staining
ha 7 8-82). In the earlier stages of the daughter star, con
sat are again favorable for counting the chromosomes (fg. 79)
fae smaller size is quite evident when compared with the bivalent
oa. of the mother star. Delicate radiations are usually
chromatin at ontraction stage of the incipient daughter nuclel,
Dodia-lik gins to expand, the chromosomes putting out pseudo-
is fo € branches which become more extended until a coarse net
hee ed (figs. 83-85); but even in the oldest daughter nuclei dis-

hable before the beginning of the following division, a con-

208 BOTANICAL GAZETTE [MARC

siderable part of each chromosome persists as an irregular compact
mass (fig. 85). There is thus in these figures an indication that the
individuality of the chromosome is preserved even in the chromatin
network,

GENERAL CONSIDERATIONS ON REDUCTION

The important facts brought out in the present investigation con-
firm a number of conclusions put forward by the writer and others
during the past ten years, most of which have been the subject of
continual controversy. In a science like cytology so much depends
on the manipulation of the material and the interpretation of the
figures, that one need not be surprised at the diversity of views held
in respect to all the more important cytological problems. In the
present paper, by leaving out certain figures in the series, one can
produce several of the “reduction processes’’ heretofore published.

The writer appears to have been the first to present a definite series
of observations to show that the first division after pseudo-reduction
is the real reduction division. A few previous reports had been pub-
lished, which were, however, largely guesses or assertions, without
definite evidence and sometimes even without drawings. =

In 1897, the writer presented his views on the reduction division
in the ovules of Lilium philadelphicum, advancing the definite con
clusions that the spirem is continuous and contains a single TOW .
chromatin granules which later undergo transverse fission; that the
continuous spirem doubles up and twists into twelve loops
reduction number, which then break apart at the inner ends opposite 5
heads of the loops to form the twelve chromosomes; that during
metakinesis the two limbs of the chromosomes are pulled apatt, finally
breaking at the middle; and that, therefore, there is a transverse division
in the first reduction karyokinesis, or a true qualitative division of -
chromatin. In that paper figs. 1, 2, 2a, 4, 4a, 8, 8b, 11,1 1b, 12, sy
214, 22, 23, 23b, 34, 35 formed a series for which only one interpre
tion was possible. Only by leaving out fig. 4 could another interpre
tation be given, in which case the double spirem appearing later Me"
be considered as conjugating instead of dividing.

In T9OI practically the same results were obtained for Eryth
and in 1906 for the microsporocytes of Lilium tigrinum.

ronium,

1909] SCHAFFNER—REDUCTION DIVISION IN AGAVE 209

PAULMIER in 1899 showed that in the spermatogenesis of Anasa
fristis the first division is the reduction division, and more recently
Montcomery, in a series of important investigations, has come to the
same conclusion. Griccs found a reducing division in Ascaris, and
observed that the chromosomes are not entirely separated until they
are drawn into the equatorial plane. :

Mortier after a long-continued study of Podophyllum, Lilium
Martagon, and other plants, has come to conclusions for the most part
similar to the writer’s, although for many years he held opposite
views.

GarEs, in a recent article, finds that in the reduction nucleus of
the microsporocytes of Oenothera rubrinervis the spirem segments
transversely into the 2x or sporophyte number, and that the members
of a pair are thus at first arranged end to end on a single thread.
Later the univalent chromosomes are separated, usually in pairs.

It is needless to review the extensive recent literature of reduction,
for in many cases the results appear radically different from those
presented in this report, and in examining the drawings and conclu-
‘tons based on them there seems little possibility of harmonizing or
explaining the differences.

F inally, it may be said that if any individuality whatever is ascribed
‘0 the chromosomes, it becomes evident that they should be arranged
~ to end to form the spirem, since this is the method in somatic
divisions, It ig not probable that the cell would develop a funda-
Mentally hew method of division in reduction, but rather that such
changes would be developed in the process as would suffice to

ting about the separation of the two sets of chromosomes. The

-eaiheg 8 described in this paper appears to the writer to be the only

Possible explanation of the figures. As has been stated, however, by

— Suitable selections, one could represent almost any of the
°us reduction karyokineses that have been described.

SUMMARY

=e une Testing nucleus in the microsporocytes of Agave virginica

a a linin network in which small chromatin granules are held,
separate or in lumps.

7 te beginning of division, the chromatin granules are massed

210 BOTANICAL GAZETTE (wince

together through the massing of the linin into a number of lumps cor-
responding approximately to the reduced number of chromosomes.
These masses probably represent bivalent protochromosomes.

3. The masses are all united and elongate greatly until a very
delicate, continuous spirem is produced, holding a single row of
chromatin granules.

4. After the delicate spirem stage the nuclei in killed material are
usually in synizesis. There is no union of two spirems in synizesis.

5. In living material no synizesis is evident, and the nucleoli are
not crowded against the nuclear wall, but usually have a central posi-
tion in the nuclear cavity. Synizesis at this as well as at later stages
is an artifact.

6. The spirem shortens and thickens while the chromatin granules
undergo transverse division. It finally orients itself into twelve loops
of different shapes and sizes.

7. The loops are pressed close to the nuclear membrane, forming 4
rather definite wreathlike circle, and do not radiate from a closely
entangled central mass as in Lilium.

8. The twelve loops break apart, forming the twelve chromosomes
—four very large, long, twisted chromosomes; three ring-sha
chromosomes; and five smaller, irregular, more or less bean-shaped
chromosomes.

g. The chromosomes are united by connecting fibers, which appar”
ently contract and draw the scattered chromosomes into the equatorial
plane while the spindle is elongating.

to. One or two nucleoli are usually present, which are st
in appearance after the spindle is far advanced in development.
nucleoli are sometimes thrown out bodily into the cytoplasm

11. The spindle originates as a more or less fibrous layer
surface of the nuclear membrane before that body dissolves,
this stage decided synizesis of the chromatin is often present
12. The spindle is bipolar from the first, with no accessory smaller
poles, the poles appearing at first as two, more or less pointed, dome-
shaped caps, much the same as in vegetative karyokinesis. _ .

13. The spindle fibers are usually most prominent and thickest ite
the middle, even in the early stages. There are commonly defin!
asters at the poles.

+1 normal
e

over the
and at

1900] | SCHAFFNER—REDUCTION DIVISION IN AGAVE 211

14. The multipolar spindles observed are explained as artifacts,
mostly produced by cutting.

15. The chromosomes divide transversely during metakinesis.

16. In the daughter nuclei, irregular masses of chromatin persist
into the resting condition. ‘These masses represent parts of the
twelve daughter chromosomes.

17. In the second division the chromosomes divide longitudinally. _

Onto STATE UNIVERSITY
CoLuMBus

LITERATURE CITED

ROSENBERG, O., Ueber die Individualitat der Chromosomen im Pflanzenreich.
Flora 93:251-259. 1904.
Overton, J. B., Ueber Reduktionsteilung in den Pollenmutterzellen einiger
Dikotylen. Jahrb. Wiss. Bot. 42: 121-153. 1905.
Miyake, K., Ueber Reduktionsteilung in den Pollenmutterzellen einiger Mono-
kotylen. Jahrb. Wiss. Bot. 42:83-120. 1905.
ARMER, J. B., AND Moone, J. E. S., On the maiotic phase (reduction division)
in animals and plants. Quart. Jour. Mic. Sci. 48:489-556. 1905.
Mormer, D. M., The development of the heterotypic chromosomes in pollen
mother cells. Annals of Botany 21: 309-347. 1907.
R, J. H., The division of the macrospore nucleus. Bor. GAZETTE 23:
430-452. 1897.

——, A contribution to the life history and cytology of Erythronium. 55
GazeTre 31: 369-387. 1901. :
~——; Chromosome reduction in the microsporocytes of Lilium era

‘ Bor. Gazerre 41: 183-191. 1906.
sAUIMER, F. C., The spermatogenesis of Anasa tristis. Jour. Morph. 15:
223-272. 1899.
— R. F., A reducing division in Ascaris. Ohio Nat. 6:519-527- 19°°-
7 & 1. The development of the microsporangia and microspores of
C Hemerocallis julva. Bor. GazEtTE 28:81-88. 1899. :
ates, R. RK, A study of reduction in Ocnothera rubrinervis. Bot. GAZETTE
46: 1-34. 1908,

EXPLANATION OF PLATES XII-XIV
‘ The plates are reduced five-eighths in reproduction. Figs. 2a, 11¢, 149, 16a,
ie va 204, 20b, 20c were drawn with a compensating ocular 18 and oil immer-
obj. objective v3 all the rest with a compensating ocular 12 and oil immersion
Hective xy, the latter combination having a magnification on the table of 2259.

212 BOTANICAL GAZETTE (mance

PLATE XII
Fic. 1.—Microsporocyte showing resting chromatin network.
Fic. 2.—Microsporocyte at a later stage showing masses of chromatin
granules in the net.
Fic. 2a.—A small portion of the chromatin net showing the linin and massing
of the granules,
1G. 3.—A nucleus with prominent massing of the chromatin into rather
definite adc. protochromosomes.
1G. 4.—Later stage; the chromatin masses stretching out into a definite
spirem.
Fic. 5.—The delicate slender spirem complete.
Fic. 6.—Somewhat older than fig. 5; synizesis of the spirem in the middle
of the nuclear cavity.
Fics. 7-10.—Other types of synizesis of the same stage as fig. 6; in fig. 9
the nuclear cavity is expanded.
Fic. 11.—Nucleus with spirem becoming thicker
Fic. 11a.—Single chromatin threads from fig. 11, showing the light-staining
linin with a single row of chromatin granules
IGS. 12, 13.—Types of synizesis in a later stage than those of figs. 6-10.
G. 14.—Continuous spirem beginning to fall into loops, showing chromatin
palin enlarged and partly double.
Fic. 14a.—A short piece of the spirem from fig. 14, showing the double nature
of the chromatin granules.
Fic. 15.—Same stage as fig. 14, showing one-sided synizesis of the continuous
gars d
16.—Beginning of the looped spirem, showing further thickening ”
ae of the chromatin granules.
Fic. 16a.—Pieces of a spirem from fig. 16, showing double rows of chromatin
granules and distinct lini ‘ciel
Fic. 17. so eleaciias spirem, much thickened and thrown into

ps.
Fic. 174.—Pieces of the spirem showing the method of looping and pe
Fic. 18. —Microsporocyte somewhat later than fig. 17, showing further thi

ing of the thread and development of the chromatin loops.
Fic. 18a.—A number of chromatin loops before the breaking of the ied
Fic. 19.—Section of microsporocyte in which several loops ca?

out; the section represents nearly half of the spirem. caning t0
Fic. 20.—Beginning of the broken skein stage; the chromosomes beginni

break apart; three ring-chromosomes still connected. preaking
Fics. 204, 20b, 20c.—Three complete chromosomes just after the

of the spirem.

pmosomes-
Fic. 21.—Synizesis in microsporocyte at time of separation of ch

mem
Fic. 22.—Nucleus with chromosomes completely separated; nuclear
brane still present

1999] «© SCHAFFNER—REDUCTION DIVISION IN AGAVE 213

Fic. 23.—Somewhat later stage; chromosomes all crowded against the
nuclear wall with a clear cavity in the center.

Fics. 24, 25.—Nuclei showing indefinite chromosomes.

Fic. 26.—Section of nucleus showing parts of twelve chromosomes.

Fics. 27, 28.—Nuclei showing twelve chromosomes of diverse shapes and
sizes.
Fic. 29.—Section of nucleus showing the three ring-chromosomes.

PLATE XIII :

Fics. 30, 31.—Nuclei with twelve chromosomes, showing the beginning of the
appearance of delicate connecting fibers.

Fic. 31a.—F ragmenting nucleoli taken from same stages as fig. 31.

Fics. 32-36.—Microsporocytes showing synizesis after the formation of the
chromosomes; also connecting fibers between the chromosomes.

1G. 37.—Nucleus contracted away from the incipient spindle; prominent

connecting fibers between the chromosomes.

Fic. 38.—The same, but with less synizesis of the nucleus.

Fic. 39.—Nucleus showing distinctly the incipient spindle.

Fic. 40.—Incept of spindle showing as two caps on opposite sides of the
nucleus.

Fic. 41.—Nucleus showing incipient spindle.

Fic. 42.—Nucleus with incept of spindle and aster at one pole.

Fic. 43.—Nucleus showing young spindle and connecting fibers between the
chromosomes,

Fics. 44~46.—Further successive stages in the development of the spindle.

Fic. 47.—Chromosomes, connected by fibers, being drawn into the equatorial
ar spindle with aster showing at one pole.

—Spindle showing the two poles.
— 5°-59.—Successive stages in the development of the spindle and the
{ing of the chromosomes into the equatorial plane. * Figs. 57-59 on plate XIV.

PLATE XIV
me OO SPindle, showing asters and centrosomes; the chromosomes nearly
m the equatorial plane.

; a 1.—Mother star with aster at the poles.
that the = 61b, 61c, 61d, 6re—Chromosomes on the spindle fibers, showing
Fic, seg loop extends outward. cee
free end. 2.—A chromosome from the mother star of the second division w1
ends of the V proj ecting outward.
four 98 ©3—Polar view of chromosomes, showing the three ring-chromosomes;
ng chromosomes, two of which lie side by side and are very large; and
Fre va chromosomes of various shapes and sizes. roe
three Snag section of mother star, showing the twelve chrom
te Occupy a central position; also the four long chromosomes.

214 BOTANICAL GAZETTE - fwamem

Fic. 65.—A tangential section of a young spindle, showing spindle and con-
necting fibers.

Fic. 66.—Tangential section of a spindle, making a multipolar figure.

Fic. 67.—Section showing tripolar figure.

Fic. 68.—Section showing spindle torn by the knife.

Fic. 69.—Torn section, showing spindle fibers cut and spread apart by the
knife.

Fic. 7o.—Spindle with poles cut away showing two large nucleoli in the cyto-
plasm outside of the spindle.

Fic. 71.—Spindle, showing pole and aster at one end, the other pole being
cut away.

Fic. 72.—Spindle with both poles cut off.

1G. 73.—Another spindle with both poles cut.

Fic. 74.—Spindle showing first stage of metakinesis, the two large chromo-
somes being to one side.

Fic. 75.—Metakinesis stage.

Fic. 76.—First stage of daughter star, showing the separated chromosomes.

Fic. 77.—Daughter star stage.

Fic. 78.—Late daughter stars.

Fic. 79.—Daughter star, showing the twelve small chromosomes. :

Fic. 80.—Loose daughter skein stage, showing the beginning of contraction
of the chromosomes. :

Fic. 81.—Daughter skein, showing the close massing of the chromatin.

Fic. 82.—Daughter skein, showing close massing of the chromosomes below
the pole.

Fic. 83.—Beginning of formation of daughter net, showing the irregular
daughter chromosomes.

Fic. 84.—Further development of the daughter net. 1 wil

Fic. 85.—Resting stage of daughter nucleus, the chromosomes being
evident as irregular masses,

MTANICAL GAZETTE, XLVII PLATE XII

SCHAFFNER on AGAVE

BOTANICAL GAZETTE, XLVII PLATE XIII

SCHAFFNER on AGAVE

| WPANICAL GAZETTE, XLVII PLATE XIV

612 '6y, 62 |

Peat

Pat Ee
Bo ay

SCHAFFNFR on AGAVE

SPERMATOGENESIS IN DIOON EDULE’
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY 125
CHARLES J. CHAMBERLAIN
(WITH PLATES XV-XVIII AND THREE FIGURES)

In March, 1904, and again in September, 1906, the writer visited
Mexico for the purpose of studying the Mexican cycads and collecting
material for morphological work. The field study was greatly facili-
tated by the active cooperation of Governor TEopoRO A. DEHESA,
whose active interest in education has done so much to raise the
educational standard of the State of Vera Cruz. I am also deeply
indebted to Mr. ALEXANDER M. Gaw, of the State Bureau of Infor-
mation, Xalapa, Mexico. Mr. Gaw not only supervised the col-
lection of material and forwarded it to me, but on several occasions
visited localities where Dioon edule grows and sent me valuable field
notes with the collections. The material was collected at Chavar-
tillo, Mexico, the locality which furnished material for my account
of the ovule and female gametophyte of Dioon edule.

THE STAMINATE CONE
In March the ovulate cones which were pollinated the previous
September have reached their mature size, but the staminate cones
from which they were pollinated have disintegrated. Both the ovulate
1 the staminate cones which are to appear a few months later are
still hidden in buds which cannot be distinguished from leaf buds.
Until they reach a length of about 1o°™ the staminate cones are
completely hidden by large bud scales which are fleshy and very
» $0 that the whole structure looks like a moderate-sized ovulate
ane The surface of the cone at this time is densely covered with
lish hairs, which become darker when the cone emerges from the
Protecting scales,
. September the staminate cones reach their mature size and
their pollen. Tust before the pollen is shed, the cone is firm and
reriog ettion prosecuted with the aid of a grant from the Botanical Society of

uns [Botanical Gazette, vol. 47

216 BOTANICAL GAZETTE [MARCH

erect (fig. 1) and measures 10 to 20°™ in length and 7 to 11° in
diameter. At the time of shedding the pollen, the axis of the
cone elongates considerably
and becomes so weak that
it bends over until its tip
rests upon the leaves of the
crown. Insects are very
numerous in nearly all the
mature staminate cones, but
none were found on the ovu-
late cones, and although the
insects were throughly dusted
with pollen, there was nothing
further to indicate that any
pollination was being effected
through their agency.

MICROSPORANGIA

The staminate sporophylls
are long and wedge-shaped
and end in a single sharp
point which curves upW
(fig. 2). The outer, exposed,
abaxial faces of the spor
phylls are densely hairy, but
the upper faces are quite
smooth. In. the sporophylls
of the upper and lower por
tions of the cone, 4 sterile
line divides the sporangia mt
two groups (fig. 2), while 2,
the median portions the sP*
rangia cover the entire undef

-—Upper portion of plant of Dioon

Fic, 1
edule with staminate cone. Photographed

é g
“ Chavarrillo, Mexico, September, 1906. surface. The sporophyllls are
ne-third natural size. a ae i the cone that the
sporangia on the under side of a sporophyll make a distinct
below it.

pression upon the upper side of the sporophyll immediately ‘

90) CHAMBERLAIN—SPERMATOGENESIS IN DIOON 217

The number of sporangia on the larger sporophylls varies from
about 100 to 300; but the smaller sporophylls bear fewer, and the
sporophylls at the apex and base of the cone may bear only a few
sporangia, or even none at all.

The sporangia are grouped in definite sori, usually of four
or five sporangia, as shown in fig. 2. At the stage shown in this figure
the line of dehiscence is quite
obvious. As the sporangia dehisce,
the grouping into sori becomes
less obvious. The pollen does not
escape immediately, but for a short
time is held together in a spherical
mass by the scanty remains of the
hypodermal wall layers. A hand
lens shows that each sorus is sur-
founded by hairs which grow out
fom the peripheral portion of the
base of the sorus. Many hairs also
sow out from the peripheral por-
tions of the sporangia, but there
afenone in the interior of the sorus,
either upon the sporangia or upon
the sporophyll (fig. 4). £ two micro-

he sporangia are either sessile sone :
or h : sporophylls of Dioon edule. X$

ave short massive stalks. ‘The
si and even the lower portions of the sporangia of a sorus may be
cl Seam but there is not so much crowding or S° much
“a shown by Miss Suri (8) for Zamia and Ceratozamia. The

: - im surface view and in section, is about as figured by

— (8) for Zamia and Encephalartos. :

. eg neral appearance of a sporophyll in transverse section,
canes oo vascular bundles, and sori, is shown in jig: 3
which ohn view of two of the sporangia of a sorus is seen 1n fig. 45
; om the usual amount of union at the base of the sorus. The
oo those which come from the sporangia, are olen
ae and crowded that in transverse section they look more like
yma tissue of angular cells than like sections of tubular hairs.

with

218 BOTANICAL GAZETTE [arcu

The hairs do not branch and seldom consist of more than two cells,
which are sometimes empty and sometimes filled with a deeply
staining substance like that in the epidermal cells of the sporangium,

The wall of the sporangium is composed of three distinct regions,
the epidermis, the tapetum, and the intervening wall layers. The
epidermis, which is thicker in the upper half of the sporangium than
in the lower, consists of thick-walled cells with rather scanty proto-
plasm, but with an abundance of suberin and tannin. The tapetum,
in comparison with the size of the sporangium, is very insignificant,
consisting of a single layer of small cells, with occasional patches two
cells in thickness (fig. 5). The portion of the sporangium wall facing
the center of the sorus is noticeably thinner than the wall of the
opposite portions, there being four or five layers of cells between the
tapetum and epidermis in the former case, while in the latter there
may be as many as eight layers (fig. 4). The structure of the sporan-
gium from the epidermis down to the sporogenous tissue is shown in
more detail in fig. 5.

MICROSPORE MOTHER CELLS

The microspore mother cells of Dioon present some peculiarities
which are worthy of mention. Upon becoming dissociated, they
seldom assume the usual spherical contour, but remain more of less
angular, and are nearly always elongated. The chromatin is abun-
dant, but not always well defined, and it is not unusual for the entire
nucleus to stain a dense homogeneous black with iron alum hematox
ylin, as if chromatin had gone into solution in the nuclear _ in
such cases, there are in the cytoplasm irregular masses of sl
_ Staining material which take the spherical form and begin to resemble
nucleoli as the homogeneous staining of the nucleus ceases and the
chromatin becomes definitely outlined. When these spherical 7
Were first observed, an effort-was made to connect them with the
blepharoplast, but it was easily determined that they were formed
by the rounding-off of the irregular masses, and that they at ie
surrounded by radiations. They vary in number and sth eS
this may be true of young blepharoplasts. While the om of th
Masses was not determined absolutely, there is little doubt that they
represent a portion of the deeply staining material which has
from the interior of the nucleus into the cytoplasm.

1909] CHAMBERLAIN—SPERMATOGENESIS IN DIOON 219

The behavior of the chromatin during the two mitoses by which
four spores are formed from the mother. cell will be described at some
future time; at present I merely note that the number of chromo-
somes in both mitoses is 12. A hasty examination might lead one to
suspect that the number is much larger, since it is not difficult to find
prophases of the first mitosis, just before the disappearance of the ~
nuclear membrane, showing any number of chromosomes from 12 to
24. But when there are more than 12, some are always about half the
full size. Any number beyond 12 is due to the early separation of the
two parts of the chromosome, which, in most cases, are separated only
after the chromosomes have become arranged in the equatorial plate.
Even in prophase, there are occasional indications of the second
splitting which is to be completed at the metaphase of the second
mitosis.

These mitoses are not simultaneous throughout the whole sporan-
glum, but begin at the periphery and proceed toward the center, so that
there may be a zone of dividing cells surrounding mother cells which
are still in the resting condition.

POLLEN BEFORE SHEDDING

The young pollen grains are not quite spherical, there being a
flattened portion which might be called the base of the grain (fig. 6).
The exine is thickest in this basal region, while at the opposite end of
the grain where the pollen tube is to emerge, it is very thin. The
intine is thinnest in the basal region where it is in contact with the
thickest portion of the exine, On the sides of the spore, the intine is
very thick, often thicker than the exine. ‘There is no trace of a
third spore Coat, as described by FeRGusON (4) for Pinus.

rhe microspore germinates while still in the sporangium. A single
Persistent Prothallial cell is formed, lenticular in shape and closely
applied to the base of the spore. WEBBER (2) described two prothal-
oe in Zamia, the first formed being evanescent and the second
c but a reexamination by Miss Grace SmirH (8) showed
Encential We find only one prothallial cell in Dioon, Zamia, 6
eh. “ae and it is persistent. In later stages, after the aon
tla, ee to form, it would be easy to misinterpret, for the line
eis th the stalk cell with the persistent prothallial cell often
Impression of a small prothallial cell beneath the large

220 BOTANICAL GAZETTE [MARCH

persistent one (jigs. 10-12). The illusion is emphasized by the fact
that deeply staining granules simulating a broken-down nucleus
are sometimes found at the base of the prothallial cell (fig. 15). It
seems probable that a misinterpretation of this sort was responsible
for the description of an evanescent prothallial cell.

The nucleus in the main body of the grain now divides again, the
mitosis resulting in the formation of a tube cell and a cell which
resembles the prothallial cell and becomes so closely applied to it
that the two look as if they had arisen by the division of the prothallial
cell (jigs. 7-9). This cell, so closely associated with the prothallial
cell, has been called the generative cell. It soon divides, giving rise
to the stalk and body cells (fig. ro). The tube nucleus, even before
the formation of the tube, increases greatly in size, and the cell which
is to form the pollen tube becomes filled with large starch grains.
Late in September the pollen is shed in this three-celled condition.

The output of spores can be estimated with reasonable accuracy
by the formula ¢7R3=the number of spores in a sporangium.
To apply the formula, it is necessary only to count the number of
spores in a line from the center of the sporangium to the tapetum,
substitute this number for R, and then make the calculation. Of
course, this assumes that the mass of spores is spherical and that all
spores develop, both of which assumptions are more or less incorrect
but the error is easily less than the variation in the output of individual
sporangia of average size. In a few cases, spores were actually
counted in a series of sections and the results were practically identical
with the estimates by the formula. In the larger sporangia there a
about 20 spores in a radius, and consequently the output 1s abet
33,507 spores. The largest sporangia, with a radius of 22 spores,
would have an output of 44,600 spores. In some of the s
sporangia, which nevertheless produce good spores, the outp weeps
fall as low as 8000 spores. About 30,000 spores may bet
a typical output for the average sporangium of Dioon edule. :

Late in September or early in October, when the pollen 1 - on
large pollination drop, of the appearance and consistency of glye a
oozes from the micropyle of the ovule. As the pollen passes
the drop into the pollen chamber, at least a portion of the drop beco
brownish and so hard that it adds to the difficulty of sectioning:

1909] CHAMBERLAIN—SPERMATOGENESIS IN DIOON aet

THE POLLEN TUBE

Whether in artificial cultures or in the nucellus of the ovule, the
pollen tube begins to grow at once, emerging from the apex of the
spore and growing out into the sugar solution or into the tissues of the
nucellus. The tubes are irregular in diameter and sometimes have
short branches, but they are nearly straight and lie so close to the
surface that their position is indicated by brown lines radiating from
the beak (jigs. 19, 23). Since the haustorial portion of the tube
teaches a length of 2 or 3™™, the brown lines are easily visible to the
naked eye. As the tube begins to form, the pollen grain end is pushed
into the pollen chamber before the haustorial end has penetrated far
into the tissues of the nucellus. The tube is formed from the intine,
which breaks through the exine and increases greatly in thickness,
as may be seen by noting the comparative thickness of exine and intine
in jigs. 8 and 18. The difference is even greater than is indicated by
the figures, because fig. 8 is more highly magnified than fig. 8. The
tube stains a light brown with iron alum hematoxylin, contrasting
ae with the brilliant red which the exine takes when stained with

Starch is abundant in the pollen tube, and filaments looking like the
Tadiations about the blepharoplast, only much longer, are conspicuous,
“specially in the vicinity of the nucleus.

THE BODY CELL
The division of the generative cell, giving rise to a stalk cell and a
body cell, takes place soon after the pollen is shed, all material col-
lected later than the middle of October showing this division already
completed. 7
In characteristic cycad fashion, the prothallial cell now pushes up
0 the stalk cell (figs. 10, 11, 12, 15, 18). Stages between jigs. 9
é& 70, which might show the cause of this peculiar and remarkably
astant behavior of the prothallial cell, were not available.
es body cell, which is to produce two sperms, does not divide
the or following spring. The division usually takes place about
Week in of April, but may occur a week earlier, or as late as the first
gradual May. During this period of about half a year, there 1s a
Stowth and differentiation of the body cell.

int

222 BOTANICAL GAZETTE [MARCH

At first the cytoplasm of the body cell seems homogeneous,
without any vacuoles or conspicuous granules (fig. 10). The
chromatin and nucleoli of its nucleus stain sharply with iron alum
hematoxylin. Most of the chromatin is in the form of deeply
staining granules.

In a short time it is noticed that the nucleus no longer stains
sharply, the reticulum appearing very faint, and even the nucleolus
and larger chromatin granules taking scarcely any stain. But while
these changes are taking place within the nucleus, many granules,
staining sharply with iron alum hematoxylin and apparently identical
with the chromatin granules, appear in the cytoplasm. For the sake
of reference we may call them the black granules (bg, figs. 13, 14):
I believe that they have come from the nucleus. Whether the granules
pass through the membrane bodily, or become dissolved and a
through by osmosis, might be a question. Living chromatin 1s
semi-fluid and the nuclear membrane at this time is extremely thin.
If the nuclear membrane is formed by the condensation of cytoplasm
about the nuclear vacuole, the “breaking-down” of the membrane
in the prophase of mitosis may be merely the return of the condensed
cytoplasm to the ordinary alveolar condition; and as this condition
approaches, but while the membrane is still recognizable, it is reasoo-
able to suppose that particles may pass from the nucleus to the
cytoplasm without becoming soluble. The black granules migh ;
pass from the nucleus to the cytoplasm in this way. There is litle
doubt that chromatin is more or less soluble. In solution, the
granules could pass by osmosis through a membrane with such &
ete ee as a physiologist might imagine the nuclear membrane '

ave.

The black granules are very small at first and are more —
near the nucleus. They increase in size by imbibing liquid from : de
surrounding cytoplasm, until the granule becomes 4 thin pelli
inclosing a liquid. As the pellicle stretches, granules pass throug
it into the watery interior, and the color with iron alum suet
gradually changes from black to gray. For reference, these glo fe
may be called gray bodies (gb, figs. 14, 17). Both the black gr
and gray bodies are found not only in the body cell, but also
stalk and prothallial cells, and even in the cells of the nucellus.

1909] CHAMBERLAIN—SPERMATOGENESIS IN DIOON 223

THE BLEPHAROPLAST

The origin of the blepharoplast is not easy to determine. In
Zamia, Cycas, and Ginkgo, the blepharoplast, when first recognizable,
isa small sharply staining granule in the cytoplasm of the body cell.
We must admit that the same is true of Dioon, but a study of the black
granules led us to surmise that blepharoplasts, in their origin, are
simply these granules derived from the nucleus. It would follow
that not two but several blepharoplasts might begin to develop. Why
only two should become differentiated is not clear. The blepharo-
plast, in all the early stages of its development, takes an intense homo-
geneous black color with iron alum hematoxylin, never behaving like
the gray bodies. Sometimes the cytoplasm about one or more black
granules becomes dense and homogeneous, quite unlike the usual
alveolar structure, and resembling the archoplasm which surrounds
young centrosomes (fig. 13). It is possible that blepharoplasts may
begin their growth in this way.

After enlarging considerably, two blepharoplasts become unmis-
eo through their influence upon the surrounding cytoplasm,
which takes on a radiate arrangement with the blepharoplast as a
ecsoe (figs. 15, 16). At first, the radiations are nothing more than
the intersections of alveoli (fig. 16), but as the walls of the alveoli
Pecome less distinct, the radiations become definite granular filaments,
sending from the blepharoplast to the periphery of the cell. Many
of the filaments are simple, but branching is very common (figs. 77; 22).
: The appearance of the filaments in preparations indicates a stream-
ws Movement, especially toward the blepharoplast. That the
a ig in sections are streams of cytoplasm is indicated by
where th a ny similar structures are found in the pollen nae
a “re Js certainly a strong streaming movement. An 2088
by —. that the filaments are streams of cytoplasm is furnis s
its ae S In cutting out the top of the female gametophyte x
Of the Sonia, the least pressure will cause some of the cytoplasm
This — = be squeezed out through the necks of the archegonia.
arch oes cytoplasm, streaming out through the neck of the
o.: shows wey numerous filaments with a structure iden-
2 ene at of the radiations about the blepharoplast, and in case

artifact there is no doubt that the filaments are nothing but

224 BOTANICAL GAZETTE (MaRcR

streams of the egg cytoplasm. In many cases the connection of the
radiations with the blepharoplast also indicates a streaming (fig. 21).

Soon after the stages shown in figs. 16 and 17, the filaments appear
very granular, some of the granules being almost certainly the black
granules. The gray bodies become attached to the filaments and
give the radiations about the blepharoplast a striking appearance
(figs. 18, 20, 21, 31). The general topography of the pollen tubes, pol-
len chamber, and nucellus at this time is shown in fig. 19.

The watery gray body runs along the filament, usually in both
directions, so that it becomes spindle-shaped, but often it spreads
only in one direction, and consequently becomes top-shaped. As the
gray bodies spread along the filament, depositing granular matter and
giving up their watery content, the filaments become smoothly and
sharply defined and have much greater density (jigs. 21, 24). hig
growth of the blepharoplast is due, in great measure, to the acquisition
of granules, and perhaps other matter, brought to it by the streaming
filaments. |

During the early stages of its growth, the body cell elongates and
the two blepharoplasts with their conspicuous radiations lie in the
plane of the long axis, one above and the other below the nucleus (jig.
18). The pollen tube, at this time, is very narrow, and this fact may
account for the elongation of the body cell, which fills nearly the entire
diameter of the tube. _

In March the pollen tube has become very large, especially ~
free end of it, which projects into the pollen chamber, and with this
increase in the diameter of the tube the body cell changes from #*
elongated to a nearly spherical form, the blepharoplasts at the ~~
time rotating go°, so that they become transverse to the long 515
the tube (fig. 25).

Even before assuming the transverse orientation, the a
Plasts may begin to show vacuolation (fig. 22), but after the se
verse orientation has become established, the vacuoles Be -:
large and so numerous that they occupy nearly the entire body in
blepharoplast (jig. 25). They are scarcely affected by eee
nearly all preparations showing a dirty-white or pale-yellowish tain
The ground substance of the blepharoplast still continues =e
black with iron alum hematoxylin, or red with safranin-

1909] CHAMBERLAIN—SPERMATOGENESIS IN DIOON 225

The blepharoplast, when nearly mature, is spherical and measures
16 to 18m in diameter. Just before breaking up into granules, it
becomes somewhat elliptical in section, with its longer axis parallel
with the longitudinal axis of the body cell. The longer axis of the
blepharoplast then measures about 20 m.

THE SPERMS

The body cell divides longitudinally, giving rise to two sperm
mother cells (fig. 26a). At about the time of this division, the
blepharoplast breaks up into a large number of granules, which at
first occupy the elliptical area of the blepharoplast. The granules are
derived not only from the rim of the blepharoplast but from the por-
tions between adjacent vacuoles. It is possible that granules may
also be formed from the radiations, for these begin to disappear at
this time. The area of granules soon becomes elongated, and the
spiral band begins to appear (figs. 26, 26a). By this time the pollen
chamber has extended until it has destroyed all that part of the nucellus
lying above the archegonial chamber, so that there is no obstruc-
ton between the ends of the pollen tubes and the necks of the arche-
gona (cf. figs. 19 and 2 3). If the nucellus be removed, the numerous
Pollen tubes protruding from the pollen chamber are easily visible
tothe naked eye, and a little later the sperms may be observed without
Noo the aid of a pocket lens (figs. 27). In this figure, the star-shaped
afea 1s a portion of the tissue of the nucellus, exposed by the rupturing
of the megaspore membrane. The evenly dotted portion represents
the megaspore membrane, which in this region adheres to the nucellus
Tather than to the female gametophyte.

As we have said, the spiral band begins to appear as soon as the
area of granules elongates. The band is closely applied to the nucleus
shown in fig. 26. The nuclear membrane is very weak in this
Tegion, and the nuclear structure indicates a movement of material
foward the point of attachment. The connection between the nucleus
oa the band is maintained, even after the band has come to te
BS fe (fig. 28). In the mature sperm the band is 4 spiral of oa
t ‘ix turns, the direction being, almost without exception, from /¢
° Nght, as viewed from above. The radiations, which were SO

226 BOTANICAL GAZETTE [marc

conspicuous during the growth of the blepharoplast, disappear as
the blepharoplast breaks up into granules.

The two sperms are formed within the two cells resulting from the
division of the body cell. That this is the case is readily seen in
sections (jigs. 28, 29, 32). The relations are particularly clear in
fig. 29, in which ¢ is the pollen tube, and w the wall of the sperm
mother cell. The figure shows the apex of the sperm with a small
portion of its large nucleus still surrounded by the mother cell. The
two sperms within their mother cells are shown in the photomicro-
graph, fig. 32.

That the sperms are formed within mother cells is also clearly seen
in living material, where the mother cells enlarge considerably after
the sperms are ready to move. The peripheral portion of the partition
between the two sperms breaks down, thus allowing the sperms to

“Move about within the old body cell. At this stage the term body
cell is not strictly correct, because the cavity now consists of the com-
bined areas of the two sperm mother cells. Since, however, the outer
wall is still the wall of the original body cell and there is no name for
the new cavity, we may refer to it as the old body cell. The wall of
the old body cell soon breaks down and the sperms escape into the
main portion of the tube.

Sperms within the pollen tube measure about 200 # in diameter
and about 275 # from apex to base. After leaving the tube, they
increase somewhat in size, reaching a diameter of 230 # and 4 length
of 300 w. Consequently, they are easily visible to the naked eye.

The living sperm, as seen under the microscope, has a large gran-
ular nucleus, surrounded by a thin and almost colorless sheath of
cytoplasm, which is somewhat thicker at the spiral end. The nucleus
usually shows a large depression just beneath the apex of the sperm
(fig. 32).

The movements of the sperms are easily observed
mounting a piece of the nucellus with its pollen tubes. re
to the lighting from above, some light may be reflected up through
pollen chamber. The upturned ends of the pollen tubes (fig 27)
so transparent that they scarcely obscure the view. The cilia so
to move while the sperms are still fast together and more OF ©

attached to the stalk cell. The movement of cilia is accom

by simply
In addition

1909] CHAMBERLAIN—SPERMATOGENESIS IN DIOON 227

by pulsating and amoeboid movements which continue for an hour
or more before the sperms separate. After the separation, they may
swim for half an hour or more in the old body cell before they escape
into the general cavity of the tube. Occasionally, the sperms are
still attached to each other after they have escaped into the tube, and
in such cases their movements are awkward, because they naturally
try to move in opposite directions. When free from each other the
principal movement is straight ahead, with a rotation on the long
axis. The sperms swim up and down in the tube, going up as far as
the diameter of the tube will permit, and then coming back. The
amoeboid movements both of the cytoplasm and the nucleus are quite
noticeable, especially while the sperm is changing its direction. At
the apex, where the cytoplasmic sheath is thickest, the amoeboid
movement is most conspicuous, and may be so rapid that it is more
like a twitching. How long the sperms might swim in the pollen tubes
under natural conditions, one could hardly guess. When a nucellus
Is inverted in a drop of sugar solution on a slide, and is still further
protected by a bell jar, the movements have continued for five hours.

After the sperms begin to move, there is a rapid increase in the
turgidity of the tube, which sooner or later ruptures at or near the
fxine of the pollen grain. Most of the starch and liquid contents
of the tube escapes with a spurt, unless one of the sperms is immedi-
ately drawn into the opening. The first sperm may escape in two or
three seconds, but the other may be half a minute in getting out,
Probably because there is not so much pressure behind it. The
Tupture is often not more than 50 » in diameter, while the average
Sperm is four times as broad. But however much the sperm may be
a in getting out, it promptly regains its form and begins to

Sg to keep the sperms alive after their escape from the pollen
diate = me very successful. In weak sugar solutions they imme-
shat 24 pieces, almost explode. In a 1o per cent. sugar
unti] “eh quickly die. Sugar was added to a Io per cent. solution
sperms sti perhaps, a 12 or 15 per cent. solution, and in this the
continued to swim for several minutes. No experiments were

mace to determine whether the sperms are chemotactic or not.
The material would have allowed a more detailed account of the

228 BOTANICAL GAZETTE [MARCH

later history of the blepharoplast and the development of the sperm,
but these features, particularly the relation of the blepharoplast to
the spiral band, are shown so much more clearly in my preparations
of Ceratozamia, that I have refrained from any extended description
at this time.

The further history of the sperm of Dioon edule will be considered
in a forthcoming paper on fertilization and embryogeny. '

DISCUSSION

With TurEssen’s (14) paper on the seedling of Dioon edule, my
own paper on the ovule and female gametophyte, the present paper _
on spermatogenesis, and a study of fertilization and embryogeny
now nearly completed, considerable information concerning this
form is available. The temptation to draw conclusions is strong,
but studies on Ceratozamia mexicana and Dioon spinulosum ate well
under way, and Sister HELEN ANGELA (12) has already completed
an investigation of the anatomy of Ceratozamia. Since any theories
will be more likely to be well founded if based upon a comparative
study, I shall reserve speculations for a safer opportunity. At pee
I shall merely consider a few points suggested by the foregoing de-
_ scription. :

The largest staminate sporophylls in the Cycadales are found bg
Cycas and the smallest in Zamia. The number of sporangia weil
sponds, roughly, to the size of the sporophyll. The sporangi rs
Cycas, Encephalartos, Dioon, and all the forms with large ma
sporophylls, so far as I have been able to examine them, have the
_ Sporangia in definite sori with 3, 4, or 5 sporangia in a Sorus:
forms with smaller sporophylls, like Zamia, Ceratozamia, and Micro-
cycas, some of the sporangia are single, but most of them are on
with 2 or 3, or occasionally 4 sporangia in a sorus. CALDWELL 9
states that in Microcycas the sporangia are not arranged 10 oe
but his photograph shows that even on the smaller sporophyll g
of the sporangia are in sori; the number of sporangia is about
on the larger sporophyll. The number of sporangia on 4 pot
according to Miss Grace Suri (8) is as follows: Cyeas 0”
700, Encephalartos Caffer 700, Macrozamia Miquelit 600, Encep We
lartos villosus 500, Dioon edule 200, and Zamia floridana 24-

1909] CHAMBERLAIN—SPERMATOGENESIS IN DIOON 229

can now add that the number in Dioon edule often reaches 300, the
usual number in Ceratozamia mexicana is 250 to 300, and in Stangeria
paradoxa, about 260. In both Ceratozamia and Stangeria the sporan-
gia are very small.

If this is a reduction series, as we believe it is, Microcycas, so far
as this character is concerned, has scant claim to the position of
“the most primitive cycad yet described,” especially since LAND (10)
from a consideration of the female gametophyte, and Sister HELEN
ANGELA (13) from an investigation of the vascular anatomy, have
concluded that Microcycas is far from primitive. As far as micro-
sporophylls and microsporangia are concerned, Dioon is nearer the
Cycas condition than any other western cycad.

The output of spores is very easily estimated. BoweEr’s studies
on spore-producing members have shown that the more primitive
ferns have a large output, which is gradually reduced as we pass to
the highly specialized recent forms. We believe the same is true of
cycads. The output in Dioon edule is about 30,000 spores per Spo-
rangium. According to Miss GRACE SMITH (8) the output in Encepha-
lartos villosus is 26,000, in Ceratozamia mexicana 8000, and in Zamia
idana 500 to 600. It would be interesting to know the output in the
other cycads. I should expect an output of 30,000 or more in Cycas,
and should be surprised if the output in Microcycas reached 20,000.
Judging from CatpweELt’s (9) account, I should look for an output
of about 10,000 spores.

In our study of the spore coats, I looked for the third coat described
by FeRcuson (4). The fact that only an intine and an exine are
present in Dioon led me to reexamine Pinus Laricio. 1 found only
re exine and an intine, as in Dioon, the third coat described by Miss
FRGUSON (4) being merely the usual intine which her excellent
aa ¢ had sharply differentiated from the exine. As ..
‘ “ee her own figures indicate, the intine, which she mistakes tor
Coat, grows out to form the pollen tube. >
me blepharoplast in Dioon edule is probably of nuclear origin.
ey (1) was inclined to believe that the blepharoplasts of Cycas
wa came from the nucleus, although when first recognizable they
gr bodies just outside the nuclear membrane. Wannee (2)
at in Zamia the blepharoplasts originate de nove the cyto-

230 BOTANICAL GAZETTE [MARCH

plasm of the body cell, but no convincing early stages are given.
CALDWELL’s (g) material was all too far advanced to show the origin
of the blepharoplasts. ;

It seems probable that the manner in which the spiral band is
formed from the blepharoplast is similar, in its main features, in all
the cycads. The solid blepharoplast becomes vacuolated and then
breaks up into a group of granules from which the ciliated band is
formed. CALDWELL (g) describes in Microcycas a band, already dis-
tinct during the division of the body cell, and says that this band
becomes broken up into fragments upon which the beginnings of
cilia may be seen. His fig. 25 indicates that the band is a sec-
tion of the rim of the much vacuolated blepharoplast, while the
“fragments” in his fig. 27 are sections of the spiral band, which has
already made several turns. The cilia which he figures on the inside
of the fragments need confirmation.

The origin of the blepharoplast in pteridophytes has been con
sidered by several investigators, all of whom agree that it first appears
in the cytoplasm. Some find it present even from the early sper-
matogenous divisions, while others find it first in the cell which is to
give rise to two sperm mother cells, or, occasionally, one generation
earlier than this. In a very detailed account of spermatogenesis ™
Nephrodium, Yamanovucut (11) finds that two blepharoplasts first
appear in the cell which is to give rise to two sperm mother cells.
The blepharoplast in pteridophytes simply elongates and forms the
band directly, there being no radiations, no vacuolation, nor break-
ing-up into a group of granules which subsequently give rise toa spiral
band. While the blepharoplasts of ferns and cycads are doubtless
homologous structures, no intermediate conditions have yet been f
which would explain the behavior of the blepharoplasts of cycads.

In Dioon edule, as in nearly all gymnosperms, only two 5 cp
are formed in the pollen tube. In a few instances I have noted four
sperms in the pollen tube of Ceratozamia mexicana. JUEL (3) fo
four to twenty sperms in the pollen tube of Cupressus es
naturally regarded the condition as primitive. CALDWELL (9) f of
sixteen or twenty sperms in icrocycas calocoma, and on the wee
this character claimed Microcycas to be the most primitive ©
yet described.

1909] CHAMBERLAIN—SPERMATOGENESIS IN DIOON 231

That the two sperms represent a reduction from a larger number
is so evident that there is no need for discussion, but a word in regard
to the so-called “body cell” from which they are produced may not
be out of place. In the cycads, in Ginkgo, and in Coniferales the
cell whose division gives rise to the “stalk cell” and body cell is
often called the generative cell, probably because the stalk cell is
regarded as a spermatogenous cell which has ceased to function.
In Cupressus, Microcycas, and Ceratozamia all the sperms come
from the body cell, the stalk cell being entirely sterile. It might be

i Fic. 33.—Diagram illustrating the homologies between the parts of an ordinary
— fern antheridium and the pollen tube structures of a cycad: A and C, the
antheridium, in C the dome cell being represented as elongated into a tube; B and

: .
toga Structures: , prothallial cell; s, stalk cell; sp, spermatogenous cell; 4,

ei ested that the stalk cell corresponds to the basal cell of the
a fern antheridium and that the dome cell corresponds to the
Fed woes hile the primary spermatogenous cell corresponds to the
te cell. If the dome cell, without forming the usual cover cell,
‘ ould become elongated, we should have very much the same situa-
on as that found in Cupressus Goweniana, Microcycas calocoma, and

cc mexicana. The diagram (fig. 33) indicates these
in . es in an ordinary polypod fern and a cycad. The fact that
the “tern the dome cell is sister to the spermatogenous cell, while in
“Yad the stalk and spermatogenous cells are sisters, might be

232 BOTANICAL GAZETTE [MARCH

regarded as an objection by those who lay great stress upon the
importance of a rigid sequence of cell divisions.

I have shown that in Dioon edule the sperms are formed within
sperm mother cells from which they are subsequently discharged.
IKENO (1) does not state whether the sperms of Cycas are formed
within mother cells or not. WEBBER (2) claims that in Zamia they
are not formed within mother cells, but that the two cells resulting
from the division of the body cell become ciliated. CALDWELL (9)
does not mention this feature in his account of Microcycas and his
figures are noncommittal. MrvaKe (6) saw the sperms within the
body cell in Ginkgo, but could not make a definite statement for
Cycas. I have examined Zamia and find that the sperms are orgal-
ized within definite sperm mother cells, and have found the same
situation in Ceratozamia. This condition is probably general in
cycads. :

In bryophytes the final division of the spermatogenous cells results
in conspicuous pairs of sperm mother cells, or perhaps, as has bees
claimed, two sperms are formed in a single mother cell. In pterido-
phytes the pairs are not so conspicuous, but the feature is just as def
nitely present. In nearly all gymnosperms the spermatogenous
tissue has become reduced to a single pair of. spermatogenous cells,
which in some cases, as in Juniperus, bear a striking resemblance 10
the sperm mother cells of cycads. In Juniperus, however, no vests?
of a blepharoplast has been reported, and it is assumed that the
mother cells function directly as sperms. In forms like Pinus, there
is merely a nuclear division within the body cell, giving rise 10 ka
sperm nuclei, no sperm mother cells being formed. In Pinus there
are structures which may be vestiges of blepharoplasts. It cong
interesting to know whether a cytological study of some form of
Juniperus or Thuja would show non-ciliated sperms with vestig®
blepharoplasts within mother cells. ed

Enough is now known of the sperms of the four genera of 0 cen 1
cycads to identify the genera by this character alone. A comparatt
study will be made later, but a few features will be mentioned we
In Cycas the spiral makes five and a half to six turns, the direch®
being from right to left, as viewed from above. In Zamia, accom
to WenBer (2), there are five to six turns, always from left t¢ right.

1909] CHAMBERLAIN—SPERMATOGENESIS IN DIOON 233

In Dioon also, the direction is from left to right. In Microcycas
CALDWELL (9) figures three cases, one from right to left, and the
direction of the other two uncertain. What causes the direction of
spiral is not known.
_ Zamia has the largest sperm yet described, measuring 222 to 306 ps
in diameter and 222 to 332 “in length. In Cycas the diameter is 180
to 2104. In Dioon edule the living sperms measure about 230 #
in diameter and 300 » in length. In microtome sections, the sperms
are smaller, perhaps on account of plasmolysis, but more probably
because the sperms increase in size after leaving the tube. In section
the sperms measure about 200 » in diameter and 275 in length.
CALDWELL (9) gives no measurements, but, judging from his figures,
the sperms are comparatively small, with a diameter of about 60 p.

In all the cycads which have been studied, the movements of the
sperms are very similar, a forward movement with a rotation upon
the long axis. While experiments have been made, no chemotaxis
has yet been noted. When the sperms are mature, the neck canal cells
are very large and turgid. It may be that they exert a chemotactic
influence.

If the foregoing account seems to consist rather largely of data, I
can only say that I prefer to reserve any discussion of phylogeny
until the investigations in which I am already engaged shall have
been completed.

SUMMARY

The sporophylls of the staminate cone are rather large, and bear
about 250 sporangia, a larger number than in any cycads except
Cycas, Encephalartos, and Macrozamia. The output of spores per

Tangium is about 30,000, a larger output than in Zamia, Cera-
‘ozamia, or Encephalartos, the only genera in which this feature has
been noted.

There are twelve chromosomes in the pollen mother cell, but they
often split so early that the number may appear larger-

ere is only one prothallial cell and that is persistent. The
"eport of an evanescent prothallial cell in Zamia is probably due to a

misinterpretation.
: adiong Ph OPlast are very probably of nuclear wa The
are streams of cytoplasm, which, in early stages, have 4

234 BOTANICAL GAZETTE - [MaRcH

peculiar appearance on account of the granules and globules which
adhere to them. The solid blepharoplast becomes vacuolated and
then breaks up into granules from which the spiral band is formed.
The ciliated band makes five or six turns from left to right. The
sperms are larger than those of Cycas or Microcycas, but not quite
so large as those of Zamia.

The sperms are formed within sperm mother cells, from which they
are discharged. The same is true of Zamia, Ceratozamia, and
probably of other cycads. 3

In addition to the movement by cilia, there is a vigorous amoeboid
movement of both nucleus and cytoplasm.

Discussion of phylogeny will be reserved until investigations now
in progress have been completed.

THE UNIVERSITY oF CHICAGO

LITERATURE CITED
1. IKENO, S., Untersuchungen iiber die Entwickelung der Geschlechtsorgane
und die Vorgang der Befruchtung bei Cycas revoluta. Jahrb. Wiss.
32:557-602. pls. 8-10. 1808.
- WesseER, H. J., Spermatogenesis and fecundation of Zamia. U. S. Sp
Agric. Bureau Pl. Ind. Bull. No. 2. pp. 1-100. pls. 1-7. 190l. |
- Juet, H. O., Ueber den Pollenschlauch von Cupressus. Flora 93: 36-2.
pls. 3. 1904. ee "
4. FERGUSON, MARGARET C., Contributions to the knowledge of the life history
of Pinus. Proc. Wash. Acad. Sci. 6: 1-202. pls. I-24. 1904. Deutsch
- Mrvaxg, K., Ueber die Spermatozoiden von Cycas revoluta. Ber.
Bot. Gesells. 24:78-83. pl. 6. 1906. 80.
6. ——, The spermatozoid of Ginkgo. Jour. Appl. Micros. §:1173-178
1906,
CHAMBERLAIN, C. J., The ovule and female gametophyte of Dioon.
GAZETTE 42: 321-358. pls. 13, 14. 1906. f the
- SirH, Frances Grace, Morphology of the trunk and development °
microsporangium of cycads. Bor. GAZETTE 43:187-204. pls. 10. 1997 re
9. CALDWELL, QO. W., Microcycas calocoma. Bot. GAZETTE —
bls. 10-13. 1907. . |
Io. Ail W. J. G., Fertilization and embryogeny of Ephedra ne a
AZETTE 44:273-292. pls. 20-22. 10907. F 3
II. YAMANOUCHT, Sy Spermatogenesis, ss and fertilization 10 Neo
dium. Bor. Gazerrr 45:145-175. pls. 6-8. 1908. ig. Bot.
- Dorety, Sister HELEN ANGELA, The seedling of Ceratozami
GAZETTE 46: 205-220, pls. 12-16. 1908.

eet 3:

1909] CHAMBERLAIN—SPERMATOGENESIS IN DIOON 235

13. DoreTy, SisteER HELEN ANGELA, Vascular anatomy of the seedling o
Microcycas calocoma. Bot. GAZETTE 47:139-147. pls. 5, 6. 1908.

14. THIEssEN, R., The vascular anatomy of the seedling of Dioon edule. Bor.
GAZETTE 46: 357-380. pls. 23-29. 1908.

EXPLANATION OF PLATES XV-XVIII
Fics. 1, 2.—In text.
PLATE XV

Fic. 3.—Transverse section of a young microsporophyll, showing sori, 5,
mucilage ducts, m, and vascular bundles, b. X10.

Fic. 4—Two microsporangia of sorus. X55.

Fic. 5.—Section of a part of the microsporangium, showing epidermis, wall
layers, tapetum, and some sporogenous tissue. 250.

Fic. 6.—Microspore, Aug. 14, 1905. 1260.

Fic. 7-—Microspore showing the * rothallial cell, p, generative cell, g, and
tube cell, #. X 1260.
«Fa. 8.—The pollen tube is beginning to form. X 1260.

Fic. 9.—Pollen tube somewhat later. Oct. I, 1907. X1260.

Fic. 10.—The generative cell has divided to form the stalk cell, s, and body
cell, b. Oct. 21, 1907. X 1000

Fic. 11.—A later stage; black granules and gray bodies in the cytoplasm of
the body cell. X 1000

Fic. 12,—Nearly the same stage as jig. II.

Mic. 13. —Body cell showing black granules. Oct: 23, 1907. X 1890.
i 14.—Body cell with black granules and gray bodies. Oct. 23, 1997:

eee 15.—Beginning of radiations about the blepharoplasts. Oct. 30, 19°7-
5.
Fic. 16.—The blepharoplasts and radiations of the previous figures. X 1899.

PLATE XVI
me 2 17. Sagar with radiations and gray bodies, gd; the figure shows
veola.

Fic. 8 Pollen tube structures ‘with oaly aig on the radiations sur-

“
Po 21-—Later stage of inh ea and radiations. 1890.
“iy 22 —Still later stage,
len ete "View of nucellus oe eke the division of the body cell; the pol-
T has extended entirely through the nucellus. X°8.

236 BOTANICAL GAZETTE

PLATE XVII

cla 24.—Body cell with two blepharoplasts. 945.
25.—Pollen tube structures after the transverse orientation of the
aa May 5, 1906. 237
Fic. 26.—Beginning of the oa band. X18
Fic. 26a.—Topography of pollen tube atvuctures at the stage shown in jig. 26.
Fic. 27.—View of nucellus with pollen tubes at the stages shown in jigs.
26-29. X8.
Fic. 28.—Connection of the nucleus with the spiral band. X945.
Fic. 29.—Apex of a sperm, showing that the sperm is organized within @ :
mother cell: ¢, pollen tube; w, wall of sperm mother cell. 800. F
Fic. 30.—Transverse section of spiral band. X45.

PLATE XVIII

Fic. 31 Fe ecroerach of pollen tube showing blepharoplast vit
on the radiations. 800.
feng 32.—Photomicrograph showing that the sperms are formed within
X 800, te

Fic. 33—-Th text.

A ASS ES IEDR pases eaten

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CHAMBERLAIN on DIOON

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CHAMBERLAIN on DIOON

PLATE XVIII

BOTANICAL GAZETTE, XLVII

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CHAMBERLAIN on DIOON

BRIEFER ARTICLES

THE MOUNTING OF ALGAE
In a personal communication from the well-known phycologist Pro-
fessor G. S. West, of the University of Birmingham, England, I received
an outline of a method of fixing, mounting, and preserving algae which,
as he tells me, has not been given the attention that it perhaps deserves.
The fluid used serves at the same time as a killing, fixing, preserving, and
mounting medium, and for delicate structures like desmids and other algal
forms, it perhaps cannot be surpassed. It has the advantage, moreover,
that it keeps the natural coloring of the green algae, something which the
instructor in elementary laboratory work will appreciate better than anyone
else. The fluid is a 2 per cent. solution of potassium acetate, just made
blue with a small amount of copper acetate. The substance reduces
plasmolysis of the cell contents to a minimum. The algae can be put into
the solution and kept init. Ifa permanent mount is wanted a small amount
of the material is put on a rather thick slide and sealed with old gold size
several times after each drying. The mounts are permanent, but it is
Usually necessary to take great care in sealing, and to this end to use a thick
‘lide. A thin slide will bend considerably in handling, and the sealing
may be separated in this way from the slide, so that the preparation will
dry up as the result.
_ For some reason the fluid presents considerable difficulty with Vauche-
eva Plasmolysis is hard to avoid. I have found that the best way
oo. especially the zoospores before or just after germination,
Sg plant is particularly delicate, is to kill it rapidly with 3 OF 4 a
: - ormalin, : The formalin must be completely and quickly remove
‘ Preparation will turn black afterward. Fixing for half an ciaad 2
i 3 Per cent. formalin will not be injurious. Remove the formalin by
so, Mashing with water. If the Vaucheria thus treated is rapidly
‘ought into glycerin to which a little thymol is added, the preparation

abel as perfectly green as when alive, and will retain its green color
tely. The method may be extended to all small green forms like

the .
smaller liverworts, fern prothallia, and moss protonemata.

5 my ® Set the material into glycerin, add first a considerable quantity of
2 *° Per cent. glycerin in water, and put the dish near but not on a
[Botanical Gazette, vol. 47

238 BOTANICAL GAZETTE [MARCH

radiator. In a few days the evaporation will leave the fluid thick. Once
the preparation is in thick glycerin the color will not change, if the formalin
has been completely removed.

The potassium-copper-acetate solution will not keep the natural color
of diatoms. It has the property of removing the diatomin or yellow color-
ing matter from the diatoms and leaving the plants perfectly green. The
solution can thus be used in demonstrating the presence of chlorophyll in
these plants. The diatomin is removed or absorbed in a few minutes after
application.

As I have found some difficulty in keeping the microscopic mounts
made in the potassium-copper-acetate solution because of drying, I have
evolved a modification of the glycerin method in combination with it. The
mounts made by this method are perfectly durable, and when carefully
prepared are superior to ordinary glycerin mounts, as all green algae treated
with it keep their natural colors indefinitely. Glycerin jelly can also be
used at the end to make the mount even more durable than the ordinary
glycerin mount would be. The procedure is as follows.

The algae to be used are fixed in the potassium-copper-acetate 2 pe
cent. solution, After they have been killed and fixed in this fluid (the
time varying according to the specimen treated), add to the above solution
an equal part of ro per cent. glycerin solution and allow to concentrate
by evaporation in a warm dry place protected from dust. The algae must
be thoroughly separated from dirt and soil or the concentrated solution
will precipitate a reddish-brown cloud of reduced copper. In nearly @
cases the preparation when thickened will be covered with a film of acetat®
which can be removed from the top of the fluid without injury to the oye
rial. The concentrated solution should be perfectly clear, of a light gree
color, and the chromatophores of the algae as perfect a green peste?
I have often been asked by students, and in fact by those well acq =
with algae, whether the plants thus given them for examination wand
really alive. The advantage of having plant material, espech y for -
mentary students, in a condition as near as possible to the live ae see
explanations about stains. I have found it very undesirable to giV* Tie
ners any material other than alive or such as looks like the live stag®
plant studied.—J. A. Nreuwianp, University of Notre Dame, Int

1900] , BRIEFER ARTICLES 239

PAUL HENNINGS
(WITH PORTRAIT)

Professor PAuL HENNINGS, the well-known mycologist, died after a
short illness on October 14, 1908.

In the botanical circles of Berlin he was a welcome and esteemed per-
sonage, having won the sympathy of his colleagues by his extensive learning
as well as by his kindly and un-
assuming nature. He was a dis-
tinguished collector and preparator,
an authority on the world’s fungi,
a faithful and conscientious official
of the Museum, and last, but not
least, a gifted and humorous dialect

As his personality and his whole
hature were made up of a multi-
tude of contradictions, understood
only by those who knew him inti-
mately, so the course of his de-

unusual man, who was, in the
best sense of the word, an original.

: PauL HENNINGS was born on :
November 27, 184r, in Heide, Dithmarsischen, Holstein. He grew up in
Provincial Surroundings, attending the gymnasium at Meldorf until circum-
stances compelled him in 1860 to give up the scientific career to which oof
aspired, and to leave school when only a third-form boy.

He became an assistant in the Kiel botanical gardens and soon an
acknowledged authority on the endemic flora. Professor cae re
lime director of the gardens, gave much attention to the aspiring young
man, and ever afterward looked out for his interests in the most fatherly
manner,

_ Urged by his older countryman, the Low-German poet KLAvs pie
mith whom he was always on the most friendly terms, he was goer
n Kiel in the winter semester of 1863-1864. The breaking-out of the war
1864 obliged him to give up his work in Kiel, and he secured an official
PoSition in the post-office at Augustenburg. After many changes of residence

240 BOTANICAL GAZETTE [uance

he was transferred to Hohenwestedt where he remained until 1874. His
official work was repugnant to him, and during this whole time he remained
faithful to his love of science, teaching in the agricultural school of Hohen-
westedt and soon taking a prominent position as a collector. In addition
to all this work he began to issue not only his herbaria for agricultural
purposes, but also the first hundred of his comprehensive seed collections.

In 1874, he was called by ErcHLER, who at that time was director of the
botanical garden at Kiel, to be his assistant. Here he put in order the
Lucas herbarium and devoted himself with great zeal to the cryptogamous
herbarium. EICHLER was called to Berlin in 1879, and in 1880 he invited
Professor HENNINGS to join him, and confided to him the arrangement of
the newly established exhibition museum of the cryptogamous herbarium.
While doing this he was also busy in the gardens. His power of application
made it possible for him to complete speedily the work assigned to him.

From about 1885 he devoted himself almost exclusively to fungi. Itis
true that during this period he issued two fascicles of the algae of the Mark
Brandenburg, but his interest centered in the mushrooms of this region,
and later, when the museum received abundant collections from tropical
regions, he devoted himself to the fungi of the whole world. His fine feeling
for form enabled him in a short time to become an authority in all systematic
questions regarding fungi. When in 1890 he was appointed assistant
Custos, and in 1891 Custos of the Botanical Gardens, he had already brought
together in Berlin one of the best collections of fungi in the world.

In 1902, as a well-deserved recognition of his work, he was appointed
royal professor. Until his death he continued indefatigably at his work,
the division of the fungi assigned him in the great museum. as

Twelve months ago the death of his son paralyzed his energie =
stole the pen from the busy hand.

Henwics in his special domain was self-taught, and his entire
must be judged from this point of view. He possessed a fine sense 0°"
which made it possible for him at once to put every newly discovered sper
in the right place in the system. By this his work was greatly facie
and this explains his easy command, not only of the fungi of the we
but also of tropical regions. He published in twenty years 25° ee
which dealt with the fungi of innumerable tropical regions. He = i
specialty of the mushrooms of the German colonies and of Braz =
dominated the difficult domain of the Hymenomycetes in @ masterly ma
Berlin, that he discovered many unexpected treasures even at the gee?

erlin
etimes almost

particu

activity
of form,

Few except those who stood near to this reserved—som
repellent—man, suspected that he had a really childlike soul, one

1909] BRIEFER ARTICLES 241

larly responsive to lyric poetry and to the dialect of his home. He wrote
many humorous poems, revealing a rich poetical power, a deep comprehen-
sion of life, and a faithful devotion to his home.

He showed rare courteousness to his friends and even with strangers
he was not parsimonious of his great knowledge. Helpful, modest, retiring,
aman of the old stamp, of the right sort, has passed away with this scholar.
Honor to his memory!—Translated from the German of Linpav, by
J. Perkins, Berlin.

PURE CULTURES OF FUNGI

The Association internationale des botanistes, founded some years ago,
has an office where pure cultures of fungi can be obtained either in exchange
for others, or on payment. Although this fact is probably not unknown
to the readers of this journal, we wish to remind them of it and state its
exact purpose, trusting that more use will be made of the office than has
been the case hitherto.

This office proposes to become a living register of the described fungi.
Large numbers of species are mentioned in the handbooks, which are said
to be insufficiently described and cannot possibly be identified. The
number of identical species described under different names is immense.
This evil may be avoided in future if every mycologist, when describing
a new fungus, will send a culture to the office of the Association. The
author not only is thereby relieved of its cultivation, but everyone who
is studying kindred species may procure material for comparison.

Rather frequent applications are made to the office, but the collection
does not grow in proportion to the description of new species. It has often
happened that upon requesting a person to send us a culture of a certain
Tecently described fungus, the author is obliged to reply that the cultures
have been lost. Who can be sure ever to find again his fungus ? The
little trouble of sending it to the office, however, would have saved the origi-
nal material to posterity.

But the office does not desire the new species only.

.. “cceptable of which you have pure cultures and which are not men

= ~~ list, published regularly in the Botanisches Centralblait; because many

pecies are asked for which we do not possess. You are requested to tell

us whether the species left to our care need frequent renewing. The greater

ma of our cultures are transferred once every three months, but many
them need particular care.

Further information, and details of terms for the propo
8 adly supplied on application.—Dr. JOHANNA WESTERDIJK,

cherstraat, Amsterdam, Holland.

Those also will
tioned

sed service will
1 Roemer

CURRENT LITERATURE

BOOK REVIEWS
Physiological education

In putting out the second edition of his laboratory course in plant physiology,’
Professor GANONG directs attention to its threefold purpose in these words:

First, it aims to lead students through a good laboratory course in plant physiology.
Second, it seeks to provide a handbook of information upon all phases of plant physi-
ology having any educational interest. Third, I venture to hope that it may find serv-
ice as a guide to self-education by ambitious teachers or students. .... The book
is not a compendium of physiological knowledge, nor yet, except incidentally, a hand-
book of investigations; but it is a guide to the acquisition of a physiological education.
It is designed as a contribution to educational economy, and as such I wish it to be
judged.

No one who examines the work can fail to see that it fulfils the threefold
design of its author, so far as is compatible with success in making it, what
distinctly is, a contribution to educational economy, and by no means the first that
Professor GaNnonc has produced. The strong pedagogical spirit which runs
through the book is suffused by the even stronger scientific spirit, and the com-
bination will make it of the highest service to teachers, as well as to those who,
without other guidance, seek to gain a first-hand knowledge of plant physiology:

t must be pointed out that the book plans vastly more work than can 3
ted to any elementary course in plant physiology in colleges. In the collateral
lines of reading and inquiry that it suggests, there is opportunity for some se
of labor, and for acquiring a wide knowledge of certain topics. Doubtless the
author had it in mind that it is far better to err in this direction than to leave the
student with the false notion that the book presents fully the whole subject. a
a less informed and considerate teacher might heedlessly use such assl,
to overburden seriously the conscientious student. ;

If, however, a self-taught student had this book alone as his guide, his know
edge of plant physiology might not be well balanced, because

th ue of different parts of the subject, and hence the attention paid eat
© author, are not always proportionate to their importance. Thus,
’ YS proportionate to thei po: d about an equal

this is occupied .
the description of apparatus. On the other hand, a little more than 20 pase
Siven to transpiration alone, and nearly twice as much to photoes i it is
this allotment may be defended on pedagogical grounds, it is not clear

j +2656
*Ganone, W. F., A laboratory course in plant physiology. 8v0- PP: s
Sigs. 68. New York: Henry Holt & Co. 1908. $2.00.

242

1909] CURRENT LITERATURE 243

justifiable from the standpoint of the student or of the subject. Obviously, the
author expects that by lectures or other reading and experimentation such inequal-
ities are to be corrected.

One very helpful feature for the teacher is that vari Pr
of conducting experiments are either described in full or are referred to, so that
they may be available. In most cases, however, it will be found that the method
adopted in the book is clearly the most suitable for the elementary student, taking
all things into consideration.

Much of the “normal apparatus,” devised by Professor GANoNnG and now
put upon the market, is highly convenient and useful. In some cases, however,
it is doubtful whether the game is worth the candle, e. g., in the quantitative deter-
mination of transpiration; and often laboratory funds are more limited than the
time of the student.

As to the particular course outlined in this guide, one must inquire whether,
from the point of view of convenience, training, and knowledge acquired, the
selection of experiments is the best that could be made, and whether appropriate
attention is paid to the various topics. On these matters each teacher will have
to form his own conclusion, because his own attitude toward the whole subject,
and the special conditions under which his work is carried on, must be determining
factors. Consequently, it is possible that no two would precisely agree. But
we fancy there will be small disagreement with the statement that no one will
fail to find this book of the greatest service in conducting elementary courses,
even if he hesitates to adopt it formally as a laboratory manual; and for that pur-
pose it is, in many respects, far and away the best that has appeared in any
language—C. R. B. ;

+. fe | +thade

MINOR NOTICES

Javanese fresh-water algae.—Various works have contributed to make known
the algal flora of Java, of which the nearly simultaneous ones by DE WILDEMAN
and by Gurwinsxr are best known and most comprehensive. BERNARD, though
"Ot a professed phycologist and modestly decrying the value of his work, adds
very materially to the knowledge of the Protococcaceae and Desmidiaceae, in a
— voluminous paper published by the Department of Agriculture of the Dutch
rm inning the work’of collecting almost accidentally, the beauty and
coat Of the unicellular forms and the necessity of examining them in the living
co. determined his study of them. In an introduction (45 pp-) the author,
“ * giving briefly the history, bibliography, methods of study and collection, and

— of the localities explored, discusses the variations, —

* BeRNarp, Cu., Protococcacées et Desmidiacées d’eau douce,. recoltées & Java

et é fs
Par C. B. Imp. 8vo. pp. 250. pis. 16. Batavia: Lan erij. 1908.

244 BOTANICAL GAZETTE [uarce

two orders, of which BERNARD has collected and described 202. Of these 4 are
new to Java, 79 others are new to the East Indian region, and 81 are described

as new species or varieties. By 580 carefully drawn figures, rather crowded on
the plates, the author represents all species of his collection, so that later workers
can see what plants he has actually been working with. The evident care and
thoroughness of the work indicate that this is no mean contribution to the
knowledge of the Javenese flora—C. R. B

- Folk names of Brazilian plants.—For some years there has been running
through the Pharmaceutical Review a series of articles by Dr. THEODOR PECKOLI,
giving the vernacular names of Brazilian plants and plant products, including
both the Portuguese names and those adopted from the Tupi language. This
material is now brought together in book form,3 as monograph no. 15 of the
Pharmaceutical Science Series, under the editorship of Dr. EDWARD KREMERS.
The vernacular names appear in alphabetic order, with the German equivalent
where it exists, the scientific equivalent, including the specific name and family
name, when known, and brief comments in German on the use made of the prod-
ucts. Itis rather unfortunate that there is not an index to the scientific names,
for this would undoubtedly greatly increase the usefulness of what has been 4
difficult and time-consuming task. The volume will be of special assistance to
taxonomists, to dealers in es drugs, and to manufacturers who call for
Brazilian products.—C.

German South-polar Expedition—The second part of the eighth volume
(Botany) of the sumptuous report upon this expedition has just n issued
with an account by REINBOLD of all the seaweeds except the Lithothamniaceat,
which are elaborated by Fostie. The collections were not extensive and no ns
species were found by RemnBotp. FostrE, however, recognized and descr
several new unsegmented corallines from the material obtained by this sp
and here presents again the descriptions with photographic illustrations —C-

NOTES FOR STUDENTS

A primitive type of seed.—OLIver has made a most inte

resting contribution’
to our knowledge of the structure of paleozoic seeds. In 1875

Pro-
3 PECKOLT, THEODOR, Volksbenennungen der brasilianischen Pa oe
dukte derselben in brasilianischer tevetngie sischer) und von der TupisP Co. 7
tenNamen. 8vo. pp.252. Milwaukee: Pharmaceutical Review es tes des
eutsche ag ae a 1901-1903, im Auftrage op Baie TI vin.
ined beau von EricH von Drycarsk!, Leiter der Expediti a
Band, Botanik, Heft II. (1) Retnsotp, Tu., Die eae are
Fosur, M., Die “sean ie pp. 203-220. pl. 20. figs:
Reimer. tock. Ms.
F. W., On Physostoma elegans Williamson, an are ae $. a -
from the Palaeozoic eck’ Annals of Botany 23:73-116. P vs. 57° om”

1909] CURRENT LITERATURE 245

described Physostoma elegans from the Lower Coal-measures, but a little later
placed it in his new genus Lagenostoma as L. physoides. OLIVER now proposes
to revive the genus Physostoma as distinct from Lagenostoma, replacing the
name L. physoides by the original P. elegans, and associating with it ARBER’s
L. Kidstonii as P. Kidstonit.

Physostoma elegans is remarkable in several particulars. It is a small (5.5-
6™"), narrow, usually ten-ribbed seed, with the integument free only in the region
of the nucellar beak, as in Lagenostoma; but in this free region the integument
consists of ten distinct lobes (‘‘tentacles” the author calls them). These lobes
are the direct prolongation of the ribs below, and represent the units of the “canopy”
of Lagenostoma. Both lobes and ribs are clothed with long, club-shaped hairs.
The testa shows no stony layer, and therefore is homogeneous, consisting of five
or six layers of close-fitting, thin-walled, elongated cells. The vascular system
enters the narrowed base of the seed as a single strand, which at once breaks up
into a ring of strands, each one of which traverses a rib and continues on through
the corresponding free lobe. The vascular strands lie along the inner limit of
the integument, in what would be the ‘‘inner fleshy layer” had the usual three-
layered differentiation of the testa occurred. On the outside of each strand there
isa lacuna; and in the base of the seed a continuous lacuna surrounds the single
large strand and the group of separating strands. It is suggested that these lacunae
fepresent the position of disorganized phloem.

The nucellus is remarkable in its peripheral “secretory zone,” which extends
from the chalaza to the tip of the nucellar beak. The “‘secretory sacs” are thin,
oblong, tabular cells, separated by a tissue of smaller cells. They are most abun-
dant and crowded in the funnel-shaped region of the chalaza bounded by the diver-
ging vascular strands and the base of the embryo sac. The author sees in the
Presence of these secretory sacs the retention in the ovule of a feature present 1n

this may represent the active nutritive zone developed in many gymnosperms
rdinary nucellar tis- -

mordi ;
ordial tent pole,” as found in Cordaitales and Ginkgoales. SS un
len grains,

cell-complex now known in several paleozoic seeds. The author thinks that

246 . BOTANICAL GAZETTE [MARCB

at least the larger of these cells produced sperms, and asociated bodies, probably
representing sperms, were also found.

The author concludes that an integument of free segments is more primitive
than that of “‘coalesced” segments, and that probably intermediate stages of coales-
cence occur in the Lagenostoma group. In this group, therefore, the ongin of
the integument is multiple, but the nature of the units is the residual question.
OLIvER is not inclined to accept Miss BENSON’s suggestion that the integument
(as illustrated by that of Lagenostoma Lomaxii) has arisen by the sterilization of
the peripheral sporangia of a synangium; but prefers to regard it as a new struc-
ture, arising contemporaneously with the seed habit, and related in some way
to the “encasement” that so often accompanies reproductive activity.

The reasons for regarding Physostoma as a member of the Lagenostoma
group are given in detail, and the conclusion is reached that it is the “most primi-
tive seed yet come to light,” the plant to which it belongs probably being one of
the Lyginodendreae. The reasons for the conclusion quoted above are not quite
clear, and seem to contradict some rather convincing conclusions reached by the
same author in his study of Stephanospermum and other paleozoic seeds.—J. M. C.

Sterility in hybrids. —TiscHiEeR® has a lengthy treatment of the subject of

Sterility in hybrids. A preliminary paper, summarizing his conclusions, has

dy been reviewed in. this journal.?/ The present paper is in two parts, |

first presenting the cytological data and the second dealing with the theoretical
conclusions. The use of charcoal in drawings can scarcely be recommended for
clearness, many of the figures being mere smudges, and they furnish no sufficient
evidence of such cytological matters as the pairing of threads in synapsis. P Ne '
formation, and in some cases megaspore formation, is described in hybrids 0
Mirabilis, Potentilla, and S$ ringa. A variety of irregularities, such as ate com
mon during the reduction divisions in hybrids, are described, including gee
of extra nuclei by chromosomes left in the cytoplasm and failure of one oF 00"
reduction mitoses. In other cases the reduction processes were normal, but there
was a lack of cytoplasm and the pollen grains failed to grow. Pantie
cultivated in dry and hot conditions, matured good pollen, but after ite :
the young embryos died. In all cases a paucity of cytoplasm was 0 —
ning during or after reduction.

TISCHLER concludes that the cause of sterility is not any lack of sigsie
between the chromatic elements. However, it seems necessary to aera
‘incompatibility ” of the chromatins or plasms, which makes itself € sn
during the formation of reproductive cells, for otherwise there is 10 €XP" m

_Why a plant continues to show vegetative growth and yet fails to mature - ee
cells. Instead of a chromosome incompatibility, perhaps we may have ss
cee

orschung
°TISCHLER, G., Zellstudien an sterilen Bastardpflanzen. Arch. Zellf
1333-151. figs. 120. 1908,

7 Bor. Gazetre 45:68, 1908.

1909] CURRENT LITERATURE 247

cytoplasm some process which is symbolized by the pairing of chromosomes in
synapsis, and which, owing to differences in the composition of the parental idio-
plasms, leads to derangement and finally cessation of the metabolism that had
previously been carried on successfully. Some such hypothesis is necessary to
explain why failure of growth usually begins with germ cell formation, and the
necessity is not lessened by the fact that sterility is a purely relative phenomenon
produced also by other conditions than hybridization.

TISCHLER agrees with Jost that the increased luxuriance of some hybrids is
probably due to a “‘poisoning” effect of one species on the other. Some of the
cases of self-sterility bear a similar interpretation.

Three classes of facts are cited to show that there is not a segregation of charac-
ters during reduction in Mendelian hybrids: (1) Cases of vegetative splitting,
as in Syringa correlata and Cytisus Adami. (2) Certain cases of latency or cryp-
tomery (TscHERMAK); e. g., the crossing of two white forms having certain other
characters gives a violet hybrid. But such cases have been otherwise explained
by the Mendelians. (3) Characters mendelize which cannot be represented by
distinct portions of the idioplasm. Here are cited annual and biennial races of
Hyoscyamus niger, immunity and non-immunity to rust in certain grains, and

dition in the germ cells, a hypothesis which will undoubtedly have to be given
and in
This

DR having long and round pollen, in which all the F, had long pollen
re F, gave long: short in the ratio 3:1. From this it appears that it is pos-
to have Mendelian behavior without segregation of characters during
reduction.
PG frequent sterility in mutants, accompanied by similar irregularities to
elke: hybrids during reduction, as the reviewer has shown,® TISCHLER also
to some disturbance of the idioplasm.—R. R. GATES.
t .
pa Temperature and growth.—Beginners in research will do well to study this
— by Batts.° It is of a type really too rare. It shows how a keen scientific
‘Timent is alert to appreciate the significance of a casual observation in its
upon a fundamental problem. It shows how difficulties may —
8 :
and . — R. R., Pollen development in hybrids of Ocenothera lata XO. Lamarckiana,
"is to mutation. Bor, GAZETTE 43:81-115. pls. 2-4. 19°7-
soe « reps LaWRENcE W., Temperature and growth. Annals of Botany 22557

248 BOTANICAL GAZETTE [Marcy

by ingenuity in devising efficient apparatus and illustrates the potency of logical
inquiry. One unfavorable criticism is that the scientific name of the organism
is not given. During a study of a pest of the cotton crop in Egypt, the author
noted that cultures of this ‘‘sore-shin’”’ fungus showed a notable diff bet
the thermal death-point and the temperature inhibiting growth. ‘This observation
suggested an analysis of the temperature factor in its effects upon growth. Itis
stated that the hyphae of this fungus are morphologically and physiologically
equivalent, in that spore-formation, sexual or asexual, does not occur.

this statement is not to be taken literally, as it would be very difficult to say that
all the hyphae of a given fungus are physiologically equivalent. As a matter of
fact, the author himself states that in liquid cultures at 20° C. resting cells are
formed in abundance. If the cultures are grown at 34° C. growth ceases (culture
becomes stale) much earlier than at lower temperatures. This feature of “stale
ness”’ or of discontinued growth was found to be caused by the accumulation of
substances which retard and if sufficiently concentrated stop growth. The sub-
stance or substances which originate in the organism as a result of the effect of
temperature, and whose influence is to inhibit growth, have been isolated from
the organism as products of katabolism, though they have not been chemically
identified. To such katabolites the provisional name of “X”’ is given. em
a large number of tests whose results are tabulated, illustrated by appropriate
Curves and verbally discussed, it appears to be demonstrated (a) that with ae
ing temperature there is a regular acceleration in the rate of growth up to oe
and this acceleration approximately fulfils the expectation based upon V
Horr’s law; (b) above 30° C. the growth-rate acceleration decreases as the factor
of time becomes limiting; (c) later growth stops at a fairly definite ener
which the author proposes to call the ‘‘stopping point;” (d) the optimum . effects
fore not a definite temperature but a status of the organism in which the se
of the factors of time and of temperature physiologically balance. As ~ sal
expected the style and composition of the paper are consistent with the 108!
development of the investigation—_RAyMonpD H. Ponp.

tly.
thirteen gene
nted, the P odo-

G., AND FRAINE, E. pE, The seedling structure of gym

os. L
‘Ena: fT.
Annals of Botany 22:689-712. 1908.

1909] CURRENT LITERATURE 249

cotyledon. The cotyledonary bundles contain either centripetal xylem or its
lineal descendant, transfusion tissue, the pronounced mesarch bundle occurring
in greater proportion in the Taxineae (Taxus and Cephalotaxus). With respect
to the number of cotyledons, only members of the Pinaceae (Cryptomeria and
Sequoia gigantea in the Taxodineae, and Libocedrus and some species of Cupres-
sus among the Cupressineae) have more than two. T he presence of resin ducts,
likewise, was observed only in the Pinaceae, Juniperus having them in the leaves,
and the Taxodineae having them in the cotyledons in all the forms examined,
except Widdringtonia. Two instances of fusion of cotyledons are reported: in
Widdringtonia Whytei, the two cotyledons unite laterally to form one, recalling
the leaf of Sciadopitys; in Cupressus torulosa, the cotyledons fuse near the base
to form a tube. In every case the number of root poles corresponds with the
number of “‘whole’’ cotyledons.

The authors believe that dicotyledony is the more primitive condition, and
that the polycotyledonous form has been derived from it by splitting; but the
statement of their reasons for this conclusion is deferred to a future paper.

It is a hopeful sign that the authors did not intrust this valuable collection
of seedlings to the mercy of a razor; to read that the sections were made in an
orderly fashion with the help of a microtome gives doubl that t .
vations are accurate—HELEN A. DorETY.

Root excretions. —SroKLasa and Ernst* report a conclusive piece of wor,
syste excretions. The excellent technique these workers have developed in
dling similar problems makes their contribution in this much disputed field

‘nusually valuable. They find that no acid (organic or inorganic) except H,COs

Potassium phosphate, contrary to the claim of CzaPex. The organic acids appear”
Buckwheat and

the beet, oxalic.

rien authors are to make an exhaustive investigation of the question oe
acids Ydrogen is produced in the aerobic respiration of roots, and what —
seni, excreted by the roots of many other species of plants under limited oxygen
The authors also determined the amount of CO, excreted by the root systems
amoun » Oats, rye, and wheat. The barley root-system gives off the grea =
tof CO, and produces the greatest dry weight. The quotient arising a.
Pea oe

t2 :
Stoxtasa, J., ano Ernst, A., Beitrige 2iir Lésung der Frage der —

Natur
des Wurzelsekretes. Jahrb. Wiss. Bot. 46:55-102- 1908.

250 BOTANICAL GAZETTE [Marc

dividing by the dry weight of the root system the weight of CO, produced is least
in barley and greatest in oats. This quotient is considered as the indication of
the specific energy of respiration.

The authors point out that the injury to farm crops by insufficient aeration
of the soil probably arises from the accumulation of the highly toxic organic acids
due to the incompletely oxidized products of respiration. A

It should be mentioned here that the authors have studied only the aliphatic
excreta and make no mention of any of the aromatic series—Wa. CROCKER.

Graft hybrids.—The question of the occurrence of graft hybrids has long been
undecided, and the possibility of their existence has even been denied. WINKLER
undertook an extensive series of experiments on this subject, using as material
two species which will not hybridize in the ordinary manner, namely Solanum
migrum and Solanum lycopersicum. In an earlier paper?s he dealt with the
production of what he calls chimeras, that is shoots, one side of which resembles
either parent, the cells of the two parents growing in juxtaposition without modify-
ing each other. He has finally succeeded*4 in producing a true graft hybrid
between the same species by the same method.'5 In all, 268 grafts — made,
which after decapitation produced over 3000 adventitious shoots. Five of the
latter were chimeras and a single one a graft hybrid, which came from

S. lycopersicum on S. nigrum. After decapitation the cut surface of one graft
Produced 14 adventitious shoots, 8 of which were pure S. nigrum, 5 S. lycopersicum,
and r the graft hybrid. ‘The latter was detached and rooted, finally producing
flowers. It is intermediate in character between the parents, though somewhat

nearer S. nigrum. The purity of both parents was assured by using
“pure line” cultures, WINKLER names the hybrid S. tubingense, and | pa
to use the sign + for graft hybrids instead of X, the sign for a sexual pee ne
other adventitious shoots are also probably intermediate in character, one
being nearer the |S. Lycopersicum.
Adami, found the number of chromosomes to be the same as in eact | are
Several interesting cytological questions, which WINKLER hopes to determin
involved in the nuclear and chromosome behavior of his graft hybrid. ae
ently there must be a union of cells, nuclei, or chromosomes, or perhaps °
three, in the production of this form.—R. R. GATES.
ee
*3 WINKLER, Hans, Ueber Pfropfbastarde und pflanzliche Chimaren. Pe
tsch. Bot. Gesells. 25 2568-576. figs. 3. 1907.
%4——.. Solanum tubingense, ein echter Pfropfbastard zwischen
Nachtschatten, Ber. Deutsch. Bot. Gesells. 26a:595—608. figs. 2. 1908
*5 Briefly described by the reviewer in Bor. GAZETTE 47:84. 1999
‘ ” STRASBURGER, E., Ueber die Individualitit der Chromosomen und die
briden-Frage, Jahrb. Wiss. Bot. 44:482-555. pls. 5-7. fig. I. 1907:

Deu
Tomate und

Pfropihy-

TRASBURGER,'® in a cytological study of the reputed graft hybrid psc

Rie ia ee eS eee 8 a ite a

1909] CURRENT LITERATURE 251

Germs in the air.—Sarro has determined, by agar and gelatin plates, 10
in each experiment and exposed by the ancient method (cf. SepGwick and TUCKER,
rath Ann. Rep. Mass. State Bd. Health, Boston, 1889), the number of bacteria,
as counted by colonies, and the kinds found in the University garden and the street
airof Tokyo.?? Several of these tests were made in each month of the year, with
observations as to temperature, humidity, wind, etc. He found a high street
average in July and August during a period of dust and dryness following a wet
season. The garden average increased in November and December in proportion
to a large number of windy days. Snow or rainfall always cleansed the air of
germs. Of the 55 Bacteriaceae and 17 ‘Coccaceae found, Saito described 18
as new species, without making much attempt to relate these to forms already
known. It is difficult to see, for instance, wherein his B. rujulus differs from
B. rubiginosus described by CaTarno in Comn’s Beitrage 7:538. 1896; and Sarcina
agilis appears to differ from M. agilis as described by Migula in 1897 only in the
easily lost character of pigment formation. Motile sarcinas, however, are rare
and should be carefully studied Mary HEFFERAN.

Anatomy of Saxegothaea.—This monotypic conifer, restricted to the wet
woods of the upper slopes of the Andes of Chili, has long been in demand for
morphological investigation. When LinpLEy described it in 1851, he called
attention tg the transition characters it exhibits between Taxaceae and inaceae.
Allied with the Podocarpineae in what are regarded as the more important charac-
ters, it shows even more resemblance to the Araucarineae than do the other podo-
carps, especially in its wingless pollen grains and distinct cone of spirally arranged
Sporophylls. Srrzzs*® has examined the anatomical structure of two specimens
in cultivation in England, and concludes that the genus is relatively primitive.
The structure of the wood of the stem and of the medullary rays is simple; and
taken together with the simple arrangement of the sporophyllls, has convinced
real that Saxegothaea is at least the oldest of the podocarps, and shows

tivation from a common ancestral stock with the araucarians.—J. M. C.
the Potato breeding.—Easr"? considers the extensive record of the history of
i and future methods of its improvement. Many interesting biological
«Of variation and hybridization in the potato are brought together, as well
Practical suggestions for the improvement of varieties. The cultivation of

ES, W., The anatomy of Saxegothaea conspicua Lindl. New Phytol. 7:
4. 1908.

Ill, “psy E. M., A study of the factors influencing the ——— oe
~ *xpt. Sta. Bull. 127. figs. 10. 1908.

252 BOTANICAL GAZETTE [MARCE

include (1) crossing under controlled conditions, (2) selection of fluctuations, and
(3) selection of wider variations and a study of ways of causing them, graft hybrids
being a possible method.—R. R. Gates.

Salts of aluminum.—F uri’? finds that aluminum salts cause the disappear-
ance of starch from Spirogyra and other water plants, even when they are well
illuminated. It likewise renders the protoplasm permeable to ordinary plasmoly-
tic agents. The disappearance of the starch is probably due to the joint action
of three effects of these salts: loss of sugar through the permeable protoplasm,
the increased diastatic action, and the slower photosynthetic activity. If glucose,
glycerin, or isodulcitol is mixed with the aluminum salt, the salt does not then
render the protoplasm permeable.—Wwa. CROCKER.

Transpiration and water storage.—SHREVE? finds that Stelis ophioglos-
sotdes, a water-storing epiphyte which grows at mid-height in the rainy forest of
Jamaica, is less able to resist continued drought than Guzmania tricolor, an epi-
phyte which does not store water, but grows at the top of the rainy forest. Stelis
reduces its transpiration almost to nothing when forced to draw upon the stored
supply in the leaves. With an abundant external supply, it shows rather rapid and

transpiration WILLIAM CROCKER.

Branch cankers of Rhododendron.—Von SCHRENK?? describes some interest-

ing swellings of the branches of Rhododendron maximum that are produced where "4
dead twigs have fallen. The healing layer starts to develop at a distance from os

- base of the fallen twig, and in a few years a large swelling results. The chief
interest in the phenomenon arises from the fact that a gall-like structure 1s pe
duced without the intervention of a parasite.—H. C. CowLes.

Virescence in Oxalis.—Hvs?3 has been studying plants of Oxalis _— ser
bright green petals, but which are otherwise representative of the species.
€ green petals approach sepals in structure in some respects, they retain
characteristics of the ordinary petals. The new form, which has been nam
Oxalis stricta viridiflora, is constant from seed, and is to be made the object
careful study.—H. C. Cowes.

*° FLurt, Max, Der Einfluss von Aluminiumsalzen auf das Protoplasm4
99 :81-126. 1908. ae i

** SHREVE, Forrest, Transpiration and water storage in Stelis op — a
Plant World 1I:165-172. 1908. ard. Rep

22 SCHRENK, H. von, Branch cankers of Rhododendron. Mo. Bot. GA ae
18:77-80. 1907. : a

* Hos, H., Virescence of Oxalis stricta. Mo. Bot. Gard. Rep. 18:99° 1" “”

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‘-CONTENTS FOR APRIL 1909 No. 4
RIBED PLANTS FROM GUATEMALA AND OTHER CENTRAL AMERICAN
PARATIVE HISTOLOGY OF FRUITS AND SEEDS OF — SPECIES OF

CUCURBITACEAE (wiri FIFTY-THREE FIGURES). Kate G. Barbe 263

ANATOMY OF ISOETES. ContTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY 126
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VOLUME XLVII NUMBER 4

BOTANICAL (GAZETTE
APRIL 1909

UNDESCRIBED PLANTS FROM GUATEMALA AND OTHER
CENTRAL AMERICAN REPUBLICS. XXXI*
Joun DONNELL SMITH
(WITH ONE FIGURE)

Magnolia guatemalensis Donn. Sm.—Glabra. Folia breviter
petiolata elliptica apice acuta basi obtusiuscula concoloria subtus
glabrescentia crebre penninervia. Sepala oblonga tenuiter coriacea
ynervia. Petala spatulato-obovata coriacea. Stamina numerosissima.
Gynophorum fusiforme.

Arbor 6-8-metralis. Folia supra cum ramulis petiolis [
pilis sparsis castaneis aspersa vel denique glabra 12-16 longa 5.5-8.5°™ —

enue abrupte breviterque cuspidata, costa subtus tuberculato-rugosa, nervis

‘ralibus utringite 13-16, interjectis minoribus vix ullis, petiolis 1.5-2™ longis
apice canaliculatis ceterum teretibus. Pedunculus 3°™ longus, stipulis foliorum
nascentium §—y5™m longis passim castaneo-pubescentibus. Sepala 5 e
ie a lata ima basi 6-8™™ lata humectate pellucido-punctulata. Petala
oes * tad 29™™ lata ima basi 2-3™™ lata opaca cum sepalis obtusa glabra
oe eceptaculum staminale 6™™ longum 4™™-diametrale, staminibus oF
Paige Gynophorum 25™m longum medio 9™™-diametrale, carpellis circiter
ang Sulcatis in stigma 4™™ longum recurvatum attenuatls. Fructus

i =. . portoricensi Bello proxima differt praesertim glabnitate. ‘
1550 ag profunda prope Tactic, Depart. Alta Verapaz, Guatemala,

' - 1998, H. von Tuerckheim (n. II. 2165).

: Verapazensis Donn. Sm.—Folia supra glabra subtus
. eo puberula et diaphano-punctulata ceterum opaca oblonga acute
mie er cusPidata basi acutiuscula. Racemi folia subaequantes
oboy. 5p Sepala ovata vel ovalia pedicellum aequantia petalis
Ati -oblongis paulo breviora. Antherae in appendicem linearem

—. Stylus ovario vix brevior, stigmate minimo.

nunued from Bot. Gazette 46:117. 1908.

j ln olahra subtus
oO

alt.

253

254 BOTANICAL GAZETTE [pri

Ramuli glabri. Folia coriacea 18-20°™ longa medio 6-6.5°™ lata, nervis
subtus elevatis et pallidis, lateralibus parallelis utrinque 13-15 sub ipso margine
arcuatis, venis transversis undulatis, venulis minute reticulatis, petiolis 12-15™
longis. Racemi fere a basi floriferi prope basin ramum nonnunquam emittentes,
pedicellis inordinatim dissitis puberulis, floribus pentameris. Sepala in alabastro

uberula post anthesin deflexa 7-8™™ longa, 3 exteriora late obtuseque ovata
interioribus suborbicularibus paulo angustiora. Petala 9™™ longa apice 3-4™"

i 1™™ lata. Stamina 4-5™™ longa, antheris appendice eis paulo breviore
computata 2™™ longis. Ovarium glabrum elongato-oblongum, stylo 5™™ longo,
stigmate pyramidato 1™™ longo. Capsula ignota—M. macrophyllae Benth.
proxima.

In silvis montanis ad praedium Cubilguitz dictum, Depart. Alta Verapaz,
Guatemala, alt. 350™, Oct. 1904, H. von Tuerckheim, n. 8660 ex Pl. Guat. ete.
quas ed. Donn. Sm.

Leandra (§CARASSANAE Cogn.) Tuerckheimii Donn. Sm
Ramuli teretes uti petioli paniculae calyces strigillosi. Folia ovate
oblonga vel -lanceolata incurvo-acuminata basi acuta septuplinervia
discoloria supra tuberculato-setosa subtus foveolata pubescenta.
Flores ebracteolati. Calycis lobi interiores brevissimi, segmenta
exteriora filiformia tubum aequantia.

Ramuli dichotomi. Strigillae densae arcte appressae purpurascentes. F olia
leviter disparia rigida inter bullas diaphana supra intense viridia setis cra
curvatis armata subtus pallida nervis strigillosa venis pubescentia Ti pet
infra medium 3-4.5°™ lata basi inaequalia, petiolis in eodem jugo inaequali a
T.5~3°™ longis. Paniculae terminales singulae vel 2-3-nae trichotomae 6-7-5.
longae, bracteolis lineari-lanceolatis 1. 5-2™™ longis, pedicellis 1-5~?
floribus 5-meris. Calyx longe strigillosus, tubo campanulato EE ee ‘s
limbum non constricto, lobis interioribus vix o.5™™ longis semiorbiculan “
Petala oblongo-ovata 4-5™™ longa. Stamina so, antheris 1.5" longis eat
compressa aequantibus, connectivo infra loculos haud producto. Ovarium ver"
convexo glabrum 5-loculare, stylo 4-s™™ longo. Bacca non adest-~
methodum clari Cogniaux juxta L. strigillifloram Cogn. locanda.

In summo monte silvestri prope Coban, Depart. Alta Verapa;
alt. 1600", Jun. 1908, H. von Tuerckheim (n. II. 2369).

Secundum

Guatemala,

Hoffmannia Tuerckheimii Donn. Sm.—Pubes undique wee
monili-formis. Folia inter minora longiuscule petiolata ae Me
Ovalia contracto-acuminata basi acuta vel rotundata supra ao b-
subtus prasertim ad nervis pubescentia. Flores fasciculah *

is bis
sessiles pubescentes. Calycis tubus obovatus lobis subulat

1909] SMITH—PLANTS FROM CENTRAL AMERICA 255

longior corolla partita 6-plo superatus. Filamenta antheris triente
breviora.

Ramuli dichotomi teretes lenticellati fusco-pubescentes, stipulis triangulari-
bus. Folia coriacea 5-6.5°™ longa 2.5-4°™ lata subtus pallida et inter nervos
subtilius pubescentia, nervis lateralibus utrinque 5~7, petiolis 15-22™™ longis.
Pedunculus vix ullus nodiformis glandulosus, pedicellis 3-10-nis 1™™ longis,
floribus tetrameris 14™™ longis. Calyx tetragonus 3™™ longus sinubus glandu-
losus. Corolla sparsius pubescens 12™™ longa, segmentis linearibus tubo 5-plo
gioribus. Stamina fauci inserta 5™™ longa, antheris lineari-oblongis. Stylus
corolla paulo brevior, stigmatibus 2 linearibus 2™™ longis. Bacca deest.

In silvis supremis montis prope Cobdn, Depart. Alta Verapaz, Guatemala,
alt. 1600, Mart. 1908, H. von Tuerckheim (n. II. 2160).

Guettarda (SULoLosus DC.) cobanensis Donn. Sm.—Folia ter-
hatim verticillata longe petiolata oblongo-elliptica sursum subsensim
deorsum contractius acuminata glabrescentia. Stipulae internodiis
superioribus breviores. Flores 4-5-meri. Corolla inter longiores,
tubo calycem 7-plo. lobos proprios 3-plo stylum dimidio superante.
Drupa glabra, putamine 3-4-loculari atque -quetro, lobis intermediis
saepius adjectis.

loculis rectis—G, crispiflorae Vahl proxim

alt. ts
100", Jan. 1908, H. von Tuerckheim (n. IT. 2096).

Chomelia brachypoda Donn. Sm.—lInermis. Folia glabrescentia
vente lanceolata utrinque acuminata paucinervia needs
. “ings Pedunculi petiolis parum longiores, cymis laxe paucitions

Tibus
Culatus, Corollae tubus anguste cylindricus calyce 4-plo lobis pro-
Prlls 5-6-plo longior. ?

paene glabris. Calyx subsessilis oblongus minute denti- -

256 BOTANICAL GAZETTE [APRIL

Ramuli petioli stipulae pedunculi appresse pilosi, cymis calycibus corollae
lobis sparsim pilosiusculis. Folia 7-10°™ longa medio 2.5-3°™ .lata sursum
tenuiter acuteque incurvo-angustata deorsum subsensim acuminata chartacea
in sicco viridia supra glabra subtus pilis raris aspersa et axillis barbata, nervis
lateralibus utrinsecus 4-5, venulis erga lucem inspectis manifestis creberrimis
undulatis, petiolis 5-6™™ longis, stipulis filiforme linearibus 6-8™™ longis. Pe-
dunculi 5-8™™ longi medio bibracteolati, cymis 4—8-floris, pedicellis brevissimis
crassis. Calyx 2.5-3™™" longus, denticulis triangularibus vix 0.5™™ longis.
Corollae tubus glaberrimus 11-12™™ longus, lobis ellipticis 2™™ longis. Antherae
inclusae vix 2™™ longae. Ovarium calyce dimidio brevius, stylo ramis 2”
longis computatis 7™™ longo, ovulis linearibus. Drupa desideratur—Ob folia
C. filipedi Benth. valde affinis differt praesertim pedunculis perbrevibus.

Ad ripas fluminis Ogewaj prope Sasis, Depart. Alta Verapaz, Guatemala,
alt. goo™, Maj. 1908, H. von Tuerckheim (n. II. 2253).

Satyria meiantha Donn. Sm.—Folia juniora lanceolato-oblonga
tenuiter acuminata basi acuta, provectiora ovalia bis longiora quam
latiora utrinque subaequaliter angustata praetermisso utroque nervulo
basali mox evanido triplinervia. | Corymbi sessiles subsimplices,
floribus minimis. Antherae tubo filamentorum paulo longiores
apice acutae.

- Frutex grandis congeneribus habitu similis. Folia in ramulis~ :
13-17°™ longa 4.5—6°™ lata, in ligno vetere 18-19°™ longa 9°” lata, nervis ger
bus utrinsecus 5-7, petiolis 1-1.5°™ longis. Corymbi ad nodos defoliatos 2. si
longi 8-r4-flori, axibus confertis plerumque simplicibus 1°™ longis, bracteolis
lanceolato-ovatis, basalibus 2™™ longis, medialibus 2 suboppositis yam es
Calycis tubus r.5™™ longus 2™™-diametralis basi intrusus, limbus dentatus 1
altus. Corolla r1-r2™™ longa etiam in sicco laete rosea minu issime
lobis ovatis apiculatis membrana connexis. Stamina 5.5-0"™ longa, bis
torum tubo 2.5-3™™ longo, antheris majoribus 3-3.5™™ longis, omnijum tt
Sursum dilatatis apice acuto discretis, poris oblongo-ellipticis. Stylus paulo
exsertus. Baccae deficiunt. alt

In silva montana prope Cobdn, Depart. Alta Verapaz, Guatemala,
1600™, Jan. 1908, H. von Tuerckheim (n. II. 2101).

Gonolobus (§Monosremma K. Schum.) patalensis Don? oe
Folia oblongo-ovata acuminata leviter cordata sparsim minut?
strigillosa. Flores umbellatim cymosi. Corolla calyce 3-plo longior
intus glabra colorato-reticulata lobis ovato-oblongis alte fiss4- —
cyathiformis gynostegium subincludens ab eo libera 4
minutissimis bifidis s-denticulata intus infra marginem 44a”
minuta cum marginalibus alternante instructa.

1909] SMITH—PLANTS FROM CENTRAL AMERICA 257

Suffrutex volubilis, ramis petiolis cymis calyce patenter pilosis. Folia 7-11°™
3.5-6°™ lata, sinu aperto acuto, nervis lateralibus utrinque 5-6, petiolis
15-30™™ longis. | Pedunculus 1o-13™™ longus, ‘pedicellis plerumque 5-nis
15-20™™ longis. Calycis partiti segmenta lineari-lanceolata 5"™ longa. Corolla
14™™ longa rotata extus pilis raris aspersa, lobis 10™™ longis obtusis. Corona
2" alta 3™™ lata. Pollinia pendula obovata. Folliculi ignoti—Secundum
conspectum a cl. Schumann ad coronam ordinatum gregi “Bb” ascribendus.

Patal, Depart. Baja Verapaz, Guatemala, alt. 1600™, Jul. 1908, H. von
Tuerckheim (n. II. 2371).

Gonolobus (§MonostemMA K. Schum.) araneosus Donn. Sm.—
Folia oblongo- vel lanceolato-ovata acuminata leviter cordata molliter
pilosa. Cymae umbelliformes petiolum paulo superantes, floribus ~
inter minimos. Corolla calyce dimidio longior colorato-reticulata
usque ad medium lobata intus praeter lobos triangulari-ovatos gla-
bros niveo-arachnoidea. Corona cyathiformis gynophorum aequans
ab eo libera callis bipartitis 5-denticulata, squamula interna minuta
cum callis alternante.

Suffrutex volubilis, ramis petiolis cymis calyce patenter pilosis.
sericea lineari-lanceolata, provectiora 6-8°™ longa 2-5-3: 5c™ Jata, sinu aperto
acuto, nervis lateralibus utrinque 4-5, petiolis 1o-20™™ longis. Pedunculus
35™™ longus, pedicellis 4-5-nis 3-7™™ longis. Calycis partiti segmenta linear
lanceolata 5™™ longa. Corolla 8™™ longa rotata, lobis 4-4.5™™ longis acutis
extus parce pilosiusculis apice ciliatis. Corona 1™™ alta 2™™ lata. Pollinia
pendula compresso-orbicularia. Folliculi desiderantur—Ad speciem praeceden-

= foliis calyce corona arcte accedens inflorescentia atque corolla insigniter
recedit. j

Folia juniora

on . inter Tactic et Cobian, Depart. Alta Verapaz, Guatemala, alt.
» Jul. 1908, H. von Tuerckheim (n. II. 2332)-

Merinthopodium campanulatum Donn. Sm.—Pedicelli filiformes
Hloribus permagnis paulo breviores. Corolla sepalis bis fere longior €
fundo subcylindrico abrupte lateque dilatata, lobis elongato-triangular-
ibus, sinubus acutis. ;
ogee, foliis coriaceis ovalibus acutis (ex cl. repertore in literis). seit

ngi. Sepala acutissima apiculata 5. 1-5 .3°™ longa basi 1-6-1-7

oS -2°™ longa in sicco tam lata quam longa,
lete plicatis. Antherae 1. 7em Jongae. Stylus gem longus.
_cuanquam folia non vidi, tamen ob flores ab eis M- neuranthi Donn. Sm.

—" Pantes plantam publici juris faciendam puto. eee
V “iS primaevis superioribus montis haud procul a Coban, Depart.
Lerten 1600", Mart. 1908, H. von Tuerckheim (11. 2391): -

258 BOTANICAL GAZETTE [APRIL

Neotuerckheimia Donn. Sm., nov. gen. BIGNONIACEARUM ¢€
tribu CRESCENTIEARUM.—Calyx coriaceus glaber primum clausus
denique in lobos 2 ovales inaequaliter ruptus. Corolla infra medium

tubuloso-campanulata medio antice ad plicam transversam deorsum
flexam geniculata ventricosa, limbo obliquo vix lobato crispatim
dentato. Stamina 4 didynima inclusa paulo supra basin corollae
inserta, loculis oblongis pendulis. Discus pulvinatus. Ovarium
conicum costis 8 angulatum perfecte uniloculare, placentis 2 parieta-
libus valde intrusis, ovulis pluriserialibus, stylo angulato, stigmate
bilamoso.—Arbores glabrae. Folia singula vel 3-fasciculata oblan-
ceolata subsessilia. Nodi floriferi laterales vel terminales bracteis
numerosis obtecti, pedunculis 1~s-nis. Corollae tubus_ plurinervis,
limbus colorato-reticulatus.

In honorem nominavi liberi baronis H. von Tuerckheim floram Guatemalensem
ad cognoscendam viginti tres per annos collaboratoris amicissimi.

Fic. 1.—Neotuerchheimia megalophylla; jructus.

Neotuerckheimia megalophylla Donn. Sm.—Folia 3-fasciculal®
a
incurvo- et falcato-acuminata deorsum longissime attenuata. P pre

rii ¥

culi laterales squamis permultis cartilagineis circumdatl solitar
ni near ‘ s OVO!
bini. Corolla calyce dimidio longior, tubo angusto. Fructus 0
utrinque acuminatus octangularis.

deus

; : ae juniorum
Arbor ro-metralis cortice suberosa rimosa. Fasciculi foliorum arcte
oblanceolato-linearium 20-28¢m longorum ad apicem versus ease * hana.
; : : la :
conferti. Folia provectiora usque ad 76°™ longa 11-13°™ lata chartaces :

a a a a

1909] SMITH—PLANTS FROM CENTRAL AMERICA 259

nervis lateralibus fortioribus utrinque 28-36, petiolis 6-8™™ longis tumidis cor-
ticatis. Nodi in ligno vetere floriferi squamis (seu bracteis pedunculos olim ful-
cientibus) concavis lanceolatis acutis armati, pedunculis 2-4°™ longis. Calyx

™ Jon orolla 28™™ longa, plica retroflexa. Stamina 14-16™™ longa,
loculis 4™™ longis. Discus 2™™ altus 3™™ latus. Ovarium 4™™ longum, stylo

Cubilquitz, Depart. Alta Verapaz, Guatemala, alt. 350™, Sept. 1904, H. von
Tuerckheim, n. 8723 ex Pl. Guat. etc. quas ed. Donn. Sm. Sub Crescentia olim
distributa—Ad ripas rivuli Chit prope Cobdén, Depart. Alta Verapaz, Guate-
mala, alt. 1350", Maj. 1908, H. von Tuerckheim (n. I. 2278).

Neotuerckheimia gonoclada Donn. Sm.—Folia singula cuspidato-

acuminata deorsum attenuata. Pedunculi terminales 2-5-ni bracteis
pluribus linearibus comitati. Corollae tubus late oblongus, limbus
glanduloso-punctulatus.
_ Ramuli angulati, cortice exfoliato. Folia superiora approximata, inferiora
internodiis 2-4°™ longis remota opaca 19-30°™ longa 5-7°™ lata, nervis lateralibus
fortioribus utrinque 20-24, petiolis vix 2-3™™ longis crassis. Pedunculi 2.5-4°™"
longi, bracteis 6-9™™ longis canaliculatis deciduis. Calyx 22" longus. Corolla
nm longa e schedula Tonduziana pallide flavicans, plica retroflexa 2.5™™ lata.
Stamina 16-18™™ longa, loculis 4™™ longis. Discus 2"™ altus 4™™ latus. Ova-
Tum 4™" longum costis 8 angulatum, stylo 28™™ longo. Cetera desunt.

In silvis ad La Palma, Proy. San José, Costa Rica, alt. 1460™, Sept. 1898,
Ad. Tonduz, n. 7384 ex Pl. Guat. etc., quas ed. Donn. Sm. (n. 12563 herb. nat.
Cost.). Sub Crescentia olim distributa. ;
Justicia (§TyLocLossa Lindau) multicaulis Donn. Sm.—Folia
inter minora thomboideo- vel oblongo-ovalia vel lanceolata utrinque
acuta. Flores axillares sessiles singuli vel subterminales et >
interdum breviter spicati. Corollae tubus cylindricus limbo ampliato
dimidio longior. Antherarum loculus inferior ab altero remotus
“ppendicula ovali herbacea munitus.
tea nanus, caulibus e basi pluribus 25-33°" longis, ramis ogg
a a bifariam pubescentibus lineolatis. Folia pa nite an
rath lata, interdum 34™™ longa 7™™ lata, cystolithis farcta preeter waded

es ee los glabra, nervis laterlibus utrinque 4-5, petiolis bifanam Lge 3,
ae longis. Bracteae spatulato-obovatae 7”™ longae cum bracteolis linear!
baie 5"™ longis herbaceae glabrae lineolatae. Calycis segmenta 5 is
labie 7 . longa. Corolla 16™™ longa alba, tubo gracile igi none? cee
a bidentato violaceo utrinque pubescente, antici paulo Se hs
ity * intermedio maximo 4™™ longo atque lato. Stamina summis a

6™m longa, antherarum loculo altero usque ad o.5™™ inferius affixo periecto.

260 BOTANICAL GAZETTE [APRIL

Capsula oblonga acuminata 6"™ longa abortu disperma, retinaculis acutis, semi-
nibus disciformibus rugosis alato-marginatis.—Corollae indole anormalis.

Ad ripas rivuli, Pansamal4é, Depart. Alta Verapaz, Guatemala, alt. 1250”,
Jun. 1885, H. von Tuerckheim, n. 741 ex Pl. Guat. etc., quas ed. Donn, Sm.
In silva montana prope Coban, Depart. Alta Verapaz, Guatemala, alt. 1600”,
Jan. 1908, H. von Tuerckheim (n. II. 2091).

Ruprechtia (§ PseupoTRiPLaRIs Benth.) Kellermanii Donn. Sm—
Folia subsessilia elliptica acuminata basi obtusiuscula subglabra
minute inconspicueque reticulata. Racemi fasciculati, bracteis pedi-
cellos superantibus. Perianthium fructiferum inter minora totum
cinereo-atque-cano-sericeum, segmentis exterioribus linearibus, inte-
tioribus dimidio adnatis. Achenium substipitatum tenuiter elongato-
conoidale.

Folia coriacea 11-13.5°™ longa medio 5.5-6°™ lata, venulis retiformibus
tantum ope lentis manifestis et puberulis, petiolis vix 2™™ longis. Racemi fructi-
feri densiflori pilosi, rhachi 1. 5—3.5°™ longa, bracteis orbiculari- vel oblongo-ovatis
5-6™™ longis fuscis puberulis, pedicellis 4™™ longis cano-pilosis prope apicem
articulatis. Perianthii 18-21™™ longi tubus oblongus 7-9™™ longus, segmenta
exteriora apice 3™™ basi 2™™ lata obscure trinervia et reticulata, interiorum pars
libera 3™™ longa. Discus staminum sterilium 1™™ altus g-denticulatus. Ache-
nium 9™™ longum 2.5™™ Jatum in imam basin stipitiformem 1™™ longam ange
Statum prope apicem acute triquetrum et ciliatum ceteroquin rotundato-trilobum,
lobis sulcatis glabris, stylis 1™™ longis stigmata acquantibus. Cetera cee
R. Deamii Robinson ejusdem loci incolae valde affinis.—Beato KELLERMAN
florae Guatemalensis exploratori indefesso dicata.

Gualin, Depart. Zacapa, Guatemala, alt 122™, Dec. 1995; W. A. Kellerman
n. §985.—Typus in herb. Musei Nationalis servatur.

SPECIEM AMERICAE AUSTRALIS INCOLAM LICEAT HIC ADJUNGERE

Ruprechtia (§ PseuporripLarts Benth.) colorata Donn. Sm.—-Folia
oblongo-ovata vel -ovalia apice acuta vel obtusiuscula basi pases
vel obtusa glabra subtiliter reticulata. Racemi .singuli, pear ;
gracillimis. Perianthium fructiferum rubiginosum, segmenns apes
oribus lineari-spatulatis glabrescentibus, interioribus liberis- Ac
mum lanceolatum acuminatum basi obtusum.

em Jata, venulls

fructiferi solum visi sublaxiflori pubescentes, rhachi 2-4°™ longa, DI
dio articulatis.

* Obiit in Guatemala m. Mart. ann. 1908 febri confectus.

1909] SMITH—PLANTS FROM CENTRAL AMERICA 261

anthii 27-29™™ longi tubus obovatus 5—6™™ longus pubescens, segmenta exteriora
apice 5™™ basi 2™™ lata pulchre trinervia et reticulata, interiora 4™™ longa.
Discus staminum sterilium o.5™™ altus g-denticulatus. Achenium glabrescens
g™™ longum 3™™ latum triente superiore acute triquetrum inferne rotundato-
trilobum, lobis leviter sulcatis, stigmatibus 2™™ longis stylos 3-plo superantibus.
—R. juscae Fernald et R. Cumingii Meisn. affinis perianthis etiam in sicco laete
colorato insignis.

In collibus sabulosis prope litora oceani, Savanilla, Republica Colombiana,
Febr. 1896, J. Donnell Smith.

Daphnopsis ({NoRDMANNIA Benth. et Hook.) monocephala Donn.
$m.—Folia sessilia oblanceolata apice rotundata infra medium sensim
attenuata supra glabrescentia subtus cano-sericea. Pedunculus sub-
terminalis solitarius simplex gracilis cum capitulo unico cano-sericeus.
Perianthium masculinum subsessile anguste infundibuliforme, lobis
obtuse ovatis tubo 3-plo brevioribus intus glabris, squamulis 4 ovalibus.
Rami dichotomi ad apicem versus cano-villosi et confertim foliacei. Folia
juniora utrinque villosa, provectiora (saltem superiora) 5~6.5°™ longa 1.5-2°™

coriacea, nervis lateralibus utrinsecus 5-6 longe ascendentibus et reticulis

rudimento pistilli glabro triplo brevioribus. Perianthium femininum ignotum.
El Rancho, Depart. Baja Verapaz, Guatemala, Jan. 1906, W. A. Kellerman

Euphorbia (§ ALEctoroctonum Boiss.) adinophylla Donn. Sm
Glabra. Folia in quoque verticillo indefinite compluria rhomboideo-
cliptica apice obtusa vel rotundata basi acuta petiolos subaequantia.
gaa axillares et terminales corymbiformes petiolis subaequilongae-
Wolucrum graciliter pedicellatum, glandulis 5 appendice subcre-
nulata paulo angustioribus. Styli brevissime lobulati.

: ramidatae pedunculo 5-7™™ longo computato 2275 mm Jongis
mi me repetitus dichotomae, foliis floralibus oblanceolatis 3-7 : 8 ,
otomiis glandula magna instructis, axibus uti pedicelli 2-5" = se?
uer i . per uatidlais 16-2 longed inf Icatus, lobi minut! obovatl
briati, glandulae transverse ellipticae ab appendice arcuata margin ayer
R “a longe stipitatum, stylis indivisis 1.5™™ longis. c — a ue
Schlechtendalii. Boiss. proxima differt foliis pro rata longioribus ad nocos

262 BOTANICAL GAZETTE | [aren

fertioribus, inflorescentia evoluta, pedicellis elongatis, stylis indivisis—Ab incolis
Carana (Latine Resina) vocatur.

Republica El Salvador, loco natali haud accuratius addicto, ann. 1905, ase
Rénson (n. 187).—Typus in herb. Musei Nationalis servatu

GUZMANIA BRACTEOSA André.—Folia auctoribus adhuc ignota
linearia 30-55°™ longa acute attenuata medio 13-25™™ lata in
vaginam 25-5o™™ latam sensim dilatata tenuiter coriacea utrinque
praesertim subtus pallide lepidota vel glabrescentia, nervis crebris
utrinque prominentibus.

In silvis ad Las Vueltas, Tucurrique, Costa Rica, alt. 635™, Mart. 1899, Ad.
Tondusz (n. 13291). —Epiphyta (ex cl. repertore) in silvis montanis prope Purulé,
Depart. Verapaz, Guatemala, alt. 1800™, Apr. 1907, H. von Tuerckheim
(n. If. 1826).

eee MARYLAND

COMPARATIVE HISTOLOGY OF FRUITS AND SEEDS
OF CERTAIN SPECIES OF CUCURBITACEAE

KATE G. BARBER

(WITH FIFTY-THREE FIGURES) ’
Introduction

This investigation was undertaken with the view to supplementing
: the work already done on the histology of the fruits and seeds of
| Cucurbitaceae. The literature, although not meager, treats only of
the most conspicuous elements of the spermoderm, ignoring, with a
few exceptions, the less important tissues of the seed and the whole
of the pericarp.

In the following paper are included additions to the histology of the
spermoderm and new descriptions of the pericarps of the common
species; also brief descriptions of six interesting seeds previously
undescribed.

I wish to acknowledge my great indebtedness to Dr. A. L. Win-
TON, under whose guidance and inspiration this work was carried on,
and also to Professor A. W. Evans for advice in its preparation.

Résumé of the literature
In 1833 BiscHorr™ published two cuts showing cross-sections of
the seed coats of Cucurbita Pepo and C. Lagenaria, which, according
(0 FIckEL, leave much to be desired with regard to detail.

Von HéuNeL; was the first to publish a detailed description
seed coats of the Cucurbitaceae, the species studied being Cue
¢po L.., Lagenaria vulgaris Ser., and Cucumis sativus L.
a ae ugh investigation of the integuments, from Be .
inner j e flower until maturity, led him to conclude that: :
pe cpent (three to four layers thick) develops but s
oming the collapsed, thin-walled inner parenchyma of the see

figs. 1872, 18734.
menschalen einige

of the
urbita

re fertiliza-
(x) the
lightly,

Handbuch der bot. Terminologie und Systemkunde. seed
die Anatomie und Entwickelungsgeschichte der s
n. Bot. Zeit. 34:738. 1876.

Mo ; ‘
und ein; hologische Untersuchungen iiber die Samens
einiger

263]

2 Ueber
Cucurbitacee
chalen der Cucurbitaceen
verwandter Familien. Sitzb. Akad. Wiss. Wien, Mathem.-Naturw-
6.

[Botanical Gazette, vol. 47

264 BOTANICAL GAZETTE [APRIL

coat; and (2) the outer integument (eight to ten layers thick) forms
the greatly differentiated outer layers of the coat.

At maturity he found that the spermoderm consists of a compli-
cated structure with five distinct tissues, namely: (1) inner epithelium
of the carpel; (2) outer and ( 3) inner integument; (4) perisperm;
and (5) endosperm. These five in turn are subdivided into ten
layers, each one to many cells thick. About the edge of the seed,
between the fifth and sixth layers, runs the small raphe.

This author divided the family into two large groups. In the
first belong those seeds with the epithelium of the carpel firmly
attached to the spermoderm, represented in his work by Cucurbita
Pepo and Lagenaria vulgaris. The second includes the species with-
out this epithelium, illustrated by Cucumis sativus. Otherwise there
are no great differences in development.

In the same year, one month after the appearance of von HOHNEU'S
paper, FickEL? published his inaugural dissertation, describing the
seed coats of Cucumis sativus L., C. Dudaim Lae. myriocar pus
Naud., Cucurbita Pepo L., C. melanosperma A. Br., Lagenaris
vulgaris Ser., Citrullus vulgaris Schrad., Benincasa cerifera Savi.
Bryonia alba L.., B. dioica L., Ecballium agreste Rchb., Sicyos ang
latus L., Cyclanthera explodens L., C. pedata Schrad., and Bryonopss
erythrocarpa. He gives descriptions and cuts of the mature seed
coat and its development from the two integuments, and reaches the
following conclusions: (1) the seed coat has five layers; (2) the oe
dermal cells are radially elongated and, with the exception of ore
Sicyos and Cyclanthera, have thickenings of various kinds on t ¥
radial walls; (3) the second layer consists of one or more siete
cells varying in size and thickness of walls; (4) a third nye :
either radially or longitudinally elongated cells; and (5) the remain
ing layers are of compressed cells without definite structure. — .

GopFrin‘ briefly notes the seed coats of Cucumis eee
Cucurbita maxima Duch., Lagenaria vulgaris Ser., Sicyos “ es

on t
L., and Cyclanthera pedata Schrad. In his general description oe
Spermoderm is represented as having six layers, as follows: oS
mis of prismatic cells; (2) small cells differing as to number, 51%

; ; es.
: 4 Etude histologique sur les téguments séminaux des angiosperm
Nancy 1880: 160,

1909) BARBER—FRUITS AND SEEDS OF CUCURBITACEAE 265

thickness of walls; (3) a single layer of sclerenchymatized cells vary-
ing but slightly in all the species; (4) outer and inner parenchyma;
(5) inner epidermal cells, often separated from the preceding layer
by longitudinally elongated cells; and (6) inner row of thick-walled
cells with granular contents, the author evidently including the
endosperm in the spermoderm. ‘This is followed by a description of
each of the species already mentioned.

Von Héunet’s division of the family into two groups, distinguished
by the presence or absence on the spermoderm of the inner carpellary
layer, is criticized, as GODFRIN always found this layer present in
fresh material. He used the following grouping, based on the elonga-
tion of the cells of the “protective” (third) layer: (1) tangentially
elongated cells (Cucumis, Lagenaria, Cucurbita, Citrullus); and (2)
radially elongated cells (Sicyos and Cyclanthera). The individual
descriptions are brief, many details being omitted.

Fiscer’, in his classical work on sieve tubes of the Cucurbitaceae,
describes a detailed study of the elements in all parts of the plant.
The tubes are of four kinds: (x) bundle sieve tubes (usual type found
in fibro-vascular bundle); (2) ectocyclic (isolated tubes of hypoderm) ;
(3) entocyclic (isolated tubes of the inner tissues); and (4) commisural
(connecting those of the bundle with the isolated tubes). In the
young isolated tubes the contents are thickened but in the old there
Sa watery slime secreted by the neighboring cells. These isolated
tubes, being free from callus plates, were mistaken by earlier writers
for latex tubes,

Six plates are given illustrating these tubes in Cucurbita Pepo,

smaria vulgaris, Sicyos angulatus, Cyclanthera pedata, Melothria
ee Bryonia alba, and Luffa pentandra. : aa
C Met, HANAUSEK® gives chemical analyses of Cucumis sativus L.,
Du ms 9L., C. Citrullus L., Cucurbita Pepo L., and C.- mer

» and mentions briefly the histology of each.
Citra 7 describes and illustrates cross-sections of species of Cucumis,
divj a Lagenaria, and Cucurbita. The numerous —— on
Into races, varieties, and subvarieties, with long lists of syno
Berlin. 1884.
Kassel. 1884-

oe iiber das Siebréhren-System der Cucurbitaceen-

? a Nahrungs- und Genussmittel aus dem Pflanzenreiche 195-
hdwirthschaftliche Samenkunde 767. Berlin. 1885.

266 BOTANICAL GAZETTE [APRIL

nyms under each. The author briefly notes the macroscopic appear-
ance of the fruit, then gives short histological descriptions of the seed.
According to his division the spermoderm has five layers. Some of
the details of structure of the outer layers, and nearly all of the inner
ones, are omitted.

VocL* devotes but a single paragraph to a microscopical descrip-
tion of Cucurbitae.

BRAEMER?® in 1893 studied the stem, leaf, pericarp, and seed of
Bryonia dioica Jacq., Ecballium Elaterium Rich., and Citrullus
Colocynthis Schrad., but confined his attention chiefly to a micto-
chemical investigation of the contents of the sieve tubes and latex
tubes, both of which he regards as latex tubes, accepting the
descriptions of the tissues given by earlier writers. According -
his introduction, the pericarp has an outer and inner epidermis with
a sarcocarp between. Within the spermoderm, having hard and
lignified layers, are inclosed an embryo, reduced to a small radicle
and two cotyledons, rich in oil and aleurone grains, and the remains
of the perisperm and endosperm. ab

The main part of the paper is devoted to the long, ramifying,
straight or sinuous tubes containing a yellow semi-fluid, finely aa
lar, refractive substance which entirely fills the cavity. Both =.
and cross walls are composed of cellulose without any indienne? .
sieve or callus plates. In their morphological and chemical char-
acters they resemble the latex tubes of Convolvulaceae and Campanv-
laceae, and the isolated sieve tubes of FiscHER. In these
BRAEMER found the three “active principles” bryonin, i) il
and elaterin, - .

PLANCHON and CoLLin'® treat the species used as drugs: al
scopic and microscopic sections are figured, but no cacti
details are given, of the fruits and seeds of Cucurbita P id Blat
Citrullus Colocynthis Schrad., Cucumis sativus L., Ecballium
rium Rich., and Fevileae.

Vittters and Coxuin'® use the bicollateral bundles

* Pharmakognosie 196. Wien. 1892. -

° De la localisation des principes actifs des Cucurbitacées. Tou
'° Les drogues simples d’origine végétale 2:292. Paris. 1896.
" Traité des altérations et falsifications des substances alime

and sieve

louse. 1893-
ris.

ntaires 454 ,

TQgoo.

1909] BARBER—FRUITS AND SEEDS OF CUCURBITACEAE 267

tubes (or latex tubes) in the detection of pumpkin pulp as an adulter-
ant of preserves. They have an original cut, but their description of
these elements is taken from FiscHERS and BRAEMER.®

T. F. HaANAusEK”? gives one figure illustrating the characteristic
elements of the seed of Cucurbita Pepo L., C. maxima Duch., and C.
moschata Duch. ‘These elements are the Shleriden (resembling the
epidermis of Spanish pepper), spongy parenchyma, starch grains of
the inner parenchyma, endosperm and cotyledon tissue with aleurone
grains.

ARTHUR Meyer’: refers briefly to the aleurone grains of Cucurbita
Pepo as having a diameter of 1-4 # (mostly 3 #) and small crystalloids.
In his table taken from LuptKE,"4 the seeds of Citrullus and Colocyn-
this are stated to contain numerous aleurone grains 1-7-5 # in diam-
eter, each having one globoid (o. 5-1 #) and one crystalloid (1-3 #)-

BOHMER'S gives a chemical analysis of the seeds of the Cucurbitae
and one figure showing a cross-section and surface view. His brief
histological description is taken from other authors.

COLLIN and Perrot"? describe briefly and give a cross-section of
the seed of Cucurbita Pepo Duch.

According to MoELLER'? the fruits of Cucurbitaceae are all
large berries with hard shells, soft fruit-flesh, and many seeds. The
seed coat is divided into four tissues: (1) palisade cells with thicken-
ings on radial walls; (2) stone-cell layer; (3) stellate parenchyma;
and (4) thin-walled parenchyma. A brief description follows of
Cucurbita Pepo L., C. maxima Duch., C. moschata Duch., Cucumss
‘alious L., C. Melo L.., and Citrullus vulgaris Schrad., with cuts of the
last two.

'? Lehrbuch der technischen Mikroskopie 370. Stuttgart. 1901. Translation,

INTON, The microscopy of technical products 369. New York. 1907-

‘S Die Grundlagen und die Methoden fiir die mikroskopische Untersuchung von
Or.

Pflanzenpulyern g2.° Jenar

me Ueber die Beschaffenheit der Aleuronkérner einiger Samen. Ber. pee
Tm. Gesells. 1891 :56—s0.

ont ae

lichkej

Kraftfuttermittel, ihre Rohstoffe, Herstellung, Zusammensetrane Ve
a eg Verwendung, mit besonderen Beriicksichtigung der ‘omni arial
ie opischen Untersuchung 508. Berlin. 1903-
Les résidues industriels 270. Paris. 1904.

: Mikroskopie der Nahrungs- und Genussmittel 479. ?- Aufl. Berlin. 190°5-

268 BOTANICAL GAZETTE [APRIL

BARBER,*® in a chapter on Cucurbitaceae, gives brief descriptions
of Cucurbita Pepo L., C. maxima Duch., Cucumis sativus L., C.
Melo L., and Citrullus vulgaris Schrad., illustrating the first-named
species by three original cuts.

Geographical distribution

The Cucurbitaceae are scattered over the greater part of the
earth’s surface, but reach their highest development in the tropics.
Representatives are, for the most part, absent in the colder regions
of the temperate zone, the two exceptions being Sicyos angulatus,
found in Canada, and Echinocystis lobata in New England.

The total number”? of genera is eighty-five, the Old World claiming
fifty-four and the western hemisphere thirty-eight, while seven are
common to both. Under each there are long lists of species, sub-
species, and varieties, authors differing greatly as to the number.

EDIBLE sPECIES.—Many species have long been cultivated for
food. Probably one of the oldest on record is Citrullus (watermelon),
a favorite with the ancient Egyptians. Other important species 47°
Cucumis sativus (cucumber), eaten in China as a vegetable more
than two centuries before the Christian era; Cucurbita Pepo (pump-
kin), grown by the aborigines in America; and Cucumis Melo (musk
melon), greatly prized in Asia and Africa. These have now spread
to all the warmer regions of the earth.

ORNAMENTAL SPECIES.—There are many varieti
curious form and vivid coloring of the fruit, among which are a
mordica balsamina (balsam apple), Lagenaria vulgaris Ser. (common
gourd), Cucumis erinaceus (hedge-hog gourd), Cyclanthera explodens
(squirting cucumber), and Lu/fa cylindrica (dish-cloth gourd). both

PHARMACEUTICAL sPECIEs.—Among those used for drugs, -
here and in Europe, are Bryonia dioica (bryony), Citrullus oe
this (colocynth), and Ecballium Elaterium (elaterin).

es grown for the

General characters
MACROSCOPIC ae
FLOWER.—The axillary flowers, yellow or white 1 color, |
borne either solitary or in groups of various kinds. They are
8 In Winton, Microscopy of vegetable foods 4or. New York. 1906.
10 ENGLER UND PRANTL, Pflanzenfamilien IV. 5:9-

1909] BARBER—FRUITS AND SEEDS OF CUCURBITACEAE 269

monoecious, epigynous; calyx and corolla actinomorphic, adnate at
the base. The stamens are five in number, four of which frequently
cohere in pairs, or more rarely they all unite to form a column. The
ovary is one- to five-, usually three-celled.

Fruir.—In the Cucurbitaceae are found the largest fruits of the
whole plant kingdom. From the enormous berry of Cucurbita, reach-
ing a maximum of several kilos in weight, there are all gradations
down to the small burr of Sicyos angulatus, only slightly larger than
the single inclosed seed.

The fruits are nearly always fleshy berries; they may, however,
become membranaceous and dry, preserving at the same time their
original shape. There is no characteristic form for the family, the
shape differing as much as the size. All possible variations of spheri-
cal, elliptical, greatly elongated, and curious unsymmetrical forms are
tepresented. They are smooth, warty, or covered with spines, or
various other kinds of emergences. In addition to these outgrowths,
the young fruits of all the common species bear one or more forms of

aits which, in most cases, persist at maturity. The color varies from _
white, green, and yellow to red, with spots and stripes on some of the
Varieties.

All the fruits described in this paper are indehiscent with the
exception of Echinocystis, which bursts irregularly at the top.

The pericarp, or rather the pericarp and adherent receptacle,
ae in thickness (thin or thick rind, or solid fruit-flesh), color
(white, green, yellow, or red), and texture (watery or dry, sclerenchy-
matized or soft). The central placentae extend to the outer wall
and divide, turning back so as to give a parietal appearance.

SEED.—The seeds, borne either singly or in great numbers, are
‘natropous, large, ovate, and flattened, with or without a border
\formed by elongated epidermal cells) on each side at the edge. They
Vary in shape from the narrow and pointed to the broad, rounded, or
Tectangular forms, and in color from white to brown and black.

MICROSCOPIC
The pericarp may be divided into six, more or less
tinct tissues as follows:
1. Epicarp.—The cells are for the most part polygonal, forming

270 BOTANICAL GAZETTE [APRIL

a palisade layer, those about the stomata being frequently elongated.
The outer and radial walls are cuticularized and occasionally colored.

2. Hypoderm.—Few or many layers of isodiametric cells form the
tissue beneath the epicarp. The cells are somewhat thickened and
occasionally pitted.

3. Outer mesocarp.—Isodiametric cells with either cellulose, or
sclerenchymatized and pitted, walls (stone cells, etc.) form a sharply
defined zone varying from a few to many cells in thickness.

4. The middle mesocarp consists of large, usually isodiametric,
thin-walled cells, often turgid with a watery cell sap and containing
a small amount of starch. The starch grains are small, with an
average diameter of tom. They are truncated, frequently occurring
in aggregates of two and three, with slightly eccentric hilum and faint
rings. Polarization crosses are very distinct.

5. Inner mesocarp.—Several layers of thin-walled cells, forming
this tissue, closely resemble the preceding layers. The cells are
small and have no visible contents.

6. Endocarp.—Very small, thin-walled, tangentially elongated cells,
arranged side by side in groups, form a thin transparent tissue. With
the exception of Cucumis, this layer remains so firmly attached to
the dry seed that some authors describe it as the outer layer of the
spermoderm.

The anastomosing bundles occurring throughout the mesocatP are
bicollateral. They are either small and soft, or large and stiff, form-
ing a conspicuous network. ‘The elements consist of spiral, annular
and reticulated vessels, and sieve tubes having large sieve plates
evident without staining.

Isolated sieve tubes and latex tubes.??—FISCHERS claims there are
no true latex tubes, those known by this name being simply siev*
tubes that have ceased to function; BRAEMER? and other Tater
authors, however, use the term “latex tubes.’ I have seen the pet

2° According to DE Bary (Comparative anatomy of the vegetative organs =
phanerogams and ferns 198. Oxford. 1884) DrppEt finds the septa and a ween
’ articulated tubes provided with sieve plates and thinks them intermediate ont
Steve tubes and latex tubes. De Bary himself finds not the plates but il bes.
Scattered over the whole wall, and further states that there are two kinds of latex a
which do not correspond in function: (1) those secreting tannin, etc-, and (2) those
milky plants) which are closely related to sieve tubes.

1909] BARBER—FRUITS AND SEEDS OF CUCURBITACEAE 271

forated plates in some tubes and have not found them in those con-
taining granular milky contents which harden in alcohol, therefore
shall mention both in the following descriptions. These elements
occur in considerable numbers throughout the middle tissues of the
pericarp.

Each seed has a firm spermoderm of many layers, a thin collapsed
perisperm and endosperm, and a large embryo consisting of two large,
flat, leaf-like cotyledons and a small radicle.

SpeRMODERM.—Authors variously state the number of layers ip
this leathery coat or shell as four’? to ten.?* I myself consider it as
consisting of five distinct tissue layers, the second being occasionally
differentiated into two, and the fourth often into two or three forms
of cells arranged in as many layers. It is developed from the two
integuments (fig. 10), the outer integument forming the three outer
and part of the fourth layers, the remainder of the coat developing
from the inner integument.

1. The epidermis consists of a single layer of prismatic palisade
cells, polygonal in surface view. They are usually of equal height
over the flat surface of the seed, increasing in height at or on both sides
ofthe edge. The radial walls of a few species (as Echinocystis lobata)
are uniformly thickened; in all the other seeds they have either straight
or branched thickenings running from the inner to the outer tangential
walls. The outer walls, and frequently the inner, are thickened.

Voct,® the only author that notes the presence of starch in this
layer, gives no description of the grains. According to my ag
observation they are small, globular, reaching a maximum diameter
of 7 #, the larger ones showing a central hilum but no rings. They
polarize very indistinctly.

2. Subepidermal layer.—One or more layers of sclerenchymatized
cells, varying greatly in size and shape, make up this layer. The
cells are either small, pitted, without intercellular spaces (Cucurbita),
or longitudinally elongated, arranged end to end in rows, with numer-
etc intercellular spaces (Cucumis), oF form 4 oe

ot greatly thickened irregularly arranged cells. No contents
are evident.

3. Sclerenchyma.—This consists of a layer of exceedingly thick-

* ENGLE Pe :
NGLER UND PRANTL, Pflanzenfamilien IV. 5:5-

272 BOTANICAL GAZETTE [APRIL

walled cells elongated either tangentially and arranged end to end in
longitudinal rows, or radially forming a palisade layer. The walls
are sinuous with numerous pits, and after maceration the margins of
the outer and inner tangential walls are seen to be lobed, each lobe
branching with the ends overlapping and fitting together to form a
flat compact layer (fig. 18). The empty cell cavities are compara-
tively small and present a striking oval appearance in median
section.

4. Parenchyma.—Many cell layers of spongy parenchyma, dif-
fering greatly in size and shape, make up this layer. The cells of the
outer layers are usually small and frequently sclerenchymatized and
pitted. Within these small cells are one or more layers of either large
or small stellate cells having very large intercellular spaces. Thett
thin walls are usually sclerenchymatized and often pitted or reticu-
lated (Cucurbita). The remaining layers consist of small, thin-
walled spongy parenchyma cells containing chlorophyll in some
species. :

The small raphal bundles are found imbedded in this parenchyma
usually about the edge only; a few exceptions, however, show branches
on the flattened surface of the seed.

5 Inner epidermis.—A single layer of small, polygonal, and incom
spicuous thin-walled cells forms the inner tissue.

PERISPERM.—This thin coat is collapsed in the mature fruit, but
treatment with Javelle water brings to view several layers of small,
very thin-walled cells covered with an epidermis having cuticularized
outer and radial walls. No contents are evident. a

ENDOSPERM.—This consists of one layer of very thick-w
polygonal cells, containing oil and protein granules. A few excep"
tions (as Citrullus vulgaris) have within this layer several more layers
of empty thin-walled cells. :

MBRYO.—The leaf-like cotyledons have an epidermis 0 ©
cells below which, on the inner side, are two sharply defined p vil
layers. Procambium bundles run through the small-celled mesoP ning
All of the cells are filled with oil and protein granules . ally
globoids and crystalloids which are very minute and of gree
no diagnostic importance.

s of small

1909] 1BARBER—FRUITS AND SEEDS OF CUCURBITACEAE 273

Specific characters.
Cucursira Pero L.

Cucurbita Pepo L. (pumpkin), according to the earlier writers,
was introduced from southern Asia, but WITTMACK, in his recent inves-
tigations of prehistoric remains in Peru, claims it to be a native of
America. Early explorers also recorded the cultivation of this
fruit by the aborigines in their maize fields.

The fruit is one of the largest of the Cucurbitaceae, occasionally
reaching a weight of 200 kilos. It is a smooth, apple-shaped berry,
with about twenty, more or less pronounced, longitudinal grooves,
the color varying from yellowish green to orange. There are many
subspecies, differing greatly in size and shape, of which Harz enumer-
ates thirty. Among them are the small to enormously large spherical,
ellipsoidal, flask-, egg-, J-, and curiously-shaped forms. Most of the
larger ones are cultivated for food, the smaller as ornamental fruits.

At maturity the fruit consists of a hollow, yellow rind, 2-3°™ in
thickness, containing a tangle of slimy fibers,
among which are the numerous _ flattened

The white seeds (fig. 1) are 1.5-2.5°™ im
length, elliptical, flattened, and have a narrow
border about the edge on both sides. Fic. 1.—Cucurbita

PERICARP.—This makes up the bulk of the PepoL. Seed. Xt
fruit. It includes the rind and fibers, the con-
necting parenchyma breaking down before the fruit reaches maturity.

1. Epicarp (figs. 2, 3, epi).—The prismatic cells form a palisade
layer about 50m in height, with outer and radial walls greatly thickened,
cuticularized, and colored bright yellow. In surface view they are
polygonal (14m in diameter), except at character i
about which they are elongated and curved. These white spots con”
sist of a stoma of the common type, from which radiate TOWS of
tangentially elongated epidermal cells. The stomata art —o
formly distributed and occasionally two are found in the same group
of Tadiating cells,

On the very young fruit, even before the fertilization of the ovary,
two forms of hairs are found (fig. 3). Both arise from 4 foot-cell

differing from the neighboring epidermal cells only in the more

274 BOTANICAL GAZETTE [APRIL

rounded shape. The hairs dry up and disappear while the fruit is
still very small, leaving the foot-cell intact. Later the epidermal cells
divide, thicken their walls, and press against the foot, which thus be-
comes polygonal. As both kinds of cells are of the same size, the
two are indistinguishable, which accounts for the absence of hair
scars on the mature fruit.

The first form of hair is jointed (¢*) and of great size, often reach-
ing a length of 1.5-2™™. The diameter increases rapidly (up to
about 85 ) for a short distance from the foot, then gradually tapers,
forming a long, thick-walled (5 “), conical hair. The first few cells

are two or three times as broad as long, while the following cells
become elongated as the hair increases in size. .
The second form is capitate (¢?), and like the first increase ig
diameter from the foot, forming a jointed stalk of four or hye
It ends with a large spherical head of one or more cells, often 54 Mg
diameter, : .
2.-H ypoderm.—Many cell layers of exceedingly small, —
cells, 20 in diameter, form this layer. The cells have thick W zi
occasional pits, and small intercellular spaces. The visible conte a
consist of numerous protein granules. This layer closely es™ ).
the corresponding one of C. Pepo var. verrucosa Naud. (f é pies
3. Outer mesocarp.—Within the preceding layer is 4 ca erm
Posed of cells graduating in size from the small cells of the hypod

1909]

BARBER—FRUITS AND SEEDS OF CUCURBITACEAE

275

tothe large ones of the middle mesocarp. They are isodiametric, have
thick walls and intercellular spaces, and but few contents.

4. Middle mesocarp (fig. 4).—
In this layer the cells become
gradually larger, thicker-walled,
and more loosely arranged. They
contain small starch grains (a7),
reaching a maximum size of 10 #.
The outer cells contain a con-
siderable number of these grains,
but the increase in size of the cells
is accompanied by a decrease in
amount of starch.

5. Inner mesocarp.— This
layer consists of rather large-
celled parenchyma with no evi-
dent contents.

Bundles and isolated sieve
tubes, together with anastomos-
ing latex tubes (Jaé) are found
throughout the mesocarp, while
in the center of the fruit there
IS a great mass of tough fibers,

mmature
epi, epicarp
#2, capitate

Fic. 3.—Cucurbita Pepo 1.1

view;
hair,
160.

epicarp in surface
with ¢1, jointed conical
hair, and sto, stoma. x

surrounted by the remains of the broken-down parenchym2.

iN
F ;
ring co Pepol.. Cross-
Starch of mesocarp showing am,
X 160 Stains, and Jat, latex tube.

6. The endocarp appears on the
seeds as a thin membrane of long}-
tudinally elongated cells, arranged
end to end in rows. The inner wall
is about the same thickness as the
outer wall of the epidermis of the
to which it is quite
from the dry seeds,
be readily separated
as a thin, transparent, colorless skin.

SpERMODERM (fig5 5: 6).—The
five layers making UP the sper-
moderm are as follows:

spermoderm,
firmly attached;
however, it may

276 BOTANICAL GAZETTE [APRIL

1. Epidermis (ep).—The prismatic cells form a continuous layer
without intercellular spaces. They are radially elongated, on the
flat side of the seed to 270 m, on the edge to 50 m, while the border con-
sists of a ridge of cells several times as high as those of the flattened
‘surface. The outer wall is thickened, but has no cuticle, and the radial
walls are very thin, with peculiar branching thickenings of cellulose.
The thickenings run from the base of the cell, one on each wall, as
single straight rods until near the outer wall, where they branch pro-

. . . . i .
fusely, Siving rise to a beaded appearance in tangential ser oa

Maceration, or in scrapings of the spermoderm, the rods f ‘6
their sides, Presenting the appearance shown in fig. 5; ef f
seeds the walls of this layer are frequently broken down, and i . ;
few of the thickenings remain, which may be mistaken for the
surface of the walls themselves. already
Round starch grains (am), up to 7 u in diameter, of the Be :
described, occur in considerable numbers in the epidermal ce"

mae
2. Subepidermal layer (hy).—Below the epidermis is 4 !#Y°

1909] BARBER—FRUITS AND SEEDS OF CUCURBITACEAE 277

small, polygonal, somewhat elongated cells, about 25 in diameter,
with numerous minute pits, giving the walls a beaded appearance.
This layer, without intercellular spaces, is three to five cells thick,
the number increasing toward the edge of the seed.

3. Sclerenchyma (scl)—Firmness is given to the spermoderm by
one layer (over the edge two or three) of longitudinally elongated
(250) cells, arranged end to end in rows. In surface view, both side
and end walls are sinuous, often reaching 204 in thickness. After
maceration and by careful focusing, the outer and inner surfaces of
these cells may be seen to send out remarkable ramifications, whose
dichotomous branches overlap one another. In transverse section
the oval appearance of the cell cavity is a characteristic feature. A
few pits may be seen with a high power.

4. Parenchyma.—There are three quite distinct layers, the paren-
chyma showing the greatest differentiation in this species. In contact
with the sclerenchyma is the first layer of small pitted cells, with few
or no intercellular spaces (m*). Resting on this compact layer,
either singly or in groups, are larger cells, which stand out like
branches in the intercellular spaces of the next layer.

The cells of the second layer are characteristic of the genus (m?).
In this species the layer is two cells thick, with intercellular spaces 5°
large that they form a great cavity in which the cells are suspended.
In form the cells are somewhat stellate, joining the preceding layer
by _ branch or a similar but smaller cell. The walls are peautifully
teticulate, the thickenings following a definite course around the
curved surface of the cell. The cross walls are netted in a similar
manner but with a somewhat larger mesh. F

The true spongy parenchyma consists of @ number of layers ©
somewhat collapsed cells, but treatment with Javelle water expands
ne so that they may be easily studied. The Sey ost a

ed, decreasing in size toward the inner epidermis (p"; P*)-
surface view the contour of the branches appears, 11 the center of ~
Cells, as rings nearly as large as the cells themselves. A green color Is
Siven to these inner layers by chlorophyll. | | tied
e . The inner epidermis (p3) is composed of small, thin-w

PertsPeRM (figs. 5, 6, N).—In this coat there are about six layers

278 BOTANICAL GAZETTE [APRIL

of thin-walled cells with longitudinally elongated, cuticularized,
epidermal cells.

Ca
ee /

REM

SS be)
ySSTers OPS Yee
p3 Daten

SES Se
canes Sarsy

Sas

se S
Dap ame i
J

moderm consisting of

Pho —— i Wi Seed in cro seca, S, sper hy pitted subepidet-

ep, ri , starch grains,

mal io seen m, iad parenchyma, m2, reticulated spon Sx
chyma, hee nchyma, 2, spongy parenchym aoe p3, inner cide cay aleurone
sperm; £, sn consisting of aleurone Ae C, cotyledon containing

grains. ae

: : ells
Estocini (figs. 5, 6, E).—This consists of a single layer een
with thickened walls, containing granular protoplasm and
central nucleus.

i909] #BARBER—-FRUITS AND SEEDS OF CUCURBITACEAE 279

Empryo.—The cotyledons (C) have small epidermal cells on both
surfaces and two layers of palisade cells within the inner epidermis.
Aleurone grains (al) up to.6 #, containing globoids and crystalloids,
also oil, are found in all the cells.

C. PEPO VAR. VERRUCOSA NAUD.

This variety, the crook-necked squash, has a rather large flask-
shaped fruit, with the neck showing a distinct curve or crook. Longi-
tudinal grooves occur frequently, but are not so well marked as in C.
Pepo. There is a hard rind, yellow or orange in color, covered with
very pronounced warts and filled with bundle fibers. The central
parenchymatous tissue breaks down only in the swollen base, the
neck not expanding enough to tear the tissues apart.

The seeds, of a dirty white color, are not over 2°™ long, smooth,
and flattened, with a border at the edge.

: PeRICARP (fig. 7).—The pericarp of this variety differs very
little from that of C. Pepo, the chief difference being in the greater
hardness of the rind.

he Epicarp (epi).—The palisade layer is about 36 in height, the
thickened, prismatic cells having cuticularized outer and radial walls.
The outer wall is not uniformly thickened; instead it has a depression
over each cell, giving it a wavy contour in cross-section.

There are two forms of hairs, resembling those of C. Pepo in size
and shape; instead of falling off, however, they persist on the mature
fruit. Sunken stomata (sto) are present in considerable numbers.

; 2. Hypoderm (hy).—The cells of this layer resemble those of C.
¢po, the description given for that species applying to this.

3- Outer mesocar p (st).—This characteristic layer consists of many
layers of stone cells which differ considerably in size. The cells are
polygonal, small in the sharply defined outer layer, increasing in size
"ward until they are lost in the next cell layer. About spherical
Fae (x), occurring at the junction of this layer with the hypoderm,
Reg elongate radially to about twice their transverse diameter.
ee ies are really large intercellular spaces appearing 1n the
Young fruit and showing no evidence of secreting cells in any stage of
vic ba Before the walls thicken the cells bulge out into these

Vities, Among the smaller polygonal cells are also large spherical

280 BOTANICAL GAZETTE [APRIL

ones with cell cavity, frequently 150 » in diameter, about the size of the
intercellular cavities, both being conspicuous. The cell walls vary
from 8-14 » in thickness and have numerous pits.

: f : Z q jon; epi,

Fic. 7.—Cucurbita Pepo var. verrucosa Naud. Pericarp 10 o aoe

epicarp with ¢, hair, and sto, stoma; hy, hypoderm; s#, outer mena a

layer) with x, spherical cavity; mes, middle mesocarp with fv, bundle, ané am
60.

4. Middle mesocarp (mes)—The middle layers ‘ “ truncated

consist of rather thick-walled cells containing numerou
starch grains (am).

1999] BARBER—FRUITS AND SEEDS OF CUCURBITACEAE 281

5. Inner mesocarp.—The cells of the inner layers are the same
size as those of the middle mesocarp, but have thinner walls, larger
intercellular spaces, and no evident contents. The fibrovascular
bundles, sieve tubes, and latex tubes, found throughout the tissues,
are of the usual type.

6. Endocarp.—This layer corresponds with that of the species
already described.

SPERMODERM.—This differs from that of C. Pepo only in the first
and fourth layers, as follows:

1. Epidermis—The thickenings on the radial walls, besides
branching profusely at the outer end, send out occasional fine branches
the whole length of the rod.

4. Parenchyma.—The second layer of this parenchyma is four or
five cells thick, with somewhat smaller intercellular spaces than in C.

Pepo,

C. Pepo var. MELOPEPO L.

The popular name (scallop) for this variety suggests the shape of
the fruit. Itisa large depressed berry with ten to fifteen quite deeply
cut longitudinal grooves. Like C. Pepo it is firm, smooth, and yel-

With a rind and a central cavity containing fibers.

The seeds are elliptical, 1-1.5°™ in length, smooth, flattened,
tordered, and of a yellow-white color.

PERICARP.—The cell structure does not differ essentially from
that of . Pepo,

SPERMODERM.— This coat differs slightly from the corresponding
“at ofC. Pepo. The palisade layer, with thickenings like those of the
iC verrucosa, is thicker, reaching o.3™™ on the flattened surface

€ seed.

Phe 9, We which in the species previously described is found sot
oy = edge of the seed, here sends out a few ee ae ase
evide ne “ross-section the cut surfaces of these branches

€nt in the inner parenchyma.

C. PEPo VAR. OVIFERA NDN.

T : -
whi he fruit of this variety is egg-shaped, 8-10°™ long,

Ke color, with hard rind which dries up and persists in its original

of a yellow-

282 BOTANICAL GAZETTE [APRIL

form. The rind is 1°™ thick surrounding the central mass of fibers,
which do not break down. A cavity is formed, but it is not so evident
as in the other species of this genus, since the tissues, although separat-
ing, retain their original position.

The seeds are of the usual type for this genus.

PERICARP.—The layers are essentially like those of the variety
verrucosa, having a wavy-walled epicarp, thick-walled hypoderm,
outer mesocarp of stone cells with the characteristic cavities and large
spherical cells, middle mesocarp with starch, etc.

SPERMODERM.—The layers are similar to those already described.

C. PEPO VAR. ORANGINA SER.

The fruit of this variety differs from that of the type only in macro-
scopic appearance, having the size, shape, and color of the orange.
Rigidity is given the thin rind by the outer mesocarp of stone cells.

The small seeds are histologically like those of the preceding
variety.

CucURBITA MAXIMA DUCH.

This species (winter squash) is a native of southern Asia and, like
C. Pepo, has many varieties differing in size and shape. h

The fruit, the largest of the family, varies from 15 to gos in lengt-
It is rounded ovate, with warts and occasional longitudinal —
and is of a yellow, orange, or green color. In the so-called Zeus :
variety, the top of the fruit projects beyond an encircling line, OF od
striction, which marks the margin of the adherent receptacle.
hollow rind with central cavity is filled with bundles and seeds. an

The white seeds are 1.5-2.5°™ long, smooth, and —_
a border.

Both fruit and seed closely resemble in structure
varieties. nchy-

PeRIcarP.—The thin epicarp, small-celled hypoderm, sae f
matized outer, thin-walled middle, and inner mesocar p are sae
C. Pepo var. verrucosa (fig. 7). d that of

SPERMODERM.—The chief difference between this eco chyma,
the species already described is in the middle spongy paren
which in this seed is five to six cells thick.

cs Pepo and its

00) | BARBER—FRUITS AND SEEDS OF CUCURBITACEAE 283
Cucumis sATivus L.

Cucumis sativus L. (cucumber). occurs native in the East Indies,

and from this species have been derived many varieties which have
long been cultivated in gardens.
_ Thefruit varies in size, but is always elongated or oval, and rounded
triangular in cross-section. It is fleshy and solid, without a central
cavity, and of a yellow-white color with white fruit-flesh. Numer. us
warts cover the surface, each capped with a short
blunt spine which readily becomes detached dur-
ing growth or on handling.

Numerous white seeds. (fig. 8), borne within oo
‘the three locules, are 1-1.5°™ in length, flattened, aa ae
seldom over 2™™ thick, and are not, like those of — xx.

Cucurbita, provided with a border.

RICARP (figs. 9, 11).—This consists only of the solid fruit-flesh,
eae occurring throughout the mesocarp and not in a central

vity,

_ 1 Epicarp (epi).—The prismatic cells form a palisade layer 75 #
m width, with strongly thickened outer and radial walls, and very
thin inner ones. The intense color
of the fruit is due, not to the cell
contents, but to the yellow walls.

The warts, which. appeat before
fertilization of the flower, have the
same cell structure as the outer
layers of the pericarp. Each ‘beats
an emergence (jig. 11) consisting of
large cells with thickened, sparingly
pitted walls. They grow rapidly for
a short time, but lose their contents
soon after their walls begin to

zi (38 Im-
TP In cross-section, with
airs. X 160.—FIG. Io.
ature seed

9. Cucumis .
Ky
Mature epica $ Sativus

thicken.

At its apex the emergence bears
a long, jointed (up to ten cells),
conical hair, with thickened walls.
The cross walls and inner sunken
foot, also thickened, are pitted (i).

284 BOTANICAL GAZETTE [APRIL

Occasionally a second hair, similar in structure but of smaller size, is
developed at the side of the terminal one. The hairs usually disap-
pear in the carly stages of growth, but the emergence, unless rubbed
off, persists as a brown hyaline spine.

In addition to the hairs of the wart, numerous small capitate hairs,

rf , consisting of a four-
celled head and
stalk of three cells,
cover the immature
fruit, but disappear
early, leaving no
scars. Stomata are
not present.

2. Hypoderm
(hy) A aes
of layers of small,
rounded, loosely ar
ranged cells form
the subepidermal
tissues. In the

oy LN at maturity.
Tc t the base of
SW ites the emergence

oo ol pohagage sativus I. Pericarp in cross-
h ? PM, epicarp with emergence bearing ¢, hair; /y, hypod
ypoderm; st, sclerenchymatized cells at base of hair. X55- whic h becom e
This thicken”
vaporation, or
In

thickened, sclerenchymatized, and pitted (st).

the cell walls probably serves either to prevent ©

entrance of fungi, or both, after the removal of the emergenc® a

leguminous seeds there are sclerenchyma cells immediately belo

the hilum groove which serve this same purpose- ¢
3- Outer mesocarp—MoELLER’? finds a weakly developed ae
22 Mikroskopie der Nahrungs- und Genussmittel 473-

90] | BARBER—FRUITS AND SEEDS OF CUCURBITACEAE 285

céll layer. In the many fruits I have examined, however, the meso-
carp does not show any great differentiation. The outer layers consist
of rather small thin-walled cells with intercellular spaces.

4, The middle mesocarp differs from the outer only in the greater
size of the cells.

;. Inthe inner mesocarp the cells become smaller, resembling those
inthe outer mesocarp. The bundles, isolated sieve tubes, and latex
tubes, found throughout the mesocarp, are of the type already
described.

6. Endocarp—The elongated, thin-walled cells of this layer
remain with the mesocarp and are not attached to the seeds as in
Cucurbita.

SPERMODERM (figs. 10, 12-14) Seeds for study should be taken
— from the fruit, as in drying they usually lose their outer
Walls,

1. Epidermis (fig. 12, ep):—The prismatic cells are radially
dlongated to 160 # on the sides of the seed and 260 # at the edge. In
surface view they are polygonal, transversely elongated, and arranged
side by side in rows. The outer and inner walls are thickened, the
outer having in addition a cuticle, while the radial walls are very
thin, with the characteristic thickenings on only the side walls.
Bach thickening consists of a single straight rod, broadened at the
base and tapering rather abruptly into a blunt almost rounded point
8 within the cuticle. Von HOaNEL,?3 in the year 1876, described
ar as consisting of two different layers; PICKED, the aon

“ar, lound three; and Harz,?5 although quoting Von HOHNEL,
oo gia My own observations corroborate FICKEL'S state-

ae he three layers can be distinguished by the difference 1n
eg Power and by staining. The inner layer, which gives the
si . for lignin, shows the strongest refraction; this ah
ripe ae thin layer of cellulose (blue with iodin and ae
bin; er weak refractive power; and this in turn G tee
Oe ihe ve scarcely visible, second layer of cellulose ( # ah
es ssi aa _ Tangential sections show the arrangemen
4 e elliptical appearance of the cut rods.

23 L >
24 oe cit., footnote 3, Pp. 330. 25 Loc. cit., footnote 7, P- 773:

cit., footnote 2, p. 742.

286 BOTANICAL GAZETTE [APRIL

2. Subepidermal layer (sub and fig. 13).—On the flattened surface
of the seed this tissue consists of one layer of longitudinally elongated
cells, arranged end to end in rows, while at the edge they are some-
what shorter and form several layers. They are quite large, ranging
from 50-175 /# in length, and have thickened and sclerenchymatized
sinuous walls. The most striking feature of the layer, best seen in
surface view, is the intercellular spaces which occur in great numbers

; between the side and end walls.
They are very small, several
occurring in every turn of the
sinuous wall, which is greatly

=) Se — hare
76 Bee a
na eee SY
1 WetSeest aaet
ane JS
Vi

Fic. 13.—Cucumis sativus L. Iso
N lated subepidermal cell of spermoderm
in surface view. X300-

Fic. 12.—Cucumis sativus L. Seed
in cross-section. .S, spermoderm consist-
ing of ep, epidermis, sub, subepidermal
layer, scl, sclerenchyma, /?', stellate

parenchyma, p?, spongy parenchyma; os Oe
» pe > ie, endos Tm; coty- Fic. 14.—Cucumis ann ae
ledon consists of ep, epidermis and me- half of isolated cell of sclerenc yma ©)

sophyll with a/, aleurone grains. X 160. in surface view. *39°
thickened about the space. Between these spaces ont z
outer and inner walls are small pits.

3. The sclerenchyma (scl and fig. 14) consi
(more toward the edge) of longitudinally elongated
narrow cell cavities and very thick lignified walls.
potash brings to view the middle lamella and striations

sts of OM ye

(220 #) cells 4
Heating

of the wall, but

1909] BARBER—FRUITS AND SEEDS OF CUCURBITACEAE 287

only after maceration can the overlapping branches of the ramifica-
tions be seen. Pits are numerous.

4. Parenchyma.—The outer layer (p") is one cell thick at the side,
increasing in number toward the edge, and consists of small cells
more or less stellate in form. This layer is followed by two or three
cell layers of thin-walled parenchyma (7), which are usually rup-
tured in dry seeds. - :

5. The inner epidermis consists of small, elongated parenchyma
cells.

PerisperM (N).—A number of layers of collapsed, thin-walled
ells form this layer. The longitudinally elongated epidermal cells
have a cuticle.

Enposperm (E£).—The cells of this layer are polygonal, 22 # in
diameter, have thickened walls, and contain small proteid granules.

The Epryo (C) lacks distinctive features.

Cucumis MEto L.

All the varieties of musk melon are derived from a single species
indigenous to Africa and Asia. Harz divides the species into nine
groups, each with one to ten subspecies. gn ee

The fruit varies somewhat in size and shape, but is usually spherical,
oval, or occasionally elongated, and has eight to ten longitudinal
stooves, differing in depth in different specimens, to which is due the
characteristic “melon shape.” The surface is green, yellow, or red,
with a gray network of corklike tissue.

A tind 2.5 to 5°™ thick, with yellow to reddish fruit-flesh, incloses
4central cavity containing fibers, seeds, and a considerable amount of
Watery fluid.

The seeds are like those of C. sativus, except that their color is
Yellow,

PERICARP (figs. 15-17).—This consists of the hard rind, and the
soft, sweet watery fruit-flesh with bundles.

1. Epicarp (epi).—The cells of this layer vary im size, shape, and
t ickness of walls. On the ribs there is a palisade layer of greatly
mickened, pilygons] cella with a cutide. tn me eee =O
netease in size, the walls become thinner, and pits make their appear-
‘nce. In cross-section the cell cavity appears flask-shaped, the

288 BOTANICAL GAZETTE [APRIL

neck running out to a point. Fig. 15 shows the radiating cell cavity
in tangential section.

According to MOELLER,”? cork cells occur here and there below the
epicarp. This I find to be true of the young fruit, but at maturity
they break through the epicarp, forming the corky ridges or netted
thickenings previously mentioned. The cells are small, thin-walled,
and radially arranged, forming a dense mass of cork tissue, which,
together with the ruptured epicarp, closely resembles lenticels, but no
complementary cells are formed. This likeness is very marked in
cross-section (jig. 16).

Stomata (fig. 17, sto), with guard cells of the usual type, are present
in the grooves and, :
less frequently, in
the depressions be-

(~iK
tween the corky (eas =e
ridges. The wall jegeerssses
2 he ty + $7

8 aa PS SRD oni

of the accompany-

hy---* vem Bi
Nigeeccasss=

od La me

Fic. 15.—Cucumis Fic. 16.—Cucumis Melo Ll. Ribol paar

Melo L. Epicarp in tan- section; epi, epicarp; su, cork; jy, hypoderm; gr oy
gential section. Xx 160. mesocarp. X50.
ing cells have few or no pits and are irregularly thickened. In st
instances the whole wall is thickened, while in others large protuber-
ances are sent out into the cavity.

In the depressions and grooves are jointed (three or ‘- the
conical hairs up to 375 win length (¢). The diameter at the base 1S “a
same as that of the neighboring cells, but gradually diminishes to
apex. The thickened walls have additional minute local eso
or warts, visible only by the most careful focusing. About wes os
= the cells are distinguished by the smaller cavities and thie
walls,

The immature fruit bears small capitate hairs like those of .
Salivus,

more cells),

1909] BARBER—FRUITS AND SEEDS OF CUCURBITACEAE

289

2, Hypoderm (hy).—The cells of a number of layers beneath the
epicarp are thickened and pitted. Below the thick-walled epidermal

cells they are of
medium size
with intercellular
spaces; under
the thinner epi-
dermis of the
grooves, through
which they are
visible, they are
larger and thin-
ner-walled with
very large pits.
Chlorophyll is
present in greater

FM Fic. 17.—Cucumis Melo L. Pericarp in surface view;
epi, epicarp with /, hair, sto, stoma; /y, hy

poderm. X 160.

or less amount even though the fruit appears yellow.
34,5. The outer, middle, and inner mesocarp of this species con-

F Ae,
s @& Aine
Dick

“Cheah 8.—Cucumis Melo L. Seed in
&p, ae S, spermoderm consisting of
Ssclerenc 18, sub, subepidermal layer, scl,
Prec’ “ig p', sclerenchymatized spongy

tae Z gy parenchyma; JN,
ep, epidermis, endosperm; C, cotyledon with
aleurone grai, and mesophyll containing a/,

grains. X 160.

. sist of cells similar to those of

the corresponding layers of C.

sativus. The placentae, how

ever, instead of persisting 1n-

tact, are almost if not quite

n.
6. Endocarp—Asin C. sati-
vus, the cells are thin-walled
and longitudinally elongated.

Epidermis
face view the cells are polygonal
and in cross-section
a length of 200 #-
wall is thickened and the thin
ql walls have the character-
Harz” fig-
th rods

radi
istic thickenings. *™
ues eo

26 Loc. cit., footnote 7, P- 777°

290 BOTANICAL GAZETTE [APRIL

having small side branches at right angles, and MOELLER?? describes
them as simple rods without branches. Like the latter author,
I find single straight rods, one on each radial wall, which stain
yellow with iodin and sulfuric acid, thus showing them to contain
lignin.

2. The _subepierinal layer’ ae) is the most characteristic layer
of this seed. Whereas in C.
¥ | *“"\ °. — sativus it consists of but one
ep Ay = cell layer, in this species it has
a % five or six layers of cells lon-
gitudinally elongated (except
at the edges) and arranged end
to end in rows. Both species
have sinuous walls and char-
acteristic circular intercellular
spaces visible in cross- a5 We
as in ~longitudinal _ section.
The cells of the outer layer
are small, increasing inward in
size and thickness of wall with
every layer, until the: inner
one is distinguished from the
sclerencliyma — only Fick the
somewhat more’ irre ar-
rangement “and the slightly
smaller size of the cells.

We
|
a

Sy
eo
xe

G
AD
‘iy

: Ny Gi
ws
ONY
Gi

ax
5
=
oe

v
\)
5 rh
1 a
Oe rare
K Qu
O r

Fic. 19.—Cucumis Melo L. Isolated fig f C.
sclerenchyma cell of spermoderm. X 300. layer differs from an
sativus only in the irregular
contour of the outer surface, where the layer conforms to —
of the inner surface of the preceding layer. ne
4. Parenchyma (p* and p?).—Beneath the sclerenchyma are cA :
sometimes two, layers of small cells, which are thin-walled, page m
matized, and frequently pitted. This tissue gradually pase! ope
without inward to larger-celled parenchyma with bei: ree
spaces.

1909] BARBER—FRUITS AND SEEDS OF CUCURBITACEAE 2g1

s. Inner epidermis.—A layer of small parenchyma cells makes up
this layer.

PERISPERM (fig. 18, N)), ENDOSPERM (fig. 18, E), and EMBRYO
(fig. 18, C) have the same structure as the corresponding parts in C.
sativus.

CuCUMIS ERINACEUS L.

The seed of Cucumis erinaceus L. (fig. 20) is 6-8"™ long, 2-3"

broad, and 1-1.5™™ thick, flattened, pointed, smooth, and yellow.
SPERMODERM (fig. 21).—This is quite similar
] to that of C. Melo.

1. Epidermis (ep).—The prismatic cells, are
ee 20.—Cucumis radially elongated to 70#; increasing in length
maceus L. Seed. : : i
<. but slightly if at all over the edge. Each radial

wall has one rod broadened at the base and
bluntly pointed at the outer end. :

2. A subepidermal layer (sub), about six cells thick, consists of
small, tangentially elongated cells, increasing in size Over. the edge
of the seed. They are thick-walled, pitted, striated, and increase in
= from without inward. Occa- SYR
sional small intercellular spaces are °P "3G a
seen, especially at the edge.

“3 Sclerenchyma (scl). —This
“ingle-celled layer, 40 » in width,
differs from the subepidermal layer
Principally in the greater size of
the cells, which are longitudinally
elongated and have very thick
uous walls containing pits.
bap - arenchyma (p).—The outer
i a made up of very small cells me, : L
thin walls and small pits seen Seed in cross-section; Ss spermoderm
only with very high magnification. consisting of @P
epidermis, 54s

Tn the mi pb, subepid
€ middle layers the cells are 5), sclerenchyma_ layer, p, paren
jsperm; Z, endosperm;

somew :

Mewhat larger, also thin-walled, chyma; N) pert -
with intercellul C, cotyledon, consisting of ep, &P’

of th ea gata the angles mi and mesophyll with al, aleurone
aes rains. X 160

292 BOTANICAL GAZETTE [APRIL

5. The inner epidermis consists of one layer of very small paren-
chyma cells.

PERISPERM (jig. 21, N), ENDOSPERM (fig. 21, E), and EMBRYO
(fig. 21, C) present no new features.

CITRULLUS VULGARIS SCHRAD.

Citrullus vulgaris Schrad. (watermelon), now extensively culti-
vated in the tropics and warmer regions of the temperate zone, is
a native of South Africa. ENGLER and Pranti emphasize the fact
that on its native soil the fruit forms a considerable part of the food
of both the natives and the larger animals. This view is substantiated
by LIvINGSTONE in his Travels in Central Africa.

The fruit is spherical, or more often ellipsoidal, of a dark green
color, frequently mottled with white in ragged, longitudinal stripes
several centimeters in width. The rind is firm but not hard, green at
the outer surface, white further inward, chang-
ing gradually to the glistening pink, red, or yellow
inner fruit-flesh, which contains g1-95 per cent.
of water. ge

In the inner colored fruit-flesh are imbedded — * ew
the numerous white, brown, or black mottled me te
seeds (fig. 22). They are flat, without a border,
lustrous, and smooth except when mottled, in which case the
slightly rough.

PERICARP (figs. 23, 24).—The great bulk of the fruit is the solid
fruit-flesh, and it is this sweet, watery portion that is usually eaten

; and not the rind as in Cucurbita.
The placentae are beautifully out-
lined, the bundles being wie ng
what lighter color than the Hes”.

ipe th

y are

separate along th

no true cavity is formec fesh
individual cells of the Inn
: e:

: are visible to the naked ey wry
Fic. 23.—Citrullus vulgaris Schrad. Epicar p (epi).—Inst

Pericarp in surface view. Epicarp with — * Ils there is a

“to, stoma; hy, hypoderm. X 160. the true palisade ce

1909 BARBER—FRUITS AND SEEDS OF CUCURBITACEAE

293

layer, 33 » in thickness, of cells often broader than high. The outer

wall is very thick, the thickening

running down on the radial walls.

In surface view the. middle lamella is visible and the hypoderm is
quite evident through the thick-walled but transparent epicarp.

The ovary and very young fruit bear jointed hairs, which often
reach a length of 32™™ but are only 84 » broad. They are borne on
stalks of several joints, the foot becoming no larger than the surround-

ing epidermal cells in which
it is imbedded. All the walls
are only slightly thickened.
These hairs disappear soon
after fertilization, leaving scars
which are evident only on the
immature fruit.

Stomata (sto) are very nu-
merous, occurring singly or
m groups. They have rather
large guard cells and thin-
walled accompanying cells.

2, Hypoderm (hy).—A
layer, many cells thick, con-
taining chlorophyll, forms the
green tissue of the rind. The
outer layer consists of short
tells, resembling those of the
“picarp in size and shape; the
following layers are made up
of cells smaller in size and
Sodiametric. Numerous pits
are present, the walls appear-

bee dts: : :
§ distinctly beaded in surface view.

occur throughout the tissue.

‘ re Outer mesocar p (st).—In the very young fruit the
. et but after a few weeks single cells
icken their walls and become pierced with pits:

Fic. 24.—Citrullus vulgaris Schrad.
Pericarp in cross-section; epi, epicarP with
sto, stoma; hy, hypoderm;
(stone-cell layer); *;
groups of stone cells; mes
X 160.

Small ‘ntercellular spaces
cells are small

or groups of cells
These groups are

‘distributed about the fruit just within the hypoderm. As growth

of t . ; , .
he fruit continues, these cells increase nu

mber, the groups

204 BOTANICAL GAZETTE [APRIL

approaching each other until there is eventually, in the mature fruit,
a distinct zone of stone cells surrounding the inner tissues. This
stone-cell layer is not quite continuous, the groups being separated in
many places by a few cells which retain their thin cellulose walls and
afford easy communication between the hypoderm and middle
mesocarp (x). The inner contour of the layer is very irregular.

4. Middle mesocarp (mes).—The cells are thickened, pitted, and
gradually increase in size from the small outer layers inward.

5. Inner mesocarp.—This layer is the great central mass of pink
(or yellow) tissue. The cells are of enormous size, often 1.25™™ in
diameter, and, as previously stated, they can be easily distinguished
with the naked eye. The walls are thin and are separated at the
angles by intercellular spaces. A sweet, watery liquid fills the cavity.

Bundles, sieve tubes, and latex tubes are found scattered through-
out the mesocarp.

6. The endocarp consists of one layer of small, very thin-walled,
elongated cells as in C. sativus.

SPERMODERM (fig. 2 5).—This coat, consisting of the usual number
of layers, is thin but very firm.

» I. Epidermis (ep).—Prismatic cells form a palisade layer covered
with a thick cuticle, which occasionally reaches a thickness of 35
- The outer and inner walls have wavy contours, that of the inner wall
being much more pronounced. Each radial wall, which is thicker
than in the species previously described, has one thickening > the
form of a Straight rod pointed at the outer end. Occasionally It
branches once dichotomously, the two branches running straight
out to the cuticle. These rods are sclerenchymatized, responding 1
the test with iodin and sulfuric acid. In the colored seeds the ae
ete appearance is due to the colored contents of the epiderma
cells.

rs of

2. The subepidermal layer (sub) consists of a number of laye et

sclerenchymatized cells, increasing in number over the edges.
outer cells are small and isodiametric; those. of the middle ait
large and radially elongated; while the inner cells are ™ —
small. The cell walls increase in thickness from without oa aie.
those of the inner layer having scarcely any cell cavities. Allo
Walls are sinuous and deeply pitted.

1909]

BARBER—FRUITS AND SEEDS OF

CUCURBITACEAE — 295

3. Sclerenchyma (scl). —This layer differs somewhat from the

corresponding layer of the other

cucurbitaceous seeds. Instead of

conspicuously elongated cells arranged end to end in rows, they are

butslightly if at all elongated
and very irregularly distri-
buted, showing only the
faintest indications of an
end-to-end arrangement. In
other respects, that is as
regards thickness and sinu-
ousity of the pitted walls,
they are similar.

4. Parenchyma.— One
layer of small and some-
what spongy cells forms the
outer tissue (p*). They
are thin-walled, pitted, and
sclerenchymatized. Below
are several other layers of
more or less spongy paren-
chyma, the cells decreasing
i size toward the inner
epidermis (p?).

5: Inner e pider mis.—
This layer consists of small
thin-walled cells. :

PERISPERM (fig. 25,

‘)—An epidermis of lon-
situdinally elongated cells
with a cuticle covers a num-
ber of layers of thin-walled
parenchyma.

ENDOsPERM (fig. 25,

E).—This is the only spe-

cies ee .

esa in which this layer is more than 0

the mocystis lobaia there are occasional bro
thick-walled protein layer, but in the present

Fic. 25.—Citrullus pulgaris Schrad.
( oderm consisting of

in cross-section ; S, sperm

’
, epidermis, sub, sub idermal layer,
sclerenchyma, ? scle nchymatized —
chyma, #?, inner pa N, perisper™

y enchyma;
E, endosperm; C, cotyledon, with ep, epidermis,
and mesophyll, co
X 160.
ne cell thick. In
ken-down cells below
species the cell

296 BOTANICAL GAZETTE [APRIL

layers are numerous, forming a tissue as thick as the perisperm.
The outer polygonal cells are of the kind described for other species,
and the inner layers consist of extremely thin-walled parenchyma,
distinguishable only after treatment with Javelle water.

The Empryo (fig. 25, C) corresponds to that of the general descrip-
tion.

SICyOS ANGULATUS L.

Sicyos angulatus L. is a native of northeastern United States,
occurring as a weed in damp places, and is also occasionally cultivated
for arbors.

The fruits, each consisting of a small ovate pericarp (up to 2°" in
length), filled with a single seed, are borne in capitate clusters on a

SMOLIN =

Je 7x Q
ONAN a ea
SOON WSS
OA\\ ree nwest

RON
aes
ae

rss
ee

F ing chat
20.—Sicyos angulatus L. Base of prickle in cross-section, showing

Fic.
acteristic small cells. X 160
long peduncle. They are dry and covered with deciduous barbed
prickles, 8-ro™™ long, which give them a burrlike appearance.

The brown seeds are 12™™ in length and 10™™ broad, flattene?,
ovate, smooth, and lustrous.

PERICARP (figs. 26-29).—This coat forms only a very th
ing for the seeds and does not make up the bulk of the fruit,

in cover

199} BARBER—FRUITS AND SEEDS OF CUCURBITACEAE 297

other species described. The surface is roughened and covered with
spines and hairs, of which, as described below, there are four forms.

1. Epicarp—This layer is composed of small flattened cells 13 #
high, with slightly thickened walls. In surface view they are polyg-
onal and 20 # in diameter (fig. 29).

Very striking are the prickles (emergences), each being a long
($10™™") outgrowth borne on a swelling of the pericarp (fig. 26).
They have a constricted base, the epidermis of which consists of thin-
walled cells. Above this base the epidermal cells elongate and become
thickened. Over all the surface of the prickles are borne slightly
curved, single-celled, conical hairs (often 135 # in length), with
extremely thick walls (fig. 27). They are very stiff and turn sharply
backward. Doubtless they play an important part in the dispersal
oithe seeds, acting as barbs to fasten securely the prickles, and thereby
the fruit, to passing animals. Similar pointed but not deflexed hairs
occur on the epicarp, either singly or in pairs.

At the end of the prickle there is a thin-walled capitate hair, con-
sisting of a stalk of several short cells and a single-celled glandular
head (fig. 27).

Among the prickles there are also large glandular hairs, reaching
a length twice that of the emergence (jigs. 28, 29, i), The foot con-
‘sts of one or two enlarged epidermal cells and is surrounded by
ree but slightly smaller. From the foot the cells are elongated and
oe aie the hair tapering gradually and ending with short glandu-
8 cells. ‘The prickles become entangled in the hairs and do not fall

even though detached from the pericarp.

a form of hair (fig. 29, t?) on the pericarp is thin-walled,
Th ular = several-jointed, reaching a maximum length of 549 B-

— cell is conical and not glandular as in the preceding hair.

“umerous stomata occur among these hairs. :
j_oboderm—One or more layers of very small iseciamestic
ie, an inconspicuous layer just within the epicatp- cht
Sines tmal cells of the constricted base of the prickle are also very

we and thin-walled. These together with the thin-walled ep!-
'S permit the prickle to be readily detached at maturity. In the

ri ‘
Prickle the cells are elongated and sclerenchymatized, with numerous
Pits (fig. 26),

298 BOTANICAL GAZETTE [APRit.

3. Outer mesocarp.—Several rows of small rounded cells with
intercellular spaces make up this tissue. In the vicinity of the
prickles and hairs they are sclerenchymatized and tangentially elon-
gated.

4. The middle mesocarp consists of several layers of large-celled,
loosely arranged parenchyma with intercellular spaces.

5. Inner mesocarp.—The cells of this coat, although somewhat
smaller than those of the middle mesocarp, form several layers of
similar tissue.

29
Fic. 27. Sicyos angulatus L. Apex of spine. X 160.—Fic. 28. Sat a
- Apex of long jointed hair. X160.—FiG. 29. Sicyos angulatus L. cena
surface view; ¢, base of glandular hair; #2, short pointed conical hair; 5, eee
conical hair. X 160

The bundles and latex tubes, few in number, are scattered throug?
the mesocarp. .
6. Endocarp.—as in Cucurbita Pepo, the single layet of longitudi
nally elongated, thin-walled cells remains on the seed. pee
SPERMODERM (fig. 30).—1. Epidermis (ep). Prismatic a
a palisade layer only som in width. In cross-section no thicke
are apparent on the radial walls, but in surface view, what ap ing
in the cross-section to be walls thickened at the angles are rods runn! is
one on each wall, from the inner to the outer wall which they 5°

1909]

BARBER—FRUITS AND SEEDS OF CUCURBITACEAE

299

io join. No cuticle is present; the thickening, however, is continued

on the inner wall.

2. Subepidermal layer (sub).—
Small thin-walled cells, 13 # in dia-
meter, with intercellular spaces,
make up one layer. Additional
cells of the same type are present
below the radial walls of the
epidermis, making the tissue two
cells thick in those places.

3. Sclerenchyma (scl).—As in
Echinocystis, there is one layer
of radially elongated cells form-
ing a palisade layer, 230 in
width, which is best seen in cross-
section. The cell cavities are
hatrow, constricted about 70
from the outer wall, from which
constriction branches radiate
toward the outer wall. This is
Tepeated at the other end, the
‘onstriction being slightly nearer
the inner wall. In surface view
the cells are sinuous and indefi-
hitely arranged.

4. Parenchyma.—The outer
cells are small and thin-walled
(?*) and the several middle layers
“onsist of large stellate cells and
'ntercellular spaces (p?). All of
te layers are sclerenchyma-
—. About ten layers of
Pad parenchyma, the cells
ages In size inward, form the
—. (p3). These cells
i. chlorophyll, which gives

sue a distinct green color.

°? 49.8, | ak
: __ sieves angulatus L. ee
IG. 30- ph i oderm consist-

enchyma; N, pe
c. don Ww :
mesophyll containing 4, aleurone grains.
1005

300 BOTANICAL GAZETTE [APRIL

5. The inner epidermis with small, thin-walled cells resembles that
of Cucurbita Pepo.

PERISPERM (jig. 30, N’).—Five or six eke of ee with
outer epidermis and cuticle, have the same str
ing layer of otherspecies. The most charattaea festa | is the
transversely elongated epidermal cells.

ENposPERM (fig. 30, E).—This consists of one layer of thick-
walled cells containing protein granules.

Emprvo (fig. 30, C).—No characteristic features are evident.

EcHINOCYSTIS LOBATA Torr. & GR.
nis plant, Echinocystis lobata Torr. & Gr. (Sicyos lobatus Michx.,
Momordica echinata Muhl.), is one of the two Cucurbitaceae indige-
nous to New England. It has no value as a food, but is quite exten-
sively cultivated for arbors.

31 33

G. 32 Echinoess*

31. Echinocystis lobata Torr. & Gr. Fruit. X1 tis lobata Torr. &
XI.

Fic.
lobata Torr. & Gr. Fruit in cross-section. 1.—FIG. 33- pte
Seed. X1.—Fic. 34. —-Echinocystis lobata Torr. & Gr. Seed.

The oval fruit (jig. 37) is 5°™ long, light green in color, and pi
with soft spines 12™™ in length. At maturity it bursts gape
at the top, showing the fruit-flesh separated into an outer af oe
tissue consisting of a fibrous network. This inner part form"
large oblong cells, ag divided at the base into two ic

1909} 1 BARBER—FRUITS AND SEEDS OF CUCURBITACEAE 301

locules, as shown in fig. 32. Soon after dehiscence the pod dries
up.
The flattened ovate seeds are 20™™ in length, 8"™ broad, and 4"™
thick. The color is gray brown with beautiful brown. markings
(jigs. 33, 34) varying greatly on different specimens. On the flattened
surface of some seeds these markings are in the form of circles, on

ai0ey
AYR

- 36
Fic. 35. Echinocystis lobata Torr. & Gr. Apex of emergence from immature

hints a view; ep, epidermis; f, fibers of hypoderm. x 160.—Fi6. 38. Echinocytis
ort. & Gr. Base of emergence from immature pericarp. X 160.

e edge they generally
ged plotches. The
he greater thickness

sia as various shaped spots, but about th
emb; i border of elongated diagonally arran
tyo differs from the typical embryo only int

of the cotyledons.
PERICARP ( figs. 35-39).—The outer part, OF prickly rind, consists of

302 BOTANICAL GAZETTE [APRIL

the epicarp and outer mesocarp, the inner of the hard, stiff bundles
with only vestiges of ground tissue.

1. Epicarp (figs. 37, 39).—The cells of this layer are slightly
thickened, in surface view appearing polygonal and distinctly
beaded. On the spines the epidermal cells are elongated and have
thick walls.

The tip of each spine (jigs. 35, 36) curves decidedly, bearing at
the very extremity, or more frequently just below it, a long, jointed,
capitate hair. There is great variation among these hairs. They
- point downward, upward, or in an intermediate direction; the joints
may be few or many, the length of the hair varying accordingly.
Each cell is about as long as broad, with moderately thickened outer
and slightly thinner cross walls. The head consists of one cell
(fig. 35), two ( fig. 38), arranged side
by side, or three (jig. 36); arranged
about a common axis. They have
free rounded ends and are elongated,
with longitudinal diameter twice that
of the transverse.
hy Similar hairs, but much shorter,
: with one- to three-celled stalks, -
Ss 7 an aon. present on the sides of the spine

Fic. 39.—Echinocystis lobata Torr. ( jig: 38). Allof the hairs frequently
&Gr. Pericarp in surface view; epicarp disappear before the fruit reaches
with hy, hypoderm. X 160.

maturity.

Interspersed among the spines are the stomata with small
accompanying cells.

2. Hypoderm (fig. 39, hy).—Several layers of pitted cells gee
this layer. They are polygonal, thick-walled, and. DAY® EF ery
intercellular spaces at the angles, about which the walls = ib
thick. In the spines the walls are also thickened and pitted, ee 3
addition very much elongated longitudinally. The ype et
through the transparent epicarp, the circular intercellular pane ee
strongly thickened surrounding walls, being especially aoe

3- Outer mesocarp.—Several layers of jsodiametric paren¢
cells form the tissue below the hypoderm. The cells are sm?
thin-walled.

pitted

Fa we wheter et, PCW hd ed eee a

1909] | BARBER—FRUITS AND SEEDS OF CUCURBITACEAE — 393

4. The middle mesocarp cells are large, thin-walled, and mostly
broken down in the mature fruit, thus forming the cavity between the

outer rind and inner fibrous
tissue,

5. Inner mesocarp.—
Groups of small parenchyma
cells, with structure similar to
that of the middle mesocarp,
are found between the bundles.

The numerous large bundles
make up the bulk of the ma-
ture mesocarp, the anasto-
mosing branches forming the
fibrous network. Into each
of the spines passes a small
bundle which connects with
this netted system.

Latex tubes are present in
the young fruit, but very few
or none are found with the
-. bundles of the mature
pericarp.

6. The endocarp (fig. 41)
persists on the seed as a single
layer of very thin-walled cells,
longitudinally elongated and
arranged side by side in groups.

SPERMODERM (figs. 40, 42,
43).—This thick, hard coat is
best Studied after bleaching
with Javelle water.

1. Epidermis (ep).—The
cells differ markedly from those
of the seeds previously de-
scribed in that th
‘ ey vary greatly
height throughout, owing to
regularities, not in the sur-

ep- —=_ =) ama
oO ~8 } 4 at
PYROS Yor
OK) ») OS

eV OAIC

Wee I
JO) @ ‘e)
A\ ‘sf
SOORL tar
SOary Ty!)

TN)

quae:
oT SORES

__Echinocystis lobata Torr. &
Fic. 40.—Echinocy tm

._ Seed in cross-section; > spern ,
consisting of ef, epidermis, sub, ps
layer, sel, sclerenchyma, pi, sma ‘

renchyma; N, peti-
s ndosperm; ©; coty edon with .
epidermis, and mesophyll containing 4,
aleurone grains. x 160.

304 BOTANICAL GAZETTE [APRIL
face of the seed, but in the subepidermal layer. A cross-section
best illustrates this characteristic feature. The cells are much
shorter than broad, except where they suddenly elongate, running

down into the grooves of the subepidermal layer to form palisade
In surface view the whole radial wall

cells about 54 in height.
is thickened, without the thickened rods so characteristic of other

seeds.
2. Subepidermal layer (sub).—This coat, which varies in thickness,
as above described, always consists of numerous (ten to fifteen)

\ )

hil

<
SSves)), |

‘Mee

Y pra
WiC

—
Ss i,

IAN

bate
ees
YY
=
LS

\)
:
Ss
Sa
SNES
3

er
y
i

j S

fa ee
\; | | weeeed,< \
NEC poe
Lewy

A if t)

1G. 41. Echinocystis lobata Torr. & Gr. Endocarp in surface :
. 7 tan:
rr. & Gr. Subepidermal layer in mee in sur-

F
Fic. 42. Echinocystis lobata ‘To
X300.—Fic. 43. Echinocystis lobata Torr. & Gr. Parenchyma W!

view. be ge
jal section.

face view. X80.
layers of sclerenchymatized cells. They are very loosely —
with large circular spaces surrounded by thickened walls. In cros
mass of cel 1 ™
distinct.

alls is Very

section the tissue appears only as a confused
e outer cell layer

tangential section (fig. 42) the contour of the w
e seed owes its color to the dark contents of th ll layer

3. Sclerenchyma (scl).—In cross-section this single a being
appears as a palisade tissue, the cells, while only 4°/ ee sd the
radially elongated to 1804. The walls are extremely thick a

1909] BARBER—FRUI TS AND SEEDS OF CUCURBITACEAE 305

cavities narrow, each sending out branches to the layers above and
below. The folds of the sinuous outer and inner walls are also
conspicuous in cross-section. In surface view the cells have sinuous
walls and are irregularly arranged.

4. Parenchyma.—The outer layer (p’) consists of small, thin-
walled, and sclerenchymatized cells without pits. It bears no resem-
blance to the corresponding layers of Cucurbita. Adjoining this layer
are two or three layers of true spongy parenchyma with large cells,
thin walls, and large intercellular spaces (p?). From this tissue the
cells decrease in size inward for about fifteen layers (p3).

The branching and anastomosing raphe ramifies through this
inner parenchyma. Although each branch contains but few bundle
elements, the branches are so numerous that they form a conspicuous
tissue, best seen in surface view (fig. 43)-

+ The inner epidermis corresponds in structure to that of Cucur-
bila Pepo, consisting of a layer of small cells with thin walls, the cir-
cular contour of the radial walls being very noticeable.

PerisPerM (fig. 40, N).—The structure of this tissue is the same
as that of the seeds already described, consisting of several layers of
thin-walled parenchyma and an outer epidermis with cuticularized
outer and radial walls. The epidermal cells are elongated, in many
places transversely.

ENDOSPERM (fig. 40, E).—One layer of uniformly thickened cells,
Pontaining protein granules, forms this coat. In one or two seeds
‘xamined there were indications of broken-down parenchyma.

The empryo (fig. go, C) corresponds to the general description,
= cotyledons having minute epidermal cells and mesophyll filled
with oil and protein granules.

LUFFA CYLINDRICA ROEM.

hy smooth, flattened seed (fig. 44) is 12"™ long, 6-7

~3™" thick, of a dark-brown or black color.

SPERMODERM (figs. 45-47).—This 1s thick and consists of the

tollowing layers:

* epatage (ep).—The palisade layer varies from

the mg the outer surface having a sinuous contour.
Tadial walls is a single rod (fig. 45) which, instead ©

mm broad,

30-60 in
On each of
f ending free

306 BOTANICAL GAZETTE [APRIL

at the outer end, joins the thickened outer wall. The contents are of
a dark-brown color.

2. The subepidermal layer (sub) consists of two quite distinct
tissues. Within the epidermis "a
is a layer, one to five cells
thick, consisting of small reti-
culated or pitted cells.
tangential section (fig. 46) the
zig-zag walls are seen to pro-
ject into the cavity, with an
intercellular space in the angle
of each projection.

The second tissue consists
of one layer of cells, 20 » in
thickness, polygonal and regu-
larly arranged in cross-section,
and slightly elongated tangen-
tially.

3. A sclerenchyma layer
(scl) of cells radially elon-
gated up to 235 u forms this

ndrica Roem

| walls of epidermis.
I rmoderm, show-
ing radial walls of subepidermal layer. X 300.—Fic. 47. Luffa cylindric Roem
in Cross-section; S, spermoderm consisting of ep, epidermis, Su
sel, Palisade sclerehncyma, p, parenchyma; N, perisperm; , en ti
with ep, epidermis, and mesophyll containing a/, aleurone grains: X 160.

. * ; yous,
characteristic layer. The outer and inner walls are fh sec

. a : . ‘oOss-
sending out peculiar branches. The cavity, best seem 12 @

Oe Bit, ite See eee Eee ae a fer epee RN

1909] BARBER—FRUITS AND SEEDS OF CUCURBITACEAE 307

tion, is extremely narrow, with a, globular enlargement just without
the center, the two ends branching and running into pits at the outer
and inner tangential walls. In tangential section the polygonal cells
are 30-45 # in diameter.

4. Parenchyma (p).—A number of layers of large stellate cells form
the spongy parenchyma. All the walls are thin, the cross walls being
pierced with pits. In the outer layers the walls are sclerenchymatized.

s. Inner epidermis.—This layer of small parenchyma cells is also
somewhat stellate.

The PERISPERM (fig. 47, N)), ENDOSPERM (fig. 47; E), and EMBRYO
(fig. 47, C) are of the usual type.

ABOBRA VIRIDIFLORA COGN.

The seed of Abobra viridiflora Cogn. (fig. 48) is 7-8"™ ene, AY
slightly flattened, the enlarged apex being 2-37" broad and 1. 5-27"
thick, of a deep-brown color with ———
longitudinal dashes of a darker
rown.

SPERMODERM (fig. 49).— Thin

ut firm. a ¥ a §

1. Epidermis (ep).—The _ pali- a , ne oe (8
sade epidermis varies in thickness “f \ ces

from 16-45 #. On the thick radial
walls of the larger cells are one or
more straight rods joining the outer sel--1 :
wall, which is extremely thick. P

2. Subepidermal layer (sub).—
This broad layer, six to
ten cells thick, consists of
‘ety thick-walled, pitted p
cells irregularly arranged.
oS and frequently
: jMner, cells are small, on; S, §

ut the middle ones are of ep, epidermis, sub, subepidermal layer,
enormous Be Few orno ™cbyma, & parenchyma; N, perisperm, E,
interc endosperm; C, cotyledon with ep, epiderm!s,

ellular Ss
ac a
present, a ae mesophyll containing al, aleurone grains

160

ep--- SOM age ne

‘me

Fic. 48. Abobra viridiflora Cogn. Seed.
%1.—Fic. 49. Abobra viridiflora Cogn. Seed
derm consisting of
scl, scler-

308 BOTANICAL GAZETTE [APRIL

3. Sclerenchyma (scl).—Small, thick-walled, pitted, irregularly
arranged cells make up this layer, which has no sharp line of demarka-
tion from the preceding layer.

4. The parenchyma (p) consists of several layers of small thin-
walled cells.

5. Inner epidermis.—One layer of parenchyma cells makes up
this indistinct inner tissue.

The PERISPERM (fig. 49, N) of several cell layers, ENDOSPERM
(fig. 49, E) of one layer of thick-walled cells, and EMBRYO (fig. 49,0)
are like those described in the introduction.

MELOTHRIA SCABRA COGN.

0 The yellow-white seed (jig. 50) is 4-57 long,
—Melo- 9mm broad, and 0.75™™ thick, flattened, pointed,

50
thria scabra Cogn.
-

and smooth.

SPERMODERM (fig. 51)-—Com-
sists of five layers, and has a
microscopic structure resembling
that of Cucumis sativus.

1. Epidermis (ep) —A palis-
ade layer, 270 » in width, forms
the outer layer, which, a5 ™
Cucumis, is frequently absent
on the dry seeds. The walls are
thin except for a single, straight

a AV pointed rod on each radial me
sub Sear Ae 2, Subepidermal layer (stl)
601 —— SON Dy This single layer is compo e
Se - thick-walled cells having pits @
distinct laminations. At the ed
of the seed the layer —
from two to five cells in thickness,

gn. the outer ones being elongat
Seed in cross-section; y spermoderm as dially.
consisting of ep, epidermis, sub, subepi- J) Small
dermal layer, sci, sclerenchyma, p, paren- 3. Sclerenchyma ss ne
chyma; N, perisperm; E,endosperm; C, pitted cells, longitudinally €

cotyledon with ep epidermis, and meso- ird layer:
? > r
phyll containing al, aleurone grains. X 160. ated, make up the thi

1909] BARBER—FRUITS AND SEEDS OF CUCURBITACEAE 309

4. Parenchyma (p).—Several layers of thin-walled parenchyma
without characteristic features form the inner tissue of the spermo-
dem. Only with high magnification can the pits in the outer layer
be seen.

5. Inner epidermis.—As in the other species, this tissue consists
ofa single layer of thin-walled
cells.

The PERISPERM (fig. 51, N), ss
ENDOSPERM (fig. 51, E), and
EMBRYO (fig. 51, C) correspond
with the general description. sub----@

TRICHOSANTHES COLUBRINA L.

The seeds are 15™™ long,
$9™" broad, and 4-5™™ thick,
of a gray-brown color. The
megular contour and beautiful
dark markings on the flattened
surface are illustrated by fig. 52. ee eeoes ak

SPERMODERM ( UR | cee A aes ay, ‘area
very ha . : anes a ees

ty hard and thick seed coat with - ; Bites
‘wo distinct subepidermal layers
is characteristic of this seed.

: 1. Epidermis (ep) —This layer
irregular in width (12-135
: - The larger cells, running
own into grooves of
the second layer, bear
Straight rods on their
radial walls. These
Tods join the thick-
ned outer wall and in
cr ti vag Fic. 52-
eae lon are indis- _ Fis., 53- utes brina
: inguishable from the ection 5: omer
Walls at h subt, outer and sub?, inner subep! noo
Al the angles. éclerenchyms, p, parenc ae . ee epi-
of the oe ee 6 oe ea . oi grains.
1 containing
Seiwih s So

¥ ae
zak: wags \ asset
TY
ae arises
ae use
a

= pee ete Se Gis

310 BOTANICAL GAZETTE [APRIL

brown coloring matter and starch is present in considerable
quantity.

2. Subepidermal layer.—As previously stated, this layer is divided
into two distinct tissues: the outer (swb*) consists of cell layers increas-
ing in size from the small outer cells inward, with numerous pits in the
thickened walls; below this tissue is a second (sub?), many cells
thick, consisting of very thick-walled cells with small branching
cavities and small intercellular spaces at the angles. The walls of
both tissues are brown.

3. The sclerenchyma (scl), consisting of large, longitudinally
elongated cells of the general type, forms a third sclerenchyma tissue
which is impregnated with a brown substance.

4. Parenchyma (p).—Medium-sized thin-walled cells, the outer
layer somewhat stellate, make up this layer. Chlorophyll is present
in the inner layers in considerable quantity.

The raphe (R) branches out over the sides of the seed, the small
bundles being found throughout the inner parenchyma. ;

5. An inner epidermis of thin-walled parenchyma cells completes
the spermoderm.

The cells of the PERISPERM (fig. 53, N’), ENDOSPERM (fig. 53; 2),
and EmBryo (fig. 53, C) are not characteristic in this species.

YALE UNIVERSITY

THE ANATOMY OF ISOETES
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY 126
ALMA G. STOKEY
(WITH PLATES XIX—XXI)

There is no group of plants among pteridophytes whose anatomy
has occasioned so much discussion as the genus Isoetes. The most
recent writers, FARMER on J. lacustris, R. Witson SMITH On i
echinospora, and Scott and Hit on J. Hystrix, disagree in many
points. In view of the lack of harmony, both of observation and
interpretation, it has seemed advisable to make a comparative investi-
gation of the anatomy of several American species.

Historical

The literature on the anatomy of Isoetes began in 1840 with a
paper by von Mount (24). Although he recognized the lycopod
aflinities of Isoetes, he noted several important points of difference,
inthe arrangement and structure of the roots, and in the nature of the
cambium products. Ever since that day the question of the nature
of the cambium activity has afforded a fertile field for discussion, and
tis with that subject that this paper is chiefly concerned. VON
Mout regarded the whole of the secondary growth as parenchymatous,
and states that, as in other vascular cryptogams, there is no increase
in thickness of the wood. The next significant work was that of
HorwersreR (16) in 1857. In discussing the cambium products ne
“ays (p. 36r): “The effect of the yearly renovation of the cambial
wi €f is not only to increase and renew the cortical tissue, but new
‘pital cells also become added, although only sparingly, to the wood
sh vigorous plants. Individual cells of the cambium; ee
aes or three cambium cells from the older principal ~ a t i
na a exhibit thickenings of the walls, which seg oni 6 a0
ee, of color betray their undoubted recent orig! “ Te “
Krypto Russow’s Vergleichende Untersuchungen der Let :
ihe gamen (18), with a discussion of the anatomy of J — iy a
ae Spora, and I. Hystrix. This paper has had great influeace

€quent work, and to Russow’s work may be attributed the cur
atr] [Botanical Gazette, vol. 47

312 BOTANICAL GAZETTE : [APRIL

rent conceptions of the nature of the cambium products, i. ¢., that the
cambium produces cortex externally to itself, but internally it pro-
duces a tissue which he calls the “ prismatic layer,’’ and this he regards
as being made up of phloem cells, tracheids, and parenchyma.
HEGELMAIER (12), writing in 1874, is inclined to question Russow’s
interpretation, but adds that no decisive grounds can be given against
the interpretation of the Davuerzellen as soft bast. FARMER (9)
published in 1890 a very full discussion of J. lacustris. He seems
reluctant to accept Russow’s theory of the nature of the cambium
products, but does not offer another. After quoting Russow he
adds: “Quite apart from the fact that it is produced internally to
the cambium and would, from the point of view of its position, be
anomalous, its structure is remarkably complex and heterogeneous.”
In discussing the structure of the “prismatic layer” he says:

The zone-like arrangement consists in alternations of tubular thin-walled
cell-rows of varying thickness, whose cell contents are clear and watery, with
others, whose cells are wider in the radial direction, and filled densely with starch.
Occupying a middle position in the latter zone is embedded an irregular =
of cells whose walls are thickened like those of the tracheids, but these to, unlike
the latter, often contain protoplasm and starch.

In 1900 R. Witson Suir (22) discussed the morphology of I.
echinospora, with some incidental work on its anatomy. His obset-
vations do not agree with those of FARMER on J, lacustris, and he
proposes “to drop the term phloem until its justification is established
on physiological grounds.” In the same year there appeared a very
exhaustive paper on J. Hystrix by Scott and Hitt (19). Tee.
of the cambium products was taken up, and Russow’s concepo® ie
indorsed, in that they regard the secondary tissues internal to te
cambium as consisting of secondary tracheids, parenchyma; .
phloem. Their statement is as follows:
of the xylem
phloem, 4
ry cortical

The cambium in I. Hystrix, arising in the tissues just outside
cylinder, continues its activity indefinitely, producing parenchyma,
a variable amount of secondary xylem on its inner side, and seconda
parenchyma only, towards the exterior.

Each of the last three writers has based his conclusi
of a single species, while Russow, who investigated three 3
hampered by the fact that he worked on herbarium material.

on on the study
forms, W4°

SRC eee eee wie ee eT reer ney ee

1909] * STOKEY—ANATOMY OF ISOETES 313

desirability of a comparative study of several species embracing a
considerable range of habit is obvious.

Material

Lam indebted for material of I. Tuckermani var. H arveyi (A. A.
Eaton) Clute (I. Harveyi A. A. Eaton) to Dr. Leroy H. Harvey,
who made a collection at Pushaw Pond, Maine, of plants in various
stages cf development; to Mr. WILHELM SuxsporF for material of
I. Nuttallii Engelm. (I. Suksdorfii Baker) collected at Bingen,
Wash.; and to Dr. H. C. Cow.es for material of J. echinospora var.
Fleitii which he collected at Spanaway Lake, Wash., in the summer
of 1907. The latter material has been kept in cultivation at the
greenhouses of the University of Chicago and has afforded abundant
material for the study of a close series of stages in the development of
the stem. 1 am also indebted to the late Mr. A. A. EATON of the
Ames Botanical Laboratory, North Easton, Mass., for the determina-
tion of I. melanopoda Gay.

For killing and fixing the stems a medium solution of chrom-acetic
acid was used. The material was embedded in paraffin and cut in
serial sections 7 to 12 thick. Some of the series were transverse
and others longitudinal, both parallel and perpendicular to the fur-
ow. A considerable number of stains and combinations of stains
es used, of which the most satisfactory were the following: 2 com-
bination of safranin and anilin blue devised by Dr. Lanp; iodin
green and Bismarck brown; iodin green and eosin.
I — four species studied present a considerable range in habit.

: Tuckermani var. Harveyi is a submersed form which grows 1D
Naler 30-150°" deep. The trunk is deeply two-lobed, occasionally
three-lobed. The leaves are short, strongly recurved, without
‘tomata, and said by Eaton (8) to be relatively the stoutest of any
‘orth American species, being 5-6°™ in length and 23°. 8
ee Isoetes echinospora var. F lettii grows On the gravelly
— of mountain lakes in Washington. In the spring it may be

mersed, but during most of the year it is out of the water. The
a water so that the substratum is never Very dry. .
fo * two-lobed and the leaves are moderately stout. The larges

Tm studied is J. melanopoda. It hasa large two-lobed trunk, with

314 BOTANICAL GAZETTE : {APRIL

very long, moderately stout leaves. It grows in inundated fields
and shallow ponds, where it is emersed during most of the summer.
I, Nuttallii has a three-lobed trunk and long, very slender leaves. It
never grows under water, but near springs or springy places, and on
low wet grounds or meadows, where the ground becomes very dry in
the latter part of the season.

Investigation

The complicated structure and arrangement of the old stem is more
easily understood by a study of the sporelings. The stem of a spore
ling is a mass of undifferentiated parenchyma, traversed by leaf
traces, which come together to form the short flat stem plate and
continue outward and downward into the roots. All observers
agree that in the young plants there is no cauline portion in the stem
bundle, but whether or not there is a cauline portion in the older
stems has long been a disputed point. HOFMEISTER, DF Bary,
CAMPBELL, and Farmer look upon the stem bundle as being made
up of leaf traces; while HEGELMAIER, BRUCHMANN, and SCOTT 4
Hit maintain that there is a cauline portion. In the short and com-
pact stem there are no indications of a procambial strand, and as
there are never any tracheids present above the last leaf trace, the
evidences of a cauline portion are not satisfactory. In following the
cross-sections of leaf traces in a series of longitudinal sections UP to
the place where the traces coalesce, it is easily seen that there 8 *
sufficient amount of tracheary tissue in the leaf traces to account for
all the xylem in the stem bundle. Scott raised the objection that
the tracheids of the stem are unlike those of the leaves, but 26 ie
greater difference between the tracheids of the stem and of the base ©
the leaf, than between those of the base of the leaf and of the upP®
together with oe
ences in the distribution of the thickening of the walls is —
ds are foune:
regions, al
this becomes more marked as the plant enlarges. This suggests that
the tuberous body is not wholly stem, but a contract
root. Owing to the compact growth and the consequen
of tissues, some of the root bundles in an old plant may be

SESS Oa ee tere eel megs Pre tg tee Soest ot

1909] STOKEY—ANATOMY OF ISOETES 315

up in the cortex than the old leaf traces. In fact it is not uncommon,
in sections of old stems cut at the very apex of the stem, to find in the
cortex a longitudinal section of a root making its way out. Hence,
although the stem and root regions are sharply marked off in the
central axis, it is impossible to delimit these regions in the cortex.
Inthe young plant shown in fig. 4, all the leaf traces and root bundles
are functional. In the plants shown in jigs. 19, 20, all the roots are
dead except a few at the bottom, and all the leaf traces except a
very few at the top. As the plant increases in size, the old leaf traces
are unable to keep up with the growth of the stem and are torn apart,
leaving the old stumps attached to the central axis, while the rest of
the leaf trace is carried out farther and farther by the growth of the
cortex, and in time is sloughed off.

The arrangement of the roots is described in great detail by SCOTT
and Hitt for J, Hystrix, a three-lobed form. The two-lobed trunks
show no essential variation from that type, and it is therefore not
hecessary to give a detailed discussion of the subject. Fig. 6 illus-
trates the appearance of the lower part of the stem in cross-section,
showing the relative position of the roots of various ages- The roots
near the center, which are outlined by dots, are those in meristematic
condition; next to them are the mature active roots which are indi-
cated by shaded circles; beyond the mature roots are those which are
dead and crushed. The relation of the sets of roots is indicated in
Hg. 19 also. The arrangement of the roots in the two-lobed species
"as first correctly worked out by HOFMEISTER; the youngest sets of
Toots are those nearest the furrow; and of those in the furrow, the
Youngest are at the ends.

THE ROOT

i eal of the roots has been described 50 many ae 8
cessary to take it up in detail. The most interesting feature
— y are collateral and monarch, without showing Re ee
it is a . As the root bundle passes out from the ats < fs :
ess hae or flattened bundle surrounded by parenc si : s es
sin. a from the stem, phloem replaces the Jeg so ne
a ae rom the center of the stem. The phloem 1S a aed:

ey and is small in amount in comparison wit ao

316 BOTANICAL GAZETTE [APRIL

protophloem and protoxylem are usually differentiated about the
same time, but the protophloem occasionally precedes the protoxylem.
The complete differentiation of the phloem is usually complete before
that of the xylem. The differentiation of the xylem in the roots is
much slower than in the leaf traces. Fig. 7 illustrates the structure
of a root bundle before it is completely developed and while it is still
in the cortex. The eccentric position of the bundle in mature roots is
shown in fig. 8. While the root bundle is making its way through
the cortex the endodermis is poorly differentiated, but in the mature
root outside of the stem it is well defined. The pericycle is poorly
developed. It is usually entirely absent opposite the phloem, but
opposite the xylem there is usually a small amount.

THE LEAF

The leaves of the four species studied present considerable varia-
tion, both in general appearance and anatomy.

The leaf of I. echinospora var. Flettii is moderately stout, and
quadrangular in outline (fig. g). In the cortex and lower part of the
leaf the bundle is fairly well developed, but above the sporangium it
becomes a slender strand. The bundle is collateral throughout, bat
tends to become concentric in the upper part. The differentiation
of the phloem regularly precedes that of the xylem, as Was ioe y
Krucu (17) in the species which he studied. The protoxylem - an
cases is plainly exarch, but sometimes the differentiation 1s ai
simultaneous. In this species I found no cases of distinctly —
protoxylem. It is common for the metaxylem to develo
directions from the protoxylem, but I found no case in
developed on the fourth side, i. e., on the side towa :
although Krucu and Scorr and Hr find that in their spe™”
he sporangiu™-
different
mesarch
protoxylem is sometimes present. While the protoxylem - age
xylem are both composed of spiral and a
tracheids with irregular spirals, the protoxylem ca : “
recognized by the fact that the spirals and rings arf ate are the
protoxylem cells of the lower part of the leaf are small an

1909] STOKEY—ANATOMY OF ISOETES 317

first to show signs of crushing, but those in the upper part are large
and later form canals by the loss of the thickened portions. In the
upper part of the leaf the protoxylem is usually limited to a single
large vessel, whose place can be recognized in old leaves by the large
canal formed by its disappearance (jig. 13); but in the lower part of
the leaf, as well as in the leaf trace, the protoxylem cells are smaller,
several in number, and less sharply marked off from the metaxylem.
Fig. 10 represents the section of a young leaf near the tip, in which the
only indication of the protoxylem is the large central cell, which is not
yet thickened, while the phloem is represented by several cells. The
phloem consists of very long and slender tubes whose sieve plates are
terminal. They come out clearly in sections stained in anilin blue
ot Bismarck brown. Although several preparations suggest the
presence of thin areas on the lateral walls, the evidence is not con-
clusive. In the portion of the leaf above the ligule there is a very
abrupt decrease in the amount of xylem, but not a corresponding
decrease in the phloem. The phloem is well developed throughout,
but instead of forming a band on one side of the bundle, as in the
lower part of the leaf ,itformsanarc. The phloem does not disappear
from the center of the arc, leaving two lateral groups such as KRUCH
has described in I. Hystrix, I. Duriaei, and I. velata, but the amount
in the center of the arc is variable. Near the top of the leaf the
m is sometimes most abundant in the middle of the arc. In
ig. 14 is shown a leaf trace from the cortex at the base of the leaf.
big is the type of bundle which is found in the leaf trace as it traverses
i nu part of the cortex, and in the leaf itself from its base to the
Ea €. Just above the ligule, the bundle passes abruptly me the
Shh type of fig. 13. In the former, the tracheids, which are
Hy uniform in size, are scattered among parenchyma cells that
ic € abundant protoplasm and large nuclei. Before the sporangium
a the outer vessels, the protoxylem, hegin to collapse. In
a of bundle the phloem is always in a band and see ea
aie to surround the xylem. The amount of phloem 1s a .
ian Pper part of the leaf, and by the time the leaf is ma : -
t entirely obliterated by crushing. In following the bundle
"Tough the cortex toward the center of the stem the first change seen

is a . . -
condensation of the bundle by a reduction in the amount of paren:

318 BOTANICAL GAZETTE [appr

chyma, although a few parenchyma cells usually remain in the center
as long as the leaf trace can be identified (jig. 15). For some distance
the compact bundle. remains distinctly collateral, but the phloem
decreases in amount, and near the stele disappears entirely, the leaf
trace consisting of a strand of xylem with a few parenchyma cells
in the center and a parenchyma sheath. The xylem cells become
shorter, and it is no longer possible to distinguish between protoxylem
and metaxylem. The next stage is the merging of the tracheids and
parenchyma of the various leaf traces to form the central axis.
While in general the leaf traces of the other species are similar to
those of I. echinospora var. Flettii, they present certain interesting
variations. The leaf of J. Tuckermani var. Harveyi is also quadran-
gular, but is shorter and more rigid. The bundle, however, is not
so well developed, either in amount or structure. The tissues are not
well differentiated; the phloem in particular is much less sharply
differentiated than in the other three species. The thickening of
the walls is slight and sieve plates are not evident. The amount of
xylem is noticeably less, but the phloem, although not abundant, Is
not so much reduced as the xylem. In the lower part of the leaf the
phloem is a narrow band, but in the upper part of the leaf it forms an
arc and tends to surround the xylem. The formation of sieve cells
begins in the middle of the band, or arc, and extends around three
sides of the xylem. The phloem in the center of the arc is small e
amount and more rudimentary than that on either side, so that 1
places there are indications of a tendency to form two lateral groups-
The sieve cells in the lateral groups are smaller, as well: a8 .
numerous, and they continue to function long after the first-fo :
cells have become crushed and functionless. In this species "°
sporangia are comparatively small.
I. melanopoda has very long, moderately stou
sporangia. The leaves have a greater diameter than those *
Tuckermani var. Harveyi, but they are very much longer and — in
less stout. The amount of xylem is greater, and the bundle a
general stronger than in the two preceding species. The leaf :
in the cortex of the stem contains on an average tee ea
tracheids as that of I. Tuckermani var. Harveyi, while : as
trace above the sporangium there may be three to eight 174

t leaves, with large

# Eh ESS eee fee eee || ce te

1909] STOKEY—ANATOMY OF ISOETES 319

instead of one to four, as in J. echinospora var. Fleitii or I. Tucker-
mani var. Harveyi. As in the other species, the phloem forms an arc
in the upper part of the leaf. As the phloem is abundantly de-
veloped at the sides of the arc and poorly developed in the middle,
there is a tendency for it to separate into two lateral groups.

The leaves of I. Nuttallii differ considerably from those of the
other species, both in external appearance and structure. They are
triangular in outline, long and very slender, indeed almost thread-
like. The sporangia are extremely large, while the leaf tissue, in the
sporangium region as well as above, is notably small in amount. The
bundle, however, is well developed and larger than in any of the
preceding species. It differs conspicuously from those of the other
species in the large amount of xylem present in the bundle above the
sporangium and in the upper part of the leaf. Although, as in the
other species, the bundle becomes reduced above the sporangium,
there are usually eight or ten tracheids and sometimes fifteen or sixteen
present for a considerable distance above the sporangium. In the
Tegion above the sporangium, as in the other species, the protoxylem
Consists of large cells, which are later replaced by canals. The walls
lining the canals are heavily lignified. ‘There is no trace of,an endo-
dermis, such as Scorr and Hirt find in J. Hystrix. The protoxylem
inthis region is occasionally mesarch, a few tracheids of the metaxylem
developing on the side toward the phloem, although the greater
amount of metaxylem is always on the adaxial side of the leaf,jwhere
it frequently forms a narrow band. In the region of the sporangium
the metaxylem is in the form of a crescent, with the heaviest develop-
ment often at the ends of the horns. In such cases the bundle tends
to become mesarch. The phloem also is well developed and abun-
og sieve tubes regularly separating above the ligule into stout

s. A transverse sieve plate is shown in fig. 11. ;

If the four species are arranged in a series according to the size
pet of the leaf traces, the series is as follows: ne eo ——
ia. nopoda, I. echinospora var. Flettit, I. Tuckermani var. -_
‘ ‘Species are arranged according to habitat, from ——*

quatic, the order would be the same, with J. N uttallii as the most
“cag form and I. Tuckermani var. 1: arveyt as the most aquatic.
‘series arranged according to the size of the leaves 1 as follows: J.

320 BOTANICAL GAZETTE [api

melanopoda, I. Tuckermani var. Harveyi, I. echinospora var. Flettii, I.
Nuttallii. A series according to the size of the sporangia is as follows:
I. Nuttallui, I. melanopoda, I. echinospora var. Flettii, I. Tuckermani
var. Harveyi. It is perhaps not safe to generalize from a comparison
of only four species, but it is worthy of: note that the size and
development of the bundle in these forms is not related to the
size of leaf, but follows the other two series, that of habitat and size
of sporangium.

HILL (13) calls attention to the presence of two canals in the leaves
of I. Hystrix, which he regards as representing the parichnos of the
Lepidodendreae. In none of the four species examined is there
any trace of a canal or any indication of a tendency to form canals.
In this respect these species agree with I. lacustris.

THE STEM

As is well known, the stem of Isoetes is a short tuberous body,
whose vascular axis is very small in proportion to the diameter of i
stem. The stem grows in length very slowly, and the apical rout $
left in a deep pit by the overgrowth of the surrounding region. if
the term stele may be applied to a region whose connection with a
plerome is far from certain, then the vascular axis may be defined as
a non-medullated monostele, consisting of xylem and parenchyma
forming a loose network. The xylem is made up of short spiral,
annular, and netted tracheids, whose long axis is transverse to the
stem (jig. 22). The parenchyma cells contain abundant sre
and the nuclei appear active. The xylem axis is surround )
similar parenchyma cells, one to three layers deep, but seal
young nor old plants is there a trace of phloem. A great many it
Plants of I. echinospora and a few of I. Tuckermant vat. arr
were examined, and in all cases the xylem is surrounded by undi z
entiated parenchyma. In very young plants the vascular axis :
exceedingly small, but there is a constant increase both in length am
diameter with the increase in the number of leaves. hich is

The differentiation into protoxylem and metaxylem, W Bex
usually well marked in pteridophytes, is lacking in Isoetes- : ae
neither a difference in the time of development nor in the ea to
of the elements. A difference in character of elements is no! se

BO Se ed cs CA a eee Pe ae, RR ae ay YE gras Ca
Pages Se Th | Feta 4 a re Bag oe Oh ye ee ee pee

1909] : STOKEY—ANATOMY OF ISOETES 321

occur in such a slow-growing stem, and a difference in time would
be difficult to detect in a stele whose elements run transversely.

Scort and Hit have claimed a slight differentiation in time in
certain cases, giving as evidence sections which show tracheids at the
outside and parenchyma in the center. But since the xylem is made
up of leaf traces which curve down from the leaves, as in fig. 4, it is
evident that it would be possible to obtain a section which shows
this condition without being an example of protoxylem differentia-
tion. In young stems it is not uncommon to find that at least one
section shows this apparent differentiation into protoxylem and metaxy-
lem. As was mentioned above, an examination of a series of cross-
sections of leaf traces as they approach the central axis indicates that
there is no differentiation into protoxylem and metaxylem in that part
of the leaf trace. Even if it were present in the leaf traces, the
ransverse arrangement of the tracheids would cause it to appear in
tiers rather than in vertical strands, as in other pteridophyte stems.
_ The cambium, which appears very early (fig. 3), begins its activity
in the parenchyma which surrounds the central axis, so that all the
tissues which are found outside this thin layer of parenchyma are
secondary. The secondary tissues of Isoetes have always been
described as anomalous, and so have furnished a fertile field for
observation and theorization. The cells which the cambium cuts
off externally are ordinary thin-walled parenchyma cells, which have
always been called cortex. Whether they represent ancestral phloem
, of course, an interesting question, but there is nothing in their
structure to suggest an answer. This is the great storage region of
the plant, and the amount of this tissue is much greater than at
a internally by the cambium. In the middle and outer regions
a ai the cells become rounded, often lobed,

eaves (fig. 78), and always contain large amounts 0:
_ increases its thickness from year to yeat;
— nual loss by the sloughing-off of the outer layers. In the outer
Tegion there is little or no starch.

2 € tissue formed internally to the cambium,
eS layer,” is that which has aroused the greatest interest. In a
cut section this layer is a glistening white and stands out

atply from the surrounding region. In stained sections It 15 seen

the so-called “ pris-

322 BOTANICAL GAZETTE [APRIL

that the “prismatic layer” is composed of several kinds of cells. In
the four species studied there is a considerable range both in the types
of cells and in their distribution.

In J. echinospora var. Flettii there is found scattered irregularly a
rather large number of active nucleate parenchyma cells, whose walls
are slightly thickened. Associated with them are other cells which
are almost or entirely empty and whose walls are not uniformly
thickened, but have round, oval, or irregular pits. In such cells the
thickening is not very heavy. There are usually other cells in which
the thickening is more pronounced and is arranged in irregular bands
or rings. In some cells there are heavy bands in addition to the
irregular pittings.

Of the four species, J. Tuckermani vat. Harveyi most closely
resembles the preceding species, but it differs in several points. There
is little or no active parenchyma, the entire tissue consisting of cell
whose walls are thickened irregularly. These cells are seldom entirely
empty, but usually contain a little protoplasm and small, apparently
degenerating, nuclei. The cells are for the most part of the type
shown in fig. 23, in which the pits are small and irregularly distrib-
uted, and there are but faint indications of banding. “In the older
plants, however, it is not uncommon to find the thickening forming
more or less definite bands. In the older parts of old plants the ae
become smaller and less prominent, tending to disappear entirely.
Sometimes the pits can be seen in sections stained in Bismarck brow?
when with a less transparent stain, such as Delafield’s hematoxylin,
the pits cannot be distinguished, and the walls appear to have a ee
form and rather heavy thickening. Apparently the thickening we
cell walls'goes on through a period of several years, the first peat
being irregularly distributed, leaving irregular pits, while the “a
deposits include the whole of the wall and tend to obliterate pee

In I. melanopoda we have all the types of cells which have a
described for I. echinospora var. Flettii: active parenchyma Oe
pitted cells, and cells with irregular thickened bands or ae dition
pitted and banded cells are as a rule entirely empty: In 2
to these types of cells, however, there are other cells with ae
thickenings which are slightly lignified, and also tracheids ere
or annular thickenings, whose lignification is pronounced,

ee oS ee ee

{552 ae ieee

1909] STOKEY—ANATOMY OF ISOETES 323

not as heavy as in the tracheids of the central axis or of the leaf traces
(ig. 21). The parenchyma cells, however, have thinner walls and
are richer in protoplasm than in the two species just described. The
parenchyma and the various types of thickened cells are in most cases
distributed irregularly, as is shown in fig. 26, although occasionally
in the older parts of old stems there are indications of zonation.

I. Nuttallii shows the same type of cells as J. melano poda, but there
isa difference in arrangement. This species always shows the zona-
tion which has been described by FARMER for I. lacustris and by
Hecermarer for I. velata and I. Duriaei. The parenchyma cells
form layers one or more cells thick, which alternate with layers com-
posed of the various types of thickened cells (fig. 25). The zonation
is evident in young plants and is very striking in old plants, especially
in the older regions of the “ prismatic layer.” The thickened cells are
usually entirely empty. The parenchyma cells of J. Nuétallii are
larger, contain more protoplasm, and have thinner walls than those
of the first two species. The parenchyma is more like that of J.
melanopoda, although on the whole the cells are richer in protoplasm
than those of the latter species. In the older parts of the stem the
thickened empty cells are usually collapsed, so that the zones of
thick-walled cells, which alternate with the w ell-developed parenchyma
ones, are apparently much narrower.

Almost all the writers on Isoetes have called attention to the pres-
cng of the fine-grained starch in the cells of the “ prismatic layer.”
This Was first noted by HEGELMATER in J. velata and J. Duriaei, in
Which the starch-containing parenchyma cells form zones alternating
With zones of empty cells. FARMER records the presence of starch
m the prismatic layer of I. lacustris, in which the starch-filled cells
ate also arranged in zones. The disposition of the starch was found
to Vary with the species. In J. echinospora vat. Fleitii and I. melano-
ing it is present abundantly in the cortex but not at all in the “pris-
matic layer.” In I. Tuckermani var. Harveyi, in addition to the
Starch in the cortex, there is starch in the parenchyma in the vascular
axis, and in th® layer of parenchyma surrounding the axis, but there
rei in the “prismatic layer.” J. Nuétallii shows what seems to be
_. common arrangement in the forms previously described ;

, there is abundant starch in the “prismatic layer.

324 BOTANICAL GAZETTE [APRIL

It should be noted that the only one of the four species which con-
tains starch in the “ prismatic layer” is the only one which shows well-
marked zonation, and is also the one in which the parenchyma cells are
the largest and contain the most protoplasm.

Discussion of secondary thickening

A study of the structure of the cells composing the “prismatic
layer” is of interest only as affording a basis for an interpretation of
the nature of the layer. The interpretation which is accepted in
the most comprehensive of all recent works on pteridophytes, Bower's
Origin of a land flora, as well as in CAMPBELL’S Mosses and ferns
and other current texts, is Russow’s theory, which more recently has
received the indorsement of Scott. As was mentioned above, they
look upon the “prismatic layer” as a complex of tissues, consisting of
parenchyma, phloem, and xylem. Among recent writers SMITH 1s
the only one who has even suggested that the tissue may be of a less
extraordinary nature. :

The position of this layer would naturally lead to the conclusion
that it is secondary xylem, but for the fact that its composition is not
what we have been accustomed to look upon as characteristic of that
tissue. The parenchyma is more abundant, while well-det
tracheids are not only few in number in most species but exceedingly
rare or entirely absent in others. In addition to the parenchyma
tracheids, there are the pitted cells, which have been Tega
phloem. It may be well at this point to consider the evidence upo?
which this claim is made. Scorr and HI1 say: ae

The phloem elements have an extremely characteristic structure of ae
walls which comes out conspicuously in sections stained in hematin oe
walls are much pitted, the thicker bands of membranes between the ais ssi
fine bars into
areas. Little of the nature of formed contents can usually
times small, deeply staining globules are found adhering to the walls, spat
ently localized at the pits. In the older parts of the stem the phloem is #0

walls and almost filling the cavity. The masses stain like an anti results.
Soda, but the other callus reactions tried did not give wholly ss as have
We have not investigated the more minute histology of the rete be left t°
not demonstrated the perforations of the thin-walled areas. eS nucleate
other investigators, but in the meantime, we can scarcely doubt that thes? ©

1909] STOKEY—ANATOMY OF ISOETES 325

elements, with the characteristic areolations of their walls, and their agreement
in yarious reactions with the sieve tubes of the leaf, with which we shall see they
are continuous, are best to be regarded as themselves representing the sieve tubes
of the stem.

If no other explanation of these structures were possible, the rea-
sons given above might be accepted as sufficient proof of the phloem
nature of the pitted cells, although the fact of a cambium cutting off
both phloem and xylem from the same face is so extraordinary that
one does not expect the advocates of such a theory to content them-
selves with leaving the burden of proof to other investigators. The
only analogy is in the case of Dracaena and its allies, whose anomalous
secondary thickening has been frequently referred to in the literature
on Isoetes. Before Russow’s theory made its appearance, the tissue
on the inside of the cambium had been regarded as secondary xylem.
This was the very natural interpretation given by HOFMEISTER in his
Higher Cry ptogamia and was accepted, apparently without question,
until Russow’s more critical work appeared. If one is not willing to
accept the Russow theory, the natural alternative is to regard the
“prismatic layer” as secondary xylem. This, of course, requires an
explanation of its unusual structure. The presence of a large amount
of parenchyma in the secondary wood is unusual but not without
parallel, as this is the case in certain Lepidodendreae, e. g., Le ido-
p hloios fuliginosus. The presence of unlignified pitted cells, of course,
is the situation which has led to controversy, and is the chief point
o be explained. However, the presence of pitted cells of a phloem-
like aspect does not necessitate the assumption that the cells are
Phloem, since that structure is found in cells of other tissues, ag
example, the cortical parenchyma of Helminthostachys seylanica.
FARMER and FREEMAN (10) in their description of the pits in the cells
of the cortical parenchyma say:

The pits are remarkable, forming, as they do, not merely simple depressions
J ‘ like the pores of a sieve

and they do not differ

The presence of cells of this type in tissues which do not even

petits to part of the bundle indicates that pitted walls are not neces-
sarily to be taken as an indication of phloem. The usual phloem

326 BOTANICAL GAZETTE [APRIL

tests applied to the “prismatic layer”’ do not give any positive results.
In the case of the cells in question it can be shown that not only are they
not phloem but that they are xylem.

A careful examination of the “prismatic layer’’ of such forms as
I. Nuttallii and I. melanopoda will reveal the fact that, while the
secondary tracheids are far removed in appearance and staining
reactions from the pitted cells, an almost perfect transition series
exists between the two types. Several stages are shown in jig. 21.
It is possible to trace a series from the tracheids with lignified spiral or
annular thickenings, through those with less regular thickenings and
with a smaller amount of lignin, to those in which the thickening is
very irregular and which have no trace of lignin. A combination of
safranin and anilin blue was found to be particularly valuable in
revealing slight amounts of lignin. With this variation in wall
thickening and lignification, there is correlated a variation in the
amount of cell contents. The existence of the transitional stages
leads almost inevitably to the conclusion that the various types of cells
of the “ prismatic layer” differ essentially only in their stage of develop-
ment, and that the layer accordingly consists of mature tracheids,
‘immature tracheids, and parenchyma. While the series in I. echino-
Spora var. Flettii is less perfect, it is very suggestive, but that of J.
Tuckermani var. Harveyi is usually too limited to afford much of an
indication of the nature of the pitted cells.

The recognized steps in tracheid development are as follows: -
more or less regular thickening of the wall; the loss of cell contents;
and the lignification of the wall. It should not be assumed, eee
a parenchyma cell undergoes the first steps of the changes whi ;
would lead to the formation of a tracheid, that there is any ope
necessity for their continuance. It is perfectly possible that x
course of development might be arrested at any point, and that a
or all of these changes might be incomplete, according to the en
standards of completeness. Unfortunately, the cases of secon a
growth in modern pteridophytes are so few in number and ” pit
in extent that there are very few opportunities for ia 2 esate
this very thing—the incomplete development of the tracheids ther
been described as characteristic of the secondary xylem ° ser
pteridophytes, and indeed is almost made the test of secondary X)"""

1909), STOKEY—ANATOMY OF ISOETES 327

BoopLe (1), in his account of the secondary thickening in the roots
of Ophioglossum vulgatum, figures both a cross and a longitudinal
section of secondary tracheids, which he describes respectively as a
“developing tracheid with its protoplasmic contents” and as a
“longitudinal section showing part of the xylem with one developing
tracheid containing protoplasm and a nucleus.” In both these cases
the tracheid character of the cell is too pronounced to be questioned.
Inthe case of Angiopteris evecta, HILL (14) says with reference to the
secondary growth: “Semi-lignified elements with protoplasmic con-
tents are found on the inside of the meristem.”

If seems strange indeed, that while, in the case of other pterido-
phytes, it is taken as a matter of course that tracheids may retain part
of their protoplasmic contents and appear in various stages of develop-
ment, in Isoetes the same condition has been looked upon as an jnsu-
perable objection to the tracheid character of the cells in question. It
is obviously much more natural to interpret this tissue as a case of
arrested development, than to regard it as anything so extraordinary
“s combination of xylem and phloem. The fact that we find cases
of immature and imperfectly developed secondary xylem in other
pteridophytes is more enlightening as a basis of interpretation than is
the presence of anomalous secondary thickening in the far-removed
Dracaena. :

The irregular disposition of the thin areas in the pitted tracheids,
while not usual in the pteridophytes, is probably more common than
has been supposed. GwYNNE-VAUGHAN (11), in his recent work on
the tracheae of ferns, calls attention to the irregularities in the Osmun-
— aati and others, illustrating cases of distinctly pitted

. Farmer and FREEMAN (10) describe @ Tange of structure 1n
~ xylem of Helminthostachys zeylanica from tracheids “with char-
ait bordered pits of an oval or even circular form’’ to those a
which the pits assimilate to the more scalariform type met
majority of ferns.” In Lycopodium in the primary xyle
Pa mostly long and narrow, of the scalariform type;

racheids they may be round or oval, giving the tracheid a
i pitted appearance. A study of the seal "
young SS an opportunity for a comparison oP Sak
: acheids, a comparison which is of interest as throw!ng agente

328 BOTANICAL GAZETTE [APRIL

the nature of the pitted cells of Isoetes. In the apices of Lycopodium,
where the xylem is not yet mature, it is not difficult to find tracheids
that have round or oval pits, and which before they are lignified present
a similar appearance to that of the pitted cells of Isoetes. In fact, in
L. pithyoides, the fully developed sieve cells and the pitted tracheids
in which lignification has not yet taken place differ only in the greater
regularity of the pitting of the tracheids. If the development of the
more slender tracheids of Lycopodium were arrested before lignifica-
tion had begun, it would be difficult to distinguish sieve cells from
tracheids except by position, since both have so nearly the same general
appearance and the same reactions to stains. As the disposition of
the thickening of the primary tracheids of Isoetes is much less regular
than in other pteridophytes, it might reasonably be expected that the
secondary xylem would also show irregularities.

One of the reasons given by Russow and subsequent observers for
regarding the “prismatic layer” as part phloem, is that it is in direct
continuity with the phloem of the leaf traces. I am not disposed t
question the nature of the phloem either in the leaves or roots. There
is nothing in its position and structure to cause any hesitation about
accepting it as phloem. The sieve tubes possess well-defined sieve
plates, and the tissue as a whole is so definitely marked off, both: im
position and development, from the xylem and all adjacent tissue,
that there is no apparent reason for questioning its identity as phloem.

There can be no question of the continuity of the old leaf traces and
the “prismatic layer” in such cases as are shown in jigs. 19 de :
in the case of young leaf traces the point is not so certain. age
cations are that the continuity is a result of the overgrowth a h
leaf traces by the secondary tissues. Fig. 17 is a diagram wie
illustrates a thing that occurs in some if not in all cas

ery large

continuity exists. In any old stem there are present a V
ly few at the rep
‘ : ‘tematic
are alive ive. em is a meriste
and active. The upper part of the st one whose

diagram by fine dots. The xylem strands of the various ae
that level move in together, forming a more compact TES! or four
vascular axis. The phloem ends in the parenchyma, three

STOKEY—ANATOMY OF ISOETES 329

cells from the vascular axis, in the region in which the cambium has
not yet become defined. As the meristematic region becomes localized,
forming the cambium, the parenchyma with which the leaf trace
phloem is connected is pushed out farther and farther from the
vascular axis by the secondary tissues. The phloem can retain its
continuity with this region in three ways: the xylem at the base of
the leaf may elongate sufficiently to compensate for the secondary
growth in that region; there may be a splitting-apart of the tissues
of the bundle, permitting the phloem to slide along the xylem; or
the phloem of the leaf trace will be torn apart, leaving one end con-
nected with the “ prismatic layer” while the other is carried out into the
cortex. Undoubtedly, while the leaf trace is young, there is an adjust-
ment by the first method. It should be noted, however, that the leaf
traces which are connected with living leaves are comparatively few in
number and are found in that part of the stem in which secondary
growth is scarcely observable. At some time in the development of
each leaf trace there comes a time when it is no longer capable of
extension and is unable to keep pace with the development of the
stem. At this point the tissues of the leaf trace give way, and the
outer part is carried out into the cortex and finally sloughed off, while
® base becomes more or less crushed and remains as a dead stump,
Nich in time may be completely buried in the secondary wood.
While the conditions which have led to the imperfect development
of the vascular tissues cannot be known with certainty, amons them
— aquatic habit and the reduction or shortening = =
6 omy of Isoetes does not seem to indicate, as MIT
- My 324), “that the genus Isoetes represents a more ane
view eg tophyte than any other vascular plant,” but it geil the
Some dc corr that “the group has clearly undergone reduction nai
form i. complex ty pe, and probably from some highly sci :
nee ycopod, as indicated by the secondary growth, the = .
and ie and the somewhat complex organization of the sae
aoe ech of the axis.” As a — i
rally be lee nny reduction in stem development, there would na -
of the xy] uction in the xylem. This might be either in the ere
semndary we even to the extent of the entire disappearanc® )
wood, or there might be a reduction in the development

%,
®

330° . BOTANICAL GAZETTE [sprit

of the xylem. In the other modern pteridophytes it is apparently the
former which has taken place, secondary xylem appearing rarely and
in small amounts; and, as Hit (15) infers from a comparison of the
examples of pteridophytes showing the phenomenon, it is more prob-
ably an example of reduction than a new development. In Isoetes,
however, the reduction in the bundle has not been limited to the
xylem portion, but has extended to the whole of the phloem, both
primary and secondary.

The position of Isoetes

The phylogenetic connections of Isoetes have been discussed in all
recent papers, with the great weight of evidence in favor of a lycopod
ancestry. The evidence as to its relationship afforded by its anatomy
has been taken up recently by BowER (2), on the basis of —
pretation given by Scorr and Hrxt, with the conclusion that Isoetes
is in its anatomy a lycopod, with a stem structure which can be
explained by regarding it as a stunted lycopod. In his eagerness 5g
show a unity of structure in the Lycopodiales, BOWER sect oa
following statement (p. 339): “Throughout the Lycopodiales ‘
foliar traces are inserted peripherally, and with only a slight -
disturbance upon the periphery of the cauline xylem ore
of the questionable existence of a cauline portion in the xylem
there seems to be little justification for so sweeping a ener
is in this very thing that Isoetes differs markedly from other :
lycopods, although the difference is not of such a senna
make the relationship doubtful. The difference js correlat om
the stunted habit, and such differences of body habit have nev! :
admitted to have great weight in determining the large a.
The stunted habit of stem is not limited in Lycopod a ‘ um.
but it occurs also, though of very different type, in ere of

Even among those who recognize the strong lycopod dvisable to
Isoetes, it is suggested occasionally that it might be aev’ ales-
separate Isoetes from the lycopods and establish a new -_
The present tendency seems to be toward a raultipl :
so that it may be well to consider the desirability ei ea inted

The closest connection of Isoetes is, as has been meee jae
out, with the Lepidodendreae, although it has many pol

of orders,

TSE ee ee ee Oe

1909] STOKEY—ANATOMY OF ISOETES 331

mon with modern lycopods. The spore-producing members, in
sructure and development, are unquestionably of the lycopod type,
and as such present no obstacle to the retention of the group in the
Lycopodiales. With reference to the anatomy my work would seem
to strengthen the position of Isoetes in the Lycopodiales. The irregu-
larities of its structure are not of such a nature as to isolate the group.
Aside from the possible lack of a cauline portion in the stele, the
irregularities of its anatomy are limited to the absence of primary
phloem in the stem; the absence of secondary phloem; the lack of
differentiation into protoxylem and metaxylem in the stem; the
large amount of parenchyma in the secondary wood; and the imperfect
development of the wood.

The absence of primary phloem seems to be characteristic of Isoetes.
There are no indications of it in the four species described in this
paper, and Scorr and Hix say of J. Hystrix “that it is not possible
to identify primary phloem with certainty.” Although the absence of
primary phloem is recorded for juvenile pteridophytes (e.g. Matonia
pectinata) by TANsLEy and LuLHaM, (23, P- 482), so far as is known
i is present in the stem of all other adult pteridophytes. In this
point, then, Isoetes stands alone. The presence of phloem in the leaf
traces and roots, and the collateral arrangement of the bundle in
the lower part of the leaf trace indicates a descent from a line in
which the phloem is present in the stem as a layer around the xylem.
It is difficult to tell what importance to attach to the tendency of the
leaf trace to become concentric in the middle and upper part; and to
the occasional occurrence of mesarch xylem in the leaf trace.

: In tegard to the next point, the absence of secondary phloem ina
&m with a cambium, the isolation of Isoetes 1S less certain. SCOTT

£ the Lepidodendreae

“ays: “Although the presence of primary phloem can be recogni

‘00, that there are cases among the modern pteridophytes in which
Say xylem is produced, but no secondary phloem has been
tved. This suggests that when reduction occuts in a form with

Secondary thickening, the disappearance of the phloem precedes that
of the xyl em,

332 BOTANICAL GAZETTE [APRIL

The next peculiarity of the stem anatomy, the absence of differ-
entiation into protoxylem and metaxylem in the vascular axis, is so
obviously related to the stunted habit of the stem that it contributes
nothing to a discussion of the position of Isoetes.

In the peculiarities of the structure of its secondary wood, Isoetes
finds its nearest prototype in certain Lepidodendreae, as has been
pointed out by previous writers. The forms which present the

-greatest similarity of structure are Lepidophlotos fuliginosus and
Lepidodendron obovatum. According to Scott (21) the cambium
in Lepidodendron obovatum produces parenchyma only and no tra-
cheids; but the cambium in Lepidophloios fuliginosus produces either
secondary parenchyma only, or secondary parenchyma in which
are imbedded groups of tracheids. Certain species of Isoetes present
a. close approximation to both conditions. In some species the second-
ary xylem is almost wholly parenchymatous, with no well-formed
tracheids and only a few immature tracheids. In other: cases there
are groups of tracheids associated with the parenchyma, and in addi-
tion a certain amount of immature tracheid tissue. Isoetes accord:
ingly differs from these two members of the Lepidodendreae only in
the presence of immature tracheids in the secondary xylem. =
not improbable that future work in the Lepidodendreae may bring
to light a similar situation in that group. d

In point of anatomy, then, there seems to be no adequate grown
for the separation of Isoetes from the Lycopodiales. ser

The strongest argument for the establishment of a separate © #
has been drawn from the gametophyte generation, in the .
a multiciliate sperm. While this is a character of great impor™

we should consider the extent of our evidence before attaching a
much weight to it. It must not be forgotten, moreover, that -

sperms occasionally depart from the biciliate tyP® enn

(4, p. 32) speaks of the occasional occurrence © .

cilia in Lycopodium clavatum. If we reflect that our

sperms of the modern genus Lycopodium is limited to t

species, and that we have no knowledge at all of the sP
more closely related Lepidodendreae, we may be less

regard the character of the sperms as preponderant 1?

the position of Isoetes.

erms of the
inclined t0

1909] STOKEY—ANATOMY OF ISGETES = 333

Summary

1. The vascular axis is a non-medullated monostele, composed of
tracheids and parenchyma. There is no differentiation into protoxy-
lem and metaxylem.

2, There is no primary phloem in the stem. It is found in the
leaf traces and root bundles only.

3. The cambium gives rise to cortex on the outside and secondary
xylem on the inside. The so-called “prismatic layer” is secondary
xylem. The cambium does not form phloem.

4. The secondary xylem consists of various combinations of the
following types of cells: (a) Spiral and annular tracheids. (6) Imma-
ture tracheids, slightly lignified, with irregular rings or spiral thicken-
ings. (c) Immature tracheids, unlignified, nucleate or enucleate,
with irregular rings or spiral thickenings. (@) Immature tracheids,
nucleate or enucleate, with slightly thickened, pitted walls. (e) Par-
enchyma cells, which may have little protoplasm and small nuclei, or
abundant protoplasm and large nuclei.

5. The secondary xylem of J. Nudtallit shows zonation. J. echino-
Spora var. Flettii and I. Tuckermani var. Harveyi do not. I. melano-
poda shows it occasionally in old stems. Starch does not occur in the
secondary xylem except in the parenchyma zones of I. Nuttallit.

6. The root bundles are collateral and monarch. The protoxylem
sfound on the side away from the phloem and toward the center of the
stem, i. e., it is endarch.

7. The leaf traces are collateral, but tend to be
the middle and upper part of the leaf. Thy xylem portion undergoes
great reduction above the sporangium, but the phloem is not reduc
correspondingly. The sieve plates are transverse: Ses
: 8. Near the vascular axis the leaf trace does not show differentia-
tion into protoxylem and metaxylem. In the outer part of the cortex
= in the region of the sporangium it is usually exarch. In I.
Nuttallii it is occasionally mesarch above the sporangium and in the
Tegion of the sporangium.

come concentric in

This investigation was conducted at the University tw

u Rist
nder the direction of Professor JouN M. CouLTER an

334 BOTANICAL GAZETTE [APRIL

G. LAnp, of whose advice and encouragement I wish to express my
keen appreciation.

Mount Hotyoke COLLEGE
SoutH Hap Ley, MAss.

LITERATURE CITED

. Booptr, L. A., On some points in the anatomy of the Ophioglossaceae.
Annals of Botany 13: 379-394- pl. 20. 1899.

. Bower, F. O., The origin of a land flora. London. 1908.

3. BRUCHMANN, a. Ueber Anlage und Wachstum der Wurzeln von Lycopodium

und oo Jena 1874.

’ r die Acragee und Keimflanzen mehrerer europiischer

Lo ciesae. "Gotha.

CAMPBELL, D. H., Contribution to the life history of Isoetes. Annals of

Botany 5:231- 256. pls. 15-17.

, Mosses and ferns. New Yoik. 1905.

De Bary, A., The comparative anatomy of phanerogams an

Trans. Oxford. 1884.

Eaton, A. A., The genus Isoetes in New England. Fernwort papers. oe

FARMER, 5 B. , On Isoetes lacustris L. Annals of Botany 5237-02. pls. 5,6
1890.

10. Farmer, J. B., AND FREEMAN, W. G., On the structure and affinities of

‘Helminthostachys zeylanica. Annals of Botany 13: 421-445. pls. 21-23. 1899.

lal

d ferns. Eng.

Cm wan

Ir. GwWYyNNE-VAUGHAN, D. T., On the = nature of the tracheae in ferns.
Annals of Botany 22:517-521 08. : 481

12. HEGELMAIER, F., Zur Kenntniss pris Lycopodinen. Bot. Zeit. 32:49;
497, 513. 1874. ;

13. Hi, T. G., On the See of a parichnos in recent plants. Annals 0
Botany 20:267-273. pls. 19, 20. 1906.

14. , On secondary thickening in Angiopieris evecta. Annals of Botany
16: 173, ed sete

idophytes. New Phil 5: dan totee™ oe
* a ie]

z= Horie, W, The haber Cron iaeeon della Fogle . bce
“ a sacar se Untersuchungen der Leitbiindel- Kryptous®®
19. og Gee a oe ee structure of [soetes Hystrix. —

ndar

Botany 14: 413-454. pls. 23, 24. Rei
20. Scott, D. H., The pres sent cies i paleozoic botany: Pega’

Botanicae. 1906. < of Botany
21. ———, The structure of Lepidodendron obovatum Sternb. Annals ©

20: 317-319. 1906. and spo-
22. SurrH, R. W., The structure and development of the ag

rangia of Thoetes. Bor. GAZETTE 29: . 323. pls. 13-20 ue r system
23. TANsLEy, A. G., anp LuLHaM, Miss R. B. J., A study ofthe 33» 1995:
of Matonia pectinata. Annals of Botany 19:475-519 ——

SY Oe is

ey ead a, aS ey ny aah ella aloe RT ie NA ES NN eee ns a at ed na OE

L GAZETTE, XIVU . PLATE XXT

Say

Se Aa nat
Ae a Ke INN

aS on al
re oN)

f Wy) Pi

Rice

NY =

WS
SS

a isaies -.

LAY ee
ube Wing me toes
ote BV * a
MAR. &

Von
: ayn
i

_-
=o

SHAM 7 PF HSe
eeu a

eek

ae A Is) i) CW ayy rhe Ta AS
3% DN D9 Pellet pa Le ES FASE ol ee
Ce Night ss Wis ipa

re
455 e

sTOKkY on LSORTES

1909] STOKEY—ANATOMY OF ISOETES 335

24. Von Mont, H., Ueber den Bau des Stammes von Jsoetes lacustris. Linnaea
14:181. 1840.

EXPLANATION OF PLATES XIX-XXI
PLATE XIX
Figs. 1-4.—I. echinospora var. Flettii. Figs. 5-8.—I. melanopoda

Fic. 1—Longitudinal section of the stele of young plant cut in the plane of
the furrow. X 380.

Fic. 2.—Longitudinal section of the stem of young plant cut in the plane of
the furrow. X37.

Fic. 3.—Cross-section of the stele of a young plant. X 380.

Fic. 4.—Longitudinal section of a young plant cut across the furrow. X22.

Fic. 5.—Cross-section of the stem of old plant in the leaf-trace region. x6

Fic. 6.—Cross-section of stem below the stele showing root bundles in the
cortex. 6.

Fic. 7.—Cross-section of root bundle before it has left the stem. 380.

Fic. 8.—Cross-section of mature root. X48.

ATE XX
Figs. 9, 10, 12-16.—I. echinospora var. Flettii. Fig. 11.—I. Nuttallii
Fic. 9.—Cross-section of mature leaf near the middle. X37.
Fic. 10.—Cross-section of bundle of young leaf near the tip. X380.
Fic. 11.—Sieve plate from sieve tube. 810. ea
Fic. 12.—Cross-section of bundle of a young leaf near the middle. X 380.
Fic. 13.—Cross-section of bundle of mature leaf cut just above ligule. X 380.
Fic. 14.—Cross-section of leaf trace in cortex just below base of leaf. X 260.
Fic. 15.—Cross-section of leaf trace in cortex half-way between leaf and
vascular axis. 380
Fic. 16.—Cross-section of leaf trace near the vascular axis. 380.
Fic. 17.—Diagram to illustrate the relation of leaf traces to secondary wood.
Fic. 18.—Cells of cortex with starch grains.

XI
Figs. 19, 20, 22.—I. echinospora var. Flettii. Figs. 21, 24, 26.-—I. melanopoda
Fig. 25.—I. Nuttallii. Fig. 23.—I. Tuckerman var. Harvey
Fic. 19.—Longitudinal section of vascular bundle of old plant cut across
furrow. 22.
Fic. 20.—Longitudinal section of vascular bundle of old plant cut in the
Plane of the furrow. X22.
1G. 21.—Tracheids from secondary xylem at different stages of develop-
Ment, X 810.
Fic. 22.—Cross-section of vascular axis of old plant. X175- ae
Fic. 23.—Tracheids from secondary xylem; the cell on the right is sectioned
obliquely. X8r0,
. Fic. 24.—Cross-section of vascular axis through the root region; the long
“xls is in the plane of the furrow. X22.
Fic. 25.—Cross-section of stem from the edge of the primary xylem to the
“ottex, showing zonation in secondary xylem. X450.
Fic. 26.—Cross-section of the stem from the edge of the primary xylem to the
Cortex, X 350. :

CURRENT LITERATURE

BOOK REVIEWS
Evolution of the filicinean vascular system

Professor Tansley: has done well to gather under one cover his lectures on the
vascular system of ferns, previously published in the New Phytologist. The
aitempt is made ‘‘to gather together the results that have accrued from the
researches on the morphology of the vascular system of ferns which have been
undertaken during the last few years, and to present these results from an evo-
lutionary standpoint.” The first lecture discusses various theories which have
been advanced to account for the origin of the main phyla of Pteridophyta. The
author favors the view of a direct derivation from Algae in which an alternation
of generations had already been established. Accordingly, the sporophyte of
pteridophytes would not correspond to that of bryophytes, in which an antithetic
alternation of generations seems to have been worked out. The author admits
that the presence of an archegonium in both mosses and ferns is an obstacle to

vanced

and appear to have branched dichotomously;
from a lycopod form is not credited. It is even suggested th
forms, such as Lycopodium, may have been derived by reduction from mega
phyllous ancestors. The whole scheme of phylogeny proposed is largely specula-
tive, but such attempts will be welcomed by those who find difficulty in accepting
Bower’s well-known hypothesis. ved

In the second lecture the Botryopterideae are reviewed, and much scattere
information on this group is rendered available. Proceeding from the protostelic
condition exhibited by Grammatopteris, the complications shown by Zygoptens

the
Hymenophyllaceae the mode of exit of the leaf-traces lends ase 2S
author’s view of the identical nature of leaf strand and stem stele. In ’s work

ing this family, as well as the Gleicheniaceae and Schizaeaceae, asa i Phy-
is freely drawn upon and is presented from the evolutionary standpoint. e
logeny as indicated by the stele is compared with that inferred from the 0
gia, and a general correspondence is claimed, though the latter criterion 1S
sidered to be the more reliable.

In the sixth lecture the evolution of a protostele into a sole
is considered with reference to the examples found in the foregoin

phonostele)

ostele (si

co s
« TansLey, A. G., Lectures on the evolution of the filicinean ss Author.
New Phytologist reprint no, 2. Paper. 8vo. pp. 144- Cambridge:
1908. 38s. 6d.

336

1909] CURRENT LITERATURE 337

Although no explicit statement is made, the reader is apparently left to infer that
there are two modes of origin of a hollow stele: (1) one in which the central trach-
eids are replaced by parenchyma (Schizaea), (2) one in which fundamental tissue

into “pockets” at the leaf gaps and becomes continuous with the tissue
in contiguous pockets (Alsophila). Although the latter view of the origin of
“pith” is essentially that of JEFFREY, no mention of the fact is made in the text,
but such reference is relegated to the preface, where the author disclaims adher-
ence to this view. The evolution of solenostely into dictyostely, and finally into
polycycly is clearly traced, and the complicated condition found in Marattiaceae is
adequately illustrated by diagrams from various sources.

Concerning the Osmundaceae, the conclusion is reached that the stele does
not represent a reduced type, but shows a gradual progression from the condiiion
seen in Botryopterideae, from which group the Osmundaceae have probably
been derived. The recent work of Kipston and GwyNNE-VAUGHAN on fossil

CHRYSLER,

The American Breeders’ Asssociation

The fourth annual report of the American Breeders’ Associations is, in a num-
ber of features, a decided improvement over previous volumes. The same high
Standard of matter is maintained as in previous reports, but there is more of
itis printed on better paper; and contains numerous fine half-tone engravings.
Unlike many publications which are more or less influenced by practical con-
siderations, the articles presented in the reports of the American Breeders’ Asso-
ciation appear to suffer no diminution of scientific value because of the large
contingent of practical breeders among its membership and on its programs.
Almost every phase of practical and theoretical breeding of plants and animals,
48 well as two interesting reports upon eugenics, the new science of improvement
of the human race, are included. Papers of importance from the standpoint
* the practical plant-breeder include several upon the production of disease-
resistance in various plants by W. A. Orton, P. K. Buiny, and H. L. BoLLey;
© ue eee
_ ’ Jerrrey, E. C. Are there foliar gaps in the Lycopsida? Bot. a.
46:241~258, pls. 17, 18. 1908.

8 Report of the American Breeders’ Association. Vol. IV. pp. 373- pls. 3,
Jigs. 74. 1908.

338 BOTANICAL GAZETTE [APRIL

the improvement of apples and other tree and vine fruits, by S. A. Beacu, W. T.
Macouwn, and J. A. Burton; the breeding of cereals by L. S. Kink and C, E.
SauNnDERS; the improvement of hops by selection and breeding by W. W. Stock-
BERGER; on cotton-breeding by Davip Coxer, H. J. Wepper, and D. A.
Saunpers; the breeding of fiber crops, by J. H. SHEPPARD, L. H. Dewey, Fritz
Knorr, and H. L. Botrey; the breeding of vegetables, by W. W. TRACY; roses
by PETER BIssETT, and W. VAN FLEET; tobacco by A. D. SHAMEL, J. B. STEWARD,
A. D. Setpy, and W. H. ScHERFFIUS; carnations by C. W. Warp; forage
crops by T. F. Hunt and H. S. ALLARD; and forest and nut trees by GIFFORD
Pincnor, W. L. Jepson, and G. L. Croruter. In all of these articles, as well
as in a number dealing with animal breeding, there are many facts recorded
which are of more than passing scientific interest. Papers of a more strictly
theoretical scientific character are: “Organic correlations,” by E. M. East, “Some
gaps in our knowledge of heredity,” by H. J. WessER, ‘“The composition of a
field of maize,” by G. H. SHULL, ‘Recent advances in the theory of heredity,” by
C. B. Davenport, “Color factors in mammals,” by W. J. SPILLMAN, and “Mende-
lian phenomena and discontinuous variation,”’ by W. J. SPILLMAN.
range of subjects and the almost uniform high excellence of the papers and reports
included in this volume show that the American Breeders’ Association has a large
mission to fill, and that it is filling it creditably. These annual reports are made
the treasure-house of all the best things gained in the experience of our foremost
practical breeders and students of heredity during the progress of their work.
The efforts made by the practical breeders to present their experience in as proper
scientific form as possible, and to interpret those experiences ! :
latest scientific results, and the efforts of the scientific breeders to state their
results in as simple, direct, and comprehensible a manner as possible, have 4 —
salutary effect upon all those connected with the American Breeders’ Association,
and must continue to supply us with the best annual crop { information regarding
the factors which enter into the breeder’s work, whatever may be his motive
breeding —GrorcE H. SHULL.
MINOR NOTICES

Sertum Madagascariense.t—This paper is based on two collections of ee
made in Madagascar, one by Joun Gurttor in the district of Vatomandry yee
east coast and the other by Henri Rusition on the plateau of agi :
first part of the work consists of a brief consideration of the botant
and in the second part the author in collaboration with several promine sat
specialists, gives a list of the species. Among the plants recorded 26 ae ath
4 varieties are described as new to science. The larger and mo tical gen ;
‘are accompanied by analytical keys to the species, and several is
been introduced. A complete index to the vernacular and scientific names
et Jard.

4 Hocureutiner, B. P. G., Sertum Madagascariense. Ann. Conserv

Bot. Genéve 11-12:35-135. figs. 23. 1907-1908.

§20-626,

1909] CURRENT LITERATURE 339

added. The work is a notable contribution to our knowledge of the flora of
Madagascar —J. M. GREENMAN.

North American Flora.s—Part 4 of Vol. XXII contains a continuation of Dr.
P. A. Rypserc’s elaboration of the Rosaceae. The groups treated are Potentilla
andthe related genera. In ail sixteen genera are here considered, and to these the
author refers 277 species, of which 70, approximately one-fourth, are described as
new. Potentilla leads with 176 recognized species, 44 being published as new
to science. Two new genera (Zygalchemilia and Lachemilla) are proposed.—
J. M. Greenman.

NOTES FOR STUDENTS

Longevity of seeds.—In a long paper? EWART classifies seeds according to
their duration of life under optimal conditions as: microbiotic seeds, with a longev-
sf of less than 3 years; mesobiotic, with a longevity of 3 to 15 years; and macro-
biotic, with a longevity of 15 to 100 years. Most of the paper (175 out of 210
pages) is taken up with a table, drawn from the works of various investigators,
showing the age, percentage of vitality, etc., of various stored and buried seeds.
Ewart says: “Longevity depends not on the food materials or seed coats, but
upon how long the inert protein molecules, into which the living protoplasm dis-
integrates when drying, retain the molecular grouping which permits of their
recombination to form the active protoplasmic molecule when the seed is moist-
ened and supplied with oxygen.” Longevity, however, he holds, is in general
found in seeds with seed coats impervious to water, and asserts that this imper-
meability is due to cuticular structures in almost all cases examined. In f
se digitata, on the other hand, all layers of the coats are equally resistant to

He agrees with CRocKER that seed-coat characters rather than embryo charac-
ters account for the greater number of cases of delayed germination, and he makes
considerable use of the data of this writer as evidence on this point. He believes
that the longevity of seeds in soil is far less than is generally assumed. t
maximal duration of the seeds of certain Leguminosae under optimal conditions
and of Malvaceae and Nympheaceae
gives the structure

of the coats of various resistant seeds. The body of the work is rred by a

tumber of inexcusable errors in the statement of the results of other investigators.

—Wa. Crocker

See es.—Gnriiss has suggested’ a method of capillary analysis of oa oR
or which he claims considerable value. It consists in pulverizing @ portion 0 is
’ North American Flora, Vol. XXII, Part 4, PP- 293-388- New York Botanica
tden, 1908.
6 :
2r: Ewart, ALFRED J., On the longevity of seeds. Pr
+I-20, pls. ; 2 1908. 7
’ Gris, ak Kapillaranalyse einiger Enzyme- Ber. Deutsch. Bot. G
1908.

‘oc. Roy. Soc. Victoria. N.S.

esells. 26a:

340 BOTANICAL GAZETTE [APRIL

tissue containing the enzyme in a small amount of glycerin and placing this on a
filter paper. From this mass the water circle spreads and the enzymes can be
located at various radial distances from the center. In dealing with oxidases the
whole process is performed in an atmosphere of hydrogen. It is not evident
that-this method is of any great value further than as a mere means of demon-
strating the presence of certain enzymes. Grtiss also claims by it to gain evidence
that cytase is not distinct from diastase, and believes he has shown in a number
of other cases that a single enzyme performs several catalytic functions. His
arguments against the specificity of enzymes are to a degree plausible, but are far
from conclusive.

Griiss also asserts,’ on the basis of considerable experimental evidence,
that the reducing power of fermenting yeast attributed to the action of reductase
can be accounted for by the nascent hydrogen set free by the hydrogenase of the
yeast. In the presence of fermenting yeast the reduction of sodium seleniate
and sulfur occur as they do when treated with nascent hydrogen. He finds no
evidence for postulating reductase in yeast. He believes that the fungi in general
possess hydrogenase and not reductase. If this be true the reductions carried
on by this group of plants are strikingly similar to the simplest reductions in the
chemical laboratory. He agrees that yeast and other fungi show a very slight
reducing power not due to hydrogenase, but the substance that produces this
slight reduction shows none of the characteristics of an enzyme.—WM. CROCKER.

Germination in Rhinanthaceae.—SPERLICH® believes he has demonstrated
that the germination of the seeds of the partially parasitic species, Melampyrum
silvaticum, M. arvense, and Alectorolophus hirsutus, is greatly hastened by the
presence of the host plant. These seeds show a considerable rest period ”

concludes that the favorable action of the host is evident only up to the completion
hat his conclu-

ing” he makes no mention of the general connection of del
germination with the seed coats, but attributes these phenomen
ters. He apparently has no knowledge of the literature on t
wonders if his results are not merely the measurement of seed- nh ig the first
certainly has not demonstrated dormancy in the embryo itself, which ®
step in establishing his main position. The disposition © ies of the
investigators to refer the phenomena of “after-ripening” to oo eT facts
protoplasm is to be deplored, especially when a thorough examination 0 howevel;
will often furnish a very simple explanation. It must not be forgotten,

8 Griss, J., Hydrogenase oder Reduktase? Idem: 627-630 < pflanaliche®
9 SPERLICH eo : n ein von einen ;
, ADOLPH, Ist bei griinen Rhinanthacee nar? ee Deutsch. Bot

Organismus ausgehender ausserer Keimungsreiz nachweis
Gesells 26a: 574-587. 1908

1909] CURRENT LITERATURE 341

that it has been clearly proved that the fungus of the host is necessary for the nor-
mal germination of the seeds of many orchids; but even here our knowledge is
oflittle scientific significance until we know the exact method of the action of the

whether its effect is due to the secretion of certain chemical compounds,
which aid in water absorption, or to some other influence.—WM. CROCKER. -

Chromogens.—TAmMMES"? reports a new chromogen, dipsacan, which is pres-
ent in all the genera and species of Dipsaceae examined. Dipsacan has many
points of resemblance to isatan and indican, yet it shows points of difference from
both these, as well as from the pseudoindicans of the Acanthaceae. At tempera-
tures above 35° C., in the presence of oxygen and water, dipsacan is transformed
toa blue pigment, dipsacotin. The optimum temperature for this transformation
istoo°C. At high temperatures, or at ordinary temperatures through the action
of benzin, phenol, or dipsacase, an enzyme of this family of plants, dipsacan is
transformed to a yellow-red pigment in the entire absence of oxygen. Upon
admission of oxygen this pigment is transformed to dipsacotin.

PALLADIN™' has already urged that chromogens are universally present in
actively respiring portions of plants and that they are products of respiration.

AMMES’S results agree with this conception, for dipsacan is found to be most
abundant in the most active portions of the plants and in those plants that are in
the best condition for growth; otherwise only traces of dipsacan appear.

, TAMMEs suggests that dipsacan may be a glucoside, and that the yellow-red
pigment, which originates independent of oxygen, is one the products of the hydrol-
=; but it is not known that sugar is also a product. The formation of the
dipsacotin from the yellow-red pigment is a matter of oxidation, as PALLADIN
has shown is, the case in the production of the pigments from numerous chromo-
gens he has studied. It strikes one as possible that the formation of the chromatic
materials in general requires both hydrolysis and oxidation. This would line
Up all these chromogens with indican.—WM. CROCKER.

Germination and light.—Krnzet'? publishes another paper 0® the effect
ns light on germination of seeds, confirming the results of former papers and
adding a number of species to those favored in germination by light. .

In a discussion of ‘after-ripening” he states that the several years’ delay 1n
sermination shown by the ripe seeds of Thlas pi arvense is due to the character of
= embryo and not to the character of the coat, for the coat is very delicate. In
“arte published in 1906,"3 the reviewer has shown that the very marked delay

Portas Tames, Trxe, Dipsacan and Dipsacotin, ein neues Chromogen und ein neuer
tof der Dipsaceae. Recueil Trav. Bot. Neerland 5:—- (PP- 48). 19°.
ce PALLapIn, W., Die Verbreitung der Atmungschromogens bei den Pflanzen.
—389. 1908.
aS Kinzer, WILHELM, Aad et Einige bestatigende und < eme
erkungen zu den vorliufigen Mitteilungen von 1907 und 1908. Ber. Deutsch.
- 262:631-645. 1908.

I : : AZETTE
mm, Wa., Réle of seed coats in delayed germination. Bor. GA
» 1906.

42;

342 BOTANICAL GAZETTE [APRIL

in these seeds is entirely due to the coats. The delicacy of the coat is no criterion
of its effect, for certainly few seed coats are more delicate than that of the upper
seed of the cocklebur, yet it generally secures a delay of a year or more.

It is surprising that experimenters are so slow to see that the proper test for
dormancy of an embryo is to free it from incasing membranes with aseptic
precautions and then to subject it to germinative conditions. This treatment
will probably show the cause of most cases of delay to be in structures surrounding
the embryo. If such treatment shows real dormancy of the embryo, as in the
radicle of the hawthorn,"4 it is then ne¢essary to find the particular process that
is delinquent. This is certainly possible in the light of the great progress that
is being made in studying the catalytic nature of protoplasmic activity. When
cases of delayed germination are investigated in this way, we may hope for prog-
ress. But to assume dormancy is merely marking time and leaves the physiology
of delayed germination, as it is now, more than ten years behind other phases of
plant physiology —Wn. Crocker.

Permeability.—RUHLAND'S holds entirely untenable OveRTON’s theory of
the permeability of protoplasm, both in its original form and as m ified by

ATHANSOHN. In the main RUHLAND offers the same sort of evidence as has
Ropertson*® from the animal side. RuHLAND studied the ability of vanous
organic dyes to enter the living cell. Malachite green and thionin, both almost
insoluble in lipoids, enter the live cells readily, while rhodamin, highly soluble
in lipoids, hardly penetrates them at all. He cites a number of other dye stuffs
where just the opposite behavior occurs to that expected by the lipoid theory.
Both the acid and basic phthaleins are highly soluble in lipoids. The former
penetrate living cells readily while the latter scarcely enter at all. RUHLAND
says we have no hint of a reason for this behavior. RUHLAND and Roarer#
agree that a thin layer of lipoids often exists near the periphery of the ene oe
They believe, however, that it is not continuous in any case, but only fills ee
stices of the protein matter. ROBERTSON attributes the permeable character
the nature of the outer, very sparingly soluble, protein layer.—WM. espe

Reproduction and stimuli.—FREUND"? has done a rather elaborate aie
work on the effect of external conditions upon the asexual reproduction of Uec

. : ditions determine
gonium and Haematococcus. He finds that previous culture con ees
very largely the effect of any reagent. Of the several methods he foun Aner
ducing thi two illustrati | suffice to give an idea of the work.

a |
PVttow Will

4 CROCKER, W., Longevity of seeds. Bot. GAZETTE 47 209-7? aan

*s RUHLAND, W., Beitrige zur Kenntnis der Permeabilitat der eae:
Jahrb. Wiss. Bot. 46:1-54. 1908.

76 ROBERTSON, T. B., On the nature of the superficial layer wa

to their permeability and to the staining of tissues by dyes. Journ
I-34. 1908.

ells and its relation
Biol. Chem. 4°
die
: welt auf
EUND, Hans., Neue Versuche iiber die Wirkung der Aussen
ungeschlechtliche Fortpflanzung der Algen. Flora 99:41-100- 1900.

SE AES 8 ie en ae ee

%
2
:
,
F
q
:
:
\

1909] CURRENT LITERATURE 343

Oedogonium has grown for a considerable time in distilled water in the light,
a transfer to darkness or to a dilute nutrient solution causes a development of
mospores. Resting cells of Haematococcus, kept in darkness for some time, pro-
duce swarmspores upon being illuminated or supplied with cane or grape sugar.
Freunp finds the chemical nature of the medium rather than its physical or
osmotic character the important consideration in the asexual reproduction. In
contrast to this, Lrvincston found the osmotic character of the media the main
consideration in determing the form of Stigeoclonium.—WM. CROCKER. —

Phototropic response —Braauw,'® working with the seedling of Avena
sativa, concludes that the intensity of the light, multiplied by the least time of
exposure necessary to give a phototropic response, is approximately a constant.
The intensities used varied from 0.000439 to 26,520 Hefner candles, and the time
of exposure from 1 3 hr. too.oor sec. The product of the exposure in seconds by
the intensity in Hefner candles averages about 21 and varies from 16.9 to 26.5.
This, of course, hardly looks like a constant; but the variation is attributed to the
individual differences of the seedlings. The intensity of the light was measured
with a Weber photometer, and the observation of the response was made two
hours after the end of the exposure. The author says, “The essential condition
for the production of a phototropic curvature is the supply of a definite quantity of

iant energy; whether this quantity be supplied in a very short time or extremely
slowly, is a matter of indifference.” —WM. CROCKER.

Spraying potatoes.—A recent bulletin‘? summarizes the results of the seventh
Year's work in the ten-year series of potato-spraying experiments begun in New

, in 1902. In the ten-year experiments at Geneva, six sprayings increased
the yield 39 bushels per acre and three sprayings increased it 29.5 bushels, although
both early and late blight were wholly absent and there we but few flea beetles.
zs fourteen ‘farmers’ business experiments,” including aie iaite the average
Sain due to spraying was 18.5 bushels per acre; the average total expense of
Spraying, $4.30 per acre; and the average net profit, $3.53 Per acre. In five
ie experiments spraying was unprofitable. Eleven “volunteer experimenters
ported gains averaging 66.3 bushels per acre-—F. L. STEVENS.
d comprehensive bulletin concerning
Experiment Station. Among
nial soil conditions,

alfal alfa.—An exceedingly interesting an
the has just appeared from the New York

Subjects treated are the following: Varieties grown, unconge
a

_ » WENT, F. A. F C., On the investigations of Mr. A. H. BLAAUW on the rela-

— between intensity of light and the length of illumination in the photowor:
Tvatures in seedlings of Avena sativa. Reprint from Proc. Kon. Akad. Wetens.

terdam, Sept. 26, 1908. pp. 5-

. Stewart, F. C., Frencu, G. T., and SIRRINE,

311. January, 1 909.

York. N. Y. Agric. Exp. Sta. Bull. 305. November,

F. A., N. Y. Agric. Exp. Sta.

K., Troubles of alfalfa in
New

344 BOTANICAL GAZETTE in

winter injury, failure of the seed crop, viability of the seed, impure and adulterated
seed, fodder, yellow trefoil, weeds. Among the fungous diseases discussed are:
Leaf spot, wilt, anthracnose, root-rot and damping off, downy mildew, Ascochyta
leaf-spot, Stagnospora leaf-spot, Cercospora leaf-spot, Alternaria disease (?)
of seed, frost blisters on leaves, insect enemies, and root-knot; also, as diseases
of unknown cause, white spot, yellow top, pitting of the tap-root, and bundle
blackening in the tap-root.—F. L. STEVENS.

Barium and loco.—In a bulletin on loco weeds?! CRAWFORD says: “The
inorganic constituents, especially barium, are responsible for this action, at least
in plants collected at Hugo, Colo. Perhaps in other portions of the country other
poisonous principles may be found.” Astragalus mollissimus and Aragallus
Lamberti were most fully studied, but other species of these genera, as well as
various other genera, have been reported as producing loco.—W. CROCKER.

Protection against heating.—WrFsNER holds that the distribution of oe
tissues into small leaves or finely divided leaves is a mode of protection against
overheating by the sun, and adduces observations and experiments in support
therefor.2? This protection is secured by the small size of the parts which expose
a relatively large surface for radiation and at the same time permit many light
rays to pass them by.—C. R. B
pon balanced solu-
of K, NH,

Balanced solutions.—OstTERHOUT continues his studies u
tions with plants,?3 reporting that Na ions, by reducing the toxicity
Mg, and Ca ions, are an important, and in some marine algae an in :
protection. He finds that there is no essential difference between plants }
animals in this respect—W. CROCKER. :

2t CRAWFORD, A. C., Barium, a cause of the loco-weed disease. Bur. ee i
U.S. Dept. Agric. Bull. 129. pp. 87. 1908.
22 WiESNER, J., Versuche iiber die Warmeverhiltnisse kleiner, insbes
linear geformter, von der Sonne bestrahlter Pflanzenorgane. er. Degiats
Gesells. 26a:702-711. 1908. :
43 OsteRHour, W. J. V., Die Schutzwirkung des Natriums fir Pllanzet
Bot. 46:121-136. 1908. :

Nervousness

The use of Horsford’s Acid Phos-
phate has been found exceedingly
uable in nervous oe
restoring energy, increasi
mental and physical pee ea
and as a general tonic.

Excellent results have also fol-
lowed its use in the treatment of
headache arising from derange-
ment of the digestive organs or
of the nervous system

HORSFORD’S
Acid Phosphate.

(Non-Alcoholic. )

If your dru
ggist can’t suppl d
to » Rumrorp CHEM pply you send 25 c
MICAL W
for trial size bottle, postage Sagan RIGVaICED, R. ib

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Januar 1909 ist erschienen:

lL Dortlers Botaniker-AdreBbuch.

Dritte, neu bearbeitete und vermehrte Auflage,
/III-+-478-+-296 8S. gr.

Preis, in Ganzleinen gebunden, Osterr. Kronen 16.50 (= Mark 14.—.)
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Beigebunden - eine oe uutangreiche Botanische Bibliographie (296 8.),
Hebe alle auch nur halbwegs wichtige 1 Werke und Zeitschriften der ge esamiten
betanischen Liter fac mit ausfithrlichen un tid sonst nirgends erhiltlichen Notizen be-
Tiglich Seltenheit, Inhalt etc., der Publikationen enthalt. Diese Bibliographie bildet
‘i fir jeden Botaniker wichtiges Nachschla ihe fiir sich!
Anbei Textproben aus dem ,,Botaniker Adressbuch.“

SS
By the end of January 1909 is published:

“ . 4% 2 ” 2
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Third edition, revised and augmented,
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, bound in cloth, Kr. 16.50 (= 14 s = 3.35 $.)
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ec

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te An extensive Botanie a ‘Bibliography (296 p.) has been added. It
titles of all the more important botanical publications, including periodical, “and
er notes ~ — rarity, contents ete., and con nstitutes a valuable book

reference for botan
lone will as found specimen pages from “The Botanist’s
ry.”

__<sos——
Fin janvier 1909 a paru:

Annuaire des Botanistes de I. Dérfler.

Troisiéme édition, revue . —
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Naat ajouté au volume une Bibliographie Botanique sarah ~ e la bo ae nue
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. Te détaillées que l'on ne trouvera ee ‘Bibliographic ue

pu x “9 ete,

. botanistes une source de références a
Joint un spécimen du texte |’,,Amm e de
1 “Done er,
Ill., Barichgasse 36, Wien (A
a igien: 187, Deutsches Reich: 2129, Frankreich: 2078, a Siaad: AT
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s Botanistes.*

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The Botanist’s Directory.

An Index

to the Names and Addresses of living Botanists of |
all Countries, .

of the Botanic Gardens and Institutes, of Societies concerned
in Botany and of Periodical Publications.

Edited by I. Dorfler.
Third edition, revised and augmented.

Vienna 1909.

Published by the Editor,
Vienna (Austria), IIL, Barichgasse 36.

Annuaire des Botanistes.

Recueil

des noms et adresses des Botanistes actuellement

vivants de tous les pays, des Jardins botaniques
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Etablissements, Sociétés et. Publications périodiques
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Troisiéme édition, revue et augmentée.

Vienne, 1909.
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\amen und Adressen der lebenden Botaniker
: aller Linder,
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und der

ie Botanik pflegenden Institute, Gesellschaften und

periodischen Publikationen.

Herausgegeben von
I. Dorfler.

Dritte, neu bearbeitete und vermehrte Auflage.

Wien, 1909.
Im Selbstverlage des Herausgebers,
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Textprobe aus dem ,,Botaniker-AdreBbuch“, III. Auflage.

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1820. — Schwendener.) — Dorotheenstrabe Sie

Mbkystologisches Institut der Kgl. Friedrich- Wilh ia ilniver-
sitit und Botanisches Institut der Kgl. EN ee Hoch-
Schule. (Dir.: L. Kny.) — invalidenstente 42,
- Museum der Kgl. reba pion rpc Hochse hule. " (Vorstand der vege-
__ fabilischen Abteilung: L. Wittmack.) — Invalidenstrabe 42, N. 4.
Institut fiir Landwirtschaftliches Versuchswesen und Bakteriologie aot
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} InvalidenstraBe 42,
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Pal Bere tisches Institut der Kgl. Bergakademie. (Vorsta tand:

: H. Potonié.) —— InvalidenstraBe 44, N. 4.
oe ae Biologische cae fiir _— und Forstwirtschaft. (Errichtet
— Dir.: ns, — Pub ,Arbeiten®. Mitteilungen". »lug-

bi : hlem lin
a cee Institut der Kgl. Qurtnerlehranstalt. (Vorstand:
9: Hostermann.) — Dahlem-Steglitz bei Berlin.
isches Spe sopettarlg etry (Gegr.: 1874. — Kustos: R. Buchholz.) —
Am Beioben Park, 8.0 eee aut bolo
r Girungs ewerbe Dir Delbriick. ors e .
Say Abt.: P. 88 dner, eins ML De der sabes Abt.: W. Henne-

al}

Veseisanstat fiir Getreideverarbeitung. (Leiter der pede ger
Abt, des reg ne es u. d. Versuchsmiihle: J. Bu wald.) —

Mats Bo Botanische ‘Abteilung fiir Wissenschaftliche und Sehulewoeks.
si ig. — a Institut liefert Pflanzen zur Besprechung tur
fatiche Schulen Berlins.) — Humboldhain, N. 31.

‘4b “sen der Wissenschaften (Gegr. 1700. — Pub
Berli, lungen*.) — PotsdamerstraBbe 120,

Berliner Gerdlischoft fiir Geschichte der Naturissenschafton und Medizin.

,Sitzwngsberichte®.

a — Publ.: ,,Verhandiungen* in 4 » Mitteilungen”) ag
Snathe Verein der Proving Brandenburg (Geer. t 1859. Pu Be ce
Deut sh lungen“,) — Kgl. Botanisches Museum, Datem-Stegti bei Ber

€ Botanische Gesellschaft. (Ge 1882. — Publ
- Deutse gr. :
tthe Dendrologische Gesellschaft. (Gegr.: 1892. — Publ.:

1907. — Publ: »Orchis“.)
(Geg Publ.: " ssgeaptel fiir
oS cae Rothenbargateali ms " geeiliia bei Berlin

a,
Mitteilungen. )

e i Berlin.
aft fiir Orchideenkunde. (Gegr.:

eth ‘TIT pee hae wep sue oqoudyxoy,

Spécimen du texte de la 3e édition de l',,Annuaire des Botanistes“.

135 France. (Europa) —

Viguier, René, Dr. és sci. nat., Préparateur de botanique au Muséum d'Histoire
Naturelle de Paris, 5 bis, quai de Bercy, Charenton, Seine. (Anatomie
Systématique. Paléontologie aisate :

Vilmorin*, Maurice-Lévéque de, 13, quai d’Orsay, Paris, VII. (Collections
botaniques d’arbustes. Plantes ae Chine - et Thabet

Vilmorin*, Philippe-Lévéque de, Lic. és sci. nat., — 23, quai d’Orsay, Paris, VIL.

(Phanérogames. Génétique.

Vincent, Philibert, Eléve en pharm., — Rue Bourg-Belais, Parthenay, Den-

Sévres.

Vineon, Etudiant, — 16, rue Jean-Bouchet, Poitiers, Vienne.

Violleau, abbé Eugene, Curé- na Ge — St.-Varent, Deux-Sévres. (Phanérogames.

— Collect. a nges

Viron, Dr. méd., Pharmacien en chef de l’Hospice de la Salpétriére, — 47,
‘boulevard ae Hopital, ‘Paris : a

Vitou*, Et., — 1, rue pane Caizergnes, Montpellier, Hérault. (Cécidies. Maladies
physiologiques des végétaux.)

Viviand-Morel*, Jose oh: Wie ctor, Rédacteur en chef du »Lyon-Horticole*, — 38,
cours Lafa, ayette ee Villeurbanne, Rhone. (Botanique systématique.

lec

Phanérogames. Collect. de eee gene
Voirin, Gautges, Dr. en méd.. e Rousseau, Bar-le-Due, Meuse. (Colles)
Vouaux, abbé, Professeur au Cue ae ak Malgrange, — Jarville, oie] Nancy,
Meurthe- oc ose re “f
Vuillem ary eae Whistoire naturelle médicale &
Faculté "de Médecine ce Netty, at e d’Amance, Malzéville, Meurthe-tt-
Moselle. (Anatomie. Biologie. Pps avs Pathologie. D ératologie.)
Vuillermoz, Pharmacien, — Lons-le “Saunier, Jura. (Mycologie.
Wattiez, R.-P., Professeur, — Cellule, par Riom, Puy-de-Dome po
Weiller, sory "Lieutenant au 21e Régiment d’ Artillerie, — Rue de la Font '
A éme, Charente.

ng
Wuitner, ‘E., oe oo rue Saiegreniss Levallois-Perret, Seine. (Algues)
‘, phar éres-de-Bigorre, Hautes-Pyrentes chanst
Zeiller®, Charles-René isk de CInstitut, Inspecteur ensral des Mines, Car
mi lecons de paléontologie végétale a ’Ecole Nationale Supérieure des
rue du Vieux-Colombier, Paris, VI. (Paléobotanique e.)

ABBEVILLE, Som 3 erthes;
Musées [Musée ois unal; Place St.-Pierre. — Musee Boucher de P ;
Rue B.-de-P.]. (Conservateur: A. Ledieu.) : ‘iets (onde 4
Société puaeen pour Encowragement des Lettres, Sciences @ .
en bl: ,Mémoires*. ,,Bulletin*.) ]
AGEN, Lot- i. oe ronne, Publ: Recwel
Société d’ Agriculture, Sciences et Arts. (Fondée en 1776. — 3
atGOU LT i. s Travaux. ,,Revue de V Agenais*.) .
ont-|, Gard (et Lozér -Fajeo!®

Jardin Pcie = oe — 1) Jardin du P ey ‘O
1300 m — 2) Jardin & la Moliére-du-Tréve ezel. ooh zaet

ae ri aes de-Dieu (Fondés en 1903, — Dir.

mbe

mie a Seine, adeabeasela Arts et Belles- Lettres.
Publ. : ,Mémoires“

AJACCIO, Cor rse. : h
x Musées. (Conservateur: J. F. Peraldi.) — Collége Feseh.

Maske d’Albi. — au Pare de Rochegude. ; 1877.)
aie des hag Arts et Belles-Lettres. (Fondée en
N, Orn

Musée Scientifique, Archéologique et righ Naturelle. (
. Richard et L. Brioux.) — Hot e Ville.

Conservatea™ 4

Great Britain and Ireland. 169

ler*, George, — Levens, Milnthorpe, bihees sien E. (ites: ort ee )
. William, — 93, Upper Brooke Street, Manchester, E. (Moss

eld, Frederic Wilson, M. D., | hyaleiee and Surgeon, — 120, Oxford Road,

Reading, Berks, be eae y. Phanerogams. Ferns.)

tied Ph. D., F. L. S., F. R. H. S., Corr. M. Deutsche Bot. Gesellsch.,

Secretary of the Sonean’ Society, Keeper of the Herbarium and Library, Royal

Botanic Gardens, Kew, — val Botanic Gardens, Kew, near London

(Gramineae. Taiconomy. Phytogeogra yphy.)

Sedman, C. F., — 76, High Street, Ashford, Kent, E. deioneeshices )

Meele*, A. B., — 16 me “London Gack Edinburgh, 8. of Te

Set, Charles, — Savings Bank Department, G. ee C. (Fungi.)
Edward, F. L. S. — Oakwood House, peti Wood Lane, Ashtead,

, E. ghee Ferns. Fungi.
Sevens, Mrs, En = ott. Worcester bik; Surrey, E. sisite of Surrey.)
ew Int n Entry, Dundee , 5. ‘ae

~ 10, Avenue Road, Do neaster, York, E. (Micro-botany.)

t se Se ieton, near Croydon, Surrey, E. VMiovo-A e.)

» Sir James, M. A., LL. D., — Fincheocks, Goudhurst, Kent, E. (Mosses.)

ee, M. D., F. L. S., — iy Newton Terrace, Glasgow, 8. (Mosses )

enry, M.D., M. A, — 1, Grey Friars, Chester, EB. (Diatoms.

ange: Microscopical slides or material a

Benjamin, Kt.. J. P., F. L. S., F. G. S, F. BR. GS, — The Grange,

Elston Warwick, E. _ (Geographical botany. )
erbert, F. L. S., idge Street, Birmingham, | Warwick, E.

(Fytate physiology. For ere oDeatr es to exchange specimens of wood or
purchase.)

Stopes*, “Miss Marie €., D. Se. Lond., Ph. D. Munich, University Lecturer on
: Sees speasical Dep’t., The University, Manchester, E. (Anatomy of
ant

ussex, England. e sy
Detourne Nat ral Histor y Society. (Founded 1867. — Publ. ; “T'ransactions’’.)
— Technical Tnstitu

“ona Be. (Founded se _ pir: J. J. Dobbie. — Keeper
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aM tsk Avbor (catbrad Be siaty ="(inetituted 1954. — Publ: “Trans-
= “etal of j Bainburgh thouaiel 1836. — Publ.: “Transactions and
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erocop ical. Society (Publ.: “Proceedings”.) — Philosophical Insti-

Str srg
~ “ae 1 1881. — Publ.: “Transactions .)
5, Queen Stre vy Society y. (Foundec

K10VDOAT, SISTUVJOY OBT,, JO WOMIPS pag OG} WOIZ pS

.
4

Saggio della terza edizione dell’ ,,Indicatore dei Botanici“.

185 Italia.

Brizi*, Ugo, Dott., Professore di Fisiologia e Patologia vegetale nella

e
Via Marsala, 8, Milano. (Iisiologia. Bobi: ia vegetale,
geueora eaten Diana, Dott. in Sci. nat., — Via del Lavatore, 32,
(Fisi
Bruttini, Pes, Libero Docente di Agronomia, — R. Stazione Agraria, |
Susanna, Roma.
Buscalioni*, Luigi, Dott., Professore ord. di pia Direttore del R. |
Botanico dell’ Universita, — Catania, Sicilia. (Istologia
eademali. Giovanni Battista, Professore di Storia Natarale, — R.
Caldarera, Ignazio, Dott. in Sci., Professore, — R. Liceo ,V.Eman
Sicilia. (Anatomia vegetale

* eo
Cattaneo“, — Via 8. Vittore, 47, Milano. (Biologia
Calestani*, Vittorio, Dott. in Sei., Professore, — Via Manente, 95, /
Perugia. (Sistematica. Fanerogame.)
degen Mario, Dott in Sci. agr., aed Se ~~ Coe Ambulante
— Porto Maurizio, Liguria. (Fanerogam :
Campbell®, Carlo, fopete Direttore della Cattedra ‘hiabiaaie d'Agi

a)
Camperio*, Camillo, Ingegnere, — Corso di Porta Vittoria, 12, Milano. (Si
Cannarella*, Pietro, Dott., anata “ti Scienze fisiche e e naturali nella. R.
Normale Femminile, — Via Carrettieri, 8, Palermo, Sicilia. (angie
Uemkor, Giovanni Battista, Gia Vice-Direttore del R. Orto Botanico
orzoli presso Genova
Capeder, Giuseppe, Dott., Professor re, — R. Liceo, Voghera, Pavia. (Paleo
Capra*, Sac. Giuseppe, Dott. in Sci., Professore di Scienze naturali €
tura, — Via Copernico, 9, Milano. | (Sistematica. Briologia. 2
Carano*, Enrico, Dott. in Sci. nat., Primo —— gestern: Botanio
. Universita, — Via Panisperna, 89 B, ma. (Anatomia. +
Carestia, Antonio, Abate, — Valle Vosonen Mae ive ees
. Secu :

u
Caruso*, Salvatore, Dott., Professore di Storia aired nel R. Liceo, ~
Sicilia. (Sistematica: Licheni i.)

NAPO :
ae ed SDieta Botanico della R. Universita. (Dirett.: *
Sinkione Zoologica, _ Dirett. : A. Dohrn. — Pubbl.: »Fauna und
Golfes von Neapel* Pp. Sev
sma ene Sree del Prof. P. Severino. (Dirett.:

Sett i 80. , Rendiconte
Societa Reale é i Napoli. (Fond. nel 1808. — iche eT
R. Istituto @ Iacoraggiamento alle Scienze Natur “ali, "Reonom

S06: = -P

ond. nel ubbl.: ,,Atti*.) :
Societa dei Naturalisti. (Pubbl.: ,,Bollettino*.) — Ex-Monastere ces
PADOVA, Veneto. pirett.: P. A. Sacei
R. Orto ed Istituto Botanico dell’ Université’). (Dire a are os
rto Agrario della R. Universita. (Dirett.: L. Di Me Ati
R. Accademia di Scienze, Lettere e Belle Arti. “Bubb t

»Rivista *.) av
Accademia Scientifica Veneto-Trentino-Istriana. (Gis Socks Atti* ie
i Scienze Naturali, fondata nel 1872. — Pubbl.:
gico della R. Universita.
Fy *) L’Orto Botanico di Padova fu fondato dalla Repubblica Ver eto “— +
el mondo,

THE
BoTANICAL GAZETTE

May 1909

Editors: JOHN M. COULTER and CHARLES R. BARNES

CONTENTS

T
he Megasporophyll of Saxegothaea and Microcachrys
Robert Boyd Thomson

St
Studies on the Oxidizing Powers of Roots
Oswald Schreiner and Howard S. Reed

Bo: 7 .
g robin and Their Effect upon Soils Alfred Dachnowskt
Briefer Articles
Parthenogenesis in Pinus egeey
Carnation Alternariose

. Saxton
F. L. Stevens ee G. Hail

Current Literature

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botanical staff of the University of Chi

Issued May 23, 1909

CONTENTS FOR MAY 1909 No. 5

KxV). Robert Boyd Thomson - 345
ON THE OXIDIZING POWERS OF ROOTS. Oswald Schreiner and Howard S.

- 355
INS AND THEIR EFFECT UPON SOILS (wITH Two FIGURES). Alfred Dachnowski 389
R ARTICLES
THENOGENESIS IN Pinus pinaster (WITH SEVEN FiGuRES). W. 7. Saxton - ~~ 406
NATION ALTERNARIOSE (WITH EIGHT FIGURES s). F. L. Stevens and 7. G. Hall - - 49
LITERATURE
REVIEWS - 3 Z : < : s e = - 414
ANOTHER MUSHROOM BOOK. TREES AND WOODS. MICROSCOPY OF TECHNICAL
oDucTS WORKS OF LEO ERRERA
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VOLUME XLVIL NUMBER 5

BOTANICAL GAZETTE

MAY 1909

THE MEGASPOROPHYLL OF SAXEGOTHAEA AND
MICROCACHRYS
RoBERT Boyp THOMSON
(WITH PLATES XXII-XXV)

In the Coniferae great importance attaches to “inverse orientation”
asa criterion for determining the character of the fertile scale in the
megasporangiate cone. ‘The followers of BRAUN see in the inversion
of the ovuliferous scale bundles an indication of the “brachyblast”
character of this structure. SacHs and ErcHLer, the exponents of
the “ligular” theory, look upon the inversion as 4 feature which
characterizes the vascularization of a ligule or of an appendage of the
simple sporophyll. CELaKovsKy partly combines the two views
and seems to homologize many of the diversified megasporangiate
features of the gymnosperms. In the present study some neglected
phases of the inversion of the sporangial supply bundles of both the
Staminate and ovulate cones are given prominence, and data are
advanced to show the homology of the micro- and megasporophyll in
Saxegothaea and Microcachrys.

The gross features of a fruiting branch of Saxegothaea are indi-
tated in fig. 1. The megasporangiate cones are borne terminally on
Drneties (figs. 1, 2), the microsporangiate ordinarily in the axils
a foliage leaves, but occasionally in a terminal position (7ig- 3): The

Sasporangiate cone, at the stage indicated in fig. 2, has a short pedi-
eel beset with the bracts which earlier inclosed it. Later the pedicel
Si and a lax arrangement of the bracts is evident (Ag. 4).

ter the pedicel becomes much more elongated and relatively
el slender. The bracts, sporophylls, and foliage leaves are spirally
“anged, and gradations in form are evident, the bracts gradating

345

346 BOTANICAL GAZETTE [way

apically into sporophylls and basally into vegetative leaves (fig. 4).
The microsporangiate cones may be either sessile or pedicellate. In
either case there is a series of spirally arranged bracts closely investing
the base of the cone, the lower part of the pedicel, when present, being
naked (jigs. 2, 3). In an otherwise abnormal cone I found the arrange-
ment of bracts similar to that on the megasporangiate pedicel.

The gross features of Microcachrys are indicated in a figure of a
previous article." The cones are always terminal in this form and
their sporophylls verticillate, in series of fours, the series alternating
with one another and presenting from the exterior an appearance
of a spiral arrangement (figs. 5, 6). The small concrescent foliage
leaves are opposite, in alternating pairs.

The form and structure of the microsporophyll of Saxegothaea
are indicated in figs. 7-9. Two are shown cut longitudinally
(fig. 7), and between these is one cut through the sporangium,
showing the stomium and the wall with its columnar, comparatively
thick-walled, epidermal cells. The inner layers of the wall have
collapsed, but there are indications of three or four of these.’
In the axial sections the vascular bundle with its accompanyims
resin duct is seen, and the connection of these with those of the axis
of the cone. The resin canal is expanded distally. This is apparent
in the tangential section of the cone as well (fig. 9), where the cent!
sporangia often show little trace of a canal, while in the lateral ones It
is large. In fig. 8 resin canals and vascular bundles are seen s
transverse sections of the cone, at various distances from the oe
supply. In some of the sporangia, to the right of the figure, 1
position of the stomia is indicated. ‘The microspores conli®
three cells at the stage from which these figures — ” ie

‘ fferentiat
Megasporangiate cones of the same date have not di a
the megaspore (figs. 10, 11), with the exception of some ee ;
ovules, to be described later. A. single bundle accompanied oa
resin canal passes into each sporophyll, and this giv ” ed
ovular supply just as the point of insertion of the integument Is rea

EB 47:20-2-
‘Tomson, R. B., On the pollen of Microcachrys. BOT. GazeTt
pls. 1, 2. 1909. See pl. 1.

Norén, C. O., Zur Kenntnis der Entwicklung von Saxe

2Cf gothaea conspiew?
Lindl. Svensk. Bot. Tidskr. 2: 101-122. pls. 7-9. 1908.

1909] THOMSON—SAXEGOTHAEA AND MICROCACHRYS 347

(fg. 11). Fig. 12 is a transverse section of the sporophyll a little
nearer the cone axis than the point indicated. It shows a single
vascular bundle, and below it the accompanying resin duct. The
wood (lighter in the figure) is roughly triangular in outline. Near its
center are two very darkly nucleate cells. These mark the separation
of the wood of the ovular supply from that of the main bundle. The
bast above this is but slightly differentiated, and nearer to the axis of
the cone dies out completely, as does the wood itself in its further
course through the cortex. Under the base of the ovule the bast is
quite apparent on the upper side of the supply bundle (fig. 17), which
shows a tendency to bifurcate, the branches passing laterally into the
base of the integument. Beyond the separation of the ovular supply,
the main bundle gradually acquires a new set of centripetal wood
elements, the more central ones of which are elongated, the lateral
quite typical transfusion tissue.

In material of Saxegothaea some six months older than the former,
the vascular system of the megasporophyll is further developed. It
has been recently described by Miss STILES,? with whose account my
own observations are practically in accord. The upper vascular sys-
tem may be composed, in the region of the ovule, of as many as four
stands. These, however, in all the material I examined, unite into
‘wo before passing upward into the base of the ovule. Miss STILES
speaks of there being about three bundles in this region. The pair
terminates at the level of the base of the nucellus in a considerable
‘pansion of transfusion-like tissue. The main bundle of the
‘porophyll is replaced by a series somewhat similar in the same
"gion in which the proliferation of the ovular set occurs. Above the
wule a single bundle is found, with centripetal, and on its flanks a
“onsiderable, development of xylem elements. The vascular supply
for the scale cones from the axis is a single bundle with normally
‘nented wood and bast. In the cortex gradually new xylem elements
és formed Opposite the protoxylem. Farther out wood and bast
‘oming off the sides of the main bundle supplement the ori
me elements, which up to this point appear not to have any bast 2

itown. In the further course the proliferation of the upper 4
New Phytol. 7:209-

: . .
722, ee W., The anatomy of Saxegothaea cons prcua Lindl.

348 BOTANICAL GAZETTE [utay

lower series occurs, as described above. All of the upper series of
bundles pass into the base of the ovule, the only xylem elements
found beyond the ovule in this region being a new development asso-
ciated with the main bundle. A single resin duct accompanies the
latter throughout its course. This does not branch and is the only
representative of the tissue in the scale, no canals being formed in
connection with the upper series of bundles.

The vascular system of the axis of the megasporangiate cone con-
sists of a ring of collateral bundles very similar in general to that of
the microsporangiate cone (fig. 8). The wood of these is usually of
the ordinary endarch type, but near the base of the cone there are
often a few centripetal elements associated with the bundles at the
sides of the gap left by the exit of the megasporophyll trace. In one
instance these were almost in continuity with the wood elements of the
ovular supply in the sporophyll. The’ latter, especially in its young
condition, appears very much of the nature of centripetal xylem
(fig. 12),5 and the occurrence of the isolated elements in the axis
affords confirmation of this idea. They are found also in the upper
part of the pedicel, but in my material they are rare here.

_ The scales at the base of the micro- and megasporangiate cones and
the foliage leaves both receive a single vascular bundle and a single
resin duct fromthe axis, the supply coming off in a similar way ' that
of the sporophylls. There is always a gap in the cylindrical _
opposite the “trace” to each of these members (fig. 8; for microspore
phylls, lower right hand side, etc.). F

In several of the megasporangiate cones of Saxegothaea 4 few 0
the lower sporophylls bear ovules on their under surface (fis: 10).
These have an integument, but not the epimatium or ee
second integument of the normal ovule. They stand out too fro
the sporophyll more freely, and are further developed than the pede
Ones in the same cone. A slight vascular supply goes off gt oar
these, the bast and wood showing a tendency to orient itself invers
to that of the main bundle of the scale. This supply passe sent
the resin duct. :

4 Cf. Stirs, |. c. 216,

5 Miss Stivers has'come to a similar conclusion from studying api inet a
condition.

Phe Pe ea PE ge ete eee, ee

ee ee
7 eS EE Re Ee een aE D eee

1909] THOMSON—SAXEGOTHAEA AND MICROCACHRYS 349

In Microcachrys a single vascular bundle supplies leaf and sporo-
phyll. There are no resin canals in the stem nor in the axis of either
cone. It is not until well out into the sporophyll that a canal is found
beneath the vascular bundle (megasporophyll, fig. 13). The canal of
the vegetative leaf, however, is continued downward into its swollen
base, and the leaves being concrescent, there is the appearance of
resin ducts in the stem. The vascular bundle in both leaf and
sporophyll runs closer to the upper than the lower surface. A com-
parison of longitudinal sections of the megasporophyll and vegetative
leaf shows a great resemblance between the distal part of the former
and the whole of the latter. ‘There is a distinct palisade on the lower
surface of each, a sclerotic hypoderma, and an epidermis with a very
thick cuticle and no stomata, the latter being restricted to the upper
surface, and in the case of the sporophyll, so far as I have observed,
to the region beyond the ovule. There is no hypoderma on the upper
surface, and the same kind of degenerate-looking palisade tissue
occurs in: both sporophyll: and leaf. Proximally the sporophyll is
contracted into a slender stalk, while the leaf is concrescent with the
stem. The vascular supply of the ovule originates in much the same
way as in Saxegothaea with at first no apparent bast® (fig. 14; above
the resin canal come bast and wood of the main bundle and above the
center of the latter the few wood elements of the ovular supply).
On separation the ovular supply bundle passes obliquely upward,
as in Saxegothaea (cf. figs. 11 and 13), with bast clearly evident on
its upper side. In Microcachrys, the single supply bundle bifurcates
near the base of the ovule, the divisions passing well into the basal
Part of the integument. |
In most of my material the megaspore ©
few nuclei in a parietal stratum of protoplasm
however, a prothallial tissue (fig. 15) is presen
Initials discernible. These are located at considerable distances from
oneanother. Around the prothallium the megaspore-coat is of the usual
double type.’ It is slightly thicker than in Pinus resinosa at a similar
‘lage. Like the pine coat at this young stag® +t is of uniform dis-

SCE. WorspeELt, Observations on the vascular system of t! female flowers of the
a Annals of Botany 13:538. 1899. 5
Toronto Studies, Biol. Ser. ie Gs poyergr oe :

f the ovules has only a
(fig. 13)- Tn one case,
t with the archegonial

he gymnosperms. Univ. .

350 BOTANICAL GAZETTE [May

tribution about the prothallium. It would be interesting to know the
condition in the mature seed, especially in view of the fact that in
certain species of Dacrydium the megaspore coat is very thick at a
later stage.

The integuments of the ovule are seen in their usual form in fig. 13.
Sometimes the inner one is quite open above the ovule (fig. 15), even
at an advanced stage. In the case figured it is developed laterally
into two large masses, appearing as additional ovules in gross material.
In other cases it may be uniformly quite thick (fig. 16). The two
outer cell layers of the integument are tangentially elongated in the
older condition, while the third consists of columnar cells radially
placed, many of which in the basal region become lignified and
have numerous large pits. They are in connection with the supply
bundles of the ovule through transfusion-like extensions of the latter.

Fig. 17 is of a longitudinal section of the bract and scale from the
megasporangiate cone of T'suga canadensis. The section passes to on€
side of the axis in the region of the ovule. To the lower right-hand
side of the figure the bract is seen fused proximally with the tissues at
the base of the scale. .The latter shows one large vascular bundle,
cut nearly transversely. This has the wood below and the bast above,
an arrangement the inverse of that in the bract. From this bundle IS
given off the ovular supply, which passes obliquely upward into the
base of the sporangium. The character of the supply bundle was
studied in series of transverse sections. It comes off from one ae
lateral bundles of the scale ( fig. 15; second bundle from the right
and left of the figure). Its wood and bast lie at first in a plane .
right angles to that of the scale bundles (fig. 19; the wood is the _
celled tissue in the center of the figure and the bast is to i left).
Shortly the bundle turns completely round, so that its wood lies a
and its bast below, the inverse arrangement to that of the scale bundles,
which, in turn, is inverse to that of the bract. Near the ovule the
bundle becomes concentric and finally bifurcates, the branches enter
ing the base of the integument.

In the microsporophyll of the cycads Tu1BouT® has call
tion to the inverse orientation of the sporangial supply :

ed atten-
I have
pp. 265. PS

* Tursour, E., Recherches sur l'appareil male des gymnospermes.
16. Lille. 1896.

'
:
!
/

Be hs

Peewee age eee

Sy Se ee a ee en ee

1909] THOMSON—SAXEGOTHAEA AND MICROCACHRYS 351

figured that of Ceratozamia mexicana (jig. 21). In fig. 21 the upper
bundle is one of the normally oriented main bundles—bast below
and wood above, not shown in the section—while cutting obliquely
across the field is a sporangial supply bundle with bast above and
wood below. A short ventral extension to the sporangia turns at
tight angles at first and then becomes almost inverse. Figs. 22 and
23 are transverse sections of a sporophyll. In the upper part of the
figures are the main bundles, with wood above and bast below.
Farther down and toward the ends of the normal series are some
inversely oriented bundles. One, to the left, magnified in fig. 23,
has just divided into a similarly oriented bundle, and from this
there is a strand to the sporangium. The latter has a more or less
concentric character. In some cases I: have found a second inversion
on the way to the sporangia.

The inversion of microsporangial supply bundles is of wide occur-
rence in the cycads, but is not confined to the group, being also
found in certain conifers. This feature has not been considered
by the exponents of the theories advanced to explain the inversion
in the fertile scale of the latter. hat it is of first importance seems
self-evident. Again, the double inversion in the fertile scale has not
been explained, and a glance at the figures shows how important the
second inversion is, both of itself and in comparison with the single
inversion in either Saxegothaea or Microcachrys (cf. figs. 17, 73: and
17). This and the occurrence of ovules on the lower surface of the
scale in Saxegothaea, and their vascularization from the main supply,
Would seem to be good reason to consider that inverse orientation
Cannot in all cases be relied upon as evidence of the brachyblast
character of the fertile scale. In the last case, We should have the
‘nomaly of the leaf in the axil of the shoot.

Again, in Dacrydium two bundles with or
of the scale supply the ovule. These attach directly to the axis, the
three passing close together near the upper surface of ae scale.

fre is, then, in Dacrydium, a form admittedly of a higher a
aa Saxegothaea or Microcachrys because of the Oe cee
- “one, etc., a greater amount of inversion than ” ‘
Ptimitive forms—a puzzling feature certainly 1 view of the
brachyblast theory, but possibly finding its explanation 1n the greater

jentation inverse to that

352 BOTANICAL. GAZETTE [May

importance of the ovule in this specialized fructification and in the
greater advantage which a direct connection with the axial supply
affords. This has its beginning in Saxegothaea and Microcachrys,
where the two bundles of the ovular supply have asserted their indi-
viduality only part way to the axis, the ovular supply in these forms
having possibly originated, as has been suggested, as a revival of the
centripetal development of the main bundle.

In view of the facts presented, the writer is led to look upon the
fertile scale as a simple structure, the homologue of the microsporo-
phyll-in both Saxegothaea and Microcachrys, and, ipso facto, in the
whole of the Taxaceae, if these form a natural alliance, as is commonly
held. With this group is to be associated the Araucarieae, whose
microsporophylls show an inversion of the sporangial supply bundles
of like character to that of the megasporophyll,? and which, on
other grounds, have been considered’° as having a simple mega
sporophyll. This group also has recently been shown to have many
features in common with the lower podocarps. Perhaps the most
important of these are the method of pollination and the absence
of wings on the grains themselves in Saxegothaea and the Araucaricae,
the extensive microgametophyte of the two groups; the single inverted
ovule, the wide micropyle and projecting nucellus, the nucellus ee
from the integument to its base—features which bring these forms into
intimate association.

The simple-scale groups, the Taxaceae and Araucarieae, have =
their micro- and megasporangia on opposite sides of the sporophyl
and since the homosporous forms, from which the conifers were
derived, the pteropsid series, as is shown on morphological and ani
tomical grounds, have the sporangia usually on the lower a
the ovule has probably been transferred to the upper surface int
course of phylogeny. The ovules on the lower surface of some pe
of Saxegothaea and their lateral position in Cycas are significant =
this connection, and, as well, the lateral development of the spora®
gium in Schizaeaceae. .

. ‘Society: New
9 THomson, R. B., The origin of the gymnosperms at the Linnean :
Phytol. 5:145. 1906. oe

1° Sewarp, A. C., AND Forp, S. O., The Araucarieae, recent and extinct
Trans. Roy. Soc. London B. 198: 305-411. pls. 23, 24- 1906. ,

Phil.

909] | THOMSON—SAXEGOTHAEA AND MICROCACHRYS — 353

CovLTeR and CHAMBERLAIN, after reviewing the different theories
of the fertile scale in their textbook on the gymnosperms, make the
following statement (p. 77): “Upon sifting the testimony certain
things seem to be fairly clear, and one is that the scale and its ovules
in Abieteae represent a highly modified axillary shoot, corresponding
to the characteristic spur shoot of the group.” From anatomical and
teratological evidence this conclusion seems inevitable, and since the
Abieteae, Taxodieae, and Cupresseae present such a series of natural
affinities the statement must apply to all. If, then, the fertile scale
in this group is of the brachyblast type, representing the adaxially fused
first and only two leaves of an abortive axis, the first inversion is
explained, and the ovules in this group are borne on the morphologi-
ally under surface. The second inversion is analagous to the single
one in Saxegothaea and of the nature of a sporangial supply. There
are, then, two great groups of the conifers from the standpoint of
this study, the simple- and the complex-scale series. Both forms
have the ovules on the physiologically upper surface, a position ren-
dered almost imperative by the necessities of the seed habit. This
position, however, has been attained in two very different, but pos-
sibly equally difficult, ways in the aplo- and diplosporophyllous
forms of the Coniferae.

Toronto UNIVERSITY

EXPLANATION OF PLATES XXII-XXV
PLATE XXII
Fic. 1.—Fruiting twig of Saxegothaea conspicua. Slightly enlarged.
Fics. 2, 3.—Micro- and megasporangiate cones. X4-

PLATE XXIII
Fic. 4—Megasporangiate cone with the axis more €
tay 5.—Microcachrys tetragona; micro- and megas
mature condition. x4.
Fic. 6.—Older megasporangiate cone.
_* 7-9.—Respectively radial, transverse,
Porangiate cone of Saxegothaea.
©, 11.—Megasporophylls of Saxegothaea,
ovular supply coming off the main vascular bundle.

Jongated than in ig. 2.
porangiate cones, the

and tangential sections of the

me with ovules attached; 7ig-

PLATE XXIV
12.—Transverse section of the megasporophyll of Saxegothaea, youn’

Fic,
Condition : :
dition, just proximal to the insertion of the ovule.

354 ~ BOTANICAL GAZETTE [aay

Fic. 13.—Longitudinal section of the distal part of the megasporophyll of
Microcachrys.
Fic. 14.—Transverse section of same, showing ovular and main supply
ee in transverse section
C215 ieurtipiuin of Microcachrys in t
Fic 16.—Sporophyll and megasporangium with "abacciiale thick integu-
ment, in transverse section.
Fic. 17.—Tsuga canadensis; bract, scale, and ovule in longitudinal section.

PLATE XXV

Fic. 18.—Tsuga canadensis; transverse section of bract, scale, and ovules at
the point of origin of the ovular bundles.

Fic. 19.—The same; a magnification of the ovular supply of the right ovule
in fig. 18

Fic, 20.—The same; supply bundle farther up.

Fics. 21, 22. Eis eclively, longitudinal and transverse sections of micro-
sporophyll of Ceratozamia mexicana.

Fic. 23.—A magnification of a part of fig. 22.

PLATE XXII

NWIANICAL GAZETTE, XLVII

THOMSON on MEGASPOROPHYLL

PLATE XXIII

WMNICAL GAZETTE, XLVII

- 28a Ny
* ——

ede td

THOMSON on MEGASPOROPHYLL

XXIV

PLATE

BOTANICAL GAZETTE, XLVII

e
£

‘ees.

THOMSON on MEGASPOROPHYLL

BOTANICAL GAZETTE, XLVII PLATE XXV

‘4d ..23

THOMSON on MEGASPOROPHYLL

STUDIES ON THE OXIDIZING POWER OF ROOTS!
OSWALD SCHREINER AND HOWARD S. REED

The present paper embodies a series of studies upon the oxidizing
powers of plant roots grown in aqueous extracts of soils and in
solutions of various compounds. The results, it is believed, throw
some light on the action of plants upon the soil and iadicate how
soil conditions affect certain functions of the plant.

The experiments which are presented show that plant roots are
able to carry on active extra-cellular oxidation, chiefly by means of
the enzymes which they secrete. From the standpoint of root excre-
tions the study is of interest because it has often been supposed that
the roots of growing plants excrete organic and inorganic acids which
aid in the solution of soil minerals. The idea undoubtedly owes its
Prevalence largely to the experiments of Lrepic* and of SACHS®
Which demonstrated the corrosion of polished marble plates by grow-
ing plants. The more recent investigations upon the subject made
y CzaprK,4 Kossowircu,’ and Kunze® have shown, however, that
very little acid is excreted by the roots of the higher plants, and that
the results of the earlier workers were mainly due to the action of
catbon dioxid.,

This oxidizing power of plants gives them an important action
“pon the soil. Whether they have the power to oxidize the inorganic
‘oustituents of the soil remains to be determined; but it has been
“own that they are able to oxidize organic substances, such as the
chromogens employed in these experiments, at 4 fairly rapid rate.
Tf these substances are oxidized, it is only logical to conclude that the
rganic substances occurring in the soils are also oxidized by the
‘ction of plant roots.

This oxidation of organic compounds is of additional interest in
the light of recent investigations, which show that the cause of unpro-

* Published by permission of the Secretary of Agriculture.
2
Annalen Chem. Pharm. 105:139. 1858.
*Bot. Zeit. 18:117. 1860, s Ann. Sci. Agron. II. 8:220. 1993:

‘Jahrb. Wiss. Bot. 29:321. 1896. "6 Jahrb. Wiss. Bot. 42°357- 1906.
355] | Botanical Gazette, vol. 47

356 BOTANICAL GAZETTE [May

ductivity in certain soils is due to the presence of toxic organic com-
pounds.?’ The beneficial effect of oxidation in such soils may be
inferred from the results of thorough tillage, involving sub-drainage
and cultivation, since these operations promote aeration of the soil
with subsequent increase in growth of roots and microorganisms.
Under such conditions experience has shown that the organic sub-
stances in the soil are most completely converted into substances
commonly known as humus. It is certain that the oxidizing activ-
ities of the soil and plant play a significant part in this important
process. ;
, PREVIOUS WORK

The existence of an oxidation process in soils has long been
known. Without apparently understanding the precise cause of
this phenomenon, LiEeBic pointed out its importance for productive
soils, and, according to the same author, the phenomenon had been
earlier investigated by InGeNHoUss and DE SAUSSURE. Among
the more modern investigations on the processes of oxidation In
soils the works of DEHERAIN and Demovussy,® WOLLNY,? RUSSELL,'°
and Dargisuire and RvssEtt'’ should be mentioned. These
investigations are all along the line of bacterial activities in the soils.
Recent studies by Konic,” however, furnish evidence of a catalytic
power of soils due to the presence of an enzyme. <

The literature dealing with the oxidizing power of plant Juices
is already voluminous. Within recent years, our knowledge of
processes going on within the plant has been greatly extended by the
studies which have been made upon oxidizing enzymes. Since several
comprehensive and instructive summaries of work on this cage
have appeared, among which may be mentioned those of Back an
of CzaPex’s Biochemie, it is unnecessary in the present paper et
what has been done in this rapidly developing field.
7 Jour. Amer. Chem. Soc. 30:1295, 1599. 1908. Bur. Soils, U. S. Dept. Agric:
Bulls. 36, 40, 53.

8 Annales Agron. 22:305. 1896.
ae Die Zersetzung der organischen Stoffe und die Humusbi

to Jour. Agr. Sci. 1:261. 190s.

*t Jour. Agr. Sci. 2:305. 1907.

1 Landw. Versuchsst. 63:471. 1906; 66:401. 1907-

Idung.- Heidelberg:

1999] SCHREINER & REED—OXIDIZING POWER OF ROOTS 357

The study of the oxidizing enzymes which are given off by the
roots of plants, i. e., extra-cellular oxidation, has received less atten-
tion, and it is to this particular field that the present study belongs.
Moutscu'’ appears to have been the first to demonstrate the
oxidizing power of root secretions and to show their enzymotic
nature. He found that the root secretion was capable of oxidizing
various organic substances, such as guaiacol, pyrogallol, and gallic
aid, His work showed that there was considerable active secretion
on the surface of growing roots, and that this secretion had definite
powers to effect changes in organic substances.

Czapex,'4 in making a general study of root secretions, followed
some of the investigations made previously by MoxiscH. From
experiments upon the action of seedling roots upon starch paste and
sugar solutions, he regarded it probable that the growing roots
produce only diastase or inverting ferments, although exact proof
could not be offered. He believed, however, that the experiments
of Mortscu failed to prove the production of oxidizing enzymes by
Toots.

The ideas of the oxidizing powers of roots set forth by Mo LiscH
are well corroborated by the investigations of RACIBORSKI'5 upon
the oxidizing powers of plant tissues. ;

Th his work reagents were used which were so nearly non-toxic
that they could be added to solutions in which plants were grown.
some experiments the reagents were added to water cultures
Containing the growing plants; in others, strips of filter paper which
had been saturated with the reagent were applied to the surface of
Sowing roots. The substances used for showing the oxidizing power
of growing roots were a-naphthylamine, benzidine, phenolphthalin,
ferrous ammonium sulfate, Barbadoes aloes, guaiac, phloridzin,
PyTogallol, leucomethylene blue, etc.

‘The eéxtra-cellular oxidation by the roots of the phanerogams
Studied was found to be strongly localized and limited to the absorbing
Sutface of the root. The most intensive oxidation occurs 1n the
Tegion covered by the root hairs. After the death of the root hairs,
'3 Sitzb. Akad. Wiss. Wien. Math. Nat. Kl. 96:84- 1888,

4 Jahrb. Wiss. Bot. 29:321. 1896.
*S Bull. Acad. Sci. Cracovie 1905:338, 68, 693-

358 BOTANICAL GAZETTE [May

as the root grows older, the oxidation becomes weaker (as shown by
the less intense coloration) and vanishes in basipetal order. The
short growing zone of the root between the root cap and the region
of root hairs shows very little if any oxidation. The cells of the root
cap behaved differently in different plants. In some there was a
very weak oxidizing power, insignificant in comparison with that
of the absorbing region of the root; in other plants the root cap
showed no power to oxidize. This observation is the more inter-
esting because PFEFFER’ regarded the experiments of Mo.iscH
to lack proof that the guaiac-bluing power was due to living cells
and not to the dead or dying cells of the root cap.

The oxidation which occurred in naphthylamine and benzidine
solutions first appeared on the outer surfaces of the walls of the
root hairs and epidermal cells, later in the wall itself, and finally in
the outer layer of the ectoplasm. When roots were left for a long
time in a solution of these chromogens, the entire protoplasm of the
epidermal cells and root hairs gradually assumed the dark color of
the oxidized chromogen, although it was not determined whether
this color was due to the diffusion inward of the dye formed at the
surface, or to an actual intracellular oxidation.

- MATERIAL AND METHODS
The experiments described in succeeding pages consisted in
studying the oxidizing power of wheat plants grown under various
conditions in connection with soil-fertility investigations. It --
necessary to grow the wheat plants used for experimentation 1
solutions, since in such cultures it is possible to observe the oxidation
without disturbing the roots. For the study of soil conditions 47
aqueous extract was made by stirring one part of soil with five parts
of distilled water for three minutes and filtering after 3° gee
through a Pasteur-Chamberland clay tube. It has been found
soil extracts prepared in this manner possess 4 plant-product®
power similar fo that of the soil from which they were made. ss
other words, fertile soils yield extracts which promote ay
growth, and infertile soils give extracts producing poor plant gr° a
The water used in making solutions and soil extract was

sane, 1889-
6 Abhandl. kén. sich, Gesells, Wiss. Leipzig, Math. Phys. Cl. 154375

SCHREINER & REED—OXIDIZING POWER OF ROOTS 359

=)
ordinary laboratory distilled water treated with carbon black. The
yater was distilled from a copper boiler, condensed in a block-tin
yorm, and collected in a tin-lined copper tank. This method of
distillation gives very good water for ordinary chemical work, but
does not free it from traces of volatile organic compounds, which may
wert a toxic action as was described by LiviNGsTON” and the
witers.8 It has been found that these deleterious substances May
be effectually removed by treating the distilled water with some
finely divided solid which possesses a strong absorbing power, such
as ferric hydrate, or carbon black. The procedure usually followed
was to shake up a small quantity of the carbon black in the water
and let it stand for 30 to 60 minutes; at the expiration of that time
the water was filtered through ordinary filter paper and was ready
for use. This treatment has been found to be as efficient in pro-
ducing physiologically pure water as redistillation with strong oxidiz-
ing agents, like acid potassium bichromate or alkaline potassium
permanganate.
The varieties of wheat used in the experiments were © Chul” and
“Harvest Queen.” The seeds were germinated on floating perfo-
fated plates, according to the method described by LrvincsToN”® and
in Bulletin 40 of this Bureau. The seedlings were transferred from
the perforated germinating plates to the cultures just as the first true
laf was beginning to emerge from its sheath.
The plants were held in notches cut in the edge of a cork, as

ibed by Livincston. In this way the seed in which enzymes,
Were acting upon reserve food materials were kept out of the solu-
me and the enzyme effects observed were ascribable to substances
‘tlsing from the roots.
Salt-mouth bottles, having a capacity of 250°°, were used as cul-
‘ure jars, and ten wheat plants were grown in each jar. In each
‘est two cultures containing 20 wheat plants were usually employed,
~ comparison was made with an equal number of plants pore
in pure distilled water under the same conditions. All experiments
Were conducted in a greenhouse. During the season of the year -
” Bur. Soils, U. S. Dept. Agr. Bull. 36. 19°7-

8 .
® Bur. Soils, U. S. Dept. Agr. Bull. 40. 1907-
*® Plant World 9:13. 1906.

360 BOTANICAL GAZETTE [way

in which conditions were most favorable for growth, each experiment
- was conducted for 8 to 12 days before studying the oxidizing action
of the plants, but during the cloudy winter weather the time was
some:imes extended to 14 to 16 days.

In addition to determining the oxidizing power of the plants
subjected to various treatments, their growth was estimated by
recording the green weight and transpiration of each culture.?°

SUBSTANCES CAPABLE OF SHOWING THE OXIDIZING POWER

Two classes of substances have been found useful in showing the
oxidizing powers of plant roots in solution cultures. The first class
Comprises certain soluble chromogens, which yield, upon oxidation
by the plant roots, insoluble colored compounds mainly deposited
upon the surface of the roots. The oxidation is usually rapid enough
to produce marked results before the surface extension of the roots
perceptibly disturbs the zonal distribution of the colors. The second
class of chromogens consists of certain substances which give soluble
coloring matters as the result of the oxidizing action of the roots.
The oxidizing action may be shown by the change from a colorless
to a colored compound, or by a change from one color to another and
distinctly different color.

Compounds belonging to the first class which have been used in
this work are a-naphthylamine, benzidine, vanillin, vanillic acid,
and esculin.

Alpha-naphthylamine is only slightly soluble in water, but con-
‘stitutes a good reagent for use in plant cultures because its colorless
Solution is non-toxic, or nearly so, to plants. When oxidized by the
roots of plants, or by reagents such as ferric chlorid or silver pict
a-naphthylamine is converted into the insoluble, lavender-purpic
oxynaphthylamine. When the oxidation is performed by the growing

Toots of a plant, the oxynaphthylamine is deposited upon the surface

of the roots in characteristic zones, as already described by ae

BORSKI (0. cit. 357). The root cap is slightly if at all bey a

zone of primary meristematic cells immediately back of the ene

_ 4s marked by a distinct narrow band of color; the zone of ad
20 For a discussion of the value of these criteria the reader is referred to ug

Bot. GazettE 40:178. 1905; JENSEN, Bot. GAZETTE 43:11. 1907; and

of Soils, U. S, Dept. Agric. Bull. 47- 1907.

— Xtion. Tt is only slightly soluble in water, but in

oR IE Se aa ae ig eT Tie es 22s &

199] SCHREINER & REED—OXIDIZING POWER OF ROOTS 361

sowing cells in the region of greatest elongation is not intensively
wlored; the more slowly growing portions of the root possess the
purplish color, but it becomes less intense as one passes to the upper
parts of the root.

The superior oxidizing power of the meristematic-tissues of the
plant is not only shown by the narrow zone of deep color formed
othe primary meristem of the apical portion of the roots, but also
by the small dots of color produced on that portion of the root from
which secondary roots arise. If a wheat root 8 to ro‘™ in length
is placed in a solution of naphthylamine it will exhibit, in addition
to the deeper colored zones near the apex, dark-purple spots at the
places where secondary roots are forming and are about to break
through the cortical layers of the primary root. If secondary roots
are already present they show the same zones of colors already
described for the primary roots.

The concentrations of naphthylamine used in solution cultures
ate necessarily low on account of its slight solubility in water, but
we sufficiently strong to show the oxidation. In ordinary practice
10 parts of naphthylamine to a million (10™* per liter of water)
Sa suitable concentration to use. This concentration will eventually
retard the growth of wheat plants, but is not detrimental to growth
i the length of time usually required to demonstrate the oxidizing
Powers of the plant roots. A concentration of 5 parts per million
‘Metimes acts as a stimulant to growth.

Benzidine is another chromogen, which is oxidized by plants

and May advantageously be used to demonstrate their oxidizing
weak, colorless

ble dye which
Benzidine is
gical con-

“olution it is readily oxidized by plant roots to an insolu
slves the roots a blue-black or black appearance.
‘lightly toxic to plant growth, but does not cause patholo vehi
ditions within the time required for demonstrating the oxidizing
Power of the plant roots. A concentration of 5 parts henzidine to
“million of water will give good results and does not injure erent
“bots in 24 hours, although that concentration may eventually inhibit
Stowth,

The effect of oxidation may easily be demonstrated by allowing

Toots of wheat plants to grow in a 5 parts per million solution of

362 BOTANICAL GAZETTE [May

benzidine for 12 to 24 hours. The formation of colors in distinct
zones is fully as striking as in the case of a-naphthylamine. As
before, the root cap does not produce oxidation products, the primary
meristem is marked by a narrow band of brown color, the zone of
elongation is practically uncolored, whereas the portion of the root
just above the zone of greatest elongation is entirely colored blue-
black or black by the oxidation products.

Solutions of vanillin and vanillic acid act in much the same manner
as those of naphthylamine or benzidine, but the concentrations re-
quired to demonstrate the oxidizing power of roots are quite strongly
toxic.2! Both substances are converted by the oxidizing action of
the roots into a purple insoluble dye which stains the surface of the
roots in the manner previously described. The concentration of
vanillin in the solution most favorable for showing oxidation with
wheat plants lies between 250 and 500 parts per million. A solution
of this concentration will demonstrate the oxidizing power of ‘the
roots before the plants become seriously injured. To demonstrate
the oxidizing power of roots with vanillic acid, a solution of the
latter containing 25 to so parts per million should be used.

Esculin is another chromogen belonging to this class, but was
found to be less suitable for this work. Esculin solutions, when
freshly prepared, exhibit a blue fluoresceace. After plant roots
have grown for a few days in such a solution, the blue fluorescence
is lost, and the roots themselves are colored yellow as 4 result 0
their oxidizing activity, the dye formed being insoluble an
upon the surface of the roots where the greatest oxidation occurs.
The concentrations necessary to demonstrate the oxidizing power of
roots range from soo to 1000 parts per million, and are eventually
quite toxic to wheat plants. sted

The second class of chromogens, viz., those which are conve e
into soluble coloring matters, are in many respects more useful 10
oxidation studies than those belonging to the first class, spore
the intensity of the color, and hence the amount of oxidation,
he quantitatively expressed. The substances belonging to the sig
class which have been employed in this study are phenolph

Biol. Chem

d remaining

2t Bur. Soils, U. S. Dept. Agric. Bull. No. 47- 19°7- Proc. Soe.
1:33. 1907. Bot. GAZETTE 45:73. 1908.

1909] SCHREINER & REED—OXIDIZING POWER OF ROOTS 363
loin, and leuco-rosolic acid. Alcoholic solutions of guaiac were
also used for various tests, but could not be put into solution cultures
containing growing roots.

The value of phenolphthalin, a leuco-compound prepared from
phenolphthalein, as an indicator of oxidizing enzymes, has been
demonstrated for plant work by KasTLE* and by RACIBORSKI.**
Phenolphthalin is prepared by the method described by BAEYER,”*
which consists in reducing ordinary phenolphthalein with zinc dust
ad sodium hydroxid to phenolphthalin. The latter substance
is oxidized back to phenolphthalein by the oxidizing power of the
plant roots, a change which is readily demonstrated when the solu-
tion is rendered alkaline. The following procedure was observed
in preparing this reagent. Weigh out 250% of phenolphthalein,
3" of sodium hydroxid, and 4 or 5®™ of zinc dust. Place all in a
flask and add 100 to 150°° of water. Place the flask ona sand bath
and heat sufficiently to cause a rapid evolution of hydrogen, without
causing the contents of the flask to boil violently. The heating usually
requires 2 to 3 hours to effect reduction of the phenolphthalein.
The contents of the flask, after reduction is completed, may be filtered
and rendered nearly neutral with hydrochloric acid, and may then be
used as an indicator in the plant cultures. However, better results
may be obtained by using phenolphthalin
method given by Baryer. After purification the phenolphthalin
is dissolved in N/10 or N/20 NaOH and a few cubic centimeters
of the alkaline solution put into each culture, adding equal amounts
to cultures which are to be compared. If quantitative results are
desired, it is necessary to reduce all the solution cultures to neutrality
or the same degree of alkalinity. A very slight degree of alkalinity
Snot usually harmful to plants within the duration of an experiment,
and is favorable to the process of oxidation. Phenolphthalin 1s
Slowly oxidized by mere contact with the air; therefore it is advisable
to instal controls which will allow the results to be corrected for this
atmospheric oxidation. When the phenolphthalin is added to the
Solution cultures, a like quantity is therefore added to jars of distilled

7 Amer. Chem. Jour. 26:526. 1 Hyg. Lab.
: . 262526. 1901. YB:

Hosp. Ser. Bull. 26. 1906. :

23 Bull, Acad. Sci. Cracovie, Math. Nat. Cl. 1905:33°-

74 Annalen Chem. 202:80. 1880.

purified according to the —

U. S. Pub. Health and Mar.

364 BOTANICAL GAZETTE [May

water equal in volume to the cultures. The amount of oxidation in
these blanks is subtracted from what is observed in the plant cultures,

Plant cultures usually show striking results at the end of 10 to
20 hours, depending somewhat upon the temperature and amount
of root surface. At the end of the experiment the plants are removed
from the cultures, and all are rendered distinctly alkaline with sodium
hydroxid solution, and thus the red phenolphthalein color appears.

The great advantage in the use of phenolphthalin to demonstrate
the oxidizing power of roots lies in the fact that it is capable of yielding
quantitative results. After the colors have ‘been developed in the
alkaline solution, their intensities may be estimated by the aid of a
colorimeter. In the work reported below the color intensities were
estimated by means of the colorimeter previously described,”* which
permits of rapid and accurate readings. The colored solutions may
be read against a standard phenolphthalein solution or against ‘
standard Lovibond red glass slide.2° The readings of the colori-
metric tubes are inversely proportional to the color intensity and are
easily reduced to their relative values. |

Aloin is a substance which may be used to demonstrat
dizing power of roots in the same way as phenolphthalin is used.
Aloin, or barbaloin, is the active principle of Barbadoes aloes, and
is obtained in the market in the form of a yellow powder, fairly soluble
in water and serving well as an indicator of the oxidizing power of
plants. At the concentrations used in our work it was not found
to exert any toxic action upon plants. As a result of a limited invest
gation of the chemistry of aloin, it seems that its value as an indicator
of the oxidizing power of plants depends largely upon the content
of iso-barbaloin.

When oxidized by the plant roots, the aloin solution is ne
from a pale yellow color to a permanent deep wine-red color, sim .
to that given by KLuNGe’s reaction for iso-barbaloin. KLUNGE §
reaction’? consists in dissolving aloin (containing iso-barbaloin)
2% Jour. Amer. Chem. Soc. 27:1192. 1905. Bur. Soils, U. S. Dept-

e the oxi-

Agric. Bull. 31

arm.
26 Lovigonp, Jour. Soc. Chem. Ind, 13:308. 1894; see also ScHREINER, Ph

Rev. 19:61. 1901
27 Schweizerische Wochenschr. Pharm. 21:1. 1883; also Leger
Acad. Sci. Paris 131:55. 1900.

Compt. Rend.

weg] SCHREINER & REED—OXIDIZING POWER OF ROOTS 365

ais per cent. sodium chlorid solution and adding 5°¢ of concentrated
copper sulfate solution. Almost immediately the straw-yellow solu-
tion begins to change to a permanent deep wine-red. The change is
hastened by warming the solution.
When experimenting with plant juices containing enzymes, there
appears to be a difference between the reactions to aqueous and
alcoholic solutions of aloin. As the result of experiments described
in detail in a subsequent section of this paper, it was found that an
aqueous solution of aloin is a better indicator of the presence of
oxidase, while an alcoholic solution of aloin is the better indicator
of peroxidase. :
Aloin, like phenolphthalin, should be added to neutral or faintly
alkaline culture solutions, and where quantitative results are desired
all solutions should be of the same degree of alkalinity. In all of
our work aloin was added at the rate of 100 of aloin to 250°° of cul-
turesolution. If actively growing seedlings are used in a very faintly
alkaline solution, a small amount of red color may be developed in
an hour or two, but the experiments should be continued for 12 to
20 hours for the final observation. When certain inorganic salts were
present in the culture solutions, the aloin red color was slightly mod-
ified. The addition of nitrates or previous treatment of the soil
&tracts with an absorbing agent gave the oxidized aloin a purplish
linge, resembling that of fresh fuchsin solution. The presence of
calcium carbonate gave a purer red color, resembling alkanna or
cochineal solution.

The fact that aloin is changed by oxidation from a light yellow to
a deep red solution makes it somewhat more dificult to obtain
Colorimetric readings than in the case of phenolphthalin, where there
'Sa change from a colorless to a red solution. Nevertheless, it is
Practical to use the colorimeter for measuring approximately the
Intensity of color in aloin solutions, by arranging the solutions 1
the order of their apparent color intensities, and using each solution

t as an unknown and then as a standard for the next higher.
For example, let No. 1, the weakest color, be the standard against
Which No. 2 is read. Then discard No. 1; set No. 2 at a convenient
Mark, and, using it as the standard, read No. 3. In tum No. 31s
Used as the standard for No. 4, and so on. In this way one avoids

366 BOTANICAL GAZETTE [May

the necessity of comparing a solution strongly tinged with yellow
against a solution which contains little or no yellow tint. In any two
solutions to be estimated the tints of yellow should not be greatly
different.

Leuco-rosolic acid is another reagent which is useful for demon-
strating the oxidizing power of plant juices?® and plant roots. When
a few cubic centimeters of a slightly alkaline, colorless solution are
added to a culture containing plants, the leuco-rosolic acid is oxidized
back to rosolic acid, the change being shown by the appearance of
the red color. This reagent is not so generally useful as phenol-
phthalin and aloin, since it is more readily oxidized by mere contact
with the air, as well as being more difficult to prepare.

PRELIMINARY EXPERIMENTS |

The first experiments were conducted for the purpose of ascer-
taining some general facts concerning the phenomenon of oxidation
by the roots of seedlings, as well as to learn the methods best suited
for studying oxidation in soil extracts. The experiments of Rac
BORSKI dealt with plants growing under what may be termed pure

culture conditions, and those of KaAsTLE were concerned with the
oxidizing power of plant extracts. ae

In the first experiment, wheat seedlings 4 days old were placed
in solutions of a-naphthylamine having concentrations of I, 2) 5
and 10 parts per million, and in a solution of 5 parts per million
benzidine. The experiment was set up at 4 P.M. August 6, and
observations were made eighteen hours later. At the expiration of

that time colors could be distinctly seen on the white surface of the
wheat roots. The roots in the solution of 1 part per million naph-
thylamine were pale lavender; in 2 parts per million they were oe
nounced lavender, except at the root cap; in the 5 parts per hee
solution they were violet in the region occupied by the primary
meristem, and in the region of the root hairs where growth of elonga-
tion occurs, while the root cap and a narrow zone just above the oe
mary meristem were uncolored; in the 10 parts per million solution '
roots showed the same colors as in that of 5 parts per million. their
roots in the solution of 5 parts per million of benzidine showed the

II. 26:
#8 Kast, J. H., Hyg. Lab., U. S. Pub. Health and Mar. Hosp. Set: cs
17. 1906.

1999] SCHREINER & REED—OXIDIZING POWER OF ROOTS 367

power of oxidation by the formation of brown-violet color, distributed
in the same manner as described for the roots which grew in the solu-
tions of naphthylamine.
In order to learn whether the oxidizing powers of roots were
aflected by conditions which favor growth, and also whether the
method used in the first experiment would show such differences,
the following experiment was made. Three water cultures were
made, in each of which an equal number of wheat seedlings of uni-
form age and size were employed. One culture was made with
tedistilled water, the second with an aqueous extract of a rich garden
soil, the third with a dilute aqueous extract of well-decomposed
stablemanure. After the plants had grown for one day in these liquids,
the oxidizing powers of the plants were observed by transferring
them to other bottles containing 2 parts per million of a-naphthyla-
mine in distilled water. At the expiration of 18 hours the intensity
of the purple colors showed that the roots which had previously
grown in the extract of garden soil had oxidized more naphthyla-
mine than those which had grown in distilled water, and those which
d grown in manure extract had oxidized more naphthylamine than
those from the garden soil extract. At the end of 24 hours the differ-
€nces in color intensity in the two cultures were still more marked.
The next experiment was an attempt to employ a method which
would permit a more accurate quantitative expression of the oxidizing
Power of the roots. Two cultures of wheat seedlings were grown
for 5 days in an extract of unproductive soil, under the same condi-
tions as two other cultures in an extract of rich garden soil. Each
culture contained 60°° of the respective soil extract. The oxidizing
Power of the roots in this experiment was shown by using phenol-
Phthalin. The phenolphthalin was prepared by the method —
in a previous paragraph, and o.4°° of the freshly prepared solution
Were added to each culture of plants after they had grown 5 days in
their Tespective solutions. Nineteen hours after adding the indicator
Plants were removed from the cultures and the solutions rendered
alkaline, thus producing the phenolphthalein color. The solutions
"ere brought to the same volume by the addition of distilled water,
and the relative amount of oxidation was measured by determining
the color intensities of the different cultures.

368 BOTANICAL GAZETTE [way

The two cultures of poor soil gave readings of 40 and 42 divisions
on the graduated tube against slide No. 2 (Lovibond system); the
two cultures of rich garden soil gave readings of 14 and 24 divisions
against slide No. 4 (Lovibond system). Averaging the readings and
comparing the intensity of the colors, the oxidation in the poor lawn
soil and in the rich garden soil stands in the ratio of 1 to 4, or more
exactly as 19 to 82. This result indicated that a procedure based
upon this method will give satisfactory quantitative results.

This method was further tested by another experiment in which
different beneficial treatments were applied to an extract of the
unproductive soil used in the last experiment. The results of the
last experiment showed that the oxidizing powers of plants growing
in solutions of different physiological properties vary considerably,
but left the question open as to how much of the oxidation result
might be due to plants and how much to the solution. In the pres-
ent experiment, therefore, two of the four bottles in each set of solu-
tions were left unplanted, and their oxidizing powers measured along
with those of the solutions which contained plants. The treatment
employed consisted in adding fertilizer substances in the form of
pure chemicals. Calcium carbonate was added at the rate of 2000,
and sodium nitrate at the rate of 50 parts per million. The cultures
were put up August 24 and allowed to grow until August 28, when the
amount of water transpired by each culture was ascertained and 3°°
of a freshly prepared phenolphthalin solution added to each bottle.
The color of the phenolphthalein was brought out by adding Ss few
drops of strong alkali to each culture, and the intensities of the differ-
ent solutions were compared in the colorimeter. Table I presents
the figures which give the relative amount of oxidation in the pe
and unplanted solutions. | When the phenolphthalin solution a
added to the culture jars, the same quantity was added es ed ’
distilled water, which served as a control upon the oxidation incident
to contact with atmospheric oxygen. The color intensity of the contro
was determined and subtracted from each of the other readings:

The plants used in this experiment were quite young, and the ee
iment was only continued for four days, a period rather too it
for the maximum oxidation effect, as shown by subsequent =
ments; nevertheless, the results show that the different treatmen

1909] SCHREINER & REED—OXIDIZING POWER OF ROOTS 369

TABLE I
Relative oxidizing power of cultures and unplanted solutions of Takoma lawn
soil extract with and without the addition of fertilizer ingredients. Oxidizing power
of plants grown in distilled water used as the basis of comparison (p.p.m.=parts per
illion).

No. Solutions bir sete
.......+| Distilled water (planted)
a... Distilled water (ainated> Average ep
Bee. Extract Takoma lawn soil (planted) 88

Bierce oe “ “ce “ iad 74
RS ie ce “ ef “* (unplanted) 8
a “ce “ “oe ce “ce 19
eee e's o = “+ 2000 p.p.m. CaCO; (planted) 113
Sree shies #6 “ “cc “ + 2000 ‘“ rT; “ 63
BANGa. | “ec ‘c ‘ec cc +2000 “ “ (unplanted) 19
\ ae sa nie ee * + 2000 Ss = eZ 25
fees... = - “ 4 sop.p.m. NaNO; (planted) 98
ee 63
Sn “cc “ “cc ‘cc = 50 “c “c (unplanted) 24
ao. . ¢ oe Ses . < 17

allected the oxidizing powers. The plants grown in extracts of poor
Soil possessed less oxidizing power than the controls in distilled water,
but the oxidizing power was increased by the addition of calcium
carbonate. The addition of sodium nitrate did not show any marked
increase to the oxidation in those solutions in the time of the experi-
ment, although its effect as shown in later experiments is always
eneficial to oxidation.

The point to be emphasized in this experiment, which has not
been previously brought out, is that the soil extract unplanted pos-
Ssses a comparatively feeble power of oxidation, as shown by the use
of phenolphthalin, and that the addition of calcium carbonate and
sodium nitrate slightly increased this small oxidizing power.

An additional experiment was performed, using three different
salts in distilled water. The results of this experiment, which are
sven in Table II, confirm those of the foregoing experiment in the
soil extract. The cultures were made in duplicate and the figures
epresent the averages of each pair. es

The enzymotic nature of the oxidizing processes was next investi-
Bated, using alcoholic guaiac. When alcoholic guaiac 1s added to
“solution in which wheat roots have been growing for a time, evidence

370 BOTANICAL GAZETTE [may

of the presence of peroxidase was obtained, but not of oxidase;
however, when young growing wheat roots are treated with a solution
of alcoholic guaiac they instantly give a blue color, which deepens
when hydrogen peroxid is added. This indicates that the cells of
the plant root contain an oxidase, as CzAPEK has shown.?9

TABLE II
Relative oxidizing power of cultures and unplanted solutions of three nutrient
salts. Oxidizing power of plants grown in distilled water used as basis of comparison
(p.p.m.=parts per million).

No. Solutions Ror rar ns
esis Controls in distilled water (planted) 00
sgh nea Solution, 50 p.p.m. of NO, as NaNO, (planted) 282
ee al 33 oes Af “ ~ (unplanted) 39
Chee) eee aes: K as KCl (planted) be
Series " ny th “« (unplanted) 36
Boe on iy 5° es PO, as Na,HPO, (planted) 88
7 oe as a tons «“ _« (unplanted) a

A word may be introduced at this place concerning the possible
function of bacteria in producing oxidizing ferments which might
accomplish some of the effects noted. It is, of course, possible —
such organisms existed in the culture employed, since after _
the extracts no especial precautions were taken to keep them sterile,
and microorganisms which were on the roots of the plants would
certainly be introduced into the solutions. That these microorgan
isms were responsible for any appreciable amount of oxidation 10
the experiments described in this paper is hardly possible. 2 fe :
first place, the solutions used were not well adapted for a very theity
development of microorganisms, as was shown by their gues
from turbidity, odors, or other indications. The definite zone ©
color produced when indicators like a-naphthylamine and bee .
were used, and their close correspondence to definite zones it
in the root show that the oxidation is performed only my ite
intimately connected with the roots. The colors due to —
were most intense on the regions of the root where growl *
active, whereas we would expect that the bacteria, if zonally 4
tributed, would be more abundant on the dying cells of the

29 Annals of Botany 19:75. 1905.

1909] SCHREINER & REED—OXIDIZING POWER OF ROOTS 371
or on the dismantled cortical layers of the older parts of the root.
It seems, therefore, highly improbable that the oxidizing activities
of microorganisms can be responsible to any appreciable extent
for the results observed.

OXIDATION IN SOIL EXTRACTS

Following the preliminary experiments already described, further
experiments were made, to study in more detail the oxidizing power
of plants grown in extracts of soil of different character. These exper-
iments were chiefly designed to study the oxidizing powers of plants
in extracts of good and poor soils, of extracts treated with absorbing
agents, and in distillates of soil extracts.

The difference in oxidizing power of plants in extracts of fertile
and infertile soils is shown by the following experiments. In the
first experiment, an extract of Takoma lawn soil was compared with
an extract of good Leonardtown loam. The former is a very unpro-
ductive soil, and the latter is a much better and usually a very pro-
ductive soil. The oxidizing powers of the plants were determined
by adding phenolphthalin to the cultures, after the plants had grown
in them for nine days. The growth and oxidizing powers of the
plants are shown in Table III, relative to control cultures made in
distilled water, which are represented as 100 in each case.

TABLE III
: Comparative growth and oxidizing powers of plants in extracts of Takoma lawn
soil and good Leonardtown loam. Growth expressed i f transpiration

N Relative Relative
a Solutions growth oxidation
a Oe isnest 88
oe Controls in distilled water Too ie
. --.........| Extract Takoma lawn soil 33 A é
In the comparatively short time of this experiment during cloudy
anifested

Winter weather, December 10 to 17, the plant growth as m
by the figures for transpiration did not have time enough to show the
Telative productiveness of the two extracts, since it has usually been
found that the Leonardtown loam extract produces in 14-18 days
better plants than distilled water. The figures do show, however,

372 : BOTANICAL GAZETTE [ay

a much greater oxidizing power in the plants grown in the extract
of the more fertile soil, even under these conditions.

Subsequent experiments were performed, the results of which
corroborated the foregoing. In each case where growth was good,
there was also good oxidation; where growth indicated a poor soil
extract, the oxidation was small, as will be seen from Tables IV
and V

TABLE IV

Comparative growth and oxidizing powers of plants in extracts of poor sandy
loam and garden loam. Growth expressed in terms of relative transpiration.

i Relative
No. Solutions peo oxidation
I............| Controls in distilled water 100 100
2............| Extract poor sandy loa 77 sa
3--++..+-....| Extract garden loam 125 275
TABLE V

Comparative growth and oxidizing powers of plants in extract of good and poor
oils. Growth expressed in terms of relative transpiration.

. Relative
No. Solutions peo oxidation

Peed Be
RES Gee nna Controls in distilled water ages
ue See xtract Arlington clay loam 75 ra
a Extract Clarksville silt loam 123 Bo
Ce eS Extract Stockton peat ae ;

In all these experiments where direct comparisons are made
between the extracts of soils which were so poor as to give less plant
growth than pure distilled water, and other extracts giving materially
Sreater growth than the same, it appears to be unmistakably es
that the cultures made in extracts of good, fertile soils possess ge
greater oxidizing powers than those made in extracts of soils 0
relatively less fertility.

The next question considered was concerned with the effec e
treating the soil extract with absorbing agents. Treating the exttet
of a more or less unproductive soil with carbon black or other §' es
absorbing agent is usually beneficial to growth. This response see 5
to be quite general for all poor soil extracts, although het ae ‘
to other treatments may be quite different. Previous work 1n

t of

19909) SCHREINER & REED—OXIDIZING POWER OF ROOTS 373

-laboratory3° has shown that this ameliorating action is due to the
removal of deleterious organic substances. Extracts were treated
with carbon black or ferric hydrate. The absorbing agent was
shaken with the soil extract and filtered off at the expiration of a half-
hour, in the same manner as the distilled water used in the experi-
ments was prepared. The relative effects of this treatment upon
_ growth and the oxidizing power of the plants is shown in Table VI,
where the effect in the untreated soil extract is in each case taken as
100.
TABLE VI

Effect of treatment with carbon black and ferric hydrate upon growth and oxi-
dizing power of plants grown in extracts of various soils. Growth expressed in terms
of relative transpiration.

s Soil extract ae | alee

Arlington clay loam* 100 100
: «carbon black treated 124 265
Takoma lawn soil* 100 100
«« «carbon black treated 137 100
Alloway clay* 100 100
. oY carbon black treated 116 117
4..........| Dunkirk sandy loam* <0 100
. ««" « carbon black treated 112 280
Miami silt loam* 100 100
6 Te eee ferric hydrate treated 171 198
tsss4+....| Marshall clay loamt 100 100
ea ees carbon black treated 216 130
7...+......| Clarksville silt loamt 100 100
3 rs 2 carbon black treated 450 227
Bee. Elkton silt Joamt 100 100
carbon black treated 179 317
ieee... Cecil fine sandy loamt 100 00
Io carbon black treated 112 200
eae’ -| Hagerstown loamt 100 100
é es carbon black treated 230 500
Cecil sandy loamt 100 100
en oe . carbon black treated 193 241
ay Dutchess silt loamt 100 100
3 og vs carbon black treated I10 373
oe Poor sandy loamt 100 100
14 roe 2 ferric hydrate treated 170 534
“+ +++...| Garden loamt 100 100
% “ ferric hydrate treated 136 313

Alaa phthalin used in esti spin
n estimating ee He

= Tt will be noted that in all but one of the soil extracts the effects
the treatment with an absorbing agent strongly increased the

.* Bur. oo U. S. Dept. Agric. Bulls. 28 36, 40. Jour. Amer. Chem. Soc. 30:

374 BOTANICAL GAZETTE [may

oxidizing powers of the plants subsequently grown in the extracts,
and the growth of the plants was also increased.

The increased oxidation, as well as the increased growth, points
directly to the conclusion that the soil extracts have been so improved
by the treatment given as to induce a more active functioning of
processes necessary to secure the best conditions for growth. In
the single case of No. 2 the growth was increased as a result of the
treatment with carbon black, but the oxidation was not. This
result was frequently obtained with the Takoma lawn soil; in some
cases the oxidizing power was even slightly decreased as a result of
treatment with absorbing agents, although growth was increased.
No satisfactory explanation has as yet been obtained for this appat-
ently exceptional action. It may be found upon further investigation
that the lack of response was due to the presence of matter inhibiting
oxidation, which was not removed by the carbon black. This
question seems worthy of more study than we have been able to
give it.

Extracts of poor soils sometimes contain volatile bodies of a
deleterious nature, which can be driven off by boiling and collected
in the distillate. The writers have described’* the behavior of
plants grown in such distillates. Where the deleterious bodies as
volatile, the distillate usually exhibits the same toxic properties which
the original extracts previously possessed, and the residue is corre:
spondingly improved.

To study the effects of these distillates upon the oxidizing aia
of the plants the following experiments were made. One her of
such soil extract was placed in a distilling apparatus and distille
until 200°¢ of distillate had passed over and been condensed. e
fluid was made up to 500°° by adding water and designated firs
portion. When a second 200° of distillate had been collected, 5
was likewise made up to 500° and designated second portion:
Cultures were made in each portion, together with controls 1n a
distilled water. At the end of a week the plants in the differen
solutions showed marked differences. The plants in the first ga
of the distillate were very small and were dying; those in the secon
portion were much better, in fact, were equal to the controls growin

3* Bur. of Soils, U. S. Dept. Agric. Bull. 40. 1907.

1909] SCHREINER & REED—OXIDIZING POWER OF ROOTS 375

in distilled water. One hundred milligrams of aloin was added to
each of the culture bottles, and on the following day the amount of
oxidation was noted by comparing the intensity of red color in each
culture. The cultures in the first portion showed much less oxidation
than either of the other two. The most oxidation appeared to have
gone on in the cultures in the second portion, which was slightly
in excess of that in the control cultures in pure distilled water.

The question was studied further, and in a more quantitative
manner, by the following experiment. An extract of Elkton silt
loam, having a volume of 750°°, was placed in a flask connected
with a condenser and distilled. The distillate, amounting to 500°,
was collected in two portions of 250°° each and used as a culture
medium in which plants were grown. The residue in the distilling
flask, which was diluted to its original volume, was also used for
growing plants. For comparison, cultures were also made in the
original soil extract. The wheat plants were allowed to grow in
the various solutions for 13 days, and then their oxidizing powers
were estimated by means of phenolphthalin. The growth and
oxidation are shown in Table VII.

TABLE VII

8 Growth and oxidation in distillate and residue of extract fro
towth expressed in terms of relative transpiration.

m Elkton silt loam.

Relative Relative
growth

No. Solutions oxidation
(Glenn ————
*s+++....| Original soil extract untreated 100 —
sas First portion of disti 53 =
4 ond portion of dis illate fe 19

-| Residue after distillation, diluted to original
olume 1” 180

These results show that the distillates of this soil extract were
: than the original untreated

ins favorable for growth and oxidation

~ extract, while the residue from distillation was materially
improved. ‘This seems to indicate that the original soil extract, like
Others which have been investigated, contained a volatile toxic
Substance which inhibited oxidation by the roots, and that this
Substance was driven off by the process of distillation, with resulting

32 Bur. Soils, U. S. Dept. Agric. Bulls. 28, 36, and 40-

376 BOTANICAL GAZETTE [way

benefit to oxidation in the residue. Judging from the growth of the
plants, the first portion of the distillate contained a larger proportion
of this deleterious substance than the second, although this smaller
amount appears to be just as deleterious to the oxidizing powers of
the roots as the larger amount present in the first portion. The
oxidizing power of the plants in the residue was much greater than in
the distillates or in the original soil extract.

Evidently the oxidizing powers of the roots are affected by certain
external conditions, since an improvement in the physiological
properties of the soil extract results in increased oxidation, and the
presence of deleterious bodies results in decreased oxidation.

From the experimental results thus far presented, it appears that
the oxidizing power of the soil extracts themselves can be regarded
as partly, but not mainly, responsible for the oxidation observed
in the experiments. In one of the preliminary experiments reported
in Table I, it was shown that the soil extract after filtration through
a Pasteur-Chamberland filter tube exhibited some oxidation, even
when no plants were growing. It was likewise shown by the results
in Table II that certain nutrient salts dissolved in distilled water We
able to accomplish a material amount of oxidation without the pres-
ence of growing plants. It seems unlikely, therefore, that any con-
siderable amount of oxidation was performed by microorganisms. If
we consider the result in this last experiment, where oxidation a
increased in the residue from distillation after continued boiling, it
seems that any extensive action, not only of microorganisms, Lea also
of enzymes, must be precluded. In the soil, however, it is quite prob-
able that both of these oxidizing factors would come into play, but

-it is quite certain that the oxidizing power of the roots would accom:
plish a considerable portion of the oxidation observed.

THE NATURE AND ACTIVITIES OF THE OXIDIZING ENZYMES :
Mention has been made in preceding pages of te eae
nature of the oxidizing action of the roots and consideration : aa
be given to the nature of the enzyme or enzymes which re
the oxidation. So far as known, the oxidation effects observed pe
entirely due to the action of enzymes and not to the other actiN!
connected with the growth of the roots themselves.

—— Ig0g] SCHREINER & REED—OXIDIZING POWER OF ROOTS 377

When a few drops of alcoholic guaiac are added to water, or a
suitable solution of salts, in which wheat seedlings have grown for
several days, there is sometimes a faint blue color, indicating the
presence of oxidase, but more often there is no blue color. When a
drop of hydrogen peroxid is added, however, the liquid turns blue,
siving a color varying from medium to very intense, depending
somewhat upon the age of the seedlings, and the number of roots
which have grown in the culture. The guaiac-peroxid reaction
indicating a peroxidase is confirmed by the reaction to phenolphthalin
and aloin, both of which agree in showing the presence of peroxidase.
When the roots of a young wheat plant are immersed in an alcoholic
guaiac solution, they immediately turn blue, indicating that they are
relatively rich in oxidase, although but little oxidase appears in the
water in which they grew. This may be due to the retention of
oxidase by the root cells during life, but when the outer cells are killed
by the alcoholic guaiac the oxidase escapes and becomes evident
through its reaction with guaiac. An aqueous extract of crushed
roots, shows strong oxidase reaction as well as peroxidase reaction.
In the course of a brief examination of different parts of the young
wheat plants, it was found that the partially depleted seeds showed
avery strong oxidase reaction when guaiac was used, while the per-
Oxidase reaction was relatively less than in the extract of crushed

When the solution in which wheat roots have been grown for
some days is boiled for five or ten minutes, and cooled, the oxidase —
and peroxidase reactions disappear.
: The temperature at which the peroxidase is destroyed was deter-
_tnined by heating a culture liquid which showed an active peroxidase
action. The culture liquid was heated to successively higher tempera-
tures and held at each for five minute periods. The temperature at
Which the enzymes appeared to be destroyed was 60° C. or very close
ereto.
The culture liquid was examined for enzy
lures of different ages to learn whether the enzyme reac
‘tong in all. Wheat seeds were germinated on perforated cork
Plates floating on the surface of water in crystallizing dishes of 500"
“pacity. When cultures were on hand aged two, three, four, five,

mes in a series of cul-
tion was equally

378 BOTANICAL GAZETTE [May

six, and seven days respectively, tests were made with guaiac, alco-
holic aloin, and phenolphthalin.

The tests with guaiac showed that the oxidase reaction which was
weak in the two- and three-day cultures was quite strong at four days.
The tests with alcoholic aloin and phenolphthalin showed that the
peroxidase reaction was strongest in the six- and four-day cultures,
and considerably weaker in each of the others.

Certain phenomena observed in connection with the use of aloin
in aqueous and alcoholic solutions, suggested that they react differ-
ently with oxidases and peroxidases. Experiments were accordingly
installed to test specifically the action of each solution. Two solu-
tions of aloin were prepared: I, o.250%™ of aloin in 50° of water;
II, 0.2508" of aloin in 50°¢ of 95 per cent. alcohol. One cubic
centimeter of aloin solution I or II was added to 5° of liquid in test
tubes, according to the plan shown in Table VIII. The tubes were
prepared and aloin added at 2.45 P. M., on January 10, and the obser-
vations recorded in the third column of the table were made at It 4. M.
on January 11. The culture liquid when added to the tubes showed
no oxidase but good peroxidase reaction with guaiac.

3 TABLE VIII

Comparative reaction of aqueous and alcoholic solutions of aloi
containing peroxidase.

n toa liquid

Pare
Col observed
Nos. Solution 11 A. “ss January 1
ican
Pet ee A eee

TOs 4... 5, Unboiled liquid + 1¢¢ aqueous aloin pink

St aa ee Boile ee: es ae nak

Sand 6..........| Distilled water + a iy _ Bak

y7and 8..........| Unboiled liquid+ alcoholic “ deep P

9 and 10 ....| Boile oe aati: yellow
Irand12........ .| Distilled water + Loh aciss yellow

An inspection of these results shows that when only peroxidase
present, aqueous aloin is not particularly applicable for :
the presence of that enzyme in the absence of growing par
there was the same development of pink color in the bor ed to a
unboiled liquid. Alcoholic aloin, on the contrary, Was pe like in
deep pink in the unboiled liquid, but remained unchanged 4"
the boiled liquid and in the distilled water.

in liquids
The action of the different aloin solutions was next tested in "4

900] SCHREINER & REED—OXIDIZING POWER OF ROOTS 379

which also possessed a strong oxidase reaction. The roots of 10 wheat
seedlings 12 days old were removed and crushed in a mortar with
distilled water. The filtered liquid obtained from this source gave

-astrong reaction for oxidase when tested with guaiac. As before,
1° of aloin solution I or II was added to 5°° of the root extract in
test tubes, according to the plan in Table IX. The tubes were pre-
pared and aloin added at 4.30 P. M. on January 13, and the observa-
tions recorded in the third column of the table were made at 11 A. M.
on the following day.

TABLE IX

Comparative reaction of aqueous and alcoholic solutions of aloin to a liquid
containing oxidase.

: Color observed
Nos, Solution 11 A.M., January 14
th ate SRG cats
eee Root extract + 1°¢ aqueous aloin_ rec
ee Distilled water + 1°¢ aqueous aloin faint pink
a Root extract + 1¢¢ alcoholic aloin pronounced pink
Oo" a er Distilled water + 1¢¢ alcoholic aloin faint pin
eRe enact ntact

The results of these experiments supplement those of the fore-
going in which a peroxidase liquid was used, by demonstrating that
the oxidase caused a much greater conversion of aloin to “aloin red”
with the aqueous than with the alcoholic solutions of aloin. There
Was in the root extract a distinct peroxidase reaction to guaiac, in
addition to the oxidase reaction, and it is therefore only natural that
tubes 6, 7, and 8 there should be some development of color when
alcoholic aloin was added.
It is evident, from the above results, that in the absence of living
t roots aqueous aloin is principally a reagent for oxidase and
alcoholic aloin for peroxidase. In the experiments where plants are
‘ployed it is however needless to say that only aqueous solutions
e of aloin can be used. wae
Aloin and phenolphthalin having shown their usefulness as indi-
“tors of enzyme action, several other substances were investigated
for Comparison. Leuco-rosolic acid was prepared by reducing
Tesolic acid with zinc dust in alkaline solution. When reduction
"as practically complete, the solution was filtered and neutralized
“ith hydrochloric acid, then rendered slightly alkaline with sodium

380 BOTANICAL GAZETTE [MAY

hydroxid. One cubic centimeter of this solution was added to three
different liquids: I, liquid from culture 6 days old; II, the same
liquid after having been boiled 1o minutes; III, distilled water.
When examined 24 hours later I was pronounced rose-red, while II
and III were merely faint pink; which indicates that leuco-rosolic
acid is capable of showing the action of those oxidizing enzymes.

Attempts were made to use ferrous ammonium sulfate and potas-
sium iodid as indicators of the oxidizing powers of plants by putting
small amounts into cultures containing living plants. Ferrous ammo-
nium sulfate was, in the space of time of the experiment, oxidized
by mere contact with the atmospheric oxygen, and was therefore
discarded as an indicator. Potassium iodid was not oxidized to free
iodin, as RAcIBoRSKI has also found.%$

EFFECT OF DIFFERENT CONDITIONS IN THE SOLUTIONS UPON THE
ACTIVITY OF THE ENZYMES

Mention has previously been made of instances where the variation

in oxidation appeared to be partly due to the acidity or alkalinity
of the solution used as a culture liquid. In such cases the growth
of the plant roots was affected whenever the alkalinity or acidity was
very pronounced. The effect is the more harmful when young seed-
lings are put into such solutions, because at the beginning of the
experiment, when the plants are very tender, the acidity of alkalinity
is greatest and gradually diminishes during the progress of the ed oe
ment. In investigating the effect of acid or alkaline conditions in the
the culture media, instead of using either alkaline or acid solutions at

the start, a method was used whereby the originally neutral solutions

became acid or alkaline as a result of the selective absorption of the
plant in withdrawing nutrients from the solution.s4 It has peer
demonstrated by KoHn and Czapek?s that fungi may render ed
culture media alkaline or acid as a result of their selective absorption,
whereby an acid or a basic radical is removed more rapidly than t
radical to which it is linked. REED%* has observed 4 similar action

33 Bull. Acad. Sci. Cracovie 1905 :668.

34 See CAMERON, Rept. U. S. Dept. Agr. 71:67. 199%)
Agric. Bulls. 30 and 41. 1905.

35 Beitr. Chem. Phys. Path. 8: 302. 1906.

3° Annals of Botany 21: 501. 1907.

Bur. soils Ue

1909] SCHREINER & REED—OXIDIZING POWER OF ROOTS 381

ior the higher plants and pointed out its bearing upon the composition
of nutrient solutions.

Solutions were made up from salts whose radicals are differently
absorbed by growing plants, e. g., calcium nitrate and potassium
sulfate. Where calcium nitrate is furnished the plant takes up NO,
more rapidly than Ca, with the result that the solution becomes
increasingly alkaline. In the case of potassium sulfate, the plants
take up K more rapidly than SO,, with the result that the solution
becomes acid. In the experiments which were made upon this prob-
lem, an attempt was made to determine the acidity or alkalinity of
the solutions when the experiment was terminated. A measured
quantity of solution was boiled in a platinum vessel to drive off CO,,
and then titrated. The results of these determinations are shown
with the other results in Table X.

TABLE X

. Oxidation and growth of wheat plants in solutions which became acid or alkaline
the result of plant growth. Relative growth measured by transpiration.

_ | Acidity at [Alkalinity at)
No, Solution four 9g | lage" os ) me aaa
experiment | experiment
; Bags. | Control in distilled water 100 ere ent I
‘. ae 30 p.p.m. Ca as Ca (NO3)2 i dndety tie ee AD sep aeni eo
ee 30 p.p.m. Ca as CaCl, BAS ed ee et
ye 30 p.p.m. Ca O; 2 So, 4 el aNee 175
io 66 p.p.m. SO, as (NH4)2 SOs MR eee ere oe
oc 66 p.p SO, as K,SO,4 78 n/1%0000 | «+s 100
“eeu 100 p.p.m. NO, as NaNO; Bot Lace n/5 or
ee. 35 p-p.m. K as KCl >. | Neutral | Neutral 97
ee 100 p.p.m. NO, as NaNO; n/620| 283 ee n/ 10000 38
i 63 p.p.m. K as KCl n/620
-
ee a mme n/roo00 | 256
63 p.p.m. K as K,HPO, 1/620
Poorer re OR

These results show that six of the nine solutions became alkaline,
owth and oxidation

two became acid, and one remained neutral. Gr
Were less in the acid solutions than in those which became alkaline,
although in the case of calcium chlorid the result was quite low. In
= case of potassium sulfate and potassium chlorid a part of the depres-
ed may be due to the effect of the potassium, which usually fails
'0 increase oxidation materially, but such is not the case with ammo-

382 BOTANICAL GAZETTE [MAY

nium sulfate. Neither is it probable that the sulfate radical is the
depressing factor, since calcium sulfate compares favorably with
calcium nitrate in its effect upon oxidation. The more favorable effect
of no. 2 in Table X upon oxidation over nos. 3 and 4 is probably to
be attributed to the presence of nitrate, which likewise appears to be
responsible for a material increase in growth.

The greater oxidation accomplished by no. g over no. 10 is probably
not to be attributed to the presence of Cl, but to the smaller amount of
K present in no. 9.

On the whole it appears that while oxidation is affected toa certain
extent by conditions of acidity or alkalinity arising in the culture
medium, it is more materially affected by the specific action of the
salts and their elements in the solution.

The direct effect of acid and alkaline conditions upon the activity
of peroxidase was investigated by the following experiment in which
alcoholic aloin was used as the indicator. A liquid showing strong
peroxidase action was taken from a pan in which several hundred
7-day-old wheat seedlings were growing. Various amounts of n/ 3°
HCl and 2/50 NaOH were added to a set of tubes each containing
10° of the culture liquid and 1°¢ of alcoholic aloin solution added
at 3 P.M., January 13. The following table shows the amount of
acid or alkali added in each tube, and gives the record of the colors
observed at 11 A. M. the following day.

TABLE XI
. . es pik : : absence
Effect of acid and alkaline conditions upon the activity of peroxidase 1n the
of plants.
——
Jor observed at end
No. Solution Colon 20 hours
; int pink
RPE ee eats roe culture liquid+o.1¢¢ 2/50 HCl faint ae
ee I ey - “ +0.2 : me hang
Ee nes + Fé *: 40.5 se . ovlggsa :
oe <s “
Me one CoN ee 2 a “ “ : “ rT;
5 oe a eee ew a bale 6“ “é “cc ia ee “cc “6 25 e
Ce a “ «“ NaOH :
7 i peel aoe pet “c ‘“c “c = . : n/50 “ wine-red
tS ‘“ «“ & ta 6 Pe & deep wine-T”
9 NAR oat oy au v7 “cc “cc AY ° : 7 “c sc : “ “c
IG.; “ “ “ “ te
Peo Nea at +1.0 : _ pink
ER ere . 6: “ neutral to litmus solution deep a
BM cade ow ue ec “ss “ 6c “cc “c “cc oe
Pe

1909] SCHREINER & REED—OXIDIZING POWER OF ROOTS 383

From these results it can only be concluded that a slightly alkaline
medium is most favorable for this peroxidase reaction. It will be
remembered that WoLLNy3’ found also that the oxidation processes
in the soil were distinctly favored by slightly alkaline conditions.

The effect of putrefactive processes upon oxidation is another
question which was briefly investigated. When a number of seedlings
were placed without any support in water containing aloin (the
entire root system, seed, and lower part of the plant, being thus sub-
merged), it has been observed that the red color first produced
subsequently disappeared. An experiment was accordingly planned
to learn whether oxidation phenomena would be affected when the
seeds were submerged and gave rise to products of putrefaction.
Twelve cultures of wheat plants were prepared and allowed to grow
four days in tap water. In one-third of the cultures the seedlings
were adjusted in the notched corks so that only the root systems of
the plants were submerged; in one-third-of the cultures the seedlings
were lowered so that the seeds also were submerged, and one-third
had the seedlings entirely submerged. On the fourth day 10o™
of aloin were added to each culture jar, and they were examined
twenty-four hours later with reference to the production of colors.
It was found that the cultures planted with only the root systems
submerged showed a very considerable amount of oxidation, but in
those where the seeds or entire plants were submerged there was none
of the red color produced by oxidation. In these cultures where no
oxidation was shown, there were putrefactive processes at work, a
fact which is taken to mean that the oxidation effects are not observed
when putrefactive processes occur. Whether this inhibition of oxida-
tion is caused by the products of putrefaction or by a perverted
Metabolism, since the plant must function under somewhat anaerobic
‘onditions, remains undecided.

That the oxidizing power of the plant was not destroyed is shown
by the fact that by raising the seeds out of the culture water and
refilling the jars with fresh tap water containing aloin, the character-
Stie oxidation occurred.

The foregoing experiments raised a question as to the amount of

we Die Zersetzung der organischen Stoffe und die Humusbildung. Heidelberg.

384 BOTANICAL GAZETTE [MAY

oxidation which occurs in poorly drained soils, where putrefactive
processes are known to exist. In investigating this question, two
crops of wheat seedlings were grown in Arlington clay loam in parat-
fined wire pots, giving the pots different amounts of water.

Lot I of the pots was kept at the optimum water content of the
soil. The soil in lot II was kept saturated with water from the start,
and the soil in lot III was saturated after the wheat seedlings were
up. The relative green weight of the first crop of wheat plants, which
grew from February 18 to March 14, was: I, 100; II, 111; II, 104.
The relative weight of the second crop, grown from March 17 to April
8, was: I, 100; II, 67; III, 116. Extracts of these soils were then
made and wheat plants were grown eleven days in the various extracts.
At the end of that time the growth and oxidizing power of the plants
in the different solutions were determined with the result shown in
Table XII.

. TABLE XII os
Growth and oxidation in extracts of soils of varying moisture content. Gro
expressed in terms of relative transpiration.
————
elativ Relative
No. Soil treatments yn xidation
TE ee Soil kept at optimum 1 6S
Oi ee aan Soil kept at saturation 73 ae
OP tS PR ie 6 ee be 6 8 8 i
3 Soil saturated after plants were up Pp beeen

These results show that the effects of the poor drainage condi-

tions appear to be much more marked upon oxidation than upo?
growth. The soil, which was kept at optimum and only saturated
after the plants had started, seemed to remain favorable to growth
in the pots and in the extracts, but its extract was plainly not ai
able to oxidation. In regard to the increase of growth, it should
remembered that this lot of soil was alternately very wet and CT}
during the course of the experiment. .
EFFECT OF TOXIC COMPOUNDS UPON OXIDATION

: : : d Gee h e were used
Aside from the foregoing experiments, 1n which t 2 d plants, a

extracts of soils which displayed toxic qualities tows unds

few investigations were made upon the action of organic compe

whose toxic properties had. been previously determined. aillin,
The organic compounds employed for this purposé Md

1999] SCHREINER & REED—OXIDIZING POWER OF ROOTS 385

cumarin, and santonin. The compounds were dissolved in distilled
water and the resulting solutions used as cultures, taking care that
the concentrations chosen were not so great as to be fatal to wheat
plants within the duration of the experiment. Vanillin was used
at the rate of 100 parts per million, cumarin 10 parts per million,
and santonin in a saturated solution, which was nearly roo parts per
million. The growth of the plants, as measured by transpiration
and stated in figures, taking the growth of similar plants in distilled
water as too in each case, was: vanillin, 63; cumarin, 81; santonin,
75. After the plants had grown in their respective solutions for
12 to 14 days, 100™£ of aloin were added to each and the results
noted on the following day. The results agreed in showing no color
indications of oxidation in any of the cultures where the toxic com-
pounds were present, although the roots growing in the control cul-
tures in distilled water showed by the red color produced that a
material amount of oxidation had been accomplished.

That the mere presence of organic materials did not inhibit the
oxidation was shown by an. experiment employing a solution of
leucine which was slightly beneficial to the growth of wheat seedlings
in solution cultures. Solutions of leucine containing 50 to 100 parts
per million, producing an increase in growth over distilled water of
54 and 98 per cent. respectively, were very favorable to oxidation and
ptoduced a much deeper aloin red than the cultures in distilled water.
It can only be concluded, therefore, that the toxic organic com-
pounds studied were deleterious to oxidation because of their toxic
Properties, and it appears that they were even more deleterious to
oxidation than to plant growth.

The oxidizing action of the plants upon toxic organic substances
is a phenomenon which has been pointed out by the authors cadits
Previous paper3® and will be referred to again later. The experiments
Presented in that paper also showed that the addition of sodium
nitrate and calcium carbonate to solutions of toxic organic compounds
Went far toward decreasing their harmful effects, and in some cases
overcame them entirely. That the organic salts and the physiologi-
cal activities of the plants working together had accomplished the
destruction of toxic substances, was shown by both plant growth and

3% Jour. Amer. Chem. Soc. 30:85. 1908.

386 BOTANICAL GAZETTE MAY

chemical tests. It now appears that while this destructive action
of the plant upon the toxic body is going on, the oxidizing power in
the presence of an excess, as it were, of toxic body is greatly reduced,
and may even be entirely inhibited. The conclusion drawn from
those experiments was that the plant roots are able to oxidize a
certain amount of deleterious organic material, and that the presence
of salts which favor oxidation increases the ameliorating action of the
plant.

This question was studied a little further by an experiment in
which the oxidation in solutions of toxic material was observed.
A solution of cumarin containing 10 parts per million, with and
without the addition of fertilizer ingredients, was used as a medium
for plant growth and subsequently examined for powers of oxidation.
Sodium nitrate was added to one portion of the cultures at the rate
of so parts of NO, per million, and calcium carbonate at the rate of
2000 parts per million was added to another portion of the cultures.
Wheat plants were installed in the cultures October 7 and grew until
October 17. The oxidation was estimated by means of aloin. Table
XIII gives the effect of this treatment upon growth and upon the
oxidation in the toxic solutions and in control solutions to which no
cumarin had been added. In each case growth and oxidation of
the plants in distilled water are taken as 100.

: TABLE XIII
Effect of sodium nitrate and calcium carbonate on growth and oxidation ™ ~
tions of cumarin. Growth expressed in terms of relative transpiration.
ert
‘ Relative
No. * Solutions pow ; enser™
ae cients
I........+..| Distilled water res
co co NaNO, 18 ee
2 Ee ag e +CaCO, pe I
4...........| Cumarin ro p.p.m. _ 139
. nae . s +NaNO, ee st
eS “ «= gCaCO, il ;
2 : : : jous
The results of this experiment show, in harmony with the prev!

ones, that the addition of these fertilizer ingredients overcame to 3
large extent the deleterious effect of the cumarin upon grow'™™ -

one making the cumarin solution to which it was added @ better
medium for growth than distilled water. An inspection of the figure

1909] SCHREINER & REED—OXIDIZING POWER OF ROOTS 387

expressing the relative oxidation shows, however, that the addition
of these salts produced relatively greater increases in oxidation than
ingrowth. When sodium nitrate was added to cumarin the resulting
growth was twice as great as where only cumarin was present; the
oxidizing power, however, was increased over fourfold. In com-
parison with this effect, it will be noted that the addition of sodium
nitrate to distilled water likewise increased the growth twofold and
increased the oxidizing powers two and a half times. It seems quite
evident, therefore, that the ameliorating powers observed under the
conditions of the experiment are to be referred to the cai
oxidizing powers which are thereby brought about, and the
diminution in amount and activity of the toxic material.

It may be noted that LE RENaRpD3° found that nitrates had a
greater antitoxic value than other radicals when Penicillium was
grown in the presence of copper.

SUMMARY

I. Roots of growing plants exhibit an extracellular oxidizing
power which may be demonstrated by the use of suitable chromogens
in nutrient solutions or soil extracts.

_ 2. The oxidizing power appears to be most energetic in the region
of the root where root hairs are found, and to decrease gradually in
activity as that portion of the root becomes older.

3. The oxidizing power of plants grown in extracts of productive
Soils is greater than that of plants grown in extracts of unproductive
soils,

4. Treating the soil extracts with an absorbing agent is usually
beneficial to oxidation.

5. The distillate of a poor soil extract which contains volatile
toxic compounds was less favorable to oxidation than the residue
Temaining from distillation.

6. The presence of toxic organic substances in solution was ex-
Temely deleterious to the oxidizing power of the plants. The oxi-
' power of the plants, especially in the presence of nitrates,

_ “as able to alleviate the toxicity of such solutions.

® Essai sur la valeur antitoxique de l’aliment complet et incomplet. Paris. 1907.

388 BOTANICAL GAZETTE [May

7. The process of oxidation is usually accelerated by the addition
of sodium nitrate to an aqueous soil extract. The addition of other
fertilizer salts also influences oxidation.

8. The process of oxidation by roots is largely, if not entirely,
due to the activity of a peroxidase produced by the roots. This
oxidizing enzyme is most active in neutral or slightly alkaline solu-
tions. The activity of the enzyme may be inhibited by the presence
of acid and also by the conditions in solutions where anaerobic pro-
cesses occur.

_ Bureau oF Soits, U. S. DEPARTMENT OF AGRICULTURE
WasHINcToN, D. C.

¢
i x
— _—

BOG TOXINS AND THEIR EFFECT UPON SOILS:
ALFRED DACHNOWSKI
(WITH TWO FIGURES)

In the north and in the middle west, notably in Indiana and Illi-
nois, and to some extent in Ohio and the adjoining states, there
are extensive swamp areas of vast importance to the state, some of
which are called “unproductive,” and generally are not cultivated.
The statement is made, and there is certainly much truth in it, that
these swamp lands should be naturally very rich in constituents needed
for plant food. Many of these places represent deep basins of accumu-
lated plant débris, and the drainage from hills further enriches the
accumulation in the swamps. They are noted for their dense and
luxuriant surface vegetation. Examples of swamp and bog areas in
this vicinity show clearly that bog water contains apparently all of
the constituents required for the nutrition, growth, and reproduction
of a large variety of grasses, shrubs, and trees. However, from an
agricultural point of view these muck and swamp lands seldom have
given satisfaction, even after drainage or addition of fertilizers.
Ample proof of this is seen in the reports of the various experiment
stations (7,8). Thus far the remedies proposed, as a laboratory
expedient, emphasize the fact that although some principles of soil
fertility seem well established, and can be applied with definite results,
_ there are yet many complex problems, the solution of which would
_ mnaterially enhance the economic importance of peat and swamp soils.

To the writer it has seemed probable for some time that work upon
the chemistry and upon the physiological properties of peat and humus
fompounds must result in data valuable alike to the agriculturist,
the forester, and the ecologist. Through an investigation on the
‘ause of xerophily in bogs (5), there was gained supplementary evi-
dence, of a more direct and positive sort, that the inhibiting factors of
bog are in part the presence in the soil water of injurious toxic sub-

: Contribution from the Botanical Laboratory of Ohio State University, XLII.
Paper was in part read before Section G of the A.A.A.S., at the Baltimore meeting,

This

_ “ecember 29,- 1908.

389) [Botanical Gazette, vol. 47

390 _BOTANICAL GAZETTE [MAY

stances. In that publication experimental data were given to show
that the toxicity of bog water and of the bog-soil substratum can be
corrected by various methods, and that the plants grown in solutions
thus treated show not only accelerated growth and an increase in
transpiration, but also an increase in the green and dry weight of
organic matter.

It is not known, as yet, whether the toxic action of bog water and
bog soil is determined by the action of one constituent or by the com-
bined action of several. The experiments so far completed have
given no definite evidence that the toxins are merely specific excretions
from the roots and rhizomes of bog plants. Preliminary tests, not
here detailed, which were carried on in the winter of 1907, indicate
that the toxicity may be due to a certain unstable body, of the nature
of organic compounds excreted from the roots in the absence of O,
and in heavy clay soils not adequately aerated. It is probably a
product of imperfect oxidation and decomposition of proteins and
other related substances, and it is possible that in respiration bog
plants differ from other plants. Since then an excellent account has
appeared by SToKLAsA (12), in which similar results on the excre-
tions by roots are announced. Largely, however, the toxicity of
bog water seems to be due to another cause. During the changes
which the accumulated plant material undergoes in the process of
peat-making, there are alterations and reductions leading to gaseous
and colloidal products but little known. The relative amount of
these varies with the seasons, and no doubt also with the locality,
but primarily it depends upon the stage in the progress of decompo
sition. CH, and H,S, though produced in small quantities, have
been found to constitute the principal gaseous products. They at
especially noticeable when well-corked jars of bog water remain
standing for some time. Studies on the character of the colloidal
products are still in progress. The injurious products of a mci:
flora accumulating in definite layers of the soil are, perhaps, -
additional factor to be considered. Indeed, it is a serious _—
in physiological ecology that a process must be assigned to @ nae
category or broken up into a number of what often prove _
arbitrary categories, in order to arrive at results in any Way

intelligible.
; : eo
Too often is there a tendency to lay undue emphasis upon oe’

1909] DACHNOWSKI—BOG TOXINS 391

conditions; and the more detailed responses which are due to localized
influences are thus neglected. On the other hand, even though we
decide experimentally which of the physical and chemical variables
involved is of greater influence at a given stage, it still remains to
determine how the ensemble of factors acts in the process which accom-
panies each physiographic change and serves as the functioning basis
for morphological differences.

In connection with the experiments on the presence of injurious
substances in bog water and bog soils, and their effect upon agri-
cultural plants, the question arose whether the toxins which are
harmful to plants in water cultures are injurious also to plants grow-
ing in soil containing them. This question has an added interest
just now, because facts like those cited above give indications that
the sterility of unproductive and “exhausted” agricultural soils may
partly be caused by some toxic substance of a similar physiological
and chemical origin. Different workers have observed that the growth
of plants often gives rise to unfavorable conditions. The data
obtained from various lines of experiments all go to prove that
“exhaustion” cannot always be attributed to the removal of plant-
nutrients from the soil by previous crops or by previous plant societies
(to). To attempt a review of the literature on this problem would
be out of place in the present paper. Suffice it to say that the results
: thus far obtained point strongly to the view that decreased physiologi-
_ (al activity of plants lies rather in the toxic condition of the soil.
The experimental proof is still regarded by many as furnishing nega-
" tive evidence upon the problem (6, 9), and hence a spirit of contro-
_ Yersy prevails in most of the writings upon this subject. However,
—itcan no longer be questioned that the solution of this inquiry is of
great importance to agriculture. It promises to throw new light upon
Many interrelations of soil and plants, and appears to afford a satis-
actory explanation of some of the problems connected with the
association and succession of plants, which on every other criterion
_ Would largely remain an enigma.

For the purpose of determining whether the toxins of bog water
‘te harmful also to plants growing in soils containing the injurious
Substances, it was decided to employ first of all a soil medium ‘as
Rearly hon-nutrient as possible. Quartz is one of the chief and most

392 BOTANICAL GAZETTE [May

nearly insoluble constituents of soil. It has been shown (2) that
quartz is of minor importance in the adsorption and retention of
hydroxids and various neutral salts; a knowledge of its action for
bog water seemed, therefore, of fundamental importance. The
quartz used was obtained from the Ceramics Department of the
university. To free it from possible impurity it was subjected to a
thorough washing. The air-dry quartz sand was first sifted through
a sieve having meshes of 1™™. Portions of about 250%™ of the sifted
material were each placed in a large porcelain dish containing dis-
tilled water acidulated with HCl. It was usually the practice to
boil the material for twenty minutes. After boiling, the supernatant
liquid was decanted and fresh distilled water was added. A similar
washing was carried out in agua regia and later again in dilute KOH.
The quartz was then washed repeatedly in boiling distilled water
and finally dried at 100° C until ready for use.

The bog water used in these experiments was collected from the
same central station on the bog island as described in the earlier
paper. The solution is relatively clear, the suspended particles
imparting to it a slight tinge of olive green to brown. — It is very little
acid to phenolphthalein, but alkaline to methyl orange.

Since no experiments had been made thus far to ascertain how
much of the toxic property of bog water is removed by a given quantity
of an adsorbing agent, series of ten cultures were prepared for this
purpose. Seven of the cultures consisted each of 400°? of bog water
to which was added sterilized quartz in quantities equivalent eo the
following volumes: 25, 50, 75, 100, 125, 150, and 200°° respectively;
that is, the quantities were chosen in volumes equal to a definite
fraction of the volume of bog water used. The bog water and the
quartz sand were shaken together in glass-stoppered bottles, and
left standing for several days. When ready for use the liquid was
decanted and placed in half-liter Mason jars, covered with black
paper. Three control cultures were added, consisting respectively
of untreated bog water, boiled bog water, and distilled water- The
wheat seedlings used for these cultures were germinated in $ :
until 4 to 5°™ high. In later experiments the seedlings wer ae
nated in quartz sand. They were then carefully washed in dist! “«
water and transplanted to the water cultures. Never less than 5

~ 1909] DACHNOWSKI—BOG TOXINS 393

seedlings were used in anv experiment. It should be observed also
that the seedlings were selected individuals out of a large number of
plants. The corks used were: previously sterilized and paraffined.
_ Growth was measured by transpiration and the green and dry weight
oiplants. The cultures stood side by side in the university green-
house in diffused light. The weekly atmometer readings varied
between 176 and 186°°. Below are given toxicity figures for bog
_ Water collected at two periods. Column I gives data for bog water
collected September 12, 1908, nearly at the end of one of the most
_ “vere droughts that have been experienced in Ohio; the bog water
for column II was brought to the laboratory October 16, soon after
- the first rains. The evidence derived from similar experiments with
- bog water collected at intervals of one month during the year is
_ omitted, showing, as it does, considerable repetition. It should be
_hoted, however, that the variation in the range of results for the
_ Sasons is considerable. |
: TABLE I
ADSORPTION OF BOG TOXINS BY QUARTZ SAND

TOTAL TRANSPIRATION FOR 15

CORRESPONDING DAYS, IN GRAMS
SOLUTION PLACE ON
CURVE
. | I. Sept..14 II. Oct. 16

© Distilled water 400c¢................ A 7.50 74
Bog wa O WMMTORIOI ss seas B 10. 26 14.90
3. Bog water Bear) Deiled oe fo ees Ce 54-22
Water 4 +asce SiO, Cc 22.70 25.50
§: Bog water 4ooce + sore BU i ees te ee 18.17
Water 400°¢ +7 5cc SiO........... E 39.28 13.83
Water 400°¢ + rooc¢ SiO,......... | gee neces eran 13.50
Pe quer’ -br2nce SiO... ...:.- G SUIS A fae ie eg
Water 4ooce + r50°¢ SiO,......... Peete ae 12.87
Bog water BOO’ + Z00c¢ SiO... 65... K 18.60 12.55

The results for these two dates have been plotted in fig. z. The
--SOwth-rate in terms cf transpiration is indicated on the axis of ordi-
Mates, and the progressive addition of quartz to bog water is shown on
* axis of abscissas.

_ Before taking up the facts brought out in this series of experiments,
st part of the investigation must be mentioned here. The
‘tegoing observations suggested the query whether results obtained
pg Soils of varying quality, fineness, and adsorbing surface would

304 BOTANICAL GAZETTE [way

show that the toxic strengths of the same bog-water solution have
approximately the same relation to each other irrespective of the
nature of the filter used. It was intended to use types of soil ranging
progressively through the weathering products from feldspars to
kaolin. But the feldspars are highly alterable minerals, and the
chemical products of feldspathic and granitic rock-decomposition

Fic. 1.—Diagram showing growth-rate of wheat seedlings in re =
The ordinates represent transpiration in grams; the abscissas show
—— of quartz to bog water. Unbroken lines for bog water pres rah

, 1908. Broken lines for bog water of October 16, 1908. Broken line single
boiled bog water; broken line double dotted for distilled water.

is Sei
ted

are extremely varied (4). In the residues, however, which remain
after leaching, free silica as quartz, and a number of rather es
substances known as clays, are the most abundant. In the .
case the efficiency of the following substances, characteristic of .
final residue of soil-forming rocks, and their allied substances WwW af
tested: SiO, coarse; SiO, fine; kaolin; CaCO;; SiC; and ri
the form of air-dried humus. The materials were obtained throug

DACHNOWSKI—BOG TOXINS 395

the courtesy of Mr. C. H. Kerr of the Carborundum Company,
Niagara Falls, New York. They are among the most insoluble sub-
stances known, and of great purity, which makes them of special
value in this investigation. The chemical analysis of these materials
‘is as follows:

: TABLE II

Material Quartz Kaolin Carborundum

ed by microscopic examination. Mechanical analysis was made
th the aid of a centrifuge and for the coarser components by means
their different rates of subsidence in water. The relative percent-

TABLE III
SiO. coarse SiO, fine | ‘Kaolin SiC
Io0o0
6.5 2.4 79-4
Bigatti WOE pee 80.8 61.8 16.6
Ree et 12.9 35-7 4-5

wa amount of the quantity of the sample having particles of these
q ‘n diameters. A mathematical calculation of the surface area of
— Martz flour, carborundum, or other crystalline bodies with irregular

396 BOTANICAL GAZETTE [May

surfaces, however, is not so readily obtained. It may be that adsorp-
tion of toxins and adsorption of vapors and gases are subject to the
same conditions (11). Perhaps by taking measurements upon the
rate of retention of a silver salt, one may secure an indirect method
for the calculation of the surface of these bodies. A curve showing
how the adsorption data are related to the surface presented by the
grains of the different soils used, though of interest, is not a ques-
tion at issue in this discussion, but it is hoped to continue this problem
further, and in a more quantitative manner.

The precaution was taken to allow contact between the solution
and the solid bodies for thirty minutes only, in order to reduce to a
minimum the low solubility of the materials (3) and the possible
action of the solution upon the solids. The amounts used in each
case, and the effect of these insoluble substances on the toxic action
of bog water collected January 30, 1909, are given below in Table IV.
The transpiration data cover a period of fifteen days and are for six
wheat plants in each solution.

TABLE IV
ADSORPTION OF BOG TOXINS BY INSOLUBLE SUBSTANCES

‘TRANSPIRATION IN GRAMS; “6
SIX WHEAT SEEDLINGS IN EACH SOLUTIO:

SOLUTION Se

sth day roth day | sth day —
1. Bog water 400°, untreated........... 4-30 8.55 3-65 rape!
2. Bog water 400°, filtered............- 4. 10.35 pa 3 ts
3. Bog water gooce+ 15¢¢ SiO, (coarse) 5-25 eet 7-35 20.85
4. Bog water gooc¢+ 25°¢ SiO, (coarse) 7.20 12.85 oe a
5. Bog water gooce+ socce SiO, (coarse) 7-00 13-55 10.5 ae
6. Bog water goore+ r15¢¢ SiO, (fine) 8.60 19.10 13-50 oy
7- Bog water 4ooc¢+ 25¢¢ SiO, (fine) 6.70 oe aon 16.00 Ft 318
8. Bog water gooce+ socc SiO, (fine) 7-68 16.40 1330 35
9. Bog water 4ooce¢+ 15¢¢ Kaolin... g.10 20.00 16.25 pry
Io ater 4ooc¢+ 25¢¢ Kaolin........ 9.65 19.32 21.58 re
11. Bog water 400°¢+ 5occ Kaolin........ 9-95 20.5% a #8 5
12. Bog water 400°¢+ 150¢¢ Kaolin. 11.98 22-30 pe 6.1
13. Bog water 4oo°¢+ 15°¢ CaCo ....... 10.90 30.59 gy Aa. 68
14. Bog water + asce CaCo,....... 10.48 | 20-70 | #90 52-52
rs r c me CaCoy.. ss. 10.07 19-17 23-35 58.08
16. Bog water 4oo°¢+ 15¢¢ SiC (fine)... : 14.23 21.25 aoe 48.60%
17. Bog water 4oote+ 25¢¢ SiC (fine).....| 10.90 | 17-50% | 27-2" | 6 60
18. Bog water gooce+ soc¢ SiC (fine)... . 11.25 a mare 76.80
19. Bog water gooc¢+ r1s5c¢ C. (humus)...| I-00 25-15 40:8 66.99
20. — water 4ooc¢+ 25c¢ C. (humus).. 8.30 16.70 pi - 55-5°
21 .
og water 400°¢ + r5oc¢e C, (humus).. 8.55 22.15 —

* 5 plants in culture.

1909] DACHNOWSKI—BOG TOXINS 397

Several facts seem to be clearly brought out in the above data. A
comparison of the toxicity figures of bog water collected at intervals
during the year indicates that the amount of toxic substances in
solution differs very appreciably within the year. In all cases the
physiological studies show that the adsorbing substances actually
remove definite quantities of bog toxins. In contrasting the efficiency
of the various amounts of adsorbents used, the important facts at the
outset are these. Different physiological phases result from the pro-
gressive increase of an adsorbing substance. The bog-water solution,
fatal in its effect at some seasons, gives an increase in growth-rate
when adsorption removes a sufficient amount of the toxic ingredient.
The effect is virtually one of dilution. Doubling of the amount of
the adsorbent brings the growth-rate into a physiological phase
marked bya greater functional activity. Further addition and its
consequent further dilution in toxicity carries with it a corresponding
intensification in growth-rate. The appearance of the plants, espe-
cially in the development of the root system, follows the transpira-
tion figures very closely (see 5, figs. 1, 4-5, p. 135). Stimulation and
tolerance rise to a maximum. But with successively larger amounts
of adsorbent, the optimum rate of transpiration can be neither
increased nor maintained. It falls off, regularly and rapidly at first,
_ Subsequently less rapidly, until the effect of the solution is practically

that of distilled water (13). Greater dilution and consequent
_ increase in rate of transpiration does not express therefore the whole
_ truth. Other and less injurious substances are also adsorbed; and
_ the rate of transpiration is seen to be the product of a coordination
_ of factors (r). In bog water with very slight toxicity, the course of
the experiment shows that the maximum acceleration phase deviates
_ Very sensibly toward the growth-rate approximated in the control
ie, the untreated bog water.

On account of the difference in size of particles, there naturally
‘lows a corresponding difference in the amount of adsorption.
: Compared with kaolin, calcium carbonate, carborundum, and
_ Satbon (as lampblack or humus), the adsorptive power of quartz
Stelatively low. It will be seen that the optimum rate of transpira-
tion of the tenth day, in solutions 3 to 8, Table IV, is soon succeeded
&minimum. This is due to the action of toxic substances still

So svar 2 Pah et eRe, ae a ier Oe a ns

398 BOTANICAL GAZETTE [May

present; for upon further addition of adsorbents the minimum at the
end of the fifteenth day is succeeded by a higher rate of transpiration.
Filters of finer grain are more beneficial, while the adsorptive power of
humus is very much higher than that ofany of the crystalline substances
used. The optimum and maximum rates of transpiration occur on
the fifteenth and twentieth day and lie near together. Reference
to the total amount transpired shows that the adsorption of car-
borundum and humus is about three to four times greater than that
of quartz. The transpiration data serve excellently as a basis for
assigning a limit to the magnitude of the toxic effect, and as an
expression of the amount of the unknown body adsorbed both in
terms of the total adsorption, and as a percentage of the surface factor
of grains. The results with CaCO, also show that the plants are not
affected by conditions of acidity or alkalinity, and that growth seems
to be more materially affected by the specific action of the organic
toxins present. Whatever the nature of the filter used, that the
increased tolerance of wheat seedlings to bog water is actually due to
the adsorptive power of the filters is sustained by the fact that the
decrease of the poisonous effects of bog water is apparently a function
of surface of particles and is approximately proportionate to the
quantity of the solid body used. The solution, decidedly toxic without
the solid, becomes capable of supporting a more than normal growth.

The outcome of these preliminary tests is, therefore, that the om
ditions giving rise to decreased physiological activity, to xerophily,
and to zonation of bog plants are not found in the depletion or increase
of mineral nutrients in bog water, nor in a low soil temperature, oe
lie rather in the toxicity of the soil substratum, i. e., in the production
of unfavorable soil conditions. brought about by the plants them-
selves.

However, experiments by the water-culture method may nol"
be serviceable as a safe basis for argument concerning soil conditions.
A number of life relations of the plant in a water culture become
changed when in the soil. In what particular manner the ae
bodies are held by the adsorptive force of the filters is not oe i
judged by physical or chemical analysis. The marked retention ©
the toxins of bog water observed may be due to causes other are
direct condensation on the surface. No experiments were made t0

t always

)
|
i

Ae ret

1909] DACHNOWSKI—BOG TOXINS 399

show conclusively that the retention is not due to chemical fixation or
substitution. The amount of solution thrown out of the quartz by
the centrifugal machine was too small to be tested. A priori, it
would seem that the filter used should be markedly more toxic now
than the solution, when tested by physiological criteria. The pres-
ence of the adsorbed bodies in the solid should not only reduce its
ellectiveness when repeatedly used for improving bog water, but should
replace normal growth by an abnormal retardation judged from the
growth-rate made in a similar check soil culture.

To obtain evidence on this point, and to contrast the efficiency of
the various constituents of agricultural soils for adsorption, a series
of experiments was made with quartz, river sand, field clay, and humus
soil. The air-dry soils were sifted through a sieve with meshes of 1™™,
Portions of 400°¢ of the sifted soils were each placed in glass-stoppered
jats containing 1200°° of bog water. The glassware employed in all
of the experiments cited was treated with a solution of potassium
dichromate and sulfuric acid, and repeatedly rinsed in distilled water
Previous to use. The mixtures of bog water and soil were left stand-
ing in the dark room for three days. To insure thorough contact
between the bog water and the soil, the solution was occasionally

_ shaken. When ready for use the liquid was filtered off, and portions
_ Of 4oo°° of the liquid from each soil type were used ds water-culture
_ &periments in the manner described above. For the investigation
_ the relative fertility of the soils used as filters, earthenware pots
_ Were used. The pots were new and each of about 300°° capacity
_ (8 in diameter, 8.5°™ deep). They were thoroughly cleaned and
: dried in an oven at 110° C, and later immersed in heated paraffin.
To each paraffined pot was added 200° of the contaminated soil

Well pressed into the bottom and sides of the pots. It was recognized
that difficulties of obtaining good contact between the soil and the

Walls of the pot would be probable. In the air space along the walls
_ ‘ually by far the greater proportion of plant roots are developed,
_ ind the wire-basket method as recommended by the Bureau of Soils

of the a; S. Department of Agriculture (Bull. 23) is, therefore, more
‘esirable. But the form of retainer here described was found to be

Wholly satisfactory. In no case were evidences found of roots growing
‘More freely at the sides of the pot than in the center. The exper-

400 BOTANICAL GAZETTE [May

ments were repeated later by the wire-basket method with the same
results. Six wheat seedlings were transplanted in a row in the soil
of each pot. In identically the same manner a series of duplicate
cultures with the untreated soils was prepared to serve for comparison
with the behavior of wheat seedlings in the contaminated soils. The
filled pots were then weighed and placed in the greenhouse where
they stood side by side. Direct sunlight was avoided by cloth screens.
Only one of the experiments need be given, and Table V gives a
summary of the results obtained with bog water collected September
12, 1908. The percentage increase is calculated upon the basis of
the quantities marked zero (9), considering them as unity for the
respective series. The photograph (jig. 2), which I owe to the aid
of Professor J. H. SCHAFFNER, shows these plants at the end of the
experiment.

TABLE V
ADSORPTION OF BOG TOXINS BY SOILS
AVERAGE LENGTH IN CM. PERCENTAGE INCREASE
SoLUTION Se
Tops Roots | Transpiration|Green weight! Dry weight

1. Bog water untreated....... 15.8 6.3 C ° 2

2. Bog water quartz-filtered 20.8 42. 338 134- 56
5. Bog water clay-filtered... 19.9 Ti.4 154 68. a)
8. Bog water humus-filtered 30.5 15.6 805 aici 7

2 inated soi 22 12:3 ° .

6. Contaminated clay soil....; 22.2 6.6 . °

gC inated humus soil...) 21.9 6.2 : -
4. Control quartz soil 24.7 9-6 8. ”™
#, Conteor Chay SOll, 75s. «. 26 11.7 23° 5°
ro. Control humus soil........ 30.7 13-5 - =

Laie»

Again it is evident that the addition of solids has increased the

tolerance of the seedlings to bog water. The improvement Was
marked during the entire period of experimentation. The presence
of the toxic bodies in small amounts exerted a noticeable stimulating
effect, while the plants in the control bog water gave every indication
that they would be unable to survive an exposure of a normal gone
period. The last-mentioned point has been repeatedly tested also
in field-work. It seems as if the roots, and especially the more minute
roothairs, of the plants in the untreated bog water served as adsorbing
surfaces. The roots are brownish in color and jelly-like in consist-

1909] DACHNOWSKI—BOG TOXINS 401

ency; deposited upon their surfaces are found numerous colored
bodies, as the result of the oxidizing action of the roots. The nature
of these bodies is still under investigation. A general decay of the
growing tips is noticeable, showing that the oxidizing action of the
plants upon the toxic substances went far toward decreasing their
harmful effect, but could not entirely overcome them. The effective-
ness of adding the insoluble solids proves, therefore, very conclusively

Fic. 2.—Growth of wheat plants in various cultures of bog water. Numbers
4s in Table V.

that the source of the harmful condition must logically be looked for
inthe solution and not in the condition of the plants themselves. The
difference in the tops as well as in the roots of the plants from the
Various cultures is very striking. The stimulation effect is less marked
in the solutions filtered through clay and humus, because of the
steater adsorptive power of these substances; yet the increase in the
sreen and dry weight of plants is relatively twice that in the untreated
bog water while transpiration has increased almost tenfold. The
introduced materials have had their adsorptive action, but it is evident

402 BOTANICAL GAZETTE [MAY

also that chemical reaction enters in the case of the common types of
garden soil.

We come finally to a consideration of the effects of bog toxins
upon soils. It is to be noted that the poisonous action of bog toxins
is more harmful when the plants are immersed in the solution, than
when grown in the contaminated soil cultures. That the poisonous
matter injurious to plant growth is present in the soils used as filters
is seen Upon comparison with the controls. Manifestly, the theory
of lack of O, in bog water or in bog soils as the cause of xerophily is
not satisfactory to account for the results, because water cultures
usually have less O, than any soil medium. T he transpiration data
for boiled bog water (Table I, page 393) are further evidence in this
direction. The inadequacy of the theory of low substratum tempera
ture is, for this locality, equally obvious. That the action cannot
be attributed to large amounts of dissolved substances has been
shown in the determination of the osmotic pressure of bog water in
the author’s earlier paper (J.c.5). The garden soils contain a much
larger amount of nutrient ingredients than bog water, and hence the.
presence of those salts should tend to increase the growth-rate. No
such increase in activity occurred. The length of time during which
the wheat plants were allowed to grow is palpably insufficient to
“exhaust” or contaminate the soils. The retardation seen in the
contaminated soils is lacking the corresponding normal average -
dry weight of plants to an amount of 18 per cent., 3 pet cent., and 36
per cent. for quartz, clay, and humus respectively. From the results
it may be concluded that the adsorption and retention capacity of soil
for toxins is generally higher the greater its content of humus. It was
shown elsewhere that a bog-water solution well aerated, or Upo?
long standing with exposure to air, lost its injurious properties.
When plants are grown in this oxidized solution it is found that the
solution becomes decidedly beneficial to plant growth. These results
are also obtained with the contaminated soils. When first used ae
exert a distinctly injurious effect. If the amount of water transpite
by the plants is replaced by bog water, the soils become more ek
Decrease in toxicity always follows aeration of the soil and a
and since the physical conditions mainly determine the gee
oxidation, these are of greater consequence in restoring the fertiity
to the soil.

1909] DACHNOWSKI—BOG TOXINS 403

SUMMARY

The available information of the study here reported may be
summarized as follows:

1. Many swamp and muck soils exhibit a sterility which cannot
be remedied by drainage or by the addition of fertilizers.

2. The sterility appears to be most marked where investigations
on the physiological properties of bog water and bog soils indicate
a greater amount and activity of bog toxins.

3- The production of bogs toxins is due to a number of physical

and chemical factors. One can only conclude that the chemical con-
stitution of bog water and bog soils at a given moment conditions
toxicity; and that the excretion from roots and rhizomes of plants
is one of the variables of the conditioning factors.

4. In untreated bog water there are found deposited upon the
roots of wheat plants numerous colored bodies as the result of the
oxidizing action of roots. ‘The general decay of the root-tips indicates
that the oxidizing activity is insufficient to decrease the harmful effects
of bog toxins.

5. It is possible that ecesis, association, and succession of plants
depends primarily upon respiration, and that in respiration bog plants
differ from other plants.

6. Treating bog water with an insoluble adsorbing agent is invari-
ably beneficial.

7. Different physiological phases result from the progressive
addition of an adsorbing substance. With coarser-grained materials

the low optimum rate of transpiration is soon succeeded by a minimum
Which is due to the action of toxic substances still present.
8. Finer-grained insoluble bodies are more beneficial. The
: Tésponse to toxic bodies when present in small amounts leads to
acceleration of growth. The period of growth is more prolonged,
_ 4nd the optimum and maximum rate of transpiration lie near
together. :
|. 9- The adsorptive action of carborundum and humus is about
four times greater than that of quartz; the capacity of soils for retain-
ing toxins is therefore higher the greater the content of humus.
Io. The decrease of the poisonous effect of bog water is probably
4 function of the surface of the particles; it is relatively proportionate
to the quantity of the solid body used.

act Prey te ph Ta eles

404 BOTANICAL GAZETTE [MAY

11. In agricultural soils used as adsorbents the presence of the
adsorbed unknown toxins replaces normal growth by’ an abnormal
retardation. Fertility is restored through aeration, that is, after time
enough has elapsed for the oxidation of the injurious bodies.

12. The contaminated condition of agricultural soils and the con- ©
sequent decreased physiological activity of the plants grown in them
still further indicates that xerophily cannot be due to acidity, lack of
oxygen, low temperature, etc., of the soil substratum; that is, the
factors heretofore cited are only in part the cause of xerophily.

In view of the evidence presented above, the writer believes that
these facts in the action of bog water upon soils justify the conclusion
that there are present in bog water and in bog soils injurious substances
which are, at least in part, the cause of xerophily in plants, and of
decreased fertility in bog soils.

Grateful acknowledgment is made to Professors McCatt and VI-
VIAN, of the Agricultural College of the university, for the facilities
of their laboratories, which were freely and courteously placed at the
writer’s disposal.

OxIo STATE UNIVERSITY
CoLUMBUS, OHIO

LITERATURE CITED
1. Brackman, F. F., Optima and limiting factors. Annals of Botany 19: 281.

1905.

2. Brices, L. J., On the adsorption of water vapor and of certain salts in aqueous
solution by quartz. Am. Jour. Phys. Chem. 9:617-649. 1995-

3- Comey, A. M., Dictionary of chemical solubilities. 1896.

4. CLARK, F. W., The data of geochemistry. U. S. Geol. Survey, Bull. 33°
1908.

5. DacHNowskt, A., The toxic property of bog water and bog soil. Bor-
GAZETTE 46: 130-143. 1908. 3:

6. Hart, A. D., Theories of manure and fertilizer action. Science N. S. 29
617-628. 1908. me

7. Horxins, C. G., anp Perri, J. H., The fertility in Ilinois soils. Til. Agn-
Exper. Stat. Bull. 123:251-255. 1908.

8. Huston, H. A., AND Brvan, A. H., Swamp muck. Rept. Ind. Agri. Exper
Stat. 1g00,

9. Kive, F. H., Toxicity as a factor in the productive capacity of s0
N. S. 27:626-635. 1908.

‘Is. Science

a) DACHNOWSKI—BOG TOXINS 405

10, LivincsTon, B. E., Further studies on the properties of unproductive soils.

U.S. Dept. Agri., Bureau of Soils, Bull. 36. 1907

11. Patren, H. E., AND GALLAGHER, F. E., Absorption of vapors and gases by
soils. U.S. Dept. Agri., Bureau of Soils, Bull. 51. 1908

12, STOKLASA, J., UND Ernst, A., Beitrage zur Lésung der Frage der chemischen

Natur des Wurzelsekrets. jane. Wiss. Bot. 46:55-102. 1

13. TRUE, R. H., AND OGLEVEE, C. S., The effect of the ieee of insoluble

substances on the toxic action of poisons. Bot. GAZETTE 39:1-21. 1905.

BRILEFER- ARTICLES

PARTHENOGENESIS IN PINUS PINASTER
(WITH SEVEN FIGURES)

In the course of an investigation on the life-history and development of
the embryo of the cluster pine, Pinus pinaster Soland., it has occasionally
been noticed that in some ovules containing proembryos in all stages
of development either no trace of pollen tubes could be seen in the nucellar
cap, or the tubes only extended through a part of the nucellus and no
nuclei could be found in them. This strongly suggested the occurrence of
parthenogenesis, but might have been due to imperfect preparations. There-
fore, in 1908, collections were made about every twelve hours during the
time when the archegonia mature, in the hope of obtaining more satisfactory
evidence.

Great care was used in fixing and imbedding this material, and the
following fixing agent has been found more satisfactory than any other,
including chromacetosmic mixtures. :

Picric acid, saturated solution in 50 per cent. alcohol, 100°; corrosive
sublimate 52™; glacial acetic acid 5°°. This fixing agent is mentioned by
CHAMBERLAIN,! and I have to thank Mr. A. J. BALLANTINE for suggesting
its use. Cedar-wood oil has been found much superior to xylol to precede
the infiltration with paraffin, as mentioned by Miss FERGUSON (704). The
stains used have been Delafield’s hematoxylin, much diluted and allowed to
act for several hours, and Flemming’s safranin gentian-violet orange-G
combination. The first named shows nuclear details more sharply than
the triple stain and is only equalled in this respect by Haidenhain’s 1ron
alum hematoxylin, which is more troublesome to use and in no way Supe
rior. In other respects the methods used have been those generally employed
in cytological work.

The evidence obtained shows clearly that parthenogenesis occasionally
occurs, and the most conclusive preparations are shown in figs. 1 and 2.

The points which seem to prove satisfactorily that the oosphere develops
without fertilization taking place are as follows:

1. Although the oosphere nucleus has divided or begun
pollen tube has not yet reached the archegonium and still contains bot
sperm nuclei (figs. 1, 2).

t CHAMBERLAIN, C. J., Oogenesis in Pinus Laricio. Bot. GAZETTE 277: 268-280
pls. 4-6. 1899. s
Botanical Gazette, vol. 47] Ue

to divide, the
h the

1909] _ BRIEFER ARTICLES 407

2. A careful study of the
other sections of the series
has failed to show any
other pollen tube which
might have reached the
, eo from another
direction.

3. The spindle of the

EXPLANATION OF

| FIGURES

| All sections 6-8 @ thick; cut
| with the Cambridge rocking
2 microtome; drawn with camera.
;
;

Fic. 4.—One of the sperm
- ‘huclei of fig. 1, showing that
a these are quite normally organ-
: ized, .

ie - §.—Diagram showing

: ial fertilization in one of

- the archegonia. hoe

Fic. 6.—S nucleus and

_ Part of the ihighere nucleus

= Of fig. 5. Xass.

* Wie. 7.—Part of ee. O.
X 1240

408 BOTANICAL GAZETTE [may

first division of the oosphere nucleus is parallel or oblique to the long
axis of the ovule and is approximately in the center of the original
nucleus. The normal spindle, on the other hand, is more or less transverse
to the long axis of the ovule and lies quite at the top of the original
oosphere nucleus. In both cases this spindle is entirely intranuclear,
as shown by CHAMBERLAIN,” and some of the original achromatic nuclear
material is not used up, but contracts considerably from the original
nuclear membrane. (This contraction may be due to the action of fixing
or other reagents, but as it is equally present in all preparations, whether
or not any contraction has occurred elsewhere, I am inclined to think it
normal.)

4. In normal fertilization a segregation of the chromosomes into two
groups occurs both in the first and second divisions of the oospore nucleus,
but no segregation can be seen here (fig. 3). The chromosomes are long
and rather irregular in shape and are often cut into several pieces and
distributed through as many sections. It has therefore been impossible
to count them accurately, but the number in the normal sporophytic nucleus
is certainly in the neighborhood of 24, and in the spindle of fig. 3 it is as
certainly less than that number. :

5. In normal fertilization a good deal of disorganization of the apical
part of the archegonium occurs, and the receptive vacuole is either broken
or considerably displaced (fig. 5). In the archegonium from which Sigs.
1 and 2 were drawn, no such disorganization has occurred, and the receptive
vacuole occupies its normal position.

6. In normal fertilization the remains of the second sperm nucleus and
the tube nucleus and the stalk cell can be distinguished for a time in the
upper part of the archegonium, but no trace of these nuclei can be found
_ in the archegonia of jigs. t and 2.

As far as has been seen, the abortion of the ovule frequently occurs
before the formation of a proembryo, but never after. A large number of
preparations of the proembryos and embryo in all stages of development
leaves no doubt on this point. Hence it appears that parthenogenetic em
bryos must develop as well as normal ones. It is impossible to say whee
this development is only intraseminal, or whether seeds containing such
embryos are able to germinate and produce normal plants. Ss

Fig. 5 shows, for the sake of comparison, a case of normal fertilization,
and the conjugating nuclei are shown in more detail in figs. 6 and 7. The
Preparation shows very clearly that the nuclear membranes are not in
* CHAMBERLAIN, C. J., Methods in plant histology. Second edition. The Uni
versity of Chicago Press. 1905.

1909] BRIEFER ARTICLES 409

contact, but separated by a thin layer of cytoplasm, as mentioned by Miss
FERGUSON.3

In fig. 5 the second pollen tube is evidently on its way to the smallest
of the three archegonia, and is taking its way laterally through the tissue of
the prothallus instead of down the canal leading to the neck.

The third archegonium is apparently sunken in the tissues of the
prothallus, but unfortunately the series is incomplete, and it may have only
a very obliquely placed neck. For the same reason it is impossible to say
whether this proembryo is really parthenogenetic, as it appears to be.—
W. T. Saxton, South African College, Cape Town.

CARNATION ALTERNARIOSE4
(WITH EIGHT FIGURES)

To a leaf-and-stem disease of the cultivated carnation, Dianthus
Caryophyllus L., our attention was called by local florists as causing serious
mage. ‘The disease, upon examination, proved to be one hitherto unde-
scribed and a laboratory study of it was undertaken.5

Symptoms.—The disease manifests itself as spots, mostly upon the
leaves, sometimes upon the stems, especially at the nodes. These spots
are strikingly characteristic, of ashen whiteness, with the center occupied by
_ 4n often scanty, though sometimes profuse, black fungous growth. The
_ diseased spot is dry, somewhat shrunken, thinner than healthy portions of
the leaf, approximately circular, though often somewhat elongated in the
direction of the longitudinal axis of the leaf (jig. 1). When occurring at
_ the node, the disease usually involves the bases of both of the leaves,
48 well as the stem between them (fig. 2). As these nodal spots age, the
disease penetrates through the stem, killing its tissue, which shrinks some-

3 Fercuson, M. C., Contributions to the knowledge of the life-history of Pinus,
_ With special reference to sporogenesis, the development of the gametophytes and
BS fertilization. Proc. Wash ington Acad, Sci. 6:1-202. pls. I-24. 1904.

- 4This termination was suggested by the authors in Annales ‘Myeolosici 7:49.
_ 1909, with the following explanation: ‘‘ We believe that much will be gained both in
Clearness and brevity by designating diseases in plants by the uniform termination
Ose’ (Lat. osus, signifying ‘full of’) added as a suffix to the genus of the causal
fungus, with or without elision of the —— syllable of the at name, in whole

Orin part, as may be determined by eu

5 Through the kindness of Dr. W. A. OrTON of the U. S. Department of Agricul-
» B. P. I., we learn that a Macrosporium disease of carnation was reported from
Sesburg, Pa., in 1906, and one attributed to Alternaria from Connecticut by CLINTON
© the same year.

Fic. 1,
one i rem
the dise;

BOTANICAL GAZETTE [May

ae Fic. 2. Diseased sag
Single leaf showing diseased spot near hago ts to stop the progress
oval of lower leaves by the gardener in his effor

1909] BRIEFER ARTICLES 4II

what and becomes soft and disintegrated, resulting in the death of the more
distal portions of the plant.

Variety of carnations affected.—A striking feature of this disease is its
tendency to infect to a large degree one variety, the Mrs. Thomas
W. Lawson, to the exemption of others. In all cases which. have
come to our notice, it has been this variety solely which was diseased;
moreover, the only records that we find of the disease imply the same
susceptibility.°

The causal fungus.—Throughout the diseased tissue of all spots occurs
in great abundance a characteristic, dark, branching, septate mycelium
(fig. 3). The surfaces of diseased spots in periods later than their earliest
youth present an abundance of black cespitose hyphae arising from the
stomata (jig. 4). Spores of the Alternaria type are found in abundance
(jigs. 5, 6), both in situ upon these hyphae, and strewn over the surface of the
diseased spots between the hyphal bases. The character and arrangement
of the hyphae are shown in figs. 7 and 8. This fungus was constantly
associated with the disease, and no other fungus was found. The pre-
sumptive evidence was therefore very strong that this fungus was the
cause of the disease. In view of the often saprophytic habit of Alternaria,

conclusions on this point would not be valid without evidence from inocula-
tions,

Inoculations.—The fungus was easily isolated by direct transfer of
spores from the diseased spots to carnation-leaf agar plates.

On October 27 numerous inoculations were made upon two plants
_ under bell jars, using small pieces of agar, bearing spores and mycelium.
_ One of the plants was left uninjured and the inoculum was placed in the
axils of the leaves; in the majority of these cases the inoculations resulted
in infection. The other plant was injured by the prick of a needle at the
_ Point of inoculation. In these cases about two-thirds of the inoculations
_ Were successful, Inoculations with spore suspension were also made upon
five branches each, of two other plants, and each was covered by a large

. lest-tube plugged around the stem with cotton to preserve a humid atmos-
Phere. Asin the former cases, the inoculations on one plant were at injured
Points, and those on the other plant were at uninjured points. The results
‘om these spore inoculations were the same as in the cases of inoculations

: With agar blocks. When these inoculations were made, others were made
_ "Pon six other plants from the same spore suspensions, but the plants were

tae

_ Rot covered or injured in any way. Following these last i g
_ Of the disease were seen, It seems from these experiments that the injured

° Orton, W. A., Yearbook, U. S. Dept. Agric. 1905:611.

412 BOTANICAL GAZETTE [May

plant is readily susceptible to infection, as is also the uninjured plant if kept
in humid condition, but that the uninjured plant in a relatively dry atmos-
phere is difficult or impossible to infect. In case of successful inoculation,
the diseased spots were well developed at the end of a week. The removal

ASS
Oc>
meen A

Showing mycelium
Showing catenulate
ptation and catenu-
hyphae.

Fic. 3. Mycelium showing branching and septation.— Fic. 4.
below stoma and hyphae emerging through the stoma.—FIc. 5-
spores as borne upon hyphae.—Fic. 6. Spores showing shape, se
lation.—Fic. 7. A young cluster of hyphae.—Fic. 8. An older cluster of

d usually in
Il with the
leaf axils,
ditions for

of the protecting bell jar from plants already infected resulte
cessation of development of the spot. These facts agree We
field observation that the most damaging infection occurs at the
points well adapted to collect and hold water, thus providing com
optimum development of the fungus.

a \\ebe ”

i ee ee ees we BT ieee |e a PB

ene

1909] BRIEFER ARTICLES 413

Culture characters.—The fungus was grown upon many different media.
Its characters upon these media have been noted elsewhere.7

The most important culture characters may be summarized as follows:
- Upon media poor in available carbohydrates the mycelium was nearly
hyaline, and the hyphae and spores pale; upon media rich in carbohydrates
the mycelium, hyphae, and spores were very dark. Upon the natural
medium the spores were more regular and uniform in shape and were much
larger than upon artificial media.

The species of fungus.—Of the Alternarias there seems to be only one,
A. longispora McAlph., growing upon members of the pink family (Caryo-
phyllaceae),® and the description of this does not agree with ours in size,
shape, or septation of its spores.

Therefore, unless an attempt be made to identify this form with some
one of the seven or more species of Macrosporium infecting the pink family,
a procedure which would be unjustifiable without resort to cross-culture
inoculations and extensive study in artificial media, this form had best be
designated as a distinct species, for which we propose the following name
and description:

Alternaria Dianthi, n. sp—Hyphae cespitose from stomata, amphige-
hous, dark brown, 1-4-septate, ascending, 1-25 from each stoma; conidia
26-123 X10-20 mw, catenulate, clavate, tapering, base obtuse, dark brown,
slightly constricted at the septa, transverse septa 5-9, longitudinal septa
0-5; spot ashen white, definite, subcircular.

On artificial media poor in carbohydrates mycelium = in color,
spores lighter, smaller, and with fewer septa.

Hasirat: living leaves and stems of Dianthus Caryophyllus, Raleigh,
N. C.—F. L. Stevens anp J. G. Hatz, N. C. Agricultural Experiment
Station, West Raleigh.

7 STEVENS AND Hatt, Variation of fungi due to environment, ined. Read before
the Botanical Society of America at the Baltimore meeting, December, 1908

8SaccarDo, Syll. Fung. 18:638.

CURRENT LITERATURE

BOOK REVIEWS
Another mushroom book

Mushrooms, by reason of their beauty and edibility, are almost as attract-
ive and popular as pretty wild flowers, and so we may expect to see popular
guides to their collection and consumption multiply. The latest candidate for
favor is a book by Mr. Harp, now superintendent of public instruction in Kirk-
wood, Mo., but for some years located in southern Ohio, where he became inter-
ested in collecting and studying: these plants. Under the encouragement of
KELLERMAN, ATKINSON, Lioyp, Moran, Peck, and other mycologists, he has
evidently become an enthusiastic amateur. By his camera, supplemented occa-
sionally by those of his friends, he has pictured a great number of representative
specimens. Presenting these photographs, to the number of 500 and more,
combined with descriptions, sometimes technical, but usually popular and more
or less diffuse, he has prepared a ponderous volume.!

This volume, chiefly on account of its excellent half-tone illustrations, which
include almost all of the common species, will be of good service to those who
wish a book less expensive and voluminous than McILvarne’s, and at the same
time comprehensive enough to enable them to identify the plants they pick up in
fields and woods.

It is evident that the author has no adequate technical training in taxonomy
or morphology; and in presenting such matters, neither his keen powers of obser-
vation nor his enthusiasm could prevent him from falling into errors both of form
and fact. The typography of the book shows, also, that both author and pub-
lisher are unacquainted with scientific practice, while the proofreader and the
author alike are responsible for many typographical errors. The etymology of
the scientific names, by which the author hopes to show their significance to those
unaccustomed to them, is often erroneous and occasionally ludicrous. The

ossary does not define all the technical terms that are used, no less than four 1p
a single description of eight or ten lines having been hit upon by mere chance.
The list given of authorities for generic and specific names is far from complete,
so that abbreviations used in the body of the text (which are not consistent) could
not possibly be identified.

* Harp, M. E., The mushroom, edible and otherwise; its habitat and its time of
growth, with photographic illustrations of nearly all the common species. - :
to the study of mushrooms, with special reference to the edible and poisonous be gare
with a view of opening up to the student of nature a wide field of interesting and es
knowledge. 4to. pp. x+609. pls. 60. figs. 504. Columbus, O.: The Ohio Library ©°-
1908. $4.75.

414

1909] _CURRENT LITERATURE 415

All these things show the apprentice hand; but, though they mar the book,
they do not so detract from its value that it may not be commended to the public
for whom it is intended. It will indeed be a welcome addition to public, school,
and college libraries, where there is always a demand for well-illustrated books
of this kind, and it will probably do good service in awakening an interest in mush-
Tooms. It certainly treats wisely the matter of testing the edibility of mushrooms
_ and no one who follows Mr. Harp’s advice will come to harm. Thus it has a
teal field of usefulness. But it is not for the mycologist; and, unless “of its kind”
_ is a saving clause, it is by no means what is claimed by the publishers in their
_ circular—‘‘By far the most complete work of its kind ever attempted in this
meeountry.”—C. R. B.

Trees and woods

The fourth volume of the series on Trees by the late H. MARSHALL WaRD
has recently come from the Cambridge University Press.? With a fifth now in
Press the series will be concluded, for although the author had planned another, it
Was too inchoate to permit publication. The present volume has been issued
under the editorial supervision of Percy Groom, who has left the manuscript
_ Practically unchanged, but has had the labor of selecting the illustrations, which

. th numerous and appropriate.
As the three preceding volumes have treated respectively the buds and twigs,
_ the leaves, and the inflorescences and flowers, this one presents the fruits. After
4 general discussion of the morphology of fruits (part I, 59 pp.), the second
Part (94 pp.) gives a key to trees and shrubs, based on characters derived from
fruits, and accompanied by figures of most of the species. Like the other parts

The eighth part of SCHNEIDER’s Handbook of deciduous trees (the third section
the second volume) has lately issued from the press.3 Like its predecessors,
uently referred to in this journal, it presents, in the most compact form
ible, descriptions of the species of angiospermous trees, native or planted
in central Europe, arranged in the sequence of a dichotomous key, and
: illustrated freely. It seems a most thorough and practical book, but somewhat
certing as to nomenclature. Who of our foresters will give us something
88 good, but perhaps a little less condensed ?

? Warp, H. M., Trees: a handbook of forest botany for the woodlands and the

laboratory. ‘Vol. IV, Fruits. Cambridge grec Series. 12mo. pp. 1v+161. figs.
" 147. oe University Press. 1908. New York: G, P. Putnam’s Sons. $1.50.
_, $ScHnerper, C. K., Illustriertes Handbuch der Laubholzkunde. Charakteristik
der j In Ribctcurore Sten ieicboes und im Freien angepflanzten angiospermen Gehdlz-
und Formen mit Ausschluss der Bambuseen und Kakteen. Achte eaagg et:

: dritte Lieferung des zweiten pees Imp. 8vo. pp. 241-366. figs. 166-248. Jena
Gustay Fischer. 1909. M4

416 BOTANICAL GAZETTE [May

The second part of JANssontus’ elaborate micrography of the woods of
Java* has recently come from the press. In the notice of the first part of this
works we described the plan, which is here merely extended. The part includes
in the luding section of the first volume (Dicotyledones, Polypetalae, Thalami-
florae) descriptions of 67 species, in addition to the 108 of the first section. The
second volume begins the Disciflorae and presents 54 species. The complete
work will certainly be a monument of industry and will be serviceable for the
microscopic identification of Javanese woods. That the game is worth the
candle, we may be permitted to doubt.—C. R. B.

Microscopy of technical products

Now that so much attention is being paid to the purity of foods, drugs, and
manufactured products of all kinds, it becomes of the greatest importance to
have adequately trained men in municipal, state, and national offices, to whom
can be submitted the many questions that are sure to arise as to the adulteration
or sophistication of marketed articles. Unfortunately the number of competent
persons is far short of the demand, and this state of affairs is sure to continue
for some time. In this situation the only recourse is to have accurate handbooks
in which may be found detailed descriptions of the characteristics, chemical and
microscopic, of all the commoner substances which enter into commerce. Then
one who has a reasonable familiarity with microscopic manipulation may be
able to determine the more obvious cases of adulteration, and by experience
may acquire real expertness.

To put into reach of American laboratories one of the most valuable of
foreign works, Dr. A. L. Wino, chief of the Chicago Food and Drug Labora-
tory of the Department of Agriculture, with the assistance of Dr. KATE G. BARBER,
has translated an edition of HANAUSEK’s Lehrbuch der technischen Mikroskop#,
which represents the last German edition extensively revised by the author.
The translators have also included a considerable amount of new material, an
in particular they have incorporated into the key to economic woods—a most
valuable feature of the book, permitting one to determine most of the species
from a fragment of the wood—the American species of commercial importance.
The illustrations have also been improved and augmented.

4 Janssonius, H. H., Mikrographie des Holzes der auf Java rot
javanicae auctoribus S. H. Koorpers et TH. VALLETON.
Vol. I, pp. 369-568. Vol. II, pp. 1-160. figs. 45-95. Leiden: E. J. Brill.

5 Bot. GAZETTE 43:345. 1907.

cogni
» Zweite Lieferung-
ill. 1908.

Revised by the
tion of KATE G.
& Sons. 1997:

° HanausEK, T. F., The microscopy of technical products.
author and translated by ANDREW L. WinTON, with the collabora
— Imp. 8vo. pp. xii+471. figs. 276. New York: John Wiley

4-75:

1909] CURRENT LITERATURE 417

The work opens with a section on the construction and use of the microscope,
its accessories and reagents. The important types of technical products that are
treated are the following: starches and inulin; vegetable fibers, including hairs,
with a section on the examination of paper; animal fibers, mineral fibers, and
_ textile fabrics; stems and roots, including woods (gymnospermous, dicotyledonous,
_ and monocotyledonous), barks, and rhizomes, with some practical examples of
the problems that are submitted for solution; leaves, under which only sumach
leaves are treated; flowers, with insect powder alone treated; seeds and fruits,
including a large range of oil cakes; and finally teeth, bone, horn, etc.

course in such a list there must be an end somewhere, for space is not
unlimited; but one wonders at the basis of some choices. The line between
drugs, foods, and technical products is not an easy one to draw; but if wheat
and barley appear among the fruits, why not maize and rye? If sumach leaves,
why not tea and tobacco? If insect powder, why not saffron? But it behooves
us to be thankful for what there is, rather than to complain of what there is not.
And what there is is sure to be thoroughly helpful.

The publishers’ part has been well done. The illustrations are well printed,
the text clear, and the binding substantial. The book is See for public
libraries and for governmental and university laboratories —C.R

Works of Léo Errera

We have already noticed in these pages the sumptuous republication of the
work which, under the direction of Lfo ERRERA, issued from the botanical insti-
tute of the University of Brussels. In these volumes’ his own original work takes
&conspicuous place. But he did much other writing, popular, pedagogic, philo-
ES sophic, literary, which is to be preserved by original publication or reprinting
in a series of six volumes now being issued under the title Recueil @’auvres de
_ Léo Errera. Of these three have appeared. Two deal with botanical subjects
and one contains verse and prose on a variety of topics—addresses, thoughts,
Philosophic epigrams, etc. The botanical topics of the first volumes are: A letter
0 the vegetation about Nice; Agriculture and horticulture in Norway (largely
_ &criticism of ScHtipeLeR); Structure and modes of fecundation in flowers (200
_ Pp); Secondary heterostylic characters of primroses (a posthumous work com-
_ Pleted by Miss J. Wery) (30 pp.); Progress of systematic botany; A neglected
__ field of research (the efficacy of the defensive structures of plants); ENGELMANN’S
bacterial method; Compass plants. In the second volume we find: Respiration
A of plants (one of a course of public lectures); De grace, des noms latins (a plea
is for the avoidance of vernacular names); Scientific bases of agriculture (36 pp. :
a

, sae de l’Institut Botanique Léo Errera. Bot. GAZETTE 432215, 347- 1997;
45:201. 1908.

: Ss d’ceuvres de Lféo ERrERA. 8vo. Vols. I, II, tie Général. pp.
W+341. Vol. VI, Melanges (vers et prose). pp. xiv+ 222. Bruxelles: H. Lamertin.
1908,

418 BOTANICAL GAZETTE [MAY

Descriptive text of physiological charts (90 pp., including small reproductions of
the charts which he published in conjunction with LAURENT); Letter prefatory
to DE WitpEman’s Flore des Algues de Belgique; An elementary lesson on Darwin-
ism (106 pp.; an admirably clear and brief presentation, which appeared first
in 1900 and is now printed as he had revised it for a third edition). This volume
closes with three posthumous articles: Plants in contrast with other beings;
What there is in a plant; The épopée of a ray of sunlight.

These volumes, as well as the more strictly scientific ones, will form a worthy
memorial of this distinguished savant, whose writing is always luminous and
inspiring. His bibliography, though voluminous (287 titles, as we learn from an
interesting biography just published®), is remarkable, not alone for its extent, but
for its value. To have all his work collected is a real boon.—-C. R. B.

NOTES FOR STUDENTS

Papers on mucors.—T'wo valuable papers, largely taxonomic in character,
~ have recently appeared on the mucors. In two ways they show an advance over
other taxonomic work in this confused group. In the first place the center for
fungus cultures maintained by the Association I nternationale des Botanistes has
been made use of, and the species investigated were compared as far as possible
with named cultures from this and from other sources. Provided contamination
of cultures in the source of supply is avoided, this center in Amsterdam affords
a ready method of checking up determinations and should be of increasing value
to mycologists. In the second place the differentiation of species according to
their sexual character into homothallic and heterothallic forms is recognized as an
item in the classification, and in heterothallic species the production of zygospores,
when a given strain is grown in contact with the opposite strain of a known species,
is used to establish its specific identity with the form tested.

Hacem'? announces his paper as a preliminary contrib
mucors. By exposing Petri-dish cultures to the air and allowing the
fall on them to develop mycelial colonies, he finds with Sarto that spores ©
mucors, both absolutely and relative to other molds, are une 2
quent occurrence in the air. Only seven species were thus found. In investigat-
ing the mucor flora of the soil, samples from different kinds of soils were sown
on various nutrient substrata, and the resulting growths isolated in pure cultures.
Sixteen different species, confined to the genera Mucor, Rhizopu soe
Zygorhynchus, were found, of which six are described as new, viz., Mucor sirien’s
M. sphaerosporus, M. griseo-cyanus, M. silvaticus, M. norvegicus, and
glauca. Four new forms are added to the list of heterothallic spectes:
these, Mucor hiemalis, was especially investigated as regards the distrib
Shine coaches era, membre de

ution in

9 FREDERICQ, LEON, AND MAssART, JEAN, Notice sur Lito =
Academie. 12mo. pp. 153. Brussels: Hayez. 1908. ‘densk.-
1° HaceM, Oscar, Untersuchungen iiber norwegische Macon’ oa
Selsk. Skrifter. I. Math.-Nat. Kl. No. 7. pp. 5°- 1908.

~ 1909] CURRENT LITERATURE 419

nature of the two sexual races. From a total of 52 separate isolations, 21 were of

one sex, 5 of the other sex, and 26 (50 per cent.) failed to give any reaction with the

test strains and were listed as neutral. Three of the strains that took part in

zygospore formation were very weak in their sexual activity, and one of them
under further culture entirely lost its power to take part in zygospore formation.
The distribution of the sexual races in this species is thus shown to be in accord
with the condition in Rhizopus, where out of 59 strains investigated by the reviewer,
Ig were (+), 27 (—), and 13 neutral. The large percentages of neutral races
thus established for these two species, together with the reviewer’s own experience
with neutral races in other heterothallic species, renders it probable that sexual
neutrality is a widespread phenomenon among the mucors. There is little at
present known to indicate its cause or significance.

LENDNER,"' in his studies of the Mucorineae of Switzerland, has not confined
himself to mere local species, and though he has not attempted to present an
exhaustive treatment of the whole group, he has given us a more or less critical
arrangement of the genera Mucor, Rhizopus, and Absidia. In these three genera
keys for the determination of species are given, and each form is described, either
from the original description or from A. FISCHER, with additional notes on such
species as he had himself cultivated. In classifying the genus Mucor, FISCHER’S
division into the unbranched, racemosely branched, and cymosely branched
groups is followed. Fifty-one species are recognized, of which seven, M. /ausan-
nensis, M. genevensis, M. pirelloides, M. lamprosporus, M. J anssenl, M.

:
:
i
f.

Glomerula, are reduced to the genus Mucor, as also VUILLEMIN’S Zygorhynchus.
The genus Mucor is the Crataegus among fungi_and will probably always remain
ataxonomic playground for mycologists. One might imagine that early system-
tists used the genus as a group to practice on, and their one- or two-line descrip-
tions are frequently hardly sufficient to tell us whether the form described is a
‘Mucor or a myxomycete. Such supposedly common forms as Mucor Mucedo
and M. racemosus among others, it is impossible to determine with any degree of
Accuracy, and therefore these designations can be considered hardly more than
group names. We cannot but have considerable charity toward one who feels
Inclined, in consequence, to disregard the stock names, but when each mycologist

who works on the genus gets out a list with names of his own, the result is confus-
«Iigtoa degree. Moreover, species shown to be distinct by the reaction between

a distinguish them. LENDNER has done a service in bringing together the descrip-
tions of species since FISCHER’S publication. We are grateful that he has not
a found it necessary to make new species out of more than 15 per cent. of the 51 forms

: 't LENDNER, ALF., Les Mucorinées de la Suisse. Matériaux pour la flore crypto-
Bamique Suisse. Vol. III, Fasc. I. pp. 180. 1908.

420 : BOTANICAL GAZETTE [May

listed. Seven circinellas are described, of which C. minor and C. aspera are
given as new. In the genus Rhizopus, of which 22 species are recognized,
physiological characters, such as ability or inability to grow on potato above 39°C.

and power to ferment different carbohydrates, are used in addition to the usual -

distinguishing morphological characters. Material received from the Amsterdam
center under the name of Mucor norvegicus is identified as R. nodosus. _ Seventeen
species are recognized in the genus Absidia, of which A. spinosa, a homothallic
and heterogamic species, is described as new. In addition to the forms from the
genera mentioned, Cunninghamella elegans is described as new.

In addition to the systematic part of 113 pages, an introduction of 47 pages is
devoted to methods of isolation and cultivation, and to a discussion of the sexual
reproduction in the group, together with the results of a cytological investigation
of the formation of zygospores. It might be expected that forms in which the
sexual differentiation had extended to the separation of distinct male and female
races would show a differentiation in the uniting gametes. In no heterothallic
form, however, has there been shown to be any constant difference in the size
of the gametes, such as occurs in a few of the homothallic species, where, since the
zygospores are produced between neighboring filaments of the same plant, a a
specialized sexual condition might be supposed to exist. In Absidia Orchidis,
LENDNER finds that the circinate outgrowths, which typically arise from both
suspensors, are at times produced from but one, which has been cut off from the
large progamete that he considers female. This he claims an indication of sexual
differentiation, as also the frequent inequality in the gametes of Rhizopus. From
these facts he concludes that the (+) and (—) races are potentially homothallic,
but with the opposite sex more or less completely suppressed. The suggestion that
the sexual races may be potential hermaphrodites is in line with our knowleage
of higher forms, but to formulate this as a conclusion and to claim i
and larger gametes formed by a single sexual race are male and female respectively,
as LENDNER would imply, is certainly going beyond the facts in hand. 4H€
reviewer has shown that in Rhizopus the larger gamete is derive
the (+) and sometimes from the (—) plant, and that similarly in th .
species of Phycomyces the outgrowths (which LENDNER, P- 38, wrongly says arise

times to the
(—) suspensor. The inconstant difference in size of ga Soe
growths from the suspensors in Absidia Orchidis is probably merely n bat
character and of no sexual significance. In A. Orchidis also, is figured W "
appears to be an arrested stage in the formation of a zygospore between two ou
growths from the same suspensor, and therefore belonging to the same Sf fact
race. If this is used as an argument for the contention just mentioned, the 2
should be established beyond doubt. Even if the author were not aie
the terminations of the filaments apparently in conjugation, which wou”
difficult to follow in a tangle of other filaments, these two arreste ga
be thought to have arisen adjacent to each other at the stimulus of conta :
third branch, which came from the opposite sex but had remained in only temp?
rary contact with them.

that the smaller

1909] CURRENT LITERATURE 421

No two investigators are as yet in accord as to the cytology of the zygospores
of the mucors. GRUBER, who apparently has done the most careful work on the
zygospores of Sporodinia, was unable to find a fusion of nuclei at any stage in their
formation or maturation, and several of the most experienced of American and
European cytologists of the fungi have personally told the reviewer that they also
have investigated the zygospores of this same species, but with no better results.
In 1906, DANGEARD, working with Mucor fragilis, described the uniting cells as
gametangia and saw a fusion of nuclei in pairs soon after the union of the two
sexual cells. The condition in Sporodinia was more difficult to follow, but Dan-
GEARD believed he was able to find the same condition in the zygospores of this
species. LENDNER, in the work before us, criticizes the conclusions of DANGEARD,
daiming that the figures which DANGEARD interprets as stages in fusion are in fact
stages in division, since they occur at the same time in the two suspensors as well.
What DancEarp considers as degenerating supernumerary nuclei toward the
periphery of the zygospore, LENDNER never finds in degeneration, and he believes
them to be in this position to preside over the formation of the membrane. The
teal sexual fusion, according to LENDNER, is between two large nuclei which ap-
proach the center of the zygospore. The two densely staining bodies in the fusing
nuclei, which are homologized with chromosomes, give at first four bodies in the
fusion nucleus, that eventually are reduced to two and finally unite into a single

mass. Neither DANGEARD nor LENDNER has studied the germination of the
_ 2ygospores.
Since Kiess showed that external factors are responsible for the form of repro-
_ duction in Sporodinia and many other fungi, the influence of external conditions
_ upon the growth and reproduction of individual species has become a favorite
_ Subject of investigation. As the reviewer has shown, external conditions are more
_ influential in determining the form of fructification in the two homothallic species
_ of Sporodinia and Dicranophora, found growing on fleshy fungi, than in the homo-
ee thallic species, Zygorhynchus Moelleri, recently investigated by WISNIEWSKI,"?
“Apupil of RactBorski. It seems to be generally true in regard to the influence of
external conditions, that the limits within which zygospore formation is possible
are narrower than those within which sporangial formation occurs. WISNIEWSKI
finds that, although under ordinary conditions sporangia are formed together
i zygospores on the same mycelium, extreme conditions may suppress: the
_ Production of zygospores, while sporangia are still formed. On pure agar below
: 5°C. and on the same substratum in direct sunlight, only sporangia will be pro-
duced. (It may well be the heat rather than the light effect of the direct rays
of the sun that is here influential.) Under all other conditions examined, both
-_2ygospores and sporangia were produced together, i
telative abundance of zygospores is associated with a checking of the rapidity of
growth of the mycelium. ‘The transpiration is shown to have no effect upon the
2

uchtform bei

12 WIsNIEWSKI, P., Einfluss der ausseren Bedingungen auf die Fr
Nat. 1908:

Zygorhynchus Moelleri. Bull. Acad. Sci. Cracovie Cl. Sci. Math. et
656-682,

422 BOTANICAL GAZETTE [MAY

rapidity of growth, and it is assumed that for this reason increase or decrease of
moisture in the surrounding air does not affect the proportion between zygospores
and sporangia. Upon certain substrata, zygospores are more abundant at the
junction of adjacent mycelial colonies, forming dark lines. The conditions
governing their production have not been investigated.

For several years DAUPHIN has been interested in the genus Mortierella. In
a recent paper on the genus he gives in 28 pages a systematic arrangement of the
species as an introduction to a special study of M. polycephala. The original
descriptions and figures are given for each of the 29 species and varieties dis-
cussed. Two new species, M. canina and M. raphani, and one new variety, M.
van Tieghemi var. cannabis, are described. The genus is divided into four group
species (grandes espéces) with subspecies (petites espéces) and varieties under
them. A well-arranged key would have added to the value of this part of the work.
M. polycephala is the only one of the Mortierellas which has been investigated in
regard to the influence of external conditions upon the production of zygospores.
In this species, which forms the subject of the physiological part of the paper,
Davurutn has succeeded in finding the zygospores, and since he obtains them from
sowings of single spores, he classifies the species as homothallic. This being the
case, it seems strange that they have not been found in this form by other investt-
gators, since as yet no neutral strains have been found for homothallic species.
The optimum temperature for germination of the spores is placed at 27° C., and
the optimum for formation of sporangia and zygospores, between 15 and 20° C.
Germination and growth are checked by darkness, but the fructifications are not
altered. Light increases the rapidity of development, but if too intense causes

the fructification to be confined to stylospores. The violet and ultra-violet rays of,

the spectrum seem necessary for the germination of the spores. X-rays and he
influence of inhibitive to germination and growth, the radium in addition
causing the production of cysts in the hyphae. Moisture in the surrounding air 1s
shown to be necessary for the germination and growth of the fungus. Perfect
development will take place in an atmosphere completely free from oxygen A
decrease, however, of atmospheric pressure below 150™™ causes the mycelium t
remain sterile. An increase of pressure above atmospheric checks the growth ©
the ‘mycelium without preventing the normal fructifications.
like other mucors, develops poorly in liquid media. The monosaccharids, oR
cially dextrose and levulose, were found most favorable of the carbohydrates ed
formation of zygospores and sporangia. The influence of different concentration
of the nutrient was little investigated, but stylospores and sporangia were ae
duced without zygospores when the amount of dextrose was increased as
per cent. to 60 per cent. The purpose of this part of the paper seems to have .
to find out the influence of a large number of more or less isolated external gor
ditions upon the form of fructification, rather than to work out thoroughly
influence of a few closely related factors.

: a + Nat. Bot
13 DAUPHIN, JEAN, Contribution a |’étude des Mortierellées. Ann. Sci. N
IX. 8:1-11r2. 1908

CURRENT LITERATURE 423

It is hardly necessary in a botanical publication to comment on a recent note

In Science, entitled Mucor cultures.+ The author states that “‘in the study of
the Mucoraceae for several years, some interesting facts concerning the develop-
ment or rather non-development of zygospores were observed. n a thousand
cultures of Rhizopus nigricans, made from material collected by the author or sent
him by friends, as well as in five hundred specimens found growing spontaneously
in different places, and in about five hundred other unrecorded observations of the
fungus (a total of two thousand observations), no zygospores were found. Inocu-
_ lations were made on a number of different media, including HAMAKER’Ss corn-
_ bread-muffin combination, and the growth of the cultures was tested in closed
_ jars in H, N, and CO,. No zygospores were obtained, and the conclusion is
teached that ‘‘the absence of oxygen is not a necessary condition for the growth
of zygospores.”’ It is a pity that one who has the time to make observations on
2000 cultures should not have taken the trouble to read the recent literature on
the subject, to learn in what part of a culture zygospores are produced and under
what conditions their formation is possible, and so be in a position to make a con-
tribution of some value. Additional information in regard to the relative distri-
bution in nature of the two sexual strains of this most common of molds might
have been the fruit of so extended a series of observations
Appended is a list of species, the thallic condition of which has been deter-
‘Mined, arranged according to the type of their sexual reproduction. Following
tach species is given in parentheses the name of the author who has investigated the
4ygospores and determined the sexual condition of the species in question.
Homothallic Heterothallic

Sporodinia grandis (BLAKESLEE) t. Mucor Mucedo (BLAKESLEE)

RE erte Fee es To a

4 Mortierella ae seca 4-0. Mucors TH-Vir (BLAKESLEE)15
Heterogamic ve

6 Dicranophora (BLAKESLE

‘1. Zygorhynchus Moelleri ee EE)
erm chus heterogamus (BLAKES-
4g ESists spinosa (LENDNER)

17. Cunninghamella echinulata (BLAKES-

LEE)
18. Choanephora cucurbitarum (BLAKES-

LEE
1g. He woe piriforme (BLAKESLEE)
20. Sync TT alastrum (BLAKESLEE)

21 R N, n. gen. (BLAKESLEE)

—A. F. BLAKESLEE, Storrs, Conn.

oe Sumsting, Davip R. Science N. S. 29:267. Feb. 12, 1909
'S Perhaps some of Mucors III-VIII are identical with species already in the

424 BOTANICAL GAZETTE [MAY

Mesozoic Equisetales.—One great desideratum in discussions as to the origin
of existing plants is an increase in our knowledge of those of the Mesozoic. At
the present time the Paleozoic flora is much better understood than that of the
intervening period, which gave rise to the characteristic groups of our existing

ora. A contribution by HALie’® throws a good deal of light on the organizatiun
of the equisetum-like forms of the earlier Mesozoic (Upper Triassic and Lower
Jurassic}. The author describes the vegetative stems and cones of several Equi-
setales. He establishes a new genus, Neocalamites, which has the general habit
of the Calamites, including the leaf whorl made up of ununited leaves, with the
herbaceous texture of the existing equisetums. It further resembles Calamites
in the fact that only every second internodal strand (or fewer) gives off a leaf
trace in the region of the node, and in the fact that the internodal bundles are
frequently continuous at the nodes, in contrast to the alternating condition found
in Equisetum. In Equisetites the leaves are in united sheaths as in the living
genus, but in some of the species described by the author the same continuous
bundles, and leaf traces fewer than the internodal strands, as are found in the
Paleozoic Equisetales, are described. In the smaller branches, however, the leaf
strands correspond to the number of internodal strands, thus foreshadowing the
condition found in the living Equisetum. Perhaps the most interesting feature
of this important addition to our knowled ge is the description of the cones, cone-
scales, and spores of Equisetites. The two former do not differ strikingly from
those of the living genus, but the spores, interestingly enough, show the absence
of elaters and the presence of triradiate sculpture described for the megaspores
and microspores of the Calamites. The cones are isosporous. This article COR
nects in a very satisfactory way the organization of the Paleozoic Equisetales with
that of those still living, and illustrates the important bearing of paleontological
facts on any stable scheme of evolution —E. C. JEFFREY.

Membrane of diatoms.—MAncrn presents'? an account of some extended «
observations on the diatoms, especially those of the plankton. His most impor-
tant observations relate to the membrane. This he finds to be composed of 2
Substance identical with pectic compounds, combined more or less inti
with silica; the siliceous skeleton thus formed is impregnated and invested wi
a gelatinous membrane which often hides, at least in plankton species, the charac-
teristic ornaments. He controverts the ideas of Scuiirr as to the growth of e
membrane (through agency of an extracellular plasma), which he discusses @
some length; and after describing improved methods of staining the eee
(by ruthenium red, and by an old solution of hematoxylin with annoy a
Tuthenium-alum, which may be aged artificially), he gives some detailed exam
ples in the study of certain species.—C. R. B.

: . Kung:
ALLE, T. G., Zur Kenntniss des mesozoischen Equisetales Schwedens

1
Svensk. Vetenskapsakad. Handl. 43: No. 1

‘7 Manon, L., Observations sur les Diatomées. Ann. Sci.
219. figs. 14. 1908.

Nat. Bot. IX. 8:127~

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CONTENTS

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Issued June 19, 1909

LVI CONTENTS FOR JUNE 1909 No. 6

IBUTIONS FROM THE ROCKY MOUNTAIN HERBARIUM. VIII. Aven Nelson 425
LEAVES OF PODOPHYLLUM. /. Arthur Harris . - - - - - - 438

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VOLUME XLVII NUMBER 6

BOTANICAL GaAzerTe
JUNE 1909

CONTRIBUTIONS FROM THE ROCKY MOUNTAIN
* HERBARIUM. VIII
AVEN NELSON
In 1905 Mr. LESLIE N. GooppING again made some collections
in the deserts of southern Nevada and adjacent Arizona. These
plants, like those of his collections of 1902 and 1903, were deposited
with the Rocky Mountain Herbarium at the University of Wyoming,
to be named and distributed. Interesting as the earlier collections
proved, these which came from even more inaccessible places and
from regions which represent the extremes of aridity and heat are
equally so. I submit herewith descriptions of the new species and
notes upon others that are but little known or that seem to need
generic differentiation.
PLANTAE GOODDINGIANAE

Calochortus comosus, n. sp.—Glabrous: bulb small, 8-12™™ in
diameter, covered with dead sheathing flaky scales which also invest
the base of stem: stem very slender, slightly flexuous, 1-2°™ high,
1 (rarely 2)-flowered: leaves narrowly linear: sepals lance-linear,
as long as the petals: petals pale lavender or lilac, with darker lines,
but not marked with spots or bands of other colors, broadly triangular-
ovate, 18-25™™ long; the apex subtruncate, slightly undulate-den-
tate; the gland large, ovate, inordinately densely long-bearded with
yellow hairs which are glandular-thickened toward the apex; similar
hairs are scattered over all the lower half of the petal: anthers white,
acute, 7-8™™ long; the filament as long: capsule acute-angled.

Las Vegas, Nevada, in limestone washes, May 8, 1905, Goodding 2323.

Probably most nearly allied to C. flexuosus Wats., from which it is readily
distinguished. .

425

426 BOTANICAL GAZETTE [JUNE

Mirabilis limosa, n. n—M. glutinosa A. Nels., Proc. Biol. Soc.
Wash. 17:92. 1904; not M. glutinosa Kuntze, Rev. Gen. 37:265.
1898, the latter a Bolivian plant; Hesperonia glutinosa (A. Nels.)
Standley, Contrib. U. S. Nat. Herb. 122365. 1909.

Lesquerella tenella, n. sp.—A delicately slender erect annual,
2-4°™ high, beginning to blossom when very small; uniformly but
not densely stellate pubescent throughout: stems one or more from
the summit of the slender tap root, unbranched: leaves rather distant,
narrowly oblanceolate to linear, 1-4°™ long, usually tapering to a
slender petiole: petals broadly spatulate, very obtuse or slightly
retuse, about 8™™ long, twice as long as the lanceolate sepals: pod
globose, not stipitate, about 5™™ long; the style about 3™™ long;
the pedicel 1o-15™™ long and variously curved and spreading or
even reflexed.

Moapa, Nevada, April 8, 1905, Goodding 2184.
This species belongs in the section having immarginate seeds and with the
annual species having globose pods; however, it has no known near relative.

Linum leptopoda, n. sp.—Having the appearance of a perennial
but possibly only biennial, wholly glabrous, 34™ or more high: root
stout, with white furrowed bark, the caudex-like branched crown
bearing several to many slender erect terete stems dividing into filiform
branchlets above: leaves crowded below; those of-the crowns scale-
like, very short, 2-4™™ long and half as broad, leaving a crestlike
scale when falling away; lower stem-leaves 5-12™™ long and 1-3™™
wide, becoming narrower, more distant, and finally bractlike above;
stipular glands wanting: pedicels slender, 5-25™™ long: sepals
about 5™™ long, green or brownish red, lanceolate, 1-nerved, with a
few obscure glands on the margins as have also the bracts: petals
a clear yellow, 7-9™™ long, broadly obovate or suborbicular, with
obscurely crenate summit: stamens as long as the united part of the
styles; the anthers large, as long as the filaments: styles united a
little more than half their length: capsule about as long as the sepals,
the false septum incomplete.

Las Vegas, Nevada, on stony slopes, May 4, 1905, Goodding 2276.
Except for the yellow flowers, this suggests at first glance Linum Lewis
rather than any of the yellow-flowered species.

_ 1909] NELSON—PLANTAE GOODDINGIANAE 427

_ Mortonia utahensis, n. sp—M. scabrella utahensis Coville, ex.
_ Wm. Trelease, Syn. Fl. N. A. 1:400. 1897.

: Specimens in full bloom, April 19, no. 2230, in the Muddy Range; fruiting
_ specimens May 13, no. 2369, Las Vegas Mts., both localities in southern Nevada.
q The excellent collections of this plant by. Mr. Goopprnc in 1905 leave no
_ doubt that it ought to be considered distinct from M, scabrella Gray. Not only
_ are the leaves uniformly much larger, but the flowers are very numerous and much
_ smaller. The calyx-lobes equal and in fruit rather exceed the turbinate tube;
; _they are broadly triangular, with erose scarious margins. The petals are nearly
_ oval, with erose denticulate margins, and are narrowed abruptly to a short claw-
like base; they barely equal the calyx-lobes and are never more than 3™™ long.
af e filaments are dilated downward. The leaves do not have revolute margins
a but are fleshy-thickened, looking as if bordered with cartilage.

Condalia divaricata, n. sp.—An intricately branched rigid shrub,
2-4™ high; the branchlets crowded, 3-7°™ long, strongly divaricate
(at right angles to the stem), rigid, permanently covered with a fine
close tomentum which at tip shades off to flaky deciduous scales,
_ leaving the sharp brown spine free: leaves in approximate alternate
_ fascicles, mostly on the spinelike branchlets, nearly sessile, some-
what thick and coriaceous, the margin wholly entire, permanently
lanate-pubescent, oblong to oval, or sometimes narrowed toward the
very short petiole, mostly obtuse at apex, the venation obscure, rarely
more than the midrib showing, 4-8™™ long: flowers not known:
mature fruit on very short peduncle and pedicels (each less than 2™™
long), the umbel apparently 3-flowered at most: drupe ellipsoidal,
purplish black, 5-7™™ long (in dry specimens), the stone moderately
thick-walled, completely 2-celled, with a large elliptical plano-convex
seed in each cell.

Las Vegas, Nevada, in mature fruit, May 5, 1905, Goodding 2300.

The only species to which this is closely allied is C. lyctoides Weberbauer, but
that is a Texan species with narrower thin pale leaves conspicuously veined, and
with subglobose fruit. The var. canescens of this species, at least as originally
described (Gray, Wheeler Rept. 6:82), is a shrub only half as large, with greenish
branches covered with a “gray powdery substance,” leaves on petioles 4-8™™
long, and the spiny branchlets only 12-25™™ long. Though no specimens of
the variety are at hand, it seems highly improbable that it is the same as the one
‘now described. Should such prove to be true, it certainly deserves to be separated
from C. lycioides, as is now done.

Mentzelia polita, n. sp.—Perennial, but probably short-lived:

428 BOTANICAL GAZETTE [UNE

root semi-woody, with a branched crown: stems 2 to several from the

crowns, rather slender, erect, 2-42™ high, somewhat cymosely
branched at summit, glabrous, polished ivory white: leaves all entire;
the lower narrowly spatulate-oblanceolate, sometimes abruptly acute,
all but the lowest sessile; from the middle up linear, and the upper-
most broadest at the base; all obscurely papillose, with a minute seta
from the center of the papilla, and closely covered with microscopic
pointlike hairs barbed at the summit: calyx-tube short-turbinate,
about 5™™ long: petals white, spatulate, 10™™ long: stamens numer-
ous, a few of the outer filaments dilated-petaloid but all fertile, shorter
than the petals: style stout, not cleft at apex: capsule globose,
crowned by the divaricate and now somewhat subulate calyx-lobes.

Las Vegas, Nevada, from washes in the hillsides, May 4, 1905, Goodding 2273.

Mentzelia synandra, n. sp.—Harsh and hispid throughout, dura-
tion not known (roots wanting in type collection): stems apparently
several from the base, somewhat branched, 3-42™ high, with long
whitish aciculae and the short barbed pubescence of this genus:
leaves mostly broadly elliptic-ovate, somewhat irregularly dentate,
3-6°™ or more long, the petioles usually much shorter; the upper
surface bearing long aciculae with pustulate bases, and the lower
long barbed hairs, both kinds mingling on the petioles and _ inflo-
rescence: calyx-tube 10-14™™ long, somewhat exceeded by the linear-
lanceolate lobes: petals 5, yellow, obovate-spatulate, abruptly acute,
3-4°™ long: stamens very numerous, all similar but the inner succes-
sively shorter; anthers oblong-oval; the filaments filamentous, not
longer than the calyx-lobes, all connate at base, forming a firm
thick-walled ring to which the petals cohere by their bases, the whole

deciduous together from the firm rim of the calyx-tube: style stout,

the 3-5 stigmas more or less convolute: fruit unknown.

Las Vegas Mountains, southern Nevada, May 13, 1905, Goodding 2373.

A very remarkable species, simulating the arrangement of petals and stamens

in the Malvaceae.

Chylisma hirta, n. sp.—A coarse biennial, 3-54™ high, branching
from the lower part of the stem or from the crown, densely white-

hirsute on the stems, less coarsely so on the leaves and sparsely on
the pods: leaves mostly basal (the crown-leaves wanting in the type),

‘ E ee
eit
ee eee ee et ee

~ 1909] NELSON—PLANTAE GOODDINGIANAE 429

_ very variable, 5—10°™ long or more, somewhat lyrate; the terminal
lobe large, elliptic-ovate, irregularly crenate-dentate or entire; the
_ lateral lobes irregular, small, few to many or even wanting: flowers
_ large, crowded but ultimately evolving an elongated large-fruited
_ raceme: calyx-tube turbinate, 4-5™™ long; its lanceolate lobes twice
as long, with a short hornlike appendage near the tip: petals yellow,
obovate-orbicular, about 15™™ long: stamens about equal; filament
and anther each about 5™™ long: style shorter than the petals: cap-
sules 5-6°™ long and 2-3™™ broad, on pedicels 5—15™™ long: seeds
_ numerous, somewhat ovoid but irregular and angled through pressure,

= 2-3™™ long.

- Tuly’s Ranch, thirteen miles from Las Vegas, Nevada, in stony washes, May

1, 1905, Goodding 234

Lavauxia lobata, n. sp.—Biennial or possibly more enduring, the
rather thick root with an enlarged more or less branched crown:
stemless or more usually caulescent, softly and conspicuously hirsute
throughout: leaves crowded on the crowns and short stems, 1-2¢™
long, narrowly oblanceolate in outline, irregularly and deeply pin-
nately lobed; the lobes mostly oblong, obtuse or subacute, 5-18™™
long: calyx-tube 7-9°™ long; its lobes ovate-lanceolate and about
25™™ long: corolla yellow, changing to red with age; the petals
as long as the calyx-lobes and a third broader than long: style as long
as the petals, with long-linear stigmas: capsule linear-lanceolate,
sharply angled but not winged, tapering very gradually from the base,
4-6°™ long, 4-6™™ broad: seeds 2-3™™ long, with conspicuous
tubercle.

Meadow Valley Wash, Nevada, in sandy washes, April 7, 1905, Goodding
37 and 47 (type).

This fine species might readily be mistaken for a yellow-flowered Pachylophus,

did it not have the angled capsule and tubercled seeds of Lavauxia.
Pachylophus cylindrocarpus, n. sp.—Biennial, stemless or more
often developing a stoutish stem 1-24™ high: leaves narrowly oblong-
lanceolate, nearly entire to irregularly and lacerately dentate, almost
glabrous on the faces but with a white fringe of soft hair on the mar-
gins; blade 5-15°™ long, on petioles as long or longer: calyx (includ-
ing ovary) softly white-hirsute; its lobes linear-lanceolate, nearly
3°™ long: petals broadly cuneate-obcordate, with deep sinus, equaling

430 BOTANICAL GAZETTE [JUNE |

or exceeding the calyx-lobes: capsule nearly glabrous, narrowly
cylindrical-oblong, 4—6°™ long, on a pedicel about 1°™ long; a double
row of small sharp crests on each obtuse angle: seeds light brown,:
2-3™™ long, obscurely bidentate at apex, the raphal furrow conspicu-
ous.

Carson’s, Meadow Valley Wash, southern Nevada, May 26, 1902, Goodding
g6oa. ;

Quincula lepidota, n. sp.—Very pale as if canescent, but without
pubescence except minute white pustular scales which are very numer-
ous on calyx, pedicels, and petioles, and occur sparingly on the veins:
perennial from rather thick rootstocks, the stoutish stems arising at
intervals either singly or in clusters: stems erect, short (probably
not much exceeding 14™ in length even at maturity): leaf blades
fleshy, mostly oval, from entire to undulate crenate, 12-25™™ long,
the narrowly margined petiole usually longer: pedicels slender,
1o-25™™ long: calyx-lobes triangular, shorter than the 2-3™™ long
tube: corolla 12-14™™ long and broad, campanulate-funnelform,
purple, with an orange-yellow band running from the middle of each
lobe toward the base of the tube, where there is a crown of 5 woolly
crests, the 5 slender filaments alternating with the crests: style longer
than the filaments but shorter than the corolla; ovary glabrous:
fruiting calyx and berry not known. .

Dry Lake, Nevada, in the gumbo soil of a dry lake bed, April 17, 1995:
Goodding 2232.

The appearance of this species is so different from that of Q. lobata, the only
other species of the genus, that were it not for the character of the pubescence and
the crown in the base of the corolla one would refer it, in the absence of fruit, to
Chamaesaracha rather than to Quincula.

Physalis genucaulis, n. sp.—Perennial from a stout woody tap
root, 1 or more long; densely but minutely pruinose or viscid
puberulent throughout, and with no long hairs: stems several, each
more or less branched; the branches with short zigzag internodes,
2-34" high: leaves ovate to ovate-triangular, the base varying from
abruptly cuneate to cordate, 1-3°™ long: calyx campanulate, about
5™™ long, the triangular lobes about half as long as the tube: corolla
greenish yellow, without conspicuous spots, campanulate-funnelform,
about ro™™ long: style shorter than the corolla and just surpassing

re: Pate ft ase ea es

ey eran

1909] NELSON—PLANTAE GOODDINGIANAE 431

the oblong anthers: fruiting calyx equally but not conspicuously
to-angled, noticeably reticulate-veined, ovoid, with sunken base and
connivent lobes closing the orifice.

Mesquite Well, southern Nevada, May 1, 1905, Goodding 2247.

One might think of referring this to P. crassifolia Benth. were it not for charac-

ter of the fruiting calyx, or to P. muriculata Greene, if dealing with the vegetative
characters alone.
~ — AMPHIACHYRIS FREMONTII spinosa, n. var.—lIntricately and
divaricately branched, many of the branches naked and slenderly
spinose, floriferous twigs not surpassing the foliar ones; scabro-puber-
ulent on foliage and young twigs: leaves oblong to elliptic, acute at
both ends, 5-12™™ long: heads congested-glomerate: ray sharply
3-toothed.

Moapa, Nevada, April 8, 1905, Goodding 2199.

HyMENOCLEA FASCICULATA A. Nels., Bot. GAZETTE 37: 270.
1904.

Mr. Goopp1nc again secured this species, this time at Cane Springs, Meadow
Valley Wash, Nevada. These specimens are in perfect accord with the type
collection, no. 662, Kernan, Nevada.

HyMENOCLEA FASCICULATA patula, n. var.—Slender stems (7-10°™
long) widely procumbent or drooping; branchlets assurgent from
the stems upon which they are uniformly placed, and not fasciculately
clustered at the ends; the very short floriferous twigs similarly dis-
tributed upon the branches, and the little glomerules of 3-5 heads
(staminate and pistillate) open racemosely or almost spicately arranged
upon the branches: involucral bracts of the staminate heads nearly
entire; those of the pistillate heads very broadly reniform.

Moapa, Nevada, April 8, 1905, Goodding 2178.

BAILEYA PLENIRADIATA perennis, n. var.—Stems numerous,
crowded in a dense cluster on the crown of a large indurated root,
3-59™ high, leafy almost to the summit. :

Moapa, Nevada, April 8, 1905, Goodding 2176.

Typical B. pleniradiata is an annual. This perennial form is more robust,
and has a larger number of bracts (about 40) in the involucre and more disk-
flowers (60-75). As pointed out by Hatt (Comp. S. Calif. 164), the original
B. multiradiata Harv. & Gray (Emory Report 144) is not the B. multiradiata
Gray of Syn. Fl. 1: 318, but is the var. nudicaulis of that work.

432 BOTANICAL GAZETTE [JUNE

Gaillardia pedunculata, n. sp.—Winter annual or biennial, 2-44m
high: stems few to several from the crown of the slender tap root,
leafy on the lower one-fourth only, the rest being the slender mono-
cephalous peduncle, softly cinerous-hirsute: leaves irregularly pinnati-
fied to entire, oblanceolate to linear, 2-6°™ long, more or less petioled,
slightly viscidly pubescent especially when young: involucral bracts
in about 2 rows, moderately whitened with flat woolly hairs, shorter
than the disk which is 12-14™™ wide and high: rays 2°™ or less long,
clear yellow, minutely pubescent on the outside, cleft one-third their
length into lanceolate lobes, tapering cuneately from summit to the
short very slender tube: disk-flowers also yellow; limb tubular,
densely and minutely pubescent with beaded hairs: pappus of very
thin paleae about as long as the inordinately pubescent achene, much
shorter than the disk corollas, narrowly to broadly elliptic, mostly
obtuse, and without costa or awn: fimbrillae of the receptacle nearly
obsolete, consisting of a few short slender teeth.

Moapa, Nevada, April 8, 1905, Goodding 2177.

This seems to have no near relative among described species.

Enceliopsis, nov. gen.—Enceliopsis Gray, Proc. Am. Acad. 19:9.
1883, and Syn. Fl. 1: 283. 1894, as Section I of Helianthella.—Xero-
phytic plants, perennial from an indurated branching caudex, the
crowns of which bear the rather thick simple leaves and the single
long pedunculate monocephalous scape. Leaves canescent, and the
petioles usually margined and longer than the blade. Heads large;
the -involucral bracts in 2 or 3 series. Bracts of the receptacle
chaffy, hyaline, or scarious with greenish tip, and more or less con-
duplicate. Rays (rarely wanting) yellow, conspicuous, pubescent
on the exterior, 20-40. Disk-flowers also yellow, with short narrow
tube, abruptly expanded into the longer cylindrical throat. Achenes
flat, oblong-cuneate, with narrow callous margins and the broadly
retuse summit with a wider crownlike callus, from densely to thinly
villous. Pappus of two subulate awns and in some species a narrow
fringe of confluent squamellae between them; rarely even the awns
wanting.—Plants peculiar to the “limestone clays” of the desert
Southwest (southern Utah and Nevada, and adjacent Colorado,
New Mexico, and Arizona). —

;
1909] NELSON—PLANTAE GOODDINGIANAE 433

The species for which this new genus is proposed were most of them described
under Encelia, but have in more recent years been transferred to Helianthella,
and sometimes back again. This of course indicates that they do not conform
to either genus, and since the five species constitute a very homogenous and charac-
teristic group it seems far better to give them generic rank. E. nudicaulis, though
not the oldest of the species, was the first to be correctly and completely delineated,
and may be cited, therefore, as the. type of the genus. Mr. Marcus E. JoNEs
has well called attention to the fact that these are singularly out of place in Helian-
thella so far as habitat is concerned. The true species of that genus belong in
the mountains, mostly in cold moist situations in high altitudes; while Enceliopsis
occurs only in absolutely the hottest, driest area to be found on this continent.

Enceliopsis nudicaulis, n. comb.—Encelia (§ GERAEA) nudicau-

lis Gray, Proc. Am. Acad. 8:656. 1873; Helianthella nudicaulis

Gray, Proc. Am. Acad. 19:9. 1883; Encelia nudicaulis Jones, Proc.
Calif. Acad. Sci. II. 5:701. 1895.

Enceliopsis argophylla, n. comb.—Tithonia argophylla Wats.,
Bot. King’s Rep. 5:423. 1871; £. argophylla and H. argophylla
Gray, in turn, as above; not H. argophylla Coville, Contrib. U. S.
Nat. Herb. 4:132. 1893; E. argophylla Jones, |. c. 702.

Enceliopsis grandiflora, n. comb.—H. argo phylla Coville, Contrib.
U. S. Nat. Herb. 4:132. 1893; £. grandiflora Jones, I. c. 702; H.
Covillei A. Nels., Bot. GAZETTE 37:273- 1904.

Enceliopsis nutans, n. comb.—E. nutans Eastwood, Zoe 2:230.
1891; Verbesina scaposa Jones, Zoe 2:248. Ig0I.

Enceliopsis tuta, n. sp.—The large woody root crowned with a
widely and freely branching caudex; the branches thick, 2-10°™
long, protected from desiccation by a thick felted sheath of white
wool: leaves all on the crowns, densely and minutely appressed-
cinerous or silvery white, rather small, 15-25"™ long, narrowly to
broadly elliptic-ovate, mostly cuneately subacute at both ends; the
barely margined petiole usually much longer than the blade: scape
rather slender, 15-30°™ long: involucre hemispherical, 20-25™™
broad, its pubescence similar to that of the leaves but longer; invo-
lucral bracts in about 2 series, narrowly lanceolate, the outer g-1r™™
long, the inner a little longer: rays puberulent, as in the other species,
20-25™™ long, linear, entire or 2-3-toothed at the slightly broadened
apex: chaffy bracts of the receptacle equaling the disk-flowers:
achenes softly hoary-villous, the dark body (when wet) in fine contrast

434 BOTANICAL GAZETTE [JUNE

to the white margin and crown, about 1°™ long, the slender incurved
awns fully one-third as long and wholly free from and surpassing the
hair on the achene: glandular waxlike particles occur abundantly
on the flowers and free tip of the chaff.

Las Vegas, Nevada, May 4, 1905, Goodding 2271.

Chaenactis paleolifera, n. sp—Biennial or possibly perennial:
the tap root with an enlarged indurated crown bearing few to several
freely branched stems, 1.5-34" high: leaves numerous, pinnately
parted into few to several mostly short linear entire segments, canes-
cently tomentulose as are also the stems and involucres: heads
numerous, terminating the branchlets, naked pedunculate, 12-157"
high and broad, 40-6o-flowered: involucral bracts linear-lanceolate,
slightly acuminate: receptacle convex, with numerous (as many as
the flowers?) paleae; these linear, clavellate-acuminate above, and
minutely glandular-pubescent, as are also the corollas, which exceed
the paleae but little: corollas ochroleucous, essentially alike; their
tubes a little shorter than the slightly enlarged throat: stamens
included: stigmas exserted: pappus paleae 4, usually lance-acuminate
and as long as the corolla-tube, sometimes shorter and obtuse, oF
slightly lacerate: achenes linear, slightly enlarged upward, and nearly
terete, softly pubescent.

Tuly’s ranch, 13 miles north of Las Vegas, Nevada, May 10, 1905, Goodding
2344.

Only two other species are accredited with paleae, C. carpoclinia and C.
attenuata Gray, with 10 and 5 paleae respectively. These, apart from the differ-
ences in the number of paleae, cannot be confused with the species here proposed.

LEBETINA Cass. in Dict. Sc. Nat. 25:394. 1822.—Among the
several names to which the following species have been referred,
Lebetina seems to be the earliest and the only one proposed especially
for any of them. Dysodia, as at present constituted, includes most
diverse things, and in the section Eupysopr1a extreme incongruity
seems to have been reached (see Gray, Proc. Am. Acad. 19:37- 1883;
HorrMann, ENGLER & PRANTL, Pflanzenfam. 45:266. 1890). HOFF
MANN assigns the first of the following to a section by itself, but had he
added the other species the section would still have been fairly homo-
geneous and would have relieved the section Eupysop1a. To think of
Dysodia papposa and D. Cooperi Gray as congeneric stretches one’s

‘3

eM Sets

1909] NELSON—PLANTAE GOODDINGIANAE 435

scientific imagination too far; therefore, I suggest the recognition of the
genus Lebetina, with at least the species named below. These species
in habit and habitat and in most essentials of structure are in close
accord, the first being exceptional in having an extra series of paleae,
and the first and second in that the style appendages are abruptly
instead of gradually acuminate. The characters of the genus can
be obtained from the description of the section Eupysop1A and its
subdivisions, as cited above, and in Syn. Fl. 1:356. 1884.

Rather than leave these species in Dysodia, it were better to trans-
fer them to Porophyllum Vaill, or to Nicolletia Gray, in either of
which less violence would be done so far as appearance gives any
clue to general conformity.

LEBETINA CANCELLATA Cass.—Dysodia cancellata Gray, tS
and Hoffmann, J. c.

Lebetina porophylla, n. comb.—D. porophylla Cav., Anal. Cienc.
42334; D.C. Prodr. 5:639; not D. porophylla Willd., Enum. goo.

Lebetina speciosa, n. comb.—D. speciosa Gray, Proc. Am. Acad.
5:163. 1861.

Lebetina porophylloides, n. comb.—D. porophylloides Gray,
Mem. Am. Acad. 52322. 1855; and Bot. Cal. 12397. 1886.

Lebetina Cooperi, n. comb.—D. Cooperi Gray, Proc. Am. Acad.
g:201. 1874; and Bot. Cal. 1:397- 1885.

The collection of this last species by Mr. GooDDING (no. 2246, Mesquite Well,
Nevada) led to a study of this group, which has convinced me that Dysodia
will receive further segregation, though it may at the same time be expanded in cer-
tain other directions. HorrMANN has thus referred Thymophylla Lag. (Hymen-
atherum Cass.), and has found it necessary to change the generic description in
no essential character. For that reason the following may be referred to Dysodia.

Dysodia cupulata, n. sp.—Herbaceous perennial, from slender

woody roots; puberulent on stems and leaves; foliage and involucre

more or less beset with small round oil-glands; branching below:
stems slender, less than 12™ high, very leafy; branchlets terminating
+n a filiform naked monocephalous peduncle 2-4°™ long: leaves
opposite below, pinnately parted into 3-5 filiform acerose lobes
1-2°™ long: involucres broadly campanulate or cup-shaped, about
5™™ high and broad; _ bracts about 16, completely united, 1 or 2
minute free bracts at base: rays about 12; ligule elliptic-oblong,

436 BOTANICAL GAZETTE [JUNE

2-3™™ long, fertile, yellow: disk-flowers also yellow, numerous,
slightly exceeding the pappus, one sinus more deeply cleft than the
others: pappus of 10 narrow paleae, united at base and in a single
series, obscurely bidentate at apex, the mid-nerve continued from
between the teeth as a minutely scabrous seta as long as the palea,
the alternate paleae and setae shorter: achene linear subterete,
obscurely ciliate-pubescent, 2-3™™ long, as long as the longer setae:
stigmas obtuse.

Tuly’s Ranch, Las Vegas, Nevada, May 10, 1905, Goodding 2343.

This is probably very near to Hymenatherum Thurbert Gray (Proc. Am.
Acad. 19:41 and Syn. Fl. 1:358) and it may have to become Dysodia T hurberi.
The description of that species is such as to make it difficult to settle the question
positively in the absence of the type or of authentic material, but the geographical
distribution makes their identity quite improbable.

Dysodia fusca, n. sp.—Pubescence minute, scurfy-glandular:
plants low, 1-24 high, freely branched from a woody base; the
ultimate branchlets very slender, fragile, white: leaves numerous,
opposite, crowded (the internodes short), very narrowly linear, all
or nearly all entire, mucronate, with few to several dark oval oil-
glands: heads nearly sessile, campanulate-turbinate, 5-87” high:
involucre cupulate, only the very short obtuse tips of the bracts free,
with a few subulate accessory bracts at the base and with few to several
oil glands: ligules oblong, 5-8, about 4™™ long: disk-flowers about
as many, very narrow: anthers and stigmas included, the latter
truncate, with an obscure apiculation: pappus paleae of both kinds
of flowers wholly resolved into unequal scabrous capillary bristles
as long as the disk corollas, fuscous and protruding brushlike from
the involucre of mature heads: achenes linear, very finely striate,
minutely pubescent, subterete, as long as the pappus: receptacle
alveolate, naked, or with a few soft scattering hairs.

Muddy Range, southern Nevada, in a stony wash (three plants), April ro,
1905, Goodding 2214.

This seems to fall into Gray’s section BorBera (Syn. Fl. 1:356) and possibly
may be allied to the two Mexican perennials mentioned. Those, however, have
pedunculate heads and the leaves pinnately divided as is usual in the genus:
Only the most liberal interpretation of the genus admits this species, and were 4
not for the gamophyllous involucre it were better to place it in Pectis, which it
resembles in habit and in the opposite somewhat connate leaves.

So“

1909] NELSON—PLANTAE MISCEL , ANEAE 437

PLANTAE MISCELLANEAE

Euphorbia manca, n. sp.—Annual, the decumbent base giving rise
to few to several simple, erect branches 1-22" high: leaves obovate-
cuneate, broadly obtuse, numerous, the lower reduced: primary
floral bracts ovate-reniform; the secondary broadly reniform, some-
times connate: inflorescence once or twice trichotomous: capsule
about 4™™ long: seeds short cylindrical-oblong, gray but not ashy,
nearly smooth.

Mancos, Colorado, June 23, 1898, Baker, Earle, and Tracy 23.

This has been referred to E. crenulata Engelm. by Norton in Rep. Mo.
Bot. Gard. 10:36. 1899. That species, as there constituted, however, is clearly
an aggregate, both annuals and perennials being included even when of very
diverse habits. The segregates readily discriminated seem to be as follows:
Perennial with branched stems from horizontal or ascend-

ing rootstocks; leaves crenulate; seeds with deep dark-

colored pits. 1. £. Nortoniana
Annuals:

Stems branched above; leaves crenulate; seeds ash-

colored with irregular vermiculate ridges and broad

shallow pits. 2. E. crenulata
Stem branched from the decumbent base; leaves entire;
seeds greenish gray, nearly smooth. 3. E. manca

E. crenulata is characterized adequately in the original description in Bot.
Mex. Bound. 192; as well as in Wats. FI. Cal. 2:75 (as E. leptocera, an undoubted
synonym); and in Greene, Man. Bay Region 80

Euphorbia Nortoniana, n. sp.—E. crenulata of Norton in Rept.
Mo. Bot. Gard. 10:36. 1899, as to the perennial plant, from which
the description is chiefly drawn.

pparently common in California, the type selected being Heller 6625 (San
Francisco, April 25, 1903) and 6486 (Pacific Grove, March 30, 1903).

In his key to the species of Euphorbia, Norton provides for both the annual
and the perennial plants (1. c. 8); and a very different plant from the above served
for the figure on his pl. 36.

Gaurella canescens (Torr. and Frem.), n. comb.—Oenothera
canescens Torr. and Frem., Fremont’s Rep. 315. 1845; Gaurella
guttulata (Geyer) Small, Bull. Torr. Bot. Club 23:183. 1896.

UNIVERSITY OF WYOMING

LARAMIE, WYOMING

THE LEAVES OF PODOPHYLLUM

J. ARTHUR HARRIS

An inspection of the leaves of the flowering stem of Podophyllum —

peltatum shows that they are not exactly the same size. Furthermore,
the larger one seems generally to be inserted a little lower on the axis
than the other of the pair. Hot (3) found that the two leaves do not
develop simultaneously but that one appears before the other. Con-
cerning this point he writes:

Of its two leaves, the one is developed earlier than the other. The base of the
petiole of this leaf is dilated into a pair of broad wing-like stipules which envelop
each other and enclose a small green leaf and a flower bud; thus the two green
leaves did not develop at the same time, as it might seem when we examine the
plant during its flowering period with its leaves apparently opposite.

Some of the teratological literature has an interesting bearing upon
this question of the differentiation of the leaves. Porter (8) illus-
trates one type in which the flowering stem bears two leaves, the
peduncle apparently originating from one of the petioles two or more
inches above their insertion. In another form there are three peltate
leaves with the peduncle originating between the upper two, which
are represented as about equal in size and opposite," or some distance
above the fork from one of the petioles. One of the leaves may be
much reduced in size, or but one leaf—then apparently terminal and
with the peduncle lateral—may appear. Finally both leaves may be
absent. Forrste (2), apparently unacquainted with PorTER’s
paper, redescribes these forms and adds other types, similar in a
general way. The production of a small, not peltate lamina upon the
peduncle is not very rare. The instance observed by BatLey (1)
of a flower replaced by a small erect leaf, and mentioned by PENZIG
(7) as sehr wunderlich, was probably merely due to the early abortion
of the flower bud in such a case.

The essential point to be gained from the foregoing observations
is that the flowering stem of Podophyllum, instead of producing only
two opposite leaves, may become an elongated shoot of at least three

* Here probably belongs = case described by TRIMBLE (g) which PENZIG (7)
records as not clearly express

Botanical Gazette, vol. 47] [438

1909] HARRIS—LEAVES OF PODOPHYLLUM 439

leaves. In fact Brrrron and Brown in the Jilustrated flora describe

the species as “bearing 1-3 similar leaves or sometimes leaflets.”

The third leaf is frequently small and not peltate, but it may be very
similar to the others.

Knowing that more than two leaves are occasionally produced by
the flowering stem, and that there appears to be a slight difference in
the size and position of the two leaves in normal specimens, two
questions occurred to me: (a) To what extent are the two leaves
of Podophyllum really differentiated in type and variability ? (0)
What is the degree of similarity of leaves from the same individual ?

A satisfactory measure of the area of a leaf so irregular as that of
Podophyllum is obviously out of the question. It was necessary
therefore to select some character other than size. The degree of
lobing seems to be the only practicable one, although this character
is not so definite as might be desired, and the determination is subject
to considerable error due to personal judgment.

__ The margins of the very excentrically peltate leaves are irregularly
toothed, lobed, or divided. It is quite impossible to draw a sharp
" line of distinction between the smaller lobes and the major divisions of
the leaf, but I think that personal judgment would rarely vary greatly
in the grading of an individual collection of plants. Perhaps less
confidence is to be placed in a comparison of two or more lots taken
at different times. In sorting the leaves into classes I counted as
lobes the divisions extending at least half-way from the periphery
of the leaf to the point of insertion on the petiole; divisions less sharply
marked than this were not counted. Only normal plants—that is,
those having only two leaves—were included in the collections. In
sorting material to determine whether there is a differentiation between
the upper and lower leaves of a pair, it is important that the appear-
ance of the leaf lamina does not influence the judgment in any way.
In dividing the nearly opposite leaves into upper and lower, the
insertion of the petiole alone was examined. After the relative
2 position of the two leaves was thus decided the counts were made.
‘The first lot of material examined was taken at V alley Park, near

St. Louis, Mo., in May, 1906, when the plants were with partly grown

fruit. In very few cases was there any question concerning the posi-
tion to be assigned to the two leaves. All of the countings were made

440 BOTANICAL GAZETTE [JUNE

by myself on one afternoon, so that I think there can be but little
error due to variation of judgment. All plants were taken at random,
and none were discarded except because of mutilation which rendered
the countings untrustworthy, save three which had an extra lamina
at the base of the peduncle. The data for the 400 flowering stalks
counted appears in the form of a correlation table as table I.

TABLE I
UPPER LEAF
4 5 6 4 8 Totals

& / 5 I 4 3 8
s 6 4 34 66 I I05
ae I 52 144 28 225
wy ) 8 2 32 16 ey 40
: 9 : 6 10 I 17
mq (tO 3 I I 2 x 5
Totals... 6 | 93 242 57 2 400

The second series I secured in the woods at Palos, Athens Co.,
Ohio, in early July, 1908. It was really too late in the season to work
to advantage, for many of the plants were so badly dried that they
had to be discarded. I see no reason for believing that this discard-
ing of individuals too brittle to be counted introduces any element of
error into the work, though it did considerably increase the labor.
The countings were made in as nearly the same manner as for the
first series as possible. The data are given in table IT.

TABLE II
UPPER LEAF
2 3 4 5 é 7 8 Totals.

2 4 I I S e 2
pe Ye 13 24 a 37
a 6 2 iz 80 4 I id 100
=)? I 9 121 42 8 181
of 8 2 8 8 18
H\o9 oe I -
‘Totals... 2 2 35 227 55 17 I 339

The physical constants for these two lots are laid side by side in
table III. To permit of easy comparison the differences between
the constants for the two series, and the probable errors of the differ-
ences, are given. Comparing the means for both upper and lower

1909] HARRIS—LEAVES OF PODOPHYLLUM 441

leaves for the two habitats, we notice that they differ by ten to
twenty times the probable errors of their differences. The standard
deviations differ by only one and a half to three times the probable
errors of their differences. Perhaps the differences between the
means of the collections from the two habitats are significant statisti-
cally, but I attach no biological importance to the differences, since
they may be due merely to some slight local environmental condition. —
The variabilities certainly do not differ significantly.

TABLE IIL

VARIATION CONSTANTS FOR LOBING OF PoODOPHYLLUM LEAVES

Series of materia he ee | ee
VALLEY PARK, MO.:
MAT sci Secs at's sore 6.920+ .029 .848+ .022 12.25
= Upper leat... ........5--- 5-890+ .022 .666+ .o16 L236
PeTeONCE ics. a +1.030+ .036 + .182+ .027 + 0.95
PALOS, OHIO:
doower leat, 2... .--.<--+- 6.528 .029 .792+ .021 12.13
Be CE ACRE ci ee a 5.1394 .027 -742+ .029 14.43
PICTENCE. 56 eee ees +1.389+ .040 + .o50+ .036 — 2.31
DIFFERENCES
ower leat 7 6c 54 oss 2. ky +0.392+ .041 + .056-+ .030 + 0.12
Sigper leafs. : ks. 2 oe. +0.751+ .035 — .0764+ .025 — 3.13

Taking now the question (a), that of a differentiation between the
_ upper and lower leaves, we note that the means differ in both cases
by about thirty times the probable error of their differences, and that
- the lower leaf has in both cases about one lobe more than the upper.
The standard deviations differ by an amount which can hardly be
_ regarded as significant. The relative variability as measured by the
coefficient of variation is in one case higher for the upper leaf and in
one case lower. After calculating the constants for the first series
of material, I thought that perhaps the variability of the more distally
_ placed leaf would be regularly lower than that of the more proximal
~ one, as Peart (4) found the variability of the whorls in Ceratophyllum
tobe. But the second series does not support this idea.

| It may be interesting to compare the variability in the lobing of the
_ leaves of Podophyllum with that of other leaf characters given by
- Pearson (5). From page 361 I note the following values of the
coefficient of variation for leaf characters:

442 BOTANICAL GAZETTE [JUNE

Holly, Dorgetshire, prickles on Jeaves.................-..0.-. 26.29
Holly, Somersetshire, prickles on leaves.............-..... eee
eh Drorertanire, leaficts On leaves... ....:...... 6.0... -..545: 18.65
Ash, Monmouthshire, leaflets on leaves...................... 18.57
Ceterach, Somersetshire, lobes on fronds...................+.. 18.25
Wild ivy, mixed, leaf-indices.......... Pee che. Sees Fos 17.97
Spanish chestnut, mixed, veins in leaves...................-- 15:72
Ash, Buckinghamshire, leaflets on leaves. ...................- 15-46
Spanish chestnut, Buckinghamshire, veins in leaves............ 14-31
Beech, Buckinghamshire, veins in leaves............ on cae es 10.77

It appears by these comparisons that the lobing of the leaves of
Podophyllum is rather less variable than leaf characters in general.

Turning now to the question of the degree of similarity between
the two leaves, and calculating the coefficient of correlation between
the number of lobes on the lower leaf and the number of lobes on the
upper leaf by the familiar product-moment method, we find the corre-
lations,

OR bai vee ck eee .428+ .028
ee yee ce ecek . 468+ .029
WP eee -O40 .040

I was considerably surprised when these values turned up on my
dividing machine. A priori, I would have expected considerably
higher coefficients, say about .700, for the correlation between organs
so closely associated as the leaves of Podophyllum. In thinking of
the correlation between the leaves of Podophyllum, it had always
seemed to me organic rather than homotypic in nature. The two
leaves seemed so nearly exactly opposite and the whole “normality”
of the plant seemed to depend so much upon their forming a symmetri-
cal pair that I had expected the usual homotypic resemblance plus
something more.? But instead we find values which fall directly
in line with those found by Pearson and others for homotypes in
general. Taking merely the leaf characters noted above, we find the
homotypic relationship calculated by PEarson and his coworkers
(5) to be the following: ,

Ceterach, Somersetshire, lobes on fronds..............--+++++*

Holly, Dorsetshire, prickles on leaves................002000008 .599

? Fora clear statement of the distinction between organic and homotypic correlation

1909] HARRIS—LEAVES OF PODOPHYLLUM 443

Spanish chestnut, mixed, veins in leaves.................00-5- 591
Beech, Buckinghamshire, veins in leaves... ..: 60.0... + 2200505 SAO
Spanish chestnut, Buckinghamshire, veins in leaves............ . 466
Ash, Monmouthshire, leaflets on leaves. .....--...6,225+.200+ 405
moo, orsettshite; leaflets on teaver. <:o.2 235 270 8 24 ee oe el . 396
Ash, Buckinghamshire, leaflets on leaves.................+.54: 374
fecmy, Somersetshire, prickles on leaves. 2. 62.6.5 7 i et ees cages
meing ivy, tained, leal-indices. . 6.2505 444 3 6 Ly 273

Some of these are slightly higher and some slightly lower than
our coefficients; but when the probable errors attached to all con-
stants are borne in mind, I think we cannot assert that our values
are very different from those obtained by English biometricians
for other leaf characters. Furthermore PEARSON shows reasons for
considering some of his values too high and some too low for true
homotypic relationships. For instance, ceterach is said by botanists
to be largely influenced by growth and environment.’

There still remains one possible reason for thinking that the real
correlation between the number of lobes on the leaves of the same
flowering stalk may be somewhat higher than is indicated by these
constants. The May apple spreads considerably by rootstocks. My
plants from both habitats were taken from quite a wide stretch
of woods, but a considerable number of the plants are doubtless
vegetatively related. I do not believe this has a very large influence
in my series, oe it is — to mention the point.

3F th Iso difficulti I did not apply SHEPPARD’S
correction for the second moment in calculating my standard ernie Perhaps
this should have been done, but until some mathematician works out the theory,

biologists will not know what rule to follow in ee case of integral variates. In the case
of a range of variability so narrow as we have here, SHEPPARD'S correction would
make a considerable eres raising the coefficient of correlation by lowering the
aggnitala deviations. So perhaps our _— = gee a ee aes
also the question of leave nandn+t1

lobes. Th tbe present aie I Srehully tried to throw these into the class to which they
most nearly belonged, just as one would have to do in the case of real integral variates.
But after all, the lobing of the leaf of Podophyllum is not a case of discrete variation,
and if I were repeating the work I would divide questionable cases between re
grades. Probably this would not make a very grea t difference in the end res

e reader will note, too, that I am discussing cep correlation on series
of s sactel which I have just demonstrated to be differentiated. But I think I am
quite justified in treating the material as I have done, for lower and upper leaves
“have always been kept separate. There has been no m ixing of heterogeneous
material.

444 BOTANICAL GAZETTE [JUNE

Summarizing, we may say that so far as our materials show:
(a) there is a sensible differentiation between the two leaves of the
flowering stalk of Podophyllum in the number of lobes, but apparently
not in the variability of the lobes, at least not in their relative variabil-
ity; and (0) the correlation between the number of lobes on the two
leaves of the stalk lies somewhere in the neighborhood of .45, agreeing
well with the homotypic correlations for leaf characters in other
species.

CARNEGIE INSTITUTION OF WASHINGTON

LITERATURE CITED

. BarLey, W. W., Bull. Torr. Bot. Club 13:111. 1886.
ForrstE, AuG., The May apple. Bull. Torr. Bot. Club 11:62-64. 1884.
. Hox, T., Podophyllum peltatum: a morphological study. Bor. GAZETTE
- 419-433. 1899.

ARL, R., Variation and differentiation in Ceratophyllum. Pub. 58 Carnegie
Sas Wash.
PEARSON, K., ae others. Mathematical contributions to the theory of evo-
lution. IX. On the principle of homotyposis and its relation to heredity, to
the variability of the individual, and to that of the race. Phil. Trans. Roy.
Soc. Lond. A. 197:285-379. 1

wN H

un :

I
6. , On the fundamental conceptions of biology. Biometrika 1: 320-344.
Igor.
7. PENZIG, Si Pflanzenteratologie. Genoa. 1 890-94.
8. Porter, T. C., Variation in Podophyllum peltatum. Bot. GAZETTE 2:117,

118. 897:
9. TRIMBLE, W., Teratological notes. Bull. Torr. Bot. Club 9:10. 1882.

Ee a a ie ren es See oe eee Pe

A BOTANICAL SURVEY OF THE HURON RIVER VALLEY’
VII. POSITION OF THE GREATEST PEAT DEPOSIT
N LOCAL BOGS :
GEORGE PLUMER BURNS
(WITH FIVE FIGURES)

The general appearance of the vegetation of the peat bogs in the
Huron River valley has been fully described by TRANSEAU (8) and it is
not necessary to give a detailed account in this paper. The plants are
usually growing in a more or less regular zonal arrangement somewhat
as follows: (1) open water with submerged plants, (2) water lilies,
(3) floating sedge, (4) bog shrubs, (5) tamaracks, (6) maple-poplar,
(7) willow or marginal zone.

There is wide variation in the position of the greatest amount
of peat deposit and the width of the various zones. Even a casual
survey of a number of local bogs emphasizes the fact that the open
water is seldom in the center of the original post-glacial lake. The
zonal arrangement is usually broken on one side, some zones being
entirely lacking. In fact it is not uncommon to have all peat deposit
lacking at certain places along the shore. According to some writers
(7) the greatest accumulation of peat is found on the western side,
in regions with prevailing westerly winds. In large basins which are
only partly filled it is common to find open water occurring toward
the eastern side. The wave-action produced by the westerly wind is
supposed to hinder bog plants from obtaining a foothold on the east-
ern shore. The shoreward push of the ice is also a factor of impor-
tance in this connection (8, p. 418).

A study of the bogs near Ann Arbor soon revealed the fact that the
greatest accumulation of peat on the western side was by no means
common to all bogs. In fact the greatest variation was found within
very short distances. At Dead Lake it is on the southern side (1);
at Mud Lake, about one-half mile (800™) west, it is on the northern
side (6); at First Sister Lake, it is on the western side (9); at a small
bog near Carpenter’s Corner it is toward the east (7).

t Contribution 112 from the Botanical Laboratory of the University of Michigan.

445] [Botanical Gazette, vol. 47

446 BOTANICAL GAZETTE [JUNE

It seems, therefore, that some other factor must be sought. Plane-
table maps were made of a number of bogs and the distribution of
plants was carefully plotted. Contour maps of the bottoms of the
original post-glacial lakes were then made. In making these maps
a drill was made of gas pipe, cut in four-foot (1.2™) lengths. To
the end section was welded a carpenter’s auger. With this drill,
soundings were made through the peat to the clay bottom of the
original post-glacial lake. A base line was run taking the longest
diameter of the bog. Along this and at right angles to it, cross lines
were drawn every hundred feet (30.48™) in large lakes and every
fifty feet (15.24™) in small lakes. Borings were made at the inter-
sections of these lines and every hundred or fifty feet on the cross lines.
Near the shores it was necessary to make the borings closer, often
every five feet (1.5™). In bogs with open water it was necessary
to make all borings through the ice, because it was impossible to hold
a boat steady enough to bore through the peat without breaking the
drill.

First Sister Lake.?—This bog is situated a short distance west
of Ann Arbor and has already been described by other writers (8,
9). It is surrounded on all sides by hills, except for three breaks.
To the north was.a small post-glacial lake; on the southwest corner
it connected, by a narrow channel, with Second Sister Lake. . The
whole formed a very irregular chain of lakes draining into the Huron
River.

The contour map of the bottom of the original post-glacial lake
is shown by red lines in fig. r. The heavy line indicates the margin of
the original lake as indicated by the peat deposit. The shores rise
somewhat abruptly on all sides. The lake had one basin. The
deepest part was east of the center, the deepest point being fifty-four
feet (16.5™). From this point the bottom sloped gradually to the
eastern shore. On the opposite side the forty-foot (12.17™) contour
made a wide divergence to the west; from this contour the bottom
sloped gradually to the shore.

No attempt is made to show the flora of this early time. The black
of fig. 1 shows the arrangement of things as they were last summert-
The open water occupies a very small area compared to what it did

2 Borings were made for the most part by Mr. HAROLD STEELE.

Ba ae al cc ei a a cs

990058
0 on 208"
040,007 lt
ee 2 090° )
et
oe 0%

eon

1G. 1.—First Sister Lake. Contours of the post-glacial lake in red. The heavy

line indicates the position of its shore as shown by peat deposit. Scale 1: 2880. Contour
interval of topography five feet. of hydrography ten feet.
Present distribution of plant societies in black. A open water; B bog sedge;

C bog shrub; D clearing; E tamarack; F maple-poplar; G willow; H oak-hickory.

Ashita [il OO
! i i ty fie so
il i

Ht tig: a
ie ) FES,

LIN

iri

Contours of the bottom of the post-glacial lake in red. The heavy line indicates
the ks of the shore before ‘anaes The line o indicates the water level when the map was made. Scale
1:4800. Contour interval five fee

, 3-——Dead Lake.

Distribution of plants when aie map was drawn in black. A area which had been cleared of tamarack befor
it was suitable for maple-poplar or the clearing society; B tamarack; C bog shrub; D bog sedge; E cultivated
field; F water lilies; G island; H open water; J drain; J oak woods

1909] BURNS—BOTANICAL SURVEY OF HURON VALLEY 447

originally. The volume of water, however, is much smaller than it
appears in the figure just referred to, as is seen in a profile through
this section of the lake (fig. 2). The greatest accumulation of peat
is on the western side. The zonal arrangement is also quite different
on the two sides. The bog-shrub, tamarack, and maple-poplar zones
are almost entirely lacking on the east. On the west, all the zones are
present, the tamarack zone being very wide especially in the south-
western portion.

Fic. 2.—First Sister Lake. Profile east and west. A open water; B bog sedge;
C bog shrub; D tamarack; E maple-poplar; F willow. Scale: vertical, 1:600;
horizontal, 1: 2880.

In fig. 7 it is interesting to observe that the open water now lies
over the deepest part of the post-glacial lake; and to follow the
variations in the width of the tamarack zone. The southern area
is broken by areas of bog shrubs, showing that the tamarack has not
entirely captured the area. The northern group, on the other hand,
is disappearing, and when a number of trees were cut out for wood a
few years ago an entirely different society of plants came in—a clearing
society. In other words, conditions favoring the development of a
tamarack society are found where the original post-glacial lake was
forty feet (12.17™) or more in depth, and where the depth was less
than this conditions are favorable for the development of the next
- zone of plants.

Boc NEAR CARPENTER’S CORNER.—This is a small bog east of
Ann Arbor. It is surrounded by high hills on the north, west, and
south. On the east a narrow ridge about five feet (1.5™) high sepa-
rates it from a large “drained swamp.” A ditch has been dug through
this ridge to drain the bog. The original post-glacial lake was small,

448 BOTANICAL GAZETTE [JUNE

occupying about one-tenth of an acre (400%™). It had only one
basin whose greatest depth was fifty-nine feet (18"). This point was
considerably west of the center. The slope of the shore from the
eastern margin was very gradual for a distance of nearly one-third
the east-and-west diameter, when it becomes very steep. The eastern
third of the lake was about fifteen feet (4.57™) deep. The slope from
the western shore was much steeper, and hence the greatest amount
of water was west of the center.

Today the open water has entirely disappeared. From the banks
a group of tamaracks may be seen. These are arranged around a
small central area within which are found (7) a few individuals
of the bog-sedge and bog-shrub zones which formerly occupied large
areas. Some of the plants found were Carex filiformis, Sphagnum
sp., Sarracenia purpurea, Cypripedium acaule, and Vaccinium
Oxycoccus. Outside of this area is a zone of mixed vegetation. The
width varies from several rods to a few feet, the widest area being
found on the east. The dominant trees are Acer rubrum and Populus
tremuloides. This area has a rather indiscriminate mixture of bog
and lowland plants—it is a tension zone in which conditions are not
especially favorable to either group of plants. Around this area and
following the shore, is found the marginal area so common to bogs.
The plants occupying this area are those usually found in low wet
places along rivers.

A comparison of this distribution with borings shows that the
central area is over the deepest part of the post-glacial lake. The
greatest surface accumulation is on the western side where the lake
was comparatively shallow, and over this area Acer rubrum and
Populus tremuloides are the dominant trees.

It is thus possible to tell, with considerable accuracy, the depth
of the bottom of post-glacial lakes with one basin by the distribu-
tion of the present vegetation. The order of succession is lily, bog

dra located in a zone of Carex filiformis indicates a shallow bottom
(3), but when it is found in a zone of tamaracks it indicates @ deep
basin. ;

Drap Lake.3—This lake is located about ten miles (16*™) north

3 Mr. Forest B. H. Brown assisted i making the borings.

1909] BURNS—BOTANICAL SURVEY OF HURON VALLEY 449

of Ann Arbor. Its longest diameter is nearly directly east and west.
Red contours on the map (fig. 4) show that the bottom of the post-
glacial lake was divided into four basins; a rather shallow ridge
running north and south divided it into an eastern and western half,
and each half is in turn divided into a northern and southern basin.
The central ridge was twelve feet (3.65™) deep at the lowest point,
and in the center it came above the surface, forming an island.

In the eastern half, the northern basin reached a depth of seventy
feet (21.33™) and the southern basin thirty-five (10.66™) feet.
However, a large part of this half of the lake was comparatively shal-
low. Over a large area the depth was about ten feet (3™). The
other half was also divided into a deep northern basin which reached
the depth of seventy-two feet (21.94™) and a shallower southern basin
thirty feet (9.14™). The greater part of this half of the post-glacial
lake was deep. At some time in the past a ditch was dug through
the bank on the north and the level of the water was lowered several
feet. The conflicting stories told by the older inhabitants make it
impossible to determine when this occurred. The vegetation, how-
ever, shows that it has been a number of years.

The map of the present distribution of plants at this lake (black
parts of fig. 3) shows that the peat deposit is largely confined to the
south and east. The principal vegetation of the lake is the bog
sedge. Only on the south have the tamaracks and the bog shrubs
gotten a foothold, though “islands” of these are rapidly spreading over
the bog sedge in many places. The absence of the bog flora around
the shore of the island in the center of the lake (the vegetation shown
on the map is Scirpus lacustris), along the northern side of the west
half, and on the southeastern corner, is no doubt to be explained in
part by the action of the wind, wave, and shoreward push of the ice.
However, it must be seen from the map that these places do not stand
in a definite relation to the points of the compass. Such vacant
places are found where there is shallow water. In just such places the
actions referred to above would be most intense. It seems, therefore,
that the contour of the bottom rather than the direction of the wind
is the controlling factor. The small lake in the northeastern corner
is over a very deep basin; the deep western half is for the most part
open water. The narrow channels of open water on the eastern

450 BOTANICAL GAZETTE [JUNE

portion do not follow the contours but in some places cross them at
right angles. Their existence finds its explanation in part in the
fact that they are kept open by fishermen who approach the lake
chiefly from this side. It is doubtful, however, if this is the entire
explanation (4,5). The break in the tamarack zone on the south side
is due to two factors. At this point the water was rather shallow and
this hindered occupation by bog plants for a long period of years,
as has been pointed out; during late years the tamaracks have been
cleared away and the place kept open as a watering place for stock.
The large open area south of the tamaracks is an area suited for tama-

yy 3

2 ‘I

A-— \
re en
——— — ott

een

===\\ p=

Fic. 4.—Profiles through Dead Lake. A through the eastern half; the small
body of “open water” is filled with lilies and submerged water plants. B conditions
in the western part of the lake. Scale: vertical, 1:600; horizontal, 1: 4800.

rack growth. The tamaracks formerly growing there were removed
for wood and poles before the surface had been raised enough to make
it suitable for either the maple-poplar or the clearing society. It is
occupied chiefly by marsh ferns and sedges, with a few Rhus venenata.
A narrow border of Ulmus americana is found along the southeastern
shore (jig. 4).

Mup Lake.—This lake has been described by PENNINGTON (6).
Itwas a very large post-glacial lake but has been almost entirely filled
_ with marl and peat. The greatest deposit of peat is on the northern
side, and the open water is very close to the southern shore. In

EE PEE Qrte a

1909] BURNS—BOTANICAL SURVEY OF HURON VALLEY 451

entering the bog from the north one passes through a number of large
areas of bog shrubs, wholly or partly surrounded by tamaracks. In
these areas are sometimes found small patches of tamarack and spruce.
4 A profile was run across the bog, north and south, through these areas
{} and the open water. This shows that the areas occupied by tama-
racks are located over shallow places in the post-glacial lake, and
that areas of bog shrubs are located over deeper basins. The borings
showed that this lake had a number of basins very much the same as
found at Dead Lake. It has reached a later stage of development,
however; the dominant vegetation is the bog shrub and tamarack.

ee aw

EMO AE

id
EERIE. OS
et ae
\

—<<<<<—

———

————

—_———_—

Fic. <.—Profile north and south through Snow’s Lake. The dotted line shows
the water level before drained. Scale: vertical 1: 300; horizontal, 1: 3048.

Snow’s LaKe.4—This lake is located about fifteen miles (24
west of Ann Arbor. It was formerly a very large lake, but has been
almost filled with peat and marl. The bottom of the post-glacial
lake was very irregular, as is seen in the profile (jig. 5). This profile
runs north and south. The deepest and largest basin was near the
northern shore; the southern part of the lake was comparatively
shallow. A few years ago the lake was drained at the northeastern
corner and the level of the water was lowered in the lake. Over the
shallow parts of the lake the peat had already become solid, and the
draining did not change the level of the surface. Near the open
the plants were still floating, and when the water was
drained they sank to the new level. The surface today slants rapidly
toward the water; especially is this true for the first few rods immedi-
ately next to the open water. It is very surprising from the road to
see fine large oak trees growing apparently in the peat bog. How-

4 A careful study of this lake was made by Dr. JEAN DAWSON, but the results have

not been published.

km)

water, however,

452 BOTANICAL GAZETTE [JUNE

ever, when one goes over to the oak and examines the soil in which
it is growing, the matter is easily explained. These “islands” of
oak and other upland trees are not growing in peat, but on one of the
islands formed by projections from the bottom of the lake. Fig. 5
shows such an island with an oak growing upon it.

The same relation of present distribution and depth of the original
post-glacial lake holds true in lakes with several basins, as it did in
lakes with only one: open water over the deepest and largest basins,
wide zones of plants where the bottom of the post-glacial lake sloped
gradually, a definite order of succession of plant zones. Greater
care must be exercised, however, in the determination of depths by
the present vegetation, especially in lakes with many small basins
just at the time when large areas are beginning to become favorable
for the next group of plants. The occupation will take place most
rapidly over shallow parts, because here the peat will become firm
sooner and the conditions will first be favorable for the next group of
plants.

Conctustons.—The chief factor determining the position of the
greatest amount of peat deposit and the width of the zones of plants
at the local peat bogs is the depth of the water in the different parts
of the original post-glacial lakes. The chief factor in determining
the position of the open water is depth; given time enough, the open
water will disappear from all our lakes.

In places where the water is very shallow the bog flora is unable
to get a start because of the wave-action caused by the winds and on
account of the shoreward push of the ice. Such places, however;
bear no definite relation to the points of the compass.

Different zones of plants follow in a definite orderly succession;
lily, bog sedge, bog shrub, tamarack, maple-poplar.

UNIVERSITY OF MICHIGAN

LITERATURE CITED :
I. Burns, G. P., Formation of peat in Dead Lake. Rept. Mich. Acad. Sci.
6:76. 1904.
2. , Edaphic conditions in local peat bogs. Science N. S. 29: 269. 1999-
3- Cow es, H. C., Physiographic ecology of Chicago and.vicinity. Bot. GAZETTE
31:147. 1901.

1909] BURNS—BOTANICAL SURVEY OF HURON VALLEY 453

5.
answick 5: 460

i.)

‘o

Baxane, W. F., The vegetation of the a of Fundy salt and diked marshes.
Bot. GAZETTE a6: 174 and footnote. 19
n the physical geography of Miscoo, Bull. Nat. Hist. Soc. New

PENNINGTON, L. H, Plant distribution at Mud Lake. Rept. Mich. Acad.
Sci. 8:54. 1906.
Petree, EpitrH E., Plant distribution in a small bog. Rept. Mich. Acad.
Sci. '7:126. 1905.

. TRANSEAU, E. N., The bogs and bog flora of the Huron River valley. Bor.

GAZETTE 40: 429-448. 1905. The older literature is given by this author.

. WELD, L. H., Botanical survey of the Huron River valley. I. A peat bog and

a morainal lake. Bort. GAZETTE 37:36. 1904

POLLINATION IN LINARIA
WITH SPECIAL REFERENCE TO CLEISTOGAMY
i ta IE L
(WITH FOUR FIGURES)
I. POLLINATION BY INSECTS

The genus Linaria furnishes examples of adaptations to both
cross- and self-pollination. Of the sixteen species given in KNUTH’S
Handbuch der Bliitenbiologie, nine are said to be visited by insects
and may be pollinated by them. One, L. origanifolia DC., as
observed by MacLeop, is adapted to insects, but was not seen to use
them. Four or five seem to be restricted to self-pollination, and all
can also employ it. Some produce cleistogamous flowers, and as far
as these can be of service, are compelled to pollinate in this way.
L. vulgaris Mill. (as Antirrhinum Linaria L.) was the first to be
observed and described. This was by Cu. K. SpRENGEL in his
_ book on the relations of flowers and insects, published in 1793. It
was one of the first with which DaRwIN experimented when preparing
the material for his work on the effects of cross- and self-fertilization
in the vegetable kingdom. An unexpected presentation of vigor in
one of two beds of this species, planted for the purpose of determining
some points regarding inheritance, led him to trials with this and other
plants on the results of pollination.

Linaria is called a mellitophilous genus, since bees are the princi-
pal agents in the process, though in some species several other insects,
especially Lepidoptera, share in the work. The honey, secreted by
glands at the base of the ovary, flows into the spur of the flower,
where it is stored and awaits the visits of insects with a proboscis
long enough to reach it. It is therefore well adapted to visitors of
this kind belonging to the class called Eutropic by Loew.’ The
two pairs of anthers are placed at different heights, with the slender
style and the stigma in the space between. These are brushed by the
back of a bee crowding in to get the nectar in the spur, or by the
longer proboscis of a lepidopter, and some of the pollen is removed

' Loew, Einfiihrung in die Blithenbiologie 342, 345. 1895.

Botanical Gazette, vol. 47] 454

a ee eS

}

1909] | HILL—POLLINATION IN LINARIA 455

during the operation. That which was left on the stigma of the
flowers by the entering and withdrawing of the bees was the extent
of pollination as viewed by SPRENGEL. It was an aid to the plant
in securing fertilization indispensable in the case of some, but the
full significance in the economy of its life was left for others, especially
for Darwin, to show. SPRENGEL clearly describes the process in
the text and figures illustrating Antirrhinum Linaria,’ and in reading
his book one wonders at the sagacity of the man so far in advance
of his time. The relative position of anthers and stigma, coupled
with their simultaneous maturing, can also, as stated by HERMANN
Miter, lead just as readily to self-pollination, and in the absence
of visits by insects makes it the only possible means of fruitfulness.*
The same relations hold in the case of the smaller flowers of L. alpina
Mill., which MULLER investigated.4

The common toad-flax of Europe, L. vulgaris, has been natural-
ized in this country, and is most frequently seen along roadways
or in waste grounds. Two native species are generally recognized,
L. canadensis (L.) Dumort. and L. floridana Chapm. A third, L.
texana Scheele, is made by some, but by others is considered a large-
flowered form of L. canadensis. This seems to be the only one that
has been studied with regard to its pollination. It is widely distrib-
uted, usually growing in dry locations, such as sandy or rocky ground.
Its small flowers ally it more to L. alpina than to L. vulgaris, and
like these it is adapted to pollination by insects. CHARLES ROBERT-
son observed the flowers in Florida, and found that they were visited
by bees, but more often by butterflies. He says of them: “The spur
is very slender and the tube has become so contracted that bees can
only insert their tongues, and butterflies cannot suck without touch-
ing the anthers and stigma... . - The palate, which in L. vulgaris
permits the visits of bumble-bees only, seems to have lost its function,
for it is so weak that it entirely fails to exclude butterflies or even
flies.’ Where I have noticed it in the dune region near Chicago,

2 SPRENGEL, Das entdeckte Geheimniss der Natur im Bau und in der Befruch-
tung der Blumen 317. pl. 17. figs. 5-11, 14, 18, 19. 1793-

3 MULLER, Die Befruchtung der Blumen durch Insekten 279. 1873.
, Alpenblumen 275. 1881.
s RoBERTSON, Zygomorphy and its causes. Ill. Bor. GAZETTE 13:228. 1888.

456 BOTANICAL GAZETTE [JUNE

it does not appear to be very extensively sought by insects, but species
of Syrphidae may be seen flitting from flower to flower of this and
of plants of Krigia virginica in blossom at the same time, perhaps
as much in search of pollen as for their sweets.

MULLER examined other European species of Linaria with refer-
ence to this matter, among them L. minor Desf. and L. arvensis L.
Their flowers are very small, but adapted, like L. vulgaris and L.
alpina, to pollination by bees. As a weed in his garden at Lipp-
stadt he “looked in vain” for visitors to L. minor, and L. arvensis
was repeatedly watched in favorable weather in another station with
a like result. Hence he concludes that they are restricted to self-
fertilization. As the anthers burst at the same time the stigma
matures, should a bee come for the nectar the flowers are ready for
cross-pollination. This condition lasts only a short time; the stigma
is soon covered with pollen, and self-fertilization is accomplished.
Since MULLER cannot imagine that a flower, in all the peculiarities
of its structure fitted for pollination by insects, should still be very
exceptionally visited and crossed by their instrumentality, he con-
cludes that we have in these plants a deteriorating descendant of an
ancestor with larger and more striking flowers, in whose pollination
bees as a rule took part. He considers that the same is true of
various other plants with diminutive or inconspicuous mellitophilous

flowers which are now very rarely visited by bees, citing among others
Vicia hirsuta Koch as a similar case, whose style bears unequivocal
marks of arrested development, the brush being reduced to a dozen
hairs at most.?

Il. THE CLEISTOGAMIC CONDITION

It is only a step from this reduction of floral organs mentioned
by Mix er to flowers so diminutive and constructed in such a way
that they do not open at all, or the cleistogamic stage, in which self-
pollination is the only means of securing fertility. Of the eight
types of entomophilous flowers made by DELPrNo, the sixth is that
in which the anthers and stigmas are close together and included.
Linaria answers these conditions, as must indeed be the case with
ea Mtter, Weitere Beobachtungen III. Verh. nat. Vereins Rheinl. Westf. 39:28.
Ri .

7 » Ibid. Il. Op. cit. 38:360. 1879.

I a a a ad St

1909] HILL—POLLINATION IN LINARIA 457

all cleistogamous flowers in respect of the proximity of the organs
essential to fertility. It is these cleistogamous flowers I have mainly
investigated. My attention was first called to them in 1905, when
flowers of this character were found on L. canadensis growing upon
the sandstone rocks at Oregon, Ill. They were quite inconspicuous.
The minute corolla, when pushed off by the enlarging ovary, showed
a faint tinge of violet on its upper margin, the main part being color-
less. Since it was the middle of July, all traces of the sterile radical
shoots had disappeared, as well as such flowers as are ordinarily
found earlier in the season, if, indeed, they had been formed at all.
The plants were generally small, the shortest mostly with simple
stems. Some were branched, the tallest about 42™ high. Since
the species is well represented, though not abundant, on the sand
dunes at the south end of Lake Michigan, there has been an oppor-
tunity to observe it each season since, and to note the different stages
of development. The plants begin to blossom about the first of May
and continue, in some form, the production of flowers till the middle
or latter part of July, when the heat becomes too trying for them in
the dry sand. On the larger plants there is a gradual diminution
in the size of the flowers from the earliest, 6-8™™ long, with a diam-
eter of limb of 8-12™™, till the cleistogamous stage is reached. In
some plants of this character, this may occur in the early part of June.
It is exceptional to find flowers which are relatively conspicuous dur-
ing the later part of the life of the plant, and when found they are apt
to be much reduced as compared with the earlier forms. The
inflorescence being indefinite, the lengthening of the main stem and
branches favors this progressive diminution. Plants that do not
exceed ro to 15°™ usually remain simple and are mainly restricted
to cleistogamy. Plants taller than this commonly have flowers
adapted to pollination by insects, though it must be rare in the smaller
flowers, if done at all, when the limb of the corolla is but 3 or 4"™ in
diameter, as MULLER found was the case with the small flowers he
mentions.

Cleistogamy begins on stems not more than 2°™ high, which may
be limited to a single flower at the tip, or perhaps lengthen enough
to bear two or three. Flowers will appear on these diminutive stems
as early as the larger petaliferous ones on the vigorous plants, the

458 BOTANICAL GAZETTE [yuNE

two forms being synchronous, but on stems under different circum-
stances. They continue to increase in number on plants of this
simple character until the stem ceases to lengthen or becomes
mature, various heights being reached, but rarely more than 24™,
Other slender and normally simple stems, usually not flowering at
all till 12 to.15°™ high, bear as a rule small open flowers, and may
continue to do so for some time, growing on till by progressive diminu-
tion of size the cleistogamous stage is reached. But the plants are
apt to branch when 15 to 20°™ high, and bear the larger flowers on
the main stem and branches that successively form. ‘Two or three
of these stouter stems often spring from the same root, forming a
small cluster, with larger and more abundant radical shoots, the
plant in all its features showing its greater vigor. In Brirron and
Brown’s Illustrated flora, the statement is made under L. canadensis:
“A dwarf form with no corolla is frequent.”’ This evidently refers
to the cleistogamous form. But the stage with no corolla is not
confined to the dwarf plants. It was not on such that I noticed them
at first, but on those which varied in height. In the dune region
they may rise to 64™ and bear the closed flowers in the later stages
of growth. As the taller forms often branch quite freely, great
numbers of capsules are borne, developing on the principal stem
and branches at the same time and long after the ripened capsules
lower down have opened and dropped their seeds. Since the branches
ascend rather sharply, frequently rising well up to the level of the
primary axis, a copiously branched plant, sometimes with fifteen

to twenty members, may result. These have a bushy appearance, |

but they all produce the closed flowers before the plant dies, and
manifest its ability to bear vast numbers of seeds.

The length of the corolla in the closed flowers is 1.2 to 1.6".
It is tubular, or sometimes slightly funnelform, but owing to the quite
rapid growth of the ovary soon becomes enlarged at the base, and
when pushed off is shaped more like an inverted funnel; or, if enlarged
at the same time above, it has somewhat the form of an hour-glass.
The limb is slightly irregular, the two-lobed upper lip being higher
than the three-lobed lower, and overlapping it in the bud. In the
illustration fig. r shows an ordinary chasmogamous flower, ig. 2
a cleistogamous flower, both enlarged five diameters, figs. 3 and

1909] _ HILL—POLLINATION IN LINARIA 459

4 a cleistogene corolla enlarged ten diameters, that of jig. 4 dis-
played, the division being made between the two lobes of the upper
lip. The style and stigma, if rep-
resented, would be between the
two pairs of stamens as in the
ordinary flowers, the stigma in this
case closely pressed by the anthers
when the parts are in place. The
four stamens are apt to be present,
and slightly didynamous. Some-
times a small protuberance at the
base of the tube represents the
spur. In the smaller chasmoga-
mous forms, this may be reduced to
a short sack or tooth, and usually
decreases in size as the other parts
diminish. But in some cases it
remains relatively longer in compari-
son with the lessened tube and limb.
I found no case of a cleistogene
without a corolla. But as in other cases of cleistogamy it is" easy
to imagine the flowers represented by the calyx and the essential
organs of fructification.

Depauperate as well as larger forms with cleistogamous flowers
have been noted by others. RypBERG, in his Flora of the Black
Hills, mentions a L. canadensis collected at Custer as being “slender
and depauperate, apparently with cleistogamous flowers. The
same has also been collected in Nebraska.”* The month given for
Ryppere’s collection is August. J. R. WEBSTER records cases
observed by him at Milton, Mass., August, 1898. The plants were
again noticed the next year, being “examined almost daily from April
to October, and were seen to produce flowers abundantly which were
all cleistogamous.’’ They were observed by him in other localities,
in which were also racemes which bore in addition fully developed
flowers.2 As the plants at Milton are said to reach the height of

cleistogene flower X10; jig. 4,, the
same, displayed, X Io.

8 RypBERG, P. A., Contrib. U. S. Nat. Herb. 3:517- 1896.
9 WessTER, J. R., Cleistogamy in Linaria canadensis. Rhodora 2:168. 1900.

460 BOTANICAL GAZETTE [JUNE

twenty to twenty-four inches and to produce branched racemes,
“some of which were a foot or more in length,” they were evidently
of the larger forms, such as grow in the dune region in Indiana. But
in their lack of chasmogamous flowers, they are somewhat different
from any I have noticed there. The cases alluded to in his article
as seen in “other localities” are more like those I have described.
T. S. BRANDEGEE has likewise observed the plant about San Diego,
Cal., “bearing cleistogamous flowers on the lower part of the main
and the whole length of many side branches,”'° apparently more
like the larger forms here. Yet these statements from different
sources indicate that the plant varies somewhat in its behavior in the
respective localities, due perhaps to different environment.

Cleistogamy in Linaria is not confined to our wild toad-flax.
It is one of the forty-four genera given in an article by Kuun in 1867
as producing examples with flowers of this character.*' This list
has been much increased since that date. KuHN does not give the
name of the species, but probably refers to L. spuria Mill., whose
peculiar florescence was described by EuGENE MIcHALET in 1860."?
MICHALET gives it as an example of a plant bearing hypogeous
flowers. “These flowers,” says Darwin, “may be ranked as cleisto-
gamic, as they are developed, and not merely drawn, beneath the
ground.”'3 It also has another peculiarity, according to MICHALET
“infrequent in an annual plant,” that of producing hypocotylous
buds. Its lower leaves are Opposite and much crowded. Two
kinds of branches spring from their axils. Some of these, strong and
often much elongated, spread over the surface of the ground; others
short, slender, and much twisted, with small squamose leaves, gather
in a bunch above the collar of the root, “all with a manifest tendency
to bury themselves in the ground, especially the small hypocotylous
branches which sometimes appear.”” Under suitable conditions they
may penetrate the ground to the depth of 2°™. On account of the

to BRANDEGEE, T. S., Cleistogamous flowers in Scrophulariaceae. Zoe 5:13-
1g00.

't Kumn, M., Einige Bemerkungen iiber Vandellia und der Bliitenpolymorphismus.
Bot. Zeit. 25:67. 1867.

‘2 MICHALET, E., Sur la floraison des Viola... . et du Linaria spuria. Bull. Soc.
Bot. France 7:465. 1860.

"3 Darwin, C., Different forms of flowers 325. 1877.

1909] HILL—POLLINATION IN LINARIA 461

pressure to which they are subjected, the flowers are poorly developed,
but otherwise show nothing peculiar in their structure. The corolla
is crumpled and deformed, but “preserves even its natural color,
with the two brown spots on its upper lip.” The calyx alone loses
its color. Fructification takes place regularly. MucHALET adds that
the phenomenon can be produced at will by heaping earth around
the lower part of the plant, this not interrupting the flowering of
the covered portion. The treading of cattle and the pressure of
wheels bring about the same result. As this plant of Europe and
northern Africa is now introduced into this country, being, according
to Gray’s New manual, “occasional on ballast or waste grounds,”
an opportunity is provided for observing its behavior here. Another
species of north Africa, L. agglutinans Pomel. var. lutea, belongs to
this class of hypogeous plants, as observed by L. TRasut in Algiers.
It has cleistogamous flowers on shoots which spring from the stem
near its base and ripen their fruit underground."

Ill. RELATIVE ADVANTAGES OF THE TWO MODES

It is a distinct advantage to a plant growing under the conditions
of Linaria canadensis to prolong its period of fruiting with a lessened
demand on its supply of food. The environment is xerophytic.
At Oregon it was the southern slope of a steep hill, fully exposed
to the light and heat of the sun. The soil was sandy, and soon parted
with any moisture that was supplied by rains and dews. The con-
ditions in the dunes are similar, the slopes of sand hills or along paths
and roadways in open sunny spots. The growth is usually scattered,
though many plants may form a community, but the ground is not
covered with a dense mat or bed as it commonly is by L. vulgaris. ‘The
slender stems provide but meager shade for the ground about their
roots. In the early part of the season, or if it continues wet, the
radical shoots form rosettes around the base of the stems, which
protect the roots to some extent. In ordinary seasons these soon
wither, and they may not be formed at all on plants which spring up
later, being minute or wanting as in the smaller early plants. It is also
a species poorly adapted to competition. When pressed by perennials,
or by plants disposed to form a close stand, it soon disappears. And

14 KnutH, Handbuch der Bliitenbiologie 32:113. 1905.

462 BOTANICAL GAZETTE [JUNE

the plants associated with it, even if perennials, are not very sturdy
competitors, but mostly of gregarious habit also. At Oregon they
were chiefly Lechea tenuijolia, Talinum leretifolium, Selaginella
rupestris, Silene antirrhina divaricata; in the dunes of Indiana,
Krigia virginica, Arabis lyrata, Viola pedata, Polygonella articulata,
and Festuca octoflora. But to whatever extent the time of fruit-
bearing may be prolonged by cleistogamy, it is comparatively short
in such habitats. It starts early, when there is little competition,
and being an annual or fall-biennial, soon accomplishes its life-
work.

That the cross-pollination of the earlier and larger flowers of
L. canadensis must also be much to its advantage, in increase of
vigor and productiveness, is evident from the nature of this process.
This was clearly proven by Darwin in his experimental work with
cross- and self-fertilized plants. Of two beds of L. vulgaris, raised
respectively from self-fertilized and crossed seedlings, those of the
latter were seen to be much more vigorous. This led him to trials
with this and other plants, the results of which are given in his book
upon this subject. The case of Linaria needs only to be cited. As
showing the vigor, “the naturally crossed plants were to the spon-
taneously self-fertilized plants in‘height, at least as much as 100 to
81.” In regard to fruitfulness similar results came from the two
modes of treatment, that of allowing or preventing the visits of bees.
“The number of seeds in the capsules on the exposed plants to the
average number in the finest capsules on the protected plants was
aS Too to 14,” or as expressed by him in a summary of plants so
treated, the self-fertilized were “ extremely sterile.”*5 KNUTH is
even more emphatic in stating that though self-pollination is possible
and can occur spontaneously in L. vulgaris, it is of little conse-
quence or without result.*° In cases of this kind, where pollination
from without and within takes place simultaneously, HERMANN
Miter thinks it probable that the former preponderates in its
effects, and that the desired result is secured in this way.'7

's Darwin, C., Cross and self-fertilization in the vegetable kingdom 88, 89,
363. 1877. :

‘6 KNutH, Blumen und Insekten auf dem nordfriesischen Inseln rrr. 1894.

17 MULLER, Befruchtung der Blumen 279.

1909] HILL—POLLINATION IN LINARIA 463

IV. LIGHT AND HEAT AS FACTORS IN CLEISTOGAMY

The behavior of Linaria canadensis led to the conclusion that
the gradual diminution in size of flowers was connected with the
increase of heat, and perhaps of the light, to which they are exposed.
Taking the larger plants as typical examples, the two features are
in inverse proportion. This might be taken as a coincidence, but
it seemed to be explained better as a coordination, and more in har-
mony with observations and experiments by others. In 1874 BoucHE
called attention to his observations that the diminution in the size
of flowers and the production of cleistogamy depend in some plants
on the decrease or increase of heat, in others on the decrease or increase
of the length of the day. In the behavior of some, of which Vinca
rosea L. is an example, the light acted favorably, the largest flowers
being formed during the longest days, the smallest during the season
of the shortest days. This seemed to depend on the light, since
with a higher temperature after the longest days had gone by the
_ decrease went on. In other cases cited by him, the decrease in size
and production of cleistogamy are coordinated with the increase of
heat and light, as if these acted unfavorably. As examples of this
are the malvaceous plants, Pavonia hastata Spr. and P. praemorsa
Bo Willd. They begin to bloom at the end of May and show the phe-
~ nomena of diminution and cleistogamy until the autumnal equinox,
after which the flowers gradually increase in size till the beginning
~ of winter or close of their floral season.'® The case of the pavonias
more closely accords with that of L. canadensis, as far as the floral sea-
son of the two coincide. But since, according to Boucuf, the effects
are pat uniform, and may even lead to opposite results with different
ants, there must be something in the plants themselves which
causes the different response, or other environmental conditions
must be taken into account. In the case of Linaria, I had associated
it chiefly with the increase of heat which ordinarily occurs in summer,
| and the consequent diminution or more rapid removal of the moisture
from the soil of such localities as the plants frequent. The equilib-
‘rium between absorption, either from the air or ground, and transpira-
tion is disturbed. The smaller or cleistogamic flower, requiring less
yd, permits a husbanding of resources for the production of seed.

* BoucuE, Gesells. naturf. Freunde go, 91. 1874.

A. er ae ee | oe

464 BOTANICAL GAZETTE [rUNE

The vitality of the plant is lessened, but its ability to bear seed in
abundance still remains. Economy in productive power results
in a prodigality of the means to perpetuate. The waste, seeming
or actual, is seen in the countless numbers of seeds which never have
a chance to germinate. The scattered plants which annually appear
show the need of this productiveness in order to obtain-a few that
can overcome the adverse conditions.

Aside from any effect which the increase of heat and light may
have upon a plant in augmenting transpiration, and thereby making
it advantageous to diminish the exposed surface, it is plain that the
essential organs of reproduction are withdrawn from such effects
far more in cleistogamy than in chasmogamy. As the name implies,
these organs are hidden. But there is also a further tendency in
many cases of cleistogamy to withdraw the perianth, or protective
organs, from the direct effect of the sun’s rays. L. canadensis is an
example of the former tendency, L. spuria and L. agglutinans of
the latter. These two species, as already stated, bend their peduncles
down to produce their flowers or perfect their fruit beneath the sur-
face of the ground. Other well-known examples of this are the
milkworts, Polygala polygama Walt. and P. paucijolia Willd., bearing
their flowers of this kind on subterranean runners. In the violets,
where cleistogamy is so prevalent, the peduncles of the summer
(usually apetalous) flowers are generally much shorter than those of
the large petaliferous blossoms in spring. The flowers are more
or less withdrawn from the light and shaded by the much enlarged
leaves of the summer growth, or they may be borne on stems so
shortened or declined as to be hidden under fallen leaves or buried
in soft humus. The production of the closed flowers under such
conditions may be due to a diminished intensity of light, as far as
this has a bearing on them. Experiments like those of V6cHTING
show that the perianth of flowers is affected by decrease of light more
than the reproductive organs. Chasmogamous flowers may be
made cleistogamous in this way. The violets are quite variable
in their relations to light, many of them being on the borderland
between shade-loving and light-loving plants. The majority of
our wild species bear their petaliferous flowers in the earlier part of
their season of activity, those of the woods before they are strongly

1909] HILL—POLLINATION IN LINARIA 465

shaded by the leaves of the trees and the taller plants of the forest
floor, those of the field or open places before the grass or other growth
overtops them. Their period of cleistogamy occurs when they are
not subject to the strongest light. The one exposed to the greatest
intensity of light, Viola pedata L., differs from most members of the
genus in not having such flowers. Its season of blooming as well
as environment correspond to those of L. canadensis when bearing
its largest flowers. As a perennial, the violet has the advantage
of drawing upon a supply of food stored in its much thickened root-
stock. When this is diminished or too much exhausted, it goes on
with the production of the enlarged summer-leaves, and by them
elaborates another supply of food for storage. This may be a good
explanation of its lack of the cleistogamy so general among its kindred,
since it does not seem adequate to the work of bearing flowers and
perfecting seed while producing the food for the future need of a
xerophytic perennial. Under diminished temperature and favorable
conditions of moisture its work of bearing petaliferous flowers may
be resumed in late summer and autumn, but they are mostly smaller
and much less developed than those of spring. V. /anceolata L. is
also a species frequent in our dune region. It isa light-loving plant,
often greatly exposed in the open sandy border of sloughs, but being
hygrophytic has a supply of moisture on which to draw. Hence
it passes its summer stage in the production of cleistogamous flowers,
which continues long after that of the petaliferous has ceased. Yet
it partakes of the general tendency among the violets, that of bearing
them on shorter, more hidden stems, with the additional habit of
producing them on stolens close to the ground. But L. canadensis,
being an annual subject to xerophytic conditions, cannot draw on
such resources as these two violets have. The development of its
cleistogamous flowers evidently depends on its relations to heat
and moisture more than on those of light.
Vv. DEGENERACY IN FLOWERS OF. LINARIA

In L. canadensis is found an example of a plant passing through

decadent stages to the condition of cleistogamy. The slight irregu-

larity of limb and the occasional remnant of a spur show degeneracy,
even if the smaller and varying intermediate forms of flowers were

466 BOTANICAL GAZETTE [JUNE

not present. It has already been stated that MULLER looked upon
L. minor and L. arvensis, and small-flowered species of Vicia, as
examples of plants which had descended from those adapted by
their floral structure to pollination by insects. In the plant we are
considering, this process is epitomized. Pollination by the help of
insects takes place in flowers of an inflorescence which gradually
undergoes such changes in a single season as to preclude it. The
process of reduction is seen in actual working, and it may: be that
such flowers, rather small at best, are on the way to a stage where
visitation by insects will cease. Yet one cannot regard the explana-
tion as entirely valid. By the very principle of adaptation here
invoked, the opposite might come true; that is, visits by insects,
frequently repeated and continued for a long period of time, would
finally produce flowers better suited to their work. Irregularity
of floral structure is regarded as such an adaptation; and to some
extent has been explained by it. A causal relation between the two
is traced. In Darwiy’s list of genera with cleistogamic flowers,
thirty-two of the fifty-five he gives have the flowers in their most
advanced stage irregular. He says that this “ implies that they have
been especially adapted to fertilization by insects.”?9 Without
pressing such explanations too far, it is seen in the case of the wild
toad-flax that provision for cross-fertilization is made in the structure
of flowers borne simultaneously with the cleistogamous, or at an
earlier date, on the same plant. Ini this there is insured to the species
the present means of invigorating its life, the primary benefit to be
derived from it, whether it be a waning or waxing advantage.
CHICAGO

19 Darwin, C., Different forms of flowers 339.

CURRENT LITERBAIURE

MINOR NOTICES

Botanical expedition to southern Brazil..—The recently published volume
on the results of the botanical expedition of the Royal Academy of Science of
Vienna to southern Brazil in 1901 contains an account of the Pteridophyta and
Anthophyta by Professor R. von WETTSTEIN in collaboration with several promi-
nent specialists. The conditions under which the expedition was undertaken, the
personnel of the exploring party, the detailed itinerary, and the general physical
features of the country visited are briefly set forth in the Einleitung and Reisebericht.
The major part of the volume embodies the taxonomic results of the expedition;
the larger families treated and the cooperating botanists are as follows: the
Filicineae by H. Curist, Orchidaceae by O. PorscH, Gramineae by E. HACKEL,
a. by C. Recuincer, Cyperacede by E. Parra, Malpighiaceae by
, Bromeliaceae by C. Merz, Seaicaak by L. RADLKOFER, Ver-
Diet ri A. von HaveK, Amarantaceae by A. HEmMERL, Gesneriaceae by
K. Fritscu, and the Eriocaulonaceae by W. RUHLAND. Several smaller families
are also included. More than 1300 species are recorded from the various groups

thus far elaborated, and of this number nearly 100 are new to science.
The new species are fully characterized, and the descriptions are mostly in
* Latin; the author of the Cyperaceae, however, has unfortunately chosen to describe
- the new species of this family in German, thus marring somewhat the uniformity
of the work asa whole. The text is supplemented by numerous illustrations, and
certain orchids are beautifully portrayed in color. The publication represents
the work of eminent specialists and forms a reliable and valuable addition to the
taxonomic literature pertaining to the flora of South America. —J. M. GREENMAN.
Vegetation of Java and Sumatra.—The first and second parts of the seventh
_ series of KARSTEN and ScHENCK’s now well-known Vegetationsbilder? is devoted
to a dozen plates eee descriptive text) representing the plants of the volcanic
regions of Java an matra, and especially the reoccupation of those areas
_ which have been at one ‘hoe or another devastated by the erupted solid, liquid, or
gaseous materials. Among others are three views from Krakatoa. This section
by Dr. Ernst sustains the high standard of the work.—C. R. B.

1 WETTSTEIN, R. VON, AND SCHIFFNER, V. Ergebnisse der botanischen enna
der kaiserlichen Akademie der Wissenschaften nach Siidbrasilien a9e%. I. Band.

Akad. Wiss. '79:1-313- pls. 26. I map. figs. 12. 19 rs

2 KARSTEN, G., AND SCHENCK, H., Se aes 4to. VII. Reihe, Heft 1, 2.
| Exxst, A., Die Besiedelung Malbatiachen Bodens auf Java und Sumatra. pls. I-I2.
= Jona: Gustav Fischer. 1909. Ms.

467

468 BOTANICAL GAZETTE [JUNE

NOTES:-FOR STUDENTS

Inheritance of sex.—CorReENs$ has continued his studies on gynodioecious
plants in order to discover what determines the sex of the flowers on the gynomo-
noecious individuals, and the sex of the two classes of individuals belonging to
such species. He finds that the curve of frequency of hermaphrodite flowers in
Satureia hortensis, instead of presenting two modes, as previously reported by
him, one in the mid-season and one at the end of the season, has only a mid-season
mode. ‘The mode which appeared at the end of the season was due to the repeated
counting of flowers which remained open more than one day.

uring the middle of the season no flowers open on the second day, but late in
the season the petals seem to be more resistant, and climatic conditions are less
. severe, so that the same flowers were unwittingly counted several times. When
each flower is marked as it is counted, it is found that the proportion of hermaphro-
dite flowers continues to fall till the end of the season.

He also tested the effect of environmental conditions upon the percentage
of hermaphrodite and female flowers produced on plants of Satureia from day to

day, and noted the position occupied by each kind of flower on the plant. The.

results show that poor nutrition, whether the result of poor soil, insufficient illu-
mination, or disadvantageous position on the plant, lessens the proportion of
hermaphrodite flowers, and under the combined influence of both poor soil and
poor light, only 13 per cent. of hermaphrodite flowers were produced as compared
with 79 per cent. produced under normal conditions of culture. However, the
general features of the curve of frequency of the hermaphrodite flowers remain
the same. With high nourishment the curve for the hermaphrodite flowers falls
much more gradually toward the end of the season, though during the early part
of the season it is not essentially modified.

It was found that different strains of Satureia show marked differences in the
actual percentage of hermaphrodite and female flowers, but that in,each case the
general features of the curve of frequency are the same. The conclusion is

reached that whether hermaphrodite or female flowers are to be produced by a

gynodioecious individual is dependent upon nourishment in its widest sense.
The same general results may be demonstrated in Geranium, Silene inflata and
S. dichotoma, Plantago lanceolata, Scabiosa, Knautia, and Echium

Darwin had observed that a single hermaphrodite plant of Satureia hortensis
was “‘rather larger” than the female plants of the same species, and in an earlier
paper Correns had apparently substantiated this observation, without realizing
the possibility that some of the plants classed as female might be hermaphrodite
plants, rendered apparently female by poor nutrition. He undertook to determine
the relative weights of these two classes of plants with a more careful analysis of
the material. The results show that there is no difference in weight between the

3 CorrEns, C., Weitere Untersuchungen tiber die Geschlechtsformen polygamet
Bliitenpflanzen und ihre Beeinflussbarkeit. Jahrb. Wiss. Bot. 45:661-700. figs. 17+
1908,

itp SS er.

1909] CURRENT LITERATURE 469

female and gynomonoecious plants, and therefore the difference in weight which
was assumed by DARrwIn to be a secondary sexual character has no such signifi-
cance.

CorRENS has also investigated+ the percentage of female and hermaphrodite
plants in Plantago lanceolata under conditions of controlled pollination, and has
shown that while this plant, like Satureia and Silene, shows a marked tendency
for each sex to reproduce its own kind, nevertheless there is considerable variation
in this regard in individuals of both sexes. By pollinating the same female indi-
vidual with different hermaphrodite individuals, and by pollinating different

may be readily calculated, after once the strength of the hermaphrodite tendency
in the pollen-parent and of the female tendency in the pistil-parent is known.
In other words, each individual appears to have a different strength of these two
sex-tendencies and to produce germ-cells of two kinds with respect to these tenden-
cies, the number of each kind of germ-cells produced being perhaps roughly pro-
portional to the strength of the sex-tendencies in the parents.

The theory that the germ-cells of Plantago lanceolata do not t themselves vary
in their tendency to produce a certain sex, but that they are definitely either female
or hermaphrodite, puts these plants into the class known as ever-sporting varieties,
and makes this paper also a valuable contribution to the study of this recognizedly
difficult type of inheritance.

The assumption that each germ-cell is definitely female or hermaphrodite
and that the female is dominant allowed the prediction of the actual numbers
of each sex produced in the different experiments with a fair degree of accuracy.

Several other papers have recently appeared dealing with the question of
sex-determination. DoncasTER and Raynor’ found that in crosses between
Abraxas grossulariata, a common English moth, and its rare variety Jacticolor,
reciprocal crosses are not equal, for when a Jacticolor female is crossed with a
grossulariata male, no lacticolor offspring are produced, and males and females
are all grossulariata; but when the reciprocal cross is made, all of the females
are lacticolor and all of the males grossulariata. To explain this strange situation
the authors assumed that sex is a Mendelian character, and that the lacticolor
character is coupled with the female determinant. It was assumed that
male and female individuals are heterozygous with respect to sex. In this regard
their interpretation differed fundamentally from that of CoRRENS, who assumed
that in the case of Bryonia alba X dioica and other dioecious lant the female

sex is homozygous, and the male heterozygous.
BATESON nye PuNNETT,® in discussing DoncasTER’s results, show that a

4 CORRENS, C, Die Rolle der mannlichen Keimzellen bei der Geschlechtsbestim-
mung der gynodioecischen Pflanzen. Ber. Deu tsch. Bot. Gesells. 26a:686-701. 1908.

s Proc. Zodl. Soc. 1:125. pl. I. 1906.

6 Science N. S. 27:785. 1908.

470 BOTANICAL GAZETTE [JUNE

simpler explanation may be given by assuming that the male is homozygous and
the female heterozygous with respect to sex, and that there is a repulsion between
the determinant for the grossulariata character and that for the female sex. These
assumptions fit all of the facts brought to light in the crosses of Abraxas. This

that this cinnamon canary will find an explanation essentially like that given for
raxas.

Great advances have likewise been made in the study of the determination
of sex from the cytological side, mainly through the work of McCiune, STEVENS,
Morgan, Witson, and their students. In nearly one-hundred species of insects
belonging chiefly to the Hemiptera and Coleoptera, it has been found that there
are definite chromosomal differences between the male and female, and that the
odd chromosomes, or “accessory” chromosomes as they were called by McCiuna,
are so distributed at the time of the reduction division that all the female germ-
cells are alike, while the male germ-cells are of two kinds. The chromosome group
of one of these two types of male germ-cells is like that of the egg-cell, and when
such a sperm fertilizes an egg, a female zygote is produced. The other type of
sperm has a chromosome group unlike that of the egg, and fertilization with such
a sperm produces a male zygote.

An excellent résumé of this work and a discussion of the entire problem of sex-
heredity is given by Witson,° who has been most inently engaged in these new

7 DONCASTER, L., On sex-inheritance in the moth, Abraxas grossulariata and its
var. lacticolor. Reports to the Evolution Committee 4: 537-57- 1908.
* DurHam, F. M., anp Marrvar, D. C. E., Note on the inheritance of sex in
canaries. Reports to the Evolution Committee 4:57-60. 1908.
9 Witson, E, B., Recent researches on the determination and heredity of sex.
Science N. S. 29:53-70. 1909.

1909] "CURRENT LITERATURE 471

quate treatment of such a subject. Wutson’s discussion would have been rendered
simpler and more cogent, if he had grasped the logical homologies between plants
and animals now generally accepted by students of genetics. He does not seem
to appreciate the fact that it is the gametophyte of plants which finds no clear

omologue in animals, and so fails to assign a proper degree of importance to the
parallelism between the sporophyte and the animal body or soma. Agai he
h

with Mendelian heredity. Instead of this, he presents as a ‘‘naive assumption”
what is now generally held by the students of Mendelism, and known as the “‘pres-
ence and absence hypothesis,” the assumption being that the heterozygote and
the positive homozygote differ from each other in that the former has an unpaired
unit, X, and the latter a pair of units of the same kind, XX.

CASTLE’? takes up this question and shows the perfect agreement between
the results of these cytologists, and the requirements of the presence and absence
hypothesis in Mendelian heredity. In CastTLe’s exceedingly clever discussion an
attempt is made to harmonize the apparently antagonistic results with Bryonia,
the Hemiptera, and Coleoptera on the one hand, and those with Abraxas and the
cinnamon canaries on the other, by assuming that in all cases the female possesses
one more unit than the male, this unit being called by Wirson the “X-element.”
Bryonia, and all of the insects whose male germ-cells have been found to be of
two kinds, represent a condition in which the male is a heterozygote, and the female
is a positive homozygote. Caste calls this a “dominant female,” but this is
obviously a misleading terminology, for if the female were really dominant the
heterozygote would also be a female and there could be no males. In Abraxas,
and the cinnamon canaries, and, as suggested by CASTLE, perhaps also in the
pheasant, the female is Retecnygous and the male is assumed to be a negative
homozygote, i. e., wholly lacking the X-element. This is the most promising
attempt yet made to bring all the recently discovered facts of sex-heredity in dioe-
cious animals and plants under a single hypothesis.

CasTLE attempts further, by an extension of the same hypothesis, to account
for the fact that male animals usually possess more characters than the female.
He supposes that these added male characteristics are associated with or produced
by a Y-element, the “synaptic mate” of the X-element. He also suggests that
progressive evolution may have taken place by the appearance and development
of such a “synaptic mate” for the X-element, but this, and also the attempt
to explain orthogensis on the same basis, is carrying hypothesis rather far from
empirical knowledge.

ere can be no question that the problems of sex possess many intricacies
and difficulties yet to be solved, but the results of these investigations both from
the experimental and the cytological side have placed these problems on a new

10 CasTLE, W. E., A Mendelian view of sex-heredity. Science N.S. 29:395-400.
1909.

472 BOTANICAL GAZETTE [JUNE

basis, and opened up many possibilities and suggestions for their further investi-
gation. All of the results seem to point to the truth of the view that sex is

redetermined in the germ-cells, and that therefore it cannot be modified by
environmental conditions except, of course, by such conditions, as yet unknown,
as are capable of producing mutations—Grorcr H. SHULL.

Current taxonomic literature.—N. L. Brirron and J. N. Rose (Jour. N. Y.
Bot. Gard. 9:185-188. 1908) have proposed a new genus (Carnegiea) of the
Cactaceae. The genus is based on the well-known Cereus giganteus Engelm.,
and contains but the one species. H. Prrtrer (Contr. U. S. Nat. Herb. 12: 17I-
181. 1909) has published 8 new species of flowering plants from tropical America.
The descriptions are supplemented by two full-page illustrations and several
text-figures; the types are deposited in the U. S. National Herbarium. A. THEL-
LUNG (Bull. Herb. Boiss. II. 8:913, 914. 1909) records 3 new varieties of Lepi-
dium pubescens Desy. from South America. F. STEPHANI (tbid. 941-972) has
published 43 new species of the genus Mastigobryum from various localities.
G. BEAUVERD (ibid. 986-988) has published a new Eriocaulon from Brazil and
also a new species of Tulbaghia from the Transvaal ; the same author (zbid. 993-
1007) records 8 new species and one variety of Nothoscordum from Uruguay and
gives an analytical key to the Uruguayan species. E. G. Parts (Bull. Soc. Bot.
France IV. 8:Mém. 14, pp. 1-66. 1908), under the title Florule bryologique de
la Guinée frangaise, has published 6 new species of mosses. F. GAGNEPAIN (ibid.
Session extr., pp. xxxvi-xliii) has published 4 new species of Zingiberaceae and a
new genus (A/aenidia) of the Marantaceae from Africa, and also a new species
of Calathea native of Indo-China. G. Bonatr (ibid. 509-515, 537-543) describe

25 new species and 4 new varieties of scrophulariaceous plants from Indo-China.

variety of ferns as new to science. B. P. G. HocHREUTINER (Ann. Conserv. et
Gard. Genbve 11-12: 136-143, reprint pp. 1-8. pls. 1, 2. 1908) has published a
revision of the genus Adansonia in which 8 species are recognized, one of which,
A. Stanburyana from northwestern Australia, is proposed as new to science.
ms fe Napson (Bull. Jard. Imp. Bot. St. Petersb. 8: 113-121. pl. 1. 1908)
describes a new microorganism (Rhodosphaerium diffluens) from the Caspian
Sea; the systematic position of the plant according to the author is ‘‘an der Grenze
zwischen Algen und Bakterien.” C. FERDINANDSEN and O. WINGE (Bot. Tids-

1909] CURRENT LITERATURE 473

the years 1905 and 1g06. W. Fawcett and A. B. RENDLE (Journ. Botany
47:3-8. 1909) have published diagnoses of 12 new species of orchidaceous
plants from Jamaica; these are preliminary to a monograph of the orchids of
Jamaica. E. Ure (Engl. Bot. Jahrb. 42:191-238. 1908), in collaboration with
different specialists, under the title Beitrdége zur Flora von Bahia I, has published
75 species and one variety as new to science; these are referred to families in the
Engler sequence from the Bromeliaceae to the Araliaceae and include the follow-
ing new genera: Sincoraea and Cryptanthopsis (Bromeliaceae), Heptocarpum
(Capparidaceae), and Jtatiaia (Melastomaceae). E. KorHne (ibid. Beiblatt
97:47-53) records 5 new species and 4 new varieties in the Lythraceae from Sout
America, Africa, and Siam. Different authors (Kew Bull. 1908: 432-441),
under the title Diagnoses Ajricanae XXVI, have published 19 new species and
one variety of African angiosperms, including 2 new genera (Aristogeitonia and
Androstachys) of the Euphorbiaceae; also (ibid. 445-451) in Decades kewensis:
Decas LI, 10 new species are described from various localities. E. L. GREENE
(Leaflets Bot. Obs. & Crit. 2: 1-24. 1909) proposes a series of 60 new species and
3 new varieties of flowering plants, chiefly from western United States. J. Born-
MULLER (Mitt. Thiir. Bot. Ver. 23:1-27. 1908), in continuation of his contribu-
tions under the title Novitiae Florae Orientalis, has published 23 species as new
to science, of which 17 belong to the genus Astragalus. A. SCHERFFEL (Ber.
Deutsch. Bot. Gesells. 26a: 762-771. 1909), proposes a new genus (Asterococcus)
for the alga hitherto known as Pleurococcus superbus Cienk. N. L. Britton
Bull. N. Y. Bot. Gard. 5:311-318. 1909), in continuation of his studies on the
flora of the Bahamas, has described 6 new species of flowering plants. F.S. EARLE
(ibid. 373-451), under the title of Genera of the North American Gill Fungi,
recognizes 147 genera for North America, and of these 38 are designated as
new.—J. M. GREENMAN.

Hybrids of Oenothera.—DeVrtes has recently published several remarkable
papers on hybridization in Oenothera. The results concern a new type of heredi-
behavior, which is of great interest, showing as it does that we are only on the
borderland of knowledge in these fields. Such discoveries, which open new vistas
for the future, are of special value as a stimulus to research. The first of these
papers appeared in this journal't and announced the discovery of what are called
twin hybrids, and a later paper"? dealt with triple hybrids. In certain cases, when
one of the wild species of the Onagra group is crossed with O. Lamarckiana or
one of its mutants, two types are produced in about equal numbers, both of which
breed true, the same types appearing in the different crosses. These types
DeVrres calls O. Jaeta and O. velutina. In the case of O. scintillans and O. lata

1« DeVries, Huco, On twin hybrids. Bot. GAZETTE 44:401-407. 1907-
, On triple hybrids. Bot. GAZETTE 4721-8. 1909.

474 BOTANICAL GAZETTE [JUNE

triple hybrids are produced—in addition to O. laeta and O. velutina a third type
resembling the mother (O. Jata or O. scintillans), but in its special marks inter-
mediate between its parents.

The twin hybrids of O. nanella have been worked out most completely.t3
O. muricata X O. nanella produces the two types Jaeta and velutina, about 50
per cent. of each. The Jaeta breed true for four generations, but the velutina
split in the F, and all succeeding generations, producing velutina and something
over 50 per cent. of a form called by DEVriEs O. murinella, which is a dwarf
O. velutina and breeds true. The dwarf character (but not the other O. nanella
characters), therefore, reappears in over half of each generation. The discoveries
of greatest interest follow. O. velutina X O. murinella gives the same results as
O. velutina self-pollinated, i. €., over 50 per cent. O. murinella. From this the
conclusion is drawn that the pollen of O. velutina has the same hereditary qualities
as that of O. murinella. The reciprocal cross gave Ioo per cent. O. murinella
(280 plants). The facts are all explained by assuming that the egg cells of O. velu-
tina are of hybrid (heterozygote) nature (a x 6), while the pollen bears only the
dwarf character (a). On self-pollination O. velutina would then give 50 per cent.
O. velutina (a X 6) and 50 per cent. O. murinella (a) which breed true. This

Similarly, O. laeta crossed with O. murinella, O. nanella, or O. velutina gives
5° per cent. laeta and 50 per cent. dwarfs. Therefore the egg cells of Jaeta are
also hybrid (heterozygote) in regard to the dwarf character, although when self-
pollinated Jaeta breeds true! The pollen of Jaeta therefore bears the hereditary
characters for high stature. This dominates over the hybrid nature of its own
egg cells, but is recessive to the egg cells of pure dwarfs. The remarkable situa-
tion therefore appears, that the egg cells of both velutina and laeta behave as
heterozygotes, while the pollen of the former behaves as though it carried only
the dwarf character, and the pollen of Jaeta appears to carry only the character
for high stature.

Another paper'+ shows that the hybrids of O. gigas behave differently from
those of other mutants. QO. gigas X O. Lamarckiana forms a constant inter-

'3 DeVries, Huco, Ueber die Zwillingsbastarde von Oenothera nanella. Ber.
Deutsch. Bot. Gesells. 264:667-676. 1908. :

4———-, Bastarde von QOenothera gigas. Ber. Deutsch. Bot. Gesells. 26a:
754-762. 1908.

®
Se A ae ES nN, ee ee aa eee ee

1909] CURRENT LITERATURE 475

Miss Lutz's studied forty individuals of O. lata X O. gigas and describes a
group of hybrids which probably include both the types of DeVries, and in
addition plants like O. Jata and like O. gigas, having their respective numbers of
chromosomes.—R. R. GATEs.

Nitrogen fixation by Azotobacter.— KRZEMIENIEWSKI has contributed a paper'®
that seems to throw much-needed light on the problems of nitrogen fixation by
Azotobacter in the soil. Perhaps its most valuable feature is the demonstration
of the accelerating influence of humus on the process. He finds that Azotobacter
in pure cultures in ordinary nitrogen-free media can fix little atmospheric nitrogen,
but that the addition of sterile soil or of humic acids or their calcium, potassium,
or sodium salts to such solutions multiplies the amount of nitrogen fixed many
times. It is interesting to note further that the humus derived from different soils
does not yield uniform results, and that artificial “‘humus” formed by the action
of acids on carbohydrates is of little or no value. LrpmMan’? in this country has
anticipated in part some of these results, for he found that the amount of nitrogen
fixed by Azotobacter growing in solutions to which different types of soils had
been added varied greatly. KrzEMIENIEWSKI further reaches the interesting
conclusion, as a result of repeated experiments, that humus does not serve either
as a source of nitrogen or of carbon for Azotobacter. He finds that the various
nitrogen compounds used in an effort to duplicate the stimulating influence of
humus are without such results. When these compounds were used in conjunction
with humus they were found to be even decidedly inhibitory in action. Why
the humus should thus stimulate growth of Azotobacter he fails to explain,
although he seems to have had abundant experimental evidence of the fact. The
author was able to demonstrate as much as seventeen milligrams of nitrogen fixed

in the amount of nitrogen supplied to the culture. The organic nitrogen fixed
in the culture solution was found at the close of the experiment to check very
closely with the amount which disappeared from the air.

The organism is a strict aerobe, neither alcohols, acids, nor hydrogen gas are

found as products of metabolism. The ratio 76) approaches unity. The

temperature optimum for nitrogen fixation is 28°C. Prolonged cultivation of
Azotobacter on artificial media the author finds has little influence on its ‘“viru-

1s Lutz, ANNE M., Notes on the first generation hybrid of Oenothera lata X O.
gigas. Science N. S. 29:263-267. 1909-

16 KRZEMIENIEWSKI, S., Untersuchungen iiber Azotobacter chroococcum. Beij.
Bull. Acad. Sci. Cracovie Cl. Sci. Math. et Nat. 1908: 299-1050. pl. I.

17 LIPMAN, JAcoB G., Bacteriological indications of the mineral requirements of
soils. Ann. Rep. N. J. State Agr. Exp. Sta. 2'72177-187. 1906.

476 BOTANICAL GAZETTE [JUNE

lence” or ability to fix nitrogen. The contradictory results of other investigations
are probably due to the use of different strains of organisms. The differences
between these strains are varietal or, possibly, even specific, for cultures from
various sources differ considerably in their nitrogen-fixing power.

BEIJERINCK in his work on these organisms proposed a theory of symbiotic
activity between Azotobacter and other bacteria. KRZEMIENIEWSKI concludes
that BEIJERINCK’s results are to be explained not as due to the presence of the
second organism but to the addition of humus to culture solutions.

These studies should serve to emphasize further the importance of soil humus
from the standpoint of agricultural practice. In addition to the solution of
certain puzzling questions, KRzEMIENIEWSKI has opened several very promising
avenues for successful and profitable research in soil bacteriology.—RoBERT E.
BUCHANAN

Mutability and variability.—ScHourEN™ has an extensive account of two
years’ Oenothera cultures. Seeds from DEVrrEs, as well as commercial seeds
and “wild” seeds, and rosettes of various species were used. Several new mutants
appeared, and a number of interesting combination forms possessing the char-
acters of two types are recorded. The new mutants are O. blanda and O. cande-
labrijormis, while the combination forms include O. laevifolia brevistylis, O.
laevifolia nanella, O. rubrinervis brevistylis, O. rubrinervis lata, O. gigas nanella,
and O. gigas lata (?).

He makes the suggestion, which appears rather unlikely, that the nanella
or dwarf condition in OO. Lamarckiana, laevifolia, and gigas may be due to
bacterial action. O. Lamarckiana nanella is found to exist in two forms, differing
in their bud and flower characters. O. gigas is well known to show extremely
wide variability, particularly in leaf shape, and an attempt was made to segregate
several types, but without success, since the offspring from each showed nearly
the whole range of variation.

The occurrence of a number of combination types as mutants in pure strains
gives a somewhat different appearance to the mutation phenomena in Oenothera.
SCHOUTEN concludes that mutants originate by two different methods: (1) When
both the gamete nuclei uniting in fertilization have the constitution of the same
mutant. (2) When the gamete nuclei are unlike. Of the latter he classifies two
sorts. (a) When one gamete nucleus has the constitution of the species and the
other that of the mutant. (6) When both the gametes have a mutant constitution
but not of the same mutant, thus accounting for the combination forms. Further
evidence is obtained from the fact that crossing increases the production of
mutants.

The third part of the contribution deals with statistical studies of variability
in O. Lamarckiana and its mutants, and in several wild species. The parts
measured include the length and breadth of certain stem leaves selected according
to a definite rule; the length and breadth of the petals and sepals of certain selected

*8 ScnouTen, A: R., Mutabiliteit en variabiliteit. pp. 196. Groningen. 1908.

1900] CURRENT LITERATURE 477

flowers; the number of stigma lobes; the length of style, hypanthium, and ovary;
length of main stem; number of side branches, etc. A large number of interest-
ing data of variability are here brought together. It is of interest that in nearly
all cases the modal number of stigma lobes shows a decrease from 6 or 8 or more
to 4 during the season.

The work is an extension of SHULL’s’® statistical studies. SHuLt found that
in the characters measured, the mutants’ of O. Lamarckiana are more variable
than the parent form, and hence that phylogenetically younger forms are probably
more variable than the phylogenetically older. This appears to hold for the
European O. biennis and its mutants cruciata and suljurea, but is only partly true
for O. Lamarckiana and its mutants. O. gigas and O. rubrinervis lata are more
variable than the parent in all the organs examined; but the other mutants are
more variable in some characters and less so in others. The coefficient of vari-
ability of a mutable species is not markedly different from that of a non-mutable
form.—R. R. GATEs.

Ontogenetic theory of alternation.—LANG?° has outlined an interesting
theory of alternation of generations which he calls an “ontogenetic” theory, to
distinguish it from other theories. The so-called “homologous” and ‘‘antithetic”
theories are well known, and Lanc’s work on apogamy in ferns inclined him to
accept the former. In fact, the ontogenetic theory is a theory of homologous
alternation in its phylogenetic application.

The author starts with the idea that all the cell progeny of a germ cell are
potentially similar, and that any one of them might reproduce the organism.
The development of a specific organism is regarded as due to the properties of the
germ cell and to the conditions under which the germ cell develops. The author,
therefore, reaches the conception of a specific cell corresponding to each specific
form, to which under normal conditions it gives rise. In plants with a definite
alternation of generations, germ cells capable of developing into an organism are
met twice in the life-history. The organisms developed by these two cells may
be very similar or very different. For example, in Polysiphonia the two resulting
organisms are very similar; while in bryophytes and pteridophytes they are very
different. To explain the latter case the author recognizes two alternative views:
(x) the two germ cells are so different that they necessarily produce different
structures; (2) the two germ cells are both specific cells of the same plant, but the
conditions of development are so different that the two resulting organisms are
very different. :

-9 MacDoveat, D. T., et al., Mutants and hybrids of the Oenotheras. Pub. No,
24, Carnegie Institution. | p. 57. figs. 13. pls. 22. 1905; Mutations, variations and
relationships of the Oenotheras. Pub. No. 81, Carnegie Institution. pp. 92. pls. 22.
figs. 73. 1907

20 LANG, W. H., A theory of alternation of generations in archegoniate plants
based upon the ontogeny. New Phytol. 8:1-12. 1909.

478 BOTANICAL GAZETTE [JUNE

The second alternative is the basis for the proposed ontogenetic theory of
alternation, the assumption being that the two germ cells of a life-history, althoug

one is haploid and the other diploid, have potentially the same morphogenetic
properties, and under the same conditions would produce similar structures.

fertilized egg develops in relation to the body of the sexual generation. The egg
is thus removed from all the influences acting on the spore, and is exposed to a new
set of nutritive and “correlative” influences proceeding from the parent body.
Since each stage in the ontogeny is probably determined by the preceding stage,
the general structure of the resulting organism is as fully determined by the rela-
tively short association of egg and gametophyte among pteridophytes, as by their
much longer association among bryophytes.

It is evident that this theory regards the two generations of each species as
homologous, in that they are developed from germ cells with the same morphoge-
netic powers. The really important comparisons to make, therefore, are between
gametophyte and sporophyte of the same plant; rather than between the sporo-
phyte of ferns and gametophyte of liverworts, for example.. The author promises

tails, comparing the two generations in each group in a most suggestive way.
If the chief value of a theory lies in the work it stimulates, this ontogenetic theory
should prove of great value, for it opens a large vista of experimental work.—
LAS

all its aspects, presenting under the title of morphology, not only the gross and
minute features of its morphology, but also its anatomy, ecology, and physiology.
Such a compendium of statements in reference to a single species is unusual,
for in general an investigator in these days is compelled to restrict his attention

and contrasted with those of other Potamogetonaceae; Ecology of the vegetative
organs (13 pp.), in which the hydrophytic and halophytic adaptations are pre-
sented, and the difficult problem of adaptation and heredity discussed; Repro-
ductive organs (18 pp.), in which flower, sporangia, gametophytes, fertilization,
and endosperm are described; Embryo; Fruit and seed; Seedling. The paper
closes with two summaries and a bibliography of 98 titles.
ORs RR Sh

21 GRAVEs, A. H., The morphology of Ruppia maritima. Trans. Conn. Acad.
Sci. 14:59-170. pls. I-15. figs, 33. 1908.

1909] CURRENT LITERATURE 479

Some of the ecological results are as follows. Ruppia is called a water halo-
phyte, living in salt water that would produce plasmolysis in fresh-water plants,
but unable to live in water of the open ocean. The hydrophytic responses of the
shoot are the weak and spreading form, the absence of stomata, the production of
slime, the numerous air spaces, the lack of mechanical tissue, and the reduction
of the vascular system to one axial bundle and two lateral ones in both stem and
leaf. The responses of the root are a reduction of the system to small unbranched
roots borne singly at the nodes, the presence of air spaces, and the concentric
axial bundle. The axial and cortical bundles are thought to be useless hereditary
structures.

Some of the facts in reference to the reproductive structures are as follows.
The inflorescence is a reduced spadix, and a small spathe is present, which is said
to have escaped the notice of investigators almost entirely. In the development
of the microsporangium a large archesporial group of cells is differentiated, which
later becomes septate. In the development of the megasporangium, usually
only one layer of parietal cells is formed, and in one case two functioning mother
cells were observed in a sporangium. The count of chromosomes was made in
the microsporangium and in the reduction divisions of both gametophytes, and
was found to be eight and sixteen. The male cells are produced before pollina-
tion, which is accomplished by means of the water. The endosperm is scanty,
never being more than a thin layer lining the sac. The proembryo is a filament
of three or four cells, the basal one becoming much enlarged to form the suspensor-
The three embryo-forming cells produce at first a spherical group of cells, and it
is believed that both cotyledon and stem tip are derived from the terminal cell
of the proembryo, the two other cells producing the hypocotyl, adventitious root,
and primary root, the last organ never functioning.

The paper contains a large amount of information in reference to a very
interesting form, and the plates, some of them photomicrographs, reproduce the
structures in such a way that every botanist can make his own interpretations.
—jJ.M

Orchid flowers and formative stimulii—As a product of his visit of three
months at the Buitenzorg Garden, Frrtinc published in the initial number of
the new Zeitschrift fiir Botanik an account of his experiments on the effect of
pollination and other stimuli upon the postfloration behavior of the flowers of
orchids.?? The tropical orchids, available in great abundance at this garden,
are especially suited for experimental study on this point, because the difference
in duration of pollinated and unpollinated flowers is sufficient to give opportunity
for experimentation with unequivocal results, whether the postfloration processes
are autonomous or induced. Of these processes he distinguishes four : (1) pre-
mature fading; (2) closure of the stigma and swelling of the gynostemium,
(3) swelling of the ovary; (4) greening of the perianth.

22 Fittinc, H., Die Beeinflussung der Orchideenbliiten durch die Bestaubung
und durch andere Umstiinde. Eine entwickelungsphysiologische Studie aus den
Tropen. Zeits. Bot. 1:1-86. figs. 27. 1909.

480 BOTANICAL GAZETTE [JUNE

He was able to induce premature fading by applying most various substances
to the stigma: besides their own living pollen, volcanic river-sand, spittle, dead
pollen and pollen extract, dead and leached pollen of the same species or of
other genera or even of other and remote groups, and extract of gynostemium
tissues induced it, and apparently also 5 per cent. saccharose. He was not able
to determine what the chemical agent or agents were in these reactions. Wound-
ing the stigma or the tissue at the apex of the gynostemium also caused premature
fading. Closure of the stigma and swelling of the gynostemium could be effected
by bestrewing the stigma with living or dead pollen of orchids (any genus) or
even of Hibiscus, and with the alcoholic extract of pollen. On the contrary, dead
pollen and pollen extract had no effect or the very slightest in inducing swelling
of the ovary, which occurred only when living pollen germinated on the stigma
and its tubes grew into the ovary. The greening of the perianth (peculiar to
certain species) appears only when the ovary has previously begun to swell and
to turn green.

Firtrnc considers fading as the end process of floral development, gid
released by the pollen stimulus or others earlier than it is autonomously.
stimulus, however, does not merely hasten development; it diverts its course,
for a perianth half open and quite incompletely developed may be made to fade
in twelve to twenty-four hours by a stimulus which proceeds from the distant
stigma. This also offers a new example of the separation of perceptive and
reactive regions. The closure of the stigma, etc., appears to be strictly a case

perianth, depend on the penetration of the pollen tube; but whether the stimulus
is mechanical or chemical does not appear

The prompt fading of the flowers, pesble after an insect bite on the stigma
or after stimulation by foreign pollen, and the small crop of fruit on these tropical
orchids, awaken doubts in Frrrinc’s mind as to the validity of the teleological
interpretation of the elaborate mechanisms which are believed to secure cross-
pollination. Perhaps they were effective in a past age when insect life was
richer, he adds by way of apology for his temerity in suggesting such heresy.
He will find this heresy not unwelcome, we imagine, in this country, where ecolo-
gists are questioning whether there is even adequate proof that cross- -pollination
is advantageous.—C. R. B

tology of Oenothera.—GerErts?3 published an account of embryo-sac
development and chromosome reduction in Oenothera. A row of four mega-
spores is formed, with typical reduction phenomena, the megaspore nearest
the micropyle forming the embryo sac. Its nucleus divides only twice. Both
23 GEERTS, J. M., Beitrage zur eeu der cytologischen Entwicklung von
Oenothera Lamavcltana. Ber. Deutsch. . Gesells. 26a:608-614. 1908.
———., Beitrage zur Kenntnis der i. und der partiellen Sterilitat von
Oecnothera Laidarekianie Separate (source unknown). pp. 116. pls. 5-22. 1909-

1909] CURRENT LITERATURE 481

nuclei resulting from the first division remain in the upper portion of the sac. One
of these divides to form the two synergids, and the other forms the egg and polar
nucleus. There are therefore no antipodals and only one polar nucleus. In
fertilization one male nucleus unites with the egg; the other unites with the polar
nucleus to form the endosperm

The account of reduction pliant confirms, in all the main points, the
previous accounts of GaTEs.?4 There is no fusion of parallel threads in synapsis.
The spirem later breaks into the vegetative number of chromosomes, which after-
ward become paired. The first mitosis separates whole chromosomes, and the
second separates the longitudinal halves of these. Certain critical stages during
the period between synizesis and diakinesis, which prove that the chromosomes
are formed by the segmentation of a single spirem thread, are not represented;
but these stages are the most difficult to obtain, probably because they are passed
through quickly. It seems now pretty evident that there are two general methods
of reduction in plants, as in animals, one involving a telosynapsis, the other a para-
synapsis or side-by-side pairing of chromosomes.

The question of sterility is also examined, with interesting results. In Oeno-
thera Lamarckiana 50 per cent. of the ovules are found to degenerate, and about
50 per cent. of the pollen grains—two from each tetrad of spores. A large num-
ber of other Onagraceae were examined, nearly all of which were found to exhibit
more or less sterility. GrrRTs concludes that the sterility of O. Lamarckiana
cannot be explained as the result of hybridization, cultivation, or lack of nutrition
or space, but that it has been inherited from a remote ancestor, probably from
the ancestor.of the whole sub-family. He-thinks that since this sterility is herit-
able it must have originated by a mutation, or rather two mutations, one on the
pollen side and one on the megaspore side, since they are often sterile in different
degrees in the same species!

So far from explaining anything, it seems to the reviewer that this muddies
the pool and is much worse than a flat confession of ignorance as to the cause.
It will be unfortunate if botanists acquire the habit of ascribing the origin of com-
plex conditions, such as sterility, toa sudden ‘‘mutation” in some ancestor. There
is no evidence to show that the sterility has not been gradually acquired, and for
that matter independently acquired, in the different species. To call it a mutation
helps to explain neither its origin nor its cause.—R. R. GATEs.

Seedlings of conifers.—HiLt and FRAINE?5 have published a second paper
on the seedlings of gymnosperms, the thesis being that polycotyledony is attained
by the splitting of preexisting members, which were probably two in number.

n the present investigation seedlings of Tsuga, Abies, Picea, Cedrus, Pinus,
Larix, Pseudolarix, and Araucaria were studied. The general result shows that

24 GATES, R. a ty Paces of reduction in Oenothera rubrinervis. Bot. GAZETTE
46: 1-34. pls. I-3.

2s Hitt, T. G., AND DE FRAINE, E., On the seedling structure of gymnosperms.
emma of Botany 23:189-227. pl. 15. figs. II. 1909.

482 BOTANICAL GAZETTE [JUNE

the Taxineae, Podocarpineae, and many Cupressineae have two cotyledons, and
that each cotyledon (Podocarpineae being excepted) contains one vascular strand
and the primary root is diarch. Among the Abietineae, however, in which poly-
cotyledony prevails, each cotyledon has a single vascular strand, but the number
of poles of the primary root holds no obvious relation to the number of cotyledons.

In summarizing the evidence of splitting, the authors add the following
testimony: the occurrence of partially split cotyledons, the frequent obvious
grouping of cotyledons, and the cases of transition. Pinus contorta Murrayana
may be selected as an illustration of the last case, in which form three entire
cotyledons were found, one of them much larger than the other two and contain-
ing two entirely distinct vascular strands. The authors speak of this as a case
of one whole cotyledon and two half-cotyledons. Trouble of course comes with
the higher numbers of cotyledons, and at this point the explanation offered is not
clear. It is acknowledged that in some cases an increased number of cotyledons
may result from the appearance of extra primordia, which represent the dis-
placement of foliage leaves from the first stem node to the cotyledonary node.

The general summary of facts contains the following items: the occurrence
of more or less complete cotyledonary tubes (over 20 species cited); the existence
of cases of incomplete splitting (4 species cited); the general presence of cotyle-
donary resin ducts (several in araucarians, two in 12 species cited, one in 6 species,
and none in 6 species or more); the occurrence of 4-8 vascular strands in each
cotyledon of Araucaria, and of one strand in the cotyledons of Tsuga, Abies,
Picea, Cedrus, Pinus, and Larix; the occurrence of mesarch structure in occa-
sional cotyledons of Tsuga canadensis, Pinus Pinea, and P. Gerardiana.

SHAw?° has investigated the seedling of Araucaria Bidwillii, a tuberous
species and one not studied by Hitz and Fraine. He finds that the salem:
vascular strands are very numerous and ourlable (12 to 16), that the poles of
root are equally variable . to 7), and that there is a very confused aaa 2
between the two. The protoxylem groups of ae root are gradually reduced until
the diarch condition is finally attained. on], M

The Piccard rotation experiments.—HABERLANDT?’ has repeated PiccARD’s
rotation experiments,?* for which he used the seedlings of Vicia Faba, Lupinus
albus, and Phaseolus multiflorus. He characterizes Piccarp’s conception as
good, but the execution of the experiments and the interpretation of the results
as faulty.

He claims to. have eliminated all Piccarp’s technical errors by devising a
very substantial and accurate centrifuge, and by securely fastening the seedlings

26 SH , ee 4. 8. The a structure of Araucaria Bidwillit. Annals of
ssi 23: Pais pl. 21. figs.
27 HABERLANDT, G., Ueber die Yr der geotropischen Sensibilitit in der
Wurzel. Lh Wiss. Bot. 45:575-600. I9g09.
CARD, AuGust, Neue Versuche iiber die geotropische Sensibilitat der Wurzel-
spitze. Jahrb. Wiss. Bot. 40:94—r102. 1904.

1909] CURRENT LITERATURE 483

with plaster of Paris, at the same time aeoae' them with sufficient moisture.
The roots were placed at an angle of 45° the axis, as were PIccARD’s.
While PiccarD used 20-40 rotations per seoendl HABERLANDT found it neces-
sary to use only 5~20.

If the tips of the roots extended 1.5™™ or more beyond the axis, they always
bent in the direction according with the irritability of the tip; if 1™™ or less, the
curvature was determined by the irritability of the growing zone. HABERLANDT
points out (what he says Piccarp and his critics have failed to notice) that where
the root tip extends r.5™™ beyond the axis, the growing zone receives on the aver-
age greater stimulation than the root tip, the centrifugal acceleration of the grow-
ing region, by reason of its greater length, being 2.8-3.9 times that of the tip.
This, of course, is due to the considerable length of the growing zone. HasEr-
LANDT never gets the S$ curve described by Piccarp. He concludes that 1.5-
2™™ of the root tip, in the forms worked with, is very sensitive to gravity and to
centrifugal acceleration. The growing zone is likewise sensitive, but far less so
than the tip. The marked geotropic sensitiveness of the tip corresponds to the
well-developed statolith apparatus of the cap, while the slighter sensitiveness of
the growing zone is due to the rather poorly developed statolith starch of the
periblem in that zone.

He finds that the geotropic irritability of the growing zone is manifested with
acclerations as low as 0.25 gravity, and therefore that it comes into play in ordinary
geotropic response, exactly opposite to NEMEC’s conclusion.

ABERLANDT also conducted a set of decapitation experiments, making full
allowance for the shock effect of decapitation, which accord fully with the results
by the Prccarp method. He concludes that all these results are quite in harmony
with the statolith theory —Wm. CROCKER.

Plant proteases.—VINES has now for more than ten years devoted his atten-
tion to the proteases of plants and he has made the field practically his own. The
conclusions he has from time to time announced mark periods in the develop-
ment of the problem. The last paper by this author?® should be considered in
two parts, the first of which deals with his latest results, while the second consti-
tutes a review of the earlier investigations, together with final conclusions.

The papain or papayotin of the latex of papaw, which has long been known
to digest proteins, was shown by Martin to be both peptic and peptolytic. It
was therefore designated a tryptic enzyme. The discovery that other vegeta
extracts (germinated lupin, castor-bean, some fruit juices, malt, yeast) had a
like action, led to the notion that plant proteases in general are tryptic. This
conception, although a generalization from too limited data, was an advance, as
it supplanted the prevalent idea (also resting upon an insecure foundation) that
plant proteases are peptic. Following up his work on tryptic extracts from vari-
ous sources, VINEs has finally come to believe that the proteases of plants are of
two sorts, the peptases and the ereptases. This conviction has been further fixed

20 Vines, S. H., The proteases of plants. VI. Annals of Botany 23:1-18. 1909.

484 BOTANICAL GAZETTE [JUNE

by his latest results, which are that from papayotin both peptase and ereptase
may be obtained. The former is soluble in dilute NaCl and little soluble in dis-
tilled water, while the latter is easily soluble in pure water. That the demon-
stration of these two proteases in papayotin might be more complete is admitted
by the author. ead preparations from both fresh and dry yeast show the
presence of peptase tase.

The proposal of fae. after reviewing the subject, to supplant the ‘‘ vegetable
trypsin” idea by the conclusion that the proteases of plants belong to two main
groups, the peptases and the ereptases, and his further classification of the former
into endopeptases and ectopeptases, appeal to the reviewer as unnecessary and
unwarranted, inasmuch as the new may prove to be as incomprehensive as was
the “trypsin” idea. Further, if a name must be given to something of which
little is known, that name should have some reference to the qualities marking
individuality, rather than to the mere incident of its occurrence. So far, the
“ectopeptase” is confined to Nepenthes. The anticipation of the author that
“ectopeptase” is of wide occurrence may be justified, but in one case, namely,
the pitcher-liquid of Sarracenia, peptic action has not been found .3°—RAYMOND
H. Ponp.

Morphology of Pseudotsuga.—The investigation of the North American
representative of this interesting genus by Lawson3" has filled a gap in our
knowledge. In general it conforms to the well-known characters of Abietineae,
but it presents some interesting peculiarities. The pollen grains are wingless,
and the mechanism for receiving them is most unusual. There is a stricture of
the integument above the nucellus, which results in two distinct micropylar
chambers. The outer chamber is partially inclosed by the infolding tip of the
integument, from whose inner face numerous hairlike processes are developed
as outgrowths from the epidermal cells. Within this chamber the pollen grains
are received and germinate, a tangle of tubes passing down through the inner
chamber to the nucellus.

At the time of pollination (April-May in California) the pollen grain contains
the two disorganized prothallial cells and the generative and tube nuclei. Just
before tube-formation the generative cell divides to form the stalk and body cells,
both with distinct membranes, but soon becoming very unequal. Before the
tip of the nucellus is reached by the tube, the nucleus of the body cell divides
to form two unequal male nuclei. Fertilization takes place within 60 days
after pollination, and the entire nuclear contents of the tube are discharged
into the egg.

The functioning megaspore is surrounded by a distinct tapetal zone, and the
magaspore membrane becomes conspicuous. The development of the female

BINSON, WINIFRED J., A study of the digestive power of Sarracenia purpurea.
eS 8: 2181-194. 1908.

3: Law of A., The reer em and embryo of Pseudotsuga Douglasii.
Annals of Boleny 23: 165-186. pls. 12-14. 1

1909] CURRENT LITERATURE 485

gametophyte proceeds as usual, filling the sac with primary endosperm cells,
in which free nuclear division occurs before cross-walls form the permanent
tissue. The archegonia are usually four in number, and a distinct ventral canal
cell is cut off, the membrane persisting until fertilization occurs. The proembryo
is the usual one of Abietineae, walls appearing at the eight-nucleate stage, and
four tiers of cells being organized, the uppermost tier being open.—J. M

Embryo sac of Pandanus.—CAmPBELL3* has published the details of the
development of the embryo sac of this interesting form, the preliminary account
having been published last year.33 e stage showing fertilization was not
obtained, so that it is not certain that the fourteen-nucleate condition described
is the fertilization stage. The megaspore mother cell is overlaid by several
layers of parietal cells, which are thought to be derived from a single cell. The
division of the mother cell is followed by the direct production of the embryo
sac by the inner daughter cell. At the first division, the two nuclei assume the
polar positions, and subsequent divisions result in two micropylar nuclei and
twelve antipodal nuclei. If two megaspore nuclei are supposed to enter into the
structure of this sac, there is a single division of one of them, and a succession of
divisions from the other one. From any point of view, such a sac would be
unusual, and the author is inclined to regard it as ‘‘a new type with its nearest
analogue in. Peperomia,” a type which is probably more ancient than the prevail-
ing eight-nucleate sac. He dissents from the idea that the reduction division
necessarily determines a megaspore in angiosperms, believing that this event
may so shift in the life-history that a megaspore may be defined regardless of it.
After all, this is merely a matter of definition, and that is a matter of agreement.
Shall a megaspore be defined as the product of the two reduction divisions or
as the cell which produces the embryo sac? Which definition will have the
greater morphological stability >—J. M. C.

Diffusion of CO, in leaves.—That CO, does not diffuse extensively through
the mesophyll has been known for more than thirty years from the researches
of Mott, and experiments to show the localization of photosynthesis have become
common in every physiological laboratory. Under Mo t’s direction, Z1yLSTRA
has investigated the extent of diffusion in leaves of different structures.3+ He
finds that in all leaves the CO, formed in the leaf itself suffices to produce a line
or zone of starch at the edge of the darkened region. In net-veined leaves the
movement of CO, supplied from the air is prevented by the larger veins which

32 CAMPBELL, D. H., The embryo sac of Pandanus. Bull. Torr. Bot. Club
36: 205-220. pls. 16, 17. 1909.

33 CAMPBELL, D. H., The embryo sac of Pandanus. Preliminary note. Annals
of Botany 22:330. 1908.

34 ZIJLSTRA, K., Kohlensaduretransport in Blattern. Proefschrift ter verkrijging
aan der graad van Doctor in plant- en dierkunde aan der Rijks-U atten te Gro-
mungen... . . 8vo. pp. 128. pls. 2. figs. 2. Groningen: M. de Waal. 1909.

486 BOTANICAL GAZETTE [JUNE

stretch from surface to surface without intercellular spaces. Consequently the
width of the lines of starch produced at the edge of the darkened region is not
widened even though the unlighted area is supplied with CO, under abnormal
pressure. If the net be coarse the zone of marginal starch will be wider than if
it is fine. Diffusion of self-produced CO, to 2.5°™ at most is thus possible; for
in parallel-veined leaves of Triticum, Hordeum, and Zea, though the veins do
not prevent diffusion, the intercellular passages are so narrow as to limit it to 3°™.

n Tradescantia and Acorus the transverse anastomoses prevent more extensive
movement. In Ejichhornia, Pontederia, and Eucomis the leaves have spacious
intercellular passages, and so the movement is much more free. But even here
the diffusion scarcely surpasses 3°™, unless through a region of the leaf that is not
in condition to act on the CO,. In nature, therefore, movement of CO, may be
considered practically nil—C. R. B

Seedling of a graft-hybrid.—Certain branches of the graft hybrid, Cytisus
Adami, revert, producing flowers having the characters of the reputed parents
C. Laburnum and C. purpureus respectively. The C. Adami flowers are ordinarily
sterile, while those borne on reverted branches reproduce their respective parents.

In May, 1904, HILDEBRANDS observed that several flowers of a C. Adami
branch of a cultivated specimen in the Freiburg botanical garden had set seed,
and was able to obtain three fruits from them, which had chiefly the characters of
C. Laburnum, but in certain respects resembled C. purpureus. It is not known
whether these flowers were self-pollinated, but it is not unlikely that the pollen
came from C. Laburnum flowers, since the C. Adami flowers are usually sterile.
Two of the seeds germinated. Both were very similar in character to C.
Laburnum and in 1907 one of them produced hundreds of flowers, all having the
characters of C. Laburnum. No conclusions can be drawn regarding the heredi-
tary bearing of these facts, in the absence of a knowledge of the manner of pollina-
rn of the flowers and the nature of the next generation of offspring.—R. R.

ATES.

Chlorophyll.—The discussion as to the phosphorus content of chlorophyll
waxes warm. STOKLASA replies vigorously3° to Tswert’s criticisms? and takes
issue with WILLSTATTER’s results.3° The question is yet in the stage of polemic

35 HILDEBRAND, FRIEDRICH, Ueber Saimlinge von Cytisus Adami. Ber. Deutsch.
Bot. Gesells. 26a:590-595. 1908.

3° STOKLASA, J., BRALIK, V., UND Ernst, A., Zur Frage des Phosphorgehaltes
des oneeuia Ber. Deutsch. Bot. Gesells. 27: 10-20. 1909.

37 'T: , M., Ist der Phosphor an dem Anbau des Chlorophylline beteiligt ?
Ibid. a Sek suc 1908.

38 WILLSTATTER, R., Zur Kenntniss an Zusammensetzung des Chlorophylls.
ore bu ne der Chemie 350: 48-82.

., UND Benz, M. tae Lipiialliciertes Chlorophyll. Ibid. 358:
ances ee

1909] CURRENT LITERATURE 487

discussion and will doubtless require further and most careful work. STOKLASA
is particularly emphatic, declaring that ‘‘we have determined—not only I, but
my collaborators, at different times—by definite researches, which we are ready
to repeat in any forum, that preparations of chlorophyll all actually contain
phosphorus.” And again: ‘‘Our new investigations, carried out in my labora-
tory both on crude and pure chlorophyll, prove that the phosphorus is bound up
in the chlorophyll complex and does not occur in ionic form. We have recog-
nized with complete certainty glycerophosphoric acid and cholin. Consequently
the assertion of EuLER and of ScuutzeE, that the chlorolecithin hypothesis is
Say refuted by WILLSTATTER’s work, is at least premature.”—C. R. B.

rr
finds that ale root and a ae a axis soon escape from the seed coats,
leaving behind, in close connection with the reserve food, a foot or sucker. The
cotyledons are at first small, but later enlarge somewhat and do photosynthetic

work. An older specimen shows the ers rodlike foot in the center o

the endosperm. The foot contains vascular tiss

Hitt remarks that the foot develops to a se extent in Gnetum than in
either Tumboa or Ephedra. Comparison with the last genus is certainly astonish-
ing. The reviewer can speak for many of the ephedras of the western world.
They do not have a structure in the remotest degree resembling the foot of Gnetum
as figured by Hr1t, nor is there even a rudimentary trace of such a structure.—
W. J. G. LAND.

Sex of Sphaerocarpus.—Acting upon a suggestion by STRASBURGER, DOUIN*°
has carefully examined 81 groups of Sphaerocarpus terrestris, taken by chance
from material collected at Chavannes. He finds that about 75 per cent. clearly
show 2 male plants and 2 female arising from the spore tetrad, whose mem
cohere usually until germination. The others were mainly explicable by non-
germination of one or more spores of a single or double tetrad, or the accidental
dissociation of the members of a tetrad. Several cases clearly anomalous were
found: one group (from 2 tetrads) of 54 and 39; another of 3é and 19; and two
others, 14 and 3%. He corrects certain earlier misstatements regarding sporelings,
and now specifies differences between juveniles of S. terrestris and S. californicus,
which before he declared indistinguishable—C. R. B.

Necrosis of the grape.—-Vines killed by this disease*t have usually, in the
opinion of the owners, ‘just died,” yet the writer regards it as a serious disease
causing a large percentage of damage. In one young vineyard of 14 acres it is

39 Hitt, T. G., The germination of Gnetum Gnemon L. Jour. Roy. Hort. Soc.
S4:1, 2 1008.

4° Doutn, Cu., Nouvelles observation der Sphaerocarpus. Rev. Bryol. 36:37-41.
ROOD:

4: Reppick, D., Necrosis of the grape vine. Cornell Univ. Agric. Exp. Sta. Bull,
263. Feb. 1909

488 ' BOTANICAL GAZETTE [JUNE

estimated 4000 to 5000 vines succumbed. The symptoms are: A trimmed and
tied vine that has failed to put out shoots; a vine that has sent forth shoots, the
latter dying after a few weeks; shoots and leaves that exhibit dwarfing; blanched
and chlorotic leaves; leaves and fruit shriveling and dying in the summer; the
presence of fleshy or corky excrescences on the stem, of minute black pimples on
a dead spur, or of small reddish-brown spots on the green shoots. The disease
is attributed to Fusicoccum viticolum, which is described as a new species.—-F. L.
STEVENS.

Origin of plastids.—Without giving any adequate evidence, even in outline,
SCHILLER propounds the idea in.a preliminary paper? (which can have no other
purpose than to secure priority, and this ought to be denied in such cases even
if the guess proves correct) that the plastids of plant cells arise by the extrusion
and fragmentation of nucleoli, whose fragments subsequently grow and change
their structure. He “is inclined to the view” that the plant cell is therefore to be
looked upon as binucleate, in the sense that the chromatophores correspond to a
macro- or yolk-nucleus, a view which has fapely been expressed by Mororr for
animal cells—C. R. B

Leaf blight.—Stevens and Hatt describe*3 a disease of apple, pear, and
quince, whose prominent symptom suggests the name leaf blight. As it is due
to Hypochnus ochroleucus Noack, they propose the name hypochnose, in conform-
ity to a scheme for making names of diseases by combining euphonically the name
of the fungus with the termination -ose. The disease resembles fire-blight (bacil-
lose), but only the leaves are affected (no twigs), and they stand erect instead of
drooping. The disease prevails in the mountain section of North Carolina, West
Virginia, and Alabama, but is probably much more widespread.—C. R. B

Geoglossaceae of N. A.—The attention of those interested is directed to an
elaborate monograph4+ of this family of Discomycetes, allied to the better-known
Helvella and Morchella types, as represented on this continent. There are
eleven genera, and the original fifty-three species DuRAND reduces to forty-two.
We have no competence to review the work critically.—C. R. B.

42 SCHILLER, Jos., Ueber die Entstehung der Plastid dem Zellkern. Oesterr.
Bot. Zeits. 59:89-91. figs. 3. 1909.

43 STEVENS, F. L., AnD sg J. G., Hypochnose of pomaceous fruits. Ann.

ng oe: 7249-59. - 8. 19

DURAND, as: j., ae Geoglossaceae of North America. Ann. Mycol.

ees pls. ae 1908.

GENERAL INDEX

Classified entries will be found under Contributors and Reviews.

New n

mes
and names of new genera, species, and varieties are printed in bold-face ai

synonyms in italic.
A
Abobra, histology of fruit and seed 307

iner on 4
rown and Stapf on

163; ore O ; Schinz on 161
ce singe, "eduction division j in 198
Air, Saito on germs
eagle Stewart et ng on ‘diesen of 343
Algae of 237
A pecnacth ‘Dianthi 413
Alternariose, carnation 409
Alternation, Lang on lipetiene theory
of

America, plants of noe tage 472

Amitosis in Synchytr

Amphiachyris F remontia ioe

Anato: aang of, fru re eeds of soa te
tacea 263 S$ 301; Microcycas
ican ig anaes Stiles on

Androstachys 473
ynaceae, Wildeman on 163

\ragallus, b

— A sag og agra sis in 50

os a, Shaw apa of 481
asad neat 72

sogetonia 473

Occus, Scbertfe lon 473
saga, barium and loco 344; Born-
miiller on 473

Aspidium -acrstichoides, bane in 46
Ataenidia, Gagnepain on

> > >> > > > >
a
cE

aaah rium, ao — a on Ths Filix-foe-
bs

At sions tgs ar

Austalan plants, * iden and Betche

Ave net Blau on phototropism in 343
Azotobacter, Krzemieniewski on nitrogen
fecaton i 475

B
Blaauw, A. H., work of 343

Bacillariactee, Schroeder on 163
Bagnisia, Cheeseman on 163

Bahamas, Britton on flora 473
Baileya, multiradiata 431; perennis 431
Balls, L. W., work of 247
arber, Ae
baciee and oes pon ere on 344
Barnes, C. R. 2 244, 344, 414, 415,
416, 424, Sh, 7 “8s 486, 487, 488
Bateson, work of 4
Beauverd, G., work of 472

A
Blight, ata ad Hall on leaf 488
Bogs, peat deposits in local 445; toxins

3°9
Boissieu, H. de, work of rtd
Bo nese os = work of 162, 4
Bornmiiller, J., work of oo 163, 473
Bornacges Bisset lae Botanicae” 160
of 161

a7, 473

475
soon i "work of 170
- 445
Ree, Prain on 163
C

Calathea, Gagnepain on —
Calceolaria, Krinzlein on
alochortus, comosus 42 33 Scud 425
Calorimeter, respiration
— of R dada as. von Schrenk

Capparidacae, Gagnepain on 162; graft-
hybrids
Foe. bigs dioxd i in leaves, Zijlstra on diffu-

riose
Carnegiea, Britton - ‘pias on 472
Carya, Donde on 1

490

Castle, W.-E., work o

Lonely labiata, homocosis of 44
Centaurea, Gugler on 163

c an on 162

ee amia, extrafascicular cambium
Chabert. Alfred, work of 1

Chaenactis, attena ta ae carpoclinia

434;

Chamberlain 5, Charles pie 215

Chee T. F., wor 163

China plants of, Dunn on oe Komarov

Chip ec “T F., work of 162

perms is Stoklasa on 486; Tswett on
Willstatter on 486

Chomatia vat ga 255; filipedis 255

Christ, H., work of

Chromogen bee pnatetigh

Chrysler, M. A. 169, a se work of

172
Chylisma hirta 428
Circinella, phe on 420
Cirsium, Petrak on 162

i “2 on I

ee
ma
-
QO

a Chipp on or euanie on

Golds Crataegus in digefe
Compannia, een

: ts
ber, K. G. 263; Barnes, C. R. 242, 243,
44, 344, 414, 415, 416, 424, i, 479;
A+ 486, 487, 488; Blakeslee F.

8; Buchanan, R. E. s.
P. 445; Chamberlain, C. J. 172

15; Chrysler, M. A. 169, 171, 337;
land, W. F. 9; Coulter, J. M. 15

159, 170, 172, 244, Sys 477; 478, 481,

484, 485; Cowles, H. C. 73, Se i 159)

170, 252; Crocke r, W. 69, 8

240, 252, 339, 340) 341, 08s se 44,
482; Dachnowski, A. 389; 3 3
; De

153, 157, 158, 161,
162, 338, 339, 467, 472; Griggs, R. F.

136; Harris, J. A. 438; Hefferan, Mary

75,76, 251; Hill, E. J. 454; Holm, T.
167; Jeffrey, 424; cd W. ].
G. 487; nite R. G. 30; ovens a
241; Nelson, A. 425; Nieu nd, J. A

a37; Olsson-Seffer, a Hite ated By

INDEX TO VOLUME XLVII

Wi j.N. 248; Peirce, G. J. 725 Per
kins, J. 230; ae R. H. 247, 4733
Ramaley, 22 6ed.— Ti. -S:

Saxton, 1s 0 Schreiner, Os-
wald 355; Shull, G. He 337: 468;
Smith, 2535. Stevens, -F 15
172,343,400, 487; ree Alma
Go Arts omson, R. 3453
Westerdijk, Johanna 241; Varian:

chi, S$: 173
Convolvulus, House on 161
Copeland, E. eo ane of 164
nd

156 » 159; ae 172, 244,
478, 481, sie
, Henry C. 73, 74, 81, 159, 170, 252
Crataegus in Capac (Ee gre le on 163
Crescentia 259
Crocker, William 69, 82, 164, 170, 249,
252, 339, 349, 341, ose Me 344, 482
Crocus sativus, homo:
Crucianella, M sree dele on oe
ryptanthopsis, Ule on 4
Cucumber, histology of fruit and seed 283
Cunninghamella elegans, Lendner on 420;
Jeffrey on 171
Citcurbitaceae histology of fruit and
a03
Caran histology of fruit and seed 287
Cylindrosporium, Brooks on 162
Cyperaceae, Palla on 162
Cytology, Agave 198; asa basis for Men-
delism, Grégoire on 80; of Fucus 173;
of Oenothera, Geerts on 480

clea si es 380
Daucus, Carota, homoeosis in 51

Dicksonia Big wage deen ula, homoeosi

Disease: Dianthus ‘caryophyllus 409;
Sivas a nd Hodgkiss on plant 172

Dioon edule, spermatogenesis in 215

487
Dro: rmedia, homoeosis in 40
Drude see ‘ae

1909]

Drygalski’s “Deutsche Siidpolar-Expedi-
tion 24

camp’s ‘‘Du colibacille et du bacille
typhique”’ 76
nn, S. T., work of Sie

Durand, E. J., work of 4

Durham, F. M., work 3
ysodia, cancellata 435; Cooper 434»
at cupulata 435; fusca 436; pap

osa 434; porophylla bake porop hyl-
ibitles 435; speciosa 435; Thurberi 436
E
Earle, F. S., work of 473

East, E. M., work of 168
Echinocystis, ear ie of fruit and seed

Roplogy of sea shores 85
Edaphic factors prt ooo: 94
Elmer, °

paar sac of Pandanbas Camisbell on

495
celia, nudicaulis 433; nutans 433
Reecelieis 432; grandiflo * 433; nudi-
caulis 433; nutans 433; tuta 433
Engler, ‘‘Das Pflanzenreich” 157; and
D i r 733

Epi , Komaro

Equisetales, TE on Mesozoic 424
“iat “Recueil d’ceuvres de” 4

Erioc ro n, Gnaiverd on 472; Lecomte

Er rat oe work of os
Eryngium, Wolff on 472
Euphorbia, Reramilier on por —

phylla spt crenulata 437; m 4373
Nortonian gli rerige chew =
Evans, A Nic
mRacditcs arg Connecti Sat? I i
Se age ya owe n vegetation go

A. J., w

Ew 69
Excretion ne pe: Ernst on root

Extracts, soil, coon ah in 371

Eylesia, Moore

F
Fawcett, W., work of 473
Ferdinandsen, C., work of

Ferns, acminn nd o
MeNichol on evielies in 109% mutation
in the Pierson 32; Rosens ock on 472

Fitting, i: “work of 479

INDEX TO VOLUME XLVII

491

Fischer, E., work of 76

k o
ork of 343
Freycinetia, Merril on 164

f 1

Friuli Davide 163

Fruit of Cuc arhiearen comparative his-
ios a of 2

Fucus, mitosis

Fungi, Fe nian. and Winge on 472;
Peck on 163; re cultures of 241;

subterranean 75

Gagnepain, G., work of ts 163, 472
oe peduncula ata

anong’s “Laboratory course in plant

physiology’’ 242
ae R. R. 79, 83, 84, 154, 168, 169,
250, 251, 473, 476, 480, 486

ee Sages 4375 fatiebets 437
Geerts, I. M., work of 4
Gemmation, Binet in Synchytium 129
Gentiana crinita, homoeosis in 39
Geoglossaceae e of North Maes, Durand

on 4
Gens , Fischer on 78
Geotropism, Buder on 170; Haberlandt

on 482
ee of Gnet — so - nate
d light, Heinricher o nzel

rs 36, 341; of Rh Pe ea rigs

lich on
Germs in -~ wi yy on 251
Gill fungi, E 473
Gleditsia Brgietet omoeosis in 49
Gnetum, Hill o eae of 487
Gonolobus, aran 7; patalensis 256
Gooddingianae, plantae 425
Graft-hybrid 84; Hildebrand on —.

486; of Solanum, Winkler on 250
Grape, Reddick on necrosis
Grasses, Hackel on 472
Graves, A. H., fe)
paced “New manual of gin 153
, E. L., work of 472

ce J. M. 153, 157; se . 162,

Griggs,

Grossweilera, Moore 161

Growth an sai temperature, Ball on 247
of 339

#
7
ag

Griis
Guatemala, pec of 253
Guettarda, cobanensis 255; crispiflora 255

492

Gugler, W., work of 163
Guraniopsis, Urban on 162
Guzmania bracteosa 262

i ct tt Kern on 162

uinqueseta, homoeosis in 58
-, work of 482

erste ‘M of technical
” 416

Hes The ‘eager gs 414

Harris, J. Arthur 4

Hayata’s ‘Flora

: 164

Helianthella, ‘argophylla 433; Covillei
433; nudicau lis 433

Hema “

He ennings, Paul 239

Hepaticae, en on 163; Stephani on
161, 163

ucurbi-

‘Has HL, work o 252
Huthia, Urban on 162

INDEX TO VOLUME XLVII

[fUNE
Hymenatherum Thurberi
Hymenoclea fasciculata es
I

Impatiens, Hooker on 161
Inheritance of sex, Bateson and Punnett

’ petcia 431

jon and mutation, Leavitt on 170
Itatiaia, Ule on 47

— “Mikrographie des Holzes”
Jasmin officinale, homoeosis in 50
effrey, E. C. 424; work bs 171
ohansson, K., wor .
Justicia multicaulis a5
K
Kern, F. D., work of 162

Kranzlein, F., work of 162, ”
rzemieniewski, S -» work of 4

Lesquerella tenella 426
veillé, H., agp a ee 163

INDEX TO

Lindau, G. 241

Linum leptopoda 426

Loco, Crawford on ation and 344
Luffa, aig 3 of fruit and seed =
Lutz, Anne M., work of 475
Lycopodium Selago, homoeosis in 61
Lythraceae, Koehne on 473

i ap gautemalensis 253; portoricen-
sis

aides , J. H., work of 163

Maize, ‘Shull on 169

} owski, Edmond, work of 161
aaron amg sh on 163

Malvales, Fries 164
Mangin, ar este of
— macrophylla nee verapazensis
a rryat , work of 470
eee Stephan on 472
Matt ., work of 163
Pn Bars ion
McNichols, M., wor

61
rk of 169
Melon, musk, histology of fruit and seed
287; water, histology of fruit and seed

292

Melothria, histology of: fruit and seed
30.

‘Soria Grégoire on cytological basis
of 79

eaten polita 427; synandra 428

Merintho apa campanulatum 257;
neuranthe

257
Merrill, E. D., work of 164
hrys, megasporopbyl of 3453

Microcac
tetragona, pollen
Mic rage as eticekec: vince anatomy

Mirabilis glutinosa 42 426
Mitosis, in Fucus 173; Oes on solutions

of 1 172
Moisture of air, influence on vegetation go
‘urpurbac pipers 75

oore ork of 161
Morphology ng Fischer on 76
— Dauphi ee 22

Mortonia, scabrella tahentts 427; utah-

2
Mosses, Paris on 472
Mounting of algae 237
Mucilage ducts in Pipereae 167
Mucor, papers on 418
Mucuna, Léve ine on 162
aerate ility and variability in Oenothera

Mut ee a vegetative 30
Mutation and genes Leavitt on 170
Rivcinecoeyutis, Fischer on 77

VOLUME XLVII

493

*
Myrmecophily, Nieuwenhuis-Uexkiill on
81

N
Nadson ig A., work of 472
Na nese , work of 162
Necrosis of the e grape, Reddick on 48

7
Nectares Nieuwenhuis- Uexkiill on extra-
ral 8x

aeaaes Aven 4
Neocalamites, ‘alle 0
Neotuerckheimia 258; ‘hiegalockytin 258;
gonocla
Nephrodium Satie, ee paceanely in 42
Nicholas, set
Nieuwenhuis- Vexkil work of 81

gen fixation mn by pene) Krze-
mieniew: eso
Norwegian s, Hagem on 418
Neatoecuneite: Bewuveet on 472
O
Oaks, Merrill on 164
Oes , Ado lf, work of 172
O ra, s 437; Geerts on
cytology © 480; DeVries on hybri
Of 3 Aga: mutability and variability

4
Oliver, F. W., work of 244
O

Orchidaceae, Fawcett 4 Rendle on 473;
ranzlein on 472

Orchid, flowers and formative stimuli,

ie n 479; Finet’on 162; Rolfe

Osiris in root ‘> Hill on 170 :
Osm sna cin mea, homoeosis in

Osterhout W. J. V. 148; — of 344
us on virescence in 252
saree by roots 355

Palms, Beccari on 472

ringers oe Canigioell on embryo sac 0
4

Paris, E. G., work of 4

Parthenoge enesis “a n Pins pipastir pe

Peat deposits in 445

, wo
es ‘olsbenenniogen der bra-
flanze

achat
Pedi peratim Bonati on Poles

494 INDEX TO VOLUME XLVII [yUNE

fctnre George J. 72
Pelargonium, homoeosis in 51
Periodicity in Spirogyra 9
-erkins,

ermeabily, Sage on 342
etrak, F., work o

mu
Pivant ane clegans, Oliver on 245
Piccard, Haberlandt on rotation experi-

ments 482
Pierson fern, mutation in 32
Pinus pinaster, So in 406
Pipereae, van Tieghem on 167

472
Platanthera viridis, abn eit os of 36
Plastids, Schiller on origin of 4.
Podophyllum, caer of 43
Pollen of Microcachrys tetragona 26
—— in ‘pees 454
Polygonaceae, Nakai on 162
Polystichum, acrostic hoides, homoeosis in
i angulare, homoeosisin 43; Braunii,
oeosis in pinna 4
Pied: Raymond H. 247, 483
Potato, bre reeding, East on 251; Stewart
ying 343

roteases, Vines on es 483

>seudoabu tilon, Fries on 164

seudobalsamia, Fischer on 78
ia, inz on

hand Aceil ‘food bak teed
7 i)

: suga, Lawson on morphology
of 484

Pueraria, Léveillé

Pumpkin, amines of a and seed 273
Punnett, R. C., work of 469

Quincula, lepidota 430; lobata 430
R

ph Sige pas on 164
aley, Francis

Ranales, Worsdell on 171

Ranunculus bulbosus 41

Reaumuria, ieee on 163

Reddick, D. 487

Reduction ¢ mn division i a Agave 198

ard S. 355
Relcke’ 3 * Pinementacinig 9 in Chile”

73
Reinbold’s “ Meeresalgen” 244

is in 59
Sader soya a 431; & Scum 430;
culata ‘

Rendle, A. B., work of 473
“Report of American Breeders? Associa-

Re meet ction, Freund on stimuli 342
Beep reon, calorimeter 72; chromogens

3
Ruviews Bernard’s “‘Protococcacées et
Desmidiacées 243; Borntraeger’s
*Tabulae Botanicae”’ 160; DeVries’s
“Naar Californie II” 159; Drude, see
ae ee Dryga oe ski’ s Bot ae a ice

polar-Expedition”’ 244; D

tolibacille et ee bacille pace 76
a spel s “Das Pfla seeing 157; Eng-
and ig 3“ Vegetati n der Erde”

Pog Eng and Prant iF “ Natiir-
viran Planzentaniin” 161; ‘Errera’s
vres’’ 417;
Nicholse’s Besouhyis es of Connecticut”
159; Fernald, see Robinson; Foslie’s
“ Lithothamnien” ong’s
“Laboratory course in plant physi-
ology” 242; Gray’s nu
of botany” 153; Hanausek’s ‘Micro-
scopy of technical products” 416;
Hard’s “Th shroom”’ ; a
ata’s “Flora montana Formosae’’ 158;
Hochreutiner’s “Ser rtum epee iT
iense”’ 338; Molisch’s urpurbac-

terien” 75; Peckolt's “Wolksbenen-
nungen eS. ‘ghar eee Pflanzen”

aa4;. Pre Engler; Reiche’s
gy Ehweaan ponte in Cbhile?. 735
Rei ort is “Die Meeresalgen”’ 244;
“Re of American Breeders’ Asso-
ciation” 337; Robinson and F ernald’s

“ Gray’ nu tany” 153;
Rydberg’s “ Rosaceae”’ 339; Schiffner’s
“Erge e der bota n Expedi-
tion nach Siidbrasilien” 467; ite
der’s “Handbuch der Lau .
415; Simroth’s “ Pendulationstheorie”’
v4; ’s “Guide erby’s

Smit to So
models of British Pricg 161; Smith’s
" ieee — of Bri ala oe oa mycetes”

cinea

der botanisch
Sudbrasilien” por inton’s Hanau-
s ** Microscopy of technical pro-
duc ts”
aa emma Sperlich on germination
of

Rhinarrad nigricans, Sumstine on 4

23
Rhododendron, von Schrenk on py ae
of 252

1909]

Rhodosphaerium diffluens, Nadson on
n, T. B., work of
sea F cal
ual of b Vere
Ro ocky 8 Mountain plants Bs

342
% Gray’s new

Root, pore ror | I Ses, ees excretions,
Stoklasa and Ernst on ; hairs, Hill
on — properties of aes oxidizin ng
a, bowet

eee N,, work of 472

Rosenstock E., work Ne a thy

Ruhland, W., work o

Ruppia, Graves on be of 478

ss: ewes colorata 260; Cumingii 261;
Dea 260; fusca 261; Kellermanii

Rydberg’s ‘‘ Rosaceae” 339

5
cn ieee Pane 61
Salts um, Flurl on 252
Send. ‘one on a IOI, 102
°

argent, C.

Setyria meian 25

Saxegothaea, pias he as OF 34k
Stiles on anatomy of 2

Saxton, W. T. 4

Scallop, histology of fruit and seed 281

Schaffner, John H. 19

Scherffel, A., work of 473

Schiffner's “Ergebnisse der botanischen
Expedition ond Siidbrasilien” 467

Schiller, sh work of 4 i

Schinz, ‘Hans, ae of 1

Schneider's “Handbuch ee Lanbhok

kunde’

eae peas ld

eae dts osis of 44
Scrophulariaceae, Bon n 4
Se , of Ara

oe Hill and Fraine on 248, 481;
of graft-hybrid, Hildebrand on

ds, oy Cucurbitaceae, comparative
of 263; Heinricher on germi-

zel on germination and
= mone of 69; Oliver
on iti ve of Rhinanthaceae,
Sperlich on germination 340
esa ea renee je homoeosis in 40

ne wor
ee og ea tt of, Bateson and Punnett
on 468; Castle on 471; Correns on
468; Doncaster ea Raynor on 468;

INDEX TO.VOLUME XLVII

495

Durham on 470; Marryat on 470; of
Bane are gand Douin on 487
Shaw, F. J. F., work greed
Shreve, Forrest, Sek of 2
Shull, George H. 337, 468; lh of 82,
9
Sicyos, oe of fruit and seed 2096
are, aaa sain 74
eo

pores of 34
Smith’s “Guide to Sowerby’s ni ls of
ritish Fungi” 161; — sis of
British Basidiomycetes” 16
Smith, John Don 253
Soils, effect of bog | spent upon 389
racts, oxidation in 371; a vegetation,
relation of on sandy shores 85
Solanaceae, Winkler on graft hybrids 84,

: cltions Osterhout on balanced 148, 344
Somali land plants, Mattei on 163

So ars American plants Urban on 162
Spanish plants, rat ay on 164

Sperlich, A., work of 3

S ermatogen esis in Diosn edule 215

. arpus, Douin on sex 0

oO
onl

Belew: si tea ra frit and seed, crook-
ed 2 chu winter 282
Stapf, Oz k of 6:
Statolith theoky, Buder on 170
Stem
Ste epha ni, F., work of 161, 163, 472
Steril in hybrids, Tischler on 246
. L. 172, 343, 409, 487
oy C., be: of 172, 343
251
g pe formative 479; to
reproduction, or d on 342

Sofa 249, 486
on transpiration and

rk of 424

s mu
Heat Moore on 161

es
Tammes, Tine, work of 3
Tansley’ 5 “EN volution of Picea vascu-
lar system” 336

496

Taxonomy 161, 472
Temperature, and

rt B. 26, 345; J. Arthur,
it ity” 15
* Thonner’s “ Die Bliitenpflanzen Afrikas”’
I

Tischler, G., work of 246
y, influence on

Topo; ‘ph vegetation 111

Toxins, bo effect on oxidation 384

Tracheids, of conifers, sler on 172;
Cunn ja 171

spe tao influence on vegetation

ce
water storage, Shreve on 252
Pichomanshice histology of fruit and seed

og
Tswett, M.; work of 486

ms, Kirsch on 169;
n 169; in tracheids of
conifers 172

U
* pea ty 473
Urban, I., of 162
Uropedium 1 Linden, homoeosis in 40

Van Tie eghem, P., work of 167
Variability i if Oenothera 476

Vascular system of Ranales I7I
Vegetiton a soil on prea sea shores 85
Violaceae, Boissieu on 163

Ule, E.

INDEX TO VOLUME. XLVIIi

[JUNE 1909

Vines, Ss. H., work’ o

1; ‘Naar Californie IL”
159; work of 473

Wes terdi ilk T 24

W = van plants, ‘Britton on 162;
pla ee on

Wettstein’ s “Egebnis oe botanischen
editi 7

Wo k v4
Worsdell, W. C., work of I7I

Yamanouchi, Shigéo 173
Z

Zingiberaceae, Gagnepain on 163, 472
abe oe hus Moelleri, Wisniewski on

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Medical Department of the University of New York, wrote: “For the past four years I
have used BUFFALO LITHIA WATER in the treatment of Chronic Bright's
Disease of the Kidneys, occuring in Gouty and Rheumatic subjects, with marked
benefit.”’

: DR. G. A. FOOTE, Warrenton, N. C., Ex-President State Medical Society, ‘backs
: Member of the State Board of Medical Examiners: and also of the State Board of Health
“In Bright’s Disease of the Kidneys I have in many cases noted the disappearance
of Albumin and Casts under the action of BUFFALO LITHIA WATER, which
I regard as the most citichdlens of known remidies in this distressing malady.”

DR. JOS. HOLT, of Wew Orleans, Ex- Party’ the State Board of Health of Louts-
tana, says: “I have prescribed BUFFALO LITHIA WATER in affections of the
Kidneys and Urinary Passages, particularly in de subjects in Albuminuria, and
in irritable condition of Bladder and Urethra in females. The results satisfy me of
its extraordinary value in a large class of cases usually most difficult to treat.’

GRAEME M. HAMMOND, M.D., Professor of Diseases of the Mind and Nervous
seek in the New York Post-Graduate Medical School and Hospital: “Yn all cases of

right’s Disease I have found BUFFALO LITHIA WATER of the greatest
service in increasing the quantity of Urine and in eliminating the Albumen.”

MEDICAL TESTIMONY ON REQUEST
FOR SALE BY THE GENERAL DRUG AND MINERAL WATER TRADE

uffalo Lithia Springs Water Co.

BUFFALO LITHIA SPRINGS, VIRGINIA.

HIGHEST IN HONORS

BAKER'S Cocos

HIGHEST sponge sprinkled
AWARDS Wl occasionally wit
IN Platts Chlorides.
Wash the spon gs
EUROPE. ice a week | 7a
AND

Bacinerst, AMERICA
A perfect food, preserves

heaith, prolongs life CAI Or )/ eh es
WALTER BAKER & C0, Lid | || Ze Odoriess

Disintfecta aim tf.
Established 1780 DORCHESTER, MASS.

Druggists Everywhe

treet, New York, for free book.

d by
Write to the pean: r, Henry B. Platt, 42 Cliff

eee Baer eB Sd
THE FOUNDATION OF“
‘GOOD HOUSEKEEPINGS.

PIANOS S222
Prime ev
cumstances rag own a V
exchange and ae he
and explanations.

im your home free of expense, wea. for r Catalopue D
VOSR & SONS PI

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