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Rin oh A ie BS
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ara M
THE
JOURNAL
OF THE
COLLEGE OF SCIENCE,
IMPERIAL UNIVERSITY OF TOKYO,
JAPAN.
VOL. XIII.
a U
mR tf BK *® A GT
PUBLISHED BY THE UNIVERSITY.
TOKYO, JAPAN.
1900—1901.
MEIJI XX XIII—XX XIV.
31690
Dec, 191%
CONTENTS.
Pt. I. Published June 2nd, 1900.
Notes on the Geology of the Dependent Isles of Taiwan.
By B. Korô, Ph. D. Rigakuhakusht, Professor of Geology,
Science College, Imperial nb pds (With Plates
ER): ne uns
Change of Volume and of Length in Ton! ‘Steel, aad Nickel
Ovoids by Magnetization. By H. Nagaoka, Rigakuhakusht,
Professor of Applied Mathematics, and K. Honpa, Rigakushi,
Post-graduate in Physics, (With Plates VI & VII)...
Combined Effect of Longitudinal and Circular Magnetizations
on the Dimensions of Iron, Steel and Nickel Tubes.
By K. Honpa, Rigakushi, Post-graduate in Physics. er
Plates VIII & IX) a, Se
Studien uber die Anpa ssangsfahigkeit siniger ‘Infusorien an
concentrirte Losungen. Von A. Yasupa, Rigakushi, Pro-
fessor der Naturgeschichte an der Zweiten Hochschule zu
Sendai. (Zierzu Tafel X-XII)...
Ueber die Wachsthumsbeschleunigung einiger Algen And
Pilze durch chemische Reize. Von N. Ono, Higakushi.
(Hierzu Tafel X-XLID) ... . sae cee ce ne oe
Pt. II. Published July 25th, 1900.
Ammcnium Amidosulphite. By E. Divers and M Ogawa.
Imperial University, Tokyo ..
Products of heating Ammonium Sulphites, Thiosulphate,
and Trithionate. By E. Divers and M. Ocawa, jr
University, Tokyo ... ...
Pottassium Nitrito- hydroxvinidosulphates and the "Non-
existence of Dihydroxylamine Derivatives. By E
Divers, M.D., D. Sc, F.R.S, Emeritus Prof., and T. Haca,
D. Se., F. C. 8., Professor, Tokyo Imperial University ...
57
77
101
141
187
201
211
Identification and Constitution of Fremy’s Sulphazotised
Salts of Pottasium, his Sulphazate, Sulphazite, etc. By
E. Divers, M.D., D. Sc., F. R. S., Emeritus Prof., and T. Haca,
D. Sc., F. C. $S., Professor, Tokyo Imperial University
On a Specimen of a Gigantic Hydroid, Branchiocerianthus
imperator Allınan), found in the Sagami Sea. By M.
Miyasima, Rigakushi, Science College, Imperial ne
Tokyo. (With Plates XIV & XV).. ;
Mutual Relations between Torsion and Magnetization i in m
and Nickel Wires, By H. Nagaoka, Rigakuhakushi, Pro-
fessor of Applied mathematics, and K. Honpa, Rigakushi,
Post-graduate in Physics. (With Plates XVI) ... … ose
The Interaction between Sulphites and Nitrities. By E.
Divers, M. D., D. Sc., F. R. S., Emeritus Prof., and T. Haaa,
D. Sc. F. C. 8., Professor, Tokyo Imperial University
Pt. III. Published Dec. 28th, 1900.
Contributions to the Morphology of Cyclostomata. IL—The
Development of Pronephros and Segmental Duct in Petromyzon.
By 8. Harta, Professor in the College of Peers, Tokyo. (With
Plates XVII-XXI) … … ie
Beitrage zur Wachstumsgeschichte der Bambungewachse,
Von K. SaiBaTA, Rigakushi. (Mit Tafeln XXII-XXIV) .
Decomposition of Hydroxyamidosulphates by Copper
Sulphate. By E. Divers, M. D., D. Sc, F. R.S., Emeritus
Prof., and T. Haaa, D. Sc, F. C. S. RER RE Imperial
Univerity CRUE a O80 Se? eee
Pt. IV. Published Oct. Sth, 1901.
Observations on the Development, Structure and Metamor-
phosis of Actinotrocha. Iwası un N (With
Plates XXV-XXX) u cies RUE”
PRINTED AT THE “TOKYO TSUKIJI TYPE FOUNDRY.”
225
235.
263
281
311
427
497
508
Publishing Committee.
aa ep ——
Prof. K. Mitsukuri, Ph. D., Rigakuhakushi, Director of the College
(ex officio).
Prof. B. Kotö, Ph. D., Rigakuhakushi.
Prof. T. Haga, Rigakuhakushi.
Prof. S, Watasé, Ph. D., Rigakuhakushi.
All communications relating to this Journal should be addressed to the
| Director of the College of Science.
RTrABRBPEE
ze FF
Rr Re FR Be R
THE
JOURNAL
COLLEGE OF SCIENCE,
IMPERIAL UNIVERSITY OF TOKYO,
JAPAN.
VOL. XIII, PART I.
KR HH kK %Æ HE fr
PUBLISHED BY THE UNIVERSITY.
TOKYO, JAPAN.
1900.
MEIJI XXXIII.
Publishing Committee.
— nm
Prof. K. Yamagawa, Ph. B., Rigakuhakushi, Director of the College
(ex officio).
Prof. J. Sakurai, Rigakuhakushi.
Prof. B. Kotô, Ph. D., Rigakuhakushi.
Prof, |. ljima, Ph. D., Rigakuhakushi.
All communications relating to this Journal should be addressed to the
Director of the College of Science.
Notes on the Geology of the Dependent
| Isles of Taiwan.
By
B. Kotö, Ph. D. Rigakuhakushi,
Professor of Geology, Science College, Imperial University, Tokyo.
With Plates I-V.
THE HOKO GROUP (PESCADORES).
I. Introductory.
Between the, geologically neglected, south-east coast of China
and Taiwan, the expanse of sea is studded with a great number
of islands, collectively called the Höko or Pescadores Group.
It consists of islands, islets and rocks, great or small, altogether
numbering 57, besides countless hidden rocks under the water.
The waterway on the continental side of the Pescadores is the
shallow Fokien Strait, only a hundred miles wide, and on the
Taiwan side, is the still narrower Höko Channel,—the only pas-
sages which allow free communication to the waters of the de-
2 KOTO : NOTES ON THE GEOLOGY
pressions of the North and South China Seas. The region
is alternately subjected to strong ebbs and floods through the
influence of the branch currents of
the swift Æuro-shiwo from the
north and south, creating foamy
and turbulent waves, in conjunctin
with the steadily blowing heavy
north-easters,—the dread of coast-
ing navigators for ship-wrecks and
other deplorable accidents.
I have not yet had oppor-
tunity to learn by my own inspec-
tion the geology of the Pescadores
Group; but through the kind-
Fia. 1.—Index map of Taiwan to show
ness of Messrs. Y. Saitö and the position of the islands described,
T. Tada, I have obtained about forty specimens of rocks, which
no doubt fairly represent the types that build up the crust of the
islands. In anticipation of a fuller report by Prof. Yokoyama,
who has made the islands the subject of his special study, I
may give here brief notes on the descriptions of rocks and the
inference drawn as to the probable geologic structure of this
interesting volcanic group.
The islands are, broadly speaking, distributed within an
elliptical space. On the north of the Tropic of Cancer lie main-
ly the larger islands which are arranged after the manner of
Santorin. They resemble the latter not merely in general out-
lines, but they owe their very existence to the same cause; both
are of volcanic origin. ‘These Santorin-like islands are Gio-6,
Höko, Hakusha, and Chü-don, the latter three fuse together,
especially during low tide, into one mass with the intervening
OF THE DEPENDENT ISLES OF TAIWAN. 3
coral-reefs which stretch from one island to the other, making the
shape very much like Thera. The single island of CGio-6, then,
corresponds in shape and position to that of Therasia. Here, how-
ever, we look in vain for the active centre of Kaimenis of Santorin.
Taking into account the general distribution of the above-men-
tioned islands, and also the bathometrical condition, which the
chart, Plate IV plainly shows, it is likely that they form an
independent centre of effusion, in contrast to the Southern group
(the Rover group), from which this Northern is separated by
the Rover Channel, though both sit upon the eastern end of the
so-called Formosa Bank, which stretches out hither from the
coast of Fokien. The same type of topography seems to prevail
throughout the whole group. It is simple, monotonous, flat-
topped and low; the highest prominence scarcely exceeds 56 m.
(located at the south-west point of Gio-6), and the land can
only be recognised from the sea within few miles. The islands con-
sequently are wanting in wind-protected harbours, being constantly
exposed to the north-east stiff gales during full three-quarters
of a year. The land surface is bare, desolate and barren, being
entirely destitute of green covering, due, it is said, mainly to
the savage violence of the wind, against which even hardy
shrubs can not maintain their footing.
The rain-fall, which the south winds occasionally brings
thither during the summer season, is soaked up as soon
as it falls on the craggy ground; and there are scarcely any
rivulets that properly deserve the name. The erosive actions
of running water thus become totally suspended, and valleys
and dales are scarcely to be seen in the interior, but only the
butte-like table-land capped with the Basalt-sheet. The deflation
alone is instrumental in modelling the topography, and here we
4 KOTO: NOTES ON THE GEOLOGY
have a quasi-desert, and not an oasis, amidst the green island-
world of South-eastern Asia.
Forty of Mr. Tada’s specimens of rocks, on which I base
my petrographical descriptions in the present paper, were collect-
ed from the following islands :—
1) Höko island, the largest of the whole group.
2) Haku-sha-té,” lying north to the foregoing.
3) Impai-sho.
4) Ché-sho, the eastern neighbour of Hakusha-tö.
5) Kippai (Bird Island of English Admirality chart), the
northernmost of the whole group.
6) Gi-ö-tö (Fisher Island), west of Höko-tö.
7) Hattö-sho, lying farther to the south of the main group.
In addition to these, I have received lately a few specimens
collected by Mr. Y. Saitö.
1) The words ‘té’ and ‘sho’ recurs frequently in the geographical name of Taiwan, the
former signifying an island, the latter an islet or rock.
OF THE DEPENDENT ISLES OF TAIWAN. 5
II. Stratigraphical Characteristics.
HOKO ISLAND.
Höko or Tai-san-sho” is the largest among the forty-seven
islands of the Höko group, having an area of 62.7 square kilo-
métres. Its general outline is k-shaped, curving in at three
points in the coves, Fükibi,” Giû-bo-ken,” and Kétei.” The re-
lief is simple, low and flat-topped, the maximal elevation being
Mount Tai-bu,” located nearly at the centre, with a height of
only 48 m. The coast is cliffy, interrupted often by sandy flats
fringed with coral reefs.
Mr. Y. Saitô has geologically reconnoitered the principal
islands of the group during last winter, and has kindly placed at
my disposal the written account of his observations, which I am
here following in its main points.
The island is essentially composed of the Tertiary Basalts,
of which three different flows, poured out after long intervals,
in sell marked by the intervening tufaccous sedimentaries ofa _
4 KOTO: NOTES ON THE GEOLOGY
have a quasi-desert, and not an oasis, amidst the green island-
world of South-eastern Asia.
Forty of Mr. Tada’s specimens of rocks, on which I base
my petrographical descriptions in the present paper, were collect-
ed from the following islands :—
1) Höko island, the largest of the whole group.
2) Haku-sha-tö,” lying north to the foregoing.
3) Impai-sho.
4) Chö-sho, the eastern neighbour of Hakusha-tö.
5) Kippai (Bird Island of English Admirality chart),
northernmost of the whole group.
6) Gi-6-t6 (Fisher Island), west of Höko-tö.
7) Hattô-sho, lying farther to the south of the main group.
In addition to these, I have received lately a few specimens
collected by Mr. Y. Saitö.
1) The words ‘6’ and ‘sho’ recurs frequently in the geographical name of Taiwan, the
former signifying an island, the latter an islet or rock.
CORRIGENDA.
Page 4, 12th line,
„ 6, the last line,
» 18, 18th line,
w. 21, the last line,
» 28, 9th line,
=> “20; 13th line,
» 42, 2nd line,
» 45, 14th line,
» 653, 24th line,
57, 20th line,
for Gi-ô-tô
for Cholnecky
for there lics
read Gio-6-t6.
read Cholnoky.
read the relics.
for basite read bastite.
for crystals read crystal.
for Büking read Bücking
delete the word! macroscopically.’
for ‘leached’ read ‘ leached or percolated.’
I pyrites read pyrite.
for granual read granular.
read quadrant.
: Explanation Pts. I&II, for octant
OF THE DEPENDENT ISLES OF TAIWAN. 5
II. Stratigraphical Characteristics.
HOKO ISLAND.
Höko or Tai-san-sho” is the largest among the forty-seven
islands of the Höko group, having an area of 62.7 square kilo-
métres. Its general outline is k-shaped, curving in at three
points in the coves, Fükibi,” Giû-bo-ken,” and Kötei.” The re-
lief is simple, low and flat-topped, the maximal elevation being
Mount Tai-bu,” located nearly at the centre, with a height of
only 48 m. The coast is cliffy, interrupted often by sandy flats
fringed with coral reefs.
Mr. Y. Saitö has geologically reconnoitered the principal
islands of the group during last winter, and has kindly placed at
my disposal the written account of his observations, which I am
here following in its main points.
The island is essentially composed of the Tertiary Basalts,
of which three different flows, poured out after long intervals,
are well marked by the intervening tufaceous sedimentaries of a
considerable thickness. The dopmost flow caps the surface of
butte-like elevations, or makes the flows of extensive ‘ mesas,’ the
surface being covered with its eluvial products—a fine, ferrugi-
nous loam which gradually passes downwards into a blocky
loam and then the massive lava. The flow is rather thin, and
characteristically columnar. It is frequently wanting in some
parts of the island.
In the irregularly formed strip of land—the Fükibi-Jiri®
1) Tai-san-sho (Kl), signifying ‘great mountain islet,’ is by no means literally true,
thongh undoubtedly it is the largest of the whole Pescadores.
2) MER 3) FR 4) ER 5) A 6) eR.
6 KOTO : NOTES ON THE GEOLOGY
tongue, which projects out from Sci-shi-an” towards the citadel
of Bako, thus enclosing within it a safe harbour,—we see the second
sheet of flow, beautifully exposed along the steep declivity all round
the shore under the uppermost lava-flow, from which it is
separated by a thin bed of tuffite This is a most extensive and
strong sheet, aggregating about 10 m. In its upper portion, the
lava is porous, whitish, and much decomposed, while the lower
portion is fresh and compact. It is the one which we usually
see along the sea-shore on whose trappean floor the rollers
break and recoil in tumultuous waves.
The third is the lowest, consequently the oldest flow visible
in the Pescadores, and frequently forms the floor of the coast, when
the second sheet, already referred to, makes its appearance higher
up the precipice. It is likewise doleritic and porous as in the
above flow, and this Basalt is well seen at the environs of Jiri,
already referred to, where it is underlaid by a meagre lignite-
bearing bed. It rarely happens to come to the surface not because
of its absence but that it is hidden under the level of sea.
Tertiary strata, often accompanied by lignite seams, occur
inserted between the first and second flows, and also below the
third sheet. An undeterminable cast of gasteropod together with
an Arca were secured by Saitö from the corresponding bed at
Run (Lun) point in the Island of Gio-ö. The sure proofs of their
being of the Tertiary age are not at hand; but from the analogy
of the occurrences of Basalts in the neighbouring regions, I
conjecture the sedimentaries, here referred to, to be of later Terti-
ary. According to Cholnecky”, two volcanic lines are said to be
1) FIR.
2) ‘ Vorliiufiger Berichte über meine Forschungsreise in China.’ Pelermanne Mitth. 45,
1899, S. 8.
OF THE DEPENDENT ISLES OF TAIWAN. 7
distinguished in Eastern Asia; the one has served for the well-
ing out ofan enormous quantity of Basalt in later Tertiary age, the
other has given rise to the chains of modern (Andesitic) volcanoes.
In the north of the Chang-pei-shan, in Korea, he announced
recently the discovery of an extensive Basaltic mesa more than
60,000 square km., which extends from Mukden through Kirin to
Ninguta, forming the water-shed of the Sungari River and the
Tumen-kiang. I have been informed, verbally by Mr. Nishiwada,
of the occurrence, outside of Manchuria, of a trappean plateau,
of small extent, along the eastern water-shed of the Korean
Peninsula, and the island of Quelpart; and Vénukoff” cites a
number of localities where Basalts make their appearance on the
plateau of Mongolia. Furthermore, the Basalts occur sporadically
in Liau-tung, and Shang-tung as far down as Nanking, ap-
proximately in a straight line, and v. Richthofen” brings the
line in connection with the tectonic movement which has created
the ‘ great plain ’ of China, and he assigns the age of this crustal
movement to the Tertiary period. The Basalts of the Pescadores”
seem to me to be included in this petrographical province of
Eastern Asia.
Since the beginning of the Diluvial epoch, a subaerial condition
has prevailed over Höko, as well as in all the islands of the whole
group, and erosion and disintegration have been at work, thereby
carrying off the greater part of the uppermost flow, and gradually
diminishing the area of the islands, and finally reducing them to
ruins, as we see at present. Consequently, no record is left of
the deposit representing this period, unless we take for it the
1) ‘Les Roches basaltiques de la Mongolie Bulletin de la Scciélé belge de Géologie, ete.,
tome II, p. 441. . |
2) ‘Shantung und seine Eingangspfort Kiautschou, 1898, 8. 66.
8 KOT) : NOTES ON THE GEOLOGY
thin superficial covering of ferruginous loam which is in part
at least the product of decay of Recent epoch, though a certain
portion may have been defladed away and lost during dust-storms.
Along the shore free from escarpment, white sandy beaches
stretch from one point to another. They are the Alluvial deposits,
into whose composition enters a special element which we are not
accustomed to see in our own coast. Nearly all round the island,
coral reefs grow upon the Basaltic shelf, and the detritus derived
from them is driven up to form low sand dunes, leaving behind
them, if the coast-line is deeply indented, as it is in many
places, muddy shallows filled with the residual clay of decomposed
Basalt.
Such is the general outline of the geology of the Island,
and of the rest of the group as well.
Looking more into the details, we find that at Baké”
Point, on which is situated the town of the same name, the second
flow extends in a great sheet, covering all but a few points of eleva-
tion which are capped with there lics of the young columnar lava,
being separated from it by a blue rock. The last is a fuller’s
earth, which is a bluish-grey, dull, compact mass of greasy
lustre, splitting, when dry, into angular clods with sub-conchoidal
fracture. It adheres to the tongue, and falls readily to a muddy
state on placing in water, and is not plastic. Under the micro-
scope, the whole mass consists of brownish, double-refracting
particles, and seems to have been derived from the decomposition
of a Basaltic glass. It crops out for a short distance, and on shore
a poor bed of Tertiary lignite occurs associated with it.
1) 484.
OF THE DEPENDENT ISLES OF TAIWAN. 9
The same state of things prevails throughout the tract
southward as far as Sei-shi-an”, the surface being covered with
thick ferruginous loam mixed up with Basaltic fragments, and
the upper and middle flows coming in direct contact, distinguish-
Fio, 2,—Isolated erosion hill Sha-bö-san, near Jiri, showing two upper flows with
interbedded sedimentaries.*®
able only in the difference of structures. At the last-mentioned
locality, a ‘ haul-over’ of base-levelled middle flow, masked with
coral sand, separates the tongue of land Jiri”, on which stands
a Basaltic, hat-shaped Sha-bö-san”, 47 m. high (Fig. 2).
A good section may be seen along the shore, west of Jiri,
as is shown in Fig. 3. The columnar, upper (No. 1), and doleritic,
porous middle flows (No. 2), aggregating about 6 m., cap the
cliff, 20 feet high. That the two flows are separated by long
time intervals can be clearly shown elsewhere (Fig. 2) by a bed
inserted between them. I may cite the case of a lignite bed at
Bakö, occurring in company with fuller’s earth. Another instance
may be given of it just east of Jiri, where an ash bed makes
its appearance. This ash bed is a fine, greyish-white, pulverent
Darm 2) 9) ROL.
* All the figures in the following wood-cuts, not otherwise mentioned, are originally
sketched by Y. Saitö.
10 | KOTO : NOTES ON THE GEOLOGY
earth, wholly consisting of the microscopic particles of plagio-
clase, a few fragments of pleochroic hypersthene, and little
magnetite, but no glass splinters are seen. It reminds me of
the felspar sand that cover the
RAT SEEN flat and form the ground of
ER Pampanga, north of Manila”.
After this short digression, I re-
No, 2, Basalt.
turn to the former subject. Now,
. D * a=
L rs “a ne A | =
a oe
ER a yellowish-brown, loose sandy
al Sa pines, Felspar sand with
‘| limonite rodules,
bed, 3 m. thick, comes below the
middle flow, locally with limonitic
SS ——"
22] Banded felspar nodules (Fig. 3). Thisis succeed-
ee ee sands and clays.
di
EE, sei
-—- 3m. — 3 m.—— -——6 mi, —
ed by another complex bed, 3 to
-—— + Per
Se RR se, 4 m. thick, made up of multi-
À farious alternations of clays and
Sandy clay with
lignite.
sands, all retaining the original
Fi. 3.—Section exposed at the west coast
of Jiri, TTöko. horizontal position. Then comes
the third sheet of porous lava
of variable thickness, underlaid by a lignite bed, the last one
can be only seen at low tide. The whole seems to me to be
one complex bed belonging to later Teritary ; and this profile
serves as a type of the stratigraphical order of the island. After
passing over the second ‘ haul-over ’ to the Fükibi point (Plate V),
opposite to Bako, nothing but the two upper flows is exposed.
A table island, named Ko-sei-sho”,off the coast of Jiri,
already referred to, is an erosion relic of the Basaltic mesa,
surely connected in former times with the main island of Höko.
The adjoining wood-cut shows clearly the geological structure
oe
1) B. Koté, ‘Geologic Structure of the Malayan Archipelago.’ This Journal, Vol, XT.,
p. 113.
2) RAM.
OF THE DEPENDENT ISLES OF TAIWAN. 11
and the general view, as seen from Jiri, exhibiting the two
upper flows, mainly hidden by debris cones. This island served
for the Chinese in former times for the strategic base against
Fic. 4.—A view of the Isle of Ko-sei-sho, au erosion relic of Basaltic mesa,
as seen from the const of Jiri.
the Hollanders and Koku-sen-ya (Koxinga), in maintaining the
sovereignty over her supposed vassal domain of Taiwan.
Starting again from Sei-shi-an, already referred to, and
going round the south coast along the points of Kan-on-san”
and Kô-kaku”, Basaltic cliffs with underlying sandy bed, and
sandy coves repeatedly occur as far as A-kan”. At Sa-kan”, a
little south of the last-mentioned locality, fuller’s earth similar
to that of Bakö, is said to occur according to Tada and Ishii.
Upon the walls of the cliff at the recesses of the coves are found,
attached, according to Saitö, apparently recent shells, telling the fact
that at no geologically remote period, probably Diluvial, a negative
shifting of sea-level has taken place in this tract. We are,
however, not informed of the height of the former level, as
compared with the present; but at any rate it is of paramount
importance for us to have been acquainted with this movement
in view of the fact that on the opposite coast, 2e. on Front
Taiwan, there are not wanting evidences tending to prove the
negative change on the shore.
1) Mis 2UAO OHR 4) MAE.
12 KOTO : NOTES ON THE GEOLOGY
Between A-kan and Ri-sei-kaku”, the easternmost point of
the island, a white sandy beach bounds the south shore.
All: along the coast from Ri-sei-kaku to Hoku-ryo”, coral
reefs limit the eastern shore, and the detritals derived from them
form the beach-flat. It is a noteworthy fact that on the north side
the coast is very deeply indented in the north-south direction, and
the lowland, partly marshy, is covered likewise with coral sand.
I may here mention an occurrence of coal which was once con-
sidered to be a very important natural resource of the island,
though afterwards it turned out to be almost worthless and unworthy
of public attention. At one of the points, called Kotö or dragon
head, that stretches out northwards, a butte of Basalt, 22 m.
high, elevates itself from the shore, and at its northern foot
a seam of lignite, 5 feet thick, crops out with a sandy rock be-
tween the first and second flows, corresponding to the Arca zone
in Gio-6 Island, already referred to. The exposure is meagre and
soon disappears under the rubbish to be scen no more. This
mineral combustible is but imperfectly incarbonized, and the
woody structure is said to be yet well preserved.
From Sei-kei? through Kö-tei”, and Sha-k6” as far west as
to the oft-mentioned Bako, along the north coast, the two upper
flows are the sole rocks that can be seen, being covered with
an incoherent brownish, coarse and craggy earth.
HAKU-SHA ISLAND.
Haku-sha-t6,” or the white sand island is bodily connected
with Höko through the intervening islet of Chü-don”, at the
1) REAR 2) RR SUR 4) GER Bd Ti
OF THE DEPENDENT ISLES OF TAIWAN. 13
two narrow necks of the abraded second flow of Basalt, and
forms a part of the geological unit, differing from them only in
that here the interstratified sedimentaries seem to be wanting.
The other features that strike the eyes of observers are firstly,
the lowness of its relief, the highest point being Ké-don-san”,
36 m. high, and secondly, a considerable development of Alluvial
accumulation of the shells and skeletons of low organisms, hence
the name of the island. Cliffs, however, can be seen in its northern
shore, exposing the youngest flow with its usual columnar structure
at the water’s edge. White sandy flats prevail throughout the rest
of the lonely island, especially towards the Bay of Höko, and
the residual product of considerable thickness, derived from
the Basaltic decomposition, covers the interior.
One thing worthy of mentioning is a sporadic occurrence
of lapilli that had run aground on the east shore, probably
from one of the Indonesian volcanoes. The pumiceous fragments,
worn and rounded, belong to a Hypersthene-andesite with a
highly pleochroic, rhombic augite, and this rock either massive
Or pumiceous can be seen in no other parts of the group.
The islets, Impai? and Chö-sho® or Bird Island, off the
east coast, seem to be geologically identical, representing the
erosion-relics of the Diluvial epoch. A luxuriant growth of coral
reefs fringes the latter, as well as the neighbouring islets, just
as in Haku-sha.
KIPPAI ISLAND.
Farther away in a northerly direction lies the islet of
Kippai”, which is a low Basaltic flat, covered with halt-
1) 2G DAR AM NHR. NS +
14 KOTO: NOTES ON THE GEOLOGY
hardened foraminifer sand (Pl. II, Fig. 6.) of Recent age; frag-
ments of corals, bivalves and serpula mixed with other components.
The foraminiferal rock consists of millions of discoidal and
spiral, water-worn shells. Rarely they have spines well-preserved.
Viewing a section of the shell under the microscope, it is seen
that the test consists of the tubulated proper walls of chambers,
besides the canaliculated intermediate skeleton which forms
spur-like marginal appendages, characteristic of Calcarina, and
its external form and microscopic details agree well with C.
Spengleri, Linné”, dredged for the first time near the coast
of Amboina at the depth of 1,425 fathoms. This species seems
to be quite asabundant in the East Indian Archipelago, as we
find here in the Pescadores. By wear and tear of rolling waves,
the surface of the test becomes smooth, and the presence of spines
can be usually only recognized in examining the structure of the
supplementary skeleton which points to the former existence of
some sort of prominence.
GIO-0 ISLAND.
Gio-6, or Fisher Island, lies to the west of Hôko, and
encloses with the latter the head-less Bay of Hoko, or rather
an arm of sea. What has been said of other islands as regards
the geology and the topography, holds true also of Gio-6, with
the differences, that the island is really table-shaped, bounded
on all sides by cliffs, leaving no space for Alluvial deposits,
excepting the shore and fringing reefs; and that the igneous
sheet as well as the interbedded sedimentaries are developed to
their full advantage, thus affording the best opportunities for geolo-
1) Challenger Report, ‘F orafninifera.’
OF THE DEPENDENT ISLES OF TAIWAN. 15
gists to get insight into the geological structure, and to study
the stratigraphic details, of the whole Pescadores.
The oft-mentioned three flows and interstratified tuffites as
well as the underlving bed are likewise present, and well scen
Soil.
OO
ered No. 1. Basalt.
et
ee an Sandy tufüte.
Se dl
<q No. 2. Pasalt.
U
et.
»
a. »" LAN
er
ote
ard ©
ETS IS Ta Fel par san.
.
ete eee EPS
= 079%,
No. 3. Basalt.
A Sandy clay.
Fia. 5.—General profile ar seen in the southern
part of Gio-6.
especially in that portion that lies
southwards of Shö-chi-kaku.”
Between the last-mentioned
locality that is situated in the
middle of the island and Shü-
ba-wan,” good sections may be
traced, as in fig. 5, in descending
order. Under the superficial
covering of the ferruginous soil
of decomposition from Basalt
comes the No. 1. Basalt-flow,
with its usual columnar struc-
ture, of about one foot, and
sometimes disappears altogether.
The third in the series
consists of pelitic sand and
loose sandstone, the latter being
made up of muscovite, plagioclase,
and Basalt-glass. Concentric
nodules of hematite are frequent-
ly found in them. Saitd is
fortunate enough to find in this complex bed casts of an Arca and
gasteropod (Turbo) in the matrix of ferruginous felspar sand
with a little magnetite. Judging from the cast, the shell of the
D'hAf 2ER:
16 KOTÖ : NOTES ON TEE GEOLOGY
Arca is thick, egg-shaped, the ends of the margin obtuse-angled ;
the margin anteriorly rounded, posteriorly sloping; the beak
prominent, anteriorly inclined, widely separated and inflated ;
coarse radial ribs more than 20 in number. Our specimen
apparently resembles A. subcrenata, Lischke, though in details
they may differ, if perfect samples are taken in comparison.
The next in the series is the porous, No. 2. sheet, under-
laid by a fine felspar sand bed. Then the lowest, No. 3. sheet
of 6-7 feet, often Agglomeratic; and lastly, the bluish-grey
sandy clay, consisting of clay, muscovite, plagioclase and brownish
opaque grains probably of Basaltic glass together with carbona-
ceous matter. It is remarkable that muscovite is more or less
intermixed with in all the sedimentaries,
Before quitting Gio-ô, it should be remarked that the area
north of Shö-chi-kaku, as well as the whole east coast is com-
posed of the two upper flows only with or without interstratified
beds; while the rest of the island, as may be seen in fig. 5, are
built up of the second and third flows, accompanied with sedi-
mentaries, unsurpassed in complexity and in thickness,
According to Tada, the islands of the Southern Group (Pi.
IV.) of the Pescadores, are geologically of the same type.
Counting southwards, they are:—Hattô,” with the dependent
isle of Shô-gun-0” ; the Smaller and the Larger Biö-sho”, so
named cat islands from their appearance as seen from a dis-
tance; Tai-sho” and Shé-hei® with columnar Basalt; T6-kitsu®
and Sei-kitsu”, likewise Basaltic; all being encircled by coral
reefs,
)AE DEM SAN AM A OO) RA TEE:
OF THE DEPENDENT ISLES OF TAIWAN. 17
III. Petrography of the Effusives.
The groundwork of the Pescadores is essentially built up of
Basalts, making extensive flows to the water’s edge, and the whole
is encircled by the fringing reefs of corals, which, in parts raised
above the water, connect many ofthe detached rocks with the shore,
thus contributing greatly to the enlargement of the areas of the
island. Each and every island visited by Tada and Saitö, presents
the same physiognomy, and consists of the same black rock. The
specimens, brought back from moat of the islands, and of which
descriptions will be given in the sequel, have a certain common
feature which stamps them as genetically identical, and their
field relations in different areas seem to point to a common
centre of volcanic activity. They exhibit, however, a considerable
variation of character. Thus from the same island, I have
specimens at one place perfectly massive and compact, at another
vesicular and porous, and sometimes Doleritic. Colours vary
from black to bluish-grey in fresh ones, and through weathering
the Doleritic and vesicular varieties become whitish or grey,
while the compact rocks acquire a reddish brown tinge.
We are indebted to Mr. Y. Saité, for characterising the
different flows, and for tracing their vertical as well as horizontal
distributions in the Northern group. According to him, there
are three distinct Basaltic flows of nearly the same distribution,
separated by long time-intervals which are represented in inter-
bedded sedimentary rocks. Judging from the nearly perfect
horizontality which the beds and flows keep in all the islands,
it seems probable that there existed a lava field or volcanic mesa
of considerable extent. But, on account of its remote age, pro-
18 KOTÖ : NOTES ON THE GEOLOGY
bably later Tertiary, and of its insular position, waves gnawed
the ground in time, finally reducing the once wide volcanic
field into the ruins of islands, as we see at present. It is not
easy to know the former extent, and the ancient surface
feature, of this lava-flat; but, generally speaking, the relief
becomes higher as we go southwards from one to the other
in the islands of the Northern group. Saitö recognises, as I
have already said, three lava-flows in the -Northern group,
viz., the uppermost or youngest being of columnar, the middle
porous and vesicular, and the lowest also partly vesicular, and
Agglomeratic. After the comparative study of the Basaltic rocks,
to which the effusives exclusively belong, several important facts
are brought to light, and now I am able to say, that the young-
est flow (a, b and ? d types) contains the iddingsitized olivine, at
least in one type, and violet brown titan-augite; the second
(c and ? d types) the brown augite, olivine sometimes lacking,
being often replaced by hypersthene ; and the third (e type) the
analcime-bearing. I will record first my observations on the
component-minerals, and then give the special description of rocks.
A. Component-minerals of Basalts.
OLIVINE.
Olivine is rarely automorphic, but mostly xenomorphic,
being the remains of resorption by magma. The olivine in the
Basalts of the Höko Islands seems to be of several varieties.
Automorphic ones show vivid polarisation-colours, and alter
usually into some red minerals. The xenomorphic type shows com-
paratively a low degree of polarisation, and suffers deep corrosion,
OF THE DEPENDENT ISLES OF TAIWAN. 19
often being reduced to a mere grain, and is also traversed with
fractural lines, from which the mineral begins to form a ser-
pentinous substance. The olivines are undoubtedly the intra-
telluric products, being sometimes enclosed by an automorphic
augite, and large individuals are habitually surrounded by
heaps of the crystals of augite (in Andesites, instead of it often
hypersthene). Inclusions of gas and liquid are not rare, and
the octohedra of magnetite are also found in the olivines.
Zonal structure of olivine is, as is well known, of rare
occurrence, and if it really exist, this could only be discerned
either by measuring the optical angles at different portions, or
by finding the altered zones in a crystal in consequence of the
formation of the mineral rouge. The zone of the red mineral is
not constant in position, for, it makes its appearance sometimes
on the periphery, at other times in the interior; but, so far as
my experience goes, the recurrent zones are never found. The
condition under which the isomorphic shells of different chemical
compounds are formed in the olivine, seems to depend, as
Lagorio” and Morozewics” say, mainly on the Massenwirkung,
that is, the degree of saturation of magma in certain temperature
and pressure. In my slide, in which olivine has a red central
zone (the Kippai Island specimen), magnetite is scarce, and large
in its size and rod-shaped ; while the magnetite-rich rock (the Héko
specimen) has an olivine with an external red zone. Here the
magnetite occurs in small isometric crystals and grains.
The red mineral, that forms the periphery (Pl. J. Figs. 1,
4 and 5) and the kernel (Pl. I. Fig. 6), differs in habit. The
1) ‘Ueber die Natur der Glasbasis, sowie der Krystallisationsvorgänge im eraptiven
Magma’ T. M. M. Bd. VIII, 1887.
2) Ibid. Bd. XVIII, 1898.
20 KOTO : NOTES ON THE GEOLOGY
first has a facile cleavage but brittle, and consequently becomes
lamellar like a brittle micaceous mineral. It is probably identical
with the so-called biotite, which we occasionally find mentioned
in petrological literature, as being formed from an alteration of
olivine, just like as schillerspar has been considered to be a mica,
as an alteration-product of enstatite. Recently, Iddings” and
*) described a similar mineral and the latter author
Lawson
named it iddingsite. In the second, we fail to find such a
distinct cleavage, and it seems to me to be the same body which
Michel-Lévy called the mineral rouge”. Now, a question suggests
itself to me, whether the red micaceous mineral is identical with
the mineral rouge or not? It is true that the former confines
itself to the margin, and in the case where the entire substance of
olivine has been transformed to this mineral, the process of
alteration has started from the periphery, and it not infrequently
happened to me to find every stage of progress from the very
beginning to the complete alteration. The latter, on the contrary,
starts from the centre in irregular patches, and gradually attacks
the whole body but the clear and granulated, thin margin. The
formation of the red lamellæ begins with the development of a
fine parting which appears like stripes, and which runs parallel
to the vertical axis (Pl. I. Fig. 1); while cracks on the margin
favour the olivine being changed into the red mineral in the
centre.
In my opinion, there may be a slight difference in the
i ee nn um
1) U. 8. Geol. Surv., ‘Monograph’ XX., p. 388. Iddings identifies this mineral to
thermophyllite, a foliated mineral having the composition of serpentine.
2) ‘The Geology of Carmelo Bay.’ Bulletin of the Department of Geology in the
University of California, Vol. I., p. 51. See also Pirsson’s paper, Amer. Journ. Sci, XLV,
1893, p. 381.
3) ‘La Chaine des Puys et le Mont Dore,’ Bull. Géol. Soc. France, 3me Serie, XVIL,
1890.
OF THE DEENDENT ISLES OF TAIWAN. 21
chemical composition of the two alteration-products, yet on tlıe
whole they must be practically identical. The lamelle are
oriented parallel to one of the pinacoids, as may be deduced from
the position of the optic plane (in the fresh substance of olivine),
which stands at right-angles to the easy cleavage (Pl. I. Figs. 1
and 5). Pleochroism is distinct; it is brownish-green in the
direction of facile cleavage, but greenish-brown when at right-
angles to it. Hence, c>a or 5b. Mügge”, however, says that
the absorption is stronger in the direction perpendicular to the
‘ Längsrichtung’ than in that parallel to it. Zirkel” and Rosen-
buch? interpret the above statement in the terms, that the rays
vibrating parallel to c absorb far less than those parallel to a
and 6. The observers, however, seem to have examined the
mineral rouge. My observation, therefore, accords well with that
made by Lawson for iddingsite; but it is not known to which
pinacoid, 010 or 100, the lamelle are parallel, though it is
probable that the brachypinacoid is the lamellar plane, as may
be inferred from the fact that the elasticity perpendicular to
the lamellæ is greater (X=6) than that parallel to the c-axis, the
latter corresponding to the mean axis of elasticity.
With HCl, the iddingsitic mineral becomes bleached, and
then acquires a greenish-yellow colour, with corresponding decrease
of pleochroism. Bearing in mind the fact of the brachypina-
coidal lamellar cleavage, of the colour, and of the chemical com-
position which is a hydrous non-aluminous silicate of iron, lime,
magnesia, and soda, J am rather inclined to consider the iddingsite
to be a mineral approaching to basite. Prof. Rosenbusch‘,
1) Neues Jahrbuch, 1883, II, 8. 205.
2) ‘ Petrographie,’ Bd. I, 1893, S. 353.
3) ‘ Physiographie,’ Bd. I., 1892, S. 469.
4) ‘Physiographie,’ 1892, Bd. IL, S. 461.
22 KOTO : NOTES ON THE GEOLOGY
in speaking of bastite, says ‘die Umbildung scheint in
hohem Grade durch die gleichzeitige Anwesenheit des Olivin
und dessen Umwandlung zu Serpentine befördert zu werden.’
Chemically speaking, there exists a close resemblance between
iddingsite and the ‘ crystallised diallage’ of Baste”, considering
out of question a trace of alumina. Optical schemes differ, of
course, in the two minerals, but I could not make out surely
the optical orientation of iddingsite in my slides, on account
of its extremely fine lamellar structure.
PLAGIOCLASE,
Plagioclase has, generally speaking, crystallised out in a
single generation of the flow period. Differing from the Ande-
sitic plagioclases which present various dimensions, the felspar
of Basalt is uniform in size. It is, however, not wanting in
large, phenocrystic crystals in some slides, which also belong
to the products of the effusive period, slightly earlier in crys-
tallisation than the ones in the general mass; for, the small
laths of plagioclase are partly embraced by the phenocrysts,—
a fact which also leads me suppose that the plagioclases have
grown in a comparatively motionless magma. They show no
signs of corrosion, so common in the olivine of the intratelluric
origin, though the effects of tossing and fracturing of crystals
are by no means seldom observed.
The phenocrystic plagioclase (Pl. II, Fig. 2) has a tabular
form on M, somewhat elongated towards the vertical axis. Zonal
structure is rare in contrast to the Andesitic felspar; the same
1) Hintze, ‘Handbuch der Mineralogie,’ Bd. II., S. 972.
OF THE DEPENDENT ISLES OF TAIWAN. 23
is the case of glass-enclosures, save that ground-mass which fills
up the rectangular space between the lamellæ, showing as if the
larger crystals have grown out by the apposition of numerous
flowing lamellæ. Penck” holds the same view, as given here.
In consequence of this lamellar composition, which is one of
the causes of the paucity of inclusions, both terminations of the
ledges became indented and forked, after the manner of a
parapet (Pl. I, Figs. 4 and 6., Pl. II, Fig. 5), a characteristic com-
mon to all the plagioclases of Basalts. Jt seems more rea-
sonable to consider these monstrosities as incipient forms of growth,
having simultaneously many centres of crystallisation in space,
which in later stages have grown together to make up one ın-
dividual with but internal complex compositions. Morphological
and optical homogeneities are, however, frequently disturbed
through the flowing motion and sudden cooling of the consolidat-
ing magma. Several stages of similar kind in crystallisation
may be frequently observed under the microscope in the forma-
tion of artificial crystals.
Symmetrical but contrary extinction takes place at the
maximum angle of 33°-35°, with reference to the suture of the
albite-twinning, and the extinction with regard to the pericline-
lamelle amounts to —16°, showing that the plagioclase is of a
basic labradorite. It is easily acted on by HCl. The Baveno
twins were once observed.
The plagioclase in the ground-mass is lath-shaped, extremely
slender, and polysynthetic ; termination being also a parapet-like.
The habit of crystals is prismatic, and such a form is said to
be elongated parallel to the a-axis. This is indeed true; for,
1) ‘Studien ueber lockere vulcanische Auswiirflinge.’ Zeitschr. d. d. geol. Gesell. Bd.
XXX, 8. 101. Taf. V., Figs. 3, 5, and 7.
24 KOTO : NOTES ON THE GEOLOGY
along the longest extension lies the axis of greatest elasticity,
and there are tens of thousands of laths visible in microscope
slides, with but a few tabular sections. Symmetrically opposite
extinctions make the maximum angles of 23° to 25° with the
suture of lamellæ. Microlithic sections twinned on the albite
type extinguish at the angles from 0° to 26°, with reference to
the longer dimension. According to Michel-Lévy, labradorite
and albite have similar optical deportment, but as they do not
usually come together, and as we are dealing now with a basic
rock, the nature of the microlite should be considered to approach
that of an acidic labradorite. In a few slides, the poles of
the laths resolve themselves into a number of prisms, and such
fine slender needles are scattered through the whole groundmass.
AUGITE.
Augite, so says Morozewicz” belongs to one of the ‘ ver-
hangnissvollen minerals. It does not obey Fouqué and M.-Lévy’s
rule of crystallisation of silicates in the reversed order of
fusibility, nor Rosenbusch’s scheme of crystallisation according
to acidity. It is rather subjected to the influence of masses, 7.e.
a degree of saturation under certain temperature and pressure.
Under such circumstances, augite may form crystals before
plagioclases, and at other times, just the reverse may occur,
while in the third case they may individualise at the same time.
According to the priority of secretion of either of the two
minerals, in other words, the relative idiomorphism of one to
1) ‘Experimentelle Untersuchungen ueber die Bildung der Minerale im Magma.’
Tschermak’s Müth., Bd. XVIII. 8. 84.
OF THE DEPENDENT ISLES OF TAIWAN. 25
another, various structures may be brought about under vary-
ing conditions, and these we find in fact in my slides.
As regards the forms of augite, it is sometimes idiomorphic,
bounded by faces © P®, o Pd, o P, and P&, the first being well
developed, consequently the crystals become tabular; at other
times granular, needle-shaped, in ophitic plate, and in partial
crystals. Prevailing colour is either violet or yellowish-brown.
It is to be. expressedly remarked that the typical Basaltic augite
with a tinge of violet occurs only in the Pescadores, and in the
dykes of Basalt near Taihoku and Taikokan, in Formosa. My long
experience forced me to conclude that, in Japan proper, the
Basalt with the violet augite is confined to the northern Kiû-
shia, and Chiu-goku, in Hondô, as far east as the provincial
boundary of Tajima and Tamba. The same type of Basalt is
also known to be wide-spread in Korea, Liau-tung, and Mongolia.
Thus the distribution of the Basalt with the violet tilaniferous
augite marks a definite area, being, so far as my knowledge goes,
confined to the inner side of the festoon islands and the adjoining
continent in Eastern Asia, constituting the well-defined Japan-
China petrographical province. Larger crystals show a zonal
structure, coloured intensely on the periphery, and the hour-
glass structure occurs frequently with deeply-coloured, additive
cones in the prismatic zones, which have at the same time a
greater angle of extinction. Pleochroism is stronger in the
direction parallel to the c-axis. Polarization-colours are generally
weak in comparison to those of the Andesitic augite. Twins on
oP have a suture, running just along the middle of the body
of the crystals, Crystals often form stellar aggregates; they are
generally free from foreign interpositions, excepting the larger
ones which have sometimes enclosures of glass and magnetite.
26 KOTÖ : NOTES ON THE GEOLOGY
HYPERSTHENE.
Hypersthene takes the place of olivine in some Basalts of
the Pescadores ; consequently the presence of one totally excludes
that of the other, —a state of thing quite exceptional to the modern
Japanese Andesite of a glassy, black, porphyritic type, in which
both minerals appear always concomitantly. We have then the
Hypersthene-Basalt, in lieu of the Basalt proper. It is a note-
worthy fact that this stray variety of rock seems to be wide-
spread, at least in my specimens, in the out of the way islets,
such as Impai, Kin-sho, and Hatté, the only exception being
the one from Sei-kei (West Valley) in Höko, though I could
not find a sufficient reason accounting for the special dis-
tribution of this hypersthene-bearing rock.
It is usually a comparatively easy task to discriminate
hypersthene from olivine, but in the present case some difficulty
is experienced in making out for certain the presence of the
former.
In regard to the form, the (1) hypersthene is extremely
slender, being about six times longer than broad, and, as being
of the intratelluric origin, it has a marginal zone deeply corroded
and partly granulated, and has indefinite faces at the poles
of the crystals (Pl. II, Fig. 3). I observed once a morphotropic
growth of a highly-polarising, monoclinic pyroxene around a
hypersthene, just as is the case in Andesites. Cleavage is
developed along the longest extension of the crystals. In a patch
of a coarse aggregate which appears as an endogeneous or homoge-
neous enclosure in the finer general mass, the (2) hypersthene
comes together with plagioclase and augite, and in this case the
hypersthene occurs in broad plates (Pl, II, Fig. 4), with only a few
OF THE DEPENDENT ISLES OF TAIWAN. 27
traces of cleavage, but with numerous fissures; and has an ap-
pearance exactly like olivine.
The hypersthene possesses a brown colour, and its pleochroism
is scarcely discernible. In favourable cases, the ray vibrating
parallel to the c-axis is slightly green. Sections present a rough
surface, owing to its having a high index, approaching to that
of olivine; its polarisation-colour is grey.
From the brief diagnosis, given above, of the hypersthene,
its cleavage, colour, non-pleochroism or very weak if present, high
index, but low magnitude of refraction, extinction-direction, and
similar chemical composition,—these several physical properties
afford no means of discriminating it from a fresh olivine. Olivine
has, however, a lighter colour, and has usually but one trace of
cleavage in a section. The hypersthene on the other hand pos-
sesses the characteristic traces of prismatic cleavage, which in
a random section gives scarcely a clue to distinguish it conosco-
pically from monoclinic pyroxene. A basal section, once observed,
presented a square outline, truncated little at the four corners.
From the combined evidence of more slender section, of the
want of decomposition-products, of indifferent behaviour towards
common acids, of the presence of comparatively numerous
traces of cleavage, I infer, in the Basalts, the presence of a hypers-
thene. It is to be remembered that the prismatic sections of
olivine show also a low colour of polarisation, exactly like that
of a hypersthene. It seems to me that the hypersthene in
the Höko Basalts stands in its chemical composition near to
that of dronzite. The want of a distinct pleochroism may be
attributed to the same cause. Axial angles, therefore, become
large, and the axial poles were not observed in any of the
pinacoids by ordinary methods. u ay
98 KOTO : NOTES ON THE GEOLOGY
APATITE.
Apatite occurs in the Doleritic or Anamesitic rocks in the
form of extremely fine needles, devoid of terminal faces,
being colourless, and always traversed by transversal fissures.
Its crystals sink almost to a minimum size, and are not, com-
paratively speaking, so large as those found in the typical
European Dolerites; and for this reason they might be easily
mistaken for the microlites of felspar which often resolves from
the poles of a larger crystals in Basalts. The apatite is typi-
cally found in the three slides only, which are in my pos-
sesion (Kippai and Höko), and both are magnetite (not ilmenite)-
bearing rocks. The crystals are dark-margined, owing to the
total reflection of light on the prismatic faces; and sometimes a
single brown-coloured axis entirely or partially runs through the
crystal. A grey or light-brown variety, so often found in An-
desites, is entirely absent, though a dark-brown crystal of an
apatite-like mineral was once observed with strong absorption
parallel to the prismatic axis. The sure criterion of the presence
of apatite can only be found in its hexagonal cross-scction.
ANALCIME AND NATROLITE.
A cave-rock in the southern Gio-ô, presents an anomalous
habit ; a slide made of it contains a colourless mineral in angular
or polygonal interspaces between the crystals of plagioclase
(Pl. II, Fig. 5). It shows no signs of any crystallographic face,
nor cleavage, but only has a frittered appearance, being traversed
with irregular cracks, and also being pierced through in all
directions with the needles of apatite which is excessively rich
OF THE DEPENDENT ISLES OF TAIWAN. 29
in this rock. The polysomatic mineral has a smaller index of
refraction, when compared with that of the accompanying pla-
gioclase, as may be easily experimented upon by Becke’s method.
These colourless patches, as a rule, behave optically isotropic ;
at times, however, faintly double-refractive, and separate into
several optical fields. They readily dissolve in HCl, with the
formation of the cubes of rock-salt. The same patches frequently
resolve themselves into a radial-fibrous, somewhat brownish and
highly double-refractive body (well seen at the margin in the
lower, left quadrant in Pl. II, Fig. 5) with the positive sign along
the axis of the needles.
The polygonal base-like mineral, moulded upon plagioclase
and augite, seems to be identical with what Biking” calls the
‘ Basis zweiter Art,’ and is allied to the pitchstone-glass of Hunter
and Rosenbusch.” Recently, this base was studied with great zeal
by the American petrologists, Lindgren,® T. F. Williams,” Kemp,”
Fairbanks,” Cross,” Coleman® and Pirsson”; the last author
especially paid closc attention to this subject, in making care-
ful analyses and also recalculating the analytical result, ob-
tained by Hunter. From his study, Pirrson is forced to the
conclusion that the so-called colourless base has exactly the
1) ‘ Basaltische Gesteine, etc.,’ Jahrb. K. K. preuss. geol. Landesanstalt. 1880, S. 153, und
1881, S. 606.
2) ‘Ueber Monchiquite, ein camptonitisches Ganggestein aus der Gefolgenschaft der
Eleolithsyenite,’ Tschermark’s Min. Mitth. XI, 1890, 8. 445.
3) Proc. Cal. Acad. &ci., Vol. III, 13%.
4) Cited in Pirsson’s paper.
5) ‘Trap Dikes, Bull. 107, U. S. G. 8. 1893.
6) ‘On Analcite Diabase from San Luis Obispo County, California,’ Bull. Geol. Depart.
Univ. Cal, Vol. I. p. 272.
7) ‘An Analcite-Basalt from Colorado,’ Journ. Geol. Vol. V. p. 684.
8) ‘A new Analcite Rock from Lake Superior’ Journ. Geol. Vol. VII, 1899, p. 422.
9) ‘The Mochiquites or Analcite Group of Igneous Rocks,’ Journ. Geol., Vol. IV. 1896,
p 879
30 KOTO : NOTES ON THE GEOLOGY
same chemical composition as that of analcime, and the physical
properties observed give no hinderance to the assumption that
this component actually is that mineral. He thinks the analcime
is primary, having been formed from the magma, containing
water and much soda, under pressure with considerable rapidity.
From what has been stated before, I have also, to all ap-
pearances, the primary analcime in the interspaces of the com-
ponents in the Basalt from Gio-6, and the radiating bundles of
a strongly birefringent natrolite are formed secondarily from the
analcime through a molecular rearrangement. Both components
make their appearance with the dodecahedric networks (Pl. II,
Fig. 5) of the skeleton magnetite which occupies the other portion
of the slides.
THE IRON ORES.
Both ilmenite and magnetite are present, and they usually
belong to a single generation, and indeed the product of the
effusive period, as the iron ores were not found enclosed in the
olivine of the intratelluric crystallisation. Both ores, especially
the ilmentite, have crystallised Zafer than plagioclase, but slightly
prior to, or contemporaneous with, the monoclinic pyroxene.
The ilmenite and magnetite are, under the microscope, not easy
to be distinguished, as every petrographer will agree, if crystal
forms are not serviceable for their diagnosis.
The ilmenite is, however, tabular and needle-shaped in sec-
tion in "the Basalt with a strong lustre and a violet tinge, when
seen by reflected light, on the flanks corresponding to the
thickness of slide. The laths are slender, appearing merely as
lines, and cross several crystals of felspar and augite, mean-
OF THE DEPENDENT ISLES OF TAIWAN. 31
while the substance of the ilmenite entirely disappears when
traversing other crystals, and comes again into view in the same
direction as a continuation of the interrupted crystals. Unfortu-
nately basal sections were not frequently observed, and this
was the greal obstacle in ascertaining the presence of ilmenite
n microscopic analysis. The ore with above-mentioned lamellar
habit occurs exclusively in a coarse-crystalline type of intersertal,
or ophitic structure, irrespective of hypersthene or olivine-bearing
Basalt ; and this fact lends evidently a strong support to the view
advanced by K. Hofmann,” that the ilmenite accumulates in the
lower portion of lava-flows, and in that which has crystallised
under high pressure, while the magnetite is rich in the upper
part that has consolidated under a low pressure. Fr. Sandberger”
says also that Basalts may be classified into Dolerite and Basalt
proper, by the presence of ilmenite in the former and magnetite
in the latter. These fruitful ideas inaugurated by both authors,
now unfortunately passing into oblivion, deserve the careful
attention of petrologists.
A slide of the Basalt from the islet of Hattô was treated for
a considerable length of time with a strong hydrochloric acid
without any appreciable result. A large quantity of the pul-
verised sample of the same specimen was then digested in
boiling HCl with the addition of tin-foil, and the solution was
coloured slightly violet, showing the presence of titanium in the
dissolved portion of the ore. Ilmenite also occurs, according
to Vénukoff®, very abundantly in the Basalts of Mongolia, and
even transparent lamelle were found by him, just as in the Pes-
cadores rocks. The ilmenite is fresh and leucoxene not noticed.
1) ‘ Basalt von Bakony,’ Zeitschr. d. d. geol. Ges., XXIX., 1877, 8. 191.
2) Rosenbusch, ‘ Mikroskopische Physiographie, II., 3te Auflage, S. 1015.
3) ‘Les Roches Basaltiques de la Mongolie,’ Bull. Soc. belge de Geologie, ele. T. D., p. 448.
32 KOTO : NOTES ON THE GEOLOGY
In my few slides of Basalts, bearing the iddingsitised olivine,
ilmenite seems to be wanting, though the rocks approach to a
Doleritic type, being replaced by magnetite. I cannot say
positively that this rule holds true for all the iddingsite- bearing
Basalts.
The magnetite is, on the other hand, the prevailing ore in
the compact Basalt, and in the Limburgitic type, in the form
of isometrie crystals and dust, occurring either in the general
mass, or else enclosed in augite and olivine. The face of the crystals
shows a metallic lustre with a tinge of blue by reflected light.
The dust is sometimes peripherally altered into a blood-red
iron-glance. A slide made of a chip from Höko, was digested
in HCl with the addition of KI; and then the black ore,
therein contained, was entirely removed, and the solution not
coloured when tested with tin-foil, proving thus the presence of a
pure magnetite. As it is already stated above, the magnetite-
rich, compact type seems to make up the upper portion of the
thick flows of the Höko Basalts.
In the Anamesitic type from the islet of Gio-6, we find beauti-
ful networks of the skeleton-crystals of magnetite in a devitrified
mesostasis within the polygonal spaces between crystals. They
are the dodecahedric dendrites, consequently the skeletons inter-
sect each other at the angles of 60° and 120°, and are said to
consist of garnetohedrons. They all gointo solution by treating
with HCl. Morozewics” tells us that the spire and filigree-work
of the skeleton magnetite, crystallising out of the magma rich in
iron oxides, consist of minute octahedra, arranged rectilinearly —
in the direction of the crystallographic axes with secondary and
1) ‘Experimentelle Untersuchungen über die Bildung der Minerale im Magma,’
Techermak’s Mittheilungen, 18, 1898, S. 90,
OF THE DEPENDENT ISLES OF TAIWAN. 33
tertiary offshoots. This mode of growth, the octahedric dendrite,
so called by Morozewics, is well known in petrographical
literature, since the publication of Prof. Zirkel’s’ work. On
the other hand, it is said that the dodecahedric dendrite, as is in
the present case, is formed out of the magma poor in iron-
oxides.
B. Special Description of Individual Occurrences of Basalts.
A, THE GRANULAR TYPE.
(Pl. I, Figs. 1 and 2.)
As seen by the naked eye, it is greyish-black and compact,
with the dots of olivine which is the only visible component
of the whole mass. This type is represented by two specimens
from Höko, and one from Hakusha.” Microscopically it is
holocrystalline with the smaller phenocryst of olivine, imbedded
in the still finer aggregate of the ground-mass.
The fineness of the ground-mass, however, varies in different
specimens, and even in the same slide. Some portion of the
same slide is, therefore, extremely rich in idiomorphic augite
to the total exclusion of felspar and olivine, but with small
patches of brown glass. Were this portion independently devel-
oped, it would be fitly called the Augitite (Fig. 2). It is the
local assemblage of augite within the rock, and that mineral es-
1) ‘ Die mikroskopische Beschaffenheit der Mineralien und Gesteine.’ Leipzig, 1873, 8. 244.
2) Collected at Ryö-Lösan (ll); and, according to Mr. Sait6, it appears in I. horizon,
2. e., the uppermost sheet, consequently the youngest of all the lavas of the Hôko Group
(Pi. I, fig. 1).
34 KOTÖ : NOTES ON THE GEOLOGY
pecially accumulates near the margin of the secretionary mass,
the augite being sometimes arranged along the linear common
base, with the free ends of the erystals toward the interior. These
phenomena indicate that the lava had consolidated in a quiet
state.
The relative proportion of augite and plagioclase is also vari-
ous, and in the cases where the former outweighs the latter, tlıe
olivine increases in its quantity and comes also in the ground-
mass, as a product of the crystallization of the effusive period ;
and at the same time the texture of the rock becomes finer.
If, on the other hand, the plagioclase becomes predominant over
the augite, then, the texture gets coarser and more crystalline,
and the distinction between phenocrysts and ground-mass is not
then commonly well marked. Apatite and ilmenite seem to occur
in the latter variety only, the ilmenite is sometimes transparent
with a deep brown colour.
The only mineral that serves as the phenocryst is olivine.
Its forms are various, owing to the various degrees of resorption.
Most have partial crystallographic faces with deep indentations
of corrosion, and a drop-like black zron-ore and felspars were
formed in those spaces. Sometimes the act of corrosion has ad-
vanced so far that there remain but patches as the relics of a
large crystal, and the eating away of the body by the magmatic
menstruum proceeds always from the lateral pinacoids. As usual,
the crystals of the olivine are not fresh ; but the routine of change
is the same in all. They become fibrous and lamellar, parallel to
one of the lateral pinacoids, the altered portion being yellow or
brown, according to the degrees of transformation. The mode
of change is similar to 2ddingsilizalion (Migs. 1 and 2).
The ground-mass consists, first of all, of the crystals and
OF THE DEPENDENT ISLES OF TAIWAN. 35
grains of augite, all of a violei-brown colour, besides the grains
of olivine, and the laths of the multiple-twinned plagioclase, the
octahedra and dust of magnetite, and ilmenite. The texture of
the rock is crystalline and typically granulitic. In a coarse
variety, the idiomorphic augite with hour-glass structure forms
stellar aggregates, and these aggregates closely resemble the
glomeroporphyritic phenocryst.
B. THE TYPE OF THE IDDINGSITE-BEARING BASALT.
(Pl. I, Figs. 4, 5 and 6; Pl, II, Fig. 1.)
Megascopically this type is greyish-black Anamesitic-looking,
and finely uniform-granular, owing to the nearly equal size and
form of the laths of plagioclase which predominates quantitatively
over the other components.
The characteristic featurcs of this group are firstly, the pre-
sence of large phenocrysts of olivine which is more or less iddingsi-
tized; secondly, the majority of the augite is xenomorphic or
granular, and of small size, and these grains are grouped together
intersertally with the devitrified glass between the laths of
plagioclase. The structure is typically intersertal. The promi-
nent characiers distinguish this group from the rest of the Basalts.
This type is represented in my slides from the Pescadores by
three specimens, one from Kippai, and two from Höko, one of
which was struck off at the locality Tai-san ;” according to
Y. Saitö, it forms the uppermost flow there. ‘The same may
be said of the specimen from Kippai, since the youngest
1) FAW, see, WH. The rock eflerve:ces with acid. The microscope discloses the
fact that the radially arranged fibres of calcite fill up the polygonul spaces between other
components, showing bars which correspond to the position of crossed nicols.
36 KOTÖ : NOTES ON THE GEOLOGY
lava-flow is the only effusive that can be met with on that
island.
Lhe olivine is the sole phenocryst; it is variable in size (the
largest one measures even 5 min.), irregular in distribution, and
multifarious in form, some having partial crystallographic faces,
while others have none of them. The iddingsitization is pecu-
liarly inherent in the olivine of this rock-group, and I refer the
readers for further details to the topic: “ component-minerals ”
p. 18 et seg. By the way, I have only to mention that the name
iddingsite may conveniently be applied to a special transitional
form of the alteration of an olivine which, after passing this
stage, changes into dirty-green spherulitic fibres of an optically
positive character.
In the felspar-rich rocks (PX. J, Fig. 6), which are prevalent
in the group under question, the plagioclases arc all approximately
of the same size, and surpass the augite both in dimension and
quantity; while in the augite-rich rocks (Pl. J, fig. 4), the
plagioclases are of two generations, and the larger ones behave
porphyritically towards the minor ones. ‘They are lath-shaped,
and multiple-twinned, the terminations being imperfect and
sheafy, and these laths are thrown together in an orderless
plexus, which eminently characterises the structure of normal
Basalt in contradistinction to that of Andesite.
Lhe augite is all of a single generation, consequently uni-
form, but inferior in size to the plagioclase and olivine. Some
are rudely idiomorphic, but by far the most of it is granu-
lar, occurring in groups, and filling the angular spaces left
by the laths of plagioclase. The augite is, as usual, of a violet-
brown colour, but in the specimen from Tuai-san, it is almost
colourless in sections. It is free from foreign inclusions, and
OF THE DEPENDENT ISLES OF TAIWAN. 37
the hour-glass structure is faintly indicated in some individuals.
In the coarse, felspar-rich specimen, the iron-ore is present only
in small quantity (Pl. J, Fig. 6), but comparatively large, lamel-
Jar and flat with glittering bluish lustre on the perfect cleavage-
surface. It looks rather more like ilmenite than magnetite. Stiff,
slender apatite-needles, sometimes with a brown canal traversing
the whole length, are particularly abundant, being scattered
through the. whole mass,
In the dark fine specimens (Pl. I, Figs. 4 and 5), small
regular crystals of magnetite are plentiful, and in these slides,
I found abundantly the small laths of twinned plagioclase, which
resolve at the ends into slightly diverging columns (Pl. I, Fig. 5),
and these may be easily mistaken for those of apatite, if needles are
found detached from the waist. Optical properties are not in-
dependtly shown in them, on account of their extreme thinness.
Similar bodies are noticed by H. 8S. Washington in the sanidine
of some Ischian Trachytes and named by him keraunoid.”
He and also Lehmann” attribute the splittings and ramifications
from the main crystal to the existence of internal tensions in
felspar, but the cause of the existence of such tensions remains
to be solved.
The glass together with the augite fill up the polysynthetic
space left by the laths of plagioclase. The glass-base is coloured
bottle-green, sometimes dirty brown, and devitrified in various
ways. It consists of polarizing scaly aggregates of vermiform,
spherulitic, or, irregular shapes. Sometimes fascicular and
radiating needles, which are colourless and birefringent, are
imbedded, in the green base as a product of devitrification. The
1) ‘On some Ischian Trachyte.’ Journ. Amer. Sci, Muy, 1896, p. 350.
2) ‘Molecularphysik,’ I, 1888, S. 378.
38 KOTO : NOTES ON THE GEOLOGY
needles may possibly of a felspathic nature and such a structure
is termed variolitis by Harker,” though in the present case those
circular, whitish spots, called varioles, are wanting. Green,
fresh base is here and there also found between the angular spaces.
Thin lamellae of rugged outlines, with violet-brown colour,
may be frequently noticed in all of my slides, and they closely
resemble those found as interpositions in a hypersthene. I
conjecture the mineralogic nature of these plates to be ilmenite.
C. THE OPHITIC TYPE.
(Pl. II, Fig. 2)
This type is represented by a single specimen from Höko,
and Gio-6” respectively, and two from Haku-sha, though the
‘lie’ is not known to me exactly ; but it is highly probable that
samples are taken from the second sheet which is separated
usually from the uppermost columnar flow by an ash bed ofa
certain thickness. Itisa greyish-black, Anamesitic rock, with the
brownish, lath-shaped phenocrysts of plagioclase (4 mm. length).
This is the coarsest type of the Höko Basalts, and is the one rich
in plagioclase in comparison with ferro-magnesian silicates; it
seems to have solidified in the lower portion of the lava flow.
Under the microscope, it shows a porphyritic, hypocrys-
talline, diabasic structure (Fig. 2) with the ophitic plates of
augite of considerable dimensions, enclosing the laths of plagio-
clase which lie in all possible directions. The augite is of a
1) “The aggregates of felspar-microlites or fibres with fan-like or sheaf-like groupings.
They may be closely packed to make up the entire mass of a portion of the rock (Basalt).”
Petrology for Students.’ 2nd. Edit., p. 191 and 201, Fig. 41 A,
2) The exact locality being Sho-chi-kaku, (hf) at the middle of the island.
ON THE DEPENDENT ISLES OF TAIWAN. 39
kind of light-brownish colour, and its plates are often multiple-
twinned, and enclose, besides plagioclase, a number of round and
corroded crystals of olivine which is for the most part changed
into green, pleochroic fibres ; the iddingsitization of the olivine was
so far not observed. The plagioclases are of two generations (Fig. 2),
the larger, probably intratelluric, species has fissures (see Fig. 2)
filled with films of brown hydrous sesquixide of iron, which
cause the phenocrystic feldspar to appear macroscopically like an
olivine. The plagioclase is partially embraced by the ophitic
plate, while the smaller laths became entirely enclosed in it.
The polygonal interspaces, when not occupied by augite, are
otherwise filled up with the fibrous devitrified glass, the latter
containing globulites, sometimes dendritic, and apatite; and the
thick lamelle of c/menite traverse the base, but not the plate of
augite, consequently the crystallisation of the ore must have
taken place posterior to that of the pyroxene. Sometimes the
greenish-yellow augite is coarse-granular, and in this case the
structure approaches to that of anfersertal. Magnetile seems to
be wanting. Owing to the coarseness of the structure, the rocks
are often porous, and the polygonal, angular spaces are often
filled up with banded, purplish chalcedony.
D. THE TYPE OF THE OLIVINE-LESS BASALTS.
(Pl. I, Fig. 3; Pl. IL, Figs. 2 and 3.)
The olivine-lese, hypersthene-bearing Basalts are represent-
ed in my collection by two specimens from Wampai", and one
from each of the following islands, Höko”, Kin-sho”, and
Hattô-sho*. They are megascopically wet-grey, and fine-granular,
MH 2) Sei-kei WR in Hoko DE 4) ABS
40 KOTO : NOTES ON THE GEOLOGY
the general microscopic aspect being a crystalline Andesite-like.
They are all extremely rich in augite, and the structure is granu-
htic. The felspars are of two generations (Pl. II, Fig. 2 and 3),
and the rock is consequently porphyritic, owing to the presence
of a few large crystals of plagioclase, though this structure could
not be easily recognised as such in the present group.
The phenocrystic plagioclase is narrow-tabular with a few twin-
ning lamelle (see Fig. 3), and is remarkable in its being traversed
through by sets of cracks which run approximately parallel with
each other. In one instance, only one lamella, out of many twin-
ned parts after the pericline law, is provided with close/y set fissures.
This anomalous feature can be seen in all the specimens of the
present type, but not common in others, and the same peculiarity
recurs also in augile whose granular aspect is due in great measure to
the same cause. I cannot offer at present a satisfactory explanation
to account for this phenomenon; but, as Judd says, it might in
part have been caused by a slow but constant movement of a
crystallizing magma, and also chilled suddenly, perhaps by the
access of water at the final stage of consolidation. I may here
adduce in support of my ground a fact of the special distribution
of the Hypersthene-basalt which, so far as I am aquainted with,
occurs only in the outlying islets, excepting the locality Sei-kei,
on the north coast of Héko, which also lies not very far from the
present sea-shore.
Hyperthene occurs exclusively, though insignificant in quan-
tity, in the form of phenocryst (Pl. 11, Figs. 2 and 3) and takes
the place of olivine in the present rock-group. It is sedge-like
in general shape, and granular in its margin (especially in
Fig. 2), being fringed with grains of common augite, whose
presence becomes strikingly apparent between crossed nicols,
OF THE DEPENDENT ISLES OF TAIWAN. 41
on account of their vivid colours of polarisation in contrast with
the grey tint of the hypersthene in the interior. Pleochroism is
scarcely perceptible. Traces of a few rough cleavages run
through the hypersthene lengthwise, and as in the case of the
plagioclase, it is trasversed with many fisaures. The hypers-
thene is of intratelluric origin, and has the general aspect of its
having been worn out caustically and frittered, and the peri-
pheral accumulation of augite, already referred to, seems to have
some genetic relation with the act of degeneration.
Large, monoclinic augile sometimes makes its appearance in
company with the hypersthene and plagioclase, forming local
patches of secretional origin, with the Ayperitic structure.
The ground-mass, which constitutes the main bulk of
the rock, consists of laths of plagioclase and grains of the
frittered and corroded augite, together with rugged clumps
of magnetite. The relation of the first two components cannot
be told in a few words. In one instance, the mutual relation
is such that we could almost say it is ophitic; in another, it
is intersertal in company with a little remnant of brown glass,
while in the third, no such arrangement could be discovered,
but a simple aggregate of felspar and grains of augite, there-
by calling forth the structure which is termed granulitic. The
augites of both generations are of yellowish brown and not violet-
brown.
Shingly iridymite fills up polygonal spaces, and the loose
brushes or tufts of either plagioclase or apatite are thrown
through the whole mass. A doubtful iddingsite (Pl. I, Fig. 3)
was once observed, and some rocks are calcareous too. The
stratigraphic position of this type is not known to me. It may
be the lava of either the first or the second flow.
42 KOTÔ : NOTES ON THE GEOLOGY
E. THE TYPE OF THE ANALCIME-BASALTS.
(Pl. II, Fig. 5.)
This to the naked eye is macroscopically deep-grey, and fine-
granular. Under the microscope it is hypocrystalline and more or
less porphyritic, either the xenomorphic olivine or the aggregate of
the automorphic augite being the phenocryst, or sometimes both.
The texture varies within a wide range, but generally speaking is
coarse (fig. 5). The porphyritic elements, however, differ general-
ly not much in size from the crystals of the ground-mass, and
the mode of arrangement of the several components is granulitic.
Plagioclase predominates over augite in quantity ; and magnetite
is not plentiful, and completely soluble in HCl. The paucity of
iron-ore causes the rock to appear of a grey shade.
Olivine occurs as a phenocryst in the xenomorphic grains, a
few of which have been reduced even to mere flecks through gradual
resorption. Cleavages are not noticeable in contrast to other
olivines, but in stead of them there are curvilinear cracks, con-
forming approximately in their direction to the boundary of
resorption. The substance of the olivine is colourless, and usually
more or less altered into a greenish or yellowish, fibrous sub-
stance (not iddingsitic). Brown decomposition is quite foreign
to the olivine of this type. The present olivine seems to belong
to a variety rich in magnesia. Phenocrystic pyroxene is scantily
present in some, but none in others. The augite is of the typical
Basaltic variety, with a violet-brown type, possessing the hour-
glass structure, and idiomorphic, flattened on the orthopinacid.
It occurs singly or in stellar aggregate. There is no felspar-
phenocryst. |
The ground-mass consists of laths of plagioclase, crystals
OF THE DEPENDENT ISLES OF TAIWAN. 43
and crystalloids of violet-brown augite, magnetite, and xeno-
morphic olivine, with the interstitial mass of analcime and
base. The laths are multiple-twinned with the parapet-like
terminations (Pl. ZI, Fig. 5) produced by the shifting of lamelle
to the one end or the other with reference to the adjacent plate.
The slide treated with HCl shows a considerable corrosion of
the interior lamelle of the laths, while the exterior remains
intact and fresh, as if a frame is enclosing the hollow space.
The crystals of a violet-brown augite of the short prismatic habit,
rather flattened towards the ortho-axis, are freely developed, or
occur in clusters. The augite and plagioclase must have, there-
fore, crystallized simultaneously, and at their contact the one
is partially penetrating the other and vice versa. Magnetite is
idiomorphic, but frequently possesses irregular outlines, owing to
the penetration of the crystals of plagioclase, augite, and apatite,
and the larger crystals are anhedrons, as they are moulded upon
the neighbouring laths of the plagioclase. The magnetite is
comparatively large and few, excepting its dendritic skeleton
crystals which are found abundantly in the specimen from Gio-6,
in company with devitrified glass. In the specimen, which is
wanting in dendritic magnetite, there are brown, biotite-like
lamellæ usually in association with the hexahedral iron-ore. The
lamelle are anisotropic, and distinctly pleochroic, and the
mineral is conjectured to be «mente.
It is of no small interest to note the presence of analcime.
It occurs sporadically rather in large patches in the cuneiform
spaces left by other crystals. It is generally fresh and colourless,
and isotropic, but often shows the optical anomalies so common to
this mineral. At times, the analcime resolves into a dirty, fib-
rous nalrolite (as in the left, lower margin in Fig. 5). The
44 KOTO: NOTES ON THE GEOLOGY
analcime seems, so far as my experience goes, to be exclusively
confined to this type, though it is possible that the colourless
base in minute interspaces of other Basalts of the Héko Group,
might turn out to be that mineral, if the means are at hand in
ascertaining its presence.
, Another accessory to be mentioned is apatite in colourless
prisms, which is especially plentiful in this type.
The colourless base and analcime are rather unexpected
guests in the basic, black rock, such as we have here, and the
mode of occurrence is that they fill up the polygonal interspaces
left by the crystals of other components of the rock. If we
accept the primary origin of the analcime, as Pirsson” would do,
it is all the more very striking to see that the residuum of
a Basaltic magma should have an exact composition of
N2,0'Al,0,'4 SiO,2H,;0. Yet the analcime seems to all appear-
ances to be of primary origin, if we take into account the
perfectly fresh state of the rock in which it is found, and not
only in the Basalts of the Höko Group, but in the Teschenite
of the Nemuro promontory in Hokkaidö, I had several occasions
to observe the same mode of occurrence of the analcime, so that
it could not be attributable to a mere accidental circumstance
to find it in such state, as several foreign writers also noticed
the same. It excludes the idea of its having replaced the base
which formerly occupied the place of the now-existing analcime.
The present mode of occurrence of the analeime may perhaps
be explained by supposing that, when the Basalt was consolidating
on the surface in a quiet state, carrying in it the intratelluric oli-
vine, the newly created crystals, such as those of plagioclase, augite,
1) ‘The Monchiquite or Analcite Group of Igneous Rocks.’ Journ. Geol., Vol. IV., 1896,
p- 679.
OF THE DEPENDENT ISLES OF TAIWAN. 45
magnetite, apatite, together with the olivine had then sunk down,
and formed the heap of crystals at the bottom, meanwhile the un-
consolidated residuum of the magma was actually slowly flowing
through the sieves of crystal-heap, or changed its chemical com-
position through diffusion, after the manner of liquafion as
in a metallurgical process. And, then, the solution having the
composition of the hydrous alumino-sodium-silicate has finally
crystallised out in the interspaces of the meshes of crystals.
Similar process can be frequently observed during the formation
of crystals on the stage of the microscope. If this be the actual
condition under which the Analcime-Basalts have consolidated,
considerable leaching and percolation must have taken place
during the formation of rocks, and the structure of such a rock
should better be called the ‘leached.’ This structure is therefore
properly seen only either in the granitic or in the granulitic
rock, consequently it is wanting in the family which has
a fluxional arrangement of the components.
The Analcime-Basalts are represented in my collection by
three specimens from Gio-6, and one from Höko. The hand-
specimen from Nai-an” in Gio-ö, is, according to Y. Saitö, said to
occur at the water’s edge, the main portion of the flow usually lies
under the level of sea, and constitutes the third sheet of lavas,
and is the lowest, consequently the oldest of the accessible lava-
flows of Gio-6.
Other Analcime-Basalts of the Höko Group no doubt belong
io the same horizon.
nn nn m nn
1) Nai-an AR, MER
46 KOTÖ : NOTES ON THE GEOLOGY
THE ISLAND OF KOTO! (BOTEL-TOBAGO).
Starting from Makian”, one of the Spice Islands, a long
chain of the Moluccan volcanic system runs upwards, and joins
at the solfataric volcano of Api, in Mindanao, with that of the
Sangirs, that comes from the north end of Celebes. The united
system of volcanoes in the Philippines, then, receives the name
of the Mayon system. It goes right through the whole breadth
of Mindanao, and enters Caminguin, Leyte, Biliran, and, after-
wards, the peninsula of Camarines of south Luzon. It is in
the last-named region that the volcanic activity of the Philippines
is fully displayed. Albay or Mayon stands foremost in rank
among the mighty cones. For a while, we lose sight of the chain
northwards under the Pacific bottom, and it reappears in full force
at the crater of Cagua near Cape Engano, in north Luzon.
The northern prolongation of the Mayon system may still
be traced through the little-known Babuyans,® the Batans, and
the Bashi islands. All are said to be of volcanic origin. Among
the Bashi or Vasshi”, the five larger islands, going from the south
to the north, are Liayan, Mabudis, Tanem, Maysanga, and Tami,
the last being the largest of the forlorn isles. An active volcano
is said to exist in the southern region (?), spreading fire and
destruction.
The Balintang Canal at 20° N. lat. separates the Japanese
1) SCH.
2) B. Kotö, ‘On the Geologic Structure of the Malayan Archipelago. This Journal,
vol. XI, pages Ill and 118. Wichmann calls the chain the ‘North Moluccan bow.’ ‘ Der
Wawani auf Amboina und seine angeblichen Ausbrüche, III’ Tijdschr. r. h. Kon. Nedert.
Gen., Jaargang 1899, S. 32. This bow is now said to start from Batjan, lying to the south
of Makjan. loc. ci. 8. 14.
3) Kotö, loc, ei. p. 118.
4) The Japan Mail, August 10th 1897, ‘ Forlorn isles.’
OF THE DEPENDENT ISLES OF TAIWAN. 47
domain from that of the United States. Within the Japanese
waters lie the Batans, the Bash islands, the rocks of Gadd and
Forest Belle, the islands of Shö-Kötö (Little Botel-Tobago) and
Kötö (Botel-Tobago), and, lastly, Kashö (Samasana), as the conti-
nuation of, I conjecture, the Mayon system of volcanoes (Fig. 1).
The smaller isle of Kötö is, geologically speaking, entirely
unknown, but the Larger Kötö has been several times visited
by the Japanese, since the first landing of a staff of the gover-
norship of Taiwan, in April, 1897. Among our University
men, Mr. Tada stayed there a week collecting zoological speci-
mens, and, lately, Mr. Torii remained longer in this lonely island
among the aborigines for his anthropological study. I myself
have not had the opportunity of visiting it, though the island
has been within my sight for a week long, while travelling the
pathless beaches of south-eastern Taiwan.
The island ofthe Larger Kötö (Fig. 1) lies in a south-eastern
direction about 50 miles off the coast of Pinan, and 35 miles
north of north-east from the Cape of Galambi in Taiwan. Its
north-south extent is 3 rz and the breadth 14 ri, with the
circumference of 9 ri. It is the abode of 1,500 nude aborigines.
Seen from a distance, this scapula-shaped island appears plateau-
like in general profile, crowned by a prominence of 120 m.,
somewhat excentrically situated in the north ; and is bounded by
steep declivity all round the coast, so that it leaves only a nar-
row patch of lowland on the south-western shore, which serves
at the same time for the chief anchor-ground of this islet.
Being situated amidst the stormy and swift Kuro-shiwo cur-
rent, the narrow beach is highly cobbly, as may be seen from
Mr. Torii’s photographs; and the steep cliff undoubtedly owes
its present form to the abrading action of dashing waves,
48 KOTÖ : NOTES ON THE GEOLOGY
Fringing reefs are said to skirt the shore, some portion attain-
ing double the man’s height above the water’s edge, indicative
of a recent negative shift of the relative levels. It seems to me
probable, that they ure not the reefs of Neocene time, which
usually attain a considerable height of more than 200 m., as in the
Apes Hill of Tukao, but those of a comparatively recent date,
possibly representing a Dailuvial formation. The plateau-like
elevation, which faces the sea in cliffs, seems in parts at least
in the north-east point to consist of volcanic agglomerate. A
greater part of the interior seems to be built of volcanic rocks
with a gabbro-like plutonic mass as the foundation of the island
exposed at the west coast, but their mutual relations and area
of distribution are quite unknown to me.
In the following, I will give a succinct account of rocks,
kindly placed at my disposal by Messrs. Ishii and Torii.
A, FELSPAR-BASALTS.
(Pl. III, Figs. 3 and 4.)
My slides of Basalts and Andesites are prepared from chips
of water worn gravels, used as weights attached to a fishing net
of the aborigines.
The Basalt is rather porous, greyish-brown mass with a few
phenocrysts of a brown olivine (1-2 mm.) and black common
augite. Under the microscope, the olivine occurs in two gene-
rations (Fig. 4). Its forms are acute six-sided, sometimes
nearly square, truncated at corners, but mostly corroded and
disfigured, with a few traces of basal cleavage. The crys-
tals are slightly decomposed in their margin, being either: yellow
OF THE DEPENDENT ISLES OF TAIWAN. 49
or brown ; but as a whole the interior is fresh. It is the iron-
rich variety—hyalosiderite, as is proved by the micro-chemical re-
actions, which show only a trace of magnesia. The olivine
encloses a large quantity of regular octahedra or elongated
crystals of the brown, transparent picotite, mixed with the crystals
and dust of magnetite.
Plagioclase, as a phenocryst, is observed only once in my
three slides ; it is long-rectangular in form, with negative crys-
tals, filled with a gas. The crystal is multiple-twinned, extin-
guishing light symmetrically with the maximum angle of 32°;
consequently it is the calcium-labradorite. The augite is rather
automorphic, showing, however, a slight corrosion marginally.
This character is common to all of the specimens. The crystal
occurs in polysynthetic twins ; the colour yellowish-green and non-
pleochroic. As usual, it has glass-enclosures with air- pores.
Sometimes, the augite is internally and nucleally resorbed, leaving
an accumlation of grains of the same in its place. The augite
is of nearly the same size as the olivine.
The ground-mass is seen, under the microscope, to make the
main bulk of the rock: The micro-phenocrysts of olivine and
augite are the same in habit as the macro-phenocrysts. The
augite is in a few cases fringed with skeleton-crystals. They are
inbedded in the plexus of the felspar-laths and clumps of mag-
netite, rudely showing a flow structure. The laths are twinned
simply or polysynthetically, and in many case hollow, with the
very thin external rim, partially or entirely filled up with glass.
So far as I am aware”, such skeleton-crystals of felspar seem to
1) The same skeleton laths are observed_by E. Elich in the Amphibole-pyroxene An-
desite from the Rio Blanco, West Cordillera, Ecuador. Reiss u. Stübel, ‘Reisen in Siid-
Amerika, Das Hochgebirge der Repubik Ecuador, I.; Petrographische Untersuchungen, L
West Cordillere,’ S. 163.
50 KOTO : NOTES ON THE GEOLOGY
be of extreme rarity. The laths extinguish light symmetrically
but in the contrary direction at an angle of 26°-27°, proving
the felspar to be more acidic than its phenocryst. Interstitial
space is occupied by a brown glass which contains globulitic
and rod-shaped bodies. From the foregoing descriptions, it 18
evident that the Basalt of the Island of K6l6 does not properly
belong to the category of normal Basalt with violet augile, present-
ing the interserlal structure, which is so common in the rocks of
the Höko group, already described. Here exclusively monoclinic
augite presents the character of diopside. Both the olivine and
augite, all being equally corroded, present so great a variation
in size from the macroscopic to the microscopic dimensions that
I could not discriminate the products of the intratelluric from
the effusive period of consolidation. The ground-mass, as I have
said, is highly felspathic, and the structure is Andesitic and
hypocrystalline-porphyritic, somewhat resembling a pilotaxitic
type. Richness in olivine and paucity in iron ores, as well as
globulitically granulated mesostasis make the rock approach
to a navitic structure (Fig. 4), the only difference being the presence
of feldpar-laths in the ground-mass. Tlie rock seems to me to
be a lava-flow, consolidated rapidly, accompanied by a brisk
liberation of gas from the cooling magma. |
B. HORNBLENDE-ANDESITES.
(Pl. III, Fig. 5.)
I have three specimens of rocks in Torii’s colleetion, belong-
ing to the same category. They differ in colour consequent on
the various stages of decomposition. A fresh one is greyish and
porous, speckled with phenocrysts of hornblende (2 mm. in
length).
ON THE DEPENDENT ISLES OF TAIWAN. 51
Plagioclase is long-rectangular along the zonal axis at right-
angles to 010, and tabular when parallel to that face (Fig. 5).
It varies in size so that between the phenocrystic felspar and that
of the ground-mass we could find a series of dimensions.
Zonary structure is typically developed in almost every indivi-
dual, especially on the tabular section of 010. It contains, as
usual, glass arranged in zones; sometimes encloses crystals of
augite and hornblende, parallel to base and the positive dome;
it extinguishes light in symmetrically opposite directions with the
maximum angle of 30°-34°. The extinction observed on 010
amounts to 15° with reference to P/M, the trace of the peric-
line twins making 1.5° with P/M on the same face. These rough
observations all point to the labradorite-nature of the felspar.
Hornblende occurs only as the phenocryst and small in quantity.
It is a brownish-green variety of optically normal character.
The crystals are all corroded and enclosed by the opacitic
margin (Fig. 5) which is composed of confused aggregate of crys-
talloids and grains of monoclinic pyroxene, and clumps of mag-
netite. The pyroxene appears in tolerably large size that it could
be optically ascertained. Sometimes the substance of the margin
has been replaced by brownish, double-refractive fibres. In one
slide the body of the hornblende is impregnated with countless
swarms of black dots which lend to the crystal a darker shade.
With high powers, they resolve into glass-enclosures with bubbles.
Augite occurs sporadically as a phenocryst. Its coarse dis-
tinct cleavage, pale colour, and small angle of extinction (less
than 32°) prominently characterise this pyroxene, and contrast
pronouncedly with the brown, Andesite augite. That it is diop-
side is highly probable, but not proven. In one slide, I observed
porphyritic aggregate of needles, producing the glomeroporphy-
52 KOTO : NOTES ON THE GEOLOGY
ritic structure, and they look more like a druse than like a mass
of crystals, having a mutual relation, characteristic of plutonic
rocks. Thus our augile is remarkable in many respects.
The ground-mass consists mainly of the idiomorphic plagio-
clase, long-rectangular or square in shape, and of various sizes, with
some degree of parallel disposition. The square sections of the
microliths show occasionally truncation of corners by domal
face and at other times slightly diverge from rectangularity on
the edge 001:010. The traces of cleavage run parallel to the
same edge, and the sutures of twins run vertically. Symmetrical
extinction takes place at 30°-32° with reference to the same
trace, showing that the plagioclase stands just at the middle of
the series between the sodium and the calcium labradorite”.
According to Becker, these square sections, which are prismatic
sections in vertical positions, are very convenient for the deter-
mination of the microlithic plagioclase. Intermixed with the
felspar, we find the less idiomorphic crystals of pale augite,
together with rounded magnetite and the crystals of apatite. The
cuneiform space left by minerals being filled with the brown glass,
densely charged with transparent augite. The structure of the
rock is therefore that kind which we call the ‘ orthophyric.’ In
the ? variety, minute felspar-needles make the greater part of
the ground-mass, exhibiting the typical pilotaxitic structure.
C. APOANDESITES.
(Pi. III, Figs. 1 and 2.)
One variety is whitish, bleached and compact, the other is
green through the presence of a chloritic mineral, having a por-
1) Becker, Amer. jour. Sei., May, 1898.
OF THE DEPENDENT ISLES OF TAIWAN. 53
phyritic structure with the phenocrysts of plagioclase and horn-
blende. They are much speckled with glittering iron-pyrites
(a large black spot in ig. 1), which likely attracted the atten-
tion of Mr. Narita, who had brought back the specimens to
Tai-hoku.
The phenocrystic plagioclase has a tabular form being nearly
equidimensional. It has a distinct zonary banding, like the pre-
ceding rock. - Contrary symmetrical extinction of about 33° on
both sides of the trace of the albite twins shows the plagioclase to
be a labradorite of a similar coniposition as in the rocks, just des-
cribed. Hornblende is entirely decomposed (in the right halves
of figs. 1 and 2) into an aggregate of pistacite, chlorite, and
calcite-films, which together form the pseudomorph after the
hornblende of a prismatic habit with the combination of 010,
as may be conjectured from the original outlines of the now
altered mass. The chlorite possesses the normal character, and
pleochroic, showing a green shade parallel to the axis of fibres,
which corresponds to X. The epidote occurs in tufts and in rugged
plates.
The ground-mass consists of very fine laths, simply twin-
ned, and they are arranged in more or less parallel disposition
around the phenocrystic felspar. ‘These minute crystals of fels-
par swim within the chlorite-lamellæ, mixed with the felspar-
microlite, magnetite and the pyrites, the last does not contain
any trace of copper. This Apoandesite is no doubt derived from
the § variety of the Hornblende-Andesite, already described, by
the pneumatolytic process which caused the impregnation of the
pyrites in the rock-mass.
54 KOTO : NOTES ON THE GEOLOGY
D. AMPHIBOLIZED GABBRO.
A dark-greyish, coarse rock of gabbroitic aspect, in which
a cleavable hornblende lies after the manner of plutonics, and
a plagioclase is moulded upon the amphibole. Patches of
epidote and iron-pyrites complete the list of megascopic elements.
Under the microscope, the greenish-brown hornblende is for the
greater part altered into a nearly isotropic lamellæ of chlorite,
calcite-films, and common epidote. The hornblende has been so
highly altered that the original substance remains but in few
stripes. The plagioclase-anhedra possess only a few twinned
lamell®, besides the Carlsbad type of twins, Suitable section
could not be found for ascertaining the nature of the plagioclase.
The general deep-greyish appearance of the felspar is due to
the presence of a pennine-like chlorite in the fissure of it. Com-
mon epidote occupies the place of the felspar and hornblende
in rugged plate. Crystalloid of apatite, full of air-pores, was only
‘once observed.
I conjecture this rock to be a metagabbro, though a diallage-
like augite was never seen in my slides. This gabbroitic mass
probably makes the foundation of the island, and crops out on the
west coast, together with the Apoandesite and Serpentine.
E. SERPENTINE.
Associated with the above rock, there occurs a Serpentine
which is yellowish-blue in its general appearance. Under the
microscope, the whole mass presents between crossed nicols a
beautiful lattice-work, which is a characteristic feature of its having
being derived from an amphibole. There are found intermixed
with the Serpentine a little quantity of iron-ore.
OF THE DEPENDENT ISLES OF TAIWAN. 55
THE ISLE OF KASHÖ (SAMASANA).
(Pl. III, Fig. 6.)
Kashô is a forest-covered, conical volcanic island (Fig. 1),
only 8 km. in circumference, skirted by fringing reefs. The
inhabitants are of the mixed blood of the Chinese and the
Malays. According to Mr. Ishii, who gave me a rock-specimen,
the island is Andesitic, consisting of Pumice and lava-flows, and
carries two craters. My slide shows the rock to be the Hypers-
thene-hornblende-Andesite.
To the naked eye the rock resembles very closely those of
Héradaké, in Shinano, and Hakusan in the Kaga province. It
is greyish-looking, with the only phenocryst of hornblende,
measuring 5 mm. by 2. The hornblende is the largest of
phenocrysts (on the right half of Fig. 5), broad-columnar in
form in combination of 110 and 010, and has always thick
margin of opacite. ‘Ihe hornblende has dark-brown colour, and
optically normal. It encloses the grains of felspar after the
fashion of poikilitic plate, especially on periphery. This fact
conclusively shows the simultaneous crystallisation of the
hornblende in its later period with the forerunner of plagioclase.
The formation of these crystals might have taken place at the
close of intratelluric period of the magma. The opacitic margin
consists, as usual, of the grains of monoclinic pyroxene and
magnetite. They seem to have been formed by resorption and
re-combination through the gradual caustic action of the surround-
ing magma upon the already existing hornblende, at a slightly
lower pressure and in the upper column of effusive lava than
the situation in which the original amphibole has crystallized
out. The majority of crystals seems to have been eaten up by
56 KOTO : KOTES ON THE GEOLOGY
the magma, so that there remains nothing but the accumulation
of magnetite-dust in the place of the hornblende.
A brown pleochroic hypersthene occurs in few quantity, and
small in size and less idiomorphic when compared with the
amphibole. Its base shows no axial poles, but symmetrical
hyperbolas ; it forms penetrating twins upon a domal face, and
often moulded upon plagioclase. The plagioclase is of tabular
or long-rectangular shape; extinguishes symmetrically in the
direction at about 30° against the trace of the albite-twins, and
the trace of pericline lamellæ makes -5° to -10° on 010 with P/M,
indicating the presence of labradorite. The albite-lamalle are
clear and definite, but the width varies much from one lamella
to another, and even in the same the width varies from one
point to another,—these are also said to characterise labra-
dorite. Zonary banding is pronounced, the interior abounds in
glass-enclosures, with the clear shell of different optical orien-
tations. We meet often with the broken crystals, from which it
may be inferred that the rock is a lava-flow. The ground-mass
consists of a plexus of augite-needles in a colourless base, inter-
mixed with a somewhat larger plagioclase of a tabular, or long-
rectangular form, after the manner of a micro-phenocryst. Twin-
ned slender sections show symmetrically the opposite extinction
at an angle of 20°, indicating that the felspar in the ground-
mass is andesine in lieu of the larger, phenocrystic labradorite.
Magnetite abounds in the glassy base. Tridymile fills free spaces
in imbricated scales. ©
OF THE DEPENDENT ISLES OF TAIWAN.
CONTENTS.
The Höko Group (Pescadores)
I. Introductory ss
II. Stratigraphical Charmeterieties.
Höko Island ...
General outlines of the ‘colby of the island ...
Local details of seology
Haku-sha Island ... Bis
Kippai Island
Gio-6 Island .
III. Petrography of the Effusives
A. Component-minerals of Basalts
Olivine
Plagioclase
Augite =
Hypersthene ...
Analcime and Natrolite |
The Iron Ores
B. Special Description of Individual en of Basalts ..
a. The granual type
b. The type of the Tddingsite, arte ‘Baaalts es
c. The ophitie type Fr
? The type of the Olivine-less Basalts
e. The type of the Analcite-bearing Basalts
The Island of Kötö (Botel-Tobago)
a. Felspar-Basalts ...
b. Hornblende-Andesites
c. Apoandesites de
d. Amphibolized Gabbro
e. Serpentine ... =
The Isle of Kashö (Samasana)
eee
57
PLATE I.
(PHOTOGRAMS, )
PLATE 1.
Fig. 1.—A fine compact basalt, with comparatively large
phenocryst of olivine which is more or less iddingsitized. The
ground-mass consists of small crystals and grains of augites,
granular olivine and the laths of plagioclase, with the structure
typically granulitic. Bö-ryo-san, Haku-sha Island. P. 33.
Fig. 2.—The same rock-type as the preceding, but rather
coarse. On the right side in the figure is a augititic patch,
composed of exclusively the crystals of augite in the base. Höko
Island. P. 55.
Fig. 3.—Olivine-less basalt from lHattö, Southern Group,
and it probably belongs to the same type as Figs. 3 and 4 in
Plate IT. A doubtful olivine is present in the form of chloritic
patches, but no visible hypersthene. General mass consists of a
plexus of fine grains of augite and fine laths of plagioclase in
the base. This is quite an anomalous rock. P. 39.
Fig. 4.—Iddingsite-bearing basalt with a large idiomorphic
olivine, externally changing into iddingsite. Magnified 65 diame-
ters. Höko Island. P. 35.
Fig. 5.—Rock belonging to the same type as the preceding.
It is also from Höko Island. Olivine on the left side of the
figure shows various stages of iddingsitization.
Fig. 6.—Also iddingsite-bearing basalt, with olivines chang-
ing from the interior, as may be seen on the lower side of the
figure. Magnified 38 diameters and not 65, as is stated in the
Plate. Nicols crossed. Kippai Island.
Koto, The Höko Group, etc. ‘en % Jour. Se, Coll. Vol. XI. PI. 1.
r 4
x65 -+nicols
Kotö phot. imp. Lokyo Printing co, Lid. C
PLATE EL
(PHOTOGRAMS.)
PLATE I.
Fig. 1.—Iddingsite-bearing andesite, magnified 65 diameters,
showing the typical intersertal structure. Olivine is here changed
internally into a red mineral, which the writer believes to be
iddingsite, as is well seen on the lower right octant in the
figure (pp. 19 and 35). Kippai Island.
Fig. 2.—The slide of ophitic basalt (p. 38). Shö-chi-kaku,
the Island of Höko. |
Fig. 3.—Olivine-less hypersthene-bearing basalt, with two
large crystals of hyperthene in the centre of the figure. The
structure is granulitic. The Isle of Wam-pai. P. 39.
Fig. 4.—The same rock-type as the preceding, but with
intersertal structure. Local patches of hypersthene, augite and
plagioclase, with the hyperitic structure. Sei-kei, the Island of
Höko. Pp. 39 and 41.
Fig. 5.—Analeime-basalt from Nai-an, Gio-Ö. It has granu-
litie structure. White patches are filled with analcime, and a
dirty portion at the middle of the field is the secondary natrolite.
P. 42.
Fig. 6.—Foraminiferal rock, consisting of discoidal and spiral,
water-worn shells of (alcarına Spengleri, besides fragments of
corals, bivalves and serpula. In natural size. Kippai Island.
P. 15.
Koto, The Hoko Group, etc. Jour. Sc. Coll. Vol. XIII. PI. I.
Fig. 3. X24 Fig. 4. x88 +nicols
Fig. 6. Nat. size
Kotô phot. imp. Tokyo Printing Co., Ltd)
PLATE Ill
(PHOTOGRAMS.)
PLATE II.
The Plate III illustrates the type-rocks from the Isles of
Kötö (Botel-Tobago), and Kashö (Samasana).
Fig. 1—Apo or altered andesite, magnified 65 diameters,
showing a phenocrystic hornblende, with opacite margin (on the
right side of the figure). The hornblende is entirely decomposed
into an aggregate of pistacite, chlorite, and calcite-films. Plagio-
clase is much decomposed. A dark spot (in the lower left octant)
is the iron-pyrite (p. 52).
Fig. 2.—The same slide under crossed nicols.
Fig. 3.—<A porous, greyish-brown basalt, with a rather large
corroded olivine (on the left of the figure). The ground-mass,
which encloses a corroded diopside-like augite, is highly fels-
pathic. The structure is hypocrystalline-porphyritic, approaching
to the pilotaxitic type (p. 48).
Fig. 4—Another basalt, with abundant olivine of various
dimensions. It contains globulitically granulated mesostasis, and
the structure is navitic (p. 50).
Fig. 5.—Hornblende-andesite, with dark hornblende-crystals,
surrounded by opacitic margin (on the upper and the lower end
of the figure). The structure is orthophyric (p. 50).
Figs. 1-5 are all from the rocks of Koto.
Fig. 6.—Hypersthene-hornblende-andesite from the Isle of
Kashö, with a large phenocryst of hornblende (on the right
half of the figure). It is enclosed by a thick margin of opacite,
but enclosing the grains of plagioclase after the fashion of
poikilitic plate (p. 55).
Koto, The Hoko Group, etc. Jour, Sc. Coll. Vol. XIII, PI. I,
Fig. I. x05 Fig. 2. x88 +nicols
Fig. a x any Fig. Ö, 4 YN
eo > T
Kotö phot. imp. Tokyo Printing Co. Ltd»
PLATE IV.
(MAP,
PLATE IV.
The Plate IV shows the bathometric condition of the neigh-
bouring seas of the Pescadores or Höko Group. It seems to me
that the North Group forms itself an independent centre of
extravasation of magma, in contrast to the South or Rover Group,
from which the Northern is separated by the incurve of the
forty fathom-line,—the position indicated by the Rover Channel.
Both groups are, however, located at the north-eastern end of
the Formosa Bank, which is disconnected on the east from
Taiwan by a channel of the same name (p. 3).
Kotô The Höko Grou
SSS SS nr
—a no —
ri # y
Pid HH F
F F Z
f I = +
1 mt —_ —__"
! |
; | | 1
CET 2 = '
in nn — RER — pe Ze =
4
3 #4 he =
P- re ys.
f Fa #4, = "a
i | ren x, =
7 aq =
|
South Group } "7.
+ À | # F
\ | | N N
ie
SS] HE
= = | ;
THE HOKO GROUP.
Jour. Sc. Coll. Vol. XM. PI. IV.
p, etc. | Je
= = Le ae es an LT AUS à RER TAN SEEN | —— 772
c se 4
‘
À
L 1
|
|
r 1
P| 5 !
| 1
LT
- |
si
=}
t
;
ad
|
2
sf
Pi
=
LA = ™~ 3 u .
zi — Hokuto.
= ea os a
f f * ZA
i = f Fi \ \ \ ie *
à # | | | 1.
+
“e
CP
4
%
fille. ”
v
é
'
ah
Pl
#
or
0
c
N;
j LS \ is
F4 ! ) | Pr
De ites / ee a m
# te” 4 =
# RTE Er M . -
> 717 D Mehl, \
F # F le u «
oe
2
| Gee ‘ E
| Paflo-ö N Pr
OT » de
us # Po. 2 |; —————,
| u
j FE pe
# { Uasw-sde = ran 5
—
= 3
<= =
Rover Chantel 27
l'A a - u LL" *
| Er / 77777, Le =:
Es 7 PN =
= Si T1 JE
„Riö sh SA |e 7 \
4).
N
;
|
i
N | + / ~~
| BAR
ie
| pi Taf et) ;
. |
4
1 | a : m
a Be : Sa
| Serks cr ak ine
D 'a | — ,
D ' Tat—she nn — u
Fi | Po
4
= — | i /
;
; À i
] i
r #
|
é
|
|
|
|
|
=.
1950
Er ER Re RE RE BERN
Srale 1:680,000.
PLATE V.
(GEOLOGICAL MAP.)
PLATE J.
The Plate V is intended to show the geographical dis-
tribution of the Tertiary basalts with intercalated sedimentaries
and several Recent formations, one among the latter being the
coral-reefs which fringe the coast all round. The topographic
basis for the geological map is compiled by myself from various
sources, the data being supplied chiefly by Mr. Y. Saitö, who
also offered me assistance in colouring the geologic elements
represented on the map.
Kotö, The F
Change of Volume and of Length in Iron, Steel,
and Nickel Ovoids by Magnetization.
By
H. Nagaoka, Rigakuhakushi,
Professor of Applied Mathematics.
AND
K. Honda, Rigakushi,
Post-graduate in Physics.
With Plates VI. & VII.
1. In our former paper,” we described some effects of
magnetization on the dimensions of nickel and iron, as well as
those of hydrostatic pressure and longitudinal pull on the mag-
netization. We then showed that there is a reciprocal relation
between the two, and that the Villari effect in iron is a natural
consequence of the observed changes of dimensions. Unfortunate-
ly on that occasion the range of the magnetizing field was
limited to a few huudred C.G.S. units, so that the investigation
of the behaviour of these metals in high fields was reserved for
further experiments. In addition to this, the ferromagnetics were
not of a shape to be uniformly magnetized with the exception
of the iron ovoids. It was therefore thought desirable to repeat
1) Nagaoka and Honda, Journal of the College of Science, 9, 353, 1898; Phil. Mag.
46, 262, 1898.
58 H. NAGAOKA AND K. HONDA.
the experiment on ovoids of ferromagnetic metals, and so to
extend the investigation into still stronger fields.
2. In his well-known researches on the changes of dimensions
of iron and other metals by magnetization, Bidwell” pushed the
field strength to 1500; in the present experiment, the field
strength is greater than that of Bidwell by 700. In addition to
ordinary soft iron and steel ovoids, wolfram steel from Bohler
in Vienna was tested with a result which showed a remarkable
difference from ordinary steel as regards the change of dimen-
sions wrought by magnetization. As was generally supposed, the
change of volume is very small in iron and nickel in weak
fields, but with strong magnetizing force the effect becomes
generally pronounced.
3. The apparatus already described was used in measuring
the change of length and of volume. A small alteration was
made in the arrangement of the magnetizing coil. Owing
to the strong magnetizing current, special arrangements were
made for keeping the interior of the coil at a constant tem-
perature. A double walled tube of brass was inserted in the
coil, and a constant stream of cold water was passed in the
interspace for more than an hour before each experiment. As
the resistance of the coil was only 0.562, the rise of temperature
was so small, that the ferromagnetics placed in its core were
scarcely affected. The change of length was measured by an
optical lever, as before described.” For measuring the change of
volume, the ovoid was sealed in a glass tube with a capillary neck
(internal diameter about 0.4 mm.) and so placed in the tube
1) Bidwell, Phil. Trans. 179, 205, 1889.
2) Nagaoka, Phil. Mag. [5] 37, 131, 1894; Wied. Ann. 53, 487, 1894; Nagaoka and
Honda loc. cit.
CHANGE OF VOLUME AND OF LENGTH. 59
that it rested in the axial line, and never came in contact with
the wall of the tube. The magnetizing coil and the tube were
placed in a horizontal position. The motion of the meniscus
was measured by a microscope provided with a micrometer ocular.
For more minutely detailed particulars, we must refer the reader
to the former paper.
3. The following are the dimensions of ovoids used in the
present experiments :
|
en Metal a (cm.)| c (cm.) ae,
Nickel 0.750 | 12.50 | 31.50
i 0.500 10.00 | 10.48
Soft iron | 0.750 12.50 | 31.45
3 0.500 | 10.00 | 10.53
Ordinary steel} 0.750 | 12.50 | 31.60
” 0.500 | 10.00 | 10.57
Wolfram steel | 0.750 | 12.50 | 31.82
re 0.500 | 10.00 | 10.53
1
2
3
4
5
6
7
8
a gives the semi-minor axis, c the semi-major axis, v the volume,
p the density, and N the demagnetizing factor of the ovoids. The
volume of each specimen was measured by weighing the ovoids
in water,
The elastic constants of the metals were measured by flexure
and torsion experiments on rectangular prisms made from the
same gpecimens as the ovoids. The prisms were 14.6 cm. long
and 0.896 cm. square in cross-section.
60 H. NAGAOKA AND K. HONDA.
E (C.GS.) | X (0.G.8.)
Nickel | 2.07x10% | 0,771 x10" | 1.082
Soft iron | 2.10x10% | 0.800x10" ' 0.844
Steel | 204x108 | 0.838x10" | 0.384
Wolfram steel | 2.02 x 10% | 0.849 x 10" 0.306
E gives Young’s modulus, À (=n. Thomson and Tait) the
modulus of rigidity, and @ a constant defined by the equation
+ (4330)="
The magnetization of each of these ferromagnetics was deter-
mined by the magnetometric method, after the ovoids had been
carefully annealed, with the following results :
Nickel (2) Soft iron (4) | Steel (6) Wolfram steel (8)
HT | a IH I | H T
07 | 242 | 10] 62 | 1.9 | 3| 27] 18
14 | 49.8 2.51 160 | 44 6.8 | 65
30 | 1386 | 43) 291 | Go) 183] 126] 193
54 | 2380 9.5, 587 : 97| 279| 202] 498
10.9 | 3368 | 12.7, 750 13.1 m 25.8 | 748
37.8 | 395.7 | 199] 948 ge 651 44.5) 992
74.1 | 420.0 37.2, 1111 30.3! 815! 836! 1116
1253 | 4345 |. 9961 1255 | 502] 984! 1180] 1170
171.6 | 438.7 | 155.5 | 1309 | 116.3) 1196 | an 1224
240.3 | 440.7 | 270.3 | 1400 on 1260 | 3446 | 1301
4814 |, 443.4 | 493.6! 1479 345.0! 1379 | 5123 | 1348
6742 , 4445 | 584.6 | 1520 ' 520.2 ia = 1373
9140 | 4468 7928) 1546 | 873.7| 1489 | 940.3 | 1400
1233.0 447.7 , 992.6 | 1562 | 1149.8} 1512 | 1213.3 | 1423
1747.0 | 4487 ' 1585.8| 1607 | 1822.6] 1549 1674.9 | 1452
CHANGE OF VOLUME AND OF LENGTH. 61
Change of Length.
4. Iron (Fig. 1).—The change of length experienced by soft
iron is too well-known to need any description. The ovoid
elongates in weak fields till it attains a maximum, being longer
by about 3- to 4-millionths of its initial length ; it then decreases
in length and becomes shorter than in the unmagnetized state.
The contraction goes on gradually increasing, and, in the present
experiment, it does not seem to reach an asymptotic value,
even in fields of 2200 C.G.S. units, where the contraction
amounts to about 100,005" The present result agrees qualitatively
with Bidwell’s experiment, but the contraction is much greater.
The discrepancy is perhaps to be chiefly accounted for by the
difference of shape.
5. Steel (Fig. 1).—Ordinary steel behaves just like iron, the
difference being the smallness of elongation and contraction, while
the field at which the elongation vanishes lies in the stronger.
The field of maximum elongation in wolfram steel is greater
than in ordinary steel or iron, that of no-elongation in the unan-
nealed state being several times greater than in iron or ordinary
steel. Such a field lies in H=1200. When the wolfram steel
is annealed, the retraction after reaching the maximum takes
place very slowly and the characteristic as regards the field of
no elongation becomes exceedingly pronounced. From the curve
of length change, it does not appear that it will ever cut the
line of no-elongation even in intense fields.
6. The curve of elongation (in dots) plotted against the in-
tensity of magnetization is given in Fig. 1. The change of length
62 H. NAGAOKA AND K. HONDA.
at first takes place very slowly, but on reaching saturation, the
rate of decrease becomes very rapid. So far as the present ex-
periment goes, the rate does not diminish except in annealed
wolfram steel, in which we notice a slight flattening.
7. Nickel (Fig. 1).—The behaviour of annealed nickel ovoid
as regards the length change is nearly the same as that already
observed by one of us. With an increasing field, the contraction
reaches an asymptotic value, which in the present case is greater
than that obtained by Bidwell from experiments on a nickel wire.
The explanation of this discrepancy is to be sought for partly in
the difference of shape, and partly in the difference of treatment,
as will be clearly illustrated by experiments on the change of
volume. We have also reason to believe that repeated annealing
alters the elastic behaviour of ferromagnetics as regards the
strain wrought by magnetization. Plotting the curve of length
change against the intensity of magnetization, we find a slight
bend when the magnetization becomes saturated and the con-
traction approaches its asymptotic value.
Change of Volume.
8. Experiments by several physicists prove that magnetiza-
tion produces change of volume in ferromagnetics, in contradiction
to the popular belief which is based on Joule’s experiment. The
alteration of volume accompanying the magnetization of ferromag-
netics is generally very small in weak fields, but as will be seen
from the present experiment, the phenomenon becomes more marked
as the field is made stronger. As we have already remarked, the
change of volume as measured by Cantone” in an iron ovoid must
1) Cantone, Mem, della R. Accad. dei Lincei 6, 487, 1891.
CHANGE OF VOLUME AND OF LENGTH. 63
have been exceedingly minute as the magnetizing field was very
small. Dr. Knott has published several papers on the change
of internal volume of ferromagnetic tubes, showing that iron,
nickel, and cobalt are subject to the change by magnetization.
As our former result regarding the same question was somewhat
different, especially in the case of nickel, we have thought it
advisable to settle the discrepancy by fresh experiments.
9. Iron and Steel (Fig. 2).—Preliminary experiments on soft
iron and steel ovoid showed that considerable increase in the volume
change takes place as the ovoids are annealed. The increase
becomes more significant as the field is made stronger. In steel,
the effect of annealing is greater than in iron. In strong fields,
the volume change of the annealed steel ovoid is nearly twice as
great as in the unannealed state. Wolfram steel is very little
affected by annealing as regards the volume change, but the
change itself is much greater than in nickel or iron. The
motion of the capillary meniscus in the dilatometer can be easily
followed by the naked eye. The curves in Fig. 2 have been
plotted from measurements made on annealed ovoids.
10 Nickel (Fig. 2).—As specimens of nickel almost always
contain traces of iron, the change of volume will probably depend
on the chemical nature. In addition to this, the mechanical
process which the metal had to undergo before it could be brought
to a form suitable for experiment, must have substantially altered
its elastic behaviour.
The nickel rod, which we used in the former experiment,
was hammered from a nickel plate to a prism of square cross-
section. It contained 1.75 % of iron, besides traces of man-
ganese and carbon. The ovoids used in the present experiment
1) Knott, Trans, Roy. Soc. Edinb. 88, 527, 1806; 99, 457, 1808.
„am. = .
64 H. NAGAOKA AND K. HONDA.
were prepared from a thick plate, and were nearly pure nickel, the
quantity of iron present as an impurity being inmeasurably
small. As the material is likely to become homogeneous by
repeated annealing, the ovoids were carefully annealed for
about 50 hours. The ovoid was wrapped in asbestus and
placed in a thick metal tube, the interspace between the ovoid
and the wall of the tube being filled with fine charcoal powder.
The tube was then placed in charcoal fire. When the ovoid was
annealed in this way, there were some traces Of surface oxidation.
The change of volume after each annealing was examined with
the result that it became evident that the process of annealing
increases the effect. It therefore appears that the previous history
of the specimen exercises an important effect on the magnetization
and on the dimensions of ferromagnetics as affected by magneti-
zation. ‘The anomaly in the length change noticed by Bidwell
in two specimens of nickel wire is probably not the effect of
temperature, but is perhaps to be ascribed to the cause above
stated. In contradiction to our former result with a square
prism, the ovoid showed increase of volume. The amount of in-
crease was small compared with the decrease noticed in the
previous experiment. Cantone” obtained a tolerably large increase
of volume in nickel ovoids; our former result was nearly half as
large, while in the present experiment, there is a slight increase.
The discrepancy is probably due to the difference of treatment
before the specimen can be converted into a proper shape for
experimenting, and also to its chemical composition.
11. The volume change of ferromagnetics considered as a
function of the magnetizing field takes place very slowly in weak
fields ; it then increases in a more rapid ratio till it reaches the
1) Cantone, Atti della R. Accad. die Lincei, 6, (1), 257, 1891.
CHANGE OF VOLUME AND OF LENGTH. 65
‘wendepunkt ’; after that the change becomes slower, but still goes
on increasing nearly in a straight line. Up to H=2000, the
rate of change shows no tendency to decrease. With a still
stronger field, the increase of volume will probably become
more considerable.
12. In our former experiment, the range of the magnetizing
force was confined to a few hundred C.G.S. units. In the present
experiment, the increased field strength unveiled the character of
the change of the volume considered as function of the intensity
of magnetization. As will be seen from the curves (Fig. 2.) in
dotted lines, the increase in nickel and steel takes place quite
slowly before the magnetization reaches saturation. As soon as
the magnetization reaches this state, the increase becomes very
rapid, so that the branch of the curve ascends nearly parallel to
the axis of volume increase. There we find that a slight increase
in magnetization is attended with a large increase of volume.
As the rate of increase appears to be nearly constant, it would be
very interesting, if we could push the field strength still farther to
see whether the volume change ultimately attains an asymptotic
value.
The observed changes of volume and of length are exhibited
in the following table :—
1) Cantone, Atti della R. Accad. dei Lincei, 6, (1), 257, 1891. _
66 H. NAGAOKA AND K. HONDA.
Nickel (1) Soft iron (3) Steel (5) | Wolfram steel (8)
H *|H + , H * | H “À
13 0.09x10) 8 0.10x10 7 08x10) 19 0.30x10
30 0.29 11 0.52 12 OAT 42 1.52
90 0.65 18 1.56 33 1.95 93 3.03
218 0.82 167 3.12 192 3.13 216 5.01
282 0.97 443 3.85 376 4.69 442 8.04
517 1.38 691 4.58 | 586 6.22 692 11.68
877 2.06 ds. HR, Le Si 1001 16.68
1141 24 [1115 7.18 1044 10.16 1117 18.96
1547 3.24 [1342 9.47 |1376 1407 11296 22.75
1740 3.53 |1563 1145 |1646 17.20 |1704 28.82
2253 4.12 {2089 1468 |2171 2220 |2153 32.62
___ Nickel (2) | Soft iron. (4)__ | Steel (6) | Wolfram steel (8)
EEE | ee a
4 —141xi0 6 25xid] 13 31x10 18 41x10
6 — 64.0 15 19.0 19 71 | 95 124
10 —118.2 51 316 28 11 39 91.7
23.8
3.1
17.7
52.6
62.6
13.9
18.9
82.2
86.6
89.9
91.6
— 102.0
— 163.6
— 217.9
— 264.3
— 317.6
— 343.6
— 353.8
— 356.0
— 360.0
— 360.9
— 362.2
— 362.7
— 365.3
15
33
59
124
302
561
839
1145
1289
1483
1849
2322
2180
CHANGE OF VOLUME AND OF LENGTH. 67
Kirchhoff’s Constants % and *”.
13. Starting from the formulæ
_ él (4e ( 1+0 a ER H?
l =} 3 1438) 2(1+20) 2
_ du _$ je, 3k—k) Kr) H®
pac on alt er 4 ey
which give the change of volume and of length of ferromagnetic
ovoids in terms of Kirchhoff’s constants 4’ and X”, we obtain the
following expressions for these two constants :
= an
and Vash y
where p=— ie ag) o+ 4h? + 3k,
and g=— a A+ . — — (1+N)+Kk.
These constants, as BE from a change of dimensions
of ovoids, are given in the following table, and graphically drawn
in Fig. 3:
H | Nickel | Soft iron | Steel | Wolfram steel
er on a I I’ Et ; Re TE u 14 1"
5! —229100 712800 | 21900 —22610 | 1017 —1865 | 348 —1252
10' —188900 578900 | 22520 —28450 | 2840 3322| 986 —1312
20 — 71000 216700 . 13280 -164%0 4248 —4615 | 3600 —3983
30 | — 36370 111200 7302 — 8650 | 4048 —5080 | 4881 —5440
60; — 8163 34540 215 — 2292 | 1738 —1864| 1946 —2217
80 | — 6906. 20960 1207 — 1102 | 1069 —1004! 1198 —1385
100 — 4653 14120 | 759 — 550 70L — 546 | 701 — 880
120 — 3573 10260 | 500 — 255 477 — 9279 | 557 — 595
160° — 1297 3968 | 239 18 940 —- | 35 — 317
250! — 843 2553 | 55 175 69 117 | 128 — 109
300 | - 591 1799 | 3 175 33 124] ss — 66
500; — 216 633 9 130 | —1
800 — 86 259 9 70 6 |
1200; — 39 117 = 87 4
1600 ! — 92 63 = 23 1 —3
2000 | — 14 42 -3 16 | 2
68 H. NAGAOKA AND K. HONDA.
14. The curves for % and &” present the same general feature
in iron and steel. 2 increases in}flow fields; and there attaining
the maximum value, it rapidly diminishes till it becomes less than
zero ; it then reaches a minimum, after which it again gradually
increases. The exact position of the minimum is very vague; the
curve for A ultimately coincides with the axis of H. X” is at first
negative, and attaining the minimum value, goes on gradually
increasing till it becomes greater than zero, and then reaches a
maximum. With the farther increase of the field, the value of 7”
decreases very slowly. The position of maximum for i’ and that of
minimum for %” lie nearly in the same field, which is greater for
wolfram steel than for soft iron, while that for ordinary steel oc-
cupies an intermediate position. The absolute value of X and 2”
is greater in iron than in steel. In nickel, the values of X’ and 7”
are far greater than those for iron and steel, and moreover are of
opposite signs. The maximum of k”, or the minimum of k’, seems
to lie in a weak field; the rate of decrease or increase is quite
rapid and the curves for % and %” soon approach the axis of #.
Compared with the results of former experiments, the absolute
values of X and &” are generally small for iron,—far greater for
nickel. This difference arises from the fact that for iron, the
change of length in weak fields is less in this case than in the
former experiment, and that for nickel the contrary is the case.
As regards the sign, these two experiments show fair agreement.
Consequences of the theory.
15. Effect of longitudinal pull—The change of magnetization
produced by the elongation of a wire can be easily calculated
from the formula
CHANGE OF VOLUME AND OF LENGTH.
a= H)
E
7
k
(# ty.
69
Putting 4=4.67 x 1076, 4.80 x 10-*, and 4.85 x 10° for soft iron, ordin-
ary steel, and wolfram steel respectively, each corresponding to a
pull of 0.1 Kilog. per sq. mm., we get the following results:
ber
|
800
Soft iron.
ol
0.919
1.074
0.831
0.399
0.244
0.127
0.039
| — 0.080
| — 0.164
— 0.258
—0.311
— 0.275
— 0.219
— 0.183
— 0.157
ho re SE er nn, ety eV ae | en
Steel.
ol
0.055
0.212
0.402
0.254
0.153
0.072
0.005
— 0.092
— 0.146
— 0.210
— 0.236
— 0.209
— 0.163
— 0.134
— 0.115
Wolfram steel.
— 0.028
él
(1.044
0.170
0.350
0.294
0.249
0.188
0.145
0.094
0.061
0.017
— 0.002
—0.038
—0.037
— 0.033
It will be seen from the above table that there is an increase
of magnetization in low fields, till it reaches a maximum, after
which it gradually decreases.
The decrease does not proceed
continuously, but reaches a maximum, whence the magnetization
begins to recover. Although the former result here arrived at
is the well-known Villari effect, we do not know whether the
maximum decrease due -to longitudinal stress has as yet been
experimentally ascertained. With nickel, we obtain the following
values for the change of magnetization due to elongation, 4=
70 H. NAGAOKA AND K. HONDA.
4,74 x 107%, which corresponds to a pull of 0.1 Kilog. per sq. mm.:
There is nothing remarkable in nickel. Longitudinal pull
produces decrease of magnetization, which becomes gradually less
as the field strength is increased. This is such a well established
experimental fact that we need not enter into further discussion
of the subject.
16. Effect of hydrostatic pressure. —We can easily see that
the change of magnetization ¢J due to change of volume o by
hydrostatic pressure is given by
I= -( Ps. 4, )He
If we calculate the change of magnetization due to contrac-
tions 4.68x10°, 5,88x10°%, 8,42 x 107% and 9,33 x 10 for nickel,
soft iron, steel, and wolfram steel respectively, each corresponding
to a pressure of 10 atm., we obtain the following values:
CHANGE OF VOLUME AND OF LENGTH. 71
| Nickel Soft iron | Steel Wolfram steel
aI Er | ôT | ar
0.000 0.000 0.000 0.000
0.190 0.757 0.230 0.045
0.119 0.840 0.457: 0.237
0.080 0.713 0.595 0.858
0.037 0.398 0.565 0.675
0.030 0.362 0.495 0.550
0.024 0.305 0.437 0.467
0.012 0.178 0.268 0,259
C.008 0.135 0.200 0.185
0.005 0.093 0.134 0.118
0.002 0.062 0.088 0.073
0.001 0.042 0.061 0.048
0.000 0.031 0.043 0.034
0.000 0.024 0.034 0.026
It thus appears that in nickel the effect of hydrostatic pressure
is very small compared to that of longitudinal pull. There
is increase of magnetization with the volume contraction of
the magnet. Such an increase reaches a maximum in low
fields, whence the effect gradually diminishes. Similar changes
are also noticed in the case of iron and steel. In our former
experiment, we found that hydrostatic pressure increases the
magnetization in nickel, while it decreases it in iron. The
agreement between theory and experiment is very close in nickel,
but there is a wide discrepancy in iron and steel, as we have
already noticed.
17. Effect of torsion on longitudinally or circularly magnetiz-
ed wire, There are other important consequences to be drawn
from the constant k&” with regard to the effect of torsion on
12 =H. NAGAOKA AND K. HONDA.
longitudinally magnetized wire and on ferromagnetic wire travers-
ed by an electric current. The strain caused by twisting a
circular wire can be resolved in elongation and contraction in
directions perpendicular to each other and inclined to the axis
of the wire at 45°.. Taking these two principal axes of the strain
for those of x and y, we have for the strain.
Ou
cia SU
dv
dw
age
where « denotes the amount of torsion and r the distance from
the axis. Resolving the magnetizing force which is in the direc-
tion of the axis of the cylinder, along the axis of elongation and
of contraction, we find that the circular magnetization which will
be called into play is equal to —4wrk’H at a distance r from
the axis, the mean circular magnetization being —wk’HR, where
R is the radius of the wire.
The transient current which will be thus induced in the wire
by suddenly twisting it is proportional to —k” Z.
Next suppose that the wire is traversed by an electric current
of intensity ©. Then the circular magnetizing force at a distance
r from the axis is
Hoe
By applying similar reasoning, we find that the mean longitudi-
nal magnetization is equal to —wik”C. We therefore conclude
that twisting the wire carrying the electric current gives rise to
longitudinal magnetization proportional to —k’C. Thus the
circular magnetization produced by twisting a longitudinally
CHANGE OF VOLUME AND OF LENGTH. 13
magnetized wire has a reciprocal relation to the longitudinal
magnetization caused by twisting a circularly magnetized wire.”
The view propounded by Prof. Ewing’ to account for the
existence of transient current by means of zolotropie suscepti-
bility is similar to what would follow from Kirchhofi’s theory,
but it fails to give the amount of the current or of the magnetiza-
tion which would be produced by twisting.
The theoretical inferences which we can draw at a glance
from the curves of —ik” H (Fig. 3) are as follows:
1. The transient current as well as the longitudinal magneti-
zation produced by twisting an iron or steel wire is
opposite to that produced by twisting one of nickel, up
to moderate fields.
2. The transient current as well as the longitudinal magneti-
zation produced by twisting an iron, steel, or nickel wire
reaches a maximum in low fields.
3. In strong fields the direction of the current as well as
the longitudinal magnetization is the same in iron, steel,
and nickel.
It has been established by G. Wiedemann® that the longi-
tudinal magnetization produced by twisting an iron wire carry-
ing an electric current is opposite to that produced in a nickel
one. The opposite character of the transient current in these two
metals has also been observed by Zehnder” and independently by
one of us”. The existence of a maximum transient current in
1) Voigt, Kompendium der theoretischen Physik, 2, 203, 1896, Leipzig; Drude, Wied. Ann.
63, 8, 1897.
2) Ewing, Proc. Roy. Soc. 36, 1884.
3) Wiedemann, Eletrietät, 3.
4) Zehnder,\Wied. Ann., 38, 68, 1889.
5) Nagaoka, Phil. Mag. [5] 29, 123,1890 ; Journal of the College of Science, Tokyo, 3,
335, 1890. |
74 H. NAGAOKA AND K. HONDA.
these two metals has been clearly established, although there
is some difference in the field strength between iron and
nickel. It appears from the experiments of Dr. Knott’ that the
area of the hysteresis curve in the longitudinal magnetiza-
tion produced by twisting circularly magnetized wire reaches
a maximum as the field strength is increased; but on account
of the feebleness of the current, the existence of the maximum
in the longitudinal magnetization is not well established. To
judge from the course of the curve given by the same experi-
menter, it seems highly probable that the maximum would be
reached if we could push the circularly magnetizing force a little
further. The conclusion (3) is still an open question, although
some experiments of Matteucci”) seem to corroborate the view
just stated.”
18. Looking at the curves of 1” 77, we cannot but be struck
with the close resemblance of the curves representing the amount
of torsion produced by the combined action of the circular and
the longitudinal magnetizing forces on a ferromagnetic wire. We
can no doubt co-ordinate the effect of torsion on a magnetized
wire with the Wiedemann effect. The discussion of the last men-
tioned effect we hope to lay before the public in the near future.
In spite of the qualitative explanations which Kirchhoff’s
theory affords with regard to the effect of longitudinal pull, of
the hydrostatic pressure, and of torsion, there are instances in
which the theory apparently fails in several quantitative details
that it necessarily calls for modification. We may remark that #
1) Knott, Trans. Roy. Soc. Edinb., 36, 485, 1891.
2) Matteucci, Annales de Chimie et de Physique, 1858.
5) While this paper was passing through the press, we found that the direction of the
transient current produced by twisting a magnetized iron wire is reversed in strong magneti-
zing fields,
CHANGE OF VOLUME AND OF LENGTH, io
and i” are physically functions of the strain, as is borne out by
the numerous experiments on the effect of stress on magnetization,
The present state of the theory of magnetostriction may perhaps
be compared with that stage in the history of the theory of
magnetization when the intensity of magnetization was supposed
to be simply proportional to the magnetizing force. In fact, the
theory is still in its infancy, so that there are ample grounds
for expecting further developments on further researches.
re
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Combined Effect of Longitudinal and Circular
Magnetizations on the Dimensions of
Iron, Steel and Nickel Tubes.
By
K. Honda, Rigakushi,
Post-graduate in Physics.
With Plates VIII. and IX.
1. The change of length in the direction of magnetization has
been made a subject of investigation by several experimentalists,
but few of them have measured the change in the direction per-
pendicular to that of magnetization. Joule” first observed the
diminution of length of an iron gas-piping by passing a current
through an insulated wire inserted into it, and bent over the
sides, so as to form a circular magnetizing coil of 12 convolutions.
His experiment was modified by Bidwell? who measured the
change of dimensions in an iron ring. He found that the ring
becomes thicker in a strong field and thinner in a weak one.
From the measurement of the internal as well as tle external
change of volume for iron, steel, nickel, and cobalt tubes, Knott*)
2) Bidwell, Proc. Roy. Soc. 56, 94, 1895.
3) Knott, Trans. Roy. Soc. 39, 457, 1898.
78 K. HONDA.
calculated the change of lateral dimension in these tubes. The
result for iron coincided qualitatively with that of Bidwell.
The first experiment on the change of length of an iron
wire by the combined action of longitudinal and circular mag-
netizations was made by Beatson” who observed the diminution
of length at the moment when an electric current was passed
through a magnetized wire. A similar result was afterward
obtained by Righi.” The same experiment was also repeated by
Bidwell,” who observed a large increase in the change of length
by longitudinal magnetization of an iron wire carrying a current.
2. Through the kindness of Prof. Nagaoka, his apparatus’
for the measurement of the minute change of length was placed at
my disposal. The apparatus consists of a small optical lever with
an arrangement for temperature compensation on the same prin-
ciple as the gridiron pendulum. The rod, by which the change
of length is made sensible to the lever, was slightly modified.
In the annexed figure, 7 is the tube to be tested, F'and F”
are two circular brass rings
protruding from the tube
at a distance of 1 cm. from
the ends, and soldered to a
brass rod passing through
the axis of the tube. The magnetizing coil was wound round
the tube parallel to its length extending from F’ to F” to envelope
it completely, and so arranged that the tube could slide in the
coil with little friction. Fin the lower part of the figure shows
1) Beatson, Archives des. Sc. phys, et nat. 2, 113, 1846.
2) Righi, Mem. di Bologna 4, 1, 1879; Beibl. 4, 802.
3) Bidwell, Proc. Roy. Soc. 51, 495, 1892; Beibl. 17, 582.
4) Nagaoka, Phil. Mag. 27, 131, 1894; Wied. Ann. 53, 487, 1894.
CHANGE OF DIMENSIONS BY MAGNETIZATION. 19
the front view of these rings. AR and X’ were two rods in con-
tact with the ends of the tube. The ends of these rods were bent
upwards and so filed down, that they could easily slide between
two parallel wires of the coil, which were specially fixed at a dis-
tance of 1 mm. from each other. The rod r served to communicate
the motion to the prism P. The other parts of the apparatus
remained unchanged. The apparatus was put into a magnetizing -
coil, 30 cm. long and wound in 12 layers with copper wire of
2 mm. diameter. The field at the centre of the coil due to a
current of one ampere was 37.97 C.G.S. units. The current
through the outer coil produced the change of length by longi-
tudinal magnetization and that through the inner coil gave rise
to the change of length by circular magnetization. |
To study the effect of temperature on the change of length,
the circular magnetizing coil was wound, not by a single wire,
but by double wires; thus connecting the four ends of these
Wires to a reversing key as shown in
the figure, the circular field can be
made or annulled by turning the key
one way or the other. The total
number of turns of the circular mag-
netizing coil was 44 for the nickel
tube, 40 for the wolfram steel tube and 36 for the soft iron tube.
The magnetizing currents were measured by Thomson graded
galvanometers which were compared with a deciampere balance
before each experiment.
3. The samples used in the present experiment had the
following dimensions :
80 K. HONDA.
length, external internal demagnetizing
mater (cm.) diam. (cm.) | diam. (cm.) factor. —
nickel 17.02 1.328 1.252 0.0261
wolfram steel 20.30 1.124 1.048 0.0162
soft iron 16.97 0.966 | 0.842 0.0308
The tubes of nickel and soft iron are the same as that used
in the study of the mutual influence between longitudinal and
circular magnetizations. It was found by analysis that the
nickel was nearly chemically pure, the trace of impurity being
inmeasurably small.
Results of Experiments.
1. Nickez TuBE.
4. The tube was carefully annealed, before the circular
magnetizing coil was wound round it. The change of length due
to longitudinal field alone was then measured in the usual way.
The results were compared with that obtained after the circular
magnetizing coil was wound round the tube. The comparison
showed that there was in general small difference between these
two, and that the change of length in the former case was
always greater than that in the latter, the difference amounting
to nearly 2 or 3 %. This is evidently due to the resistance to
contraction experienced by the tube, although it can easily
slide along the coil. Whether the apparatus executed its func-
tion correctly or not was tested before each experiment by
making a longitudinal field and comparing the deflection so
obtained with that in the free state; otherwise serious mistakes
would sometimes have arisen.
CHANGE OF DIMENSIONS BY MAGNETIZATION. 81
5. The experiments on the change of length by circular
magnetization, namely, on the change of dimension in a direction
perpendicular to the magnetic field, were conducted in the
following manner. The tube was first demagnetized and a
circular magnetizing current was made only for a moment and
the corresponding deflection read. The change of length due
to magnetization followed almost instantaneously, but the change
due to the heating of the coil became sensible somewhat later ;
hence these two effects were unmistakably distinguishable so
long as the magnetizing current was not strong. On this
account the highest field did not exceed 100 C.G.S. units.
The effect of the longitudinal field on the change of length
by circular magnetization was also measured. A constant longi-
tudinal field was first made and the corresponding deflection
observed ; then currents of different strength were momentarily
passed through the circular magnetizing coil, and the additional
deflection was read. These results are given in the following
table and also in Fig. 1:
TABLE I.
H=22.1 H= 182.9
ax 107 h À x 107
0.5 8.9 1.6
18.0 13.6 2.2
76.2 21.3 4.4
124.9 31.5 92
157.8 49.0 27.2
190.4 642 59.9
201.3 69.5 81.6
217.7 87.6 130.6
39 K. HONDA.
Here H and h denote effective longitudinal and circular
fields respectively, both in C.G.S. units. z represents addition-
al change of length by circular field. All these changes were
measured at a constant temperature of about 25° C.
Fig. 1 shows that the change of length by circular mag-
netization increases at first showly and then rapidly. With the
further increase of the circular field, the rate of increase becomes
gradually less. This result agrees in quality with Knott's
calculation. The circular magnetization combined with a constant
longitudinal one is always to increase the length which is first
shortened by the longitudinal magnetization. In weak circular
fields, the curve of the change of length with a constant
longitudinal field lies below the curve with no such field;
but in strong fields, the first curve lies above the second. The
point of intersection of these two curves is displaced into a higher
field with the increase of the longitudinal.
6. We shall next pass on to the change of length by longi-
tudinal magnetization with a constant circular field. The tube
was first demagnetized by reversals, and then the deflections for
longitudinal magnetizing currents of different strength were
measured. . During the experiment, the temperature at the centre
of the magnetizing coil was 18.8° C. The tube was then care-
fully demagnetized both as regards the longitudinal and cir-
cular magnetizations. Then a constant current was passed through
the circularly magnetizing coil so that the field strength became
null. Owing to the heating of the coil, the tube rapidly ex-
panded at first, but usually after an hour or two, it reached a
stationary state; when that state was reached, the measurement of
the change of length by longitudinal field alone was commenced,
which gave the length change at a higher temperature. After
CHANGE OF DIMENSIONS BY MAGNETIZATION. 83
the observation was finished, the key was reversed, so that the
circular maguetizing current was then called into play. During
this process, no gradual displacement was observed, showing that
the temperature of the tube remained unchanged during the re-
versal, but at the same time an instantaneous deflection was
noticed, which showed the change of length by circular mag-
netization. By reading the displaced position of the line in the
micrometer ocular, the deflection corresponding to the longitudinal
magnetization was noted. The tube was then demagnetized as
regards the longitudinal magnetization, the circular magnetization
remaining constant. ‘The same process was repeated for stronger
fields, till a set of observations was completed.
7. How the rise of temperature affects the change of length
by magnetization will be seen from Fig. 2. The change of
length at ordinary temperature is somewhat less than that which
Prof. Nagaoka and myself” have obtained for an ovoid made of
the same specimen. The difference may perhaps be explained
by that of annealing and of the geometrical shape of these
samples. The temperature was measured by inserting a mercury
thermometer inside the tube. Its effect is thus tolerably large;
the rise of temperature is attended with an increase of the
change of length in weak fields, and is accompanied with a
decrease in strong fields. From the same figure, we obtain the
relation of temperature to the change of length at a constant
field as shown in Fig. 3. It is well known that the magnetization
of nickel increases with temperature in low fields and decreases
in strong ones ; but under the temperature of 100° C, the change
of magnetization is too small to account for the change of length.
1) Nagaoka and Honda, Preceding paper.
84 K. HONDA.
So far as I am aware, Barrett” is the only physicist who has
investigated the effect of temperature on the change of length;
his experiment resulted in the decrease of about one-fourth of
the change of length by a rise of temperature by about 50° C.
Perhaps his field was too strong to cause an increase.
8. The results of the change of length by longitudinal
magnetization with a constant circular field are given in the
following table and in Fig. 4. The change of length was re-
duced to the temperature of 18.8° C by using the results above
obtained.
TABLE Il.
505.0
720.2 À O0 —3445 | 725.2
1) Barrett, Phil. Mag. [4] 47, 51, 1874; Nature 26, 515 586, 1882; Beibl. 7, 201.
CHANGE OF DIMENSIONS BY MAGNETIZATION. 85
The comparison of Figs. 1 and 4 shows that the general
character of the change of length is the same in these two
eases, except that the sign of the change is opposite. Hence
similar remarks as in the former hold good in the present
case.
In the experiment with nickel and cobalt wires traversed by
an electric current, Bidwell found that the effect was inmeasur-
ably small. The discrepancy in nickel perhaps arises from the
effect of temperature, which he did not take into account; the
difference in the method adopted in the present experiment for
obtaining a circular field and in that of Bidwell does not seem to
play an important part in accounting for the said discrepancy.
According to the present experiment, the rise of temperature
occasioned a comparatively large diminution of the length change
in strong fields. Hence it can not be denied that in Bidwell’s ex-
periment, the effect of circular magnetization was just as great as
that of temperature. The same remark will perhaps apply to his
experiment with cobalt; but having no cobalt tube at my dis-
posal, the experimental verification must be postponed till some
future date. However, a theoretical deduction in fayor of the
view above stated will be given in the last part of the paper.
It would not be out of place to remark that a klinging
note of the nickel tube was heard at the make and break of
cireular magnetizing current, a well known phenomenon. Even
with such a weak current as we obtain from a single Daniell’s
cell was sufficient to produce a distinctly audible sound.
9. It will sometimes happen that it is convenient to have
a simple expression for the change of length. For nickel, the
change of length is very well given by an empirical formula
of the form
Digitized by Go IQ aaa
oO uw
86 K. HONDA.
a_i”
l 1+8H" 3
where a, 3 and n are constants and # is assumed to be positive.
The determination of these constants from the experimental
curve gave the following results :
a=5.18, B—0.0164 and n=1.017.
In the calculation, only the fields H=20, 80, 320 were
chosen to simplify the calculation. Using these values of the
constants, the change of length due to fields of different strength
was calculated and compared with the experimental value as
shown in the following table:
TABLE Ill.
3 (cal.)
— 46x10
— 81
— 108
— 148
— 185
—215
— 230
— 247
— 258
— 269
— 278
— 285
— 292
— 293
CHANGE OF DIMENSIONS BY MAGNETIZATION. 87
Thus except in weak fields, the coincidence between these
two is very close; the difference does not amount to 1%.
This formula applies, not only for the change of length, but
also for every curve which has only one inflexion point and
becomes asymptotic when one of the co-ordinates increases
indefinitely, such as the curve of magnetization.
2. WoLFRAM STEEL TUBE.
10. The method of procedure with the steel tube was exact-
ly the same as in the corresponding case of the nickel tube.
The result of the change of length by circular magnetization,
e.g., the dilatation in a direction perpendicular to the field, as
well as the effect of longitudinal field on the change of length
by circular magnetization are given in the following table and
graphically shown in Fig. 5. These observations were taken
at a constant temperature of about 17° C.
TABLE IV.
H=0 H=15.1 H=31.8 H=81.3
h * x 107 h + x 107 h x 10° h 42x10
14.0 — 0.0 13.2 — 0.4 13.6 — 0.4 12.9 — 0.4
20.8 — 8.6 31.7 —12.9 30.9 — 21 31.2 — 0.9
35.2 —20.2 41.9 —32.2 41.9 —14.6 416 — 5.2
31.3 — 22.8 51.7 —40.4 51.1 —25.8 51.1 —12.9
65.1 —26 6 63.7 —45.1 63.7 —38.7 63.7 —20.8
78.7 —27.9 74.7 —47.3 14.7 —48.1 15.6 —30.9
99.5 —27.9 88.6 —49.4 91.4 —58.0 88.6 —40.0
88 K. HONDA.
Here we observe that circular magnetization produces con-
traction which increases very slowly at first, but afterwards
quite rapidly, till it reaches a nearly constant value. The
existence of the field of maximum contraction is still a question.
The result is somewhat discordant as compared with that of Bid-
well with an iron ring, in which case the diminution vanishes
in a field of about 86 C. G. S. units. Since the behaviour of wol-
fram steel as regards the change of dimensions by magnetization is
very different from that of soft iron, the cause of the discrepancy
is probably to be sought for in the difference of the specimens.
That the effect of longitudinal field on the change of length
by circular magnetization is of the same nature as in the case
of nickel, except that the sign of the change is opposite, is also
apparent from the same figure. As we have remarked, Beatson
and Righi observed the same phenomenon.
11. The middle curve in Fig. 6 represents the change of
length by longitudinal magnetization at the temperature of the
room. The lower curve was obtained at 80.2° C, and the upper
curve at the same temperature by reversing the key so as to
produce circular field. From the figure, we see that the be-
haviour of wolfram steel as regards the change of length is
widely different from that of other sorts of iron. It is remark-
able that the length of the tube, after reaching the maximum
elongation, diminishes very slowly as the field is increased, a
fact already noticed by the experiment’ referred to. In that
case, the maximum elongation was somewhat less than in the
present experiment. The discordance between the two is pro-
bably due to the difference of annealing and also of the shape
of the specimens. |
1) Nagaoka and Honda, loc. cit.
CHANGE OF DIMENSIONS BY MAGNETIZATION. 89
effect of temperature is to decrease the change of length;
inution increases with the field, till it reaches a maxi-
nd then decreases very slowly. Barrett” did not find the
the case of iron and cobalt. The upper curve shows
influence of circular magnetization on the change of
s large for steel.
The effect of circular field on the change of length by
inal magnetization is shown in the following table and
=
’. The results are reduced to the temperature of
TABLE V.
us the longitudinal magnetization combined with a constant
one is always to increase the length which is first
cit.
90 K. HONDA. |
shortened by the circular magnetization. In weak longitudinal
fields, the curve of the change of length with a constant circular
field lies slightly below the curve with no circular field; but
in strong fields, the first curve lies markedly above the second.
The point of intersection of these two curves shifts into a high
field as the circular field is increased. The field of the maxi-
mum elongation seems to increase with the circular field.
3. Sort Iron TvuBE.
13. The experiments of the change of length by circulai
magnetization and of the effect of longitudinal field on th
change of length led to the following results, which are graphi.
cally shown in Fig. 8. The observations were taken at the
temperature of 18° C.
TABLE VI.
H=0 H=5.7 H=25.8
h = x 107 h Le x 10° h &x10 h = 10!
53 — 78 5.3 — 42 5.3 — 0.0 5.3 — 0.5
140 —13.0 13.8 —11.9 140 — 52 13.8 — 10
21.4 —15.6 20.7 —16.6 21.4 —10.4 210 — 42
37.9 —15.6 30.7 —20.8 37.3 —20.8 37.3 — 9.9
53.2 —14.5 51.8 —22.3 53.3 —26.0 53.2 —14.0
69.4 —12.5 66.6 —22.3 69.2 —28.0 67.8 —18.2
81.6 — 9.3 81.3 —20.8 81.6 —26.0 80.5 —20.8
98.8 — 7.8 97.7 —19.7 | 98.0 —23.4 98.1 —20.8
By circular magnetization, the length of the tube diminishes
rapidly at first, till it reaches a minimum, then it gradually
CHANGE OF DIMENSIONS BY MAGNETIZATION. 91
recovers. The. field at which the tube returns to its former
length is not yet reached so far as the present experiment
extends. The result agrees qualitatively with that of Bidwell and
the calculation of Knott.
The general form of the curve does not change by the
application of a constant longitudinal field, but the field of maxi-
mum contraction shifts into high field as the longitudinal field
increases. The amount of the maximum contraction increases
mith the longitudinal field, till it reaches a maximum, and then
t gradually decreases. In weak circular fields, the change of
ength diminishes with the increase of the longitudinal.
14. As in the case of wolfram steel, three curves in dotted
ines are given in Fig. 9, two of which correspond to the change
flength at the temperatures of 18.7°C and 76.1° respectively.
Vhen the key in the circuit of the circularly magnetizing coil
as reversed so as to produce a field, the change of length
orresponding to the third curve was obtained.
The change of length by longitudinal magnetization at ordi-
ary temperature is somewhat less than those obtained by previ-
us experimenters. The difference is probably to be ascribed to
he well annealed state’ of the tube; also, the resistance to the
longation experienced by the tube due to the friction of the
ircular magnetizing coil was found to affect the result slightly,
‘he general feature of the change of length is so well known
nat farther remarks are unnecessary. It is only to be noticed
hat here the field of the maximum elongation is greater by 20
»G.S. units than that of the minimum contraction due to
ircular magnetization. |
The rise of temperature is to diminish the change of length
1) Bidwell, Phil. Mag. 55, 228, 1894,
92 K. HONDA.
in weak fields and to increase it in strong ones. The field at
which the temperature produces no effect is about 52 C. G.S. units.
In the case of wolfram steel, this field, if it exists, seems to be
pushed to an intensely strong field. We also observe that the
effect of circular field on the change of length by longitudinal
magnetization is tolerably large, as observed by Bidwell.
15. The results of the experiments on the change of length
by longitudinal magnetization with a constant circular field are
summed up in the following table and graphically shown in
Fig. 10, these results being reduced to 18.7° C.
TABLE VIL.
Thus the nature of the change of length is the same as in
the reciprocal case already mentioned, except that the sign of
the change is opposite. As shown in the figure, in strong fields,
the curve corresponding to the change of length with a constant
circular field lies always above that with no circular field.
CHANGE OF DIMENSIONS BY MAGNETIZATION. 93
In weak fields, the curves nearly coincide with each other. The
field of maximum elongation slightly increases with the circular,
and the amount of the elongation, after reaching a maximum, begins
to decrease with further increase of the circular field. Though
Bidwell did not observe this point, the present experiment agrees
quite well with his result.
16. So far the experiments made on the tubes of nickel,
steel and iron show that the effect of circular field on the change
of length by longitudinal magnetization is of the same nature
as the effect of longitudinal field on the change of length by
circular magnetization.
From the results of the change of length by longitudinal
and circular magnetizations, the change of volume by magne-
tization can easily be calculated, provided we assume the material
to be isotropic, as was already done by Bidwell. If « and v
represent these two dilatations respectively, the volume change o
is given by the formula c=u+2v.
Assuming the isotropy of our specimens, we find the :
calculation leads to the following results:
TABLE VIII.
Nickel Wolfram steel! Soft iron
ôt Sy ov
: |
~18.5x10
~21.0
0.0
18.0
46.5
54.0
ze
94 K. HONDA.
We thus obtain incredibly large values for the change of volume.
In nickel and soft iron, there is at first decrease of volume,
and then follows an increase; in wolfram steel, the diminution
of volume reaches a maximum and then gradually decreases.
The above result for soft iron agrees fairly well with that of
Bidwell” for unannealed iron ring. But in the experiment with
ovoids? made of the same specimens, there was always small in-
crease of volume for nickel, steel and soft iron. The amount of the
change at the field of 100 C.G.S. units was 0.7 x 107”, 3.1x 107
and 2.8 x 10’ for these metals respectively. Hence the question
now arises whether the change of volume is so influenced by the
shape of these metals. To settle this point, fresh experiments on
the change of volume were undertaken with a dilatometer. The
answer was in the negative, the result being in rough agree-
ment with that for the ovoids. The initial decrease of volume
was never observed, but the volume always increased with the
increase of the magnetizing field. The discrepancy between the
calculated and the experimental result is perhaps due to the
æolotropy of the materials. For, if it were not isotropic, the
lateral dilatation by longitudinal magnetization would not coin-
cide with the change of length by circular magnetization. It will
also be explained by the æolotropy of the specimens that in weak
fields, Bidwell’s calculation resulted in the large diminution of
volume of iron rings in contradiction to the experimentally
established fact.
1) loc. cit.
2) Nagaoka and Honda, loc. cit.
CHANGE OF DIMENSIONS BY MAGNETIZATION, 95
Concluding Remarks.
17. From the experiment on the relation between magnetism
and twist, Knott” concluded that the pure strain effects on a
ferromagnetic wire caused by tension and longitudinal current
through it are of an opposite character, and also, on the ground
of Maxwell's explanation for Wiedemann’s effect, that in an
iron Or nickel wire carrying an electric current, the change of
length by magnetization must be greater than when there is no
longitudinal current. Since the change of length for cobalt is
scarcely affected by tension, the same must also be the case for
longitudinal current. The consideration is partially verified by
the experiment of Bidwell and also by the present one.
The same phenomenon may also be more concisely explained
in the following manner. Suppose our samples to be isotropic
and to have no residual effect. Let Z and ¢ be two magnetic
forces acting longitudinally and circularly in two perpendicular
directions. When these two forces act simultaneously, we have
a resultant force 7; this force occasions the change of dimen-
sions in our ferromagnetics. The dilatation in the direction of
the resultant force, as well as that in the direction perpendicular
to it, can be expressed by /(77) and F(H) respectively, which
are even functions of ZZ To obtain the dilatation in the longi-
tudinal direction, we have simply to construet a strain ellipsoid
at any point of the ferromagnetics and to find the change of
length of the radius vector in this direction. The simple cal-
culation gives
ol,
L
7? EX. -
AH) —— + F(H)—
(He FH TER
1) Knott, Trans, Roy. Soc. Edinb, 36, pt. IL, 485.
—_—
rer
* oe
Hu ae ie) Pe ee eer > ia
96 K. HONDA,
In the case of nickel, the change of volume is negligibly small
compared with that of length; hence we may put with tolerable
accuracy f(H)+2F(H)=0. With steel and soft iron, the
change of volume is not very small compared with the change of
length. But if ¢ does not exceed 50 C.G.S. units, the effect of
volume change on the change of length by combined action of
Zand ¢ is negligibly small, for in these strong fields at which the
change of volume is pronounced, the ratio ’/H’ in the above
expression beeomes very small. Hence even for these metals,
we may neglect the change of volume, provided the circular
field is not very large, and the expression for li becomes,
in all cases,
DL EE
L =, SA).
Since the material is supposed to be isotropic, f(H) is the same
as the ordinary change of length by longitudinal magnetization.
Thus the change of length by longitudinal magnetization with
a constant circular field can be calculated from the change of
length by longitudinal magnetization alone. The same expres-
sion can also be used for the calculation of the change of
length due to circular magnetization with a constant longitu-
dinal field.
In order to compare the above result with that of the
experiment, it is obviously necessary to subtract from a the ex-
pression /’(¢) for the change of length by longitudinal magne-
tization with a constant circular field ¢, and f(Z) for the reciprocal
case.
Assuming for the expression /(Z) a suitable empirical for-
mula for iron, steel or nickel, a simple analytical discussion of
CHANGE OF DIMENSIONS BY MAGNETIZATION. 97
BY
the expression a or numerical calculation of it for different
values of / and ¢ from the experimental curve of the ordinary
change of length leads to the conclusion that for iron, steel
and nickel, all the points, which we have remarked in connection
with the curves shown in Figs, 1, 4, 5, 7, 8 and 10, are involved
without exception in the expression of a
It may be observed that the behaviour of cobalt with regard
to the change of length is just the reverse of that of iron, and
therefore every result which we have obtained for iron is also
applicable to the case of cobalt, provided the sign of the length
change be properly reversed. ‘Thus in strong fields, the length
of a cobalt tube should, by the combined effect of longitudinal
and circular magnetizations, become shorter than when acted
upon by the former alone. In weak fields, the result should be
just the opposite. The field of maximum contraction should
increase with the circular, and the amount of the contraction, after
reaching a maximum, gradually decrease. The circular field at
which the maximum contraction occurs should be far greater
than that for iron.
19. The comparison above made is qualitative; how the
calculated and the experimental numbers agree with each other
is seen from the following table:
98 K. HONDA.
TABLE IX.
Nickel ans t= 10.7
I 4 (cal.) L (exp) difference
10 | — 20x10 | — 25x10! 5xi0
2) ate JE | M
30 | —112 | —125 13
50 162 | —175 13
so | —204 216 |! 19
120 | -237 | —250 13
200 | —270 285 | 15
300 | —291 — 305 14
| ~309 | —33 | 14
|
— 391
Here L' denotes aid — (4), and its value was calculated from the
experimental curve for the ordinary change of length. A glance
in the above table shows a fair agreement between the calculated
and the experimental values. The difference between these num-
bers is not of a serious nature, if we remember that one scale
division of the micrometer ocular corresponds to the change o
5.12x10~ for nickel, and that the correction for temperature
amounts to 11x10~ in the most significant case.
The discrepancy is probably due to the residual effect and
also to the wolotropy of the tube. ‚If the tube, after it is
magnetized both longitudinally and circularly, is demagnetized
by reversals with regard to the longitudinal magnetization,
the circular field remaining constant, as was actually the case
in the present experiment, the elongation due to the circular
field alone is usually increased by one or two scale divisions,
CHANGE OF DIMENSIONS BY MAGNETIZATION. 99
a phenomenon which is perhaps to be attributed to the residual
effect noticed in my former paper”. The constancy of the
difference in the above table furnishes additional evidence in
support of this view. The æolotropy of the tube as regards the
change of length evidently influences the experimental values.
Moreover the change of the intensity of longitudinal mag-
netization due to the mutual interaction of longitudinal and
circular fields is not taken into account in the calculation of the
effective field. These causes, I believe, are sufficient to account
for the said discrepancy.
In steel and soft iron, there are comparatively large diffe-
rences between the calculated and the experimental numbers, as
will be seen from the following table:
TABLE X.
Wolfram steel, t=17.7 Soft iron, t=26.2
L’ (cal.) L’ (exp.) I/ (al) Z' (exp.)
2x10 1x10 9x10 8x10
22 18 23 30
31 37 29 38
39 51 31
46 57 29
48 62 21
48 64 12
47 63 me,
45 62 _17
For iron and steel, the sensibility of the apparatus was about
2x10” and the correction for temperature amounted to 5x 10”
1) K. Honda, Jour. Sc. Coll. XI., 311, 1899.
100 K. HONDA.
in the most significant case. I believe that the principal causes
of discrepaney above enumerated are sufficient to account for
the difference between the calculated and the experimental
numbers.
19. Thus qualitatively the above result and the experiment
are in complete agreement with each other, although there are some
discrepancies in quantitative details ; there are, however, probable
causes to account for the discrepancies. According to Knott, the
change of length in cobalt by longitudinal magnetization is very
little affected by the presence of a circular field, but the above
consideration leads to a result which contradicts his anticipation.
Hence a single experiment on this point for cobalt will deci-
dedly establish the correctness of the one explanation agains
that of the other.
In conclusion, I wish to express my best thanks to Prof
H. Nagaoka, and also to Prof. A. Tanakadate for useful advice
and kind guidance.
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Studien über die Anpassungsfähigkeit einiger
Infusorien an concentrirte Lösungen‘).
Von
Atsushi Yasuda, Rigakushi,
Professor der Naturgeschichte an der zweiten Hochschule zu Sendai.
Hierzu Tafel X-XII.
Einleitung.
In der Natur finden wir Thiere und Pflanzen stets den
obwaltenden Bedingungen angepasst. Diese Anpassung an die
Umgebung ist in der That erst im Verlaufe langer Generationen
hervorgebracht worden. Wenn dann aber die Lebensbedin-
gungen sich ändern, wie werden diese Organismen dadurch beein-
flusst? Unsere bisherigen Erfahrungen lehren uns, dass den
Organismen im Allgemeinen die Fähigkeit innewohnt, sich diesen
veränderten Verhältnissen genau anzupassen und so dauernd
leben zu können, nur unter der Bedingung, dass die Veränderung
nicht plötzlich stattfindet, oder, wenn sie rasch vor sich geht,
sie doch verhältnissmässig unbedeutend ist.
1) Die vorläufige Mittheilung dieser Arbeit erschien in The Botanical Magazine.
Tokyo 1897. Vol. XI, No. 121. pp. 19-24. und Annotationes Zoologice Japonenses. 1897.
Vol. I, Part I et I. pp. 23-29.
102 A. YASUDA : ANPASSUNGSFAEHIGKEIT
Bekanntlich giebt es in der Pflanzenwelt die Wasserformen
der amphibischen Gewächse, wie Polygonum amphibium und
Ranuneulus aquatılis, die sich in ihrer morphologischen und
anatomischen Beschaffenheit ganz anders als ihre Landformer
verhalten. Auch sind die Hydrophyten in Bezug auf Gestali
und Struktur einer grossen Metamorphose unterworfen, die sie
zum Leben im Wasser befähigt. Aehnliche Thatsachen finder
wir auch in der Thierwelt. Hierher gehören z. B. unter der
Amphibien die Anuren, deren aus den Eiern ausschliipfend
Larven durch Kiemen athmen, aber im erwachsenen Zustand:
durch Lungen, während die Thiere, welche zu den Perenni
branchiaten gehören, fortdauernd Kiemen besitzen, weil si
lebenslang im Wasser wohnen und niemals ein oberirdische
Leben führen, so dass sie sich jenem Medium völlig accommodir
haben.
Es dürfte daher von Interesse sein, wenn wir die Beschaf
fenheit der in der Natur gefundenen Medien verändern, künst
liche Nährmedien anfertigen und die Anpassungsfähigkei
gewisser für diesen Zweck ausgewählter Organismen an dies
künstlichen Medien studiren. Es liegen bereits Untersuchunge:
mancher Forscher über derartige Kulturversuche bei niederen
Organismen vor. Im nächsten Abschnitt fasse ich die Resultat
der wichtigsten einschlägigen Arbeiten zusammen.
Vorausschicken will ich noch, dass ich die Anpassung der In
fusorien an solche künstlichen Medien studirt habe, die aus hetero-
genen Lösungen von höheren Concentrationen bestanden. Fol.
gendes sind die Fragen, die ich zu beantworten versucht habe :—
1) Welche Grade der Concentrationen der Aussenmedien könner
die Infusorien ertragen ? 2) Welche relative Widerstandsfahigkeit
haben sie im: Vergleich mit Algen und Pilzen ? 3) Welche
EINIGER INFUSORIEN. 103
Veränderungen ihrer äusseren und inneren Gestalt werden
dadurch hervorgebracht, und wie wird ihr Bewegungs- und
Vermehrungsvermögen durch diese heterogenen Medien beein-
flusst? In Folgendem werde ich der Reihe nach die diesbezüg-
liche Litteratur, die Methodik meiner Versuche, die Beschreibung
der Experimente und endlich die Zusammenfassung der Resultate
mittheilen.
Die vorliegende Arbeit wurde grossentheils im botanischen
Institut der kaiserlichen Universität zu Tokyo unter freundlicher
Anregung des Herrn Prof. Dr. M. Miyoshi ausgeführt, wel-
chem ich für seine liebenswürdige Rathschläge meinen verbind-
lichsten Dank ausspreche. Herrn Prof. Dr. J. Matsumura
sage ich auch an dieser Stelle für das Interesse, welches er meiner
Arbeit entgegengebracht hat, meinen besten Dank.
Litteratur.
Ueber die Accommodation niederer Pflanzen giebt es ziem-
lich viele Versuche; so beobachtete Stahl'), dass Aethalium
septicum sich allmählich an Traubenzuckerlösungen anpasste und
der Einwirkung einer 2%igen Lösung widerstand. Richter’)
experimentirte mit Cyanophyceen und fand, dass Rivularia 3%
ige, Gloeocapsa 6%ige, Anabaena 6%ige und Oscillaria 10%ige
Kochsalzlösung ertragen konnten. Auch gelang es ihm, Dia-
tomaceen in einer 7%igen Kochsalzlösung ein Jahr und in einer
10%igen einen Monat lang leben zu lassen. Er zog ausserdem
1) E. Stahl. Zur Biologie der Myxomyceten. Bot. Ztg. 1884. Nr. 11. p. 166.
*) A. Richter. Ueber die Anpassung der Süsswasseralgen an Kochsalzlösungen. Flora.
1892. pp. 18-56. -
104 A. YASUDA : ANPASSUNGSFAEHIGKEIT
verschiedene andere Algen in den Bereich seiner Untersuchungen,
darunter Zygnema, Mougeotia, Spirogyra, Cosmarium, Chlorella,
Tetraspora, Chaetophora, Vaucheria, Oedogonium, Chara u. 8. w.
eine gewisse Anzahl von ihnen vermochte sogar in 13%ige
Lösung zu existiren. Ferner theilte Klebs') mit, dass einige in
den Lösungen organischer Verbindungen kultivirte Süsswasser-
algen anfänglich eine Plasmolyse zeigten, die aber nach einiger
Stunden vollständig ausgeglichen war, worauf sie ohne Schaden ir
den neuen Medien fortlebten. Nach demselben Autor gedieh Zyg-
nema in einer 10 bis 20%igen Glycerinlösung eine Woche lang
Auch 10-50%ige Rohrzuckerlösungen vermochten dieselbe Algı
im Leben halten, aber mit verschiedenen Wirkungen je nacl
der Concentration: 10%ige Lösung veranlasste lebhafte Kern.
theilung, 20-25%ige Längenwachsthum, 30%ige Zellhautbildun;
und 40%ige Assimilation und Stärkebildung, während in 50%
iger Lösung die Alge nur wenige Tage lebte.
Unter den Meeresalgen nahm Janse’) eine ähnliche Er
scheinung bei Chaetomorpha wahr, und zwar hervorgebracht durel
Kalisalpeter- und Kochsalzlösungen. Er fand nämlich, dass
wenn man diese Alge in eine solche Lösung legt, in Folge ihre
Anpassung an dieselbe nach kurzer Zeit ihre Widerstandsfähig-
keit bedeutend gesteigert wird. Oltmanns?) machte Experi-
mente über den Einfluss der Concentrationsinderung de
Meerwassers auf Fucus, der bei niedriger Concentration sich dem
neuen Medium gänzlich accommodirte. Eschenhagen‘) kulti-
1) G. Klebs. Beiträge zur Physiologie der Pflanzenzelle. Berichte der deutsch. bot
. Gesellsch. 1887. Bd. V, Heft 5. p. 181.
3) J. M. Janse. Plasmolytische Versuche an Algen. Bot. Centralbl. 1887. Bd. XXXII
p. 21.
3) F. Oltmanns. Ueber die Bedeutung der Concentrationsiinderung des Meerwassers fii
das Leben der Algen. Sitzb. d. Königl. preuss. Akad. d. Wissensch. zu Berlin. 1891. p. 193
¢) F. Eschenhagen. Ueber den Einfluss von Lösungen verschiedener Concentration
auf das Wachsthum von Schimmelpilzen. Stolp. 1889,
EINIGER INFUSORIEN. 105
virte Aspergillus niger, Penicillium glaucum und Botrytis cinerea
in verschiedentlich concentrirten Lésungen von Traubenzucker,
Glycerin, Natronsalpeter, Kalisalpeter, Chlornatrium und Chlor-
kalium, und wies sowohl die Grenzpunkte ihrer Accommoda-
tion als auch ihr Wachsthumsverhältniss zu diesen Substraten
nach. Bachmann’) bewies durch zahlreiche Experimente, dass
Thamnidium elegans durch veränderte äussere Bedingungen
gezwungen werden kann, diese oder jene Art von Sporangiolen
zu bilden oder die Bildung derselben gänzlich zu unterdrücken.
Ray’) site die Sporen von sSterigmatocystis alba in Medien,
welche aus Zucker, Stärke, Möhren, Kartoffeln, Gelatine und
mineralischen Salzen bestanden, und erhielt verschiedene aus
diesen Sporen entwickelte Pilzformen.
Auch für das Thierreich fehlt es an diesbezüglichen Unter-
suchungen nicht. Als Beispiele seien folgende angeführt :—
Schmankewitsch?) beobachtete, dass Branchipus stagnalis, der
immer in Süsswasser gefunden wird, sich, wenn man ihn in
versüsstem Meerwasser züchtet, der Form von Ariemia Milhau-
senii, einer das Brackwasser bewohnenden Art, nähert, und,
wenn man das Brackwasser so lange concentrirt, bis dasselbe
den Salzgehalt des Meerwassers erreicht hat, sich in Artemia
salina, eine Meerwasser-Art, verwandelt. Herbst?) ziichtete die
Larven einiger Seeigel in verschiedenen Lösungen von Lithium-,
1) J. Bachmann. Einfluss der äusseren Bedingungen auf die Sporenbildung von
Thamnidium elegans Link. Bot. Ztg. 1895. Abt. I. p. 128.
3) M. J. Ray. Variations des Champignons inférieurs sous l’influence du milieu.
Revue générale de Botanique. 1897. T. IX. pp. 193-259 et pp. 283-304.
3) W. Schmankewitsch. Zur Kenntniss des Einflusses der äusseren Lebensbeding-
ungen auf die Organisation der Thiere. Zeitsch. f. wiss. Zool. 1887. Bd. XXIX. p. 429.
*)C. Herbst. Experimentelle Untersuchungen über den Einfluss der veränderten
chemischen Zusammensetzung des umgebenden Mediums auf die Entwicklung der Thiere,
I. Theil. Zeitsch. f. wigs. Zool. 1892. Bd. XV. p. 446.
106 A. YASUDA : ANPASSUNGSFAEHIGKEIT
Natrium- und Kaliumsalzen, und fand, dass die Wirkungsstärke
auf die Entwicklungsstufen derselben in einer Reihe der Salze
von demselben Radical ihrem Molekulargewicht umgekehrt pro-
portional ist, d. h. ihre Wirkungsstärke nimmt um so stärker
ab, je mehr ihre Molekulargewichte zunehmen. Nach ihm
wurde in einem Falle die Gastrulation der Seeigel bedeutend
verzögert, in einem anderen Falle wurde die Pluteusorganisation
entweder mit der runden und gedrungenen Gestalt ohne Fortsätze
oder sogar ohne eine Spur des Kalkgerüstes gewonnen. Cohn‘)
bemerkte, dass eine plötzliche Concentrationsänderung des
Aussenmediums der Infusorien eine schädliche oder tödtliche
Einwirkung ausübt. Fabre-Domergue’) beobachtete auch das
Verhältniss der Ernährung in den Körpern einiger Infusorien,
und gelangte zu folgendem Schluss: „Dans des conditions par-
faites de nutrition prise dans l’acception la plus large du mot
il se produit des aliments de réserve qui disparaissent quand
des conditions deviennent défavorables à la vie.“ Weiter
studirte Bokorny?) die Veränderungen der Bewegung, der Ge-
stalt und der Grösse der Vacuolen von Paramaecium unter dem
Einfluss gewisser Basen, wie Coffein, Ammoniak und Kali, deren
1 promill. oder noch dünnere Lösung im Allgemeinen die Bewe-
gung verlangsamte, die Gestalt abrundete und sowohl Vergrösse-
rung der Vacuolen als auch das Auftreten von neuen verursachte.
1) F. Cohn. Entwickelungsgeschichte der microscopischen Algen und Pilze. Nova
Acta Akad. Caes. Leopold. 1851. Bd. XXIV, Th. 1. p. 132,
*) M. Fabre-Domergue. Recherches anatomiques et physiologiques sur les infu-
soires ciliés. Ann.d. Sc. nat. Zool. 1888. Sér. VII, T. 5. p. 135.
5) Th. Bokorny. Einige vergleichende Versuche über das Verhalten von Pflanzen
und niederen Thieren gegen basische Stoffe. Pflüger’s Archiv. 1895. pp. 557-562.
Bokorny gab auch in einer anderen Schrift (Vergleichende Studien über die Giftwir-
kung verschiedener chemischer Substanzen bei Algen und Infusorien. Pflüger’s Archiv.
1896. pp. 262-306.) eine genaue Untersuchung über die Giftwirkung von Basen und Säuren
unorganischer Natur, Salzen, Oxydations-Giften, Phosphor, organischen Säuren, Alkoholen,
Alkaloiden u. a. m, aufdas Leben der Infusorien und anderer Organismen,
EINIGER INFUSORIEN. 107
Endlich müssen noch die Resultate der Untersuchungen von
Davenport und Neal!) Erwähnung finden; sie züchteten Stentor
2 Tage lang in einer 0,00005% Sublimat enthaltenden Kultur-
lösung; die Thiere liessen sich sehr wohl acclimatisiren und
erwiesen sich gegen eine 0,001%ige sonst tödtliche Sublimat-
lösung ca. vier Mal länger widerstandsfähig als diejenigen, die
im Wasser kultivirt worden waren.
Ueberblickt man die Ergebnisse dieser Untersuchungen, so
ersicht man, dass sowohl den niederen Thieren als auch den
niederen Pflanzen die Fähigkeit innewohnt, sich geänderten
Aussenmedien leicht anzupassen. Da aber diese Fähigkeit bei
verschiedenartigen Organismen verschieden stark ausgeprägt ist
und unter Umständen mannigfaltig auftritt, so muss jeder specielle
Fall genau erforscht werden. Meine vorliegenden Studien sollen
in dieser Hinsicht einen kleinen Beitrag bringen.
Methodisches.
Als Versuchsmaterial wählte ich solche Infusorien aus, die
in Gräben und Teichen steis gefunden werden können. Da aber
die in der freien Natur vorkommenden Infusorien nie in reiner
Kolonie vorhanden sind, so liess ich sie in einem Gefisse
sich massenhaft entwickeln und unter Vorsichtsmassregeln eine
längere Zeit fortleben.
Genau nach den Angaben von Miyoshi’) kultivirte ich
die Infusorien in einem mit Spirogyra gefüllten Gefässe. Sobald
1) C. B. Davenport and H. V. Neal. On the Acclimatization of Organisms to
Poisonous Chemical Substances. Archiv für Entwicklungsmechanik der Organismen. 1895.
Bd. II, Heft 4. p. 581.
»)M. Miyoshi. Physiologische Studien über Ciliaten. The Botanical Magazine.
Tokyo 1896. Vol. X, No. 112. p. 43.
108 A. YASUDA: ANPASSUNGSFAEHIGKEIT
die grüne Masse der Alge sich allmiihlich zu verfärben begann
und die ursprünglich klare Flüssigkeit immer mehr getrübt
wurde, bemerkte ich die Entwickelung verschiedener Arten von
Infusorien, die sich oft mit einer erstaunlichen Schnelligkeit
vermehrten und sich auffallenderweise in bald fadenförmigen,
bald netzartigen Kolonien gruppirten. Die microscopische
Untersuchung ergab, dass diese Kolonien nur aus wenigen Arten
bestanden, die im Kampfe ums Dasein den Rest überwunden
hatten. Aus dieser Mischkultur isolirte ich einzelne Arten, indem
ich mittelst einer Pipette eine kleine Menge der Kolonie zusam-
men mit Wasser herausholte und in ein ebenfalls mit Brunnen-
wasser und der Alge gefülltes Gefüss versetzte. Bei gewöhnlicher
Zimmertemperatur zeigten diese Kulturen ca. in einer Woche üppige
Entwickelung ; nach vier oder fünf Wochen aber nahm die
Vermehrung ab, und endlich nach sechs Wochen konnte nur
noch eine ausserordentlich kleine Anzahl in der Flüssigkeit
gefunden werden. Um eine und dieselbe Art immer in üppiger
Kultur zu halten, legte ich deshalb alle drei Wochen neue
Kulturen an und trug dafür Sorge, dass sie nicht etwa durch
Bacterien infieirt wurden.
Nachdem ich die gewünschten Arten auf solche Weise in
Kultur hatte, wurden die Experimente auf zweierlei Weise
ausgeführt: einerseits prüfte ich die Anpassungsfähigkeit der
Infusorien in dem Zustande, wie sie in der Natur vorkommen,
d. h. in ihrem Zusammenleben mit Bacterien ; anderseits wandte
| | ich zu demselben Zwecke die Reinkultur jedes Infusors an, also
Der grösste Theil meiner Experimente wurde mit unreinen
|
| frei von Bacterien.
| Kulturen ausgeführt; in einigen Fällen wiederholte ich die
Experimente an Reinkulturen, um zu wissen, ob die Gegenwart
|
Digitized by G oogle
| EE Oe
EINIGER INFUSORIEN. 109
der Bacterien etwa das Ergebniss der Experimente modificirt
hätte. Die Resultate stimmten aber bei beiden Kulturen vollkom-
men überein, wie wir nachher sehen werden.
Als äussere Medien verwendete ich Lösungen von Rohrzucker,
Traubenzucker, Milchzucker, Glycerin, Kalisalpeter, Natron-
salpeter, Chlorkalium, Chlornatrium und Chlorammonium in
verschiedenen Concentrationen. Diese Stoffe waren chemisch
rein und wurden yor dem Gebrauch vollständig getrocknet.
Folgende Infusorien wurden bei meinen Studien ausschliesslich
verwendet : Æuglena viridis, Chilomonas paramaecium, Mallomonas
Plossliv, Oolpidium colpoda und Paramaecium caudatum. Alle
Kulturen, sowohl unreine als reine, wurden bei Zimmertem-
peraturen von 25°-30° C gehalten und in den . Wintermonaten
in einen Thermostat von etwa 30° C gestellt.
Für unreine Kulturen benutzte ich eine grosse Anzahl der
5 cm hohen und 3 cm weiten cylindrischen Glasgefässe, deren jedes
30 ccm der Versuchslösung und etwa 1 Gramm Spirogyra-Füden
enthielt. Da die Infusorien im Brunnenwasser weit besser gedeihen
als in destillirtem Wasser, so wendete ich bei der Zubereitung
der flüssigen Versuchsmedien das letztere als auflösendes Mittel für
verschiedene Substanzen an, wobei die Menge der darin gelöst
vorhandenen Stoffe mit Ausnahme von Kochsalz!) so unbedeutend
war, dass ich sie ohne grosse Ungenauigkeit ausser Acht lassen
konnte. Bei vielen Kulturen, die gleichzeitig gemacht wurden,
nahm ich keinen Anstand, eine Kontrollkultur, in welcher nur
Brunnenwasser und Spirogyra-Fäden angewendet wurden, anzu-
fertigen und zum Vergleiche dienen zu lassen.
In Bezug auf die Reinkultur der Protozoen im Allgemeinen
1) Die chemische Analyse des Brunnenwassers zeigte, dass es 0,095% Kochsalz
enthielt.
110 A. YASUDA : ANPASSUNGSFAEHIGKEIT
haben viele Forscher') in neuerer Zeit versucht, sie entweder auf
festen Substraten oder in flüssigen Medien zu züchten. Bei der
Isolirung der Infusorien befolgte ich genau die Methode von
Ogata’) mit positivem Resultat. Ich liess mir nach seiner Vor-
schrift feine Glascapillarrôhren anfertigen, deren Durchmesser je
nach der Grösse des Versuchsobjects variirten. So war z. B. bei
Chilomonas paramaecium, dessen Körper 25-30 « Länge und
10-12 «# Breite hat, das Capillarrohr etwa 0.1 mm in innerem
Durchmesser und 10 cm in Länge, während es bei Calpidium
colpoda, dessen Körper 60-70 # lang und 25-30 « breit ist, einen
inneren Durchmesser von 0,15 mm und eine Länge von 10 cm
hatte.
Sobald das Capillarrohr nach dem Eintauchen in eine
sterilisirte Nährlösung mit der letzteren grossentheils gefüllt
war, brachte ich das nämliche Ende desselben in die Mischkul-
turflüssigkeit von Bacterien und Infusorien und liess das Rohr
sich mit der Flüssigkeit völlig füllen. Untersucht man ein
solches Capillarrohr unter einem Microscope, so findet man an
der Capillarrohrmündung eine grosse Anzahl von Infusorien in
Bewegung. Einige streben sich ins Innere zurückzuziehen, bald
aber kommen sie nach der Mündung zurück. Wegen der starken
Aörotaxis und schwachen Chemotaxis der Organismen’) gelingt
1) Während Beijerinck (Kulturversuche mit Amöben auf festen Substraten. Centralbl.
f. Bak. u. Parasit. 1896. Bd. XIX, No. 8), Celli (Die Kultur der Amöben auf festen Sub-
straten. Centralbl. f. Bak. u. Parasit. 1896. Bd. XIX, No. 14/15.), Schardinger (Reinkulturen
von Protozoen auf festen Nährboden. Centralbl. f. Bak.u. Parasit. 1896. Bd. XIX, No. 14/15.),
"Gorini (Die Kultur der Amöben auf festen Substraten. Centralbl. f. Bak. u. Parasit. 1896.
Bd. XIX, No. 20), Tischutkin (Ueber Agar-Agarkulturen einiger Algen und Amöben.
Centralbl. f. Bak., Parasit. u. Infekt. 1897. Bd. III, No. 7/8.) und andere Forscher Amöben
auf festen Substraten künstlich züchten konnten, ist es Ogata (Ueber die Reinkultur gewis-
ser Protozoen-Infusorien. Centralbl. f. Bak. u. Parasit. 1893. Bd. XIV, No. 6.) auch gelun-
gen, Polytoma uvella in füssigen Medien rein zu kultiviren.
3) M. Ogata. loc. cit. p. 168.
3) M. Miyoshi. loc, cit. p. 48.
EINIGER INFUSORIEN. 111
es nicht immer, die Infusorien auf diese Weise hervorzulocken, und
ihrer habhaft zu werden. Nur wenn sie zufällig in die sterilisirte
Flüssigkeit tief eindringen, kann man unter dem Microscope das
Capillarrohr an der betreffenden Stelle abbrechen und dann das
Ende des Rohrs zuschmelzen. Sodann impft man den infu-
soriumhaltigen Capillarrohrinhalt, indem man das Rohr mit
einem sterilisirten Pincet abbricht und den Inhalt in ein mit
sterisirter Nährlösung gefülltes Reagensglas hineinbläst.
Bei meinen Versuchen impfte ich wenigstens zwei Individuen
in ein und dasselbe Reagensglas, um erstens den Effect der Inocu-
lation zu sichern, und zweitens mit der Hoffnung, dass sie, wenn
alle beide in dem neuen Medium unversehrt fortlebten, durch
Copulation sich vermehren könnten. Bei der Zimmertemperatur
von 20° C blieb die geimpfte Nährflüssigkeit nach zwei oder
drei Tagen vollkommen klar, und erst naeh ungefähr zehn Tagen
erschienen Hunderte von Individuen, die nahe der Oberfläche
der Flüssigkeit als sehr kleine weisse Pünktchen hin und her
schwammen. Die Zahl dieser weissen Pünktehen nahm hernach
allmählich zu, und dieselben waren nicht allein am oberen Theil
des Reagensglases, sondern auch am mittleren und unteren Theil
desselben zerstreut sichtbar. Eine solche Erscheinung bedeutet,
dass die Reinkultur gut ausgefallen ist, und dass man in jener
Nährlösung nichts anders als die isolirte Art der Infusorien
findet. Wenn aber die Nährlösung während der Impfung von
Bacterien inficirt wird, so tritt immer eine starke Trübung zu
Tage, und man bekommt in diesem Falle selbstverständlich
keine Reinkultur der Infusorien.
Die auf diese Weise hergestellte Reinkultur gedieh 4-5
Wochen lang, vorausgesetzt dass die Nährstoffe in der Kultur-
flüssigkeit nicht völlig erschöpft waren. Durch erneuerte Wieder-
112 A. YASUDA : ANPASSUNGSFAEHIGKEIT
impfung konnte ich die Organismen in reinem, gutem Kultur-
zustande eine lange Zeit erhalten.
Die Nährlösung, die ich für die Reinkultur gebrauchte,
stellte ich nach der Vorschrift von Ogata an, und zwar war
ihre Zusammensetzung folgende :
BISRERBSETSC nen 1g
RATION ea 20 5,
Concentrirt gekochte Lösung von Porphyra vulgaris. 250 ccm
DAMES. WASSER... eek 729 „
Beschreibung der Versuche.
Wie gesagt, stellte ich die Experimente hauptsächlich mit
unreinen Kulturen an, in der Absicht, den Einfluss, welchen die
äusseren Bedingungen auf die Infusorien in ihrem natürlichen
Vorkommen ausüben, festzustellen. Dabei versäumte ich aber
nicht, Kontrollversuche mit reinen Kulturen zu machen und die
beiden Resultate zu vergleichen.
Bei allen Versuchen mit unreinen und reinen Kulturen liess
ich die Beschaffenheit des Mediums sich plötzlich ändern und
prüfte die Anpassungsfähigkeit unserer Organismen an das neue
Medium. Hatte ich unreine Kulturen, so verglich ich gewöhn-
lich im Verlauf von 1-7 Tagen, zuweilen aber erst nach einem
Monat, die Wirkungen der verschiedenen Mediumsconcentrationen
auf das Reproductionsvermögen und die Gestaltänderungen der
Versuchsorganismen. Speciell bei den Versuchen mit Rohrzucker
war es nöthig, eine Reinkultur anzuwenden, weil bei unreiner
Kultur der Rohrzucker durch vorhandene Bacterien oder Pilze
nach und nach invertirt und schliesslich gespalten wurde. Um
EINIGER INFUSORIEN. 118
zu erkennen, nach wie vielen Tagen der Rohrzucker zum Trau-
benzucker invertirt wird, prüfte ich mit der Fehling’schen
Lösung und fand, dass in meinen Versuchen nach etwa 4 Tagen
eine kleine Inversion stattgefunden hatte. Meine unreinen
Rohrzuckerkulturen waren daher binnen der ersten drei Tage
doch noch brauchbar.
Dass diese Inversion ausser durch Bacterien und. Pilze auch
unter Mitwirkung der Infusorien stattfände, ist schon von vorn
herein unwahrscheinlich. Um dies aber experimentell zu con-
statiren, stellte ich einige Versuche mit Rohrzuckerreinkulturen')
an, und gelangte beim Prüfen der fraglichen Flüssigkeit wie
erwartet zu negativem Resultat. Die Kontrollkultur mit Asper-
gillus glaucus zeigte eine starke Inversion, |
Bei allen Kulturen mit verschiedenen Stoffen machte ich
immer Kontrollkulturen, und bei den kritischen Versuchen, wie
z. B. der Bestimmung der Concentrationsgrenze einer Flüssigkeit,
in welcher die Organismen sich mehr oder minder anpassend
leben können, wurden dieselbe Kulturen einige Male wiederholt.
Ich gehe nun zur Beschreibung der einzelnen Versuche bei
jeder Art meiner Versuchsorganismen über.
(a) Huglena viridis Ehrbg.”)
Dieser Organismus hatte in der Kontrollkultur folgende
1) Da: der Rohrzucker bekanntlich bei langem Kochen zum Theil in Trauben- und
Fruchtzucker verwandelt wird, so kann bei den Rohrzuckerreinkulturen die gebräuchliche
Sterilisirung durch Hitze nicht ohne Vorsichtemassregeln angewendet werden. Ich sterilisirte
deshalb den Rohrzucker mit absolutem Alkohol und brachte ihn dann in die vorher
sterilisierte Nährlösung ein.
3) Figuren in Friedrich Ritter v. Stein, Der Organismus der Infusionsthiere.
Leipzig 1878. Abt. III, Heft I. Taf. XX., W. Saville Kent, A Manual of the Infusoria.
1880-81. Vol.L PL XX. und O. Bütschli, H. G. Bronn’s Klassen und Ordnungen des
Thierreiches. 1883-87. Bd. J. Protozoa, Abt. IL Taf. XLVIL
114 A. YASUDA : ANPASSUNGSFAEHIGKEIT
Merkmale : Gestalt gewöhnlich spindelförmig, Hinterende schärfe
zugespitzt, aber wegen der Metabolie sich mannigfaltig verä
dernd. Aus dem Schlunde entspringt eine lange Geissel. Chr
matophoren zahlreich vorhanden, klein, scheibenförmig und rei
grün gefärbt. Eine contractile Vacuole nahe dem Vorderend
gelegen. Dicht bei demselben Ende befindet sich auch ein rothe
Augenfleck. |
Versuch 1. Rohrzucker, C:H20,.—Von einer 1%ige
Lösung anfangend liess ich in anderen Kulturen die Concentré
tion um je 1% steigen. Obgleich die Accommodation schwe
wurde, als die Concentration zunahm, so lebte das Infusor doc
bis zur 15%igen Lösung, welche die Maximalconcentration fü
den Organismus war. 1%ige, 2%ige und 3%ige Kulture
zeigten keine wesentliche Veränderung am Körper des Organis
mus. Bei einer 4%igen Lösung aber begannen die Chromato
phoren an Grösse zuzunehmen. Von 1% iger Lösung bis z
7%iger war die spirale Bewegung des Organismus lebhafl
dagegen über 8% wurde sie allmählich langsamer, während di
Chromatophoren selbst sich merklich ausdehnten ; als die Con
centration des Mediums zunahm, wurde auch die Vermehrun;
verhindert. Bei 12%iger Lösung konnte das Thier nicht meh
normal gedeihen, bei 13% überlebte eine kleine Anzahl, di
jedoch nach einer Woche alle zu Grunde gingen ; bei 14% lebter
noch einige Individuen, aber nicht länger als 4 Tage, währenc
sie bei 15% kaum einen Tag lebendig blieben. Da der Organis
mus metabolisch ist, so konnte keine deutliche Veränderung aı
seiner äusseren Gestalt beobachtet werden.
Versuch 2. Traubenzucker, C,.H,O;—Unser Organismu:
konnte 1-11%ige Concentrationen ertragen. Bei 1%- und 2%-
Kultur war noch keine merkliche Veränderung wahrzunehmen,
EINIGER INFUSORIEN. 115
aber schon bei 3%iger Lösung dehnten sich die Chromatophoren
ein wenig aus, und bei der Concentration über 3% wurden sie
noch etwas grösser. Die Bewegung des Thierkörpers schien bei
1-6%-Kulturen normal zu verlaufen ; erst bei 7% wurde sie
langsamer mit gleichzeitiger Verminderung der Vermehrungs-
fähigkeit. Bei einer 9%igen Lösung vermehrten sich die Thiere
überhaupt nicht mehr, und nach einer Woche war nur noch
eine kleine Anzahlam Leben. Alle Individuen des Infusoriums
gingen bei einer 10%- Kultur nach einer Woche, und bei einer
von 11% schon nach einigen Tagen zu Grunde.
Versuch 3. Milehzucker, CeH20u1+H0.—Unter den oben
erwähnten Zuckerarten schien unser Infusor sich an Milchzucker
am besten anzupassen. Die Maximalconcentration, welcher es
widerstehen konnte, war eine 17%ige. Von 4% an aufwärts
schienen die Chromatophoren sich zu vergrössern. Bei 1-11%-
Kulturen nahm die Multiplication rasch zu, aber über 12%
wurde sie etwas vermindert und auch die Bewegung wurde
einigermassen träge. Eine 17%ige Lösung erwies sich als die
Grenzconcentration für das Versuchsthier.
Versuch 4. Glycerin, C;H;0,.—Die Versuche lehrten uns,
dass sich unser Infusor an Glycerin weit schlechter anpasste als
an eine der oben erwähnten drei Zuckerarten, denn das Thier
konnte nur 1-6%ige Lösungen ertragen. Bei einer 2%igen
Lösung erweiterten sich die Chromatophoren ; bei 3%- Kultur
lebte eine kleine Anzahl noch am fünften Tage, und bei 6%
blieben nur wenige Individuen noch einige Tage am Leben.
Die Bewegung wurde bei einer 4%igen Concentration schon
vielfach retardirt, und bei 6% hörte sie fast gänzlich auf.
Ferner war in der letzteren Lösung eine pathologische Erschei-
nung wahrzunehmen, indem die Cuticula des Körpers um die
116 A. YASUDA : ANPASSUNGSFAEHIGKEIT
Chromatophoren etwas einschrumpfte, sodass ihr Umriss im
optischen Schnitte gesehen zickzackformig aussah, und die
Chromatophoren selbst verschmolzen mehr oder weniger mit
einander.
Versuch 5. Schwefelsaures Magnesium, MgSO,—-Unter den
unorganischen Substanzen erwies sich das schwefelsaure Magnesium
als dem Leben des Organismus am besten zusagend. Der
Organismus konnte die Concentration von 1-6% vertragen. Die
Chromatophoren nahmen in ihrer Grösse fortwährend zu, als die
Concentration von 1,5% bis auf ihr Maximum stieg, In einer
8,4%igen Lösung zeigte das Thier eine sehr träge Bewegung,
und schon bei 4-6%igen Lösungen ging es beinahe zum
Stillstand über. Betrug die Concentration nur 1-2,5 %, so gedieh
unser Thier einen Monat lang vollkommen normal, aber von
einer 2,6%igen Concentration an aufwärts büsste es seine Ver-
mehrungsfähigkeit ein, und endlich bei 5-6% blieben nur
vereinzelte Individuen am Leben.
Versuch 6. Salpelersaures Kalium, KNO,.—Der Organismus
widerstand einer 2,4%igen Concentration. Von 0,8% an fingen
die Chromatophoren an sich zu erweitern ; über 2% wurde die
Bewegung sehr langsam. Im Allgemeinen schien das vorliegende
Salz auf die Mulptiplication des Infusors hemmend zu wirken,
da selbst bei verdünnten Lösungen das Thier eine unbedeutende
Vermehrung zeigte.
Versuch 7. Salpetersaures Natrium, NaNO,—-Dieses Salz
verhielt sich fast wie das vorige. Eine 2%ige Lösung war das
Maximum, welches unser Thier ertragen konnte. Die Chromato-
phoren dehnten sich schon von 0,8% an aus. Die Bewegung
war bei 2%iger Lösung nach zwei Tagen sehr träge.
Versuch 8, Chlorkalium, KCl.—Nächst dem Magnesium-
EINIGER INFUSORIEN. 117
wies sich unter den unorganischen Stoffen das Chlor-
ür die Vermehrung des Organismus am günstigsten.
%ige Lösung verursachte sowohl Zahlvermehrung als auch
erweiterung der Chromatophoren. In 0,2-1%igen
ationen gedieh der Organismus noch nach 40 Tagen.
. Concentration auf 2,3%, welches die maximale Grenze
Organismus war, so hörte die Bewegung fast gänzlich
rend die Chromatophoren sich theilweise zu grösseren
verschmolzen.
such 9. Chlornatrium, NaCl.—0,2-1,8% waren die Con-
nen, bei welchen das Thier am Leben blieb. Die
ophoren schienen bei einer 0,8%igen Lösung an Grösse
nen, und bei einer 1,6%igen Concentration zeigte der
aus noch eine langsame Bewegung.
such 10. Chlorammonium, NH,Cl.—Dieses Salz wirkte
en oben genannten Stoffen am ungünstigsten auf das
s Organismus ein, sodass die Anpassungsgrenze hier am
en war. 0,2-0,6% Kulturen gediehen noch am Ende der
Voche, aber über 1% vermehrte sich das Thier nicht mehr,
1,4% lebten kaum noch einige Individuen. Bei 0,6%
ung nahm die Grösse der Chromatophoren zu, und bei
shmolzen sie sich zu wenigen grösseren Körnern. Eine
Lösung verursachte immer die Verschmelzung der
ophoren und hob gleichzeitig die Bewegung des Orga-
uf.
wiederholte dieselbe Versuche zehnmal mit Reinkulturen
lich die Resultate mit denjenigen bei den unreinen Kul-
Jie Ergebnisse stimmten in beiden Fällen völlig überein.
eobachtete ich, dass bei den Versuchen mit Reinkulturen
elligkeit der Multiplication für die Lösungen verschiede-
| | }
Digitized Me ( IX [
|
118 A. YASUDA : ANPASSUNGSFAEHIGKEIT
ner Concentrationen eines und desselben Stoffes nicht allzu
gleich war, obgleich sie gleichzeitig geimpft worden waren
Im Allgemeinen nahm die Vermehrungsenergie in dem Masse
ab, wie die Concentration des Mediums stieg; so war zum
Beispiel bei einer Traubenzuckerkultur, im Verlaufe von 2
Wochen nach der Impfung, die Vermehrung eine starke bei
2%, eine mässige bei 4%, eine sehr unbedeutende bei 6%, eine
noch spärlichere bei 8% und keine bei 10%. Ferner war die
Vermehrung bei derselben Kultur nach 4 Wochen bei 2% eine
sehr starke, bei 4% eine starke, bei 6% eine mässige, bei 8%
eine unbedeutende und bei 10% eine höchst spärliche Vermehrung
zu beobachten, während sich am Ende der sechsten Woche bei
2-4% eine sehr starke, bei 6% eine starke, bei 8% eine mässige
und bei 10% eine spärliche Vermehrung zeigte. Auch beim
Milchzucker wurden ähnliche Thatsachen constatirt. So gediehen
nach 2 Wochen 2-4%-Kulturen ausgezeichnet ; 6%-Kultur zeigte
eine starke, 8% eine mässige, 10% eine spärliche, 12% eine noch
schwächere und 14% gar keine Multiplication mehr. Nach 4
Wochen aber vermehrten sich die Organismen bei 2-6% sehr
stark, bei 8% stark, bei 10% mässig, bei 12% spärlich und bei
14% sehr spärlich. Endlich nach 6 Wochen gedieh die Multi-
plication stark bei 10%, mässig bei 12% und spärlich bei 14%.
Auch für schwefelsaures Magnesium, salpetersaures Kälium,
Chlornatrium u, s. w. habe ich ähnliche Erscheinungen wahr-
genommen. |
(b) Chilomonas paramaecium Ehrbg.')
Der Organismus in Kontrollkultur hatte folgende Charak-
teristika : Körper nicht metabolisch, sondern plastisch. Gestalt
1) Figuren in Friedrich Ritter v. Stein. loc. cit. Taf. XIX, W. Saville Kent.
loc. cit. Pl. XXIV und O. Bütschli. loc. cit. Taf. XLV.
EINIGER INFUSORIEN. 119
länglich oval, seitlich comprimirt. Vorderende breiter und schief
abgestutzt, Hinterende dagegen rundlich zugespitzt. Zwei
Geisseln am Vorderende, eine derselben rollte sich, wenn der
Organismus im Ruhezustande war. Zahlreiche sphäroidische
Amylumkörner dicht unter der Körperoberfläche. Eine contractile
Vacuole am Vordertheil des Körpers.
Versuch 1. Rohrzucker.—Mit einer 1%igen Lösung anfangend
liess ich die Concentration um je 1% steigen, wie es bei Euglena
wiridis der Fall war. Von 2% an aufwärts dehnten sich die
Körnchen fortwährend aus. Ueber 6% nahm der Organismus
an Dicke und Breite zu, nicht aber an Länge, und sah so
einfach oval aus. Die Vermehrung wurde schon bei 4% verzögert;
bei 7% lebten einige Individuen noch eine Woche lang. Die
Individuen aus der 7%igen Kultur waren so träge, dass sie an
einem bestimmten Platze still lagen und nur eine zitternde Be-
wegung zeigten ; vor ihrem Tode hüpften sie einige Male
rückwärts. Die letztere Erscheinung wurde auch bei den con-
centrirten Lösungen anderer Stoffe beobachtet.
Versuch 2. Traubenzucker.—Der Traubenzucker wirkte stärker
als der Rohrzucker. Die höchste Concentration, die der Orga-
nismus vertragen konnte, war 6%. Bei 4%-Kultur ging die
Vermehrung nicht mehr gut vor sich, und bei 5% blieb nur
eine kleine Anzahl der Individuen am Leben. 2%ige Con-
centration bewirkte, dass die Körnchen sich vergrösserten, und
bei 5% wurde die Bewegung sehr langsam. Bei 6% kam der
Organismus fast zum Stillstande, und wurde eine Unebenheit des
Körperumrisses hervorgerufen.
Versuch 3. MMilchzucker.—Der Organismus widerstand 1-8% -
igen Concentrationen. Ueber 3% vergrösserten die Körnchen ihr
Volumen, bei 6% wurde die Multiplication verhindert und end-
120 | A. YASUDA : ANPASSUNGSFAEHIGKEIT
lich bei 8% lebten nur noch einige Individuen eine Woche lang
weiter, mit dem erwähnten Unebenwerden der Körperumrisse.
Merkwürdig war, dass der Körper an Dicke und Breite zunahm,
als die Concentration stieg.
Versuch 4. Glycerin. —Eine 4%ige Lösung war. die Maxi-
malconcentration für die Accommodation des Thieres. Bei 2%
erweiterten sich die Körnchen, bei 3% hörte die Vermehrung auf.
und bei 4% lebten nur noch einige Individuen, deren Körpe:
unregelmässige Umrisse zeigten.
Versuch 5. Schwefelsaures Magnesium.—In 1-3%igen Lö
sungen lebte der Organismus fort. Von 0,8% an aufwärts ver.
srösserten sich die Köruchen, und in 3%iger Lösung wurder
dieselben auffallend gross. Eine 1,4%ige Lösung verhinderte di
Multiplication. Bei höheren Concentrationen trat bei einigen In-
dividuen ein unregelmässiges Aussehen zu Tage. Diese Gestalt
änderung wurde bei einer 2,5%-Kultur besonders gut beobachtet
indem alle Individuen noch 2 Wochen mit einer ungewöhnlicher
Unebenheit ihrer Körpergestalt fortlebten. |
Versuch 6. Salpetersaures Kalium.—Eine 2%ige Concentra-
tion schien die obere Grenze der Anpassung zu sein. Eine
0,5%ige Lösung veranlasste eine Vergrösserung der Amylum-
körner, die bei einer 1%igen Lösung nach einer Woche einen
sehr grossen Durchmesser zeigten. Die Vermehrung ging nut
bei niederen Concentrationen gut vor sich.
Versuch 7. Salpetersaures Nairıum.—Der Organismus konnte
in 0,2-1,2%igen Lösungen leben. Die Volumenzunahme der
Körnchen fand von 0,6% an statt. Bei höheren Concentrationen
gedieh unser Organismus nicht. Im Uebrigen fast dieselben
Erscheinungen wie beim vorhergehenden Versuche.
Versuch 8. Chlorkalium.—Eine 0,8%-Kultur am Ende des
EINIGER INFUSORIEN. 121
Tages nach der Impfung untersucht zeigte sowohl Ver-
ng der Körnchen als auch Abrundung des Körpers. Nach
einer Woche besassen einige Individuen in einer 1%
ung eine fast scheibenförmige Gestalt. Ueber 2% konnten
; mehr leben. |
such 9. Chlornatrium.—Eine 0,4%ige Lösung liess die
n sich erweitern. Bei 0,8-1%-Kulturen dehnte sich
ımen bedeutend aus, indessen der Körper des Organismus
kürzte und rundlich wurde. Bei 1%iger Concentration
äusserste Grenze der Accommodation erreicht.
such 10. Chlorammonium.—Der Organismus konnte sich
0,6%ige Concentrationen anpassen. Bei einer 0,2%
sung trat schon Körnchenvergrösserung ein, und bei
igten einige Individuen unebene Umrisse, mit gleichzei-
sschwächung ihrer Bewegung. Wie bei Euglena viridis
bei Chilomonas paramaecium übte der vorliegende Stoff
n augewandten Chemikalien die stärkste Einwirkung aus.
(c) Mallomonas Plessliv Perty.')
" Organismus in der normalen Kultur zeigte folgende
le: Gestalt oval, am Vorderende etwas schmaler. Die
Cuticularoberfläche mit langen, biegsamen, borstigen
n bekleidet; am Hinterende mit einer langen Geissel
. Anstatt der Amylumkörner war eine Anzahl von
n vorhanden. Eine contractile Vacuole befand sich nahe
teren Ende. Das Thier schwamm mit lebhafter Bewegung,
; oft plötzlich stillstand.
rsuch 1. Rohrzucker.—Der Organismus vertrug Anpas-
uren in W. 8. Kent. loc. cv. Pl. XXIV.
122 A. YASUDA : ANPASSUNGSFAEHIGKEIT
sungsconcentrationen von 1-7%. Auch bei höheren Concentra-
tionen nahm die Grösse des Körpers mehr oder minder zu,
während die Multiplication und die Lebhaftigkeit der Bewegung
allmählich sanken. Ferner wuchs die Zahl und Grösse
der Vacuolen mit der Conceutrationserhöhung bedeutend an;
diese Erscheinung trat schon bei einer 2%-Kultur ein. Erst
von 4% an wurde die Multiplication schwächer und bei 7%
erlitt die Bewegung eine Retardirung, welche zu der sehr
schnellen, normalen Bewegung in grossem Contrast stand.
Versuch 2. Traubenzucker.—Eine 6%-Kultur war das Maxi-
mum der Anpassung des Organismus. Die Vergrösserung des
Körpers und die Zunahme der Vacuolen bei stärkeren Concen-
_trationen waren wie beim Rohrzucker. Eine 2%ige Lösung
erweiterte die Vacuolen einigermassen, und von 3% an nahm
die Vermehrung des Organismus ab. Die Bewegungshemmung
war schon bei 4% zu beobachten, noch stärker bei 5%.
Versuch 3. Milchzucker—Das Infusor ertrug 1-9%ige
Lösungen. Das allgemeine Resultat stimmte mit dem bei den
anderen Zuckerarten überein ; der einzige Unterschied war der,
dass der Milchzucker eine schwächere Einwirkung auf den
Organismus ausübte als die anderen Zuckerarten. Die Vacuolen
fingen erst bei einer 3%igen Lösung an sich zu vermehren, und
die Multiplication wurde erst von 7% an etwas verlangsamt.
Versuch 4. Glycerin. —Die Grenze der Accommodation war
eine 4%ige Lösung. Bei einer 2%- und noch auffallender bei
einer 3%-Kultur fand Zunahme der Zahl und Grösse der Vacuolen
und Anschwellen des Organismus statt.
Versuch 5. Schwefelsaures Aagnesium.—Der Organismus
vermochte sich 1-3,4%igen Lösungen anzupassen. Eine 0,8%
ige Coneentration verursachte sowohl Vermehrung als auch
EINIGER INFUSORIEN. 123
Vergrösserung der Vacuolen, und bei höher concentrirten Lö-
sungen war Verhinderung der Multiplication und Retardation der
Bewegung wahrzunehmen.
Versuch 6. Salpetersaures Kalium.—Eine 0,7 ren Lösung
vergrôsserte die Vacuolen etwas. Die Bewegung begann bei
1-1,5%igen Lösungen sehr langsam zu werden. Bei den Kul-
turen höherer Concentrationen pflegte der Organismus sich nicht
zu vermehren, und im Verlaufe einiger Tage ging der grössere
Theil der Individuen zu Grunde. Eine 1,5%ige Lösung bildete
die Grenze der Anpassung.
Versuch 7. Salpetersaures Natrium.—Für diesen Stoff besass
der Organismus eine besonders grosse Resistenzkraft. Er ver-
mochte sich sogar einer 2,6%igen Concentration anzupassen,
wenn auch mit grosser Schwierigkeit. Die Bewegung war bei
1,5% noch lebhaft.
Versuch 8. Chlorkalium.—Bei einer 0,8%-Kultur vergrösser-
ten sich die Vacuolen. Die Grenze der Anpassung des Organis-
mus war bei 1,4%iger Lösung zu beobachten. Mit der Con-
centrationssteigerung trat Körperabrundung ein.
Versuch 9. Chlornatrium.—Die Maximalconcentration war
1,5%. Bei 0,8%-Lésung schien der Körper nach 5 Tagen sich
abzurunden. In einer 1%igen Lösung konnte das Thier 3
Wochen lang gedeihen, aber in 1,5% starb es schon am Ende
des vierten Tages.
Versuch 10. Chlorammonvum.—Für diesen Stoff besass der
Organismus die kleinste Anpassungsfähigkeit, ganz wie es bei
den anderen Infusorien der Fall war. Eine 0,8%ige Lösung
war das Maximum. Die Cuticularoberfläche des lebenden
Organismus zeigte in dieser Lösung nach einem Tage einige longi-
tudinale Falten.
124 A. YASUDA : ANPASSUNGSFAEHIGKEIT
(d) Colpidium colpoda Ehrbg.')
Merkmale des Organismus in der normalen Kultur :—
Körper mittelgross, nierenférmig. Rückenseite mässig gewölbt;
Bauchseite in der Nähe des Mundes etwas eingebuchtet. Vor-
derende viel schmaler als das abgerundete Hinterende. Cuti-
cularwimpern auf der ganzen Oberfläche des Körpers reichlich
vorhanden und an Grösse alle gleich. Mund in mässiger Ent-
fernung vom Vorderende, in einer die Bauchseite querenden
‘ Einbuchtung. Eine contractile Vacuole und einige Nahrungs-
vacuolen vorhanden. Bewegung lebhaft.
Versuch 1. Rohrzucker.—Die Concentrationsdifferenz der
Versuchsserie war 1%. 8% wurde als das Maximum erkannt.
Schon bei einer 3%igen Lösung begannen die Vacuolen sich
etwas zu vermehren und zu vergrössern. Diese Erscheinung
wurde mit der Concentrationserhöhung immer mehr merklich.
Ueber 4% sah der Körperumriss rundlich aus und die Grösse
nahm merkwürdig zu. Multiplicationshemmung schon bei 6%.
Versuch 2. Traubenzucker.—Der Organismus lebte in 1-7%
igen Lösungen. Vermehrung und Vergrösserung der Vacuolen
schon bei 2% und Abrundung des Körpers bei 3%. Bei einer
4,5%-Kultur wurde die Multiplication sehr verzögert, bei 6%
lebte am Ende des fünften Tages noch eine kleine Anzahl der
Thiere; bei 7% waren nur noch vereinzelte Individuen am
Leben, welche schliesslich nach 5 Tagen abstarben.
Versuch 3. Milchzucker.—1-10%ige Lösungen wurden ver-
tragen. Vacuolenvergrösserung von 3% an aufwärts und Kör-
perabrundung über 4%. Bei einer 7%igen Lösung wurde die
Vermehrung verzögert, und bei 10% konnten nur einige Indi-
1) Figuren in O. Bütschli. loc. cit. 1887-89. Abt. III. Taf. LXII.
EINIGER INFUSORIEN. 125
viduen 10 Tage lang leben. Auch hier fand mit der Concentra-
tionssteigerung Grössenzunahme des Körpers statt.
Versuch 4. Glycerin. —Maximalconcentration 5%. Vermeh-
rung und Vergrösserung der Vacuolen bei 2% ; Körperabrundung
bei 3%. Bei 5% lebten nur noch vereinzelte Individuen wenige
Tage lang.
Versuch 5. Schwefelsaures Magnesium.— Anpassungsconcen-
tration: 1-5%. Vacuolenvergrésserung und Körperabrundung
begannen bei 2%. Bei schwächeren Concentrationen gedieh der
Organismus gut, aber über 3% schlecht.
Versuch 6. Salpetersaures Kalium. —Maximalconcentration
2%. Das Thier gedieh bei 0,8%iger Lösung nicht mehr.
Zahlzunahme und Vergrösserung der Vacuolen waren wie ge-
wöhnlich.
Versuch 7. Salpetersaures Natrium.— Anpassungsconcentra-
tion: 0,2-2%. Ueber 0,8% nahm die Vermehrung stufenweise
ab. Gestaltänderung u. s. w. waren ähnlich wie in den vorher-
gehenden Fällen.
Versuch 8. Chlorkalium.— Anpassungsconcentration : 0,2-
1,6%. Die höheren Concentrationen über 0,8% verursachten
Multiplicationshemmung. Körperabrundung von 0,6% an.
Vacuolenvergrösserung fand auch bei stärkeren Lösungen statt.
Versuch 9. Chlornatrium.—Maximalconcentration 1,5%.
Volumenvergrösserung der Vacuolen wie gewöhnlich.
Versuch 10. Chlorammonium.—Der Organismus konnte sich
nur äusserst verdünnten Lösungen anpassen. Schon bei 0,2%
trat Vacuolenvergrösserung ein, bei 0,8% Bewegungshemmung
und Unregelmässigwerden der Körperumrisse. Bei 1%, der
höchsten Concentration, welcher das Thier widerstand, waren
einige Individuen nach 2 Tagen noch lebendig.
126 A. YASUDA : ANPASSUNGSFAEHIGKEIT
(e) Paramaecium caudatum Ehrbg.')
Merkmale des Organismus in der normalen Kultur : Körper
verlängert, spindelförmig, biegsam. Wimpern überall an der
Oberfläche des Körpers, dicht und gleichmässig. ‘Trichocysten
senkrecht zur Oberfläche in der unter der Cuticula unmittelbar
befindlichen Rindenschicht gelegen. Mund nahe der Mitte der
Bauchseite. Schlund ziemlich lang. Zwei contractile Vacuolen
am vorderen und hinteren Ende, mit strahligen zuführenden
Kanälen. Nahrungsvocuolen vorhanden. Bewegung lebhaft.
Vermehrung langsam.
Versuch. 1. Rohrzucker—Anpassungsconcentration 1-7%.
Von 3% an aufwärts bis 7% Zahl- und Durchmesserzunahme
der Vacuolen. Ueber 4% Dickwerden des Körpers; bei 7%
blieb das Thier noch viele Tage lang lebendig.
Versuch 2. Traubenzucker.—Anpassungsconcentration : 1-5%.
Vacuolenvergrôsserung bei ca. 2%, Kôrperabrundung bei 3%.
Sonst wie beim vorhergehenden Versuch.
Versuch 3. Melchzucker.—Maximalconcentration 8%. Ver-
mehrung und Vergrösserung der Vacuolen bei 3% u.s.w. In
höheren Concentrationen erreichten die Durchmesser der Vacuolen
bedeutend grössere Dimensionen, und der Körper erhielt ein
fleischiges Aussehen.
Versuch 4. Glycerin.—Anpassungsconcentration 1-3%. In
diesem Medium konnte das Versuchsinfusor nicht lange am Leben
bleiben. Vacuolenvergrösserung und Körperabrundung wie bei
den vorigen Versuchen.
Versuch 5. Schwefelsaures Magnesium.—Maximalconcentration
2,4%. Obgleich der Körper bei 0,2% verlängert war, so wurde
ı) Figuren in O. Bütschli. loc. cit. 1887-89, Abt. IH. Taf. LXITI.
EINIGER INFUSORIEN. 127
ei 2,4% viel fleischiger, wobei sich auch die Vacuolen
ten.
uch 6. Salpetersaures Kalium.—Anpassungsconcentration
Die durch dieses Medium hervorgebrachten Gestalt-
en waren fast dieselben wie die von Colpidium colpoda
]ben Medium.
uch 7. Salpelersaures Natrium.—Maximalconcentration
ickenzunahme des Körpers von ca. 0,7% an. In 1,2%
ng lebten nur noch vereinzelte Individuen mit schwacher
uch 8. Chlorkalium.—Aupassungsconcentration 0,2-1%.
1%-Kultur starb das Thier nach 3 Tagen gänzlich ab.
uch 9. Chlornatrium.—Anpassungsconcentration 0,2-1%.
1 mit Concentrationssteigerung auch Abrundung des
statt.
uch 10. Chlorammonium.—Maximalconcentration 0,5%.
sor accommodirte sich an dieses Medium am schwersten ;
Kultur blieb es lange am Leben.
Allgemeines und Schlussbemerkungen.
den oben angeführten Versuchen ergiebt sich, dass mit
erung der Concentration unabhängig von der chemischen
nheit die Cuticularoberfläche der Infusorienkörper ein-
t, wenn die Organismen plötzlich in das Medium
werden, weil durch concentrirtere Medien das Wasser
Thierkörper herausgezogen wird. Zugleich wird ihre
128 A. YASUDA : ANPASSUNGSFAEHIGKEIT
Bewegung, die bisher lebhaft gewesen war, immer langsamer, un
nach einem kurzdauernden Zittern an einem Platze kommen di
Thiere endlich zum Stillstande. Wenn aber die Concentratio
des Mediums nicht zu stark ist, so können es die Infusorie
ohne grossen Schaden ertragen, und die einmal gebildeten long’
tudinalen Falten der Cuticularoberfläche verschwinden nac
einiger Zeit wieder. Sogar bei concentrirteren Lösungen find
man nicht selten einige Individuen, welche mit der contrahirter
unebenen Körperoberfläche noch einige Tage lang fortlebe
können. |
Je höher die Concentrationen der Medien sind, dest
schwerer wird selbstverständlich die Anpassung, und wenn sic
schliesslich die Maximumgrenze nähert, so stirbt der grössere The
der Individuen ab. Im Falle gelungener Anpassung an ei
gewisses Medium sieht man stets Volumen- sowie Zahlzunahn
der Chromatophoren, Amylumkörner und Nahrungsvacuole
Gleichzeitig nimmt der Körper selbst an Dicke und Breite z
dagegen an Länge etwas ab, so dass er ein einigermassen abgı
rundetes Aussehen erhält. Zugleich ist ausserdem Grössenzunahn
des ganzen Körpers wahrnehmbar, wie ich dies ausschliesslic
mit Zuckerarten bei Colpidium colpoda, Mallomonas Plosshi un
Chilomonas paramaecium nachgewiesen habe.
Als eine allgemeine Regel gilt auch, dass die Vermehrung:
fähigkeit bei höherer Concentration stark beeinträchtigt win
Unsere Versuche mit den Reinkulturen von Zuglena viridi
Chilomonas paramaecium und Colpidium colpoda bieten hierfü
unzweideutige Beweise dar. Zum Vergleich führe ich einige de
bei Schimmelpilzkulturen gewonnenen Erfahrungen an. Eschen
hagen') constatirte, dass das Wachsthum einiger Schimmelarte
1) F. Eschenhagen. loc. ci. p. 56.
EINIGER INFUSORIEN. 129
rch stärkere Concentrationen des Substrates stark ver-
; Klebs') beobachtete, dass das Auftreten der Konidien-
und die Perithecienbildung von Zurotium repens, die
g und die Sporangienbildung von Mucor racemosus durch
gerung der Concentration des Mediums retardirt wurden.
tlich fand ich’) auch, dass Aspergillus niger, der in Mag-
ulfat-Nährlösungen von verschiedener Concentration
t wurde, nach 4 Tagen verschiedene Grade der Ent-
ng zeigte. Der Pilz wuchs in einer 5%-Kultur am besten,
gut bei 10%, während bei 20% und 30% nur noch eine sehr
1e Entwickelung zu beobachten war. Die weisse Anlage
nidienfrüchte trat bei 5% und 10% nach 4 Tagen, bei
ch 5 Tagen und bei 30% erst nach 6 Tagen ein.
ter den zehn von mir angewendeten Stoffen— vier organi-
nd sechs unorganischen Verbindungen—passten unsere
ien sich den Zuckerarten am besten an, und wieder unter
ckerarten erwies sich der Milchzucker als das beste An-
smedium. Ihm folgt der Rohrzucker in seiner Concen-
höhe, während der Traubenzucker schon in weit verdünn-
ösungen auf die Organismen schädlich einwirkt. Glycerin
s Anpassungsmedium den Zuckerarten sehr nach. Unter
organischen Verbindungen, deren Einwirkung stets viel
ist als die der organischen Substanzen, ist schwefelsaures
ium zur Vermehrung der Infusorien am geeigneststen,
1 Chlorammonium für ihr Gedeihen das unpassendste
Klebs. Die Bedingungen der Fortpflanzung bei einigen Algen und Pilzen.
pp. 446-535,
Yasuda. Ueber den Einfluss verschiedener unorganischer Salze auf die Fort-
rgane von Aspergillus niger. The Botanical Magazine. Tokyo 1898. Vol. XII, No.
),
130 A. YASUDA : ANPASSUNGSFAEHIGKEIT
Medium ist, und unter den übrigen Chlorkalium eine mittlere
Stellung einnimmt. |
Die verschiedenen Infusorien zeigten in Bezug auf ihre
Anpassungsfähigkeit grosse Unterschiede, und zwar wohnte unter
unseren Infusorien Huglena viridis die grösste Widerstandsfahig-
keit inne, während Paramaecium caudatum die kleinste Resi-
stenz besass. Folgende Tabelle zeigt die Grenze der Concentra-
tionen, bis zu welcher die Infusorien am Leben blieben :
amt
5 | = | ke 3 | 4
“| K | 3 ale ale ula B| 2/8
Stoff Sels) BER EEE a |88
Gm ger ~~ EL
™ | a) a| 2) ÉlSRSlS 2] § 2"
Al) SSH)
Sald'élsls sis) sls
Formeln am | LÀ sm OS le © |
| © sel 20 e & N 7, fr
ge oye te Z zZ
Concentrationen der % % % % % % ar, él 6g
mit 0,1 Aeq. KNO, | 5,40 | 5,13 | 2,70 1,38 1,80 | 1,01 | 0,85 | 0,75 | 0,59 | 0,54
isotonischen Lösungen’ ) |
ee | | se | nes. | ans | nes — —— FA
a à Euglena
2: on, \17 |15 jı |6 |6 124 |2 28 |18 |1,4
CE ei ee el ES eee
a | Chilomonas
i 3 paramae- | 8 | 7 | 6 | |3 |2 [12/2 |1 [0,6
D cium
5 — |} —
= ® | Mallomonas
Be | po | 9 | 7 | 6 ja [34 [15 2,6 114 |1,5 108
D à
à 4 Fea es eee de ee. 1
D & Colpidium
rE "10 |8|7|5 |5 |2 |2 (16 [15 |1
= .2 —— [ll
Ko Paramae-
SS cum | 8 | 7 | 5 |3 124 |1 1,2 1 |1 [05
*& caudatum
1) Hugo de Vries. Eine Methode zur Analyse der Turgorkraft. Jahrb. f. wiss. Bot.
1884. Bd. XIV. pp. 536—537.
pen Om es
„AT ee ee
EINIGER INFUSORIEN. 131
Wie aus der vorstehenden Tabelle ersichtlich ist, muss die
Wirkung der angewandten Substanzen auf die Infusorien nicht
allein dem Grade ihrer Concentration zugeschrieben werden,
dagegen zeigt sich ein annäherndes Verhältniss zu den isotoni-
schen Concentrationen jedes Stoffes. Ich sage ausdrücklich ,, an-
nähernd,‘‘ weil die wahre Beziehung zwischen beiden durch unsere
Versuche noch nicht sicher gestellt ist. Folgende Tabelle
dient zum Vergleiche der isotonischen Concentrationen mit den
entsprechenden gefundenen Werthen der maximalen Anpas-
sungsconcentrationen') :
1) Die bisherigen Untersuchungen ergaben in Bezug auf den Zusammenhang zwischen
isotonischen Concentrationen und Reaktionsgrösse durchaus negative Resultate. Man ver-
gleiche hierüber B. Stange. loc cit. Nr. 22. p. 364, und C. B. Davenport and H. V.
Neal. loc. cit. p. 579.
132 A. YASUDA ! ANPASSUNGSFAEHIGKEIT
|
be | | =
ns le | £ LAN Ble |S
sı8|<|= 18:22, 2.3218
s|&|ı8 |: |i2|38135:|3 | 8 | 2
= - = LR cé | a
Stoffe. RI8|2|32 28 52 E2 82|2|38
= a 5 > |\sh|se|/3s| % pe =
= = = - ea | =} Rie | = a =
= Le 3 | © |4e/4 | Br =
Ka PL Dr | =
Fi = L i , ac D =
| | D)
Covcentrationen der
Stoffe, die mit 15%5 F Fo
Robrzucker iru hax 1 J 1 ad 5
2 .S nisch sind
We mes —
fx Gefundene Werthe
der maximalen 15 17
Anpassungscolioen-
tration
por centra tone der
toffe, die mit 7°5
3 5 Robrzucker isolo- 7 7,4
Il = g nisch sind .
= | mm | 00000000 ln
= —————
= 8 | Gefundene Werthe
a 2 der maximalen T g
& Anpassuugsconoen- |
tration
Stoffe, die mit 7%
Rohrzucker inotö- 7 7,4 3 Ay 4 1,9 2,0
nisch sind |
Gefundene Werthe
der maximalen
Anpassungsconcen-
tration |
Mallomonas
Piosslii
|
|
|
|
Concentrationeu der
Stoffe, die mit 8%
Rohrzucker isoto-
3 nisch sind | |
3 Gefundene Werthe | | et
der maximalen 8 10 |
| Aupassungsconcen- |
| tration
Concentration der
Colpidium
Concentrationen der
Stoffe, die mit 7%
Robrzucker isoto- 7 7,4 3,1 A!
nisch sind
Gefundene Werthe
der maximalen
Paramaecium
caudatum
Anpassungsconcen- 7 8
tration
Dieselbe Tabelle zeigt auch zugleich, dass die Anpassungs-
grenzen unserer Infusorien an verschiedene Concentrationen im
Allgemeinen weit niedriger sind als diejenige der niederen Algen
und Schimmelpilze. So kann Zygnema nach Klebs') 50% Rohr-
1) G. Klebs. Beitrige mr Physiologie der Pflanzenzelle. Berichte der deutsch bot.
Gesellsch. 1887. Bd. V. p. 187.
EINIGER INFUSORIEN. 133
zucker und 20% Glycerin vertragen. Dass dieselbe Alge sich
auch einer 6%igen Chlornatriumlösung anpassen kann, ist von
Richter’) erwiesen worden. Was die Schimmelpilze anbelangt,
so zeigen sie ebenfalls eine weitaus grössere Widerstandsfähigkeit
gegen starke Concentrationen. Ich gebe hier zum Vergleiche die
von Eschenhagen?) erhaltenen Ergebnisse wieder.
Salpetersaures
Traubenzucker. Glycerin. Wat ann Chlornatrium.
Aspergillus niger 53% 43% 21% 17%
Penicillium glaucum 55,, 43,, 21; 18 ,,
Botrytis cinerea Sl, 37 ,, 16,, 12 ,,
Unsere Infusorien zeigen gegen dieselben Stoffe folgendes Ver-
halten :
Salpetersaures
Traubenzucker. Glycerin. Stumm: Chlornatrium.
Euglena viridis 11% 6 % 2 % 1,8%
Colpidium colpoda T oA; D... 1,5 ,,
Mallomonas Plosslii 6 , 4 ,, 2,6 , 1,5 ;,
Chilomonas paramaecium 6 ,, 4 „ 12,, EE
Paramaecium caudatum 5 ,, 3:5; 1,2, LÉ. 3;
Daraus geht hervor, dass die Resistenzkraft unserer Infusorien
gegen die angewendeten Stoffe hinter derjenigen der Schimmel-
pilze weit zurücksteht.
1) A. Richter. loc. cit. p. 24.
2) F. Eschenhagen. loc. cit. p. 55.
134 A. YASUDA : ANPASSUNGSFAEHIGKEIT
Zusammenfassung.
(1) Isotonische Lösungen der in Rede stehenden Substanzen
üben auf die von mir geprüften Infusorien nur eine ,, anni-
hernd‘“ gleichartige Wirkung aus.
(2) Die Grenzen der Concentration, welcher sich diese
Infusorien unter gewöhnlichen Verhältnissen anpassen können,
liegen im Allgemeinen weit niedriger als die der niederen Algen
und Schimmelpilze; selbst das widerstandsfähigste darunter,
Euglena viridis, vermag nur verhältnissmässig schwache Concen-
trationen zu ertragen.
(3) Wenn die Organismen plötzlich in Lösungen höherer
Concentrationen gebracht werden, so treten erst an der Cuticular-
oberfläche ihrer Körper longitudinale Falten auf, aber während
ihre Anpassung an das neue Medium stattfindet, dehnen sich die
Falten allmählich aus, bis sie zuletzt gänzlich verschwinden.
(4) Die höhere Concentration des Mediums verlangsamt die
Vermehrung der Infusorien.
(5) Durch Steigerung der Concentration des Mediums wird
die Bewegung der Organismen vielfach retardirt.
(6) Bei Zuckerlösungen stärkerer Concentration vergrössern
sich die Körper der Infusorien bis zu einem gewissen Grade.
(7) Die Vacuolen, Chromatophoren oder Amylumkörner
nehmen in dem Masse an Grösse zu, als die Mediumsconcentra-
tion steigt.
(8) Je mehr die Concentration des Mediums zunimmt, desto
mehr runden sich die Körper der Organismen ab, und die Kör-
perumrisse werden uneben.
(9) Wenn das Maximum für die Accommodation ein nie-
hi
kin:
0 Ap
darutt.
Cate.
hie
KS
EINIGER INFUSORIEN. 135
driges ist, so finden die Veränderungen der Körper der Infusorien
schon bei niederen Concentrationen des Mediums statt.
(10) Wenn sich die Concentration des Mediums dem Maxi-
mumpunkt nähert, so verschmelzen die in den Körpern der
Organismen befindlichen Chromatophoren oder Amylumkörner
mehr oder weniger mit einander.
Tokyo, 30. November 1898.
Inhaltsübersicht.
Einleitung unse inessnsass deasentestesis Seite 101.
Litteratur ..... See „ 10.
Methodisches ......-.sssscsosccccesscscccsccscscecsesceses » 107.
Beschreibung der Versuche ...........cscescocesses je. REZ
Allgemeines und Schlussbemerkungen ......... 55. 127.
Zusammenfassung ..............…. ET » 134.
CRT SELS
A. YASUDA : ANPASSUNGSFAEHIGKEIT
Erklärung der Figuren.
Sämmtliche Figuren wurden nach den lebendigen Thieren in
den unreinen Kulturen unmittelbar skizzirt, weil ihre Gestalten
bei den getödteten Individuen sich mehr oder weniger veränderten.
Kr,
Fig. 1-46.
1. Individuen aus einer Nährlösung mit 1
6.207 ED
bo wo WN ND YD FY bn bd bed bd bed bd be u Fi
Pen SHON Ae PWN = ©
9)
TAFEL X.
Chilomonas paramaecium Ehrbg. Vergr. 420.
29
22
„
°2
2
3
D
me OO y oo «À
to
D D ei OO Gr Ff WO RD = I oo À RP W
% Milchzucker.
29 7
29 2)
29 ”?
LE 2)
29 1
„ 33
3) 2?
22 ?
3? 1
„ 29
9) 2)
a9 „
29 2?
39 „
7) „
2 2)
29 27
„ 2)
% Glycerin.
99 39
EINIGER INFUSORIEN. 137
ig. 25. Individuen aus einer Nährlösung mit 4 9% Glycerin.
: WR: + ne 5s „ 0,5% Schwefelsaures
Magnesium,
33 27. #7 iy HE LL 3! 1 2? „
» 28. 1, 5 1 ” » 1,9, ”
cay te = See Ah 7 2. 2 ï
„ 30. ” ” ” ” ” 2,5, "
» Sl ” ” ” ” ” 34, LE
5 3. Re me abe : » 0,59 Salpetersaures
Kalium.
5 3. 2 oy te + 2.1. 5; *
„ 34 À US 5 3 15, +
„3. 2) „9 ” » 2 » LE
sis ‘i sl we = » 0.69% Salpetersaures
Natrium.
73 37 39 LE LL LE EL 1,2 „ 33
5 38 ri to Ai 7 „ 0,5% Chlorkalinm.
7 39 ” iY LE LE LE 1 9
„ 40 ae 93 a. 1.05; 7
4 ag tes r- DE r
» 42. iz a. à 7 , 0,5% Chlornatrium.
” 43 LE} LL LE 11 31 1 „ 3?
» 44 + is + js „ 0,2% Chlorammonium.
ar 45 31 31 31 33 LE 0,4 2) 31
„ 46 ” ” s! ” ” 0,6,, "
TAFEL XI.
Fig. 1-11. Zuglena viridis Ehrbg. Vergr. 420.
‘ig. 1. Individuum aus einer Nährlösung mit 1 9 Traubenzucker.
7 2. a3 37 3} 33 LE 2 29 LE
ay 3. LE a) LE LE P LL 3 „ LE à
er 4. LE LE FE LE) LE ; 4 „ ’
Digitized by
A. YASUDA : ANPASSUNGSFAEHIGKEIT
Fig. 5. Individuum aus einer Nährlösung mit 5 % Traubenzucker.
”" 6. „ ” „ CE „ 6 CE Te)
# 7. ”? 99 „ LE 29 7 33 ii
8. ” ” ” ” ” 8 4; „
T J, ” ” ” ” 5 0 #7
» 10 „ ” ” ” „10, ”
» 11. „ „ ” ” ie EG L
Fig. 12-41. Colpidium colpoda Ehrhg. Vergr. 420.
‘ie, 12, Individuum aus einer Nährlösung mit 95 Milchzucker.
7
123, ” „ ”? „ „
ww DD =
1) >?
L 14. „ ” ” „ ” CE ’
,, In. „ „ „ ” ” 4 CE ”
rt 6, F 22 29 „ „ 99 5) LE. sr
ye 3 „ 9? ” 29 6 ’ 13
vs 15, „ ” ” „ ” 1 CE
1) 19, ” „ „ ” ” 8 ” ss
Fe „ „ „ ” „ I El „
CE 21. ” „ ” ” ” 10 ” ”
we: nee a oe ‘5 » 1 % Rohrzucker.
» oe „ „ ” ” „ 2 „ ”
„ 24 „ ” ” ” ” 3 ” E
,, 20 ” ” „ ” „ 4 ’ 7
„ 20 ” ” ” ” ” 5) " ”
GS a „ ” ” ” ” 6 T ”
' 28. ” ” ” ” ” 7 „ 1)
mn 29 ” ” „ ” ” 8 El E
„30. ‘5 je u "4 „ 1 % Traubenzucker.
iy 51. 39 2) 32 2) „ 2 HE 1}
„ 2, ” ” ”? „ „ 3 'E CE
T 3. „ ” ” „ ” 4 ” "2
L 34. „ ” ” ” „ 6) ” ”
39. ” „ „ „ „ 6 ” '
EINIGER INFUSORTEN. 139
Fig. 36. Individuum aus einer Nährlösung mit 7 9 Traubenzucker.
yy le (y as on ii „ 1 9% Glycerin.
LL 38, LL " [1] pr LE 2 LL LL
0 à FRET “3 <a a
» 4 er wi. . > eg:
» 4 “ : 7 . ag >
TAFEL XII.
Fig, 1-21, Mallomonas Plosslit Perty. Vergr. 420,
Fig. 1. Individuen ans einer Nährlösung mit 1 9% Milchzucker,
LL 2, 7 > ” ” be 2 CL ”
re 3, ” LL ” ” LE 3 ” ”
rs 4 »? LL LL) " Fr 4 " ’
(2) 5 " st CE] 12) LE 2 ” ”
»? 6, pr LL ” LE ay fj LA] "”
$3 7. " LE i} | LL ul 7 9 LE
12 8. 7 93 3 ’ CE 8 ” s
ER) J, „ LE ” ” 9 " LL
“20, ss A an à „ 1 % Rohrzucker,
” 11, ” 7 [L ” ” 2 ”" ts
” 12, 4 " " " "y 3 " "
7 13, 2? " iy LE LIL 4 EL) LE
” 14, ” ” rs ” ” ” CL)
LE 15. ” LL C1} 1] si 6 1 rm
LL 16. LL ” ” #1 CE 7 „ 1
ad Wie à sham 5 Æ „ 1 % Traubenzucker,
as 18. ” 9 " „2, ”
neh, 19. LL ” ” rs CE 3 ” CE
À 4
Digitized by Ga OQIE
Fig. 22-28. Paramaecium caudatum Ehrbg. Verg. 240.
Fig. 22. Individuum aus einer Nährlösung mit 1 9 Rohrzucker.
39 23. „ 39 3 29 29 2 29 LE
32 24, 29 9) 9? 99 3 3 LE 3)
29 25. 2) 3) „ 72 2 4 „ „»
7) 26. 32 9) ” ” 3 4) 2 LE
» 27. » „ 22 2) 2) 6 I) 3
LE 28. „ 2 9) » 29 7 „ 23
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Ueber die Wachsthumsbeschleunigung einiger
Algen und Pilze durch chemische Reize.
VON
N. Ono, Rigakushi.
Hierzu Tafel XIII.
I. Einleitung und Litteratur.
In seiner Arbeit: Etudes chimiques sur la végétation'), hat
Raulin schon im Jahre 1869 darauf aufmerksam gemacht, dass
Zink- und Siliciumsalze in geeigneter Dosis das Wachsthum von
Aspergillus niger befördern. Auf dieser an sich richtigen Beo-
bachtung fussend war der genannte Autor mit Unrecht der
Ansicht geneigt, dass diese Substanzen zur normalen Entwickelung
unseres Pilzes nothwendig seien, indem er diese unter ,, les
éléments chimiques essentiels‘ rechnet und eine Nährlösung von
recht complicirter Zusammensetzung für Pilze vorschreibt.
Ueber die Mineralstoffbedürfnisse der Pilze wurden seither
von einigen Forschern Untersuchungen gemacht, von denen die
Arbeit Naegeli’s’) in der ersten Linie zu nennen ist. In
1) Ann. d. Se. nat. Bot., Ser. V, T. XI, 1869, 8. 91.
2) v. Naegeli, Der Ernährungschemismus der niederen Pilze. Sitzungsberichte d. Kgl.
Bayr. Akad. d. Wiss. Math.-phys. Cl. 1880.
142 N. ONO: WACHSTHUMSBESCHLEUNIGUNG
neuerer Zeit wurde dies Thema von Molisch'), sowie auch
Benecke’) wiederaufgenommen. Der Mühe dieser Aut
verdanken wir unsere heutigen Kenntnisse in dieser Richt
Nach den übereinstimmenden Angaben der genannten Aut
stellt weder Zink noch Silicium einen eigentlichen Nihrstoff
und kann wohl von Kulturflüssigkeit ausgeschlossen werden
Dass die wachsthumsbeschleunigende Wirkung gew
Metallradikale auf einer chemischen Reizung beruht, wurde
Pfeffer‘) in seiner im Jahre 1895 erschienenen Arbeit
ersten Male klar gestellt und später in allgemeinen Zügen i1
2" Auflage seiner Pflanzenphysiologie erörtert. Er bemerl
dem letztgenannten Werke, dass anscheinend geringfügige
stünde in der That nicht selten einen erheblichen Einfluss
Gredeihen und Wachsen haben, und sagt: ,, Vermuthlich ha:
es sich in dieser beschleunigenden Reizwirkung um eine
mannigfachen Reaktionen, die darauf abzielen, durch intensit
Thätigkeit einen benachtheiligten Einfluss thunlichst entge
zuarbeiten oder Schädigungen auszugleichen.‘“) Im vorherge
senen Jahre wurde eine Reihe Versuche von Richards’) anges
deren Resultate die Ansicht Pfeffer’s bestätigen. Er zog zu se
Untersuchungen verschiedene Schwerenmetallsalze wie Z
Kobalt-, Nickel-, Eisen-, und Mangansalze und einige ar
giftige Substanzen heran und stellte die Thatsache fest, dass
1) H. Molisch, Die mineralische Nahrung der niederen Pilze. Sitzungsberi
Wiener Akad. Oct. 1894.
2) W. Beneeke, Die zur Ernährung der Schimmelpilze notwendigen Metalle. P
Jahrb. f wiss. Bot, Bd. NXVIII, 1895, S. 487.
3) Pfeffer, Election organischer Nährstoffe. Pringsh. Jahrb. f. wiss. Bot. Bd. X.
1340.
4) Pilanzenphysiologie. 2 Aufl. Bd. I. S. 374.
5) H. M. Richards, Die Beeinflussung des Wachsthums einiger Pilze durch che
Reise, Prinwsh, Jahrh, & wiss. Bot. Bd. XXX, 1897, S. 665.
EINIGER ALGEN UND PILZE. 143
alle geprüften Substanzen mehr oder weniger die Pilzernte zu
vermehren vermochten.
Auch anderweitige Beispiele für Erhöhung der Lebens-
thätigkeiten durch geringe Zusätze giftiger Stoffe findet man in
der Litteratur. So versuchte Schulz') zu zeigen, dass die durch
Saccharomyces verursachte alkoholische Gährung bei Gegenwart
von geringeren (Juantitäten gewöhnlich als Hefegifte sich verhal-
tender Substanzen wie Sublimat, Jod, Salicylsäure, Brom, Arse-
nige Säure u. a., auf längere oder kürzere Zeit wesentlich gehoben
werden konnte. Diese Thatsache aber war bereits gewissen
Gewerben nicht unbekannt gewesen, dass Zuführung von sonst
gihrungshemmenden Stoffe, wie z. B. Kupfervitriol oder Sali-
eylsäure, unter Umständen die Hefe zu energischer Thitigkeit
veranlasse. Derselbe Autor hat früher eine ähnliche Erhebung
der Lebensäusserungen bei dem thierischen Organismus beobachtet
und folgenden Satz ausgesprochen, dass ‚Jeder Reiz auf eine
einzelne Zelle sowohl wie auch auf aus Zellengruppen bestehenden
Organen entweder eine Vermehrung oder eine Verminderung
ihrer physiologischen Leistungen bedinge, entsprechend der
geringeren oder grösseren Intensität des Reizes‘“*).
Derartigen Verallgemeinerungen begegnen wir auch bei
Hueppe?). Hauptsächlich auf die auf bakteriologischem Gebiete
constatirten Thatsache sich stützend verkündigt er die Erschein-
ung als den Ausdruck des allgemein giltigen Gesetzes für die
Wirkung von Chemikalien auf Protoplasma. Er bezeichnet dieses
als das ,, Biologische Grundgesetz,‘‘ welches er in folgendem
1) H. Schulz, Ueber Hefegifte. Pfliiger’s Archiv f. Physiologie. Bd. 42., 1888. 8. 517.
2) H. Schulz, Zur Lehre von der Arzneiwirkung. Virchow’s Archiv. Bd. 108, 1877.
8. 427.
3) F. Hueppe, Naturwissenschaftliche Einführung in die Bakteriologie. 1896, Wies-
baden. S. 55.
144 N. ONO! WACHSTHUMSBESCHLEUNIGUNG
Satze formulirt: „Jeder Körper, der in bestimmter Concentration
Protoplasma tötet und vernichtet, in geringeren Mengen die Ent-
wickelungsfähigkeit aufhebt, aber in noch geringeren Mengen,
jenseits eines Indifferenzpunktes, umgekehrt als Reiz wirkt und
die Lebenseigenschaften erhöht.“
Zu erwähnen ist noch, dass auch bei höheren Pflanzen die-
selbe oder eine wenigstens sehr nahe verwandte Erscheinung
vorkommt. Bekanntlich pflegt man seit einigen Jahren die
Weinreben mit Kupferpräparaten, der sogenannten Bordeaux-
brühe, zur Bekämpfung der Pilzkrankheit zu bespritzen. Eine
derartige Behandlung ruft ausser der indirekten Einwirkung, die
Schädigung durch parasitische Pilze herabzusetzen, auch eine Schar
auffallender Erscheinungen seitens der bespritzten Pflanze hervor,
die mit Rumm!') vielmehr als direkte Wirkung der angewandten
Chemikalien auf den Pflanzenorganismus selbst zu bezeichnen
sind. Solche sind die Steigerung der Chlorophylibildung und
daraus resultirende vermehrte Stärkeproduktion, reichlicherer
Traubenansatz, Beschleunigung der Reifung u. a. Rumm ist der
Ansicht, dass die Steigerung der Chlorophylibildung einem
chemischen Reiz zuzuschreiben sei. Im darauf folgenden Jahre
führten Frank und Krüger”) einige Bespritzungsversuche an
Kartoffeln aus, wobei sie auch eine ähnliche Thatsache consta-
tirten. Ueber das eigentliche Wesen der Wirksamkeit jener
Stoffe ist vorläufig nichts weiteres zu sagen.
Wie aus der oben angeführten Skizze ersichtlich ist, bezogen
1) Rumm, Ueber die Wirkung der Kupferpräparate bei Bekämpfung der sogenannten
Blattkrankheit der Weinrebe. Ber. d. deutsch. Bot. Ges. Bd. XI. 1893. S. 709.
Rumm, Zur Frage nach der Wirkung der Kupfer-Kalksalze bei Bekämpfung der
Peronospora riticola. ebenda S. 445.
2) Frank u. Krüger, Ueber den Reiz, welchen die Behandlung mit Kupfer auf die
Kartoffel hervorruft. Ber. d. deutsch. Bot. Ges. Bd. XII, 1894.
EINIGER ALGEN UND PILZE. 145
sich, wenn wir von der zuletzt besprochenen Thatsache absehen, alle
bisherigen diesbezüglichen Versuche mit pflanzlichen Organism-
en ausschliesslich auf chlorophyllose niedere Organismen der
Pilze. Wenn die erwähnte Erscheinung, wie vielerseits behauptet
wird, allgemeine Geltung haben würde, so sollte es a priori zu
erwarten sein, dass auch chlorophyllhaltige niedere Organismen
in gleichem Sinne reagiren. Dies aber benötigt einer experimen-
tellen Bestätigung.
Herbeigezogen wurden zu meiner Untersuchung verschiedene
Schwerenmetallsalze, wie Zink-, Nickel-, Kobaltsulfat u. a., welche
auf unsere Versuchsalgen gewisse Reizung auszuüben vermochten.
Ferner wurden eine Reihe Parallelversuche mit Pilzen angestellt,
deren Ergebnisse die oben genaunten Richards'sche Unter-
suchung bestätigt und erweitert.
Die vorliegende Arbeit wurde auf Veranlassung und unter
Leitung von Herrn Prof. Dr. Miyoshi im Botanischen Institut
der Kaiserlichen Universität zu Tokyo während des Zeitraums
von August 1898 bis Juni 1899 ausgeführt.
Es ist mir eine angenehme Pflicht, meinem hochverehrten
Lehrer für die vielseitige Anregung meinen verbindlichsten Dank
auszusprechen.
Herrn Prof. Dr. Matsumura sage ich an dieser Stelle auch
meinen besten Dank für die Belehrung und das Interesse, welches
er meiner Arbeit entgegengebracht hat.
II. Methodisches.
Bei unseren Versuchen kommen stets die Reinkulturen in
Betracht. Als Kulturgefässe wurden Erlenmeyer’sche Kolben
146 N. ONO: WACHSTHUMSBESCHLEUNIGUNG
von ca 200 cc Inhalt angewendet, und zwar von gleicher Gestalt
und Qualität in einer Reihe von Parallelversuchen benutzt. Nach-
dem die Gefässe zuerst mit Salzsäure gründlich gewaschen worden
waren, wurden sie mit Leitungswasser, dann mit destillirtem
Wasser wiederholt ausgewaschen, getrocknet und gebraucht.
Wasser.—Zur Zubereitung der Nährlösungen und zum Auf-
lösen der Reizmittel benutzte ich doppeltdestillirtes Wasser.
Das auf übliche Weise gewonnene destillirte Wasser war von
Glas zu Glas nochmals destillirt worden.
Chemische Präparate. —Die als Nährstoffe sowohl als
auch als Reizstoffe dienenden Chemikalien stammten grössten-
theils aus Merck’s ,, garantirt reinen‘ Reagentien.
Nährlösungen.—Für Algen bediente ich mich der bekann-
ten Knop’schen Lösung‘), die ich nach folgender Vorschrift
bereitete:
(A) MgSO,+7H,O 10.255 (B) Ca(NO;), 20.008
KNO, 8.00 ,,
KH,PO, 5.00,,
Wasser 175.00cc Wasser 175.00 ce
Beim Gebrauch wurden je 10 cc von A und B mit 880 cc
Wasser verdünnt. Diese bezeichne ich als Original-Nährlösung
für Algen.
Die Original-Lösungen für Pilzkulturen bestanden aus fol-
genden drei Serien:
1) Aus keinem besonderen Grunde benutzte ich hier Ca-haltige Nihrlésung. Die Ent-
behrlichkeit der genannten Metalle bei Pilzen und niederen Algen ist bekanntlich in
neuerer Zeit von Molisch und Benecke erwiesen worden.
EINIGER ALGEN UND PILZE. ; 147
(A)
KH,PO, 0508
MgSO, 0.25 ,,
NH,NO, 1.00 ,,
Eisen Spuren
Rohrzucker 5.00 g
Wasser 90.00 CC.
(B)
Wie bei A. Asparagin 0.ög statt NH,NO, 1.0 g
(C)
Wie bei A. Dextrose anstatt Rohrzucker.
Bei fast allen Kulturreihen wurde A angewendet, während
B und C nur ausnahmsweise benutzt wurden.
Kulturanstellung.—Ich goss in 5Kolben je 135cc Original-
Lösung (Knop’sche bezw. Rohrzuckernährlösung). Sodann setzte
ich zum ersten Kolben 15 cc destillirtes Wasser hinzu und liess
dies als Nährlösung für Controlkulturen dienen, zum 2°, 3°...
5" je 15 cc betreffend verdünnte Lösung von Reizstoffen, deren
Wirkungen versucht werden sollten. Bei den Algenkulturen
geschah die Verdünnung in absteigender geometrischer Reihe im
Verhältniss 1:5, bei den Pilzkulturen aber mit 1:2. Alle Kolben
wurden darauf gut geschüttelt, um die Lösungen aufs innigste
zu mischen, und dann die in jedem Kolben enthaltene Lösung in
drei Kolben gleichmässig vertheilt, so dass wir 3 Serien von
je 5 Kolben, deren jede 50 ce Nährflüssigkeit enthielt, vor
uns haben.
148 N. ONO: WACHSTHUMSBESCHLEUNIGUNG
Die auf diese Weise zubereitete Kulturflüssigkeit enthielt
bei der Knop’sche Lösung ca 2.5% wasserfreie Salze und bei der
Pilznährlösung etwa 5% Rohrzucker'). Dann folgte bei Pilzkul-
turen die Sterilisation in einem Koch’schen Dampftopf, welche
l/—1 Stunde dauerte.
Bei Pilzen fand die Impfung in üblicher Weise statt, wäh-
rend ich sie bei Algen in der Weise ausführte, dass ich mittelst
Platindraht oder Pipette eine möglichst kleine Algenmenge aus
den zuvor in Nährlösung von derselben Concentration oder auf
Agar bereiteten Reinkulturen herausnahm und in Versuchsgefässe
brachte.
Die Kulturen wurden in Zimmertemperatur (ca 15°C im
Mittel) ausgeführt, und in kälteren Jahreszeiten ins Treibhaus
(16-21° C) gebracht.
Die Kulturdauer variirte unter Umstinden zumeist zwischen
8 und 25 Tagen bei Pilzen und etwa einem Monatlang bei Algen.
Bestimmung des Trockengewichtes.—Fiir die Beurthei-
lung des Gedeihens hat fast stets die Ermittelung des Trockenge-
wichtes der gebildeten Algen- bezw. Pilzmassen Aufschluss
gegeben. Die Bestimmung wurde folgendermassen ausgeführt.
Nach Beendigung der Versuche wurde die Kulturflüssigkeit mit der
Erntemasse insgesammt durch vorher einzeln gewogene Filter
filtrirt. Dabei befreitete ich den an der Glaswand haftenden
Theil mittelst eines mit einem Kautschuk-Hut versehenen Glas-
stibchens. Dann spülte ich die Erntemasse mit kaltem destillirtem
Wasser, um dadurch etwa noch vorhandene Nährflüssigkeit
möglichst zu entfernen, und wenn sie ziemlich lufttrocken ge-
worden war, trocknete ich sie im Paraffinofen bei 100° C und
wog sie nach dem Erkalten. Das auf diese Weise ermittelte
1) Diese Lösung wurde von Pfeffer und Richards vielfach benutzt.
EINIGER ALGEN UND PILZE. 149
Trockengewicht der Ernte ist in den angeführten Tabellen als
Ernteertrag notirt. |
Bei den Algenkulturen geschah es vielfach, zumal bei
denjenigen, welche während der kälteren Jahreszeit angestellt
worden waren, dass die Vermehrung nur sehr langsam vor sich
ging, und dass nach monatelangem Stehenbleiben eine nur
schwache Entwickelung sich zeigte. In solchen Fällen musste
ich mich damit begnügen, durch das Aussehen der Kulturen die
Stärke der Entwickelung zu beurtheilen.
Was nun die specielle Ausführungsmethode anbelangt, so
wird sie an geeigneten Stellen berücksichtigt.
III. Vorbemerkungen uber Versuchsobjekte.
Für Pilzkulturen bediente ich mich der gewöhnlichen
Schimmelpilze Aspergillus niger und Penicillium glaucum.
Benutzt wurden bei meinen Algenversuchen die folgenden
Formen :
Protococcus sp.
Chroococcum sp.
Hormidium nitens.
Stigeoclonium sp.
Da wir zur Zeit über die Lebensbedingungen der Algen
überhaupt nur wenig wissen, so war es mir nicht immer ge-
lungen, die im Freien rasch wachsenden Algenarten im Labora-
torium unbeschädigt gedeihen zu lassen. Besonders schwierig war
die Aufgabe, die grösseren Formen in reinem Zustande längere
Zeit in einer bestimmten Nährflüssigkeit zu kultiviren.
Nach einigen darauf bezüglichen Vorversuchen kam ich
150 N. ONO: WACHSTHUMSBESCHLEUNIGUNG
schliesslich auf die oben genannten niederen Formen zurück, die
ziemlich leicht rein zu erhalten waren, ausserdem sicheres Gedeihen
zeigten und ferner leichtere Kontrolle der Impfmasse gestatteten.
Unsere kleineren Algen waren zumeist in den für grössere
Algen bezweckten Kulturgefässen spontan aufgetreten, und so
wurden diese durch wiederholte Uebertragung rein gezüchtet. Die
ersteren Algen liessen sich auch auf festem Nährboden, welcher
aus '/; Proc. Agar und 2.5 Promille Nährsalz enthaltender
Gallerte bestand, gut vegetiren und solcher Plattenkultur bediente
ich mich bei einigen Beimpfungen.
Hormidium nitens, welches sich auf der Oberfläche einer
Vaucheria-Kultur als eine charakteristische seidenglänzende
Decke bildete, zerfiel nach dem Uebertragen in unsere Nähr-
lösung in einzelne Zellen und vegetirte als solche weiter.
Es bleibt noch zu bemerken übrig, dass die Vermehrung
bei niederen Algen (Protococcus wurde zunächst untersucht)
während der kälteren Jahreszeit so gut wie vollständig herab-
gesetzt worden ist, wenn auch die Kulturen im Treibhaus bei
16-20° C sich befanden. Die Ursache wolle man nicht in man-
gelndem Licht suchen, da dieselben Kulturen mit noch mässigerem
Lichtgenuss vor einem gegen Norden gerichteten Fenster eine
gute Entwickelung bis Anfang November zeigten. Ob es sich
hier etwa um eine Vegetationsperiodicität handelt, beabsichtige ich
im kommenden Jahre näher zu untersuchen.
IV. Die Veranderungen in der Wachsthumsweise
und die Correlation zwischen Fortpflanzung
und Wachsthum.
Wie wird die Wachsthumsweise einer Pflanze beeinflusst
werden, wenn ihr Wachsthum bei Gegenwart von Reizstoffen
EINIGER ALGEN UND PILZE. 151
über die Norm hinaus gesteigert wird? Um diese Frage zu
beantworten, wurden bei meinen Untersuchungen einige Beobach-
tungen gemacht, um dabei auftretende Wachsthumsmodificationen
zu kennzeichnen.
Bei von mir untersuchten Algen konnte ich keine bemer-
kenswerthe Veränderung der Wachsthumsweise beobachten. Die
Zellengrösse blieb unverändert ; so lag z. B. bei Protococcus sp.
die Zellengrösse jedenfalls zwischen 7-10 x. Sie zeigte ferner
keinen Unterschied in Amylumeinschlüssen.
Bei Pilzkulturen liess sich aber die Veränderung in der
Wachsthumsweise mehr oder weniger schon makroskopisch er-
kennen. So war bei den meisten versetzten Kulturen die
Beschaffenheit des Mycels ungewöhnlich. Während bei Kontroll-
kulturen die Hyphe in Nährlösungen durchsichtig und zart
waren, bildeten diejenigen der versetzten Kulturen ein dickes,
weisses, hautartiges Geflecht, welches bei längerem Stehen sich zu
einer aufgerollten Masse umgestaltete.
Auch das Turgorverhältnis in den versetzten und den nicht
versetzten Kulturen wurde vielfach studirt, um zu ersehen, ob
hier etwa ein nennenswerther Unterschied zwischen beiden vor-
handen ist. Meine Versuche ergaben in diesem Punkte kein
positives Resultat.
Viel auffallender sind die Beziehungen zwischen Fortpflan-
zung und Wachsthum.
Bekanntlich stellen das Wachsthum und die Fortpflanzung
zwei miteinander in engster Wechselbeziehung stehende Lebens-
thitigkeiten dar. So kommt es nicht selten vor, dass bei einer
Pflanze, die in ippigem Wachsthum begriffen ist, ihre Fortpflan-
zungsfähigkeit zeitweilig suspendirt wird ; und wenn hingegen die
152 N. ONO: WACHSTHUMSBESCHLEUNIGUNG
Fortpflanzung herabgesetzt worden ist, so schreitet das Wachsthum
kräftig fort.
Es war mir daher nicht ohne Interesse, diese Verhältnisse
in unserem Falle kennen zu lernen. Ich konnte jedoch das
Studium nur bei Pilzen ausführen, nicht aber bei Algen, da
meine Versuchsalgen hauptsächlich einzellige Formen waren, die
nur durch Theilung sich vermehrten.
Bei Pilzen hingegen war die Wechselbeziehung zwischen der
Mycelentwickelung und der Sporenbildung deutlich zu erkennen.
In fast allen Fällen übten die Versuchsstoffe auf die Pilze
Sporen- bezw. Conidienbildung verzögernden Einfluss aus. Be-
sonders ausgeprägt trat dies bei Zusatz von ZnSO, und NaFl
ein. Ich konnte vielfach constatiren, dass bei normalen, in
Zimmertemperatur (ca 15° C im Mittel) gezüchteten Aspergillus-
Kulturen das Mycelium schon nach 1-2 Tagen sich ausbreitete
und mit angelegten Sporangienträgern versehen war, deren Köpfe
nach weiteren 2 Tagen durch Reifung der Sporen ganz ge-
schwärzt worden waren. Bei den Kulturen mit Zusatz von 0.005
Proc. ZnSO, dagegen habe ich selbst nach Verlauf einer Woche
vergeblich nach reifen Sporangien gesucht. In den letzteren
Fällen fand ich nur auf dem ziemlich stark angewachsenen
häutigen Mycel etwas Sporangienträgeranlage mit etwas aus-
geschwollenen Köpfchen, nebst einer Anzahl von wohl als Luft-
mycel anzusehenden Gebilden. Erst nach weiterem einwöchigen
Stehen wurden sie mit bräunlich-schwarzen Sporen besetzt.
Die Grösse der Sporen, welche gegen verschiedener Einflüsse
sehr empfindlich ist, blieb in unseren Fällen unverändert. Die
Beschaffenheit des Myceliums aber war nicht unbedeutend
beeinflusst.
Wie erwähnt, fand ich in allen von mir untersuchten Stoffen
EINIGER ALGEN UND PILZE. 153
mehr oder minder die Tendenz, die Sporenbildung zu verzögern
oder wenigstens zu verspäten. Dies gilt ohne weiteres auch für
Penicilhum sowohl, wie für Aspergillus.
Am ausgeprägtesten und am schönsten aber konnte ich
dieses Verhältnis bei Aspergillus-Kultur mit Zusatz von NaFl
kennzeichnen.
Ich nehme hier aus meinem Protokolle folgendes Beispiel :—
I. Normal Ganze Oberfläche mit schwarzen Sporen bedeckt.
II. 0.0025% NaFl Etwa !/, der Decke mit Sporen bedeckt, die übrigen
3/, steril.
III. 0.005 „ ,, Nur sehr spärliche Sporenbildung, weiss.
IV. 0.010 „ ,, Steril, hautbildend, weiss.
V. 0.021 „ , Ganz steril, weiss.
(Beobachtet nach 10 Tagen seit der Aussaatzeit der Sporen.)
Alle NaFl enthaltenden Kulturen zeigten eine üppige vege-
tative Entwickelung des Myceliums und bildeten hautartige
Decken.
Nach weiteren zehn Tagen waren die Kulturen wesentlich
unverändert im Aussehen. Bei III. sporadisches Auftreten der
Sporangienträger mit unreifen Sporen, bei IV. Sterilbleiben
der Decke mit einigen haarigen Luftmycelen, bei V immer steril.
Die Trockengewichtbestimmung nach der Beendigung der
Kultur zeigte folgendes Resultat:
I. IX. Ill. IV. V
0.314g 0.336g 0.385g 0.316g 0.274g
(Für näheres vgl. Tabelle Pilze H)
Eine Photographie am Ende dieser Arbeit veranschaulicht
das eben gesagte Verhältnis.
Die Unterdrückung der Sporenbildung in diesem Falle kann
allem Anschein nach eher dem direkt hemmenden Einfluss des
154 N. ONO : WACHSTHUMSBESCHLEUNIGUNG
betreffenden Stoffes zugeschrieben werden, als der durch stärkere
Mycelentwickelung hevorgerufenen Correlationserscheinung, welche
häufig bei günstigstem Nährboden zu Stande kommt. Um dieser
Punkt aufzuklären, stellte ich die Versuche derart an, dass ich
ein Stück Mycelium, welches für 19 Tage in 0.015% NaFl Lösung
gezüchtet worden war und keine eigentliche Fruktifikation
abgesehen von einigen schon besprochenen haarigen Gebilden
aufnimmt, nach Bespülung mit \Vasser in Normallösung brachte
Schon nach 2 Tagen kamen die angelegten Träger zur Reif
und schwärzten, während ich auf der noch in 0.015% verblei-
benden Decke vergeblich nach den reifen Sporangien suchte
Dieses Resultat spricht ohne weiteres für die gesagte Ansicht.
Es fragt sich nun, ob die stärkere Entwickelung des vege
tativen Organs der Pilze bei der Zugabe einer kleinen Dosi
giftiger Stoffe nicht eher als ein specieller Fall der Correlations
erscheinungen zwischen Fortpflanzung und Wachsthum zu be.
trachten ist, indem der direkt hemmende Einfluss der Substanzer
auf die Sporenbildung durch Correlation die vegetative Funk-
tion befördert. Es ist schon bestätigt wurden, dass Sporenbil-
dung der Pilze durch einige Gifte‘). viel empfindlicher beeinfluss
wird als die vegetative Mycelentwickelung. Fasst man diese
Verhältnis ins Auge, so ist es wohl begreiflich, dass bei eine
senügenden Verdünnung der angewandten Stoffe jene Concentratior
erreicht ist, welche an sich für die Mycelentwickelung unschäd-
lich, aber für die Sporenbildung hemmend wirkt, und dass x
infolge der Correlation das Wachsthum des vegetativen Theil:
ungewöhnlich gesteigert wird. Dieser indirekten Wirkung de
betreffenden Stoffe schreibe ich, ausser dem direkten Reizeffekt
die Wachsthumssteigerung zu.
1) O. Loew, Ein natürliches System der Giftwirkungen 1893.
EINIGER ALGEN UND PILZE. 155
V. Einfluss der Reizstoffe auf die Betriebsstoffwechsel.
Die Pflanze nimmt durch ihre Lebensthätigkeiten die Nähr-
stoffe auf und verwendet einen Theil derselben zum Aufbauen ihres
Körpers, hingegen den anderen Theil zum Oxydationsmaterial, um
dadurch die nothwendige Betriebsenergie sich zu verschaffen.
Von diesem Standpunkte aus betrachtet, lassen die im Pflanzen-
organismus sich abspielenden Stoffumsätze sich, wie bekannt, in
zwei Kategorien: Bau- und Betriebsstoffwechsel trennen. Um
die Grösse jedes von diesen Stoffwechseln zu ermitteln, hat man
einigermassen einen Maasstab. So ist bei der Betriebsstoffwechsel-
thätigkeit das Trockengewicht maassgebend, und für den Betriebs-
stoffwechsel giebt die Ermittelung der Kohlensäureproduktion, die
Ausscheidung gewisser Stoffwechselprodukte, einige Aufschlüsse.
Wie aus der in der Einleitung angeführten Skizze zu ersehen,
beziehen sich die bisherigen Untersuchungen über die Erhöhung
der Lebensthätigkeiten durch chemische Reize zumeist nur auf
Baustoffwechsel. Richards untersuchte in seiner Arbeit nur den
Ernteertrag, berücksichtigte aber nicht den Betriebsstoffwechsel.
Schulz beobachtete stärkere Entwickelung der Kohlensäure bei
Hefen. Beim ersten Anblick scheint dies auf nur erhöhtem
Betriebsstoffwechsel zu beruhen, doch blieb hier die Frage immer
offen, ob man es in diesem Falle mit der Erhöhung der Gähr-
thätigkeit einzelner Individuen zu thun hat, oder ob diese
Erscheinung von der durch die Reizwirkung verursachten Ver-
mehrung der Gährungserreger bedingt sei.
Es fehlen bisher meines Wissens einschlägige Versuche über
die Beeinflussung des Betriebsstoffwechsels in Gegenwart von
giftigen Stoffen. Irgend ein Beitrag in dieser Richtung durfte
wohl nicht ohne Interesse sein.
156 N. ONO: WACHSTHUMSBESCHLEUNIGUNG
Für solehen Zweck bieten die Algen keine geeigneten Objekt
dar, wohl aber die Pilze, welche sich dafür bequem anwenden lassen.
Einige Pilze, insbesondere Aspergillus niger, produciren eine
nicht unbeträchtliche Menge Oxalsäure als Stoffwechselprodukt
wie aus der bekannten Arbeit Wehmer’s') hervorgeht. Diese
Stoffwechselprodukt bot bei meinen Versuchen einen Angriffspunkt
und so werden eine Reihe Bestimmungen über die Säurenmeng
angeführt. Ich muss hier bemerken, dass ich die Säure nur au
titrimetrischen Wege bestimmte. Die Methode ist untauglich, wenr
Oxalsäure nicht nur als solche, sondern auch als Salz vorkommt
Wehmer zeigt aber in seiner oben besprochenen Arbeit, dass iı
NH,NO,-haltiger Zuckernährlösung Oxalsäure stets als freie Säurt
bei Aspergillus-Kulturen auftritt und da meine Kulturen haupt:
sächlich derartige waren, so war die Titration zuverlässig. Die
geschah mit Liquor Alkali Decinormalis und Phenolphthaleir
als Indikator.
Nachdem ich von der durch Titration ermittelten gesammter
Säure die ursprüngliche Acidität der Nährlösung subtrahirt hatte
rechnete ich diese als Oxalsäure um, welche in der angeführter
Tabelle gegeben ist”).
1) Wehmer, Entstehung und physiologische Bedeutung der Oxalsäure im Stoffwechsel!
einiger Pilze, Bot, Ztg. 1890.
2) Hier werde ich einige Versuche, um die titrimetrisch ermittelten Zahlen mit denjenigen
die gravimetrisch bestimmt waren, zu vergleichen, ausgeführt angeben :—
Saurenmenge in g in 10 cc Nährflüssigkeit.
Zusatz von NiSO, Titration Gravimetrisch
0 0.042 0.048
0.003% 0.052 0.060
0.007 ,, 0.050 0.058
0.014 ,, 0.066 0.075
0.028 ,, 0.064 0.060
0 0.042 0.040
0 003 ,, 0.047 0.048
0.007 ,, 0.052 0.058
0.014, 0.060 0.065
0.028,, 0.051 . : 0.054
EINIGER ALGEN UND PILZE. 157
Vergleicht man nun die Säurenmenge bei Gegenwart von
Reizmitteln mit derjenigen der Kontrollversuche, so findet man in
unseren Versuchen nur mit der einzigen Ausnahme von NisO,
stets das Minus im ersteren Falle. Dieses Verhältnis ist ersicht-
lich aus der Colonne ‚Säure pro 1g Pilzsubstanz‘“ in den Tabel-
len. Steigt der Zusatz von Reizstoffen, so wird die Menge der
Säure um so kleiner.
Man kann jedoch nicht annehmen, dass die aufgefundene
Säurenmenge die sämtliche Menge der ausgeschiedenen Säure
darstellte, da bekanntlich die Ausscheidung der Säure mit ihrer
Zersetzung Hand in Hand gehen sollte. Daraus geht hervor, dass
die Erklärung der besprochenen Verhältnisse nicht einzig in ihrer
Art sein kann. Es könnten einige Möglichkeiten, welche für
diese Thatsache sprechen, angegeben werden.
Erstens, wenngleich die Oxalsäure als normales Stoff-
wechselprodukt unseres Pilzes auftritt, ist sie doch als ein Pro-
dukt unvollkommener Oxydation anzusehen, und wenn die Stoff-
wechselthätigkeit auf einmal gesteigert wird, so wird als das
Produkt vollkommener Oxydation mehr Kohlensäure entstehen,
dagegen weniger Oxalsäure.
Zweitens könnte dieselbe Erscheinung auftreten, falls die
einmal entstandene Säure durch Wiederverarbeitung seitens der
Pilze verschwindet. Dabei könnte sie entweder als Baumaterial
wiederaufgenommen werden oder, ohne wieder den Pilzen nutzbar
zu werden, zersetzt werden. Doch, wie schon Wehmer') betont
hatte, stellt die Oxalsäure einen nur sehr armen Nährstoff für
Aspergillus dar, so dass es wahrscheinlich ist, dass sie bei
zureichendem Vorrath von guten Nährstoffen wie Zucker”) wohl
1) Wehmer l.c.
2) Bei meinem Versuche betrug der Zuckergehalt nach Beendigung der Versuche wenig-
stens 1.5 g in je 50 cc der Kulturfliissigkeit, d. h. ca 3%.
195 N. ONO: WACHSTHUMSBESCHLEUNIGUNG
intakt geblieben wäre. Ferner wurde von Wehmer') constatirt
dass weder Licht noch tote Pilzmasse allein die Zersetzung de
Säure hervorzurufen im Stande sind, wohl aber die Lebensthä
tigkeiten der Pilze. Es bleibt daher nur die Annahme übrig
dass die Säure durch Steigerung des Betriebsstoffwechsels lebhaft
erer Zersetzung unterworfen worden sei.
Die dritte Möglichkeit ist schliesslich die, dass diejenige,
Stoffe (Kohlenhydrat u. s. w.), welche bei normaler Wachsthums
energie durch Stoffwechsel z. Th. als Oxalsäure auftreten, be
der infolge der Reizwirkung über Norm gesteigerten Wachthum:
thätigkeit nicht als jene Form abgesondert werden, sondern sic
gerade in den integrierenden Theil des Pilzkörpers umwande
ten, kurz, dass sie als Baustoff verwendet werden.
Von einer anderen Seite müssen wir also dieses Problem an
greifen, um zu entscheiden, ob die eine oder andere von diesen Mög
lichkeiten für unseren Fall zutrifft. Die Ermittelung der Kohler
säureausscheidung, des ökonomischen Coéfficienten’) u. a. wird wol
an diesen Punkt anschliessend oder wenigstens rathgebend seiı
Im Folgenden gebe ich die Resultate meiner Bestimmunge
ökonomischer Coéfficienten bei den Kulturen mit Zusatz vo
/nSO,, bei denen die Erntezunahme stets am auffallendsten wa
Die Bestimmung des Coéfficienten fand in folgender Weis
statt :
Die Kulturflüssigkeit wurde zunächst durch andauernd
Kochen mit verdünnter Salzsäure vollkommen invertirt. Dan
verdünnte ich diese bis zu etwa '/,% Zuckergehalt. Eine Burett
wurde mit der betreffenden Lösung gefüllt.
In einem Kolben mischte ich genau 10cc Fehling’sch
1) Wehmer Le.
2) Man vergleiche hierüber H. Kunstmann, Ueber das Verhältniss zwischen Pilzern
und verbrauchter Nahrung. 1895. (Leipziger Dissertation).
EINIGER ALGEN UND PILZE. 159
Lösung mit etwa 40 cc Wasser und brachte es zum Sieden ; darauf
fügte ich die oben genannte Lösung hinzu, bis schliesslich durch
vollkommene Reduction des Kupfers zu Kupferoxydul die
Flüssigkeit farblos geworden war.
Da unsere Fehling’sche Lösung in 1000 cc 34.64g Kupfer-
ulfat, 174g Kaliumnatriumtartrat und 120g Natriumhydroxyd
nthielt, so sollte je 10 cc derselben durch 0.05g Zucker reducirt
erden. |
Nun kann man leicht durch die Lesung der Burette die
uckermenge in der Kulturflüssigkeit kennen lernen. Die
Jifferenz zwischen der ursprünglich vorhandenen Zuckermenge
n der Kulturflüssigkeit und der zurückbleibenden ergiebt selbst-
erständlich die verbrauchte Zuckermenge.
Kulturen mit Zusatz von ZnSO..
Aspergillus niger.
Kulturdauer 14 Tage. Zimmertemperatur.
Ökonomischer
Gehalt an ZnSO, ee Verbrauchte Coöffieient
(Gew. %) Zuckermenge d.h. Verbrauch
Erute
É.
0 0.262 1.594 6.1
0.0037 0.860 2.429 2.8
0.0074 0.875 2.429 2.8
0.0148 0.785 2.380 3.0
0.0297 0.773 2.340 3.0
IL.
0 0.386 1.707 4.4
0.0037 0.924 2.463 2.7
0.0074 0.928 2.463 2.7
0.0148 0.918 2.448 2.7
0.0297 0.837 2.480 2.8
160 N. ONO: WACHSTHUMSBESCHLEUNIGUNG
I.
(8) 0,392 1.819 4.6
0,0037 0,910 2,462 2.7
0,0074 0.908 | 2,456 27
0.0148 0.844 2.456 2.9
0.0297 0.827 2.446 | 2.8
Was sich nun aus diesem Resultate beurtheilen läss
dass der ökonomische Coéfficient in jedem Falle bei v
grösser ist in Kontrolle d.h. in nicht zugesetzter Kultur
zugesetzter. Dieses Verhältnis deutet also an, dass die Pi
Anwesenheit von Zinksulfat veranlasst wurden, mit einen
hältnismässig kleinen Verbrauch von Zucker eine bed
grössere Körpersubstanz aufbauen zu können. So scheint
wenigstens für Zinksulfat, von den oben besprochenen
Möglichkeiten die dritte die wirkliche zu sein,
IV, Specielle Besprechungen.
ZnSO.
Unter den von Richards geprüften Stoffen übt diese
die stärkste Wirkung aus.
Auch bei unseren Versuchen mit Algen wirkte ZnSO, :
FeSO, sehr günstig auf das Wachsthum ein. Schon bei |
von einer minimalen Quantität, wie 0.000016%, nahm die
etwas zu, und dies war noch deutlicher bei 0.00006% bis 0.0
Stieg die Concentration auf 0.0016%, so litten die Algen
unerheblich, ohne jedoch das Wachsthum ganz herabzu
(cf. Tabelle. Algen A. I-IV).
Unsere Versuche mit Pilzen stimmen mit denjenige
Digitized by Google -
EINIGER ALGEN UND PILZE. 161
Richards überein. Bei längerem Stehen wurde der Unterschied
zwischen den Versuchs- und Kontrollekulturen recht über-
raschend (cf. Tabelle. Pilze A. I-III).
Sehr sonderbar trat einmal bei einer Versuchsreihe mit
Dextrose es hervor, dass kein nennenswerther Unterschied in
der Ernte sowohl, als auch in der Säurequantität sich erkennen
liess. Den Grund davon kann ich aber nicht erklären (Tabelle.
Pilze. A. IV.)
Die gelbliche Färbung von Nährflüssigkeiten, sowie die
Bildung der bräunlichen Sporen in den Versuchskulturen, welche
schon von Autoren besprochen wurden, waren hier bemerklich.
Die Säurenmenge nach Beendigung der Versuche war in
Versuchskulturen viel kleiner als in Kontrollen (Tab. Pilze.
A. I-III.).
FeSO..
Richards giebt an, dass dieses Salz erst bei ziemlich gros-
sem Gehalte einen schädigenden Einfluss ausübt.
Meine betreffenden Versuche mit Algen zeigten auch, dass
lasselbe noch höhere Concentration im Vergleich zu anderen
Schwerenmetallsalzen erträgt. So lag bei Hormidium das Optimum
twa bei 0.0005% und sogar bei einer höheren Concentration
wie 0.0126% war der Ertrag noch etwas grösser als bei den
Kontrollen (cf. Tab. Algen. B. I-II.).
In einer mit Zusatz von FeSO, angestellten Penicillium-
Kultur trat merkwürdigerweise das ziegelroth gefärbte Mycelium
zu Tage.
NiSO..
Bei Algen ruft der Zusatz von NiSO, einen befördernden
Einfluss hervor. Die optimale Dosis lag etwa zwischen 0.00006
|
|
= —
162 N. ONO! WACHSTHUMSBESCHLEUNIGUNG
und 0.00012%, während 0.0028% eine beschädigende Wirk:
ausübte (Tab. Algen C. I. II.).
Säureproduktion bei Pilzkulturen war hier im Gegensat:
den meisten Fällen grösser mit der Erhöhung der Zusätzeproc«
(cf. Tabelle Pilze. C. I-III.).
CoSO.. |
Bei Algen scheint dies auch einen begünstigenden Einf
auszuüben, doch lag der optimale Punkt etwas niedriger als
NiSO, ; Optimum etwa bei 0.00012% (cf. Tab. Algen D. I. |
Säureerzeugung war wie gewöhnlich kleiner und zwar :
regelmässig in Versuchskulturen.
CusO.
CuSO, wurde von Richards nicht untersucht. Im Je
1897 constatirte Günther'), dass Kupfersalze in grôss
Mengen das Wachsthum der Pilze retardirten, in geringe
Mengen dagegen besseres Gedeihen mit sich bringen. A
bei meinen mit Aspergillus und Penicillium angestellten \
suchen beobachtete ich dieselbe Erscheinung. Hattori’) {
auch in seinen Untersuchungen über die Giftwirkung der K
fersalze eine ähnliche Thatsache.
Hier werde ich zwei Beispiele angeben ; für näheres verw
ich auf die tabellarische Zusammenstellung (Tab. Pilze. E.).
Aspergillus niger.
Gehalt an CuSO, | 0.001554 | 0.003%
0.006% | 0.01:
| Ernteertrag in g| 0.273 |0.307 | 0.313 | 0,324 | 034
1) E. Günther, Beitrag zur mineralischen Nahrung der Pilze, Erlangen 1897.
2) H. Hattori, Ueber die Einwirkung des Kupfersulfates auf einige Pi
Manuskript,
Digitized by Google
EINIGER ALGEN UND PILZE. 163
Penicillium glaucum.
Gehalt an CuSO,
0 | 0.0015% | 0.003% | 0.006% | 0.01296
Emteertrag in g| 0.213 | 0.320 0.338 | 0.359 | 0.410
Bei Algen konnte ich dagegen keine Wachsthumsbeförde-
rung nachweisen, wie aus der Tabelle ersichtlich ist. Schon bei
0.00001% steht die Ernte etwas zurück (Tab. Algen E.). Ob
bei noch weiterer Verdünnung die wachsthumsbegünstigende
Concentration erreicht sein könnte, lasse ich vorläufig unbestimmt.
Die Säurenmenge in Pilzkulturen war kleiner in Versuchs-
culturen (Tab. Pilze. E.).
HgCL.
Es ist von gewissem Interesse, dass dieses heftige Gift in
senügender Verdünnung auch das Wachsthum der Pilze befördert.
Schulz!) giebt an, dass Kohlensäureentwickelung der Hefe in
xegenwart einer kleinen Menge des Stoffes gesteigert wird. Der
ptimale Zusatz dabei ist etwa 1/500 000.
Was Schimmelpilze betrifft, so findet man in der bisherigen
uitteratur nur die Rede von dem schädigenden Einfluss des
etreffenden Stofis. Raulin’) betrachtet z. B. dies mit AgNO,,
Cl, zusammen als das giftige Salz für Aspergillus. Er gibt
1512 000 als die Grenze der Giftwirkung. In seinem Experi-
nente mit 1/819200 konnte er jedoch keinen wachsthums-
eschleunigenden Einfluss beobachten. Meines Wissens liegt
ıns zur Zeit kein Versuch vor, welcher die letztgenannte That-
ache in positivem Sinne zeigt.
1) H. Schulz, Le
2) Raulin l.c. p. 134.
164 N. ONO: WACHSTHUMSBESCHLEUNIGUNG
Meinem Versuche nach (ef. Tab. Pilze. F.) tritt scho
Verdünnung von 0.0017% oder 1/60 000 ziemlich gute Entw
lung von Aspergillus ein. Stieg die Concentration auf 1/3(
so kam die Entwickelung zum Stillstand. Die Grenze für
wirkung liegt zwischen 1/60 000 und 1/30 000.
Das Optimum war sowohl bei Peineillium als auc
Aspergillus etwas unter 0.0013%).
Hier gebe ich zwei Beispiele (cf. Tab. Pilze. F. I-VI)
Aspergillus niger.
' Gehalt an HeCl,
0 | 0.000894 0.0007 | 0.001355 | 0.0
| Ernteertrag ing
0.261 | 0.355
Penicillhium glaucum.
Gehalt an HgCl,| 0 | 0,0003% | 0.0007% | 0.001354 | 0.00
"Erntosrtrag ‘fh « | 0183 [0249 | 0213 | 0311 | 0.24
Siiureproduction ist hier wie bei den meisten Fällen klein
Versuchskulturen als bei Kontrollen (Tab. Pilze. F.).
Auf Algen übte dieses Salz keinen beschleunigenden Ei
aus, sondern wirkte nur giftig ein. Schon bei 0.00005% we
schädigende Effekt deutlich zu erkennen. Doch weitere
dünnung durfte ich nicht ausführen, da bei solchen |
Verdünnungen einige Fehlerquellen als maasgebend auf
(Tab, Algen F.).
LiNO.
Von Richards wurde LiCl! zur Untersuchung herausge
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EINIGER ALGEN UND PILZE. 165
nd in diesem Salze ein eine beträchtliche Wachsthumssteigerung
ervorrufender Stoff gefunden. Bei meinem Versuche benutzte
h LiNO, mit gleichem Resultate (Tab. Pilze G.)
Säureproduktion war hier auch kleiner in Versuchskulturen
ls in Kontrollen.
Algen zeigten auch eine besseres Gedeihen in zugesetzten
ulturen (Tab. Algen H.).
NaF.
Dieser Stoff übte auch eine beschleunigende Einwirkung auf
Igen aus. Der optimale Punkt liegt etwa bei 0.00003%, stieg
e Concentration zu 0.00016% bis 0.0008%, so nahm die Ernte
was ab, war noch grösser als bei Kontrollen. Erst bei 0.0042%
eht der Ertrag im Vergleich zur Kontrolle etwas zurück
fab. Algen G.).
Bei Pilzen beförderte dieser Stoff das Wachsthum (cf. Tab.
ilze H.). Seine Wirkung auf die Sporenbildung wurde schon
n vorstehenden Capitel behandelt.
Die Säurenmenge in der Nährflüssigkeit war wie gewöhnlich
leiner in zugesetzten als in Kontrolle-Kulturen.
Arsen.
Von Arsenverbindungen ist arsenige Säure giftig, doch ver-
agen höhere und niedere Pflanzen viel Arsensäure').
Da Arsenigsäureanhydrid nur schwer löslich ist, so bediente
ch mich des arsenigsauren Kaliums.
Bei Penieillium-Kultur war kein bedeutender Unterschied
ler Ernte sowohl als auch in der Säureproduction bemerklich.
1.) O. Loew, System der Giftwirkungen.
a
166 N. ONO: WACHSTHUMSBESCHLEUNIGUNG
Merkwürdig war hier eine eigentliche Geruchsentwickeln:
zugesetzten Kulturen’).
Auf Algen scheinen die genannten Salze etwas Wachstl
begünstigend zu wirken. (Tab. Alg. I).
VII. Schlussbemerkungen und Zusammenfassung
der Resultate.
Aus dem Vorstehenden geht zunächst hervor, das
chlorophyllführenden niederen Organismen wie Algen in :
Gedeihen günstig beeinflusst werden durch einen geringen 7
von einigen Stoffen, welche für sich nicht Nährstoffe sn
sogar giftig wirken. In dieser Reaktion verhalten sicl
Algen gerade wie die Pilze. Nur ist zu bemerken, das
optimale Dosis für Algen viel kleiner als bei Pilzen ist
Thatsache, welche vielleicht vom oekologischen Standpunkt
ihren Aufschluss haben wird. Von den geprüften Stoffen k
ich nur bei Quecksilberchlorid und Kupfersulfat die bespro
Reaktion nicht eonstatiren, indem ich bei ihnen, soweit »
Versuche reichten, stets Giftwirkung beobachtete. Daraus
aber nicht geschlossen werden, dass den beiden Stoffe
nämliche Eigenschaft nicht zukommt, da man bei ihnen
Umständen doch noch jene wachsthumsbegünstigende Einwi:
wohl erwarten kann.
Bei Pilzen ikonnte ich die früheren Versuche Rich
hauptsächlich bestätigen, dazu prüfte ich mit positiven Resu
einige bisher noch nicht untersuchte Stoffe,
Die Verzögerung oder Verspätung der Sporenbildun
unseren Versuchen ist nicht als infolge einer üppigen veget:
1) Schon von Gasio (Jahresber. über Gührungsorganismen 1893) erörtert.
Digitized by Google
| Be
EINIGER ALGEN UND PILZE. 167
intwickelung verursachte Correlationserscheinung, vielmehr als
urch Reizstoffe bewirkte Hemmung zu betrachten.
Was nun die Art und Weise der Reizwirkung anbelangt,
9 bemerke ich folgendes:
Wenn es sich hier zunächst um zeitliche oder andauernde
[yperaesthesia handelt, so muss Bau- und Betriebsstoffwechsel
leichzeitig gesteigert werden.
Wenn aber dagegen durch Zusätze der Reizstoffe die Thätig-
eiten seitens des Organismus so gesteigert werden, dass sie mit
leinerem Energieaufwand die Nährstoffe in sich aufnehmen und
ch bauen, kurz, ökonomisch arbeiten können, so kann der dyna-
ische Stoffwechsel nicht so erheblich beeinflusst bleiben. Um
her in dieser Hinsicht eine richtige Auffassung zu gewinnen, ist
n Einblick in den Betriebsstoffwechsel von Wichtigkeit. Einige
yn meinen Versuchen in dieser Richtung zeigten andeutungsweise,
iss Betriebsstoffwechsel nicht parallel mit Baustoffwechsel
steigert werden; doch sind zur Zeit meine diesbezüglichen
ersuche leider unzureichend, um in bezug auf diesen Punkt
llgemeines zu sagen.
Zum Schluss seien im Folgenden die wichtigsten Resultate
ırz zusammengestellt :—
1. Das Gedeihen der niederen Algen wird durch Einfüh-
ing gewisser giftiger Stoffe in höchst verdünnten Zuständen
»günstigt. Hierzu gehören ZnSO,, NiSO,, FeSO,, CoSO,, NaF,
INO,, K,AsO;.
2. Die Erntezunahme bei Algen muss auf die vegetative
ermehrung der Individuenzahl zurückzuführen sein, da keine
ennenswerthe Veränderung der Körpergrösse bemerkbar war.
3. Die geeignete Dosis ist bei Algen bedeutend kleiner als
168 N. ONO: WACHSTHUMSBESCHLEUNIGUNG
bei Pilzen. Schon der Zusatz von 10* Gr. Mol. Salz wirkte in
den allermeisten Fällen schädigend.
4. In CuSO, und HgCl, fand ich, soweit unsere Studien
ausreichen, keine beschleunigende Wirkung auf Algen, wohl
aber begünstigend bei Pilzen.
5. Bei Pilzen tritt durch Zusätze von HgCl, (Optimum etw:
bei 0.0013% und CuSO, (Optimum etwa bei 0.012%) die Wachs-
thumsbeschleunigung ein.
6. Die Säurequantität in Kulturen mit Zusatz von ZnSO,
CoSO,, HgCL, NaFl, CuSO, war stets kleiner als in Kontrol.
kulturen, Nur verhielt NiSO,, soweit meine Versuche en
Urtheil gestatten, sich diametral entgegengesetzt.
7. Die geprüften Stoffe (speciell ZnSO, und NaF!) neigeı
dazu, die Sporenbildung der Pilze direkt zu hemmen, wenigsten
das Auftreten der Sporen zu verspäten.
8%. Die oekonomischen Coefficienten in ZnSO,-Kultur sin
in der Kontrolle, d. h. in der nicht zugesetzten Kultur, be
weitem grösser als in der zugesetzten.
Juni, 1899. Botanisches Institut
Kaiserl. Universität
zu Tokyo.
EINIGER ALGEN UND PILZE. 169
TABELLARISCHE ZUSAMMENSTELLUNG.
BEMERKUNGEN:
Versuche mit Algen—Der Entwickelungsgrad ist entweder mit Erntegewicht oder mil
den relativen Werth gebenden Ziffern bezeichnet.
Versuche mit Pilzen—In Colonne „Acidität‘ ist die Quantität des Decinormul Allkulis
in cc gegeben, welche 10 cc der Nährflüssigkeit neutralisirte (Die ursprüngliche Avcidilit
surde natürlich vorher subtrahirt). Daraus ermittelte ich die Siurenmenge in je 50 ce Nähr-
lüssigkeit, berechnete sie als Oxalsäure und in Colonne „als Oxalsiiure umgerechnet“ angab.
In Kolonne „Säure pro lg Pilzsubstanz“ ist das Verhältnis Rue gegeben.
Obwohl bei einigen Penicillium-Kulturen die Säurenmenge gegeben sind, durfte ich doch
richt die QT ermitteln, da bei Peniciliun das Titration unzuverlässig scheint.
I. Versuche mit Algen.
A. I. Kulturen mit Zusatz von ZnSQ,.
Protococcus sp-—angestellt 5. Oct. Zimmer-Temperatur.
Ernteertrag in g.
Gram Mol.
Kultur-
dauer
Gew. %
| Gehalt in
A. IL. Kulturen mit Zusatz von ZnSO,.
Ernteertrag in g. Protococcus sp.—angestellt 11. Oct. Zimmer-Temperatir.
5
> | 3x 107 | Kultu-
a
&|Gew. %| 0 — [0.000012] 0.00006 | 0.0003 | 0.0014 | dur
I 0.035 | 0.043 | 0.038 | 0.042 | 0.024 | 44 Tage
II | 0.032 | 0.040 | 0.036 | 0.040 | 0.023 | 40
Ii 0.030 0.040 0.037 0.042 0.020 | 50 ,,
170 N. ONO : WACHSTHUMSBESCHLEUNIGUNG
A. IIL Kulturen mit Zusatz von ZnSO,.
Chroococcum—angestellt 24. Dec.
in Treibhaus 16-20°C.
4 x 10”!
Ernteertrag in g.
Gram
Mol.
107$
25x 10° Kultur-
0 0.000012 0.00006 | 0.0003 | 0.0014 | dauer
23x10
Gehalt in
sew. %
I 0.006 0.015 0.026 0.022 0.006 | 71 Tage
II 0.099 0.021 0.022 0.010
[II 0.010 0.024 0.026 0.009
A. IV. Kulturen mit Zusatz von ZnSO,.
Protococcus—angestellt 5. Feb. in Treibhaus 16-20° C.
Ernteertrag in g. .
= se | 0 25 x 1075 | 23x10 | 10° | 4x10 | Kultur-
3 Gew. %| 0 |0.000012 | 0.00006 | 0.0003 | 0.0014 | dauer
ie: 0.008 | 0.017 | 0.019 | 0.019 | 0.007 | 65 Tage
II 0.009 | 0.018 | 0.023 | 0.016 | 0.009 a
0.011 | 0.015 | 0.018 | 0.017 | 0.005 i
III
B. I. Kulturen mit Zusatz von FeSO,.
Ernteertrag in g. Hormidium nitens—angestellt 7. Oct.
(Gehalt in |
Zimmer-Temperatur.
0.042 |65 Tage
0.041 ı ,,
Gr: à À
= | 0 |x 10~| 4x 10~
Gew.% | 0 | 0.0001 | 0,0005 | 0.0025 | 0.0126 | dauer
I 0.038 | 0.072 | 0.074 | 0.080
Il 0.023 | 0.083 | 0.079 | 0.062
III 0.031 | 0.070 | 0.082 | 0.074
B. I. Kulturen mit Zusatz von FeSO,.
Ernteertrag in g. Hormidium nitens—angestellt 10. Nov.
E | Gram
= Mol.
E Gew. % 0 | 0.0001
mT 0.025 | 0.050 | 0.052 | 0.054
II 0.023 | 0.067 | 0.049 | 0.042
0.027 | 0.064 | 0.058 | 0.046
III
0.039 ii
in Treibhaus 16-20° C.
4 x 10
Kultur-
0.0005 | 0.0025 | 0.0126 | dauer
0.041 | 79 Tage
0.032 =
0.032 ‘5
EINIGER ALGEN UND PILZE. 171
C. I. Kulturen mit Zusatz von NiSO,.
Chroococcum—angestellt 2. Oct. Zimmer-Temperatur.
3x 107* | Kultur-
dauer |
Ernteertrag in g.
0.012 0.012 0.025 0.021 0.004
0.011 0.015 0.024 0.020 0.006
III 0.013 0.018 0.020 0.022 0.007
C. II. Kulturen mit Zusatz von NiSO,.
Hormidium nitens—angestellt 16. Nov. i, Treibhaus 16-24° C.
ge X 1075] 43x10 | 10° | 23x10 | Kultur-
tw G2
2 É
Grehalt in
C2
oO
=
< |
dauer
0.00006 | 0.00028 | 0.0014
0
0 0.000012
4
4
I 5 4-5 4 1 70 Tage
II 5 4 4 1 5
III 3-4 5 4-5 4 1 ‘i
N.B. Die Ziffer zeigt den Entwickelungsgrad,
D. I. Kulturen mit Zusatz von CoSO,.
Hormidium nitens—angestellt 9. Dec. i, Treibhaus 16-24° C.
0 3x 10° 3 x 10-* Kultur-
0 [0.000012 | 0.00006 | 0.0003 | 0.0014 | dauer
N.B. Die Ziffer zeigt den Entwickelungsgrad.
D. II. Kulturen mit Zusatz von CoSO,.
Ernteertrag in g. Protocoeeus—angestellt 19. Mai. Zimmer-Temperatur.
0 | 23x10) $x10-*| 10 Kultur-
Gew. % 0 [0.000012 | 0.00006 | 0.0003 dauer
=
=
©
a
D
do)
172 N. ONO: WACHSTHUMSBESCHLEUNIGUNG
E. I. Kulturen mit Zusatz von CuSO,.
Stigeoclonium—angestellt 14. Oct. Zimmer -Temperatur.
“4 en | 0 5 x 10 4x10 | 10 | 3x10- | Kultur
5 Gew. %| 0 | 0.00001 | 0.00005 | 0.00025 | 0.0012 | dauer
er 5 4 2
IT 5 4 2-3
(II 5 4 2
N.B. Die Ziffer zeigt den Entwickelungsgrad,
E. IJ. Kulturen mit Zusatz von CuSO,.
Chroococcum—angestellt 13. Dec. in Treibhaus 16-20? C.
= ie | 0 | 25x10” 3x 107 | Kultur-
= \Gew.%| 0 | 0.00001 | 0.00005 | 0.00025 | 0.0012 | dauer
71 Tage
„
N.B, Die Ziffer zeigt den Entwickelungsgrad,
F. I. Kulturen mit Zusatz von HgCl,.
Protococcus—angestellt 25. Dec. in Treibhaus 16-20° C.
x 10% 3 x 105
N.B. Die Ziffer zeigt den Entwickelungagrad
G. I. Kulturen mit Zusatz von NaFl.
Protocoecus—angestellt 24. Dec. in Treibhaus 16-20? C.
TL x 10) 2 x 10° 10° | Kultur-
| 125
Ernteertrag in g.
Gram
Mol. 0
=
| ==
=
(rehalt 1
Gew. %
I
IT
III
0.012 0.018 | 0.018
0.012 0.025 | 0.018
0.010 | 0.027 | 0.015
oo
EINIGER ALGEN UND PILZE. 17:
HA. I. Kulturen mit Zusatz von LiNO,.
Ernteertrag in g. Protococcus—angestellt 16. April. Zimmer-Temperatur.
ho 0 | ds x 107 3x10 | 10 14x10 Kultur.
À | Ger. % 0 | 0.00003 | 0.00014 | 0.0007 | 0.0034 | dauer.
I 0.010 | 0.020 | 0.017 | 0.012 | 0.009 | 24 Tage
Il 0.009 | 0.020 ! 0.020 | 0.015 | 0.010 R
III 0.010 | 0.018 | 0.016 | 0.011 | 0.008 i
I. I. Kulturen mit Zusatz von K,AsO,.
Ernteertrag in g. Frotococcus—angestellt 24. Dec. 1 Treibhaus 16-20° C.
= G
3 Sr 0 | 4x104|2,x10- 4x10 | 104 | Kultur
© |Gew.%| 0 | 0.00008 | 0.0001 | 0.0005 | 0.0024 | dauer.
T | 0011 | 0015 | 0.012 | 0.011 | 0.008 | 54 Tage
II | 0.008 | 0.017 | 0.020 | 0.012 | 0.007 F
III 0.009 | 0.015 | 0.018 | 0.014 | 0.009 .
II. Versuche mit Pilzen.
A. I. Kulturen mit Zusatz von ZnSO,.
Aspergillus niger—angestellt 24. Dec. ’98.
Geerntet 18. Jan. Kulturdauer 25 Tage. Temperatur 16-20° ©.
Gehalt in Säure Säure pro lg. |
Gram Mol. | Gew. x Acidität ae. se men Pilzsubetanz. |
0 0 0.216 15.7 0.495 2,245
32x10 | 0.003 12.9 0.406 0.470
1 x 10 0.007 13.2 0.416 0.443
3 x 107 0.014 12.9 0.406 0.430
10° 0.028 13.0 0.409 0,430
N.B, Asparagin als N-Quelle.
174 N. ONO: WACHSTHUMSBESCHLEUNIGUNG
A. II. Kulturen mit Zusatz von ZnSO,.
Aspergillus niger.—angestellt 24. Dec. ’98.
Geerntet 20. Jan. ’99. Kulturdauer 37 Tage. Temperatur 16-20° C.
Gehalt in | Ernteertrag Säure Säure pro lg.
Gram Mol. | Gew. £ in g. Aciditat | als Oxalsäurs | Pilzenbetanz.
0 O | 0.181 | 148 | 0467 | 2.580
4x 10 0.003 0.868 11.8 0.372 0.428
4x 10 0.007 0.870 11.1 0.350 0.420
4 x 10 0.014 0.858 12.1 0.381 0.444
10 0.028 0.821 14.1 0.444 0.541
N.B. Asparagin als N-Quelle.
A. II. Kulturen mit Zusatz von ZuSO,.
Aspergillus niger—angestellt 24. Dec. ’98.
Geerntet 20. Jan. ’99. Kulturdauer 27 Tage. Temperatur 16-20° C.
ies Gehalt in Ernteertrag Säure Säure pro lg.
Gram Mol. | Gew. x in g. tActaitat | als Oxaleture | Pilzsubstanz.
0 O | 0187 | 178 0.561 3.000
4x 10° 0.003 1.017 12.5 0.394 0.387
À x 10° 0.007 1.336 11.8 0.372 0.278
4 x 10 0.014 1.939 12.5 0.394 : 0.419
10 0.028 1.204 11.2 0.353 0.293
N.B. Asparagin als N-Quelle.
A. IV. Kulturen mit Zusatz von ZnSO,.
Aspergillus niger—angestellt 20. Febr.
Geerntet 11. Jan. Kulturdauer 20 Tage. Temperatur 16-20° C.
Gehalt in Ernteertrag Säure Säure pro lg.
Gram Mol, | Gew. % in g. Aciditat | als Oxalsäure | Pilzsubstanz.
umgerechnet
0 0 0.634 7.2
4x10 | 0.003 0.641 7.2
4x10 | 0.007 0.635 7.2
1x10= | 0.014 0.627 7.2
10° 0.028 0.585 7.2
N.B, Dextrose austatt Robrzucker.
EINIGER ALGEN UND PILZE. 175
B. I. Kulturen mit Zusatz von FeSO,.
Penicillium glaucum—angestellt 11. April.
Kulturdauer 14 Tage. Temperatur 16-20° C.
Geerntet 25, April.
— een: Ernteertrag Säure Säure pro Ig.
| Gram MoL | Gew, # in g. Aciditat ne Pilzsubstanz.
= sr 0.191 3.5 == +
1x 10° 0.007 0.180 3.3 — _—
4x 10"? 0.014 0.233 3.9 — —
10" 0.028 0.201 3.5 — —
2x10°° 0.056 0.181 3.4 -
N.B, NH,NO, N-Quelle. In allen eisenhaltigen Kulturen waren die Pilzmassen schön ziegelrolh
gefärbt. Schon bei 0.007% deutliche rothe Färbung bemerklich,
C. I. Kulturen mit Zusatz von NiSO,.
Aspergillus niger—angestellt 9. Febr.
Kulturdauer 22 Tage.
Geerntet 3. Mürz.
Temperatur. 16-20” C.
Säure
als Oxalsäure
umgerechnet
Säure pro lg.
Pilzsubstanz.
Ernteertrag
in g. Aciditat
0.250
0.297
0.315
0.401
11.0
11.1
10.7
14.6
0.346
0.350
0.337
0.460
1.464
1.179
1.069
1.147
0.295 14.0
N.B. NH,NO, als N-Quelle.
0.441 1.493
C, II. Kulturen mit Zusatz von NiSO,.
Aspergillus niger—angestellt 9. Febr.
Kulturdauer 23 Tage.
Geerntet 4. Mürz. Temperatur 16-20° C.
Säure Säure pro lg. |
Gehalt in | Ernteertreg
Gram Mol. | Gew. < in g- Acidität er Serene Pilzsubstanz. |
er 11.1 0.350 1.216
10.7 0.337 1.067
11.1 0.349 1.130
16.9 0.532 1,375
16.7 0.526
N.B, NH,NO, als N-Quelle.
1.500
176 N. ONO: WACHSTHUMSBESCHLEUNIGUNG
C. IH. Kulturen mit Zusatz von NiSO,.
Aspergillus niger—angestellt 9. Febr.
Geerntet 6. März. Kulturdauer 35 Tage.
Gehalt in
Temperatur 16-20° C.
Säure
als Oxalsäure
umgerechnet
0.308
Ernteertrag
in g.
Säure pro lg.
Pilzsubstanz.
0.951
Acidität
0.324 9,8
0.310
0.329
0.362
0.341
10.0
11.9
15.2
17.3
0.315
0.375
0.479
0.544
0.016
1.140
1.323
1.695
N.B. NH,NO, als N-Quelle.
C. IV. Kulturen mit Zusatz von NiSO,.
Aspergillus niger—angestellt 22. April.
Kulturdauer 12 Tage.
Geerntet 4. Mai. Temperatur 16-20° C.
Gehalt in Ernteertrag Säure en
Gram Mol. | Gew. % in g. Acidität a anse Pilzsubstanz.
0 0 0.262 en = en
4x 10° 0.003 0.390 — — a
4x10 | 0.007 0.404 RER m an
4 x 10 0.014 0.364 — er Dun
10° 0.028 0.315 aoe cn
N.B. NH,NO, als N-Quelle
C. V. Kulturen mit Zusatz von NiSO,.
Aspergillus niger—angestellt 22. April.
Geerntet 4. Mai. Kulturdauer 12 Tage. Temperatur 16-20° C.
Gehalt in Ernteertrag Säure Säure pro lg.
Gram Mo, | Gew. x in g. aciaitat | ele Oxalsare | Pilzsubetanz.
0 0 0.214 —— — ——
41x10 | 0.003 0.311 oo er ven.
4x10® | 0.007 | 0.300 — — —
3x 10 0.014 0.307 = 2S
105 | 0.028 | 0.296 — —
N.B, NH,NO, als N-Quelle,
EINIGER ALGEN UND PILZE. 177
C. VI. Kulturen mit Zusatz von NiSO,.
Aspergillus niger—angestellt 22. April.
Geerntet 4. Mai. Kulturdauer 12 Tage.
Temperatur 16-20° C.
Gehalt: in Säure
Ernteertrag Säure pro lg.
als Oxalsäure
Gram Mol. | Gen 4 | in g. Acidität an Pilzsubstanz.
0 0 0.278
4 x 10° 0.003 0.340
3 x 107 0.007 0.325
4 x 1075 0.014 0.308
10 0.028 0.324
N.B, NH,NO, als N-Quelle.
D. I. Kulturen mit Zusatz von CoSO,.
Aspergillus niger—angestellt 17. Febr.
Geerntet 16. März.
Kulturdauer 27 Tage.
Temperatur 16-20° C.
Gehalt in Ernteertrag Säure pro lg.
Gram Mol. | Gew. % ing Acidität ur Pilzsubstanz. |
0 0 0.297 9.0 0.283 0.953
ts x 10 0.0017 0.439 10.3 0.324 0.738
4 x 10 0.0035 0.565 12.3 0.387 0.685
4x 10° 0.007 0.751 11.2 0.353 0.470
4 x 10° 0.014 0.872 8.7 0.274 0.314
N.B, NH,NO, als N-Quelle,
D. U. Kulturen mit Zusatz von CoSO,.
Aspergillus niger—angestellt 17. Febr.
Geerntet 20. März.
Kulturdauer 31 Tage.
Temperatur 16-20° C.
Gehalt in Ernteertrag Säure pro Ig.
Gram Mol. | Gew. % in g. Aciditat als Oxalsiure | Pilzsubstanz. |
0 0 10.0 0.315 1125 |
ys x 10° 0.0017 12.1 0.381 0.900
4 x 10 0.0035 15.5 0.491 0.844
4 x 10 0.007 16.7 0.548 0.735
4 x 10 0.014 8.4 0.265 0.313
N.B. NH,NO, als N-Quelle.
178 x. ONO : WACHSTHUMSBESCHLEUNIGUNG
D, WL Kulturen mit Zusatz von CoSO,.
Aspergillus niger—angestellt 17. Febr.
Geernlet 17. März. Kulturdauer 28 Tage. Temperatur 16-20° C.
(rehal tin _| Ernteertrag Säure Siure pro
Gram Mol. Gen Zu in g. Aciditat na | Pilzsubsta
0 0 | 0.267 10.6 0.334 | 1.251
yy x10 | 0.0017 | 0.398 13.0 0.409 | 1.041
1x10 0.0035 | 0.561 15.5 0.488 0,870
1x10" | 0.007 | 0.742 148 0.466 0.628
1x 10- 0.014 | 0.770
10.0 0.315
N.B, NH,NO, als N-Quelle. =
0,409
D, IV. Kulturen mit Zusatz von CoSO,.
'enieillium glaueum—angestellt 20. März.
(teerntet 30. März. Kulturdauer 10 Tage. Temperatur 16-20° C
Gehalt in _| Ernteertrag Bäure _ | Säure pro
Gram Mol. Gew. = | in g. Aciditat als Oxalsiure | Pilzsubsta
| umgerechnet | os
0 | 0 0.108 5.4 0,170 _—
de x 107 0.0017 0.186 5.0 0.157 —
1x10 | 0.0035 0.225 5.4 0.170 —
1x 10 0.007 0.317 5.9 0.186 —
1 x 10 0.014 0.366 5.9 0.186 —
N.B. NH,NO, als N-Quelle. -
D. V. Kulturen mit Zusatz von CoSQ,,.
Penicillium glaucum—angestellt 20. März.
Geerntet 1. April. Kulturdauer 11 Tage. Temperatur 16-20° C
= ( jehalt in = ‘| Ernteertrag Säure | Säure pro
Gram Mol. Gow, # | in g. Acidität | en Pilzsubetaı
0 | 0 0.242 4.6 0.145 | —
Æx10 | 0.0017 | 0.469 5.7 0.179 —
1x10 | 0.0035 | 0.354 5.9 0.186 er
1x10 | 0,007 0.482 5.7 0.178
1 x 10"? 0.014 | 0.772 5.9 0,186
N.B. NH,NO, als N-Quelle,
EINIGER ALGEN UND PILZE. 179
D. VL Kulturen mit Zusatz von CoSO,.
Penicillium glaucum—angestelit 20. März.
Kulturdauer 23 Tage.
Geerntet 13. April. Temperatur 16-20° C.
Gehalt in Ernteertrag Siure
in g. als Oxalsäure
Gram Mol, | Gew. % 8 Acldität | umgerechnet
0 0 0.363 6.4 0.202
ys x 10 0.0017 0.349 6.2 0.195
4x10°
Sdure pro lg.
Pilzsubstanz.
41x10 0.007 0.649 5.7 0.179
4x 103 0.014 0.289 6.6 0.207
N.B. NH,NO, als N-Quelle,
0.0035 0.520 5.9 0.186 —
E. I. Kulturen mit Zusatz von CuSO,.
Aspergillus niger—angestellt 21. März.
Kulturdauer 10 Tage.
Geerntet 31. März. Temperatur 16-20° C.
Gehalt in Ernteertrag Säure Säure pro lg.
Gram Mol. | Gew. % in g. Acidität als: nn Pilzsubstanz.
0 0 0.307 6.2 0.195 0.635
ys x 10° 0.0015 0.305 5.2 0.164 0.538
4x 10 0.003 0.297 5.2 0.164 0.552
4x10 0.006 0.311 4.7 0.138 0.444
4x 10° 0.012 0.360 5.1 0.160 0.444
N.B. NH,NO, als N-Quelle,
E. I. Kulturen mit Zusatz von CuSO,.
Aspergillus niger—angestellt 21. März.
Kulturdauer 15 Tage.
Geerntet 5. April.
Gehalt in
Gram Mol, |
Gew. %
Ernteertrag
in g.
Acidität
N.B. NH,NO, als N-Quelle,
Temperatur 16-20° C.
als Oxalsäure
umgerechnet
Säure pro 1g.
Pilzsubstanz.
180 N. ONO: WACHSTHUMSBESCHLEUNIGUNG
E. III. Kulturen mit Zusatz von CuSO,.
Aspergillus niger—angestellt 21. März.
Geerntet 5. April. Kulturdauer 16 Tage. Temperatur 16-20° C.
— SS Ernteertrag | Säure | Säure pro Ig.
Gram Mol. Gew. % in g. Acidität | ech Pilzsubstanz.
mary 0 0.218 9.9 0.312 1.431
ps x 10 0.0015 0.252 10.1 0.318 1.265
1 x 10 0.003 0.352 9.9 0.312 0.886
1 x 105 0.006 0.358 9.3 0.293 0.818
4x 10 0.012 0.343 9.6 0.302 0.880
| > N.B, NH,NO, N-Quelle.
F. I. Kulturen mit Zusatz von HgCl,.
Aspergillus niger—angestellt 1. Febr. ’99.
Geerntet 9. Febr. Kulturdauer 8 Tage. Temperatur 16-20° C.
Gehalt in Ernteertrag | Säure | Säure pro 1g.
Grum Mol Gew. % in g. | Acidität an nie Pilzsubstanz.
0 0 0.126 12.6 0.397 | 3.176
4x10= | 0.0017 0.180 13.3 0.419 2.328
1x 103 0.0034 — — —— ——
4x10 | 0.0067 | — — oo =
4x10- 0.0135
N\A. NH,NO, als N-Quelle, 0.0017% gut entwickelt. Sporen braun. 0.0034% fast keine Entwickelung.
0.0067% u. 0.0185% keine Entwickelung.
F. IT. Kulturen mit Zusatz von HgCl,.
Aspergillus niger—angestellt 1. Febr. ’99.
Geerntet 9. Febr. Kulturdauer 8 Tage. Temperatur 16-20° C.
0.0067
4x10 | 0,0135
N.B. NH,NO, als N-Quelle. Entwickelung wie vorige.
ss mm” || _—
Gehalt in Säure Säure pro 1g.
Grum Mol, Gew. % in g. Acidität . as cu | Pilzsubstanz.
= 0 0.119 10.0 0.315 | 2.644
0.0017 0.188 12.7 0.400 ! 2.128
0.0034 — —— —
|
EINIGER ALGEN UND PILZE. 181
F. II. Kulturen mit Zusatz von HgCI,.
Aspergillus niger—angestellt 1. Febr. ’99.
Geerntet 9. Febr. Kulturdauer 8 Tage. Temperatur 16-20° C.
a Gehalt in Ernteertrag Säure ‘Siure pro lg.
Gram Mol. | Gow. % in g. _ Acidität Tanuctechace | Pilzsubstanz.
it we 0.153 11.3 0.356 2.366
ys x 10° 0.0017 0.160 13.7 0.431 | 2.568
1 x 10 0.0034 — — —- —
41x10” | 0.0067 — — rl =
en ——
4x10 | 0.0135
N.B. NH,NO, als N-Quelle. Entwickelung wie vorige.
F. IV. Kulturen mit Zusatz von HgCl,.
Penicillium glaucum—angestellt 20. April.
Geerntet 1. Mai. Kulturdauer 11 Tage. Temperatur 16-20° ©.
| Gehalt in Ernteertrag Säure Siure pro 1g.
Gram Mol. | Gow. X in g. Acidität als Oxalsture | Pilzsubstanz.
ge I © 0.203 4.5 0.142 | —
psx 10= | 0.0017 0.243 4.7 0.148 =—
1x10 | 0.0034 | 0.242 5.1 0.159 —
1 x 10 0.0067 0.473 5.1 0.159 —
|
0.0135 0.251 4.9 0.154
N.B. NH,NO, als N-Quelle.
F. V. Kulturen mit Zusatz von HgCl,.
Penicillium glaueum—angestellt 20. April.
Geerntet 2. Mai. Kulturdauer 12 Tage. Temperatur 16-20° C.
| (rehalt in Ernteertrag Säure Siure pro le.
a | | in g. als Oxalsäure | Pilzsuhstanz.
| Grom Mol. | Gew. % in g Aciditat umgerechnet ec
a Te: 0.222 4.6 0.145
4x10 | 0.0003 | 0.282 4.6 0.145
41x10 | 0,0006 | 0.264 4.7 0.148
41x10“ | 0.0013 | 0.273 4.6 0.145
10> 0.0027 | 0.301 4.5 0.142
N.B. NH,NO, als N-Quelle.
152
N.
ONO: WACHSTHUMSBESCHLEUNIGUNG
F. VI. Kulturen mit Zusatz von HgCl,.
Penicillium glaucum—angestellt 20. April.
Geerntet 1. Mai.
Gehalt in
Gram Mol.
1 x 107
ly In
4 x 10
4 x 10
1,
Gew. %
0
0.0003
0.0006
0.0013
0.0027
Ernteertrag
in g.
0.183
0.249
0.213
0.311
0.246
Kulturdauer 11 Tage.
Aciditat
N.B, NH,NO, als N-Quelle.
Temperatur 16-20° C.
Sa re
als Oxalsiure
umgerechnet
0.157
0.173
0.185
0.173
0.182
| Siure pro le
| Pilzsubstanz
—
———
F. VII. Kulturen mit Zusatz von HgCl,.
Aspergillus niger—angestellt 30. März.
(eerntet 18. April.
(iehalt in
Gram Mol. | Gew. %
0
0.0003
0.0006
0.0013
0.0027
Kulturdauer 18 Tage.
Ernteertrag
in g.
0.347
0.517
0.513
0.552
0.565
Aciditat
8.8
9.6
9.4
N.B. NH,NO, als N-Quelle.
Temperatur 16-20° C.
Säure
als Oxalsäure
umgerechnet |
0.277
0.302
0.296
0.343
0.356
F. VIII. Kulturen mit Zusatz von HgCl,.
Aspergillus niger—angestellt 30. März.
Geerntet 18. April.
Gehalt in
Gram Mol.
Ü
LCA
1x 10
4x 10
101
Gew. %
0.
0.0003
0.0006
0.0013
0.0027
Kulturdauer 19 Tage.
Ernteertrag
in g.
0.341
0.458
0.474
0.630
0.429
N.B. NH,NO, als N-Quelle.
Acidität
8.6
9.4
9.4
12.5
10.3
Siiure pro 1,
Pilzsubstan:
0.800
0.586
0.959
0.621
0.630
Temperatur 16-20° C.
Siure
als Oxalsäure
umgerechnet |
0.271
0.296
0.296
0.394
0.324
| Säure pro 1
Pilzsubstan
0.795
0.646
0.624
0.625
0.459
EINIGER ALGEN UND PILZE. 183
F. IX. Kulturen mit Zusatz von HgCl,
Aspergillus niger—angestellt 30 März.
Kulturdauer 19 Tage.
Geerntet 18. April. Temperatur 16-20° C.
Säure
x Gehalt n Ernteertrag | Säure pro 1g.
Gram Mol. | Gew. & in g. | Acidität | a nn Pilzsubstanz.
Br ıl =D 0.261 ° 84 | 0.265 1.015
1x10 0,0003 0.355 | 9.2 0.2% 0.816
41x10 0.0006 0.380 | 9.4 0.296 | 0.779
4x 10 0.0013 0.509 | 121 0.381 0.742
10 _ 0.0027 | 0.451 11.1 0.350 0.776
N.B. NH,NO, als N-Quelle.
G. I. Kulturen mit Zusatz von LiNO,.
Aspergillus niger— angestellt 1. April.
Kulturdauer 17 Tage.
Geerntet 18. April. Temperatur 16-20° C.
| Gehalt in | | Ernteertrag | Säure | Säure pro ly. |
Gram Mol. | Gew. % in g. | Acidität als le Pilzsubstanz. |
7 © | 0 : 030 9.0 0.284 | 0,946
14x10 0004 | 0.408 9.4 0.296 0.725
41x10 | 0.008 | 0.428 8.9 0.283 | 0.661
2x10 | 0.017 | 0.348 8.7 0.273 0.784
1x10 | 0.034 | 0.345 8.6 0.271 0.782
N.B. NH, NO, als N-Quelle.
H. I. Kulturen mit Zusatz von NaFl.
Aspergillus niger—angestellt 7. Febr. ’99.
Geerntet 21. Febr. Kulturdauer 14 Tage. Temperatur 16-20° C.
Gehalt in Ernteertrag
Gram Mol. Gew. £ in g. Acidität | ee
0 0 0.199 | 8.8 0.277 1,392
ds x10 | 0,0025 0.325 9.4 0.296 0.911
41x10 | 0.005 | 0,312 72 0.227 0.727
4x 10° 0.010 0.246 6.1 0.192 0.880
41x10 0.021 0.289 6.0 0.189 0.654
N.B, NH,NO, als N-Quelle,
184 N. ONO: WACHSTHUMSBESCHLEUNIGUNG
H. II. Kulturen mit Zusatz von NaFl.
Aspergillus niger—angestellt 7. Febr. ’90.
Geerntet 23. Febr. Kulturdauer 16 Tage. Temperatur 16-20 C.
_ Gehaltin | Exnteertrag Säure Giure pro 1
Gram Mol. Gew. £ in g. Acidität et Pilzsubstan
o [| ® 0.314 10.0 0.315 | 1.000
74x10 | 0.0025 0.336 10.0 0.315
4x 10° 0.005 0.385 7.6 0.239
1x 10 0.010 0.316 5.9 0.186
0.274 5.6 0.176
4x10" | 0.021
| N.B. NH,NO, als N-Quelle.
H. ITI. Kulturen mit Zusatz yon NaF.
Aspergillus niger—angestellt 7. Febr. ’99.
Ge”rntet 25. Febr. Kulturdauer 18 Tage. Temperatur 16-20° C,
} Gehalt in Ernteertrag Säure Säure pro 1
Gram Mol, | Gew. € in g- Acidität | ee | Pilzsubstan
O Ü 0.270 8.4 0.265
x10® | 0.0025 | 0.280 10.7 0.339 1.207
1x 10-7 | 0,005 0.285 7.7 0.242 | 0.849
1x 107 0.010 0.264 6.6 0.208 | 0.788
3x10= | 0.021 | 0.265 6.1 0.725
N.B. NH,NO, als N-Quelle.
EINIGER ALGEN UND PILZE.
INHALT.
I. Einleitung und Litteratur.
II. Methodisches,
III. Vorbemerkungen über die Versuchsobjekte.
IV. Veränderungen in der Wachsthumsweise und die Correla-
tion zwischen Fortpflanzung und Wachsthum.
V. Einfluss der Reizstoffe auf die Betriebsstoffwechsel.
VI. Specielle Besprechungen.
VII. Schlussbemerkungen und Zusammenfassung der Resultate.
TABELLARISCHE ZUSAMMENSTELLUNG.
I. Versuche mit Algen.
II. Versuche mit Pilzen.
185
186
Erklärung der Tafel XII.
Kulturen von Aspergillus niger mit und ohne Zusatz von NaFl.
(Photographiert 15 Tage nach der Sporenaussaat.). .
I. Ohne Zusatz ; Kontrolikultur. -
IL Mit Zusatz von 0.0025% NaFl.
III. Mit Zusatz von 0.005% __,,
IV. Mit Zusatz von 0.010% __,,
V. Mit Zusatz von 0.021% ,„
(Für näheres vgl. 8. 153 und ferner Tabelle Pilz. H.)
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CONTENTS OF RECENT PARTS.
Vol. IX., Pt. 1... yen 0.62 (Price in Tokvô).
On a Certain Class of Fraunhofer’s Diffraction-l’henomenn. By H. NaGaoxka.
Lines of Equal Intensity about the Point of Intersection of Fraunhofer's Dif-
fraction Bands. By H. NAGAOKA.
Note on Tinfoil Grating as a Detector for Electric Waves. By T. Mizuno.
The Thermo-electric Effects of Longitudinal Stress in Iron. By K. TsuRuTA.
Thermo-electrie Effects of Longitudinal Tension in Different Metals. By K.
Tsurvta.
Notes on the Topas from Mino. By T. Hıkı.
Mercury Perchlorates. By M. CHIKASHIGE.
Potassium nitrososulphate. By E. Divers and T. Haca.
Sodium nitrososulphate. By E. Divers and T. Haca.
The Constitution of the Nitrososulphates. By E. Divers and T. Haca.
Vol. IX., Pt. 2... yen 1.31 (Price in Tokyo).
The Tinfoil Grating Detector for Electric Waves. By T. Mizuno.
On the Piedmontite-rhyorite from Shinano. By N. Yamasaki. (With Plate VI).
The Atomic Weight of Japanese Tellurium. By M. CHIKASHIGE.
Das Johanniskäfer-Licht. Von H. MuRAOKA.
On the Prediction of Solar Eclipses. By SHIN HIRAYAMA.
How Mercurous and Mereuric Salts change into each other. By S. Hapa.
Imidosulphonates (Second paper). By E. Drvers and T. Haga.
Amidosalphonie acid. By E. Divers and T. HaGa.
Moleenlar Conductivity of Amidosulphonie Acid. By J. SAKURAI.
The Physiological Action of Amidosulphonic Acid. By Oscar LoEw.
The Reduction of Nitrososulphates. By E. Divers and T. HaGa.
Economic Preparation of Hydroxylamine Sulphate. By E. DIVERS and T. Haga.
On the Time-Lag in the Magnetination of Iron. By Y. Kato. (With Plates
VII-XV).
Vol. IX., Pt. 3... yen 0,65 (Price in Tökyö).
Diffraction Phenomena in the Focal Plane of a Telescope with Circular Aper
ture due to a Finite Source of Light. By H. Nacaoxa. (With Plate XVI &
XVII).
Researches on Magnetostrietion. By H. Nagaoka and K. Honpa. (With Plates XVIII
& XIX).
Vol. X. Pt. 1... yen 1.80 (Price in Tokyo).
On the Fate of the Blantopore, the Relations of the Primitive Streak, and the
Formation of the Posterior End of the Embryo in Chelonia, together with
Remarks on the Nature of Meroblastic Ova in Vertebrates. (Contributions
to the Embryology of Reptilia. V.). By K. Mirsuxurz (With Plates I-XT).
Vol. X. Pt. 2... yen 1.20 (Price in Tökyö).
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CONTENTS.
Vol. XIII, Pt. I.
Notes on the Geology of the Dependent Isles of Taiwan.
By B. Korö, Ph. D., Rigakuhakushi; Professor of Geology,
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THE
JOURNAL
OF THE
COLLEGE OF SCIENCE,
IMPERIAL UNIVERSITY OF TOKYO,
JAPAN.
VOL. XIIL, PART IT.
Rh f WK # A Mt
PUBLISHED BY THE UNIVERSITY.
TOKYO, JAPAN.
1900.
MEII X XXIII.
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[ee
Prof. K. Yamagawa, Ph. B., Rigakuhakushi, Director of the Colle;
(ex officio).
Prof. J. Sakurai, Rigakuhakushi.
Prof. B. Kot6, Ph. D., Rigakuhakushi.
_ Prof 1. Ijima, Ph. D., Rigakuhakushi.
All communications relating to this Journal should be addressed to the
Director of the College of Science.
Edward Divers and Masataka Ogawa,
Imperial University, Tokyo.
7
he. interaction of such familiar gases as ammonia and
ir d dioxide ceased to attract with any effect the attention of
gators sixty years ago and more, Yet comparatively
g had then been definitely made out about the nature of
ai en and even the few statements eoncerning it in some
t treatises on chemistry have but little experimental
The history of the subject is briefly given on p. 195.
M union of dry sulphur dioxide and ammonia.
=
=
u
en when comparatively well-dried, sulphur dioxide and
nia anite at ones and with great energy when brought
= ye they can remain mixed without combining, pro-
8 ay nt care has been observed to exclude moisture. It
yt Le necessary, however, in order to demonstrate this
phenom non, to have resort to the elaborate precautions
on,
PA J a
Digitized
188 E. DIVERS AND M. OGAWA:
adopted by Brereton Baker, in his famous experiments upon the
non-union of hydrochloric acid and ammonia (J. Ch. Soe., 1894,
65, 611; 1898, 73, 422), and we have only dried the gases
during their flow through the tubes. As we were able to dry
sulphur dioxide better than ammonia, because common phos-
phorus pentoxide could be used for the purpose, we have had
success in mixing the gases without their combining only by giving
this gas precedence. The preparation flask with its tubes having
been heated and then kept for a while in the desiccator, was
placed in ice and salt while a slow current was sent through it
of sulphur dioxide, which had passed through drying-tubes of
sulphuric acid and then of phosphorus pentoxide. The outlet-
tube dipped into mercury. Ammonia, dried first by the cold of
a freezing-mixture and then by long tubes of freshly fused and
crushed potassium hydroxide (but no Stas’s mixture), was now
also passed into the flask slowly. The result was that the in-
terior of the flask remained clear for some minutes, the mixed
gases only combining on their escape through the mereur in) |
the air. But the ammonia having, it is presumed, gra |
brought enough moisture with it through passing more 1 9 "
along the tubes than at first, the walls of the flask became
denly coated with an orange-coloured deposit, while the m
rose high in the exit tube.
Proportions in which sulphur dioxide and ammonia combine.
The proportions, in which ammonia and sulphur dioxide
combine, or appear to combine together, depend largely upon
the extent to which the temperature is allowed to rise, the heat
of union being considerable, They vary also according as one
Digitized by Google
AMMONIUM AMIDOSULPHITE. 189
er of the gases is used in excess, unless the temperature is kept
low. But the variation of the proportions and the apparent
nsation of additional sulphur dioxide by a sufficiently am-
ted product, that may be observed, are results clearly due
> secondary changes going on (p. 192). The simple union
amonia and sulphur dioxide, which can be secured by
ng down the temperature by suitable means, especially with
mmonia in excess, is that of two volumes of the former to
f the latter (p. 191). But since this union cannot be
at the ordinary temperature without being immediately
ed by a decomposition, in which ammonia is evolved, the
of the two gases can appear to take place in other pro-
ms than the above. It is pretty certain that, by proceeding
y enough and using strong cooling agencies, secondary ac-
could be almost entirely prevented and the statement just
be verified, even when working with the gases alone. We
not gone very near to getting such a result in this way,
hen we have, for good reasons, not striven much to over-
the difficulties. Our experimental work, which will be
er on referred to (p. 195), has shown that two much more
y than one volume of ammonia can be made in this way
lite with one volume of sulphur dioxide, the only propor-
which Rose met with in his experiments (p. 193), and that
jresence of much ammonium amidosulphite in the product
be established with certainty.
Preparation and analysis of ammonium amidosulphite.
In order to get the primary product of the union of sul-
‘ dioxide with ammonia in its unchanged state, ether was
|
|
|
190 E. DIVERS AND M. OGAWA :
made use of as the medium of the union, in order to keep tl
temperature under control. The ether, freed from alcohol ar
water by sodium, was contained in a small flask, fitted wi
inlet and outlet tubes, which was to serve, not only for the pr
duction of the new substance, but for its isolation and its weigl
ing for analysis. The flask was put in a bath of ice and sa
with the outlet-tube dipping into a trough of mercury, and th
the ether was saturated with dried ammonia. Having shut «
the ammonia, a very slow current of sulphur dioxide was se
into the solution while the flask was continuously shaken, n
only in order to diffuse the heat, but to prevent the produ
from caking on to the bottom of the flask and shutting in ethe
The mouth of the tube conveying the sulphur dioxide so
became filled with a yellow pasty mass (p. 192), and had to
kept open by a platinum rod, manipulated through the rubb
tubing above, but the precipitate itself was quite white and por
dery. In spite of the external cooling, the heat of combini
was sufficient to cause ammonia gas, saturated with ether-vapot
to escape through the mercury sealing the exit-tube, and wh
this escape became slight, the passage of sulphur dioxide stoppe
With the use of about 20 c.c. ether, there had then formed we
over a gram of the substance. In order to secure this undecor
posed, a second flask was put in connection with the preparatio
flask, and ammonia again passed to the saturation point. TI
ammoniated ‘ether was decanted off through the connecting-tu
into the second flask, which was then detached, the whole oper
tion being carried out in the freezing-mixture. The current
ammonia was renewed over the precipitate in the flask, a
continued for hours, until all the ether adhering to the precif
tate had been carried away, the flask being all the while sti
AMMONIUM AMIDOSULPHITE. 191
freezing-mixture, There was no other way of completely
the salt, and even this way was not sufficiently successful
he salt had been allowed to cake together. The ammonia
not be replaced by air or hydrogen for drying the salt,
uld the flask be kept out of the freezing-mixture, so long
r still moistened the salt, without the latter taking an
Colour. When dry and in an ammouical atmosphere,
; is more stable, but cannot long be kept at the ordi-
emperature without getting discoloured through decom-
1.
nalysis.—The stopper carrying the gas-tubes having been
d by a plain one, and air allowed to displace most of the
ia gas, the flask was at once weighed and left for a time
d with open mouth dipping into 100 c.c., or more, of
in a beaker. When the salt in it had become damp, it
shed into the water, and its very dilute solution distilled
lkali for its ammonia. The residue was divided into two
ed portions, one of which was acidified and heated to
nder pressure for some hours and then redistilled with
for additional ammonia, of which only a trace was got
per cent. of the salt). The other part of the solution was
with bromine, and next with hydrochloric acid and
e, after which barium sulphate was precipitated with the
precautions. The results of the analysis were :—
Ammonia Sulphur diox.
Found: — 35.09 ; 64.91 per cent.
SO.(NH;): 34.69 ; 65.31 ,,
he slight excess of ammonia indicated is safely attributable
means taken to preserve the salt till it was analysed.
192 E. DIVERS AND M. OGAWA:
Its properties, constitution, and name.
The new salt is white and apparently crystalline, an
pears to be slightly volatile in a current of ammonia.
very deliquescent and decomposes, losing ammonia, in th
It dissolves in water, giving out heat and a hissing sound
if dissolved by ice or enough ice-cold water, furnishes a so]
answering all tests for pure ammonium sulphite. In this rı
it is quite unlike ammonium amidosulphate or carbamate,
even the latter salt gives at first no precipitate with ca
chloride, which at once precipitates all sulphite from thi
salt. When the salt is much decomposed, its solution
other reactions besides those of a sulphite. In anhydrous a
it dissolves freely, evidently as ethyl ammoniumsulphite :
also slightly soluble in dry ether. It soon begins to chang
then assumes an orange-colour, even at the common temper
At 30-35° it decomposes into a liquid and a solid part,
more or less orange-coloured, and into ammonia, the liqui
undergoing further change into solid matters (p. 197)
Constitution.—The salt is more probably an amido- th
imido-compound, NH,N(SO,NH,). (analogue of normal 2
nium imidosulphate), because it can be obtained only whe
temperature is kept down and the ammonia is in excess.
still more probably a sulphuryl rather than a thionyl com
because of its feeble activity as a reducing agent and of it
easy passage Into ammonium sulphite or ethyl ammoniumsu
It has accordingly to be formulated as NH,SO;, NH, an
NH, SO'ONH..
Name.—Since the salt represents ammonium sul
NH,O'SO,NH,, in which the ammonoxyl is replaced by a
AMMONIUM AMIDOSULPHITE. 193
is properly called ammonium amidosulphite. Berglund’s
of amidosulphonate now in use, for amidosulphate is
ly based on a misconception. The name, amidosulphinate,
logy with amidosulphonate, must be rejected on the same
s, and because the salt has not the characteristic reducing
and the constitution of sulphinates. It does not seem
2, even were it desirable, to construct a term for the first
of sulphurous acid that would correspond to that of sul-
acid, the synonym of amidosulphuric acid.
ure of the decomposition by heat of the amidosulphite.
istory.—Experiments made earlier than ours on the union
hur dioxide with ammonia gave the products of decom-
1 of ammonium amidosulphite instead of the salt itself.
einer in 1826 (Schw, Jahrb., 17, 120), described the pro-
f the union as a brown-yellow vapour quickly condensing —
right brown solid mass, which the smallest quantity of
converts into (colourless) ammonium sulphite. Rose pub-
three papers on ‘anhydrous sulphite of ammonia’ in
837, and 1844 (Pogg. Ann., 33, 235; 42, 415; 61, 397),
second correcting statements made in the first, and
ing in the third the views he had expressed in the earlier
The outcome was that he had ascertained that the pro-
f the union is always one and the same single substance,
atever proportions the dry gases are taken; that it is
ed of equal volumes of the gases, is either yellowish-red
neary, or red crystalline, very deliquescent and very
/ in water without evolving ammonia; that it yields a
| solution, which is at first yellowish but soon, becomes
= — - =
194 E. DIVERS AND M. OGAWA:
colourless, and gives, when recently prepared, the reactic
mainly of a mixture of ammonium sulphate and trithionate, |
to a small extent those of a sulphite also; and, lastly, that wk
the solution is of certain concentration it gives a transient r
dish coloration with hydrochloric acid.
Forchhammer (Compt. rend., 1837, 5, 395) found that, |
sides the orange-coloured substance, crystals of ammonium sulph
are produced by the union of the gases, which can sometimes
seen apart from the other product in some spots of the m:
though often indistinguishably mixed up with it. (That
crystals observed in the product were those of sulphate, co
only have been a supposition of Forchhammer’s). The m
when moistened is alkaline and evolves ammonia, yielding oth
wise the reactions recorded by Rose. Absolute alcohol dissol
out of it a substance which takes a rose colour, soon disappe
ing. Indirectly, he represented the mass to be derived from
mols. ammonia to one mol. sulphur dioxide, as did also D
berenier. |
The views advanced as to the nature of the orange b
have been, that it is a compound of ammonia with an iso.
of sulphurous anhydride, which changes at once with we
into ammouium sulphate and trithionate, just as ammonium py
sulphite slowly changes in hot solution (Rose) ; that it is ami
gen sulphide, S(NH3),, mixed with ammonium sulphate (For
hammer) ; that it is, partly, thionamic acid, NH,SO'OH, par
ammonium thionamate, both volatile, being its colour due
impurity (H. Watts); and that it is ammonium pyrothionam
NH; SO, NH, (Joergenssen).
Interaction of the gases—We have repeated Rose’s exp
ments of measuring over mercury the volumes of the gases wl
AMMONIUM AMIDOSULPHITE. 195
ct, in which he found that always equal volumes combine,
ever gas may be taken in excess. The results somewhat
ached this when no steps were taken to restrain the rise in
rature due to the union of the gases; but when the gas-
vas immersed in a cooling-mixture and the ammonia was
ess, the volume of this gas consumed was much greater
that of the sulphur dioxide. This method of investigating
natter is, however, inapplicable, because the ammonium
sulphite, which is formed, partly decomposes with free
ion of ammonia. By letting the dried gases come together
essel agitated in a freezing-mixture and keeping the am-
‚in excess, a solid mass is obtained which consists largely
; amidosulphite, behaving as such in water, though mixed
other substances, and quantitative analysis of which shows
nuch more than three mols. ammonia to two mols. sulphur
le have gone to its formation. If, instead of examining it
2e, it is kept for a long time in a gentle current of dry
en or hydrogen, at a temperature of 30° to 35°, it no longer
ns amidosulphite or gives any sulphite to water, and contains
uch more than one atom of nitrogen to one of sulphur.
Rose’s results are explained and, at the same time, shown
of no direct significance.
Products of the decomposition.—Both Rose and Forchham-
ound ammonium sulphate to be a principal constituent of
roduct of the interaction of the gases. A sufficiently high
rature having been reached, this will have been the case ;
more, the solution of the even less heated product slowly
1es acid and full of sulphate. But when the temperature
ot been allowed to exceed 30°, or even 40°, the quantity of
ate in the product is so small that it may almost be dis-
196 E. DIVERS AND M. OGAWA:
regarded. Along with sulphate, trithionate was considered
Rose to make up most of the product, for the aqueous solut
of the mass always gives a strong reaction with silver nit
which might be that of trithionate and, in the case of his y
duct, gave other reactions of a trithionate. But when the y
duct has been carefully prepared and is free from amidosulph
its solution gives the silver reaction without the others belong
to a trithionate. Thus, the solution may be acidified and
for hours without yielding more than mere traces of sulp
dioxide and sulphur ; to get these in quantity, the solution ha
be strongly heated under pressure. Besides this, the absence
sulphate in the solution is of itself almost enough to dispr
the production of trithionate, since, as Rose himself represen
sulphate and trithionate are complementary products of the
com position.
Heating pure ammonium amidosulphite gives the same
sults as heating the coloured product of the union of sulp
dioxide and ammonia as gases. Rose’s assertion that the proc
is a single substance, even in appearance, is certainly incorr
according to our experience. By the union of the gases 1
receiver kept well cooled, the product is deposited as a &
waxy, yellow coating on the walls of the vessel and on the
tubes. Its colour varies in different parts from nearly white
orange-red somewhat irregularly but generally so as to be wh
near where the ammonia enters, the whiteness not being duc
moisture in the gases, as Rose assumed. When the product
to 30-35°, whether by its own heating or by external heat, |
decomposed at first into an obscurely crystalline white solid.
a much smaller quantity of a coloured, effervescing liquid, pa
draining to the bottom of the vessel; after a time all beco
AMMONIUM AMIDOSULPHITE. 197
vain and tenaciously adherent to the glass. When pure
ium amidosulphite is similarly heated in a dry inactive
colours, softens, shrinks together, vesiculates, gives out
ia, and becomes a mass like that derived directly from
ion of the gases. With very gradual heating, the tem-
y liquid product is much less coloured than in the other
ts colour being evidently caused by the presence of a
utter dissolved in ‘it, which gives indications of being
is orange-red substance is never formed in more than
nall quantity. It gives a yellow colour to the aqueous
1 of the whole product, which, however, slowly fades away.
|, carbon bisulphide, and other menstrua dissolve it out
1e salts, leaving them white; but the solutions are not
The yellow solution in water or alcohol takes a transient
lour when mixed with dilute hydrochloric acid, and the
ic solution an indigo-blue colour with concentrated am-
The residue left on evaporating the carbon-bisulphide-
1 becomes explosive when heated above 150°, and may
ave become nitrogen sulphide, but before being heated it
this substance.
ccept the very little sulphate already mentioned, there is
yet known substance present in the residue of the decom-
1 of the amidosulphite by a gentle heat, so far as we can
r. Alcohol of 90 per cent. dissolves out something, but
ery sparingly. By evaporation of the solution in a vacuum
tor, a very deliquescent salt is obtained in crystals, having
osition that may be expressed by 9NH;, 8SO,, assuming
sence of 2.5 per cent. moisture. The composition of the
crude residue does not differ much from this. The alco-
198 E. DIVERS AND M. OGAWA:
holic solution, cooled and charged with ammonia, gives mint
scaly crystals in small quantity. This substance, dried in
current of ammonia, has a composition expressed by (NH;),S.
and dried in the sulphuric-acid desiccator, that of (NH;).S.¢
These three substances all give the silver-nitrate reaction of t
aqueous solution of the whole residue, and on boiling with dilu
hydrochloric acid give very little sulphur and no sulphur dioxic
At higher temperatures, whether dry or in solution, they yie
sulphur, sulphur dioxide, and sulphate. Two potassium deriv
tives of these salts have also been prepared. Neither the cru
residue nor any of the above substances yields all its nitrogen
ammonia when distilled with alkali, unless it has been fi
heated with hydrochloric acid under pressure.
From the mother-liquor of the above mentioned 8,0, salt
substance was got which in composition and behaviour appear
to be sulphamide a little impure. Neither sulphamide nor amid
sulphate can be found in the fresh aqueous solution of the whc
residue, but, by heating the solid residue itself to a high
temperature, imidosulphate is obtained in considerable quantit
besides sulphur and sulphate, and imidosulphate is a knov
product of first heating and then dissolving in water, eith
amidosulphate or sulphamide. A proof-spirit extract and also
wood-spirit extract of the residue yield ammonium amidosulpha
on evaporation, no doubt generated by hydration. An aqueo
solution of the less heated residue, treated with excess of bariu
acetate and filtered, gave barium thiosulphate in crystals, «
evaporating it over the water-bath.
During the heating of ammonium amidosulphite at a ten
perature of 30° to 35°, besides much ammonia, small quantiti
of water and of sulphur dioxide are evolved, the former main
AMMONIUM AMIDOSULPHITE. 199
early stage and the latter in the late stage of the de-
sition. This remarkable production of water, though
s evident, was fully established by cooling the escaping
and testing the water thus collected. The presence of
ir dioxide later in the operation was shown by the gases
g on their escape into the air and then forming a small
deposit, slowly turning orange, and reacting as ammonium
ılphite. In the interaction of sulphur dioxide with ammo-
nd in the decomposition of the amidosulphite, no liberation
rogen could ever be discovered.
o sum up the results of our incomplete work upon the
position of ammonium amidosulphite by a graduated and
heat, ammonia and a residue consisting of a substance
bstances), which behaves as a thio-amido-sulphonic com-
, are the principal products; in much less quantities, water
n orange-red substance are also produced, and, generally if
ways, a very little sulphate; and, as secondary products,
ently sulphamide and certainly amidosulphate and thiosul-
are obtainable, as well as imidosulphate, sulphur, and
sulphate. It seems of interest to point out that we here
| the first production known of amidosulphate from am-
and sulphur dioxide, which, hitherto, has been derived
from ammonia and sulphur trioxide or from a nitrite and
ır dioxide.
Ve hope in a future paper to be able to report the com-
n of this investigation.
=
00
©
©
Q
Products of heating Ammonium Sulphites,
Thiosulphate, and Trithionate.
By
Edward Divers and Masataka Ogawa,
Imperial University, Tökyö.
‘hat has been published upon the effects of heating am-
m sulphites and thiosulphate is but little in accordance
he results of experiments we have had to make upon these
nd upon the hardly known trithionate, in connection with
estigation of the decomposition by heat of ammonium
ulphite. We therefore make known what we have ascer-
Preparation of the salts used.
mmonium sulphite, (NH,), SO,, OH:.—Statements are con-
as to whether this salt can be got from its solution by
ation (Muspratt, Phil. Mag., 1847, iii, 30, 414; Marignac,
b., 1857, 17; Forcrand, Compt. rend., 1885, 100, 245;
„ Compt. rend., 1887, 104, 1793 ; Roehrig, J. pr. Ch., 1888,
7). We find that a concentrated solution, charged with
- a
202 E. DIVERS AND M. OGAWA: PRODUCTS OF HEATING
ammonia, can be quite successfully made to deposit the salt
cold evaporation in a potash desiccator, but to get such a solut
the moderately strong solution of ammonia, which must be u:
has to be kept very cold while passing in the sulphur diox
-Dilute solutions fail to yield the salt on evaporation because
much of it suffers decomposition. Much better than evapora!
is to take advantage of the lessened solubility of the salt
presence of much ammonia. Ammonia solution, sp. gr. 0.£
containing therefore about 28 grm. ammonia in 100 c.c., is tt
treated in a flask with sulphur dioxide, while it is kept in :
tion in a mixture of ice and salt, and with the tube convey
the sulphur dioxide not dipping into the solution. The format
of a very little orange-coloured matter in the neck of the fi
cannot be avoided, but this can be easily removed afterwa
When the solution has become thick with crystals, no m
sulphur dioxide is to be added, although very much amm«
still remains. Even at the common temperature the crystals
not sensibly dissolve in presence of this ammonia. The s
drained on a tile under close cover, can be dried either
filter paper or by only short exposure in the desiccator
potassium hydroxide or carbonate, salted just before with :
monium chloride. It is equivalent in quantity to about c
fourth of the ammonia taken. By long exposure in a d
atmosphere the salt becomes anhydrous without loss of ammo
Exposed to the air, it is apparently deliquescent but in res
it evolves ammonia and thus becomes the very deliques
pyrosulphite.
Anhydrous ammonium sulphite is readily obtained from
hydrated salt by long enough exposure in the desiccator; 1
very hygroscopic.
\MMONIUM SULPHITES, THIOSULPHATE AND TRITHIONATE. 203
immonium pyrosulphite, (NH,).8.0;.—When, in the process
iven for preparing the normal sulphite, the passage of
ir dioxide is not stopped when the solution is full of crystals,
gradually dissolve up and the solution becomes greenish-
Then, as it gets charged with sulphur dioxide, in the
x mixture, the pyrosulpbite crystallises out from it, in quan-
juivalent to a little over one-fifth of the ammonia taken,
thrown out of solution by the sulphur dioxide. The salt
> obtained dry and pure in the same way as the normal
te, except that sulphuric acid, to which a little solid alkali
te has been added, is used in the desiccator, though it is
leliquescent and changeable when not carefully preserved
poisture. This salt is also easily obtainable by evapora-
s aqueous solution, but hardly free from sulphate, and not
t some decomposition, through loss of sulphur dioxide and
h oxidation. It is much more soluble than the normal
Le.
mmonium thiosulphate.—An old solution of calcium thio-
te, obtained by boiling lime and sulphur together in water
aving the solution until much of the pentasulphide had
yxidised by the air, was decanted from insoluble matters,
with ammonium carbonate in some excess, filtered, and
reely exposed to the air for some time at 50-60°. In this
very concentrated solution of ammonium thiosulphate was
ed, free from sulphate and other salts. The solution of
ery soluble salt was then dried up to a crystalline mass in
siccator. The well-dried crystals have been found by Lock
luess (Ber., 1889, 22, 3099) to be anhydrous.
immonium trithionate —This salt has apparently not hitherto
prepared by any one. Being exceedingly soluble in water,
204 E. DIVERS AND M. OGAWA : PRODUCTS OF HEATING
it cannot be prepared by Plessy’s excellent process for the :
tassium salt (Ann. Ch. Phys, 1844, iii, 11, 182), or by
slight modification by Hertlein (Z. phys. Ch., 1896, 19, 24
We therefore made the pure potassium salt by Plessy’s meth
precipitated the potassium from it by hydrofluosilicic acid, n
tralised quickly with ammonia, and precipitated the ammoni
trithionate by absolute alcohol and dried it in the desicca
This very deliquescent and changeable salt cannot be kept |
in good condition, but it was used by us when freshly prepa
and while still almost free from sulphate.
Effects of heating the salts.
The process.—The salts were heated in an oil-bath, i
subliming vessel consisting of a test-tube, 15 cm. long and ab
15 mm. in interna] diameter. The tube was closed by a cao
chouc stopper, and a very slow current of dried nitrogen throt
the tube was maintained during the heating and cooling. 1
salt, usually about 4 grm., was contained in an open slen
bottle, about 6 cm. long, having a platinum wire attached t
for lowering it into and lifting it out of the subliming tube. 1
tube was immersed in the oil to the level of the mouth of
bottle inside, so as to cause all dry sublimates to collect in
tube above this level. When, as in the case of the hydra
normal sulphite, the heating was divided into stages, the bo
was transferred between these to a second subliming-tube. 7
heating of the oil was conducted very slowly, so that the t
peratures mentioned which were those of the oil, may be acce
ed as being very nearly those of the salts at the time.
In describing the effects of heating them, the salts are tal
AMMONIUM SULPHITES, THIOSULPHATE AND TRITHIONATE. 205
> inverse order of that followed above, in accordance with
This is done because of the nature of the products.
immonium trithionate.—This salt is hardly affected until the
rature is above 150°, and at 160-170° it steadily decom-
into sulphur dioxide and a residue of ammonium sulphate
nfused sulphur. The non-fusion of the sulphur is remark-
nd only to be referred to the presence of minute quantitities
purities. It all dissolved readily in carbon bisulphide, and
llised out on evaporating the solvent.
t can hardly be doubted but that ammonium tetrathionate
pentathionate, if it can exist) would decompose in the same
3 trithionate. Ammonium hyposulphate (dithionate) has been
| by Heeren (Pogg., 1826, 7, 55), and more definitely by
3 (Ann., 1888, 246, 194) to first become anhydrous, if not
y so when heated, and then to decompose at about 130°
ulphur dioxide and a residue of ammonium sulphate.
immonium thiosulphate.—Zeise, in 1824 (Gm. Hbk) found
alt to be converted by heat into water, ammonia, and a sub-
. of sulphur, much thiosulphate again and sulphite, and a
sulphate. This result must have been obtained by rough
g. A much more weighty statement is that made by
& (Ber., 1874, 7, 1159), namely, that the dry salt can be
ned unchanged, intermediate dissociation being admitted.
ave found it to decompose very slowly at 150°, the main
cts being a sublimate of anhydrous normal sulphite and
due of sulphur unfused, as in the case of the trithionate.
also very small quantities of hydrogen sulphide and am-
| passed off in the current of nitrogen, and the sublimate
ined a very little of a salt having some of the properties
thionate and which did not strike the violet colour with
206 E. DIVERS AND M. OGAWA: PRODUCTS OF HEATING
ferric chloride given by a thiosulphate. Analysis of the sul
mate and of that part of the salt which remained mixed w
the sulphur when the progress of the decomposition was arre
ed after only half of it had been decomposed, gave results tl
showed the former to be essentially anhydrous normal sulph
and the latter unchanged thiosulphate :—
Ammonia Sulphur
(NH,),SO, 29.31 27.59 per cent,
Sublimate 27.04 27.95 __,,
(NH,) 80; 22.27 43.24 ,,
Residue 20.69 42.31 ,
The main decomposition of the thiosulphate is in full agr
ment with the relation of thiosulphates to sulphites. Very
teresting is the production of a little ammonia and hydro;
sulphide, in connection with the relation of trithionate to thios
phate as its thio-anhydride (Spring) :—2(NH,).S,0O,=2NH
SH,+(NH,).S,0,. When ammonium thiosulphate is rapidly 3
more strongly heated, ammonia is lost and sulphur sublimes; tl
as a matter of course and of no significance, thiosulphate :
even trithionate are produced on adding water to the mi
sublimates.
Ammonium. pyrosulphite—We did not get this exceedin
deliquescent salt into the tube ready for heating before it |
condensed some moisture, and to this we attribute part of the rest
obtained. Change went on slowly in the salt at 130° and sor
what faster at 150°. At first there was little else than a slight
steady evolution of sulphur dioxide, and this continued though v
feebly, to the end and while a sublimate forming. The sublim
was pyrosulphite in one experiment; in another, it was this:
mixed with a very little anhydrous sulphite. But there wa
{MONIUM SULPHITES, THIOSULPHATE AND TRITHIONATE. 207
rable residue, more than one-third of the weight of the
en, consisting of sulphate, trithionate, sulphur, and ap-
y some tetrathionate. There was no sulphite or thiosul-
The tetrathionate, the sulphur, and the sulphur dioxide
ry probably derived from decomposition of trithionate
sture. From a consideration of the results it seems almost
ry to assume that perfectly dry pyrosulphite sublimes un-
1 (with no doubt intermediate dissociation), and that the
e of a little moisture causes it to decompose partly into
e and trithionate.
hydrous ammonium sulphite volatilises at about 150°,
> a sublimate of the same salt, or rather, a pseudosub-
for the salt surely dissociates when heated.
ydrated ammonium sulphite—According to Muspratt, this
volatilises when heated, no sulphate being produced, and
water, then much ammonia, and finally a sublimate which,
, from its properties, is ammonium pyrosulphite. We
d the following effects of gradually heating it in a very
irrent of dried nitrogen. At about 90°, the salt moistened
ape of ammonia became quite evident, and at a little
100° distillation of water also took place; both water and
ia continued to escape in noticeable quantities for 2'/, hours
when the temperature for some time had been 120°; up
, a very little sublimate only had formed and matters were
most at a standstill. The quantity of the salt heated was
4 grm., and this had now lost one-fifth ofits weight, the
having the composition expressed by (NH;),,.(SO,).(OH2);,
ent to a mixture or combination of the three salts, hydrat-
phite (39.4%), anhydrous sulphite (34.1%), and pyrosul-
(26.5%), dividing equally among themselves the sulphur
208 E. DIVERS AND M. OGAWA: PRODUCTS OF HEATING
dioxide. Some repetitions of the experiment gave almost the s
results. Calculation and the results of one experiment gave
following numbers :—
Ammonia Sulphur dioxide
(NE, (80, (0) 25.00 56.47 per cent.
Found 24.65 56.20 ,
If, in the formation of this complex, no longer lo
material quantities of ammonia and water, only these prod
had been given off, the residue should have been 84'/, per «
of the hydrated normal sulphite, whereas it proved to be |
more than 79 per cent., in consequence of volatitisation of &
of the (dissociated) salt, made manifest by the production
little sublimate.
After renewing the heating in a fresh subliming-tube, al
ing the temperature to rise slowly from 120° to 150°, the res
had almost all disappeared in two hours, while an abundant
sublimate had deposited. For some time during this heat
sulphur dioxide steadily escaped, but practically ceased to d
long before sublimation was finished. The residue left v
sulphur dioxide was no longer coming off, proved on analysis t
normal sulphite again, but only half hydrated, 2(NH,),SO,, (
The sublimate, also, now and at the finish, consisted
normal sulphite, apparently anhydrous though found to
little hydrated because it is very hygroscopic and had unav
ably some exposure to the air while it was being scarped ov
the tube into the weighing bottle.
Hydrated ammonium sulphite, therefore, becomes by
dual heating to 120° converted one-third into the anhyd
salt, and one-third into pyrosulphite, by loss of water and
monia ; and then the nearly stable complex of these salts
MMONIUM SULPHITES, THIOSULPHATE AND TRITHIONATE. 209
er third of the original salt becomes converted into the
anhydrous normal sulphite, between 120° and 150°, sul-
ioxide and water escaping. The presence of water is es-
to the occurrence of both changes; dry ammonium
phite partly sublimes as such at 150° and partly changes
lphate and trithionate, as already described. Heating in
n tube, and more rapidly, Muspratt’s results will be got,
n weter is more quickly expelled, and some pyrosulphite
posit as a sublimate.
Digitizedidy.
assium Nitrito-hydroximidosulphates and the
m-existence of Dihydroxylamine Derivatives.
By
Edward Divers, M, D., D. Sc., F. R. S., Emeritus Prof.,
and
Tamemasa Haga, D. Sc, F. C. &.,
Professor, Tökyö Imperial University.
ike potassium nitrate (this Journal, 7, 56), potassium
forms double salts with the potassium hydroximido-
tes (sulphonates), the non-recognition of whose existence
lowed mistaken notions to arise about the nature and the
sts of the sulphonation of nitrous acid.
olassium nitrite and 2/3 normal hydroximidosulphate, KNO,,
S0,K),.—The sparing solubility of 2/3 normal potassium hy-
nidosulphate in water is hardly affected by the presence of
ium nitrite and when a sufficient quantity of the salt has
lissolved by heat it crystallises out again almost pure on
& the hot solution, even though the water has also dissolved
as much as one-sixth of its weight of the nitrite. When
lution of the nitrite is stronger than this there crystal lises
212 E. DIVERS AND T. HAGA:
out instead of the hydroximidosulphate itself a combination «
with a molecule of the nitrite. The same double salt is :
formed in the cold when the hydroximidosulphate is triture
‚and digested with such a solution of the nitrite. Precauti
being taken against the hydrolysis of the unstable bydroximi
sulphate this salt can be dissolved at 70° in as little as 3.8 ti
its weight of a 22 per cent. solution of nitrite and by cooling
solution the double salt be got in crystals in quantity equiva.
to about 12/13 of that of the hydroximidosulphate.
While the hydroximidosulphate itself crystallises in |
rhombic prisms with 20H), its compound with the nitrite 1:
silky asbestus-like fibres which are anhydrous. The compo
salt is also not deliquescent although potassium nitrite alon
very deliquescent. There is nothing else in its properties wh
by to distinguish it from a mixture of its component salts.
can be recrystallised from a hot solution of potassium nitrit
a strength of 10 per cent. or more nitrite. It is neutral to
mus and very soluble in water but its solution soon dep
crystals of the 2/3 normal potassium hydroximidosul phate ‘un
it is very dilute. In any case the hydroximidosulphate car
precipitated and thus separated from the nitrite by the addi
of barium hydroxide. Like a simple hydroximidosulphate (
Journal, 7, 40), the solid salt digested with a highly con
trated solution of potassium hydroxide is converted into sulp
and nitrite. When acidified its solution becomes yellowish
a short time and then effervesces from the escape of nitı
oxide, a result of the hydroximidosulphate being a sulphon:
hydroxylamine, for hydroxylamine and nitrous acid decom]
together into nitrous acid and water, the other product in
present case being potassium acid sulphate only. It decomp:
POTASSIUM NITRITO-HYDROXIMIDOSULPHATES. 218
ively when heated—more so than does the hydroximido-
te by itself—giving off almost colourless gases and white
just as might be expected and just as does a dry mixture
constituent salts in corresponding proportions or a mixture
‘ite with a little sulphite.
he compound salt can be purified from other salts or from
when these are present by recrystallising from strong
1 potassium nitrite solution. But from its own mother-
it can be separated only by draining on the tile and not
shing. Such draining however is very effective because of
ited fibrous form of the salt, its non-deliquescent nature,
ie hygroscopic character of a solution of potassium nitrite.
nalysis of the salt was made in the usual way described
r previous papers on hydroximidosulphates and other
nated-nitrite derivatives. By boiling its solution with an
10st of its sulphur appears as ordinary sulphate, but not
11; so that in estimating the sulphur the solution must be
ysed for some honrs at 150° under pressure. The results
lysis were :— |
Potassium Sulphur
Found, 33.14 17.95 per cent.
K,HN,S,O,, 33.10 18.06 ,,
here are other ways in which the potassium nitrito-
rmal hydroximidosulphate may be formed all consisting
ally in producing the hydroximidosulphate by sulphonating
Il portion of the potassium nitrite in a concentrated solution.
the following mode of working will give good results with
ity but it may be widely deviated from with due conside-
and precaution provided only that a concentrated solution
»14 E. DIVERS AND T. HAGA:
of nitrite be employed. Potassium nitrite, 30 grains; potass
hydroxide, 10 grams; water, 50 to 100 grams are to receir
current of sulphur dioxide freely until crystals begin to fo
the containing flask being all the time agitated in a coo
bath of ice and brine. The sulphur dioxide is now to be ent
more slowly for some time longer and then stopped. After
ting the flask stand for half an hour the solution should be
of the desired salt which is then drained dry on the tile.
mother-liquor is alkaline to litmus but not to rosolic acid (
sence of sulphite, absence of alkali); the well-drained salt it
is only faintly alkaline to litmus, if at all so. The double
is also produced when to an ice-cold nearly saturated solu
of potassium nitrite a similar solution of potassium pyrosulp
is very slowly added until crystallisation begins after which
solution is allowed to stand for some time. Thus prepared,
compound salt is liable to be contaminated with a little nit
sulphate and sulphite. The experiment just described was n
first by Raschig but he attached to it a significance unlike -
here presented. Discussion of his views will be found tow
the end of this paper.
There is yet another way in which this potassium nit!
hydroximidosulphate can be produced which it is of interes
mention because it illustrates the decomposibility of potass
5/6 normal hydroximidosulphate into the normal and 2/3 nor
salts. While the 2/3 normal salt dissolved in 16 per cent.
richer solution of the nitrite crystallises out only in combina
with nitrite, the 5/6 normal salt can be dissolved in a nit
solution of even 50 per cent. and yet for the most part crys
lise out again uncombined. But generally with this strengtl
nitrite solution a little fluffy. or cotton-like lustreless matter:
POTASSIUM NITRITO-HYDROXIMIDOSULPHATES. 215
ates. If now to this fluffy matter suspended in its cold
r-liquor carefully decanted from every particle of the crys-
f the 5/6 normal salt a hot solution of this 5/6 normal salt
or even 40 per cent. nitrite be poured in, a relatively large
ity of the fluffy matter is obtained and not the hard
s of the 5/6 normal salt. Under the microscope the fluffy
r proves to be crystalline and when drained on the tile it
its a silvery lustre while on analysis it proves to be the
)-2/3 normal hydroximidosulphate only slightly impure from
‘esence of a little 5/6 normal hydroximidosulphate and nitrite.
in place of the potassium 33.10 and sulphur 18.06 per cent.
und in it 33.79 and 18.35 respectively, together with an
nity equal to 1.09 per cent. potassium. Dissolved up in
2 per cent. nitrite solution it recrystallises as the pure
e salt. It is thus apparent that in a very concentrated
on of nitrite containing the 5/6 normal salt dissolved there is
ble equilibrium between the tendency to yield HON (SO, K),,
(SO;K),,OH, again and that to form HON (SO,K),,
O.
odium nitrite forms a compound with sodium 2/3 normal
ximidosulphate which has not been further examined prin-
y because of its high solubility in sodium-nitrite solution.
otassium mitrite and normal hydroximidosulphate KNO,,
\(SO,K),, 40H:.—This compound salt is only obtainable
a strongly alkaline solution. For when the normal hydrox-
sulphate is dissolved in a hot concentrated solution of the
> only the 5/6 normal hydroximidosulphate crystallises out
oling just as it would do in the absence of nitrite. In
to crystallise out either the normal hydroximidosulphate
Journal, 7, 30) or its combination with nitrite free alkali
216 E. DIVERS AND T. HAGA:
must be present in some quantity in the solution. The press
of too much alkali causes a little of it to separate with
normal salt, taking the place apparently of the water of crvs
lisation of this salt (this Journal 7, 52), and similarly
separate with the normal salt in its combination with nit
then also seeming to lessen the capacity of the normal sal
take up nitrite. The double salt is readily obtained by
solving normal hydroximidosulphate nearly to saturation i
hot (70°) solution consisting of 33-66 parts nitrite and 3-5 p
hydroxide to 100 parts water and cooling. Usually it fo
lustrous silky fibres like those of the 2/3 normal double salt
radiating from points to form voluminous soft spherical mas
When the solution is more strongly alkaline the double
separates as nearly opaque spherical granules with someti
long fibres growing out from them. Under the microscope tl
granules are seen to have also a radiating fibrous texture an
represent the soft voluminous spheres highly condensed. Proba
these always begin their growth from a minute granular nucl
The double salt can only be purified for analysis by pressin
on the porous tile, when the soft spheres become a felted lustı
cake and the hard white granules crumble down like masse
wax. Analysis of the two forms bas given us the follow
results :—
Potssm. Alk. potssm. Sulphur
Silky ; found, 30.21 9.92 16.23 per cent.
K,N;S,0,0, 44 OH., 35.14 10.04 16.43 ,,
Granular ; found, 33.99 9.20 15.90 ,
K,N;S,0,., 60H, 33.89 9.68 15.85 ,,
The varying amount of water is only the recurrence of what
POTASSIUM NITRITO-HYDROXIMIDOSULPHATES. 217
recorded concerning the normal potassium hydroximido-
ate by itself. The double salt is exceedingly alkaline, its
inity we estimated by means of decinormal acid and litmus.
Like the previously described double salt it is but little
le in concentrated nitrite solution and freely soluble in water
1 decomposes it into its constituent salts and also decom-
one of these, the normal hydroximidosulphate, into alkali
rystals of the 5/6 normal salt. When heated it decomposes
nly but gently and without fusing or scattering, and evolves
‚ red fumes only. It was by this behaviour quite distin-
able from the 2/3 normal double salt and also from any
hydroximidosulphate which, simple or combined with
e, contained less than its K, to S. By dissolving the
0-2/3 normal hydroximidosulphate in a hot concentrated
on of nitrite containing sufficient alkali the nitrito-normal
»ximidosulphate can be readily obtained by cooling the
on.
Potassium nitrite and potassium 5/6 normal hydroximidosul-
—We have obtained three compounds of the 5/6 normal
with nitrite, one being 7KNO,, 2HK,(NS8,O,),, 30H, By
an almost saturated solution of potassium nitrite con-
ig a little potassium hydroxide and dissolving in it by heat
/6 normal hydroximidosulphate there is obtained a compound
inute fibrous crystals very lustrous when dry and decomposed
ater but recrystallisable from a saturated nitrite solution.
same compound salt can be obtained also by dissolving the
o-normal hydroximidosulphate in hot almost saturated solu-
of nitrite.
Heated it proves to be mildly explosive. Its composition
jaches that indicated by the formula given above. For
218 E. DIVERS AND T. HAGA:
analysis it was only air-dried on the tile; in the desiccator
would probably have lost its 3 per cent. of water (=30H,) a
then approached in composition Fremy’s sulphazite.
Potssm. Sulphur Alk. Potssm.
Original salt, found, 36.81 14.35 4.80 per cent.
Recrystallised, ,, 36.68 14.47 451 ,,
K,;H;N 1804, 30H,, 36.87 14.20 4,34 „
A second double salt, 3KNO,, K,H (NS,O,),, OH,, was got
dissolving one mol. 5/6 normal salt and 1.4 mol. potassium |
droxide in a hot 65 per cent. nitrite solution and cooling.
appearance it resembled the other compound salt. Its anal)
gave :—
Potssm. Sulphur Alk. potssm.
Found, 36.17 15.07 4.51 per cent.
Calec., 36.81 15.06 4.61 <i
A third double salt, anhydrous, 7K NO,, 3K,H(NS,O,),, \
not prepared synthetically but by treating an almost satura
solution of the nitrite with alkali and sulphur dioxide, and :
ding alkali again after the sulphonation, imitating a process
Fremy’s. Then, filtering the heated solution from much cr
talline 5/6 normal hydroximidosulphate mixed with a little of
combination with nitrite, we got the mother-liquor, when qu
cold, almost filled with tiny prisms of a compound answering
the above formula :—
Potssm. Sulphur Alk. potssm.
Found, 36.94 16.37 4.96 per cent.
Cale, 36.99 16.51 5.05
93
This salt was quickly resolved by water into nitrite a
POTASSIUM NITRITO-HYDROXIMIDOSULPHATES. 219
als of the very sparingly soluble 5/6 normal hydroximido-
ate.
[he varying proportions in which potassium nitrite and the
ormal hydroximidosulphate unite would possess but little
st were it not for the fact that they have evidently been
ılly met with and taken to be salts of specific constitution
remy and by Raschig.
Non-existence of Dihydroxylaminesulphonates.
‘remy believed in the existence of less sulphonated deriva-
of potassium nitrite than his sulphazite (see next paper) it-
ss sulphonated than his sulphazotates (hydroximidosul phates)
ttributed his failure to find them to the fact of their possess-
xceedingly high solubility. Claus held much the same
and believed that by adding to an aqueous solution of
ium nitrite an alcoholic solution of sulphur dioxide in not
wge a quantity he had obtained an impure crystallisation
salt, ON SO,K (Ber. 1871, 4, 508): he did not prove this
the case, but what he did publish about his product is
ent to show us that he had got the compound of potassium
> with 2/3 normal hydroximidosulphate we have described
is paper. A repetition of his experiment gave us this double
gether with much ethyl nitrite. Raschig regarded Claus’s
ration as essentially the same as one of his own salts to
1 he gave the constitution of basic dihydroxylamine sul-
ate derivatives with the following formule :—
OK HO. SO.K
HON/ wad NNONS
CNSOK “" (S0,K)/ NOK
220 E. DIVERS AND T. HAGA:
These he prepared by partial sulphonation of the nitrite
known ways. They both yielded erystals of a hydroximidosı
phate when dissolved in a little water and differed in no esse
tial particular from nitrito-hydroximidosulpates. From I
solutions of nitrite and a hydroximidosulphate we obtained |
cooling an apparently homogeneous crop of crystals of alm
the same composition and properties as one or other of Raschi;
salts. Raschig gave two ways for preparing the salt having t
second of the formule just given and in these ways we ha
obtained the nitrito-2/3 normal hydroximidosulphate already <
scribed in this paper, but mixed with a little potassium sulphi
This impurity accounts for the alkaline reaction of Raschi
preparation and the presence in it of a little more than K, to
He got the other salt (K, to S) only once and in the fo
of white crusts when working unsuccessfully for hydroximidos
phate in Claus’s way, the other main product being imidosulpha
that is hydrolysed nitrilosulphate as he himself pointed o
We have obtained—also by sulphonating nitrite, following Fret
—a product qualitatively like Raschig’s salt though quantitative
a little different from it, and at the same time like the seco
salt compounded of nitrite and 5/6 normal hydroximidosulpha
described by us on page 218. The percentages found by Rascl
were potassium, 36.84, and sulphur, 15.50.
When Raschig’s salt was dissolved in water and acidified
gave nitrous oxide as the only gaseous product while ours ga
also some nitric oxide. This fact might have served to rent
incorrect the application of our formula to his salt but for t
evidence there is that this was mixed with a little sulphite whi
would have reduced any nitric oxide. Its mother-liquor
further evaporation gave, we are told, so much sulphite alo
POTASSIUM NITRITO-HYDROXIMIDOSULPHATES. 221
the next crop of the salt itself as to cause its rejection.
presence of sulphite in less quantity in the first crop of
als will have been masked by the oxidising action of the
oxide in becoming nitrous oxide. That sulphite was pre-
in Raschig’s preparation well accords also with the fact that
sium hydroxide added in excess precipitated potassium sul-
, for, although hydroximidosulphate is itself decomposed by
most concentrated solutions of potassium hydroxide into
ite and nitrite, this decomposition is slow and the sulphite
deposits after some time. Raschig’s preparation when dried
tile was only a powder, that is, presumably, was not ob-
ly crystalline, a point which also indicates an impure salt.
> the potassium and sulphur are in the same ratio in the
salts, quantitative analysis would hardly have made its
nce known.* Inspection of Raschig’s formule is of itself
ient to prevent their getting accepted as in accordance with
acts. For from these formule both salts should be strongly
ine, while in reality one is neutral. Above all it is hardly
ble that dissolution in cold water should suffice to cause
sulphonated nitrogen to become disulphonated.
Raschig held his two salts to be identical with Fremy’s
sium sulphazile and sulphazate respectively; but the nature
remy’s salts will be found, we believe, more precisely given
ıe paper following this. The point we would here insist
| is that Raschig’s preparations, judged by their chemical
viour, have no claim to be considered as dihydroxylamine
ratives, being in every way indistinguishable from synthe-
Of the 3KNO, of our formula (p. 218) only one mol. can give nitric oxide and only
extent of two-thirds of its nitrogen; the other third becoming nitric acid. Raschig’s
is indicates the presence of only 3/4 mol. active nitrite. The quantity of hydrated
te required to be present is therefore only 5.2 per cent. of the mixed salts.
222 E. DIVERS AND T. HAGA:
tically prepared compounds of nitrite and hydroximidosulphat
Dihydroxylamine salts have as yet only a hypothetical existe:
and are likely to remain so. For the double linking of 1
oxygen atom with the tervalent or quinquevalent nitrogen at
seems always experimentally to make or break itself in a sin,
act, notwithstanding its bipartite character.
Raschig in his researches on Fremy’s sulphazotised salts g
besides those we have just discussed, two other salts of undet
mined constitution, both of which were most probably a
nitrito-hydroximidosulphates. They may therefore be notic
here although Raschig did not represent them to be dihydrox
amine derivatives. Yet they were evidently closely like |
other two in properties. One was isomeric with potassium hy)
nitrososulphate (Pelouze’s salt) and also with his (K, to 8) ‘ dil
droxylamine’ salt, allowing for different hydration, and the otl
was isomeric with potassium 5/6 normal hydroximidosulphs
Each could be obtained but once and they only call for a
detailed notice because of the theoretical importance given
them as isomerides of other salts. The first referred to ab
was mistaken by Raschig for Pelouze’s salt (hyponitrosos
phate) but that salt it certainly was not (this Journal, 9, 8
It was got by dissolving nitric oxide in solution of potassi
sulphite and hydroxide and evaporating to a small volume |
crusts formed. If we assume that air or nitric peroxide was ı
excluded there were the conditions present for getting a nitri
hydroximidosulphate, for, as we show in a paper which w
shortly follow this, nitrous fumes passed into potassium sulph
POTASSIUM NITRITO-HYDROXIMIDOSULPHATES. 223
yn generate hydroximidosulphate freely together with
he other salt isomeric with 5/6 normal hydroximidosulphate
obtained in Raschig’s attempt to form 2/3 normal salt by
g surphur dioxide into a solution of potassium nitrite and
xide and letting stand for a day. These, too, are conditions
sting nitrito-hydroximidosulphate. Now, both products
in being decomposed by water in such a way as to yield
ximidosulphate and in other ways behaved as compounds
rite with one of these salts. The behaviour of the one
ic with hyponitrososulphate was indeed exceptional in
‘hen dissolved in water containing a little alkali it gave
3 normal hydroximidosulphate when according to our cal-
n it should have given the 5/6 normal salt, while it also
n hot alkaline solution a little nitrous oxide which only
<yamidosulphate is known to give. These peculiarities
y attribute to partial hydrolysis having occurred in the
instable salt before these experiments were made.
he calculated formula for the isomeride of hyponitrosul-
as a nitrite compound is 3K NO,, K,H (NS;O,),, 20H,, and
, compound we have described on page 218; that for the
ide of the 5/6 normal hydroximidosulphate treated as being
te compound is 3KNO,, 6K,HNS.O,, 5K;H(NS8,O,)., which
er should give crystals of K,HNS,O,, 20H,. This com-
salt we have failed to get but its occurrence can be
accepted as possible. Its assumed existence affords a much
satisfactory explanation of the nature of this salt of Ras-
than that we were able to offer in our paper on hydrox-
ulphates already referred to.
224
Nitroso isomer, found,
Calculated,
D, & H’s salt, found,
Oximido isomer, ,,
Calculated,
For the present, the existence of isomesides of Pelouze’s
and Fremy’s basic sulphazotate must be regarded as no longer «
probable,
Polssm. Sulphur
35.72 14.40 per cent.
36.05 1475 ,
36.17 1505 ,
33.04 21923 „
32,91 21.54 ,
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dentification and Constitution of Fremy’s
Sulphazotised Salts of Potassium, his
Sulphazate, Sulphazite, etc.
By
Edward Divers, M. D., D. Sc., F. R. S., Emeritus Prof.,
and
Tamemasa Haga, D. Sc, F. C. &.,
Professor, Tokyo Imperial University.
sufficiently concentrated solution of potassium nitrite and
‘ide submitted to the action of sulphur dioxide gave
minute silky needles of a salt which he provisionally named
um sulphazate. With slightly diminished concentration of
ution he generally obtained instead the brilliant, often
rhombic prisms of potassium basic sulphazotate (5/6-normal
imidosulphate, this journal, 7, 15). But sometimes
was obtained neither of these salts before the solution
> transformed into a starch-like jelly through the form-
f a salt which he named potassium metasulphazate, or
came filled with spangles of yet another salt called by him
um metasulphazotate. When the solution was a little too
to give any of these and when too much alkali had not
dded, there usually appeared peculiarly pointed crystals of
‚ he named potassiwm neutral sulphazotate (2/3-normal hydr-
osulphate Raschig) and, lastly, with still greater dilution
226 DIVERS & HAGA : IDENTIFICATION AND CONSTITUTION OF
the minute brilliant needles of his potassium: sulphammoi
(nitrilosulphate Berglund). Still other salts he believed te
produced in the first stages of the reaction between the nit
and sulphur dioxide, one of which he named potassium su
azite; but this he did not obtain directly, finding a reason
this in the exceeding solubility of this early formed salt.
prepared it—but only in quite small quantity and as crystal
warty granules—by the action of water upon the ‘sulphaz
whereby this was converted into ‘ basic sulphazotate’ which
posited and a solution that on evaporation yielded the ‘su
azite. ‘These two salts could together in solution be char
back into the ‘ metasulphazotate’ while the ‘sulphazite’
the ‘ sulphazate ’ could similarly often be changed into the ‘m
sulphazate’ again. These two ‘ meta’ salts he regarded there
as perhaps merely double salts of the others. The ‘ sulphaz
the ‘sulphazate,’ and the ‘sulphazotates’ he treated as b
members of a series of salts in which there were to two atom
nitrogen from one up to eight atoms of sulphur,—three in
“sulphazite, four in the ‘ sulphazate’, and five in the ‘su
azotates.’ With this conception of the nature of these salts, b
on his analyses, it was easy to understand the decompositioı
the ‘sulphazate’ into the ‘sulphazite’ and the ‘ sulphazot:
But this and other of Fremy’s interpretations of the facts
served by him have lost all importance and particular inte
through the progress of chemistry since his memoir was }
lished and only his account of the facts requires considera
now.
Subsequent work by others and ourselves in the same f
has shown that Fremy in the account he gave of the prepara
of his many salts went two little into details as to the condit
FREMY’S SULPHAZOTISED SALTS OF POTASSIUM. 227
which they were obtained,—apparently because he was
ble to be more precise. When Claus attempted to get
y’3 salts he obtained only masses of minute crystals of salts
ose individuality and nature he could make out little be-
of the impossibility of dissolving them all up undecomposed.
; experiments the ‘sulphammonate’ (nitrilosulphate) was
3 formed in considerable quantity either as a first or second-
roduct and by its presence prevented any satisfactory inves-
n of the other salts. In Fremy’s working, this most easily
d salt came only as the final product of the sulphonation
herefore gave him no trouble. Claus emphatically displayed
epticism as to Fremy’s results; yet in nearly every point
ich he differed from Fremy asto the facts we find Fremy
ye been right. When Raschig repeated Fremy’s work—
ith the modifications in procedure introduced by Claus—he
sults similar to, though less unsatisfactory than, those Claus
btained. He made an approach to Fremy’s work in so far as
1e often got very little nitrilosulphate ; nevertheless he too
in his attempts to prepare the ‘ sulphazate ’ in Fremy’s way.
n perhaps all essential points we can lay down the method
jeat Fremy’s experimental work successfully. But in some
a, little uncertainty obtains owing to the fact that the very con-
ted and complex solutions which yield Fremy’s salts are apt
osit what is virtually the same salt in different forms as well
‘umes salts quite distinct from each other under only slight and
re variations in the circumstances attending their formation.
julphazate.—This is Fremy’s first salt directly obtained in
ılphonation of the nitrite. In getting it he took approxi-
y 5 mols. potassium nitrite to 2 mols. potassium hydroxide
, little water and into the solution passed sulphur dioxide
228 DIVERS & HAGA : IDENTIFICATION AND CONSTITUTION OF
until it became almost filled with silky needles very solubl
water. So far it is easy to follow Fremy with a full measuı
success if only the water used is limited to perhaps twice
weight of the nitrite and that the heating effect of the nitri
counteracted by cooling. Claus and after him Raschig failed
then inexplicably to us they did not start with Fremy’s pro
tions of nitrite to hydroxide, though even with the proport
they took, success was possible with care. The salt thus for
by Fremy was not tested and analysed by him until after it
been changed (but without his having recognised the fact
the further treatment to which he submitted it. Before
change it is potassium nitrito-2/3 normal hydroximidosulp
described in the preceding paper, a neutral salt decomposed
water into its constituent salts. Fremy’s finished ‘ sulphaz
was strongly alkaline and very caustic and when decompose
water gave nitrite and the 5/6 normal hydroximidosulpha
not the 2/3-normal salt. Also the analysis he gave of it
nished numbers such as the original product could not have g
him. Instead of potassium, 33.10, sulphur, 18.06, and nitro
7.9 per cent., he got potassium, 34.90, sulphur, 19.55,
nitrogen, 4.9. We can learn what his after-treatment wa:
reference to other parts of his paper where he speaks of the
necessary (when sulphonating the nitrite) to maintain the &
linity of the solution by adding potassium hydroxide from tin
time and of dissolving sulphazotised salts for examination in v
containing this alkali. Certain it is he must have added :
potassium hydroxide to the solution after getting it to crysta
as a precaution to preserve the salt. Now the effect of this
dition is to change the composition of the product without n
affecting its silky asbestus-like appearance. The change in «
FREMY’S SULPHAZOTISED SALTS OF POTASSIUM. 229
ion is to deprive it of much of its nitrite and to convert the
10rmal into more nearly normal hydroximidosulphate—to
ce, therefore, potassium nitrite by potassium hydroxide.
pting Fremy’s mean numbers as accurate, what he analysed
the composition,
11K,NS,O,,0H.; K,HNS,O-, 20H;; 2(K,HNS.O,, KNO,).
Potssm. Sulphur Nitrogen Alk. potssm.
Found, 34.9 19.55 4.9 —— per cent.
Calc, 34.9 19.51 4.9 9.36 „
But his analyses have no claim to receive such close treat-
, his nitrogen seemingly being always much too low; and it
fficient to say of his ‘sulphazate’ that it was the silky
tus-like nitrito-2/3 normal hydroximidosulphate more or less
erted into the also silky asbestus-like normal hydroximido-
ate, an acconnt of it with which Fremy’s description of its
properties entirely agrees. With dilute acids it gave slowly
us oxide unmixed with nitric oxide. Fremy specially points
hat no sulphazic acid or any other sulphazates could be ob-
d from the potassium salt. There is, therefore, nothing to
y belief in this compound being the salt of a particular
e acid, the sulphazic.
Sulphazite.—W hat Fremy named potassium sulphazite he only
obtained, and then not by direct sulphonation of the nitrite,
je form of white mammillated crystalline crusts from a solu-
thickened by the other salts contained in it. That is, to
his sulphazate when dissolved in a little water containing
potassium hydroxide deposited crystals of basic sulphazotate
normal hydroximidosul phate), and left a mother-liquor which
old evaporation till syrupy yielded the sulphazite. It showed
, analogy with his sulphazate but was distinguished from it
~~
ea
230 DIVERS & HAGA : IDENTIFICATION AND CONSTITUTION OF
by having little tendency to hydrolyse and by at once evolving
some nitric oxide when its solution was mixed with a dilute acid.
Water decomposed the su/phazite, but into what products was not
ascertained.
We have sufficiently realised Fremy’s expectations that his
sulphazite might directly result from sulphonating the nitrite with
subsequent addition of alkali. The substance obtained in this
way did not differ greatly in composition from his:
Potassm. Sulphur
Fremy’s salt, 38.16 16.27 per cent.
D. & H’s salt, 36.94 16.37 ss,
and agreed with it in chemical properties, so far as is known.
At the same time it was indistinguishable from a compound of
nitrite with 5/6 normal hydroximidosulphate, and has been de-
scribed by us as such in the preceding paper (p. 218) in which
it stands as the third of these double salts and in which its
preparation is given. Other experiments of various kinds have
yielded us such ‘mammillated crusts’ as Fremy got, which,
though only in rough agreement in percentage composition with
his sulphazite, behaved like it and proved to be impure double
salts of nitrite with 5/6 normal or more nearly normal hydrox-
imidosulphate. We are therefore convinced that his sulphaz-
ite was only such a double salt.
Metasulphazate.*—In Fremy’s experience it sometimes hap-
pened, when passing sulphur dioxide into solution of nitrite and
alkali of a concentration intermediate to that giving sulphazate
and that giving basic sulphazotate, the solution set to a starch-like
jelly instead of crystallising. He obtained a similar jelly by cooling
* Often, misprinted mefasu/phazotate in the French original, but not in the German
translation.
FREMY’S SULPHAZOTISED SALTS OF POTASSIUM. 231
centrated solution of sulphazate and sulphazite; also by
- a solution of sulphazate and then cooling it. When
ly compressed the jelly became a transparent wax-like mass.
| in this waxy state to 50°-60° it suddenly changed into a
n of sulphazite and minute crystals of basic sulphazotate.
other respects it proved to be intermediate in properties
hazate and sulphazite. No other metasulphazates could be
ed from it, so that Fremy was disposed to regard it as
a double salt of sulphazate and sulphazite. Its constitution
herefore have been that of nitrite combined with normal
normal hydroximidosulphate in such proportions and with
dditions perhaps of alkali as prevented crystallisation.
Je have not had Fremy’s success in getting this salt in
of jelly and wax but have met with just such phenomena
forming barium sodium hydroximidosulphate, BaNaNS,O,,
as will be found described in our paper already frequently
d to. We have however obtained a salt, or homogeneous
re of salts, of the same composition as the metasulphazate,
ith the form of the silky radiating fibrous crystals of the
-normal hydroximidosulphate, from which it differed only
wing deficiency of nitrite, that is, it was equivalent in com-
n to a mixture of the normal salt and its nitrite com-
, both of which crystallise with the same habit. We give
Fremy’s numbers, our own, and those calculated for the
sion, 3(KNO,, 2K,NS,O,, 40H,); K,NS,0;; 30H.
Potsem. Sulphur Nitrogen Alk. potssm.
und (Fremy), 35.10 16.74 4.81 per ceut.
„(D.&H.), 35.10 16.68 — 10.47 ,„
culated, 35.06 16.74 5.23 10.23 ,,
Ve got the salt by dissolving the hydroximidosulphate in
232 DIVERS & HAGA: IDENTIFICATION AND CONSTITUTION OF
hot concentrated nitrite solution containing alkali. To 100 cc
water there were present 45'/, grm. nitrite and 1?/, grm. potassiun
hydroxide; for 66 mol. nitrite there were dissolved 10 mol
anhydrous normal hydroximidosulphate. But for the salt bein
in beautiful asbestus-like fibres, there was nothing to distinguisl
it from the jelly and the wax-like metasulphazate, which, therefore
we do not hesitate to class as a nitrito-hydroximidosulphate.
Basic sulphazotate, which Fremy considers next, has bee:
shown by us already (loc. cit.) to be the 5/6 normal hydroximi
dosulphate, and not the salt of a distinct acid, the sulphazotu
It is liable to contain a small excess of potassium when crys
tallised from a strongly alkaline solution. A solution of th
normal salt readily deposits it, as does also that of the nitrit
compound of the normal salt.
Neutral sulphazotate was shown by Raschig to be the 2/
normal hydroximidosulphate. The potassium sulphazotates wer
distinguished by Fremy from the salts previously described b:
him by their ability to form other sulphazotates by double de
composition. Fremy’s analytical results in the case of the neu
tral sulphazotates are hopelessly out of accord with its constitutio:
and properties, though those for the basic sulphazotate are satis
factory enough.
Sulphazidate, produced by the hydrolysis of the sulphazotate, 1
hydroxyamidosulphate (Claus). Sulphazilate and metasulphazilate
oxidation products of sulphazotate are ON(SO;K), and ON(SO,K),
and have been studied by Claus, Raschig, and Hantzsch.
Metasulphazotate.—Sometimes Fremy got a salt in the forn
of spangles (paillettes), in appearance like minute crystals of das.
sulphazotate, but differing from these in not being hard under pres
sure. This salt he named, therefore, metasulphazotate. Accordin;
FREMY’S SULPHAZOTISED SALTS OF POTASSIUM. 233
1 it is also obtainable by mixing (hot) solutions of the (basic)
zolale and sulphazite. It is very soluble in water, very
ne and unstable unless the water contains alkali. In pure
it becomes basic sulphazotate and sulphazite again. It
the greatest analogy with metasulphazate and is Jistin-
d in the same way as this salt from basic sulphazotate. It
je a compound of basic sulphazolate and sulphazite. So far
7. It will be evident that there is nothing in its history
yperties to distinguish it, except its occurring in the form
rkling particles and even that can be met with in the
sulphazotate suddenly precipitated ; we have also got other
e sulphazotised salts in what may be called spangles,
ı not this particular salt. In the preceding paper, page 214,
ve described an impure form of nitrito-2/3 normal hydrox-
ulphate obtained by dissolving the 5/6 normal salt in a hot
trated solution of nitrite, but still not so very concentrated
give the nitrito-5/6 normal double salt. This preparation
reless while in its mother-liquor, but when dried on the
is a fine silvery lustre. It has when dried in the desic-
exactly the composition of Fremy’s metasulphazotale and is
less alkaline than the metasulphazotate and is much less
1e than the mefasulphazate. It may be formulated as
K,NS,O,; 9(K NO,,K,HNS,O,,1'/,0H,).
Potesm. Sulphur Nitrogen Alk. potssm.
und (Fremy), 33.8 18.6 3.9 per cent.
, (D.&H), 33.79 1835: 2 1.09 ,
Iculated, 33.68 18.37 7.63 112,
ulphammonate and sulphamidale are respectively nitrilo-
te and imidosulphate (Berglund).
— SS a ee
Digitized by Google
On a Specimen of a Gigantic Hydroid,
Branchiocerianthus imperator (ALLMAN),
found in the Sagami Sea.
By
M. Miyajima, Rigakushi.
Science College, Imperial University, Tokyo.
“With Plates XIV & XV.
n the morning of January 1, 1899, quite a commotion was
ed in the Marine Biological Station at Misaki by the
ig in of a very beautiful and gigantic Coelenterate (Pl. XIV).
been caught, on the previous day, by a fishing “ long-
from a depth of about 250 fathoms near Okinose, a
rine bank 18 kilometers south of Misaki. It was an object
was calculated to raise enthusiasm in a naturalist. A large
ırmounted a long stalk which evidently fixed the animal
} sea-bottom. A circle of numerous graceful tentacles hang
from the margin of the disc, while on its upper surface
an oral tube, surrounded at its base by bushy dendritic
lages and having a second circle of slender tentacles around
per edge. The total height of the animal was 700 milli-
236 M. MIYAJIMA :
meters and the prevailing colour transparent scarlet. It
agreed on all sides that it was a New Year’s gift from Orton
and that it should be known in Japanese as Otohime
Hanagasa.
The specimen, when brought in, was entirely fresh but
not living. It was placed in 2% formalin to preserve, if
sible, something of its beautiful colour. At first the atte
seemed successful, but after a while the colour began to
gradually, until now the specimen is completely bleached to
white. For histological examination, pieces of the tentacles
the dendritic appendages were fixed in the sublimate anc
Perenyi’s fluid.
The specimen was handed over by Prof. Mirsuxvrr to
to work out its finer structure.
It was evident from the first that the specimen was
similar to the form only a short time before described by M
(98) as Branchiocerianthus urceolus. I started, therefore,
an idea that I was dealing with an Actinian.
As I proceeded in my investigation, however, it became j
that this idea was not tenable, and the conclusion was fi
reached that the animal was very closely allied to Corymor
and that it belongs probably to the species obtained by the “ (
lenger ’’ at about the same locality and named by ALLMAN
Monocaulus imperator, notwithstanding many discrepancies bet
his description and the specimen. This conclusion was comn
cated through Prof. Mirsuxurt to Dr. Mark and a request was
sent to him, that during his opportune stay in Europe, he sh
*Otohime” is a beautiful goddess who is supposed to have her palaces at the bot
the sea. ‘ Hanagasn” is the flower-sun-shade or ornamental parasol. Thus Otoh
Hanagasıı means “the ornamental parasol of Otohime.”
BRANCHIOCERIANTHUS IMPERATOR (ALLMAN). 237
sible, examine the original specimen of Monocaulus imperator
> British Museum. To the results of his examination of the
1ens I shall return in the later part of this paper.
Ieanwhile an article was published in the Zoologischer An-
by O. CARLGREN (‘99) throwing doubt on Marx’s Branchio-
thus being an Actinian, and contending that it more
bly is a Corymorpha or at least a form standing very close
rymorpha. a
n June, 1899, a correction was published by Mark (’99)
if in the Zoologischer Anzeiger. His previous preliminary
ption had been based on external anatomy, and he now.
y admitted that further researches had convinced him of the
hat the animal in question must be more nearly related to
ydroidea than to the Actinia, though its exact affinities he
ot yet determined. In a postcript he mentions our conclu-
which had been communicated to him, as mentioned, by
and thinks that both his and our specimens belong to the
senus and that our specimen is probably identical with the
aulus imperator of ALLMAN.
efore going further I wish to express my deepest feeling
ligation to Prof. Mirsuxuri for the supervision and advice
he has given during the progress of my work.
Description.
‘his hydroid is a solitary form consisting of a well marked
nth and a hydrocaulus. Its most striking feature is a
ly expressed bilateral symmetry. The hydranth is disc-
d and bears two sets of tentacles and a circle of dendritic
omes, all showing in their arrangement a well marked
238 M. MIYAJIMA:
bilaterality. The hydrocaulus, which is attached not t
center but to the edge of the hydranth, is nearly cylindrics
increases in diameter from the attachment of the hyd
towards the end which is fixed in the sandy sea-bottom.
total height of the animal attains 700 mm., as measured
the top of the oral tube to the attached base of the hydroc
In the fresh condition the hydranth was rose pink a
tentacles, both oral and marginal, were deep scarlet in c
while the gonosomes possessed light rufous colour. The E
caulus was light pink in colour, being quite pale in its m
The general features and the colours are well sho
Fig. 1, Pl. XIV, which was drawn from the preserved spe
by Mr. Nagasawa, artist of our Institute, making use a
the rough sketches I made at the time of the fresh object
Hydranth.
The upper surface of the hydranth is flattened so t
may be described as an “oral disc.” The lower surface,
ever, assumes a shallow funnel-shape, which passes down
into the hydrocaulus. This disc has an oval outline, but
from that of Branchiocerianthus urceolus, in having tts &
diameter less than ws transverse, the two diameters’ being
pectively 80 and 90 mm. (Woodcut 2).
At one end of the sagittal diameter is attached the h
caulus where the circle of the marginal tentacles is also
rupted, The plane of the disc is oblique to the long axis
hydrocaulus (Woodcut 1, I), though not to the same deg
in Branchwocerranthus urceolus MARK.
The. edge where the hydrocaulus is attached I shall des
the lower, and the opposite the higher, edge.
BRANCHIOCERIANTHUS IMPERATOR (ALLMAN). 239
W oodeut 1.
I.
is showing sagittal (1.) and transverse (II.) sections of the hydranth.
ypostomal region of the disc ; €. orifice of the diaphragm (m) in the hydranth ; © orifice
ıphragm (m’) in the hydrocaulus; cg. central, /g. lateral, globule of the gonophore ;
, HP’ lower cavity of the hydranth; A. hypostome; ic. intercalated cord; HC.
lus; mi. marginal, of. oral, tentacle; P. peduncle of the gonosome; R. outer
the disc, provided with the radial canals (r.c.).
240 M. MIYAJIMA:
The hypostome (Woodcut 1, I, II, A), the superior pro
tion of the disc, is slightly conical, diminishing gradua
its diameter from the base towards the free end whe
mouth opens. A little below the mouth the hypostome h
brush-like group of about 180 filiform tentacles (o¢.) whic
arranged in three or more closely packed verticils, the
tentacles being much larger than the inner. The out
ones attain a length of 50-55 mm., while the innermost
small and crowded that I could neither measure them we
count their exact number. Below the oral tentacles the
stome is slightly constricted, but there is no indication of sy]
glyph which is said to be present in the oral tube of Bra
cerianthus urceolus. The side of the hypostome turned t
the lower edge of the disc passes gradually to the disc,
on the opposite side it seems abruptly raised from the
so as to make an angle between. The hypostome is thus 0
to the disc proper which again is not perpendicular to th
of the hydrocaulus. Hence we can show the relation of the
parts, the hypostome, the disc and the hydrocaulus, dia
atically with three lines, of which two vertical ones, corre:
ing to the axes of the hypostome and of the hydrocaulus,
with an oblique one representing the axis of the disc, fo
obtuse angles between them (Woodcut 1, I).
The base of the hypostome (Woodcut 2, B.) occupies abc
middle of the disc, but on the side turned towards the lower
its base gradually becoming lower and lower, may be s
stretch as far as the margin of the disc, while laterally and t«
the higher edge it is distant from the margin 35 mm. and 2:
respectively. It thus assumes an ovoidal outline, the pointe
attaining the lower margin of the disc and passing directly
BRANCHIOCERIANTHUS IMPERATOR (ALLMAN). 241
W oodeut 2.
|| |! | | \\
c'e ... Wee TR NN
ty |
1)
sidi à 70777
| |
A
EL TU
MN.
»-
— P =
® | 2
+ e —
“+ e ==
a =
= =
= R
=. IN
mt.
HC
Diagram showing the upper surface of the disc.
hiatus at the lower edge of the disc. Other letters as in Woodcut 1.
of the hydrocaulus. The longer (1. e. up—down) diameter
ovoidal space measures 60 mm., while the transverse at
est middle portion is only 25 mm. This space is des-
f the radial canals which are prominently seen in the
ng part of the disc.
ound the margin of this hypostomal region there arises
he surface of the disc a row of dendritic gonosomes
ch in shape strongly remind one of the heads of cauliflowers.
242 M. MIYAJIMA :
They number in all 96 and are arranged approximately
single row, which, being interrupted at the lower edge of the:
assumes the form of a horse-shoe. (Woodcut 2, P). At the
ends of the horse-shoe are situated the smallest gonosomes w
stand at a distance of 15 mm. across from each other.
length of the stalk of the gonosomes varies from 20 mm. to 60
While the gonosomes nearer the lower edge of the disc ar
the whole shorter than those nearer the upper edge, it is t
noticed that the larger and smaller gonosomes are placed a
nately, indicating faintly the two circles in their arrangen
the larger gonosomes being placed in the outer, and the sm
in the inner, circle.
The region of the disc outside the gonosomes is marked
numerous radial canals (Woodcut 2, #.) which run from
base of the gonosomes to the margin of the disc. This re
thus assumes the form of a wide horse-shoe, whose two :
gradually diminish in their breadth towards the lower edg
the disc until they terminate at that edge. Hence this re
varies in breadth, measuring 20 mm. on the median line at
higher edge, and 35 mm. on the lateral region, while on the |:
side both arms are practically zero.
The radial canals (Pl. XV, Fig. 1, r.c.) slightly swell ou
surface of the disc thus giving the latter an undulating apy
ance. The canals are intercalated by solid cords (Pl. XV, Fi
i.c.) which appear on the surface of the disc as opaque lines.
canals and the intercalated cords are longest in the lateral re
where they run obliquely across the disc, and are longer
the breadth of this region. The canals situated nearer the |
edge are smaller and shorter than those higher up, until at
both arm-ends they are practically nid. On the other hanc
BRANCHIOCERIANTHUS IMPERATOR (ALLMAN). 243
ls on the higher side of the disc are not so long as those
he lateral, but run straight from the base of the gonosomes
1e outer margin of the disc, the length of the canal being
the same as the breadth of this region (Woodcut 2).
The radial canals and the intercalated cords increase in their
h towards the outer margin of the disc where the both struc-
are broadest. Inwards, the radial canals open into that part
ie hydranth-cavity where the cavities of the gonosomes stand
ımmunication with the latter. Outwards, the canals terminate
lly on the margin of the disc. The intercalated cords
ge suddenly near the margin of the disc and acquire a
y which forms a part of that of the marginal tentacle (PI.
Fig. 1).
The name of marginal tentacles (mt) is given to the outer-
circle of filiform tentacles arranged like a fringe around the
rin of the disc. The circle is not complete, there being a
8 (v) at the lower edge of the disc where the surface of the
ystome passes directly into that of the hydrocaulus. The
test tentacles arising from the 6th or 7th intercalated cord,
ting from the lower edge, occupy the two ends of this in-
plete ring. Whether there were any smaller tentacles nearer
lower edge, I am not sure. There is no indication, so far
can see, of any having existed. Towards the higher edge
1e disc they increase successively in length until about the
(counting from the lower edge) is reached, of which the
th on both sides is about 200 mm. After this there seems
e no special arrangement of the tentacles, which vary from
mm. to 300 mm. in length. They numbered 198 in all.
tentacles are flattened at their base, and are compressed so
ly with one another that the basal portion appears to form
244 M. MIYAJIMA:
a part of the disc. Just above the flattened base the tentac
assumes the form of a tube, 4 mm. in diameter, and tape
gradually towards its free end.
The hydranth (Woodcut 1, I. II.) contains a wide cavi
which is separated by a thin membrane (m) into two parts, :
upper (H) and a lower (7). The superior prolongation of t
upper cavity is that of the hypostome, which does not sh:
any indication of the septal partition. The lower cavity is mc
spacious than the upper, and not only occupies the whole low
part of the hydranth but extends also through the entire leng
of the hydrocaulus.
The membrane (m) separating the hydranth-cavity aris
diaphragm-like just below the upper wall of the disc. In abc
the center of this diaphragm, directly below the mouth, is
ovoid orifice (Woodcut 1, II, C) which puts the upper and low
cavities in communication with each other. The orifice
11 mm. and 15 mm. respectively in its transverse and sagit
axes.
That part of this diaphragm which corresponds to the p:
of the upper surface of the disc marked B in Woodcut 2, «.e.
the basal part of the hypostome, projects into the cavity
the hydranth like a shelf, with the aforesaid opening near
middle and with no attachment either above or below. Outsi
this portion, however, the diaphragm forms the floor of t
radial canals mentioned above, so that it is suspended, so to spe:
by the numerous intercalated cords (vide supra) to the up}
surface of the disc. At the margin the diaphragm is united
the outer wall of the hydranth (Woodcut 1).
To show the somewhat complicated relations existing betwe
the marginal tentacles, radial canals, intercalated cords, etc.
BRANCHIOCERIANTHUS IMPERATOR (ALLMAN). 245
f sections (Pl. XV Figs. 4-9) passing through the lines 1-1,
3, 4-4, 5-5, 6-6 in Fig. 1, Pl. XV is introduced. The
ction (Fig. 4) through the outermost margin of the disc,
corresponds to the line 1-1 in Fig. 1, shows that the bases
marginal tentacles (i.6.) and the blind ends of the radial
(r.c.) are arranged alternately, the former projecting out
and below more than the latter. The upper projection
onds to the enlarged end of the solid cord. The cavity
tentacle is almost filled up by the spongy endoderm which
he whole cavity of the animal, so that it remains as a
canal only in the upper and lower swollen parts of the ten-
On the other hand the radial canals contain a wide cavity
is clearly separated from that of the tentacle-base by the
eveloped mesoderm. In the next section (Fig. 5, through
e 2-2, Fig. 1) cut just inside the margin of the disc, the
canals already assume their characteristic shape in cross-
, while the intercalated cords have already lost their cavity
y. Bounded by the mesoderm the intercalated cord as-
in cross-section the form of a trapezoid. It is convenient to
uish here three kinds of the mesoderm-lamellæ, the upper,
and the vertical. The upper lamella (w./.) is situated along
face of the disc, the basal (2.2.), in the floor (z.e. in the dia-
n), and the vertical (v./.) connects these two lamelle.
traced inwards, (Pl. XV Figs. 6, 7) the intercalated cord
es thinner and thinner, until it no longer shows in cross-
ı the form of a trapezoid, but assumes the shape of a tri-
formed by two vertical and one basal lamella. Where the
yme arises (Fig. 8), the vertical lamella does not reach the
lamella; hence the radial canals communicate here with one
»r and form the upper common cavity of the hydranth.
246 M. MIYAJIMA :
Within the circle of the gonosomes (Fig. 9) the upper la
stands entirely separated from the basal, on which the ve
lamella shows itself only as a ridge-like line which in «
section is recognizable as a simple small knob.
Preserved in formalin, the fine tissues of the animal
unfortunately mostly gone. Luckily, however, the pieces ¢
gonosomes and the marginal tentacles, which were presery
sublimate, etc., helped us to ascertain something of the histol
character of the animal.
The wall of the animal-body, I need hardly say, consi
the three layers, ecto-, endo- and mesoderm as in other Cc
terates.
The ectoderm, the outermost layer, has been entirely
off from the specimen in formalin, but in the pieces fixed
sublimate was well preserved. This tissue is a single lay
cylindrical cells which in their preserved condition are mo
less vacuolised. There are present a few nematocysts whic
characteristic of the ectoderm of Coelenterata.
The mesoderm is a very firm, supporting layer whi
placed between the ecto- and endo-derm or two portions ¢
endoderm. This tissue was well enough preserved eve
formalin so that the structure of the animal could be I:
made out by this layer alone. =
The endoderm, the innermost layer, which pe
cavity of the animal, remained unfortunately only | 1e
in the specimen in formalin. From these patches it ton
made out that the endoderm lining the pane: ity iss
cells thick (Figs. 3 & 10). The cells are i la: arly a
contain but a little cytoplasm which is N ards
with the nucleus. Consequently the wall of the ie
x”
an ll
et
Digitized by Go ogle
BRANCHIOCERIANTHUS IMPERATOR (ALLMAN). 247
gy appearance. I can not think that this appearance of the
derm is caused by bad preservation, for the tentacles fixed
sublimate show also the same structure. In the preserved
, the endoderm forming the upper ceiling of the lower cavity
ie hydranth has a thickness of 3 mm.
In the cross and longitudinal sections (Figs. 11 & 12) of the
‘inal tentacle, the whole of the space inside the mesoderm
tirely filled up with a tissue which reminds one of the verte-
» notochord. It has the same structure as the spongy en-
rm of the hydranth-cavity already mentioned. Only near
Jase of the tentacle, this spongy tissue leaves in the center
all cavity which is separated by the mesoderm from the
anth-cavity. Hence the cavity of the tentacle-base is of a
ed extent, extending not farther towards the distal end, and
nunicating nowhere with the general cavity of the hydranth.
ngitudinal section (Fig. 2) through the margin of the disc
s plainly the relations of the disc and the base of the tentacle
mesoderm being drawn darker than other parts in the figures).
The gonosomes (Fig. 1, p.) as already mentioned consist
he branched tubular stalks, upon which the gonophores
yrouped in a crowded cluster. Each stalk branches dicho-
usly into about the 10th or 12th order. Each branchlet
inates in a group of small globules, of which we recognize
inds (Figs. 13 & 14). The one kind of which there is only
in each cluster is situated on the top of the terminal branch,
2 the others take a more lateral position. The former is
r than the latter, consisting of the irregularly shaped cells
ly vacuolised (Fig. 14, c.g.). In this kind of globule the
derm of the branch is no longer recognisable and the ecto-
endo-derm can not here be clearly distinguished. It seems,
SS a
m en — = a > = ee
248 M. MIYAJIMA :
however, reasonable to suppose that the centrally placed sm
cells which are continuous with the endoderm of the bran
belong to that layer. The cells which presumably belong t«
ectoderm and form the main part of this globule seem t
mostly distended. In this globule the nematocysts (Fig. 15
are found in a large number ; hence the central globule ma
regarded as the battery.
The lateral globules (Fig. 14, /.g.) are mostly spherical
consist of compactly packed cells rich in cytoplasm. Then
derm prolonged from the branchlet distinctly separates
ectoderm from the central cell-mass. After examining n
sections I was able to find a few globules which enable u
see that the clusters are true gonophores. In such glol
(Fig. 16), one is able to see that the ectoderm cells at the
are grouped into a mass forming the “ bell-nucleus” w
pushes the endoderm in as a cup. This part of the endo
is arranged into a regular layer one cell deep and is easily
tinguishable from the remaining part. Owing to the se
(Fig. 16) having been cut slightly obliquely, the cavity in
endoderm seems irregular and very limited. In reality, !
is a wide cavity occupying the whole interior of the glo
which communicates with that of the branchlet. I could
detect gonophores developed any further than this in ours]
men. January is probably not the season in which the ri
ing of the sexual products takes place.
The terminal branch thus bears two sorts of globules,
one being a nematocyst-battery and the other a true sexual o1
Hence the dendritic gonosome of this animal is a peculiar o
which bears on a common stalk the sexual and defensive elem
BRANCHIOCERIANTHUS IMPERATOR (ALLMAN). 249
In other hydroids these two elements are borne on separate stalks,
as for example in Pennalia. |
Hydrocaulus.
The under part of the hydranth is prolonged to a shallow
funnel whose neck corresponds to the hydrocaulus. At about
the point where the hydranth joins the hydrocaulus, there is
a circular constriction (Woodcuts 1 & 2). Here the diameter of
the hydrocaulus is only 9 mm. and from this part down to the
base it increases in its diameter. Within the constriction is a
diaphragm (Woodcut 1, I, II, m’) separating incompletely the
cavity of the hydranth from that of the hydrocaulus. In other
words the circular constriction is the surface expression of the
insertion of the diaphragm. In the midst of this partition there
is an opening (Woodcut 1, I, II, C’) which puts the two cavities
above and below in communication. It is about 4 mm. in dia-
meter and is almost circular. The plane of the diaphragm is not
visibly oblique to the long axis of the hydrocaulus. In the speci-
mens of Monocaulus imperator in the British Museum, this dia-
phragm is, according to Dr. Mark, distinctly oblique and the
central opening is much elongated.
The hydrocaulus is a hollow tube which has a total length
of 650 mm. including the proximal end with hair-like appen-
dages. The hydrocaulus, even when fresh, was collapsed and
more or less longitudinally folded, so that the exact measure-
ment of its diameter was almost impossible. Approximately, it
was 15 mm. just below the constriction, 25 mm. at the middle,
and 42 mm. at the terminal root.
The outer surface of the hydrocaulus is smooth. In the upper
250 M. MIYAJIMA:
half of it there are visible from outside 15-20 longitudinal
bands (Fig. 17). They stand about 2-3 mm. distant fror
another and run down to about the middle part of the |
caulus where they become obscure. From the surface the
remarkably like the mesenterial filaments ofan Anthozoon.
wavy bands anastomose here and there with one anothe
sive to the hydrocaulus of our specimen an appearance
resembling that of Corymorpha. ‘Though the bands are in tk
served state still visible, they were more conspicuous when
These longitudinal bands show themselves in cross-section (E
as dense spots (x) in the mesoderm, which have a great affin
any staining agents. From the bad state of preservation
specimen, in which the ectodermal and endodermal cells
mostly lost, I could not ascertain whether the wavy band:
the endoderm canals, a structure peculiar to Corymorpha, (
[ think it, however, very probable that they existed, anc
rise to these band-like appearances. In the published ac
of Monocaulus imperator of ALLMAN the endoderm canal:
plainly described and figured.
The mesoderm is very well developed, especially
hydrocaulus where it reaches a thickness of about 0.2-0.'
This remarkable layer shows itself in the form of a fibr
membrane, which, when macerated with caustic potash, is sep
into two layers, the outer longitudinal (Fig. 19, 72.) a
inner circular (Fig. 19, c.l.). The former is thicker and
less with any coloring matter than the latter.
In our specimen there is no sudden bulb-like expans
the lower end of the stalk, such as is described by Ma
Branchiocerianthus urceolus or by ALLMAN in Monocaulus imp
The lowest and broadest part of the hydrocaulus is enclos
BRANCHIOCERIANTHUS IMPERATOR (ALLMAN). 251
about 30 mm. from the base in a chitinous sheath which gives
an anchorage to the Hydrozoon. With the exception of the upper
edge the sheath (Fig. 20 s) bears in most parts very numerous
hair-like processes (ap) of brown color, which are so entangled that
many foreign bodies (e.g. Echinus spines, sand grains, dead
shells) are wrapped up within them. The sheath and the root
proper are united so closely that they are not to be separated from
each other without tearing. In contrast to the pink-colored
hydrocaulus the brown color of the sheath with its appendages is
very conspicuous.
At the lowest end of the hydrocaulus the wall is very deli-
cate and has an opening, the margin of which is destitute of the
hair-like appendages.
Above the root the mesoderm possesses here and there
irregular small depressions which are recognized by tolerable
magnification from the surface as clear spots. These depres-
sions are also present in the wall of the root enclosed in the
sheath.
A portion of the root cut longitudinally (Fig. 21) shows
that the sheath with its appendages is separate from the root
proper, but has an organic connection with it. The hair-
like appendage (ap.), which is scen to be a slender hollow
process of the sheath, embraces in its interior the thread-like
outgrowth (o.) of the wall, which perforates the mesoderm and
is connected directly with the endoderm of the inner cavity of
the hydrocaulus. Hence it seems to me that the above-
mentional small depressions in the mesoderm are certainly the in-
dications of the wart-like processes of the wall of the hydrocau-
lus as in Corymorpha.
252 M. MIYAJIMA :
Summary.
1. The most striking feature in our specimen is itss
expressed bilateral symmetry as shown by the excentric
ment of the hydranth to the hydrocaulus and by an ir
tion of the circles of the gonosomes, radial canals, and n
tentacles at the lower edge of the disc. Those who ha
the above account will, I think, agree with me in think
this bilateral symmetry is due, not to the primitive stat
body-organization, but rather to its elaboration and specia
We must therefore regard this remarkable case of |
symmetry in a hydriform person as very different from |
pressed for instance in the planoblast of Corymorpha and D
which is but temporary and occurs only at a certain p
development, or from the biradial symmetry as express
few genera like Monobrachium and Lar by a reduction
number of tentacles.
2. The hydranth-cavity is divided into two parts, o
the upper is in its outer part again divided into mam
canals visible even on the surface of the disc. That ren
structure is not, however, peculiar to our specimeı
example, the hydranth-cavity of Tubularia is divided s
into two parts by a peculiar ring-shaped formation” obse
several authors. In Tubularia the lower cavity is narroy
the upper, so that the former forms a slender canal in the
of the “ Wulst.’”? Gosta Grönberg (‘98) described in the h
of Tubularia indivisa slender endoderm-canals which are t
in number to the proximal (marginal) tentacles and
between every two tentacle-bases, running obliquely from t
*O. Hamann (’82) described that formation as “aboral Wulst,’? G. Grönb
“ Mesoderm-wulst.”
BRANCHIOCERIAMTHUS IMPEBATOR (ALLMAN). 253
wity outwards and downwards. ‘These canals, though not
from the surface, may be regarded as corresponding to the
canals in our specimen, since they both arise from the
cavity of the hydranth and are arranged alternately with
ginal tentacles.
The tentacles are filiform and arranged in two sets, oral
and marginal (proximal), as is characteristic of the
es of Tubularide, Oorymorphide, and Monocaulide. The
of the tentacle is mostly obliterated, being filled up with
lar tissue—a condition very frequently met with in the
e of the Hydrozoa.
The dendritic appendage is a true gonosome which bears
summit the sexual elements. Our specimen seems to be
ıre, hence it could not be decided whether the gonophore
lanoblast or a sporosac.
The hydrocaulus is marked with many wavy bands visible
he surface, and possesses a thin sheath with filamentous
lages at its lowest end.
Considerations on the Systematic Position of our
Specimen.
hose who would compare the account given above of the
ire of our specimen with that of Branchiocerianthus urceolus,
(’98) will not for a moment doubt that we have in these
ses essentially similarly constituted animals. It seems almost
luous to call attention to the points of likeness: the hydro-
with the wavy bands in its upper half and with the sheath
amentous appendages at its base, the hydranth surmounting
drocaulus, with its radial canals, dendritic gonosomes, and
ts of tentacles, all of which show a strongly expressed
254 M. MIYAJIMA :
bilateral symmetry, being interrupted at the lower (or
posterior) edge where the hydrocaulus is attached.
That our specimen and Prachiocerianthus urceolus b
least to the same genus, there can hardly be any doubt. \
they belong to the same species is another question. It
haps premature to decide this point, at present, in as 1
Marx has not yet published his full paper. Judging f
preliminary notice which deals exclusively with the :
features, the following are the chief points of difference.
a. The general shape. B. urceolus is stated to have an ex
graceful, symmetrical vase-like figure with flaring li
lateral margins of the hydranth-disc were in the
state “ folded in symmetrically from either side, so a
to touch at a point, a little below the middle of t
This bending in of the margins of the disk produc
upper end of the animal a sort of eccentric funnel
* © The fancied resembl
depression, * .
the animal to a little
which this side view pres
suggested the specific
; A adopted—urceolus.’ (M
b p. 148). The pitcher
shape of the hydranth
due to two causes: (1) t
ing in of the disc-mar;
a a
(2) the extreme obliquit
axis of the hydranth to
of the hydrocaulus. In
A. B. nexed woodcut, a repres
axis of the hydrocaulu
BRANCHIOCERIAMTHUS IMPERATOR (ALLMAN). 255
that of the hydranth. Thus these two axes make in B. ur-
ceolus an extremely obtuse angle as in A, and thus help to
produce the vase-shape. In our specimen the angle which
these two axes make with each other (B) is much less obtuse,
and moreover the folding in of the disc margin has not been
noticed from the first, either in the fresh or preserved state.
The disc lay flat and open as a disc, and never suggested the
idea of a pitcher.
b. The shape of the disc is oval in B. urceolus ; in our specimen
it is more nearly circular. Moreover in the former, the
sagittal or longitudinal diameter is greater than the trans-
verse, while in our specimen it is the ¢ransverse diameter
which is the greater of the two. The following measure-
ments will make this point clear.
Sagittal (longit.) diam. Trans. diam. Ratio of trans.
1
n mm. in mm. diam. to sagittal.
B. urceolus.
Small specimen...235..................... Less anses 60%
Large specimen...38...................…. Weisses nearly 79%
ne Colles ieee LR Vi 112.5%
c. The size :——
Length of the hydro- Maximum length Maximum length
alas in in of the marginal of the oral tentacle
ternacle in mm. in mm.
B. urceolus....... 105-200... 20er 020000 0. LA 30-35
en a cesse 650 <—..........".0000e 300 HELLE LETLNIIEIET 50-55
d. The lower end of the hydrocaulus :—Marx describes a bulb at
the lower end of the hydrocaulus. In our specimen, there
is no such sudden enlargement as deserves the name of a
bulb, although that end is, as has been stated, the largest.
256 M. MIYAJIMA:
e. The radial canal:—Marx mentions that the radial ca
B. urceolus run “ from the base of the oral tube to th
of the marginal tentacles, before reaching which many
fork, each of the branches communicating with the lun
single tentacle’? (Mark ’98, p. 150). The case is v
ferent in our specimen in which the radial canals
fork at all and do not communicate with the lumen
marginal tentacles. The latter, on the contrary,
continuations of the intercalated cords.
Whether these differences are to be regarded a
specific or due simply to the differences in size, age, ı
must leave for the present an open question. I am nh
however, to think that 2. urceolus and our specimen
different species. |
References have already been made several times in the
of the foregoing pages to the resemblance of our spec!
Monocaulus imperator of ALLMAN, a gigantic hydroid «
by the Challenger off Yokohama (stat. 327). The des
given by ALLMAN of this animal in his report of the Hy
of that Expedition (’88) is not as exhaustive us is
able. He makes no mention of any bilateral symmetry
animal, but we must remember that the specimens which
before him were extremely badly preserved, as he is ca:
mention, and that the figure of the animal which was made
spot by the artist of the Expendition must necessarily ha
made hurriedly, and as we can testify from our own observ:
the fresh object, it is very easy to overlook such a feature as I
symmetry when the disc is lying in the midst of a.
tentacles. Of course the best thing we could do under |
BRANCHIOCERIANTHUS IMPERATOR (ALLMAN). 257
cumstances was to appeal to the original specimens. At the
request of Prof. Mırsukuxı, Prof. Marx, who was opportunely
staying in Europe at the time, was kind enough to examine the
type specimens of Afonocaulus imperator, kept in the British
Museum. The results of his observation were not entirely conclu-
sive, as the specimens “ have so long been in strong alcohol that it
was quite impossible to make out anything very satisfactorily.”
He naturally made special efforts to ascertain the condition of
the hydranth—whether it was radially or bilaterally symmetrical.
In one specimen, he felt tolerably confident, though by no
means sure, that there was an interruption narrower than in
Branchiocerianthus urceolus in the marginal tentacles. In another
specimen the central opening in the diaphragm which divides the
cavity of the hydranth from that of the hydrocaulus was found
much elongated—a point which in his opinion pointed to bila-
teral symmetry.* He also thought that there is much less ob-
liquity of the hypostomal region to the axis of the Hydroid than
in B. urceolus “for the wall of the hydranth between the con-
striction and the base of the tentacles can be seen to be nearly
the same height all around, or at least not markedly different on
opposite sides.’ This last point is against the view that our
specimen is identical with Monocaulus imperator, for although
the disc is much less oblique in our specimen than in B. ur-
ceolus, as shown above, the hydrocaulus is attached at one end
of the sagittal (longitudinal) diameter of the disc. But Prof.
Marx adds, “the specimens were so much wrinkled and folded
that I have not much confidence in this conclusion.” There is
* In our specimen the opening which puts the hydranth-cavity and that of the hydrocaulus
in communication is not elongated, but almost circular as already stated.
258 M. MIYAJIMA :
one curious point of difference between our specimen and 2
caulus imperator. While the hydranth in the Challenger s
men is much smaller than that of our specimen, the sta
enormously longer, being said to reach the almost incre
length of 7 feet 4 inches. This is, however, stated to be :
stretched, and is not the normal length.
While it is not thus possible to establish absolutely
identity of our specimen with Monocaulus imperator of Aıı
there are on the whole strong probabilities in favor of
assumption. Those who read carefully ALLMAN’s descriptior
notice that the points which he brings out distinctly ir
structure of his species, such as a wide cavity extending thr
the entire length of the stalk, the presence of the stalk-meso
in the shape of a fibrillated membrane—a point which Au
emphasizes as “the most striking feature in the histology c
Hydroid ’’—and so forth, are absolutely similar in our spec:
If we remember in addition that both came from practical]
same locality, it is, I believe, within the scope of reasor
ness to conclude that our specimen belongs to Monocaulus impe
of Allmann.
If this is really the case, we must examine other spec
Monocaulus. The genus includes, besides Monocaulus impe:
two other species; M. glacialis, (Sars) (for which ALLMAN
blished the genus) and M. pendula, (Acassız). These two |
show, however, a radial symmetry, and now that M. imp
is shown to have a bilateral symmetry, can not possibly b
in the same genus with the latter. M. imperator must the
be separated from the other two species and placed in a
genus. According to the rules of nomenclature, this new ;
BRANCHIOCERIANTHUS IMPERATOR (ALLMAN). 259
must take the name Branchiocerianthus* first given by Mark,
and our specimen then ought to be known as
Branchiocerianthus imperator (ALLMAN).
P.S. We shall await with interest the full report on the
specimen of Monocaulus obtained by Prof. Cuun in his recent
deep-sea expedition (’99).
December, 1899. Zool. Institute, Science College,
Imperial University, Tokyo.
eee —
* MARK (’99) mentions that Rhizonema carnea of 8, F. CLARKE (’76) may be a form same
as, or closely related to, Branchiocertanthus. CLARKE’s original description is very brief and
it is impossible to determine whcther MARK's suspicion is correct or not. At any rate,
CLARKE makes no mention of any bilateral symmetry in the structure of the animal.
260 M. MIYAJIMA:
Literature.
Acassiz, L.
‘62. Contributions to the Natural History of the United Stat
America. Vol. IV, (Boston).
ALLMAN, G. J.
‘64, On the construction and limitation of genera among the
droidea. Ann. and Mag. of N. H., 3rd series, No. T1.
‘71, A monograph of the gymnoblastic or Tubularian Hyd
(London. )
‘83-88. Report on the Hydroids dredged by H. M. 8. Challenger d
the years 1873-76.
BRAUER, A.
‘91. Ueber die Entstehung der Geschlechtsprodukte und Entwick
von Tubularia mesembryanthemum. Zeit für wiss,
. Bd, 52,
CARLGREN, O.
‘99, Branchiocerianthus urceolus E. L. Mark, eine Hydroid ?
Aug. Bd. XXII. No. 581.
Caux, C.
‘91. Coelenteraten, in Bronn’s Klass. und Ord. des Tierreic]
‘99. Die deutsche Tiefsee-Expedition 1898/1899. (Berlin).
CLARKE, $. F.
‘76. Report on the Hydroids collected on the coast of Alaska ar
Aleutian Islands, by W. H. Dall, U.S. Coast-Survey and ]
from 1871 to 1874,
Dorteın, F.
‘96. Die Eibildung bei Tubularia ; Z. für Wiss, Zool. Vol. 62
GRÖNBERG, G.
‘98. Beiträge zur Kenntniss der Gattung Tubularia. Zool. 4
Bd, 11. Heft 1,
BRANCHIOCERIANTHUS IMPERATOR (ALLMAN). 261
wn, ©.
82. Der Organismus der Hydroid-Polypen. Jen. Zeit. Bd. 8.
Vol. 8. (Jena).
s, TH.
68. British Hydroid Zoophytes. 1868.
oc, G.
73. Vorläufige Mittheilung über Cölenteraten. ‘Jen. Zeit. Bd. 7.
‚EL. |
98. Preliminary report on Branchiocerianthus urceolus. Bull. of
the Mus. of Comp. Zool. Vol. XXXII. No. 8,
99. “ Branchiocertanthus,” a correction. Zool. Anz. Bd. XXII
No. 590.
[ANN, À.
83. Die Entstehung der Sexualzellen bei den Hydromedusen, 1883.
Explanation of Plate.
Pl. XIV.
he hydranth with the upper half and the lowest part of the hydro-
Nat. size.
PI. XV.
eference letters: B. hypostomal region of the disc ; Hct. ectoderm ; End.
rm; H upper, H’ lower, cavity of the hydranth ; ic. intercalated cord ;
nesoderm ; mi. marginal tentacle; p. peduncle of the gonosome ; 2.
region of the disc, provided with the radial canals; rc. radial canals ;
men of the base of the marginal tentacle.
ig. 1. Surface-view of a portion of the disc with gonosomes (p) and
ial tentacles (mt.). Nat. size. |
upper wall of the disc, 5 a part ofthe diaphragm in the a
ig. 2. Longitudinal section through the outermost margin of the disc
à x 4.
262
Fig. 3. Cross-section of the upper wall of the lower cavity ı
hydranth. Zeiss DDx2.
Fig. 4-9. Serial sections of the upper part of the disc. Zeiss
Fig. 4. Cross-section through the line 1-1 in Fig. 1,
Fig. 5. . i 5 2-2 fs
Fig, 6. ss | „33 j
Fig. 7. — ,, 5 » 44
Fig. 8. 5 js ss 5-5 -
Fig. 9. is ‘3 5 6-6 |
Fig. 10. Cross-section of the radial canal and intercalated cord,
BB x 2.
Fig. 11. Cross-section of the marginal tentacle. Zeiss a, x 2,
Fig. 12. Longitudinal section of the same. Zeiss a, x 2.
Fig. 13. Terminal branches of a gonosome, Zeiss a x 2.
Fig. 14. Longitudinal section of a branchlet of the gonosome.
Fx2. og. Central, !.y. lateral, globule.
Fig. 15. Central globule with nematocyst (n). Zeiss DD x 4,
Fig. 16. Lateral globule in which the bell-nucleus (b.n.) and the
derm-cup (enc.) are fairly well recognizable. Zeiss Fx 2.
Fig. A part of the wall of the hydrocaulus with the wavy
Nat. size.
Fig. 18. Cross-section of the mesoderm in the hydrocaulus, Zeiss
hd. outer, longitudinally, c./. inner, circularly, striated layer. a. spe
responding to the wavy band.
| Fig. 19. Mesoderm of the hydrocaulus, macerated with caustic :
Zeiss a x 2.
ow?
Fig. 20. Surface-view of the root and the sheath. Nat. size, ap
| like appendage ; s. sheath.
| Fig. 21. Longitudinal section of the root figured in Fig 20,
8x2. 0. outgrowth of the endoderm ; other letters as in Fir. 20,
an
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Jour. Sci.Coll. Vol. Xi. PI. XV.
b.n.
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Tutual Relations between Torsion and Magneti-
zation in Iron and Nickel Wires.
By
H. Nagaoka, Rigakuhakushi,
Professor of Applied Mathematics,
and
K. Honda, Rigakushi,
Post-graduate in Physics.
With Plate X VI.
The various effects of stress on the magnetization of ferro-
netic metals are of such a complex character that no simple
ition seems to exist among them. ‘The strains caused by mag-
izing the ferromagnetics are of no less complex a nature, so that
co-ordination of these two classes of complicated phenomena
up to the present, still a matter of doubt. Various isolated
s, such as the analogies between the change of magnetization
longitudinal pull and that of length by magnetization, the
tion between the twist caused by the interaction of longi-
inal and circular magnetizations and the circular or longi-
inal magnetization produced by twisting a longitudinally or
ularly magnetized wire respectively, were long considered as
rding a clue to the explanation of these phenomena. So far
=
264 H. NAGAOKA AND K. HONDA:
as we are aware, no attempt has yet been made to plac
these different phenomena on a common footing. Some ti
we hinted at the probable connections which exist
the twist caused by passing an electric current through
tudinally magnetized wire and the change of volume
length in ferromagnetic metals produced by magnetizatio:
said relation can also be extended to the explanation —
phenomena ; namely, the transient current produced by ts
magnetized wire and the longitudinal magnetization ca
twisting a circularly magnetized wire. It is our objec
present paper to show that these different phenomena
linked together in a common bond.
$1. Twist produced by the interaction of circula
and longitudinal magnetizations.
The subject was first studied by G. Wiedemann” w
blished remarkable reciprocal relations with the long
magnetization produced by twisting a circularly magneti:
Dr. Knott" found that the direction of twist in iron is
to that in nickel; Bidwell” afterwards discovered that tl
in iron is reversed in high fields and takes place in tl
direction as in nickel. Unfortunately some of the exp
were undertaken with wires which were longer than tha
coil, so that the magnetization was far from being unif
will suffice for qualitative tests, but we can not hope
1) Nagavka and Honda, this Journ. 18, p. 57, 1900; Phil. mag. 49, p. 341,
2)G. Wiedemann, Pogs. Ann. 103, p. 571, 1858; 106, p. 161, 1859; Elektr
3) Knott, Trans. Roy. Soc. Edinb., 32 (1), p. 193, 1882/83; 85 (2), p. 877, 18
p. 485, 1891,
Digitized by Google
TORSION AND MAGNETIZATION. 265
ite quantitative results. The position of maximum twist
ickel shows a large difference in the present from the corres-
ing experiment by Dr. Knott.
The twist produced by longitudinal magnetization of a cir-
ly magnetized wire was measured in the following way. To
xtremities of an iron or nickel wire 21 cm. long were
brazed stout brass wires, and a
ae light plane mirror was attached
— to the lower one. The end of
iN | the lower brass wire was dipped
in a mercury pool, while the
upper brass wire was clamped to
a small tripod, which rested on the
top of a magnetizing coil pro-
vided with hole, slot, and plane
arrangement. One end of the
accumulator was connected with
the tripod, while the other was
led to a mercury pool. The
hung vertically in the axial line of the coil, which was
m. long and gave a field of 37.97 C.G.S. units at the
e by passing a current of one ampere. The vertical com-
nt of the terrestrial magnetic field was compensated by plac-
mother coil in the interior of the magnetizing coil. The
part of the wire to be tested was protected against air
mt by enclosing it in a wide brass tube with a small win-
just where the reflecting mirror was attached. The twist
measured by scale and telescope method, by which the
tion of 0.3" per. cm. was easily read. The current was
ured by Kelvin graded amperemeters, whose constants were
266 H. NAGAOKA AND K. HONDA:
from time to time checked by means of an ampere bala
The experiment was conducted in the following manner :—
1. The circularly magnetizing current was kept const
and the amount of twist measured by varying the longitudin:
magnetizing current.
2. The longitudinally magnetizing current was kept const
and the amount of twist measured by varying the circul
magnetizing current.
Before each experiment, care was taken to demagnetize
wire completely either longitudinally or circularly by passin;
alternate current of gradually diminishing intensity.
Twist by varying the longitudinal field (Fig. 1).—
direction of twist in iron, so long as the longitudinal magneti
field is not strong, is such that if the current is passed down
wire from the fixed to the free end and the wire is magnetized :
north pole upwards, the free end, as seen from above, twists in
direction of the hands of a watch. By keeping the circular |
constant, the amount of twist increases at first, till it reaches a m
mum in a field of about 20 units ; it then goes on diminishing ti
ultimately changes the direction and continues to twist in the o|
site direction with increasing field. The field at which the twi:
reversed increases with the circularly magnetizing field. In nic
the direction of twist is opposite to that in iron, but the gen
feature is similar to iron, the only difference being that «
in strong longitudinal fields, the twist is not reversed. For v
of the equal thickness, the amount of twist in nickel is gre
than that in iron—the maximum twist in iron wire of 1 :
diam. by passing 6 amperes through it amounts to about
per em., while with nickel wire of 0.83 mm. diam. under sin
conditions, the maximum twist amounts to about 200.”
TORSION AND MAGNETIZATION. 267
Twist by varying the circular field (Fig. 2).—Here we
ce a slight dissimilarity between iron and nickel. In
, the twist increases with the strength of the circular field,
ne longitudinal field remains constant. Such is also the
with nickel in moderate and strong fields. In low longi-
nal fields, however, the twist does not continue to increase with
circular, but we notice a maximum as will be clear in the
e. There is great experimental difficulty in increasing the
ilar field, inasmuch as the heating of the wire becomes very
t and thus materially deteriorates the result.
The hysteresis accompanying the cyclical change of the cir-
r magnetization deserves special notice (see Fig. 3). If the
itudinal field be such that with the increase of the circularly
netizing force, the twist reaches a maximum, the curve of twist
below the former course on weakening the circular magneti-
n. The twist, however, goes on slowly increasing,. till it
ses the on-curve and then reaches a maximum, whence it
ually diminishes and ultimately vanishes in negative field.
course after passing this point is exactly the reverse of that
idy described. . The character of twist is exactly the same
iron as for nickel, when we take the opposite character of
t into account. The nature of the hysteresis is nearly the
> when the longitudinal magnetizing field is made to vary,
e the circular field remains constant.
The results thus far obtained are in accordance with the
riments of Wiedemann and Knott; we have only to notice
discrepancy as regards the position of maximum twist in
el. In Dr. Knott’s experiment, the said point occurs in
‘ably high field, while in the present experiment, it occurs
‘ly in the same field as in iron. It may partly be due to the
268 IT. NAGAOKA AND K. HONDA:
difference in the method of measuring the twist and partl
the non-uniformity of the field, as was often the case in mo
the older experiments.
The observed angles of twist in iron and nickel are exhil
in the following tables, where C denotes the longitudinal cu:
per sq. mm. in amperes, H the field strength in C.G.S. ı
and + the angle of twist per cm. expressed in seconds.
Circular Field being Constant.
Iron wire: diam.=0.98 mm.
= 1.06
— 102.1
33.6 — 85.1 53.4 — 134.1 53.2 — 159.:
84.0 — 62.9 83.8 — 101.8 84.0 — 119.
102,2 — 54.5 103.4 — 101:
225.0 — 31.7 226.0 — 59%
19,6
TORSION AND MAGNETIZATION. 269
Longitudinal Field being Constant.
Iron Wire: diam.=1.05 mm.
$2. Circular magnetization produced by twisting a
longitudinally magnetized wire.
By twisting a longitudinally magnetized ferromagnetic wire,
circular magnetization is developed. If, therefore, two ends
of the wire are connected by a conducting wire, a transient
current due to the circular magnetization appears in the cir-
cuit at the moment when the twist is applied. Some years ago,
270 H. NAGAOKA AND K. HONDA:
one” of us investigated the transient current for iron
nickel wires. It was tben found that the current due to tw
ing was opposite in direction in tbese two metals and tha
reached a maximum in moderate fields. As the magueti
current was not very strong, no conclusive measurements :
made as regards the nature of the transient current in stı
fields. In order to make this point clear and see if any
mate relation with the Wiedemann effect could be traced, fresh
periments were undertaken by the same method as before.
have to notice that the ferromagnetic wire was so placed in
axial line of the magnetizing coil that it lay in nearly uniform f
Some of the measurements of the transient current for
and nickel wires are given in the following table and in Fi:
[ron wire: diam.= 1.33 mm. Nickel wire: diam.= 1.09 m
length =20.90 cm.
0=15° | 0 = 50° 0 —15° Ü = 30°
H Qx10 | H 0x1 H Qx10 | H Qx
3.2 25.4 32 30.7 13 — El “ha qe
5.3 29.0 48 33.6 37 = M | LT =
15.6 24.9 15.8 42.0 16.0 —179 |). de
32.6 16.3 30.5 384 337 —212 102 —2
545 109 48.6 30.5 534 —21.5 25.6 —2
87.4 6.4 86.0 18.5 81.2 —20.1 449 —4
120.8 2.5 | 139.4 8.1 | 1154 -192 67.6 —4
165.6 1.5 157.0 —16.4 | 1074 —4
242.0 — 0.1 213.7 —12.6 160,2 —¢
495.4 — 1.7 319.6 —10.3 241.5 —£
650.2 — 2.9 530.3 — 6.5 316.4 —2
908.0 — 3.3 703.0 — 5.4 699.8 —1]
1298.0 — 3.1 12140 — 3.9 | 1150.0 —]
1790.0 — 2.5
1815.0 — 2.0 | 1853.0 —
1) Nagaoka, Journ. Sci. Coll., Tokyo, 4, p. 323, 1891.
TORSION AND MAGNETIZATION. 271
Here 6 denotes the angle of torsion and Q the time-integral
of the transient current expressed in C.G.S. units. The resistance
of the whole circuit was 4.5 ohms. The nickel wire here used was
made of the same specimen as the nickel prism used in our former
experiments.
As is well known, the direction of the transient current,
and therefore that of the circular magnetization, is opposite in
iron and nickel. The current for constant amount of twist in-
creases with the strength of the longitudinal field ; it, however,
soon reaches a maximum, whence it gradually diminishes. In
nickel, the transient current attains asymptotic values in strong
fields without changing its direction, while in iron, it is reversed
in & field of about 200 C.G.S. units, when the twist is small.
The increase after the reversal is not pronounced, but becomes
finally asymptotic.
$3. Longitudinal magnetization produced by twisting
a circularly magnetized wire.
The longitudinal magnetization produced by twisting a cir-
cularly magnetized wire presents the same character as the tran-
sient current above described. The experiment is very difficult
on account of the heating of the wire. To avoid the rise of
temperature, the iron or nickel wire to be tested was covered with
urushi (Japan lac) which has the special property of being a very
good insulator while, at the same time, the melting temperature
is comparatively high. The wire thus insulated was stretched
in the axial line of a secondary coil whose diameter was 1.5 cm.
and whose total number of turns was 540, and the current of
cold water was kept flowing about it to keep the temperature
272 H. NAGAOKA AND K. HONDA:
of the wire uniform. Thus maintaining the electric «
in the wire constant, it was twisted and the induced ct
in the secondary circuit due to the longitudinal magneti
thereby developed was measured by the ballistic method.
Some of the results of observations are given in the ft
ing table and graphically shown in Fig. 5.
Iron wire: diam.= 0.888 mm. Nickel wire: diam.= 0.965
length =20.74 cm. length =20.94 c
0=15° 0 = 50° 0—15° | §—50'
C Qx10 C Qx10 C Qx10 C Q:
021 0.7 | o21 16. | 01 — 45 | 0.14 —
085 3.7 | 0.69 453 | 06 —111 | 08 —
1.53 205 | 149 891 | 156 —205 | 090 —
236 314 | 219 1110 | 240 -239 | 205 —
393 365 | 3.27 1956 | 3.35 —256 | 287 —
472 33.6 | 465 1242 | 436 -265 | 442 -
6.55 292 | 59 1154 | 5.86 —272 | 737 —
7.89 219 | 829 1096 | 7.95 —269 | 1034 —
12.82 139 | 1248 962 | 1089 —256 | 1533 —
19.01 10.9 | 1708 67 | 1407 —243 | 20.85 —
2499 58 | 2437 518 | 2018 -219 | 23.04 -
28.64 58 | 2914 432 | 2646 —206 | 2613 —
C denotes the total current through the wire express
amperes; 6 and Q have the same meanings as before.
As will be seen from the figure, the quantity of in
electricity in the secondary circuit, and therefore the
tudinal magnetization developed, by twisting a circularly
|
| netized iron wire attains a maximum, when the mean ci
field is about 10 units. It then decreases, but in spite :
TORSION AND MAGNETIZATION, 273
constant stream of water, the heating due to electric current
prevented the experiment from being pushed to the point where
the direction of the current is reversed. However, to judge from
the course of the curve, the tendency is such that there is a
reversal. In nickel, the direction of the induced current is
opposite to that in iron, and the total quantity of the current
attains a maximum, whence it continually diminishes, but not
to such an extent that the current ultimately changes its
direction.
These experiments show that the twist produced by the
combined action of the longitudinal and circular magnetizations,
the circular magnetization produced by twisting a longitudinally
magnetized wire, and the longitudinal magnetization caused by
twisting a circularly magnetized wire are characterized by having
various peculiarities, which are common to all of them. This
can not be a mere chance coincidence ; we shall have to ascribe
these allied phenomena to the same common cause.
In the experiments of this and the last paragraphs, we were
assisted by Mr. 8S. Shimizu, a post-graduate in physics, to whom
our best thanks are due.
$4. Theory.
As already remarked in our last paper on magnetostriction,
Kirchhoff’s theory can be extended to the study of the relation
between torsion and magnetization, exactly, in the same manner as
was done by Maxwell and Chrystal to explain the Wiedemann
effect. There we found that the mean circular magnetization called
into play by twisting a ferromagnetic wire of radius À through
angle » amounts to
274 H. NAGAOKA AND K. HONDA:
1 Pal
in field H, and that the mean longitudinal magnetization ca
by twisting a ferromagnetic wire carrying an electric curre
amounts to
_ zw KO. | (B)
The reciprocal relation between these two phenomena is
apparent at a glance. We shall next show how the same pl
mena are reciprocally connected with the torsion produced b
interaction of the longitudinal and circular magnetizations.
The stress components in a magnetic medium as give
Kirchhoff are as follows:
x= -(z tht ét (gr +E-#) a+ +
si Ar B 2 2\4r | ‘
, 1 2 x 2 2
th+ gr glg +E-#) (a+ #4
(4
u el
: Ar 2 2 \ Ar : F
Taking the axis of z in the axial line of the wire, and two
axes in the plane perpendicular to it, we see that the compc
TORSION AND MAGNETIZATION. 275
etic forces in a longitudinally magnetized wire traversed by
lectric current are
a= — h sin 6, B=h cos 6, Y=JH,
e h denotes circular field given by
2Cr
h= Er
ing the current, r the distance of the point from the axis
e wire, R the radius, and @ the angle between r and the
of x.
The stress components in ferromagnetic medium acted upon
ne forces above specified are given by
Pe ee eee ee atte.
= - + k + +) h sin 0 + Ha + k — k ) (2 +h )
(4 +4 + Sr cos? 8 + - (= 1 +k-%) H°+ lé
4x 2\ 47 2
1
KUN... PES ee ee
(Gar tht) la rer) (us)
1
j’!
+ k + =) h ZH cos 6,
il x) A H sin 6,
=¥.=(4> Sa ) A’ sin 0 cos0
moment about the axis of the wire is given by
N=[ | (z, x—Z, y) dxdy.
=-ff (a +2+53 a pues
2/1 R
= - ul + b+-5-) CH ( "rd,
1
— 7 (a +h + >) CHIE.
276 H. NAGAOKA AND K. HONDA:
Since > and # are very small compared with %”, the tors
couple twisting the wire amounts nearly to
Sg - CHR'= ward x Cross section. (©)
Since the amount of torsion of a cylindrical wire by a ;
couple is inversely proportional to the fourth power of its ra
it is evident that for given longitudinal current and field,
angle of twist is inversely proportional to the square of
radius. This inference was approximately verified in the pr
ex perintents.
In deducing the three formule (A), (B), (C), we
not, strictly speaking, put &” outside the sign R sahen
because the strain coefficient depends on the field strength, w
is not uniform in a wire traversed by electric current. E
in these formule, we shall have to use a mean value to o
a close approximation.
The mutual relations between twist and magnetizatior
embodied in the three formule above given. There we ı
that the strain coefficient £” determines the nature of the
different phenomena studied in the above experiments.
fact that the coefficient #” is principally determined by
elongation in the ferromagnetic metal accounts for the
analogy between the said phenomena and the elongations
to magnetization. As the above result imports, the analo
not exact, inasmuch as the elongation is also affected by 1
depending on k’, which depends mostly on the change of vol
In order to test the consequences of the theory as re
the twist produced by the joint action of circular and longitu
magnetizations, we have calculated the twist by assuming
TORSION AND MAGNETIZATION. 271
3 of %”, calculated from the changes of volume and of
h in iron and nickel ovoids. Graphically represented (Fig.
he fields of maximum twist by calculation coincide nearly
that given by experiments, and the reversal of twist in iron
placein low fields as actually found by observation. The
itative differences are, however, tolerably large in iron,
n nickel the amount of twist is nearly coincident with the
imental values. Calculating, in the same manner, the
ity of the transient current produced by twisting longitudi-
magnetized wires, we find a close coincidence between the
imental and theoretical values in nickel, but the difference
rably large in iron. In using the strain coefficients, we
always bear in mind that these values are widely different
ling to the nature of the specimen; especially with wires,
e not sure of its being magnetically isotropic. The apparent
pancy would probably be lessened, if we could measure the
as well as the strain coefficients on the same specimen.
remarkable qualitative coincidence as regards the existence
ximum twist and its reversal in iron are convincing proofs
he theory, so far as we know at present, admits of con-
ig various experimental facts in a common bond.
\s regards the mutual relations among the three different
ymena above enumerated, it will suffice to state that several
em have already been noticed by G. Wiedemann in his
ches on the relation between torsion and magnetism. He
ially studied the relation between permanent torsion and the
of magnetizing the twisted wire. The principal object of
searches was to expose the different aspects of the phenomena
ved in the relation between torsion and magnetization in order
ing to light his ingenious theory of rotatory molecules.
278 H. NAGAOKA AND K. HONDA:
Elegant as it at first sight appears to be, Wiedemann’s theo
abounds with hypotheses which we are not always warranted
making.
In his work on the applications of dynamics to physics a
chemistry, J. J. Thomson has propounded a new method
investigating the mutual relations between the effects of varic
physical agencies. He showed that the existence of a cert:
phenomenon involves as a natural consequence that of anotl
reciprocating with it. As an application of his method, he shov
that if the wire be twisted by the interaction of longitudinal :
circular magnetizations, a transient current will be produ
simply by twisting a longitudinally magnetized wire anc
longitudinal magnetization will be developed by twisting a ı
cularly magnetized wire.
The peculiar feature of Kirchhoff’s theory lies in the sim
and natural way of elucidating the relations between the vari
kinds of strain caused by magnetization and the effects of st
on magnetization. Just as we can study the various elastic
haviour of isotropic bodies by knowing the bulk- and strel
moduli, we have to deal, in Kirchhoff’s theory, with the st
coefficients % and &” which play the rôles of different mo
in the theory of elasticity.
The reciprocal relations between the strain caused by n
netization, and the effects of stress on magnetization, as fo
by actual experiments, will be found to be of paramount
portance in arriving at a correct theory of magnetostrict
The strain accompanying the magnetization of ferromagnetic m
will be determined, when we know the effects of stress on n
netization and vice versa. As regards the relations between t
TORSION AND MAGNETIZATION. 279
magnetization, we may conveniently place them under the
ying parallel statements:
ins produced by magnetization.
—(Experiment and theory). A
idinally magnetized wire is
1 by circular magnetization.
—(Experiment and theory). A
rly magnetized wire is twisted
gitudinal magnetization.
— Experiment and theory). Up
lerate fields, the twist produced
> longitudinal and circular mag-
ions of an iron wire is op-
to that in nickel.
(Experiment and theory).
wist due to longitudinal mag-
tion of a circularly magnetized
r nickel wire reaches a maxi-
in low fields.
-(Experiment and theory). In
fields, the twist due to longi-
l magnetization of a circularly
tized iron wire is reversed and
place in the same direction as
kel.
Effects of Stress on magnetization.
(a!)—(Experiment and theory)
Twisting a longitudinally magnetized
wire gives rise to circular magneti-
zation.
(6')—(Experiment and theory).
Twisting a circularly magnetized wire
gives rise to longitudinal magnetiza-
tion.
(c’)—(Experiment and theory). Up
to moderate fields, the transient cur-
rent, or the longitudinal magnetization
produced by twisting a longitudinally
or circularly magnetized wire respec-
tively, is opposite to that in nickel.
(d’)—Experiment and theory).
The transient current produced by
twisting a longitudinally magnetized
iron or nickel wire reaches a maxi-
mum in low fields.
(e’)—(Experiment and theory). In
strong fields, the direction of the
transient current produced by twisting
a longitudinally magnetized iron wire
is reversed and is in the same
direction as in nickel.
n his paper on the principle of least action, Helmholtz”
laced the reciprocal relations of a dynamical system under
heads. Denoting the generalized co-ordinates, the veloci-
Helmholtz, Crelle’s Journal 100, p. 137, 1886; Abh., 3, p. 203, 1895.
280
ties, the accelerations and the forces by p’s, g’s, q's, and Ps,
the relations are generally expressible by the equations
OP, OP,
1) <= +,
Op, Ip,
op OP,
2 a ___ıb :
( ) dq, 0Qa
OP, _ 9P,
©) agi, pe
It will be easily seen that the relations above cited belong t
case (2).
The greatest difficulty that we encounter in establishing th
relations between the effects of stress on magnetization and th
strain caused by magnetization lies in the great difference «
strain coefficients according to the nature of the specimen. Ifa
the experiments be performed in a proper manner on one an
the same specimen of ferromagnetic metals, we may feel assure
of being able to discern the true merits of the theory, or |
detect its various defects, not only from qualitative points of vier
but also in various quuntitative details.
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1e Interaction between Sulphites and Nitrites.
By
Edward Divers, M. D. D. Sc., F. R. S., Emeritus Prof.,
and
Tamemasa Haga, D. Sc., F.C. S.,
Professor, Tökyö Imperial University.
The present paper gives an account of a series of experiments,
esults of which seem to leave no room for doubt as to the
of the following propositions respecting the sulphonation
itrites :—(1) normal sulphites are inactive upon nitrites ;
yyrosulphites are not active to their whole extent upon ni-
; (8) pyrosulphites are active in their entirety upon nitrous
or its equivalent of nitric oxide and nitric peroxide (nitrous
s); and (4) sulphurous acid and nitrous acid or the oxides
water equivalent to them interact of themselves and in such
y that the base of the sulphite, that may be used in place
e sulphurous acid, is needed only to preserve from hydro-
the products of their interaction. Concerning these asser-
we would point out that the first directly contradicts the
usions drawn by other workers from their experiments ; the
282 E. DIVERS AND T. HAGA:
second is novel like the first, the facts on which it is based h
been misunderstood ; the third has indeed been already enunc
but not with intention to limit it to a strict interpretation
only theoretically, without any experimental treatment of it
lastly, the fourth has been also made before and upon the
of experiment, but experiment quite inadequate to justify i
The establishment of these propositions, taken along
what we have already published as to the constitution of Fr
salts will then allow of the further assertion being made,
the interaction of nitrous acid with a pyrosulphite results in
conversion into a two-thirds normal hydroximidosulphate
water), and that all the other sulphazotised salts are seco
products simply derived. It is thus established that the
interaction between sulphites and nitrites is one of the gi
simplicity, instead of being full of complications, as hi
believed.
I—a. A Normal Sulphite inactive upon a Nitrite.
Dipotassium or disodium sulphite is quite inactive u
nitrite. In establishing this fact we have mixed soluti
normal sulphite and nitrite in proportions varying in di
, experiments, and left them in closely corked flasks, almos
for days and for weeks. No change has ever happened.
coloured with rosolic acid, a drop of dilute acid would a3
time, as at first, discharge the colour. Had any action oce
alkali hydroxide must have been generated (as to the poss
of which see sect. II. 5.). A portion of the solution to
had been added a drop of dilute sulphuric acid reacquire
pink colour of the rosolic acid when left to stand for som
INTERACTION BETWEEN SULPHITES AND NITRITES. 983
—the minute quantity of pyrosulphite which the acid had pro-
duced having slowly interacted with the nitrite, but when that
was used up, no more action occurred and, at any time added
one drop of dilute acid would again remove the colour of the
solution.
No further proof of the activity upon a nitrite of a normal
alkali sulphite is wanted, but additional evidence of the fact is
readily obtainable. Thus, in the case of the potassium salts,
while the action of pyrosulphite upon nitrite shows itself (except
in cases of high dilution) by the formation of the insoluble
nitrilosulphate, no separation of this salt occurred in the above
experiments. Again, barium chloride, added, at any time, to the
mixed and to litmus very alkaline solution of normal sulphite and
nitrite, precipitated all the sulphur as sulphite (with also a very
little sulphate) and left the nitrite in solution, neutral to litmus.
Had sulphazotised salts been present, precipitation of the sulphur
would have been incomplete, and the mother-liquor would have
preserved a strong alkalinity to litmus and when acidified soon
have deposited barium sulphate in the cold and at once if boiled.
Fremy, Claus, and Raschig all believed in the activity of di-
potassium sulphite upon potassium nitrite, although the last
named chemist recognised the value also of the pyrosulphite, as
Berglund had done before him. Fremy apparently used sulphite
neutral to litmus and took it to be the normal salt, and Claus
certainly did so. Since, therefore, they used sulphite which was,
for the most part, pyrosulphite, no evidence upon the point in
question can be gathered from their work. We would account
for Fremy’s finding sodium sulphite inactive upon sodium nitrite,
otherwise inexplicable, by assuming that the solution of sodium
sulphite which he tried happened to contain no pyrosulphite.
284 E. DIVERS AND T. HAGA:
Claus’s statement that some potassium sulphite neutral to 1
was as active upon nitrite, after he had added potash in ‘ ex
to it, as it was before only requires us to assume that the e
spoken of was large enough to give the solution a mar
alkaline action upon litmus and yet small enough to leave ı
pyrosulphite unchanged.
I.—b. Potassium Hydroxide not a Factor in the Formatior
the Sulphazotised Salts.
That a normal sulphite, potassium or sodium, remains
inactive upon nitrite, when alkali hydroxide is added, was:
tained by leaving the three substances together in solution
closed flask for some time, as in the experiments whe
hydroxide was present, and then precipitating with barium
ride after addition of ammonium chloride, and finding no su
compound left in the filtrate. (Ammonium chloride pre
precification of hydroximidosulphate, this Journal 7, 48, (
On the many occasions we have had to prepare sulp
tised potassium salts by submitting solutions of nitrite and hy
ide to the action of sulphur dioxide, taking care to kee
solution briskly agitated, we found that, even in ice-cold
tions, precipitation of these very sparingly soluble salts
began from the point at which there was no more hydr
left, and then went on freely until the solution had be
neutral to lacmoid paper. In proportion as the hydroxide
appeared, sulphite became abundant, whilst from the time
the replacement of hydroxide by sulphite was complet
quantity of sulphite steadily decreased as the sulphazotised
formed, sulphazotised sodium salts being very soluble no ]
INTERACTION BETWEEN SULPHITES AND NITRITES. 285
pitation occurs during their preparation, and with these, therefore
we made an experiment to determine quantitatively what happens
up to the point when the last portion of hydroxide disappears, a
point indicated by rosolic acid losing its pink colour.
Washed sulphur dioxide was sent in a steady stream into a
solution of 11.21 grams real sodium hydroxide and 19.34 grams
sodium nitrite in about 198 grams water, kept in active motion
and immersed in ice. ‘The sodium compounds were in molecular
proportions ; more nitrite would not have mattered. In a short
time a 10 cc. pipetteful was removed ; soon after a second; and
not very long after a third one, just when the pink colour of the
rosolic acid present had disappeared. The three portions were
treated alike. Each was mixed with excess of solutions of am-
monium and barium chlorides and the precipitate filtered off,
oxidised to sulphate, washed with dilute hydrochloric acid, and
weighed as barium sulphate. The ammonium-chloride filtrate
was evaporated to dryness, during which operation all the sul-
phur of the sulphazotised salts was converted to sulphate by the
nitrite and ammonium chloride. The soluble salts were washed
out with dilute hydrochloric acid, and the barium sulphate
collected and weighed. The solution of salts removed from the
barium sulphate was concentrated and then heated under pressure
for hours, after which it was found to be still clear and there-
fore free from sulphate, which must have formed by hydrolysis
had any sulphazotised salt escaped decomposition during the
evaporation with nitrite and ammoniuin salts.
We give the quantities of sulphur dioxide found in each
pipetteful as sulphite and as sulphazotised salts, and also state
these quantities as parts per hundred of the total sulphur dioxide
which had entered it.
286 E. DIVERS AND T. HAGA:
SOBUEMIEE _ x Tats 10:6; 2nd 10 cc. 3rd 10x.
Sulphite .0662 grms.—96.69% .1204 grms.=96.5% .3996 grms.=91.5
Sulphazot. .0023 „ = 3.4% .0047 ,, 3.9% .0365 „ = 8.5
It will be seen that all but 3.5 per cent. of the sulphı
dioxide entering the solution in the early stages of the expe
ment remained in the form of sulphite, and that even up to t
time when the last of the hydroxide had been consumed, all b
8.5 per cent. of the total sulphur dioxide was in the state of sulphi
That it must be impossible to prevent all temporary local excess
sulphur dioxide will be at once admitted, as also that it m
be difficult in the later stages to keep down this local excess
very narrow limits. Therefore it will seem in the highest degı
probable, if not certain, from this experiment that sulphur dio
ide, equally with normal sulphite, does not act upon nitrite
presence of alkali hydroxide.
Now Fremy believed that potassium hydroxide helps t
formation of sulphazotised salts and endeavoured, accordingly
keep some of it always present when passing sulphur diox
into a solution of potassium nitrite. This he did by adding
occasionally during the process, and to such an extent that
the end of the operation the mother-liquor of his salts was
ways not merely alkaline but caustic and destructive to fil
paper. Claus did not go so far as to believe that the pote
exercised any specific influence upon the action between the sulpl
dioxide and the nitrite, but, agreeing with Fremy as to
value in precipitating and preserving the sulphazotised salts,
adopted the precaution to stop passing in sulphur dioxide wh
the alumina contained in the potassium hydroxide began to p
cipitate, since this: occurs while the solution is still strong
alkaline to litmus. Raschig, in attempting to prepare Frem
INTERACTION BETWEEN SULPHITES AND NITRITES. 287
sulphazate, also used precipitation of alumina as the indication to
stop the process while yet alkali remained. —
Thus, then, it would seem, Fremy, Claus, and Raschig, the
last in less degree, have all prepared sulphazotised salts without
difficulty, under conditions which we pronounce to be in-
compatible with their production, To remove this apparent
contradiction in results it is sufficient to assume, for one thing,
that, in Fremy’s way of working, success followed only because,
temporarily and locally, the point of saturation of the alkali
was reached and exceeded again and again where the gas entered
the solution,—a state of things, never avoidable altogether, above
all at the time when the potassium hydroxide is nearly exhausted:
There is nothing to show that, to check this, he kept his solu-
tion well agitated. Secondly, we can assume, with great proba-
bility that his solution often lost its alkalinity between the additions
of the hydroxide which he made. Working as we believe he
actually did, we have found it easy to get results such as his.
So far as Claus and Raschig followed Fremy’s method, their
results are equally open to objection, while it is to be remarked
of their alumina indicator that, not only is normal sulphite
alkaline to litmus but, as we have found, any aluminium present
is precipitated as hydroxide just when the sulphur dioxide has
converted all alkali into normal sulphite. They will, therefore
in their experiments have preserved none of the alkali unchanged
and most probably have generated also some pyrosulphite. There
is besides indirect evidence in Claus’s work that normal sulphite
is either inactive or only very slowly active upon nitrite, for when,
having taken no excess of this salt, he stopped the process just
after precipitation of alumina, much of this nitrite remained in
the solution, while, as we have just pointed out, much normal
288 E. DIVERS AND T. HAGA:
sulphite must also have been present. The two salts were, then
fore, together in solution unchanged. Raschig, too, found th
sulphite and nitrite are inactive upon each other when in presen
of potassium hydroxide dissolved in only its own weight of wate
IIl.—a. Even Pyrosulphite only active upon a Nitrite till
il has become Normal Sulphite.
Pyrosulphite, neutral to lacmoid paper and containing the:
fore, neither sulphurous acid nor normal sulphite, freely sulph
nates nitrite, but is far from being all consumed in the proce
as it has been represented to be by Claus, Berglund, and Rasch
Quantitative experiments have shown us that, when pyrosulph
is left in solution with excess of nitrite in a closed vessel for
considerable time, about one-third of the sulphite remains inact
by becoming converted into the normal salt, separable, as
other cases, from the sulphazotised salts by precipitation wi
barium chloride in presence of ammonium chloride. From t
it follows that 3 mols. pyrosulphite are needed to convert
mols. nitrite into hydroximidosulphate (this Journal, 7, 1
and not 2 mols. only, as had been supposed. The third m
sulphite remains unavoidably in the solution but all the nitr
sul phonated :—2NaNO, +3Na.8,0,+ OH,=2Na,HNS,O, +2NaS
using less pyrosulphite, some nitrite remains at the end alo
with normal sulphite. That sodium pyrosulphite is not eas
all used up in sulphonating sodium nitrite was observed
Raschig.
Not only hydroximidosulphate but a little nitrilosulphate
‘formed when a pyrosulphite acts upon a nitrite, but this ne
INTERACTION BETWEEN SULPHITES AND NITRITES. 289
never be enough, with ordinary care, to cause much less than
one-third of the sulphite to remain inactive. If excess of pyro-
sulphite is used the interaction appears to be—NaNO, +2Na8,0,=
Na,NS,O,+ Na,SO;, but we have not made any quantitative deter-
mination of the sulphite remaining, the qualitative evidence being
sufficient.
The interaction between pyrosulphite and nitrite proceeds at
first very rapidly and with great elevation of temperature, but,
when the temperature is kept down by cooling, soon slows down,
so as to require many hours for its completion. The normal
sulphite seems here to inhibit the action of the pyrosulphite, just
as does the salt of a weak acid inhibit the action of that acid,
an effect now well recognised. This consideration points to the
propriety of looking upon the passage of pyrosulphite to norma!
sulphite as its action as an acid upon the nitrite, and not as the
yielding up of half of its sulphurous acid for the sulphonation
of the nitrite, the interactions being 2NaNO, + Na,S,0, + OH, =
3HNO,+2NaSO, and then 2HNO.,+2Na.S,0,=2Na,HNS.U.-
(see section III a).
II.—b. Alkah not produced in the Sulphonation of
a Natrite.
One of the most remarkable things, according to Claus, is
the production of potassium hydroxide by the formation of Fremy's
salts through the agency of a sulphite. He explained this pro-
duction by the equation—KNO,+2K,SO, + 20H, = K,HNS,0, +
3KHO. Such an equation was also published by Berglund
(Lunds Univ. Arskr. 1875, 13, 14). Raschig gave the same
equation for results obtained by himself and, in order to express
290 E. DIVERS AND T. HAGA:
other results, gave also the equation—NaNO,+2NaH!
Na,HNS,O,+ NaHO. Finding also, and again in agreement
Claus, that dipotassium hydroximidosulphate does not cor
at once or even at all with potassium hydroxide, he arguec
this salt cannot have a similar constitution to that of Fr
“basic” sulphazotate because potassium hydroxide is pro
along with it instead of being combined with it as Fr
‘basic’ sulphazotate.
Now, all this is wrong in fact on the part both of
and Raschig, as we have already shown (this Journal 7
or here show in other sections of the present paper, except
the generation of alkali hydroxide, which we now proce
deal with. Claus’s emphatic statement, supported as it
Berglund and by Raschig, that potassium hydroxide is f
when a sulphite meets a nitrite in solution, rests upon no
evidence than what we now set down in full, recalling th
(section I, a.) that between the normal sulphite and nitrite
is really no activity of any kind. A solution of sulphite
neutral to litmus and a solution of nitrite of either potassi
sodium become hot and strongly alkaline to litmus when |
together, and then contain much hydroximidosul phate and n
sulphate, both neutral to litmus, which soon crystallise |
they are the potassium salts. That is all these chemists |
evidence for the production of the hydroxide; let us add to
facts that addition of excess of barium chloride removes 2
alkalinity. It follows, since pyrosulphites are a little a
litmus and normal sulphites are very alkaline to it, th:
phenomena depended upon offer no grounds whatever fo
belief that alkali hydroxide is produced. Except by the ı
lime, baryta, or other base, there is, we believe, only one w
INTERACTION BETWEEN SULPHITES AND NITRITES. 391
which potassium hydroxide can be generated from potassium
sulphite and that is one made known by us, namely, treatment
of the sulphite first with nitric oxide and then with alcohol and
water (this Journal, 9, 106).
11l.—a. A Pyrosulphite all active upon Nitrous Acid.
As remarked at the end of section II. a, a pyrosulphite
appears to act as an acid upon the nitrite and then sulphonates
the nitrous acid itself, only indirectly, therefore, sulphonating
the nitrite of a metal or of ammonium. One third of the pyro-
sulphite should, accordingly, be replaceable by some other acid,
and so it proves to be (section III. d). It is not new to formulate
the sulphonation of HNO,, and to speak of ‘ nitrous acid’ as the
reacting substance, for (passing over Fremy) Raschig has already
done so. But, whereas we would be understood to confine the
activity to nitrous acid itself, or its acidic equivalents (sect. III. d),
such was not the thought of Raschig, who only wrote H as a
general symbol, and ‘acid’ as a general term, while representing
metal nitrites as active by generating alkali hydroxide.
Nor is it new to learn that nitrous acid can be sulphonated.
By treating a dissolved sulphite with nitrous acid (nitrous fumes)
Fremy did succeed in obtaining sulphazotised salts, but the dif-
ficulty of moderating the flow of the gas, and the presence in it
of nitrogen peroxide and nitric acid made the operation so in-
convenient, he said, that he did not use it in preparing any of
the salts he examined, and gave no further attention to it. We
have taken up the matter, since untouched and unmentioned,
where Fremy left it half a century ago. Our work has been
very simple but very effective, and has consisted in subjecting a
2092 E. DIVERS AND T. HAGA:
solution of pyrosulphite (and of normal sulphite, but of th
treat in sect, III. 6.) to nitrous fumes which act as nitrou
hydride when of the right composition. The gases were
to be fully absorbed by a concentrated solution of pota
pyrosulphite kept cold in a flask immersed in ice and brin
well agitated. Soon an abundant precipitation began of hy
imidosulphate mixed with a little nitrilosulphate. While
much pyrosulphite remained, the process was stopped anc
mother-liquor at once drained off. In this way we had
success in getting much hydroximidosulphate and only a
nitrilosulphate, notwithstanding the presence all along of so
pyrosulphite ; for, as was pointed out by us long ago, in
ciently cold solutions sulphonation to nitrilosulphate h
UCCUTrs.
The next five sections (III. 6, c, d, e, f) treat of vi
mixtures which, from the acid constitution of one of the
ponents, behave like that of nitrite and pyrosulphite, that
if each contained pyrosulphite and nitrous acid.
IIlL.—b. Normal Sulphite also all active upon Nitrous Ac
Replacing the pyrosulphite used in the last experime
the normal sulphite, it was found that again but in this
sradually, hydroximidosulphate precipitated, as well as very
nitrilosulphate. But here potassium nitrite proved to be an
product, which by gradually replacing the potassium sul
in the solution allowed the process to be carried very fi
wards completion. The reaction is expressed by the equat
SHNO,+2K,S0O,=2K NO, +OH,+ K,HNS,O., from which
seen that only one-third of the nitrous acid becomes sulphon
the rest being used up simply as an acid.
INTERACTION BETWEEN SULPHITES AND NITRITES. 293
This interaction is what, we believe, Raschig must inadver-
lly have got, when seeking to prepare Pelonze’s salt (hyponi-
osulphate) by the use of nitric oxide. The conditions are
urable to the production of the nitrito-hydroximidosulphate
s vol., p. 222).
III. —c. Action of Sulphur Dioxide upon Normal Sulphite
and Nitrite.
It has been shown in this paper (sect. I. 6) that the hydrox-
losulphate which, from the first, accompanies the normal
yhite as joint product of the action of sulphur dioxide upon
sli nitrite and hydroxide, keeps steadily to small proportions
he sulphite until nearly all the hydroxide has been saturated.
er that point is passed and when, therefore, sulphur dioxide
neeting a mixture of nitrite and normal sulphite, examination
he solution, by the method already described, shows that,
ig with a greater production of hydroximidosulphate than
re, there is pyrosulphite produced in no insignificant quantity.
s remarkable growth in the quantity of pyrosulphite, considered
ig with the fact (sect. II. a) that it is itself active upon
ite proves that much of the sulphur dioxide goes altogether
he normal sulphite. Only after the greater part of this salt
been acidified to pyrosulphite is the sulphur dioxide active
ulphonating the nitrite, which it then does by combining
1 it in conjunction with the pyrosulphite, thus :—
2K NO,+ K,S,0; + 28O,+ OH,=2K,HNS,O,;, the hydroximi-
ılphate being produced in this way with much greater facility
1 by the pyrosulphite alone because of its production not
ig accompanied here by the regeneration of normal sulphite
294 E, DIVERS AND T. HAGA:
with its inhibitory effect upon sulphonation (sect. II. a).
this change it still holds true that it is nitrous acid itself w
is sulphonated, the potassium leaving the nitrite to enteı
sulphonate radical, and being replaced by hydrogen.
Claus held that there could be no difference between
effect of submitting a nitrite to the action of a sulphite and tl
mixing it with a solution of hydroxide and then treating it
sulphur dioxide. The contents of this section and section -
show that essential difference exists between the courses
results of the two procedures.
IIL.—d. Action of Carbon Dioxide and of an Acid Carbo.
upon Normal Sulphite and Mitrite.
As would be expected, the gradual addition of one o
stronger acids to a solution of normal sulphite and nitrite
to the formation of sulphazotised salts. But even carbon di
and the acid carbonates of the alkalis are effective in excitin
tion in a solution of these salts. Concerning the activi
curbon dioxide there is nothing to add to what was publish
our first paper (J. Ch. Soc., 1887, 51, 661), that the gas is
slowly absorbed by the mixed salts in solution though n
either salt alone and at the mean temperature, and that s
azotiseu salts are then produced. Normal carbonates of the a
are inactive.
It is known that nitrites are not decomposed by c:
dioxide, and also that alkali carbonates are decompose
pyrosulphites as freely at the mean temperature as by sw
dioxide itself. Accordingly, we have found that potassiu
sodium acid-carbonate dissolved along with -excess of mc
INTERACTION BETWEEN SULPHITES AND NITRITES. 295
potassium or sodium sulphite gives off carbon dioxide to a cur-
rent of decarbonated air much to the same extent as when dis-
solved alone in water. But sodium acid-carbonate may be added
to an ice-cold solution of sodium pyrosulphite, containing also
much normal sulphite, and be only very gradually decomposed
with effervescence. Indeed, an ice-cold concentrated solution
of normal sodium sulphite will deposit some acid-carbonate when
charged with carbon dioxide.
It is, therefore, not surprising that sodium or potassium
acid-carbonate has a very marked action upon mixed normal
sulphite and nitrite. When the three salts are left together in
solution in a closed vessel for a day or two, much sulphazotised salt
is formed, so that after carbonate and excess of sulphite have
been precipitated by baryta and barium chloride in presence of
ammonium chloride, the filtrate from the precipitate when boiled
with acid gives much barium sulphate and reduces cupric hydr-
oxide freely. The interaction of the salts may be expressed
by the equation —K NO,+2K,SO,+8KHCO,=K,HNS.O,+38K,CO,
+OH,, but since the two-thirds normal hydroximidosulphate is
to a small extent converted by normal carbonate into a more
nearly normal salt and acid-carbonate (this Journal, 7, 32),
the change expressed by the above equation cannot proceed to
completion. |
III. —e. Action of Sulphur Dioxide upon Normal Carbonate
and Nitrite.
When sulphur dioxide is added to two mols. nitrite and one
mol. normal carbonate until the solution becomes acid to lacmoid
paper, the only products are hydroximidosulphate and carbon
dioxide. This was long ago pointed out by us, and also that
296 E. DIVERS AND T. HAGA:
sulphite and acid carbonate are intermediate products, the latt
of which separates for a time from concentrated solutions. VW
have made further experiments to ascertain the effect of the fir
portions of the sulphur dioxide in producing hydroximidosu
phate, which, where alkali hydroxide is used, we have shown
be insignificant.
These experiments were carried out in the same way as tho
for testing the effect when sodium hydroxide is employed (I.
but with the modification of making two pipettings each tir
instead of one, and of weighing both instead of merely meası
ing them, then in the one we determined the sodium, as sulph:
and used the result for calculating what fraction of the origir
solution the other quantity was in which we determined sulph
and sulphonates. We thus made ourselves independent of t
change of volume during the reaction caused by loss of carb
dioxide and gain of sulphur dioxide. We found in this w
admitting of no refined accuracy, that at a later sampling |
solution contained at most, as much as 3'/, per cent. less sodi
than at an earlier sampling, a difference however hardly la
enough to need attention.
The flask for receiving the portion for the sodium determ
ation was previously weighed empty but that for the otl
portion was weighed containing some concentrated solution
sodium hydroxide, placed there to arrest all action in the pipet
ful dropped into it. In the first portion could be seen, by
changes on standing, how necessary the sodium hydroxide v
for fixing the composition of the solution at the time it v
sampled ; sometimes acid carbonate was deposited by it, son
times hardly at all; sometimes the precipitated acid carbon
slowly disappeared sometimes not. The solution used contair
INTERACTION BETW EEN SULPHITES AND NITRITES. 297
1 part of sodium nitrite in 4.64 parts of water, besides the cal-
culated quantity of anhydrous sodium carbonate.
The results of the experiments showed that hydroximidosul-
phate was largely produced from the beginning, in proportion
to the sulphite also formed. Thus, in one experiment, when 25
per cent. of the sulphur dioxide required for complete sulpho-
nation had been passed in, 55.3 per cent. of it had become
sulphonate, the rest (44.7 per cent.) sulphite. When 53.6
per cent. of the sulphur dioxide required had been used,
74.9 per cent. of it had become sulphonate and 25.1 per cent.
sulphite. In another closely comparable experiment, when 33.7
per cent. sulphur dioxide of that required had been absorbed,
62.7 per cent. of it had become sulphonate and the rest sulphite;
when 44.4 per cent. of the whole had been used, 72.75 per cent.
of it had become sulphonate; and when 62.2 per cent. of the
whole had been used, 81.5 per cent. of it had become sulphonate.
That is to say, as for the last statement, when 20.2 grams of
sodium nitrite (with carbonate) had received 37.5 grams sulphur
lioxide, 23.3 grams of this had become sulphonate and 14.2
zrams had become sulphite.
Uniform results are here, however, as when hydroxide is
tarted with, only obtained by uniform working, of which the
ollowing experiment is a good example. A solution of sodium
nitrite and carbonate was divided approximately into one-fifths and
our-fifths, and both portions were treated, as nearly as could be,
like, their unequal quantities making the only difference. The
smaller portion when it had received 20 per cent. of the full amount
of sulphur dioxide was found to contain 61.8 per cent. of it in form
of sulphonate, 38.2 per cent. of it as sulphite. The larger por-
ion, having received 25 per cent. of the amount necessary for
298 | E. DIVERS AND T. HAGA :
its full sulphonation, was found to have only 53.3 per ceı
it as sulphonate and 44.7 per cent. of it as sulphite, as al
given; had we stopped here at 20 per cent. sulphur dioxi
we did with the smaller portion, the difference would have
more striking still. The difference observed was due to the sn
portion having, in relation to its quantity, received su
dioxide four times more rapidly than the larger portion ha
stream of sulphur dioxide having been steady and closely
in the two cases. The result was that local saturation wa
checked by the agitation of the flask in this case than
the much larger portion of solution was under treatment.
The lack of uniformity in the results here described,
not affect in the least the evidence they afford that the sul:
ation of nitrite in presence of carbonate differs greatly
course from that it runs in presence of alkali hydroxide.
Respecting the formation and destruction of sulphite i
process, this salt was observed to be produced rapidly un
quantity it had become equivalent to about one-eighth o
sulphur dioxide needed for sulphonation of all the nitrite.
for a time, its quantity remains nearly steady, all sulphur di
entering the solution during that time becoming sulph
Finally, it steadily lessens in quantity as more sulphur di
is added, and disappears just at the end of the sulphon
The more rapidly the sulphur dioxide is blown in at firs
less of it becomes sulphite, and the more sulphonates, as al
stated above.
One other striking thing observed in these experimen
the great variability of the point at which acid carbonate
precipitated, as well as the variability of its quantity.
quick working acid carbonate precipitated much earlier a
INTERACTION BETWEEN SULPHITES AND NITRITES. 299
much larger quantity than in slow working, proportionately, that
is, to the fraction the solution had received of the quantity of
sulphur dioxide needed for complete sulphonation of the nitrite.
Thus, while, with quick working, acid carbonate separated in
abundance when 20 per cent. of all sulphur dioxide had been
absorbed, it only precipitated, and then much less copiously,
when 44 per cent. of all sulphur dioxide had been supplied
relatively more slowly to the solution. In another experiment
it showed itself only when 53 per cent. of the sulphur dioxide
had been added. The main condition, therefore, for early pre-
cipitation of acid carbonate is rapid addition of the sulphur
dioxide at first,—the same condition as favours growth of sul-
phonates at the expense of sulphite.
Now for the discussion of the results. It becomes highly
probable from a consideration of these results, together with what
we know of the several substances concerned, that the first action
or tendency to act of sulphur dioxide when it enters the solu-
tion is to convert carbonate into normal sulphite and acid
carbonate, and to leave the nitrite untouched, and that this ac-
tion remains prominent so long as much normal carbonate
is undecomposed. Though this cannot be shown experimentally,
it is certain that this action does take place, for its products
present themselves freely, products which could not be derived
from the sulphonation of the nitrite. Both normal sulphite and
acid carbonate are active along with sulphur dioxide in sulphon-
ating nitrite.
In accordance with what is stated in III. d. the normal
sulphite and acid carbonate together slowly disappear of them-
selves from the solution when addition of more sulphur dioxide
300 E. DIVERS AND T. HAGA:
is stopped, owing to sulphonation of the nitrite and reconversio
of acid carbonate to normal carbonate—
NaNO,+2Na,S0,+3NaHCO,=N3,HNS,O, +5N2,CO,+H3,0.
such a mode of sulphonation will therefore be also in operatic
when the entrance of more sulphur dioxide has not been arreste
but it is very slow in presence of normal carbonate and m:
be disregarded as a factor in the process of sulphonating whe
sulphur dioxide is also at work. Here we would insert that on
to simplify discussion do we speak of normal sulphite and carbı
dioxide, or even acid carbonate, being together unchanged ; the
substances, as previously stated, act on each other to a lar,
extent in ice-cold solutions, and in our work we met with pr
cipitated acid-carbonate at times when it could only be there in co
sequence of carbonic acid withholding sodium from pyrosulphi
That in the earlier stages of the process, when mu
carbonate is present, the normal sulphite plays a very small ps
in the sulphonation not only follows from the observation of
rapid increase in quantity at first but is also shown by its th
nearly constant quantity for a long time though sulphur diox
is still entering the solution and forming sulphonates. Or
later, as the carbonate gets consumed, does the sulphite beco
an important factor in the sulphonation by freely becoming py:
sulphite, for then its quantity rapidly falls.
The part played by sulphite in the early stages being tk
insignificant, we have to seek in the carbonates the source of t
early considerable sulphonation of the nitrite. It would beu
reasonable to assume, with acid carbonate present, that the norm
carbonate takes part in sulpnonation ; equally so to assume tl
it remains inactive to sulphur dioxide. We are therefore co1
pelled to recognise that sulphonation goes on only after conve
INTERACTION BETWEEN SULPHITES AND NITRITES. 301
tion of all carbonate locally present to acid carbonate and
sulphite has been effected. ‘Then the reaction that ensues is—
NaNO,+ NaHCO,+2S0,=Na,HNS,O,+CO,
When all normal carbonate in the solution has been acidified by
the carbon dioxide, the sulphite becomes as active as the acid
carbonate and neither salt gets consumed before the other.
While it seems certain that first the sulphur dioxide converts
the normal carbonate into normal sulphite and acid carbonate,
and only then produces hydroximidosulphate by acting on the
nitrite along with acid carbonate in the earlier stages and on
both this and normal sulphite collaterally in the later stages, the
experimental results show that local saturation must take place
largely where the sulphur dioxide enters the solution, since so
much sulphonate is produced along with the sulphite. In con-
sequence of the activity of acid carbonate, local saturation be-
comes twice as difficult to prevent as when hydroxide is used
in place of carbonate.
If in order to impede local saturation we slacken the rate
of passage of the sulphur dioxide into the solution, we meet with
a good amount of success. Thus, it was shown by the results of
experiments already given, that the slower rate gave proportion-
ately less sulphonate and more sulphite. But the effect of
slowness in passing in the gas has its limit, in consequence of
the continuous though slow interaction which takes place between
nitrite, normal sulphite, and acid carbonate whereby sulphite
disappears to give place to sulphonate. It follows that too slow
as well as too rapid an addition of sulphur dioxide is unfavour-
able to the accumulation of sulphite, rather than of sulphonate,
in the solution, and that a medium rate of supply is best for
raising the proportion of sulphite.
302 E. DIVERS AND T. HAGA:
There remains to be explained the great variability in |
commencement of precipitation of the acid carbonate. This tal
place the sooner the faster the sulphur dioxide is blown into |
solution. When it occurs in the earlier stages of the process
is, therefore, accompanied by greater predominance than usual
production of hydroximidosulphate over production of sulph
It does not however depend upon this, for while sulphur diox
liberates a molecule of carbon dioxide in changing carbonate i
sulphite, four mols. of it are needed to liberate one mol.
carbon dioxide in changing carbonate and nitrite into hydr
imidosul phate.
An explanation is suggested by a consideration of the |
that when working the process at a moderate rate, the |
crystallisation of acid carbonate takes place long after the jx
at which the solution must contain the maximum of the salt
least potentially, the point, that is, when half the carbonate
become either sulphonate or sulphite. When it does occur
quantity of it in solution has become much less. Only wl
crystallisation is started early by a very rapid addition of sulp
dioxide, does the acid carbonate continue to separate out in m
such quantity as it could do at the stage of the process reacl
The cause in one word is supersaturation. The acid carbon
it would seem, is slow to begin to precipitate from the solut
while that is not charged with carbon dioxide. At a medi
rate of working this only happens in the later stages, any nor
carbonate and even much normal sulphite present keeping dc
the quantity of carbon dioxide, but by a rapid rate of work
local saturation occurs and the acidified portion of the solut
then crystallises. Once crystallisation has been started, it proce
unchecked. In slower working when crystallisation only beg
INTERACTION BETWEEN SULPHITES AND NITRITES. 303
te in the process, the amount of salt separating is small, and
nerally depends then for its existence upon its power to resist
ie action of acid sulphite in ice-cold solutions. The solution
hen, potentially at least, it is richest in acid carbonate, was
und by us to crystallise soon, if left to stand in closed vessel,
though sulphonation which is destructive of acid carbonate was
owly going on in it.
IIL.—/f. Primary Action of Sulphur Dioxide upon a Nitrite.
Solution of sulphur dioxide added to that of potassium or
dium nitrite produces a sulphate and either nitric or nitrous
ide, according as one or other of the interacting substances is
excess. That is the ordinary well-known result, but there are
70 ways of limiting the extent of the action so as to get either
ydroximidosulphate and nitrous acid or the undoubted products
their transformation. By these ways, the interaction of sul-
ur dioxide and a nitrite is shown to be—
2K NO, + 280, + OH, =K,HNS,O, + HNO,.
The more important way to thus limit the action is by an
‘periment first tried by Claus (Ber., 1871, 4, 508; see preceding
iper) which consists in adding an alcoholic solution of sulphur
oxide to excess of potassium nitrite in strong aqueous solution.
or this experiment gives, as we have ascertained, potassium
trito-hydroximidosulphate which precipitates and any! nitrite
hich boils off by the heat of the reaction :—
3KNO,+ 280.+ C,H,0 =KNO,,K,HNS,O,+ C,H, NO,.
y becoming ethyl nitrite the nitrous acid is rendered inactive
1 the hydroximidosulphate, which is thus saved from oxidation.
The other way of tracing the earlier action of sulphur dioxide
304 E. DIVERS AND T. HAGA:
upon a nitrite was found out by Raschig, when trying to pr
another point (sect. IV. a.). He added the nitrite to excess
sulphur dioxide, both being in very dilute and well coo
solution, evaporated down and neutralised the solution w
chalk, and again evaporated the filtered solution. After m
potassium sulphate had crystallised out, potassium amidosulpt
was finally obtained, as proof that hydroximidosulphate |
been formed at an earlier stage. Our own experiments h
yielded us an earlier product of the degradation of this cc
pound.
At the time when Raschig published his observation,
published (J. Ch. Soc., 1887, 51, 659) one of ours, that sil
nitrite and mercurous nitrite, when decomposed by sulp
dioxide solution, yield a substance answering to the copper
for hydroxylamine. This we now know to be hydroxyam
sulphuric acid, but at the time we took it to be hydroxylan
itself. We have also found that, after adding a dilute solu
of sodium nitrite to excess of a cooled solution of sulphur dio:
and then blowing out of the solution the residual sulphur di
ide by a current of air, enough hydroxyamidosul phate (hydrol;
hydroximidosulphate) is present to be easily identified by
copper test for it. A hydroxyamidosulphate is distinguish:
from hydroxylamine in applying this test by finding that
mother-liquor of the cuprous oxide (which need not be filt«
off) gives sulphurous acid when acidified (this Journal III, 2
Though less successful than Claus’s experiment, Rasch
method is serviceable for showing that the alcohol used in t
plays only a secondary part. While excess of nitrite is succ
fully used in that experiment, the sulphur dioxide must be
excess in Raschig’s method. To understand this, it has onl
INTERACTION BETWEEN SULPHITES AND NITRITES. 305
emembered, firstly, that nitrous acid would oxidise hydrox-
osulphate at once, and secondly that sulphurous acid sulphon-
the hydroximidosulphate slowly enough to allow a little of
ing secured in a hydrolysed state.
.—a. Sulphonation of Nitrous Acid by Sulphurous Acid.
Fremy helieved that certain of his sulphazotised salts are
ed in the first action of sulphurous acid upon nitrous acid.
1 this belief Claus strongly dissented, holding that the presence
base (as salt) was essential to the production of these acids.
hig considered that his experiment of treating potassium nitrite
sulphur dioxide in excess (sect. III. f.) proved the correct-
of Fremy’s belief; but that cannot be admitted since potas-
is present in this experiment playing the part of base. It
wever, quite practicable to establish Fremy’s belief and that
ase whatever is necessary to bring about the formation of
azotised acids.
When a solution of sulphur dioxide, better ice-cold, is treated
a relatively small quantity of nitrous fumes passed on to
urface while it is being well agitated in a flask, and is
deprived of remaining sulphur dioxide by a rapid cur-
of air, or even by quick boiling, it will give a good reaction
hydroxyamidosulphuric acid with the copper test. A
deviation in the composition of the nitrous gas from that
rous anhydride is not of importance. If the object is only to
midosulphuric acid, the solution of sulphur dioxide is left
ind for a day after it has received the nitrous acid without
ling what is left of the sulphur dioxide. If it is then
rated on the water-bath and further concentrated in the
306 E. DIVERS AND T. HAGA:
vacuum-desiccator, the amidosulphurie acid will crystallise o
from the sulphuric acid with which it is accompanied (tl
Journal, 9, 230). .We have purified the acid by recrystallis
tion, and have hydrolysed it at 150°, by means of hydrochlo
acid, into acid ammonium sulphate; we have also complete
volatilised the acid by heat thus proving the absence of bs
accidentally derived.
Nitrosyl sulphate dropped into much excess of cooled sol
tion of sulphur dioxide also yields the hydroxyamidosul phate.
action with copper sulphate and potassium hydroxide.
IV.—b. Influence of the Base of the Nitrite or Sulphite.
Although Fremy held that sulphurous and nitrous ac
combine together, he did not believe that the resulting sul}
azotised acids could be obtained in this way, because of their
ability to exist in absence of a base. Moreover, he conside
that a strong base is influential in bringing about the format
of these acids, even though he had had no success with sucl
base as sodium. The only hydroximidosul phates he could preps
indeed, were those of potassium, but from ammonium nitrite
got the nitrilosulphate, and also obtained evidence that caleu
strontium, and barium nitrites are convertible into amida
sul phates.
We have just shown (sect. IV. a.) that the interaction
sulphurous and nitrous acids does not require the presence
any base at all for the actual production of sulphazotised aci
although such presence is essential to preserve unchanged t
first product of the interaction. To serve this purpose so
bases will doubtless be inferior to others, and those which
INTERACTION BETWEEN SULPHITES AND NITRITES. 307
freely form soluble pyrosulphites are difficult to work with.
rwise, the nature of the base seems to be a matter of in-
‘ence. Since the time of our early publications on the sub-
we have extended our experiments to several other nitrites
those of sodium, mercurosum, and silver, with the results
iow record.
Ammonium salis— Ammonium nitrite solution was prepared
riturating silver nitrite with its equivalent of ammonium
ride dissolved in about five times its weight of water, and
ing off silver chloride over the pump. To this solution,
it had been cooled in ice, was added a little less than its
valent of ammonia-water which had just before been con-
d to sulphite by passing sulphur dioxide into it. More
hur dioxide was then passed into the mixture until it red-
d lacmoid-paper. In this way the ammonium nitrite was
st all sulphonated, without any evolution of gas having
rred till just at the last, when slight nitrous fumes appeared.
> of the solution was hydrolysed and tested then with copper
1ate and potassium hydroxide; it was thus shown to have
tined abundance of ammonium hydroximidosulphate.
ther portion of the solution not hydrolysed gave a large
pitation of dipotassium hydroximidosulphate on addition of
sium chloride.
Barium salts.—Some barium hydroxide was converted into
uite by putting it in water and passing in sulphur dioxide ;
barium sulphite was then, for the most part, brought into
ion by passing in more sulphur dioxide. The product was
d gradually to a solution of a little more than its equivalent
arium nitrite, which had been purchased of excellent quality.
308 E. DIVERS AND T. HAGA:
Having neglected to cool our solutions we had reason to fear
that our experiment was a failure; for along with very much
precipitation there was a somewhat large evolution of nitrous
gases. But for our purpose we had been amply successful. The
solution was only faintly acid to litmus and remained so for
hours. Both it and the precipitate contained large quantities of
barium hydroximidosulphate. The precipitate also contained
sulphite and sulphate, the latter being the complement to the
nitrous fumes produced. The hydroximidosulphate was extract-
ed from the precipitate by a solution of ammonium chloride.
Calcium salt.—A solution of calcium nitrate, free from
magnesium, sodium, potassium, and other ordiary impurities, was
heated with well-washed spongy lead until nitrogen oxides and
ammonia began to form. The filtered, very alkaline, solution
was freed from lead by hydrogen sulphide not used in excess,
Calcium hydroxide was then removed by carbon dioxide. (it was
interesting to find that, contrary to assertion, carbon dioxide
cannot be used to precipitate lead in presence of calcium salt,
since calcium precipitates before lead.) A. solution of calcium
sulphite in sulphurous acid was prepared just before use, in the
same way as the barium salt had been, except that carefully pre-
pared calcium carbonate took the place of barium hydroxide.
With the calcium nitrite somewhat in excess of the calcium
sulphite, the solution of the latter was gradually poured into
the former, both solutions having ice floating in them at the time.
No gas was given off and only a moderate quantity of precipi-
tate was formed, which consisted of sulphite. The filtrate was
neutral and contained the full quantity of hydroximidosul phate
expected. |
Zinc salts—Zine nitrite solution was prepared by precipita-
INTERACTION BETWEEN SULPHITES AND NITRITES. 309
ting zinc sulphate with barium nitrite and filtering. Zinc sul-
phite in solution in sulphurous acid was made from zinc oxide
in water and sulphur dioxide. The two solutions, suitably pro-
portioned and with ice floating in them were mixed. No gas
came off, zinc sulphite precipitated, and the solution proved to
contain zinc hydroximidosulphate present in it in large quantity.
Mercurous salls and silver salts—Experiments, already re-
ferred to in sect. III. f. of this paper, sufficiently establish that
mercurous and silver nitrites are readily sulphonated. It is now
evident that the sulphonation of nitrites is a general reaction,
essentially independent of the nature of the base, which only
effects the preservation of the products. It is not the salts which
are sulphonated but nitrous acid itself.
V.— What Nitrous Acid becomes when Sulphonated.
In the paper preceding this it has been established that
neither the abundant experimental work of other chemists and
ourselves nor theoretical considerations afford any support to the
view that the double sulphonation of nitrous acid into a hydrox-
imidosulphate occurs in two stages, or that a monosulphonated
nitrous acid, ON‘SO,H or (HO),N‘SO;H, must be the first product
of its change. In the present communication it is shown that
the acidity necessary for the sulphonation of a nitrite points
clearly to the fact that it is in every case the acid itself, and
not its salts, which is directly sulphonated. We are, therefore, in
the position to affirm that the fundamental action in the form-
ation of all Fremy’s sulphazotised salts is the interaction between
actual nitrous acid and a pyrosulphite, in which they unite al-
ways to form the one substance, the two-thirds normal hydrox-
310 DIVERS & HAGA : INTERACTION BETW. SULPHITES & NITRITES.
imidosulphate corresponding to the pyrosulphite acting—
HONO+(SO,K)'SO,K=HON(SO;K).. The origin of all the other
salts out of this salt has been traced, partly by others and part-
ly by ourselves, and need not be gone over again here.
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Ueber die Wachsthumsbeschlennigung einiger Algen nnd Pilze dnrch chemische
Reize. Von N. Ono. (Hierzu Tafel X-XIII).
Vol. XIII, Pt. 2, published July 25th, 1900.
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‘CONTENTS.
Vol. XII, Pt. IL
Ammonium Amidosulphite. By Enwarp Divers and Masataka
Ocawa, Imperial: University, Tokyo ... ... ..
Products of heating Ammonium Sulphites, Thiosulphate, and
Trithionate. By Epwarp Drivers and Masataka pie
Imperial University, Tokyo... ...
~ Potassium Nitrito-hydroximidosulphates and. the "Non-
existence of Dihydroxylamine Derivatives, By Eowarn
Divers, M. D., D. Sc, F, KR. S., Emeritus Prof., and Tase-
MASA, Haca, D. Sc, F. C. $., Professor, Tökyö Imperial
University
Identification and Constitution of Fremy’ 8 Sulphazotized
Salts of Potassium, his Sulphazate, Sulphazite, etc.
By Epwarp Divers, M. D., D. Sc., F. R. 8., Emeritus Prof.
and TAMEMASA Haca, D. Sc, F. C. $., Professor, Tokyö
Imperial University
On a Specimen of a Gigantic ‘Hyaroid, Branchiocerianthus
imperator (Allman), found in the Sagami Sea. By
M. Mivasuta, Pigakushi. Science College, Imperial PER
Tokyo. (With Plates XIV & XV). … …
Mutual Relations between Torsion and Magnetization in
Iron and Nickel Wires. By IL NaGaoka, Rigakuhakusht,
Professor of Applied Mathematics; and K. Honpa, Rigakushi,
Post-graduate in Physics. (With Plates XVI)... 2. soe oe
The Interaction between Sulphites and Nitrites. By Epwarp
Divers, M. D., D. Sc., F. R. S., Emeritus Prof., and Tasremasa,
Haca, D. Sc, F. C.S., Professor, Tökyö Imperial University.
PRINTED AT THE “TOKYO TSUKWI TYPE FOUNDRY.”
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JOURNAL
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JAPAN.
VOL. XIII, PART III.
Rm tf AK # A A
PUBLISHED BY THE UNIVERSITY:
TOKYO, JAPAN.
1900.
MEIJI XX XIIT,
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All communications relating to this Journal should be addressed to the
Direetor of the College of Science.
tributions to the Morphology of Cyclostomata.
1l.-The Development of Pronephros and
Segmental Duct in Petromyzon.)
By
S. Hatta,
Professor in the College of Peers, Tökyö.
With Plates X VII-XXI.
[he following pages contain the second of a series of
s on the later stages in the development of Petromyzon, the
having already been published some time since in this
al (97, vol. x, pp. 225-237).
Jur knowledge of the earliest development of the excretory
s in the lampreys is still somewhat incomplete. This
nstance is, I believe, mainly due to the want of recent
igations upon the subject. Since the appearance of the
; by Muzrer (’75), Scorr (’82), SHipLEY (’87), GOETTE
Kurprrer (’90), and others, ten years or more have
t was my intention to publish this paper shortly after the appearance of my preliminary
n 1897, (Annot. zool. Jap., vol. I. pp. 137-140) but various unavoidable circumstances
mbined to cause the delay. Meanwhile I have had opportunities of renewing my study
ous points and the results here given are different from those of the preliminary
1 several important respects.
312 S. HATTA:
elapsed, and, so far as I am aware, no important fact
added during the interval by any renewed researches.
not, therefore, apologise for the publication of the pres
which embodies the results of my study on the subjec
the last few years.
The investigation of a longitudinally stretched, 01
merically arranged, organ-system such as the neural <
chorda, the pronephros, &c., is rendered peculiarly
in Petromyzon by the fact that the longitudinal ax
embryo in early stages describes a semi-circle. Some
in a series of cross-sections of such an embryo are
unavoidably cut in planes which meet the longitud
of the embryo in variable degrees of inclination ; conse
structure stretching in the direction of this axis is cu
obliquely, as, for instance, the neural cord shown in fi:
3, Pl. xvii. The vertical dimension of the cord is not in
long as is represented in these figures. To gather accura
of the form, the position, &c., of a given structure, —
it is necessary to compare series of sections of two
embryos of as nearly the same age as possible. Fu
difficulty of observation is greatly increased by the
yolk-granules in cells, especially by their reaction agai
ing fluids. Certain fluids such as haematoxylin, borax
&c., either stain diffusely all the parts, or act on the
more intensely than on the other contents of cells, s
can not discriminate different kinds of tissues. This
was, however, obviated by employing picro-carmi
embryos were stained in loto in this fluid, decoloriz
proper degree in acid-alcohol, and then washed in 909
In the sections of specimens thus prepared, the histologi
MORPHOLOGY OF CYCLOSTOMATA. 313
are distinguishable very clearly, being almost entirely dis-
ed in all parts except nuclei which are stained intensely.
I wish here to express my warmest thanks to my former
ers, Pror. Muirsuxurt and Pror. Isıma, for much in-
ble advice and for their kindness in looking through. the
ıscripts and the proof-sheets of this paper.
To avoid confusion the present paper will be divided into two
ms, the first of which will contain mere descriptions, while
he second will be given a historical review and conclu-
I. Descriptive.
A.—The Pronephros.
The youngest stage of the embryo which I have to deal
in the present investigation, is only a little advanced beyond
lipsoidal gastrula; it is intermediate between Stage I and
the list given in my first contribution above referred to
1, À and B, of that article). The head-fold forms a pointed
ıberance at one pole of the ellipsoid, while the blastopore
ainly visible at the opposite pole. The prominent neural
extends longitudinally from the anterior end of the head-
iberance to the dorsal lip of the blastopore..
For the general relation of the germinal layers at this stage,
er to fig. 1 which is drawn from the same series as the
n represented in fig. 18 of my former work.” It represents
nsverse section though the dorsal region of an embryo in
stage above described. As seen in this figure, the solid
mn mm re nm
8. Hatta, On the form. of the germ. lay. in Petr.: this Journal, vol. V, 1891.
314 S. HATTA:
neural cord is interposed in the median line between the right and
left halves of the mesoblast, underneath the epiblast composed by
of a single row of columnar cells. The epiblast is always limited
with a sharp contour against the mesoblast. The latter consists, on
each side, of a dorsal (d.), a ventral (v.) and a median (m.) row
of celle. The dorsal row represents the parietal, and the median
and ventral rows together form the visceral, layer. Both the
visceral and parietal layers of the mesoblast show, in the prox-
imal portion, a regular arrangement of a high columnar epithel-
ium, while distally this arrangement is more or less disturbed.
The above structures are localized in a small portion on the
dorsal aspect of the ovoid embryo, the remaining larger part
being taken up by yolk-cells compactly loaded with yolk-granules.
In this early stage, we can detect, therefore, neither in the
epiblast nor in the mesoblast any structure whatever which is to
be regarded as the rudiment of the pronephros.
Period 1.
In the next following stage, which corresponds approximate-
ly to the early part of Stage 11 (doc. cit., fig. 1, B), certain alter-
ations are met with in the mesoblast. The most important of
these is its metameric segmentation. This process first begins at
the neck” and proceeds both forwards and backwards. At the
present stage, there are found 16 or more mesoblastic somites.”
At about the time when this process has extended to the
anteriormost section of the mesoblast a second change arises in
the mesoblast, wz. the first appearance of the pronephros.
1) The term neck is used for the sake of convenience to designate the slender region where
the head-fold passes over into the hind globular part.
2)The exact number of the somites can not be reckoned, for the metameres become
indistinct posteriorly.
MORPHOLOGY OF CYCLOSTOMATA. 315
Fig. 2 represents a section through the middle of the fifth
ite” of the embryo mentioned above. ‘The general features of
germinal layers and of other primitive organs are essentially
same as before. The epiblast (ep.) is a single row of columnar
; and is sharply bounded from the structures beneath it; the
ral cord (n.) remains still solid.” In the mesoblast, however,
portions are distinguishable: the proximal portion composed
high columnar cells (mt.V and a.pn.2) which undergoes
americ segmentation, and the distal portion consisting of
ose group of somewhat irregularly shaped cells (/m.) which
ains unsegmented and constitutes the lateral plate. It
oteworthy that the former takes up the largest portion of
mesoblast, while the latter is represented by a small portion ;
e two portions represent respectively the parts of the same
e in the mesoblast of Amphiorus. However, between them
e exists no distinct limit in the lamprey; the one passes
lually over into the other. Although the visceral layer shows
ign of constriction, the parietal layer is notched at about the
dle of the proximal segmented portion (x). ‘The parietal layer
11 to this notch is composed of a regular cylindrical epithelium
n.2), which is slightly arched against the epiblast, so as
ause an indentation in the latter, while the visceral layer
1e corresponding portion consists of a more or less disturbed
of high columnar cells. As the subsequent history teaches,
proximal half of this extent (mt.V) represents the myotume”
) The somites are reckoned from the anterior end. The first, ie. the foremost lies
diately behind the auditory vescicle when the vescicle comes into view.
) The vertical diameter of the neural cord in tigs. 2 and 3 is shown greater than it
; is, the sections passing obliquely owing to the bending of the longitudinal axis of
mbryo, as noted in the introduction (p. 312).
) This term here means the Sclero-myotom of German authors.
316 8. HATTA:
and the distal portion (a.pn.2) constitutes the Anlage of the
pronephros—the name I assigned to the same in my preliminary
paper (’97). To avoid tiresome reiteration, I shall often speak
of them in the following pages simply as the “ Anlage” and when
it is necessary to refer to special ones, as the Anlage first, the
Anlage second, etc. in the order of their position in the series of
mesoblastic somites, beginning from the anterior end.
In the somite next following, 2e. the sixth (fig. 5), the
mesoblast shows almost the same condition as that already des-
cribed ; but in the somite preceding the fifth, 2.e. in the fourth,
the Anlage of the pronephros is a little more advanced in
development (fig. 3). On the left side of fig. 3, the fourth
myotome is sliced only at its hind wall (m2.IV), while, on the
right, it is cut through in the middle (mi.IV). On the right
half of the section, no marked progress is visible in the meso-
blast except the separation of the myotome, which shows a pen-
tagonal outline (mé.JV) and consists of high cylindrical cells
from the lateral plate formed of a loose mass of cells (Zm.). In
the left half, however, the state of things is quite otherwise:
the Anlage of the pronephros (a.pn. 1) together with the corres-
ponding visceral layer is entirely constricted off from the myotome
(mt.IV), although it is still connected with the lateral plate (dm.).
* The cells composing the Anlage are compactly set together and
arranged more or less in a radial manner; the Anlage itself is
rounded off at the proximal end. The lateral plate, on the other
hand, still consists of loosely grouped cells of variable shape.
The Anlage is thus always (before and after its separation
from the myotome) histologically very distinct from the lateral
plate; one might therefore often be misled to suppose that there
is no Organic connection between these two structures.
ei L
MORPHOLOGY OF CYCLOSTOMATA. 317
Fig. 4 represents a section passing between the two somites
e mentioned (the fourth and fifth) and is much magnified
s, x 2) to illustrate the finer structure of this portion. The
‘tural cells are all loaded with an enormous quantity of
1 corpuscles or yolk-granules. The epiblast (ep.) consists
single row of cubical cells and shows a sharp limit against
structures inside it. The irregularly polygonal mass of
(mi.V) is the anterior wall of the fifth myotome. Two
of variously shaped cells (/m.) constitute the lateral plate
h is histologically quite like that in the somitic portion, being
9osed of irregularly quadratic cells and tapering towards
distal (ventral) extremity (compare with the lateral plate,
in figs. 2 and 3). However, in the proximal portion,
e the Anlage of the pronephros consisting of a regular row
ll columnar cells would be found in the somitic portion, we
ere a group (x) of a few cells of faint appearance, forming
proximal edge of the lateral plate. By a comparative study
vo or more series of sections, it is easily demonstrated that
cells are a piece of the somite lying in front and have
ing to do with the Anlagen. To elucidate this point still
er, I have drawn fig. 7 which represents a section through
ntersomitic plane between the sixth (fig. 5) and the seventh
te (fig. 6). In this part the Anlage of the pronephros
veloped still more weakly, and the mesoblast remains in a
primitive state. In the proximal edge of the lateral plate
10 special structure is detected, but the edge fades away without
tinct limit. By comparison with Fig. 4, we can not find any
ed difference; thus, here likewise, there is no cellular con-
on between the Anlagen in the two succeeding somites.
From the filth somite backwards for 9 or 10 somites, the
7
318 S. HATTA:
mesoblast presents almost the same feature of the Ai
the fifth somite mentioned above. Fig. 5 represent
through the sixth somite, next behind the fifth; when
with fig. 2 no marked difference is detected in reg
structure of the mesoblast. But in some segments the d
of the Anlage is somewhat weaker than in others,
fig. 6, which shows a section through the seventh sor
in a segment posterior to this somite, we find the Anla;
pronounced as in the sixth somite. However, generall;
the Anlage of the pronephros in an anterior somit
further than that in a posterior. It must be remem
the somite in which the Anlage has already become
does not pass over suddenly into the somite in whic
of it is to be seen; but its development gradually gros
less distinct from the anterior to the posterior part, u
no trace of it is perceived.
In the present stage, therefore, the Anlage of the
is detected in more than 4 somiles but is completely sepe
the myotome only in one segment, viz. the fourth somi
has no genetic connection either with the Anlage à
following somite or with the epiblast; and it must ben
we find the foremost Anlage not exactly beneath the fourt
but always underneath its hind border.
Figs. 8-17 represent sections through a still |
bryo of this stage, having about 20 somites, The epi
the neural cord (n.), and the chorda (ch,) are essential]
as before. Being cut through somewhat obliquely, the
1)Such a case is very rare. In most +peeimens examined, the Anlage
the myotome is found in many segments, an that we can hardly decide in
the separation takes place first.
Digitized by Google
MORPHOLOGY OF CYCLOSTOMATA. 319
he two sides do not exactly correspond. On the right side
x. 8, the hind border of the fourth myotome (mi.IV) is
hrough ; the Anlage of the pronephros (a.pn.1) presents in
on an oval shape, consisting of columnar cells radially ar-
ed and containing a cavity of an irregular form. The
logical structure of the Anlage is as compared with that
y, 3, more or less loose”, and the Anlage itself is there-
lso distended. The lateral plate (/m.) shows, however, no
ed progress. The left side of this figure and the right
ys. 11 and 12 represent the section through the fifth myotome
”) and the Anlage of the pronephros (a.pn.2) for that somite.
Anlage presents almost the same development as that just
ibed. The left half of fig. 12 and the right half of fig.
hows the sixth somite (m/.VJ) and the Anlage belonging to
pn.3). It can be inferred from the arrangement of its
onent cells that the Anlage has been just constricted off from
nyotome, as is shown by the fact that the cells at the point
ed with x of the visceral and parietal layers are not yet
anged to form a continuous layer,—a condition which is
'ved not infrequently in younger embryos. Fig. 14 shows
he right side a section through the hind wall of the sixth
ome ; the Anlage beneath it (a.pn.3) is, therefore, the hind
of that represented on the right side of fig. 13: it is
ely cut off from the myotome (mé.VJ), and the two layers
us point have completely fused together, enclosing a com-
tively wide cavity. The same condition is observed in the
When the pronephric Anlagen are cut off from the myotome, their structure is at
wsened, that is, their component cells become loosely set together. Later the cells
ly themselves, and are again compressed by mutual pressure; giving a compact structure to
nlage—probably the same condition observed by van WyYHE in Selachian embryos
. 476).
320 S. HATTA:
section through the anterior border of the somite. This phase
of constriction is doubtless earlier than that shown in fig. 3. The
right half of fig. 16 and the left of fig. 14 is from the section through
the mid-plane of the next following somite, 2.e. the seventh.
The Anlage of the pronephros (a.pn.4) is not yet cut off from the
myotome (mi.VII), but the process is beginning as shown by a slight
constriction and an inclination to arch out, while the suddenly
weakened lateral plate (/m.) forms the distal (ventral) continuation
of it. This feature of the mesoblast reminds us of the youngest
stage of the Anlage described above (compare with figs. 2 and 5).
In the section passing through the anterior or the posterior border
of the somite too, the same condition of the Anlage, as on the
right side of fig. 14, is observed.
From the facts mentioned above, it is easily understood that
the separation of an Anlage from a myolome begins with the
constriction which takes place at the anterior and the posterior
border of the somite, and that the middle portion is the last to
be cut off, so that the cavity of the myotome (myocoelome) com-
municates with the peritoneal cavity, during some time, through a
narrow passage at the middle point.
Myotomes when cut off from the Anlage of the pronephros
assume a pentagonal form (see fig. 3) constructed of a dorsal, a
lateral, a ventral, and two median sides, each of which is composed
of a regular row of tall cylindrical cells. The dorsal and lateral
rows of cells constitute the parietal layer of the myotome, while
the three other sides represent the visceral layer (compare with
the description on p. 314).
For about ten segments behind the seventh somite, the
Anlage of the pronephros shows the same condition as that seen
on the right side of fig. 16. Fig. 17 represents a section
MORPHOLOGY OF CYCLOSTOMATA. 321
ugh the twelfth somite; we can find no marked difference
een the Anlage in the seventh somite and in this. The
1ents lying still farther backwards are not cut through exact-
ransversely in this same series of sections, owing to the cause
d above (pp. 312 and 315), so that we can not trace the dif-
ntiation of the mesoblast from the anterior to the posterior
in this one series. But I could demonstrate from several
r series of sections that the Anlage of the pronephros is, in
present stage, found in no less than 15 somites.
Figs. 9-11 represent the contiguous sections through the
rsomitic portion, on the right side, between the first and
nd Anlagen, 2.e. between that of the fourth, and that of the
, somite. Fig. 9 is from the section next behind that shown in
8; the portion (cd.) lying proximal to the lateral plate (/m.)
ents no longer a weak appearance as in younger embryos
the statement on p. 317 and figs. 4 and 7), but is occupied
2 compact cellular structure (cd.) which suddenly passes over
the loosely composed lateral plate (/m.). Fig. 10 is from the
ion next posterior to fig. 9 and next anterior to the second
age represented in fig. 11 and shows almost the same con-
m as in fig. 9, with respect to the structure in the proximal
ion of the lateral plate. In other words, in the intersomitic
ion between the first and second Anlagen, a cellular cord
become established, which connects these two Anlagen. It
nis cord which gives rise to the collecting duct or Sammelrohr
SUCKERT (88), putting all the pronephric tubules in communi-
on.
On the left side of figs. 9, 10, and 11, the contiguous sec-
s through the intersomitic portion between the Anlagen second
third, are represented. In figs. 9 and 11, the condition of
322 8. HATTA:
the structure (cd.) at the proximal portion of the lateral plate is
almost the same as that on tlıe right side just described, although
it is here somewhat weaker in development than there. The section
represented in fig. 10 intervenes between the two mentioned above;
in this section, the structure in question (cd.) is weakest in develop-
ment, consisting of four or five cellsonly. In the left half of fig. 15
which represents the section through the intersomitic portion be-
tween the sixth and seventh somites, there is found no structure
to be compared with the cord mentioned above, the proximal
edge of the lateral plate (x) being of the same condition as that
in figs. 4 and 7. Jn fact, the cord appears after the complete
separation of the Anlage from the myotome, and when il is first
established, the nearer the plane of a section to the Anlage either
anterior or posterior, the thicker the cord. For instance, of the
above three sections (the left side of Figs. 9-11), the middle
. (fig. 10) is the weakest. But this unequal development of the
cord is soon made even by its growth as seen in the case of the
cord between the Anlagen first and second (the right side of
figs. 9 and 10).
The history of this cord as given above shows that it has doubt-
less the genetic relation with the Anlage of the pronephros. In
early stages, no such structure is found in the intersomitic portion,
but it becomes established one after the other with the develop-
ment of the Anlagen. The cord is in section, thickest near
the Anlage and weakest in the midway between two consecutive
Anlagen, when it is first established. These facts give naturally
an impression that it is growing out of the two consecutive An-
lagen backwards and forwards and these two growing ends meet
at some point in the midway between these two Anlagen, finally
to fuse together. This point of meeting is, I think, indicated by
MORPHOLOGY OF CYCLOSTOMATA. 323
part where the duct has been described above as weakest.
s also the fact that repeated cell-multiplication takes place
he outer rim of each Anlage of the pronephros. One might
pose that the product of the cell-division would contribute
to the growth of the Anlage itself and has nothing to do
ı the cord; but this is not the case: the Anlage does not
y at the outer (lateral) end, as it might seem, but by cell-
sion within its own structure. I have never observed any
of cell-proliferation along the dorsal edge of the lateral
e in an intersomitic portion, although the cords appear, in
r stages, to have some connection with that edge, when they
fully established (see the right side of figs. 9 and 10); this
nection thus is not primary, but secondary. The epiblast has,
a the first, no share in the formation of the cord, always
wing a sharp contour against the mesoblast below.
There is thus no difficulty in accepting the view that the
vecting cord 1s formed of the intersomitic cell-outgrowths which
budded out of the anterior and posterior rims of each Anlage
he pronephros and are subsequently fused together. The cord
therefore, originally brought about by the confluence of the free
emilies of the Anlagen.
Further development of the Anlage of the pronephros may
intelligible by refering to fig. 18 which represents a sec-
through a little older embryo of Stage 11. The epiblast (ep.)
sists of a single layer of cubical cells as before ; the neural cord
is still solid. On the left side of the figure, the hind border
he fourth myotome (mi.IV) is cut, while on the right, the
-plane of the fifth myotome is met with. A comparison with
corresponding parts in the younger stages (figs. 3 and 8) will
324 8. HATTA:
plainly demonstrate a progressive change undergone by the
pronephric Anlage. The Anlage on the right side (a.pn.2)
presents a feature much like that seen in fig. 3, notwithstanding
some points of progress. The Anlage on the left side (a.pn.1),
however, shows a considerable progress ; it has become much more
compact by the active multiplication of its component cells.
Owing to mutual pressure, the cells are compressed and their
nuclei are regularly arranged, describing together an ellipsoidal
figure. The inside of the ellipse encloses a comparatively large
lumen, which is standing in connection with the body-cavity
represented, at the present stage, only by the boundary line of
the parietal and the viscera] layer of the lateral plate (Jm.).
In a little more advanced embryo, the cross-sections of which
are represented in figs. 20-31, the neural cord (n.), the myotomes
(mt.), and the Anlage of the pronephros show some progress as
compared with those described in the preceding pages. The epiblast
(ep.) consists, as in the embryo just described, of a single layer
of cubical cells and is limited by a sharp line against the struc-
tures below. The component cells of the neural cord” become
arranged in two layers, leaving, in the anterior section of the cord,
a vertical fissure-like lumen in the median line of the cord, which
represents the beginning of the central canal (figs. 20-21, &c.).
The posterior part of the cord is still solid, although the position of
the central canal is marked by a vertical line produced by cell-
boundaries (fig. 26) just as described in the foregoing pages. The
myotomes are, in the anterior region, likewise enlarged, probably
owing partly to multiplication of component cells and partly to
1) Owing tothe same cause as the sections represented in Figs. 2 and 3, the vertical
diameter of the neural cord in Figs. 20-23 is shown somewhat longer than it is in reality.
MORPHOLOGY OF CYCLOSTOMATA. 325
loosening of the composition of the tissue” and assume the
e of a scalene triangle (figs. 20-23) ; the median side of the
gle (mus.) represents the visceral, and the two other sides
) the parietal, layer of the myotome. In the posterior region,
are yet of a compact structure of a pentagonal form, en-
ng a cavity (figs. 25-31, mt.VII-X).
The anteriormost Anlage of the pronephros is found as before
r the hind part of the fourth myotome, the section of which
epresented in fig. 20 (apn.1). It shows a considerable
lopment: the component cells, which are of high columnar
acter are no longer compressed, but the tissue is more or less
ned. Thus the Anlage itself is distended, and its upper
sal) angle becomes acute and grows in between the epiblast
the myotome. The internal cavity of the Anlage also be-
28 conspicuous. The Anlage of the pronephros under the
, posterior myotome (the fifth) is not so advanced as in
last somite (the fourth). In fig. 22 is shown the section
ugh the hind part of this somite and of the pronephric An-
belonging to it (mt.V and a.pn.2), a section through the
-plane being unfortunately wanting in this series of sections.
next posterior Anlage is found just under the sixth myotome
represented in fig. 24 (a.pn.3) together with the hind border
he myotome (mt.VI). The Anlage shows a compact structure
ch is probably due to a rapid multiplication of the constitu-
cells. The next following Anlage of the pronephros is found
2ath the seventh myotome (fig. 26, a.pn.4). It shows no
her development than the separation of it from the myotome
the fusion at the retrenched ends of the two layers of
oblast: it is in the same stage of constriction as that in the
)See the foot note in p. 319.
326 Ss. HATTA:
right of fig. 13 which represents the Anlage third in a younger
embryo.
The Anlagen above referred to are connected with each other
by the solid connecting cords (figs. 21, 23, and 25, cd.), which
are found between these Anlagen just as described in the younger
stages. Of these connecting cords, that between the Anlagen
second and third (fig. 23, cd.) is the thickest, while that be-
tween the Anlagen third and fourth (fig. 25, cd.) is the weakest
in development, owing probably to its having been just established.
The cord between the Anlagen first and second (fig. 21, ed.)
however, is rather weaker as compared with that in the next
posterior intersomitic plane (fig. 23, cd.). Such a case is occa-
sionally met with; but this is doubtless not normal; in most
cases examined, the cord is thickest in anterior segment and de-
creases in thickness gradually posteriorly as seen in the last
example (see pp. 321-322 and figs. 9-11).
The Anlage of the pronephros belonging to the eighth so-
mite and that of the ninth somite are not completely cut off
from the myotome to which these respectively belong (figs. 28
and 30). They are, however, constricted already at the anterior
and posterior borders of the segment: fig. 27 shows the posterior
part of the Anlage fifth, and fig. 29 represents the anterior
part of the Anlage sixth respectively. Such a case is observed in
the younger stage described in the foregoing pages (pp. 319-320).
Compare these two figures with the right half of Fig. 14 and the
description on p. 319: here the phase of constriction is a little
more advanced than there. The anterior part of the Anlage fifth
and the posterior part of the Anlage sixth, the figures of which
are omitted, have features much like those seen in figs. 27 and 29.
In these two segments, the central portion of the Anlage is
MORPHOLOGY OF CYCLOSTOMATA. 327
in the process of being constricted off from the myotome, and we
not decide by this case alone which segment (whether the an-
yr or the posterior) is the further developed ; a comparative
y of other examples shows that the separation in the posterior
nent follows that in the anterior. The state of the mesoblast in
next posterior segment, 2.e., the tenth segment (fig. 31), is quite
rent from that just described ; it isin a more primitive condition
avelopment. The Anlage of the pronephros (a.sd.) presents only
indication of constriction,—a feature which we have observed
atedly in embryos of younger stages (compare with figs. 2, 5, 6,
and 16). From this segment backwards, a few segments show
st the same condition. Still further posteriorly, the structure
he mesoblast can not be readily observed, since the planes of
ons incline by degrees in the cranio-caudal direction, owing, as
re stated, to the bending of the longitudinal axis of the embryo.
In all the segments mentioned above, the lateral plate (dm.)
ists of a loose tissue of cells of variable shape, and the Anlage
es over suddenly into the lateral plate just as in the embryos
ribed in the foregoing pages.
In this stage, therefore, the Anlage of the pronephros is com-
ly separated from the myotome in 4 somites, i.e., from the
th to the seventh inclusive; and these are connected with one
her by the intersomitic solid cord. In the following 4 or 5
tes, the constriction is just going on, while in a few of still
2 posterior somites it 18 indicated merely by a slight depression
he parietal layer of the mesoblast.
Period 2.
In the embryos which belong to Stage 111, we observe a
ded advance in several respects. Figs. 32-50 represent a series
328 S. HATTA:
of cross-sections through one of these embryos which has about 25
somites. The neural cord (n.) is reduced in size and in the anterior
part has a conspicuous canal. The myotomes which showed
before a pentagonal outline, in the anterior part of the body have
now assumed an elongated lozenge-shape” and is composed of an
outer, and an inner, layer of long cells which have begun to dif-
fentiate themselves. The inner layer (mus.) represents the median
and ventral rows of the pentagonal myotome mentioned on p. 320
and, therefore, corresponds to the visceral layer of the myotome;
the outer layer (cut.) is the product of the dorsal and lateral layers
and constitutes the parietal layer. The pronephric Anlage is com-
posed of high columnar cells which are plainly distinguishable
. from the much shorter elements of the lateral plate (/m.). The
component cells of the Anlage of the pronephros which we generally
found to be compressed in the foregoing stage (pp. 324 and 325),
are now more or less loose, and the internal cavity of the Anlage
is somewhat widened, being distended by the loosening of the cells.”
The peritoneal cavity is, at the present stage, still repre-
sented merely by the boundary-line of the parietal and visceral
layers of the lateral plate.
In the present stage, the foremost Anlage of the pronephros
is, as before, found under the hind part of the fourth myotome
(figs. 32-35, a.pn.1). The Anlage shows, in section, a circular
outline and is composed of high columnar cells arranged in a
radial manner. The internal cavity of it is confined no longer
to one section, but it is observed in three or more sections; it
is most spacious in the hind part of the fourth somite (fig. 33)
or in that part where the cavity is visible from an early period.
1) A few: myotomes in the anterior somites tend to assume this shape already in the last
stage (see figs. 20, 21, and 22).
2) See the foot-note on p. 319.
MORPHOLOGY OF CYCLOSTOMATA. 329
m this part backwards it gradually decreases in width
1 no space is perceptible. Anteriorly the cavity is also some-
t narrowed, but not as much as in its posterior contin-
on, and ends blindly rather suddenly at its anterior end
32). The anterior portion of the Anlage forms a blunt conical
(Fig. 82, a.pn.1) projecting anteriorly and lying between the
al edge of the lateral plate (/m.) and the lower surface of
fourth myotome (mé./V). The existence of this conical
" gives us a strong impression that originally there must have
ı present an Anlage of the pronephros in the anterior segment
ch was connected by a connecting cord with the Anlage
nging to the fourth somite, but had disappeared during the
logeny and that this conical tube is the remnant of this
necting cord.”
The next posterior Anlage, which is found under the fifth
tome (figs. 37-39, a.pn.2) and shows an outline much re-
bling that represented in figs. 32-35, has an internal cavity
irregularly triangular form, extending through three sec-
s, of which the foremost section contains the most spacious
ty, while in the others the lumen grows smaller and smaller.
_ pronephric Anlage in the next following somite (figs. 41-43,
3) has an outline much like that shown on the left side of
s. 18 and 24, being in the same phase of development, that is,
3 of the form of an isosceles triangle whose two basal angles
sh the myotomes. This Anlage is found under the sixth
) The internal cavity of this conical tube is not entirely closed, but there is clearly seen
all canal (x) directed towards the median side and opening below the myotome. I can
ecide, at present, whether this canal is normal or abnormal; for I can not make out the
sponding structure on the opposite side and have no other embryo of exactly the same
‚ in which the structure in question would probably be found, if it be of some definite
ing; I also can not detect any trace of such a canal in embryvus of advanced or younger
s.
2) See the description under Period. 4.
330 S. HATTA:
myotome and contains the internal cavity extending likewise
for three sections, of which however the hindmost contains the
widest cavity, while it is diminished in width anteriorly : in other
words, the width of the cavity enlarges in inverse direction
as compared with that in the preceding two somites. The
Anlage fourth under the seventh myotome (figs. 45 and 46, a.pn.
4) has an oval outline like that shown in fig. 26 and encloses
an internal cavity, which covers two sections and is anteriorly
wide and posteriorly narrow. The two following Anlagen which
are detected under the eighth and ninth myotomes respectively
(fig. 48, a.pn.5 and fig. 50, a.pn.6) show almost the same con-
dition of development as in the somite just described; the
internal cavity which they contain is likewise extended into two
sections ; the width of the cavity is about the same in these two
sections, being of a fissure-like form. |
The solid cord which is observed in the embryos of the last
stage connecting the Anlagen with one another, is also found
here. The cord in the intersomitic plane between the Anlagen
first and second (fig. 36, cd.), that between the Anlagen second
and third (fig. 40, cd.), and that between the Anlagen third and
fourth (fig. 44, cd.) are all comparatively short, so that they
are in each stretch confined to only one section, while that in
the two posterior intersomitic planes, 2.e. between the Anlagen
fourth and fifth, and between the Anlagen fifth and sixth, the
cord is extended in each case into four sections. In this latter
part, the cord is in a primitive condition ; the component cells
are actively multiplying. Hence these four sections all show
similar features. I have endeavoured to show in fig. 47 one of
these sections which is taken from one of the four sections between
MORPHOLOGY OF CYCLOSTOMATA. 331
Anlagen fourth and fifth, and in fig. 49, one between the
agen fifth and sixth.
This inequality in the length of the intersomitic solid cord
believe, due to differences in the degree to which the canali-
n within the Anlage has extended into the connecting cord.
he anterior section of the pronephros, this process has already
eeded to some extent into the interior of this cord, while in
posterior, the cavity is still confined entirely within the
age itself. The whole system of the pronephros at the present
ition may be compared to a bamboo-cane with nodes and
nodes ; in the anterior section of the system, the nodal septum
become very thin, while it has a considerable thickness
he the posterior, As will be shown further on, all these
a entirely disappear later when the collecting duct is fully
lished.
From the fact mentioned above, it will be easily seen that the
ess of canalization in the pronephric system of Petromyzon
ns in the internal cavity of the pronephric Anlage in each
rent and is extended into the intersomitic connecting cord.
direction in which this process proceeds seems, generally
king, to be from the anterior section to the posterior; for
nost cases, not only the internal cavity in each Anlage is
ious anteriorly and narrowed posteriorly, but the cavity in
rior somites is extended more, or canalization goes on further,
1 in the posterior section of the system; although the pro-
s in the opposite direction is occasionally met with.
From the tenth somite backwards, five or six segments show
same condition of the mesoblast as in the eighth and ninth
ites, after which the series can not be studied, owing to the
ination of the planes of sections, referred to above.
332 8. HATTA:
In all the segments above referred to, the lateral plate of the
mesoblast shows the same condition as in the foregoing stages, but
has become more distinct from the Anlage of the pronephros.
In the present stage of development, then, the Anlage of the
pronephros 1s cut off from the myotome in more than 10 segmenis,
and the canalization has advanced in the anterior section of the
system, lo a state just ready to put the Anlagen in the succeeding
somites in communication with one another, although the inter
somilic connecting duct in the posterior part remains still solid.
Figs. 51-58 were drawn from a series of sections through
one of the older embryos in this stage. The internal structures are
developed much more than in the embryo just described. The
cells forming the visceral layer of the myotome have been differen-
tiated into the muscle-plates, while the parietal layer is composed
of cubical cells. The Anlagen of the pronephros have acquired,
in most cases, a tubular structure and have grown dorsally, being
folded out from the body-cavity ; I will accordingly call them the
pronephric tubules.
On the right side of fig. 51, the foremost tubule (pé. 1) is
visible, which no longer contains the internal cavity but is
converted into a solid mass of cells occupying the space beneath
the fourth myotome. This consolidation is not due to retrogres-
sive changes, but is effected by very active cell-multiplication
which takes place within the tissue. The cross-section of the
collecting duct seen on the left side of fig. 52 (cd.) which re-
presents the third section behind the last, is likewise solid. The
tubule on the right side of this figure (pé.2) and that on the
left side of the third section posterior to it (fig. 53, pé.2) are
respectively the second tubule of the right and left side found
MORPHOLOGY OF CYCLOSTOMATA. 333
under the fifth myotome ; both are of a triangular form and
contain a very spacious internal cavity of the same shape. On
the right side of fig. 53, the sixth myotome and the third tubule
are shown. The section next posterior to fig. 53 (fig. 54) shows
the cross-section of the collecting duct (ed.) on the right side and
a slice of the hind wall of the second tubule on the left (pé.2).
The cells composing the duct are closely set together, although
arranged more or less radially, acquiring a tubular form. As
has been repeatedly mentioned above, the epiblast is, as in
the foregoing stage, marked off from the mesoblast as well as from
the Anlage ; but at the present stage, the second tubule (figs. 54,
pt.2) pushes against the epiblast, probably in consequence of an
enormous multiplication of its component cells, so as to cause
the latter to be a little elevated externally. It must be remarked
here that the Anlagen, especially the first and the second, when
they first assume the tubular form, are brought into an intimate
relation with the epiblast, striking against it. In some of
my sections, a mitotic figure is seen at that point of the epiblast”
(fig. 54, x). This might lead some to assume a genetic connection
between the epiblast and the pronephros in Petronyzon ; but there
is, I believe, in reality no such relation. Ifthe epiblast gives some.
cells destined to build the pronephros or a part of it, cell-prolifera-
tion or some other mode of cell-production would necessarily be
observed in the epiblast in the preceding stages or at least, in
the stage here spoken of. In the foregoing stages, the epiblast
had, as has been repeatedly mentioned above, a sharp limit against
the structures inside it. At the present stages also, it is marked off
by the boundary-line of cells from the tissue of the tubule,
1) In the series of sections, from which fig. 54 is drawn, I observe mitotic figures at
that point in several sections.
334 8. HATTA:
showing no structural alteration. Mitotic figures are ı
not infrequently in that part of the epiblast (fig. 54, x
axis lies, however, in all the cases examined parallel
plane of the epiblast, giving us an impression of the :
cells contributing to the formation of no other part than
blast itself; on the contrary, within the structure of th
the cells are rapidly multiplying (figs. 51, pt.1 and
pt.2), showing that the growth of the tubule is active
on. In fact, the connection, or rather the intimate co
the pronephric tubule with the epiblast is a tempora
dition ; the separation follows immediately afterwards,
tubule returns soon into a state similar to that seen it
(pt. 2).
According to Rickert (’88), a similar case is |
in Selachian embryos: the tubules become connected sec
with the epiblast—what caused him to believe that the latt
give some constituent elements to the tubules.
The third section behind that represented in fig. 54
shows, on the right side, the fourth (pé.4) and, on the left, ı
tubule (p£.3) respectively. The latter is not so far d
as its counterpart on the opposite side (fig. 53, pé.3), w
former presents a great progress: it consists of a definite
ium and contains a distinct cavity of triangular shape, :
the corresponding tubule on the opposite side (fig. é
which is found in the third section behind the last, is n
advanced in development. The fifth tubule, the tubul
right side of fig. 56 (pt.5), is somewhat more developed t
which belongs to the anterior somite (the fourth tubule
opposite side) ; but it has a feature much resembling th
tubule on the same side (fig. 55, pt.4) and the second
MORPHOLOGY OF CYCLOSTOMATA. 335
posite side (fig. 53, pé.2). In short, in this series of sections,
_tubules on the right side, are all more advanced than those
the opposite side. The sixth is very primitive in development ;
57 represents the section, on the left side, through the anterior
t of the ninth somite and, on the right, the posterior part of
The left tubule is sliced at its anterior wall, but the right
ule is cut through in its mid-plane. It is composed of two
ers of columnar cells, but no cavity has yet appeared in
- interior.
From the tenth somite backwards, the Anlagen are cut off
m both the myotomes and the lateral plate, and constitute the
mental duct or the posterior continuation of the collecting
tt, which is distinctly traceable for 7-8 somites. Not infre-
ntly, however, a somite is met with, in which the segmental
+ is not yet cut off from the lateral plate at the time when
separation is finished in a majority of somites, as seen in
58 which represents a section through the twelfth somite.
e left half of the figure shows the duct entirely cut off from
lateral plate, while the right exhibits the state not yet
arated. The same structure is made out in two contiguous
tions, so that one might mistake it for a pronephric tubule.
is point will be described further on.
The relation of the pronephric tubule and the peritoneal
‘ity is not so simple as in the last specimen; besides the
ynephic tubule, there is seen another structure which projects
; of the inner angle of the peritoneal cavity (figs. 52, 53, 55,
156, c.p.). This projection is originally a fold of the peritoneal
ll and gives rise, as subsequent history shows, to the radix of the
sentery, whence the gonads and the mesonephric tubules are
‘ived. It will here be called briefly the “ coelomic projection.”
336 Ss. HATTA:
At that point of the visceral layer of the mesoblast, where the
Anlage of the pronephric tubule passes over to the lateral plate,
it is always many cells deep (figs. 55 and 56, ¢.p.), and the pro-
jection in question is brought about by repeated division of these
cells. The projection formed is consequently seen in each somite
and thus shows a segmental arrangement. Its component cells
are soon re-arranged into an epithelium, and the pouches thus
formed push their way between the myotome and the hypoblast.
The coelomic projection appears, at first sight, to be homo-
logous with the coelomic pocket described by Price (’97) in
Bdellostoma. The coelomic pocket is, however, according to Price,
the product of both the parietal and visceral layers of the lateral
plate and is afterwards converted into the cavity between the
glomerulus and Bowman’s capsule of the Malpighian cor-
puscle; the floor of the pocket forms Bowman’s capsule,
and its roof together with a part of the pronephric tubule is
transformed into the cover of the glomerulus (’97, p. 213). The
coelomic projection in Petromyzon is, on the contrary, formed
out of the visceral layer of the distal half of the somite and gives
rise, as just stated, to the radix of the mesentery, from which
partly the mesonephric tubule and partly the gonads are formed.”
Figs. 59-63 are from a series of sections through an older
embryo of the same stage. In this series of sections, a further
development of the coelomic projections is clearly seen ; the first
figure (fig. 59) shows the section through the second tubule, fig.
60 through the third, and so forth. In the first 3 figures and
on the right of fig. 62, the coelomic projection (c.p.) presents an
1)I will not here further discuss this structure, as I intend to do so in a future paper
in which the development of the mesonephros in J’eromyzon will be delt with.
MORPHOLOGY OF CYCLOSTOMATA. 337
ithelial structure, forming the continuation of the peritoneum
id folding out from the peritoneal cavity. Beneath the first
bule, there is found no rudiment of the projection ; under the
cond (fig. 59) it is very weak, while beneath the third (fig. 60),
urth (fig. 61), and fifth (fig. 62), tubule, respectively it is most
gorously developed. But on the left side of figs. 61 and 62 it
again in a primitive condition, just as in the last series of
ctions (figs. 52, 53, 55, and 56).
The coelomic projections are not confined to the anterior region
here the pronephric tubules are found, but it is found likewise
the posterior part where only the segmental duct develops.
ig. 63 shows the section through the thirteenth somite ; on this
ction, the duct is cut off from the myotome and a well developed
elomic projection (c.p.) is observed; I will return once more
this subject further on.
Leaving the coelomic projection in this stage of development,
will return to the origin of the Aulage of the pronephros and
ve somewhat more exact details on the subject. Since the piece
the mesoblast called above the Anlage of the pronephros forms
r a time the proximal portion of the lateral plate, one might
esume that its whole mass will be transformed into the pro-
;phrie tubule and will not partake in the formation of the perito-
al membrane. I was at first of this opinion, but a careful
servation of sections through the embryos in each stage showed
y error.
To illustrate this point satisfactorily, I have given, in the
inexed wood-cut (Wood-cut 1), a series of semi-diagramatic
rures, which show the successive phases of the changes going
1 in the structure. A shows the first indication of the Anlage
the pronephros before the separation of it from the myotome ;
338
a-b indicates the extent of the Anlage; c-d shows that of the
coelomic projection.
called the coelomic projection.
%K E
er ” @e
5 El |F
en
|
à
Wood-cut 1.—Semidiagramatic figures to
illustrate the successive phases of the
evolution of the nephrotome.
. from the right side of fig. 16.
. from fig. 3.
from the right side fig. 18.
. from the left side of the same.
from the left side of fig. 53.
from the right side of fig. 55.
. from the left side of fig. 61.
When the myotome is cut off, the point
of the parietal layer indicated
by a becomes fused with the
point « of the visceral layer
(B, ac). This piece of the meso-
blast assumes an ellipsoidal shape
(C). The component cells of
this ellipsoid are multiplied by
active cell-divisions, and the
piece almost loses its lumen and
gets a compact consistence (D).
Meanwhile the cells in the space
d-c remain inactive. Conse-
quently the piece acquires a tn-
angular form (E ), whose upper
sharp angle, together with the two
sides enclosing this angle, gives
rise to the pronephric tubule.
The lower (median) obtuse angle
now begins to grow by cell-
mulplication (Z') and produces
a small knob (F, c.p.), which
grows further and pushes in be-
tween the myotome and the
hypoblast (the upper wall of the
enteric canal). This cellular
projection is that which has been
It is reduced into a thin
plate of epithelial cells (G, c.p.) and assumes then the form of
MORPHOLOGY OF CYCLOSTOMATA. 339
a true fold of the visceral layer of the lateral plate. At the
same time, the upper angle or the pronephric angle develops
further and assumes a tubular form composed of a single layer
of columnar cells (@).
The peritoneal cavity begins, therefore, at the point, from
which the coelomic projection starts, and the part of the layer
dorsal to this point is all appropriated to the formation of the
pronephric tubule. The nephrostome will be found, therefore, by
the point where the tubule passes over to the projection.
I will add a few words on the differentiation of the myo-
tome, so far as concerns the topographical relation of it to the
Anlage of the pronephros. The myotome consists, at the present
stage (Stage 111), of the inner and outer layers which constitute
respectively the Muskelblatt and the Cutisdblatt of German
authors (fig. 59 and 60, mus. and cut.). The cells composing the
Muskelblatt (mus.) are, simply differentiated into a transverse row
of the muscle-plates. The outer layer (cué.) undergoes, however,
subsequently a series of interesting changes: it folds in, just as
the Sklerablatt or sclerotome described by HATCHEK in Amphioxrus
(88) between the Muskelblatt and the chorda and the neural tube.’
As is well known, Rast (’88) has homologised HatcHeEK’s Sklerab-
latt with his Sclerotomdivertikel of Selachian embryos, which is
the evagination of the ventral part of the visceral layer of the meso-
blastic somite. This part of the somite (the selerotome) corres-
ponds, I believe, exactly to the ventral row of the pentagonal
myotome in my embryo (see pp. 314 and 320), which comes after-
wards to form the ventral part of the cutis-layer (see figs. 21, 22,
23, 36, 37, 43, 49, 59, 60, &c.). When the myotome is not yet
separated from the rest. of the mesoblast (fig. 2), this part of the
1) This subject will be treated of in an independent article.
340 8. HATTA:
selerotomic layer forms, as has been seen above, a direct continua-
tion of the visceral layer giving rise to the coelomic projection.
(The successive changes of the myotome are seen in figs. 1, 2, 3,
18, 21, 22, 43, 49, 60, &c.)
From the above account, it can be inferred that the ventral half
of the mesoblastic somite in Petromyzon, which gives rise to the
pronephic Anlage and the coelomic projection, is doubtless homo-
logous with the “ intermediale cell-mass’’ of BaLFour described by
him in Selachia and, therefore exactly coincides with the ‘‘ Nephro-
tom ”” of Rtckert.” So far as concerns its future destination, how-
ever, the results arrived at by me slightly deviate from their views.
A series of sections through the oldest embryo of this stage
is represented in figs. 64-76. The epiblast has undergone no
histological change, but remains, as before, one cell deep. Many
structures, however, exhibit a remarkable progress. The muscle-
layer (mus.) of the myotome is, for instance, further differentiated,
now consisting of a transverse row of long muscle-cells, although
the cutis-layer (cué.) is still composed of short cubical cells. In the
anterior region, the true coelome (pp.c), becomes conspicuous en-
closed by the parietal (».p.) and the visceral (m.v.) layers of the
lateral plate, both of which consist of a single row of cubical cells.
The ventral edges of the lateral plates on both sides do not, however,
yet meet in the ventral median line. The walls of the enteric
canal too are, in the anterior region, reduced into a single cell layer.
A great alteration is met with in the pronephric tubules.
They have assumed a cylindrical fornı composed of tall columnar
epithelium and have grown dorsally, pushing in between the
myotome and the epiblast, causing the latter to be elevated a
1) In spite of the discussion by RÜCKERT (’89, pp. 19-20) on the inexactness of the expres-
sion “intermediate cell-mass,” I homologise, with many authors, these two terms with each other.
MORPHOLOGY OF CYCLOSTOMATA. 341
little. The internal lumen of the tubules are put not only in
wide communication with the peritoneal cavity, but also in direct
continuation with one another through the collecting duct, which
consists of a regular columnar epithelium-cells arranged radially
and now encloses a conspicuous lumen.
In the foremost of these twelve sections (fig. 64), we notice that
a structure (pt.1) consisting of a few cells projects at the outer
corner of the proximal edge of the lateral plates and lies in contact
with the outer wall of the myotome on either side. This structure is
found under the anterior border of the fifth myotome and I infer
that it is a remnant of the first pair of the pronephric tubules which
begins to decline in the present stage. ‘The reason why it is found
not under the fourth myotome as in all the stages hitherto des-
cribed but beneath the anterior border of the fifth myotome,
consists probably in its shifting backwards; for we find, in this
series of sections, another pair of the tubules under this same fifth
myotome. A comparison of this figure with fig. 65 representing
the next posterior section will make the matter clear. On the
left side of fig. 65, the same remnant structure (p/.1) together with
the collecting duct (cd.), which connects the first and the second
tubules can be observed, while the cross-section of the collecting
duct in the corresponding intersomitic plane is seen on the op-
posite side (cd.). : The next following section is shown in fig. 66 ;
the tubules (pt.2) on both sides communicate freely with the
peritoneal cavity ; these are found beneath the hind part of the
fifth myotome and are the second pair of the tubules; the open-
ings (nst.2) to the peritoneal cavity are, therefore, the second
nephrostome of the pronephros. The tubule on the left side is
weaker than that on the right, since a larger part of the left tubule
is visible on the section next posterior which is represented in fig.
342 Ss. HATTA:
67 (pt.2). The shape, which the tubules of the second pair (fig. 66,
pt.2) assume at about this stage, is a characteristic triangle,
whose two angles, the one directed dorsally and the other directed
medially, are acute and whose outer (lateral) angle is obtuse
(see the left side of fig. 59 and the right side of fig. 66, pl.2); so
much so that we can easily determine by this feature the fact
of their being the second pair. This peculiar shape of the second
tubule is retained for a considerable time as will be seen further
on. On the right side of fig. 67, the collecting duct (cd.) is cut
through transversely ; on the left, the same duct (ed.) and the
hind part of the second tubule (pt. 2) are seen.
At the point where the nephrostome opens to the peritoneal
cavity, the visceral peritoneum at the median corner of the latter
projects out between the myotome and the hypoblast (figs. 66 and
67, c.p.) ; beneath the collecting duct, however, no such structure
is detected (see the right side of figs. 65 and 67). Such a pouch
is repeated in each nephrotome (see figs. 66-75, c.p.) and is what
has been called above the coelomic projection.”
The next following sections shown in figs. 68 and 69 show
the third pair of the tubules (pf.3) to be of the same structure.
In these two sections the tubules are cut through lengthwise, and
the nephrostomes (nst.3) on the two sides come into view in syn-
metrical manner. ‘The tubules are so simple as to need no further
explanation. Fig. 70 represents a section through the intersomitic
plane between the sixth and the seventh somites, and next posterior
to fig. 69. It shows on either side the cross-section of only the
collecting duct (cd.), consisting of radially arranged cells. Fig.
71 is from a section through the seventh somite and is the third
1) In this series of sections, we often see the coelomic projection on sections passing
throngh the intersomitic plane; but this is the piece of it belonging to either the anterior or
the posterior somite.
MORPHOLOGY OF CYCLOSTOMATA. 343
behind the section shown in fig. 70; the tubules of the fourth
pair (pt.4) show themselves symmetrically on both sides; they
are somewhat less developed as compared with those of the last pair.
Fig. 72 is the section next behind fig. 71 ; it shows on both sides
the collecting duct (cd.) together with the coelomic projection (c.p.)
which is a part of that of the anterior segment. Fig. 73 is the third
section posterior to that just described ; the blastoderm becomes
more flattened than in the foregoing sections ; it shows on both sides
the tubules of the fifth pair (pé.5); the condition of the tubules
and nephrostomes (nsé.5) is much like that in fig. 71. Fig. 74
is from the fourth section posterior to fig. 73; the right tubule
(pt.6) of the sixth pair and its nephrostome (nsi.6) are visible on
the right side, while the collecting duct (ed.) is. cut through on the
left. The sixth tubule and nephrostome on the opposite side are
observed in the next anterior section which is not figured. The
segments back of the ninth somite have no trace of the tubule, but
the cross-sections of the posterior continuation of the collecting
duct, the segmental duct, are repeated in each section. Fig. 75
represents a section through the sixteenth somite; the cross-
section of the segmental duct (sd.) on either side is seen; it
always occupies the space where, in the anterior region, the
tubules or the collecting duct is found.
This condition, however, is not continued to the dorsal lip
of the blastopore. As I have stated in my previous paper (’91),
many processes of development are much delayed in the hind region,
so that we are here reminded of what were seen in the anterior
region of the younger stages. Fig. 76 represents the fifteenth sec-
tion from the dorsal lip of the blastopore and passes through about
the twenty-third somite. In this comparatively late stage, in
which many mesoblastic organs have developed in the anterior
344 Ss. HATTA:
region, the neural cord (n.) is still solid; the mesoblast (ms.) is
many-cell-layered and its metameric segmentation is still going on.
On the right side of the figure, tlıe section passes through the
mid-plane of the myotome showing no sign of its separation
from the rest of the mesoblast, while on the left, which shows the
intersomitic portion, the process of separation (mi. and /m.) is going
on. On both sides, however, there is no structure that can be
recognised as the Anlage of the segmental duct.
The six pairs of pronephric tubules observed in this stage
are the maximum number for Petromyzon ; this stage ought, there-
fore, to be regarded as the highest point of development with reference
to the pronephros. Even in the present stage, the foremost tubules
show a tendency to degenerate.
Period 3.
The embryos of Stage tv, which have about 35 mesoblastic
somites, present a remarkable progress. The head-fold is much
prolonged ; in older embryos of this stage, it begins to twist
(97, fig. 1, D). Figs. 77-91 represent sections through one of
these embryos. In some myotomes, the sclerotomic fold goes
deeper between the muscle-layer and the chorda. The parietal
layer of the lateral plate (m.p.) is much lessened in thickness,
so that it is reduced, in the dorsal region (the posterior two thirds
of the pronephric extent), into a thin epithelial lining of the
body wall (see figs. 79-85). The coelomic projection is likewise
reduced into a thin plate (figs. 82-86, c.p.) except in the anterior
two segments of the pronephic region, in which it still keeps the
characters of the younger stages (figs. 77-81, c.p.), only folding in
deeper than in the foregoing stages. The visceral layer (m.v.) of the
MORPHOLOGY OF CYCLOSTOMATA. 345
peritoneum still consists of a cubical or rather cylindrical epithe-
lium. The pronephric tubules are, in general, much prolonged
and begin to coil in the dorso-lateral direction, so as to cause an
elevation in the epiblast. The walls of the tubules consist of a
regular row of cylindrical cells, which passes over suddenly into
the thin peritoneum (figs. 77-86), except in the region of the second
pair of the tubules, where the parietal layer (m.p.) of the lateral
plate still retains the character of the younger stages, being
composed of cylindrical epithelium like the tubules: themselves
(figs. 77-79). At some regions, even a few mesenchyma-cells (nch.)
appear,—for instance, beneath the chorda (see figs. 80, 82, and 84),
in the median ventral space (see figs. 81 and 82), and also inside
the lateral epiblast (see figs. 77 and 80).
Fig. 77 shows a section through the fifth somite and
therefore corresponds to fig. 66, which represents the section
through the same plane of an embryo at a younger stage. The
longitudinal section of the second tubule (pé.2), together with the
corresponding nephrostome (nsi.2), is seen on the left side of the
figure, greatly resembling the tubules of the same pair in the
younger stage (compare with fig. 66). On the right side, the
nephrostome (nst.2) alone is observed; the tubule proper is to
be seen in the two following sections which are represented in
figs. 78 and 79.
Beneath the myotome anterior to the one just described,
there is found neither a tubule nor any structure that may be
regarded as the remnant of it. In the space between the epiblast,
the myotome and the lateral plate, however, a few scattered
cells (fig. 77, mech.) are found. I at first supposed that these
might be disconnected component cells of the first pair of
tubules; but, as free cells of quite the same character are found
346 8. HATTA :
in other places, for instance, in the space between the lateral
plate and the epiblast (figs. 79-81, mch.), I have been compelled
to conclude that they have no genetic relation with the pronephros,
but are mesenchymatous cells which are destined to form the
blood-vessels and corpuscles.
As the embryo was somewhat twisted, the sections did not
pass through the lateral walls of the body in an exactly trans-
verse plane, but unavoidably obliquely, on either side, as the
continuous serial sections represented in figs. 79-86 show.
While on the left side of fig. 78 the posterior portion of the
second tubule (pé.2) is seen, the second nephrostome (nst.2) is ob-
served on the right together with a cross-section of a tubular struct-
ure (cd.). This latter might be taken as a slice of the anterior
border of the second tubule, but is, in my opinion, the remnant of
the collecting duct which once connected the second tubule with the
first and forms, at present, a tubercle in front of the second tubule,
the first tubule having disappeared ; for the second tubule on that
side is observable in the next following section represented in
fig. 79 (pt.2) showing its characteristic features stated above
(p. 342).
On the left side of fig. 79 and on the right side of fig. 81,
we see the anterior half of the third tubule (pt.3) and nephro-
stome (nst.3) of each side, their respective posterior half being
found on the left side of fig. 80 and on the right side of fig.
82 (pi.3 and nst.3); the tubules are bent laterally and dorsally,
probably caused by the prolongation of their tubular portion ; for
their nephrostomal part and dorsal blind end retain their original
position. This is the first step in the convolution of the pro-
nephric tubule.
As seen on the left side of figs. 79 and 80, the tubule (p£.3)
MORPHOLOGY OF CYCLOSTOMATA. 347
he third pair shows a new character: the dorsal blind end
the nephrostomal portion of the tubule are more or less expand-
while these two portions are united by a slender middle trunk.
n compared with the tubule of the same pair on the opposite
represented in figs. 81 and 82 (pt.3), this character of the
le will be understood more clearly : the dorsal expansion is seen
>. 81, while the nephrostomal widening is observed in fig. 82.
tubules of the following two pairs show the same feature.
the right side of fig. 80, only the collecting duct between the
nd and third tubules is found. The left tubule of the fourth
is shown on the left side of figs. 81 and 82 (pé.4); fig. 83
's a cross-section through the collecting duct (cd.) between the
land fourth tubules and a slice of the anterior wall of the
t tubule (pé.4) of the fourth pair which is prolonged and bent
the tubules of the last pair. The fifth pair of tubules is seen
he left half of fig. 84 on one side (p{.5) and on the right
g. 85 on the other (pé.5). It is not developed as much as
more anterior pairs, but shows considerable progress as com-
d with the tubules in fig. 73 which represents the younger
> of the same pair.
These four pairs of the tubules (from the second to the fifth)
ain a spacious lumen and stand in wide communication
the peritoneal cavity, which becomes, at the present stage,
picuous from this region forwards.
In fig. 86 which shows the fifth section behind the section
m in fig. 85, the space on the left side which is occupied,
he more anterior region, by the tubule or the collecting duct,
placed by the cross-section of a duct (sd.) with an oval out-
and an ovoid lumen. This is the segmental duct under the
1 myotome (mi.X). On the right side of the figure, however,
348 S. HATTA:
besides the cross-section of a duct (cd.), there is seen a pronephric
tubule (p£.6), the long axis of which is directed vertically to
the inner surface of the epiblast. It is found just beneath the
ninth myotome where the sixth tubule should be found. Al-
though the parts of it are also to be seen in two consecutive sections
(the one represented in fig. 86 and another preceding it),
the communication of its lumen with the collecting duct is not
to be found anywhere. In some embryos, the tubule loses the
connection with both the body-cavity and the duct. The structure
(pt.6) in question is, I believe, nothing else than the remnant ofthe
sixth tubule which is in a stage of degeneration, and the duct (cd.) is
doubtless the collecting duct between tlıe tubules of the fifth and
sixth pairs. Compare the segmental duct (sd.) on the left side with
the collecting duct (cd.) on the right side just described ; the latter
has a wide circular lumen, whilst in the former it is slender and
compressed. This difference of character between these two
ducts is noticeable for some time in the younger stages.
To sum up the results obtained in this stage the tubules of the
third to the fifth pairs are vigorously developed, while the second
is very weak, the sixth retrograding, and the first has entirely
disappeared.
In the present stage, a peculiar structure is observed inside
the walls of the body-cavity (figs. 77-85, pp.1-3). At some points
of the peritoneum, a thin plate which consists, in cross-section, of
one, two, or three cells, projects from the peritoneal wall into the
body-cavity ; it will be called here shortly the “ peritoneal partı-
tion.” A peritoneal outgrowth is found at the level where the
coelomic projection passes over to the visceral layer of the lateral
plate ; at the same level, another peritoneal outgrowth starts up out
of the parietal layer. These two outgrowths meet at midway
MORPHOLOGY OF CYCLOSTOMATA. 349
and cut off a long chamber from the body-cavity along the
openings of the nephrostomes (figs. 77-85, pp.1). This longitudi-
nal chamber communicates anteriorly as well as posteriorly with
the body-cavity below, which is represented, in those parts, by the
boundary of the parietal and visceral layers of the lateral plate.
This is the uppermost partition. The second partition is weaker
in development and is detected a little more ventrally, projecting
likewise from the parietal and the visceral layers of the lateral
plate. It is most obvious in the region beneath the third and
fourth nephrostomes (81-84, pp.2). We find the third partition
still more ventrally, which is weakest in development ; its extent
is almost the same as the second (figs. 81-84, pp.3).
These partitions disappear after a short existence; in a little
older embryo, none of them is detected, as will further be seen.
This is probably the same structure as the “ peritoneale Scheide-
wände ”’ or ‘‘ Peritonealbrücke ” described by GoETTE in Petromyzon
fluviatilis (90). As to the meaning of the structure I have
nothing to say.”
It is important here to illustrate the topographical position
of the pronephros and the relation of it to other parts; for these
become definite for the first time in the present stage. For this
purpose, a series of sagittal sections is instructive (figs. 112-114).”
A few anterior myotomes (m£.II-V) are seen in fig. 112, which
represents the section nearest the median line. In the posterior
part of these myotomes, four cell-layers are distinguishable; the
outmost layer (ep.) is the epiblast ; the cell cord (cd.) inside the
epiblast is the longitudinal section of the collecting duct, and
1)See the historical review under Peronyzon.
2)The embryo, from which these figures are drawn, is a little younger than that just
spoken of.
350 S. HATTA:
its caudal continuation (sd.) is the segmental duct ; while the inner
two layers (m.v. and m.p.) present respectively the parietal and
visceral layers of the lateral plate. Below these structures, the
roof of the enteric canal covers the fore-gut (/g.) and the com-
mencement of the mid-gut which forms the passage of the enteric
cavity from the anterior slender portion to the posterior wide
cavity. In fig. 113 which represents the section next outside the
last, the lateral walls of the five myotomes from the first to the
fifth (mé.J-V’) are noticed ; the cell-maes (au.) seen next anteriorly
to the first myotome (mt. 1) is a slice of the wall of the left auditory
pit. The cross-sections of the pronephric tubules from the second
to the fourth (pé.2-4) follow immediately behind the fifth
myotome ; an oblique section of the fifth tubule (p£.5) and the
nephrostomal part of the sixth tubule (pt.6) are also obvious
behind the fourth tubule. The nephrostomes of the second
(nst.2) and fifth (nsé.5) tubules are seen only in part, while a
larger part of the third, fifth and sixth nephrostomes is seen in
the next figure (fig. 114, nst.3, 5, and 6) which represents a
section still further outside. The sixth tubule, which is of weak
development and has a wide nephrostome, is visible in these two
sections (figs. 113 and 114, p4.6 and nst.6). Except the first tubule
which has already disappeared without leaving any trace, the five
tubules are all thus seen in the dorso-lateral aspect of the hind
section of the fore-gut and the commencement of the mid-gut.
The pronephros is thus situated in the neck which connects
the head-protuberance with the globular abdominal portion.
Below the pronephros, the narrow passage of the fore-gut passes
through to unite the fore-gut with the wide space of the mid-gut,
where afterwards the liver (/.) is found. Underneath the passage
of the fore-gut, a group of mesenchymatous cells (mch.), which
MORPHOLOGY OF CYCLOSTOMATA. 351
nstitutes the earliest fundament of the heart, is detected. The
sent position of the pronephros,—dorsal to the heart, anterior
d dorsal to the liver, and along either side of the chorda,—
retained by it for a comparatively long period (see fig. 97);
later stages, the liver is somewhat shifted backwards, so
it the pronephros now comes entirely in front of it (see
. 115).
In an older embryo of this stage (figs. 92-96) the median
ds of the coelomic projection, the component cells of which are
ry much flattened out, go in deeper towards the median line
meet with its counterpart on the opposite side. The second
mule (figs. 92 and 93, pt.2) has become weaker, as a com-
rison of these figures with figs. 77 and 79 will show. On the
ıtrary, the tubule of the next pair (fig. 94, pé.3) has much
ngated and is bent considerably in dorso-lateral direction,
that we can no longer observe the nephrostome together with
» tubule itself on the same section. The following tubule,
> fourth (fig. 95, pé.4), is likewise well developed; the fifth
y. 96, pt.5) is more or less weak in development as compared
th the tubules of the two foregoing pairs. In short, these
ree pairs (from the third to the fifth) make parallel pro-
ss with the development of other structures, for instance, the
senterial fold or the muscle-segments. This is a fact that
to be observed too in the younger embryo of this stage, as
ove described. At the present stage, we can find no trace of
2 tubules beneath the ninth myotome, where the tubules of the
th pair ought to be found, but only the cross-sections of the
llecting duct or the anteriormost part of the segmental duct
3 seen.
352 S. HATTA:
Thus, the tubules of the third, fourth, and fifth pairs con-
tinue to grow, while the first pair has disappeared in the early
part of the present stage (or at the end of the foregoing stage) ;
the sizth has already commenced retrogression and the second 18
also growing weaker and weaker.
In the oldest embryo of this stage, there is to be seen no
marked change in the pronephros, but the peritoneal lining is
reduced into a very thin plate of a definite epithelium every-
where except at the pericardial portion, where the cells still have
a columnar shape. The mesenchymatous cells accumulated on
the median ventral line of the body are arranged in a certain
order to be transformed into the cardiac tube. The third, fourth,
and fifth tubules are also markedly prolonged and project into
the body-cavity so as to cause the parietal layer of the perito-
neum to fold between the epiblast and the body of the tubule
(see this Journal: vol. x, Pt. xvııı, figs. 8, 9, and 10). In some
of the embryos, the tubules of the second pair undergo degenera-
tion. I have met with, in this series of sections, the same con-
dition of the sixth tubule as on the right of fig. 86,.the right
tubule having entirely disappeared.
Period 4.
The embryos in the next advanced stage (Stage v) are much
diminished in size, assuming a form of a retort or of a pistol
(97, fig. 1, #). Figs. 98-106 represent sections through an
embryo of this stage. The posterior larger section of the fore-
gut comprising the pronephric region, has been reduced into 4
slender tube (/g.) which is bounded by almost a single layer of
high cylindrical cells. The parietal layer of the peritoneum as
MORPHOLOGY OF CYCLOSTOMATA. 353
well as the coelomic projection (7...) is very much decreased in
thickness and encloses the peritoneal cavity (pp.c.) that has now
become spacious, while the visceral peritoneum is still thicker than
other parts. The mesenchymatous cells found in the foregoing stage
on the median ventral line are transformed into a thin layer of the
endocardium of the heart and its anterior continuation (A. and ér:.a.)
which are suspended by the dorsal and the ventral mensenteries,
and enclosed in the thick pericardial coat still forming a con-
tinuation of the peritoneum. I can detect, however, none of the
traces of the peritoneal partition which was developed so markedly
in the last stage that one could not possibly overlook them. I
have endeavoured to trace the mode of disappearance of this
structure, but have only found that in one lot of embryos the
whole set of the structure was present while in the other no trace
of it was perceptible. Unfortunately I have found no embryo in
an intermediate condition.
The few cells observed from the last stage beneath the chorda,
and also in the space outside the pronephric tubules on both
sides are more or less multiplied. As the former group is
transformed finally into the dorsal aorta, and the latter into the
anterior cardinal vein of either side, I shall call them the tract of
the dorsal aorta and of the anterior cardinal veins respectively.
Fig. 98 represents the section through the hind border of
the branchial region. On either side of the enteric tube a small
space (pp.c.) of the body-cavity is surrounded by the peritoneal
epithelium, still consisting, in this part, of somewhat cubical
cells. The ventral edges of the peritoneal membrane of both
sides are just meeting at the median ventral line. A few mesen-
chymatous cells (ér.a.) found in the space between this meeting
point and the ventral wall of the enteric canal, are destined to
354 8. HATTA:
form the anterior continuation of the cardiac tube or the éruncus
arteriosus. An irregular cell-structure (x) is seen on either side
above the dorsal corner of the body-cavity and inside the tract
of the anterior cardinal vein. It is this structure about which I
could not at first decide with certainty whether it was a slice of
the hind wall of the branchial chamber or a part of the pro-
nephric tubule. All the cases examined, however, point towards its
being a part of the tubule ; the structure is detected in the anterior-
most part of the body-cavity which wedges in, at about this
stage, to the branchial region with a sharp angle (see fig. 97).
The narrow space (fig. 98, pp.c.) found intervening between the
structure and the peritoneal walls is a part of this cavity. One
might suppose that the space may be the coelomic cavity of the
branchial region ; but, the space between the parietal and the
visceral peritoneum of the branchial region is consolidated already
in the preceding stage, being filled up with variously shaped cells
of mesenchymatous nature (see fig. 97).
In the next following section shown in fig. 99, a tubular
structure (pi.2) with an oval outline is seen on either side at
the place where the pronephric tubule ought to be found. Its long
axis is directed just like a tubule (compare with figs. 101, 102, &c.).
This is doubtless a part of a pronephric tubule; but the corres-
ponding nephrostome which ought to be found either in the
section in front (fig. 98) or behind (fig. 100), can not be detected
in either of them. The nephrostome must, therefore, be looked
upon as having degenerated ; and since this pair of the tubules
is, in fact, detected underneath the fifth myotome, it must be
identified as the second pair of the tubules. The section represented
in fig. 100 shows on both sides the cross-sections of the collecting
duct, (cd.). On the left side, a cellular structure connects the
MORPHOLOGY OF CYCLOSTOMATA. 355
toneum and the collecting duct; it is the posterior wall of the
le in figs. 99. Fig. 101 represents the section through the axial
e of the third tubule, the nephrostomes of which are recog-
d more clearly in the section behind it (fig. 102, nst.3). The
les of this pair are comparatively not long. The fourth pair
he tubules and their nephrostomes are obvious in fig. 104
4 and nst.4) which represents the third section behind that of
102 ; the tubules much resemble those of the pair in front,
ving the same convolutions as these. It is a peculiarity of the
ent stage that the aperture of the nephrostomes of the third
the fourth pair is not so wide open as in the last stage or
n more advanced stages! It is always nearly closed and slit-
‚so that we can hardly trace the communication between
lumen of the tubule and the body-cavity.
Fig. 103 represents the section intervening between the sections
mn in figs. 102 and 104. On the right side, the collecting
‚ alone, and on the left side, the duct together with a small
of the fourth tubule, is shown. The peritoneal membrane
he dorsal end of the body-cavity is folded far into that
ty (fig. 103, ds.).. This fold is traceable from the anterior part
Le third tubule to the hind part of the fourth (figs. 100-104, ds.).
space enclosed in this fold communicates freely with both the
| of the dorsal aorta under the chorda and the tract of the
rior cardinal vein outside of the pronephros and contains a
ber of mesenchymatous cells which probably wander in from
tract of the aorta and the anterior cardinal vein. As sub-
ent history shows, this structure constitutes the beginning of
glomerulus of the pronephros.
Figs. 105 and 106 represent two contiguous sections immedi-
y posterior to the section shown in fig. 104. In fig. 105 we
356 S. HATTA:
observe on either side the cross-section of the collecting duct
(cd.) together with a part of the fourth tubule (pé.4) ; the longi-
tudinal section of the fifth tubule (pé.5) is seen on the right
side of fig. 106, standing in wide communication (nsl.5) with the
body cavity. This is the hindmost tubule. In the sections lying
behind this, the cross-section of only the segmental duct is
repeated. |
Thus the tubules of the second pair undergo, at the present
stage, complete degeneration. This process begins, in this case,
as above seen, at the nephrostome and proceeds upwards to the
collecting duct,—a process which is just the reverse of what
is observed in the reduction of the tubules of the sixth pair and
probably also of the first pair, in both which cases the tubules are
first cut off from the collecting duct and the separation from the
peritoneal cavity follows afterwards.
Period 585.
In the Stage v1, embryos have developed so far that all
the organs have received their definite forms and proper position
with the exception of the middle and the hind portion of the gut,
whose development is much delayed on account of the yolk-mass.
Having absorbed the yolk-granules, the component cells of most
organs are much diminished in size. .
Figs. 107-110 have been drawn from a series of sections
through an embryo in this stage. The enteric canal (fg.) is much
diminished in diameter, presenting, in section, an elongated heart
shape. The peritoneum becomes very thin in all its parts with the
exception of the pericardium and the coat of the truncus arteriosus,
in which its component cells are of cylindrical or cubical shape.
MORPHOLOGY OF CYCLOSTOMATA. 357
: peritoneal membrane lining the enteric canal immediately
ind the branchial region is also thicker as compared with
sr parts (fig. 107), being composed of a single layer of cubical
3,—a peculiarity observed since the last stage (compare figs.
99 with fig. 107).
The pronephric tubules as well as the collecting duct are
posed of a regular epithelium of cylindrical cells ; the former,
eover, are much prolonged and, in some parts (fig. 108), much
ed, so that the peritoneal cavity which was almost a hollow
se in the last stage, is filled up with the tubules and the
liac tube.
Fig. 107 represents the section through the hind part of the
h myotome; a pair of the tubules (p£.3) is hanging down in
body-cavity immediately behind the hind wall of the branchial
mber. On the right side, the axial plane of the tubule is
through, while, on the left, the anterior wall of it is sliced ;
e are the tubules of the third pair. They show no bending
he antero-posterior direction, but are curved laterally and ven-
ly. The component cells are, in the nephrostomal portion,
er in comparison with those in other parts of the tubule or the
ecting duct. The fourth tubule and nephrostome are seen on
right side of fig. 108, while on its left side, the communication of
corresponding tubule on the opposite side with the collecting
t is recognizable. The left nephrostome is found in the third
ion behind this, which is not figured. This pair of the tubules
ibits, in section, constrictions at two or three points owing
heir curving somewhat in the antero-posterior direction (see the
ale on the right side of fig. 108). Fig. 109 is the section im-
liately behind the last and shows the cross-sections of the
ecting duct (cd.) and a piece of the left fourth tubule (p£.4).
358 S. HATTA:
A pair of the glomeruli (figs. 108 and 109, g/.). is seen adher-
ing on the median side of the tubule on each side and lined with
the visceral peritoneum. The glomerulus represented in the
last stage by a folding of the peritoneum which covers the tubules
from the third pair to the fourth”, is reduced, at present, into a pair
of sacs of this membrane projecting on each side between the
fourth and the fifth tubules; the other part of the folded membrane
becomes adhered firmly to the walls of either the tubules or the
body-cavity leaving no space of sacculation,—in short, a pair
of long folds, extending from the anterior part of the third
tubule to the fifth in the last stage, is reduced into a pair of sacs
found in the position just mentioned. The inside of the sacs
is compactly filled up with free-cells and communicates with
the aorta tract and with the space outside the pronephros,
where free-cells to be afterwards transformed into the anterior
cardinal vein have heen observed already from the foregoing
stage.
The section represented in fig. 110 fortunately passes sym-
metrically through a pair of the nephrostomes (nsi.5) and of the
tubules (pt.5) hanging down in the peritoneal cavity. This is
the fifth or the hindmost pair of the pronephric tubules in the
present stage. The communication of the tubules with the collect-
ing duct is seen in the section behind this. The tubules present
also some antero-posterior bendings. Posterior to this, no tubule
is found.
The pronephric tubules in the present stage are, therefore, re-
duced into the minimum number, 1.e., three pairs”, all of which are
retained so long as the organ functions as the excretory apparatus
1) See p. 355.
2) We occasionally find the four tubules to persist, and the additional tubule is the sixth.
MORPHOLOGY OF CYCLOSTOMATA. 359
during the larval life of Petronyzon. Especially it must be noticed
that the foremost pair of the persistent tubules (the third pair) is in
close contact with the hind border of the hind wall of the branchial
chamber where, in the foregoing stage, the second pair of the tubules
was found, this latter having disappeared in the course of the
last stage. It follows that the two somites, to which the first and
the second pair of the tubules have bélonged, have now entered into
the formation of the branchial region.
The development of the pronephros after this consists only
in the prolongation and the convolution of the tubules, no further
change taking place with reference to the number of the tubules
or to their histological structure, until the system undergoes
degeneration to be replaced by the mesonephros, which functions
as the excretory organ for the whole subsequent life of Petro-
MAZON.
The convolution of the tubules is hard to make out. I
have reconstructed them from a number of sections; some of
these are diagramatically given in the annexed woodcut
(Woodcut 2).
With the growth of the muscle-segments the collecting duct
is prolonged, so that the points of connection of the tubules
with that duct become farther apart from one another, while the
nephrostomal portions of the tubules retain more or less their
original positions; in this wise, the tubules are laid in oblique
positions directed anteriorly and posteriorly (A) and have no other
curvature than the ventro-lateral bending (the frontal projection
of the curvature is shown in 7’). Then the antero-posterior
bending begins to take place. The foremost tubule is curved
forwards in its whole length, while a small curvature in the distal
360 Ss. HATTA:
(nephrostomal) portion of the two following pairs is directed
backwards. The nephrostomes retain their first position (B).
Now the secondary curvatures take
a place (C). The nephrostomal part
(TT of the foremost pair is crooked just
z Vz like that of the two hind pairs in
a sd. * B; the middle tubule is bent for-
: an wards like the foremost tubule.
Nay The hindmost tubule makes a
= small forward curvature and a
c 4% large backward bending. In the
ST next stage, D, the foremost and
- : the middle undergo no marked
change, but the secondary curva-
tures of the hindmost tubule are
much more strongly expressed. In
E the foremost receives a second- —
ary curvature directed backwards
at the middle part; the middle
acquires a curvature in opposite
direction; the hind tubule undergoes
no marked change except in the
Wood-cut 2.—Diagrams
showing the convolutions
increased degree of the original
of the tubules in later stages. curvatures. It seems that the
t. pronephric tubules. ;
si. segmental duct. subsequent bendings always take
place in the curved portion until there arises a system of com-
plexly convoluted tubules filling up the chest cavity.
As has been said, throughout these phases the positions of
the nephrostomes are not markedly changed, retaining the same
condition for a considerable period. The bendings of the tubules
MORPHOLOGY OF CYCLOSTOMATA. 361
are caused, therefore, by tlıe growth of the tubule at the point
of bending.
The curvature in the ventro-lateral direction is very simple
and undergoes no remarkable change ; its projection is shown in F!
B.—The Segmental Duct and the Genital Cells.
For the sake of simplicity, the development of the segmental
duct and of the vascular system in the pronephros has been
entirely put aside in the description given above.
As already alluded to, the origin of the segmental duct in
Petromyzon is extremely difficult to make out, because its formation
goes on rapidly at a comparatively young stage. The early
process of its formation is essentially the same as in the prone-
phric tubules. In the anterior region, the intermediate cell-masa
or the nephrotome (see p. 340) behaves itself in precisely the same
manner as in the Anlage of the pronephric tubules ; the difference
is that it is cut off from the lateral plate and is transformed into
the duct, while in the case of the tubule it retains the continuity
with the lateral plate. If fig. 31, which represents the section
through the tenth somite (i.e. the somite, from which backwards
the Anlagen are converted to the segmental duct) be compared
-with the left half of figs. 2, 5, 6, 14, and the right half of
fig. 16, in which the Anlagen all develop to the pronephric
tubules, it will be found that there is no difference between them ;
in fact, they are morphologically equivalent to one another.
Such an Anlage is, posterior to the pronephros, not confined to
the tenth somite, but, as has been already repeatedly said (pp. 317,
320, and 327), 1s observed for some segments further backwards
(see fig. 17).
362 8. HATTA:
The Anlage thus pronounced in each somite soon assumes a
characteristic oval form, being completely cut off from the myo-
tome to which it belongs (compare the right side of fig. 3 with the
right side of fig. 58 and see the description on p. 335). The mode
of constriction is also the same as in the case of the pronephric
tubules; the indentation begins at the anterior and posterior
borders of the somite, and the middle portion is cut off last
(compare with the explanation on p. 320).
Here also, the coelomic projection is formgd in the same mode
and at the same point as in the case of the pronephric tubules (see
left side of fig. 63, c.p.).
Up to about this time, the Anlage shows a feature much
resembling that of the tubule, so that one who has not followed its
further history might mistake it for a pronephric tubule (compare
the left side of fig. 63 with figs. 67-74). But cell-mutiplication
which occurs almost invariably in the case of the pronephric
tubules, is not observed in the Anlage of the segmental duct
which is soon cut off from the lateral plate (including the coelomic
projection) and assumes a characteristic tubular structure com-
posed, in cross-section, of radially arranged cells of columnar shape.
Its position is always on the parietal aspect of the dorsal (proxi-
mal) angle of the peritoneum where the coelomic projection passes
over into the lateral plate (see fig. 75). This separation of the
Anlage of the duct from the lateral plate goes on, it seems to me,
on the whole from the anterior part to the posterior, but often ir-
regularly ; for not infrequently, the duct in some anterior somite is
connected with the lateral plate, while it is already cut off com-
pletely in posterior somites. In fact, there are some somites in
which the separation is very much delayed and I have often been
surprised to find what appeared like a pronephric tubule in a
MORPHOLOGY OF CYCLOSTOMATA. 363
somite (see fig. 63) far backward of the posteriormost tubule
which is found in the ninth somite.
The segmentally arranged Anlagen of the segmental duct
are secondarily united with one another just as in the case of the
collecting duct in the pronephric region. This union seems to
take place during the separation of the Anlage from the myotome
and is finished before it is separated from the lateral plate ; for,
when the Anlage first comes into view, there is no intersomitic
cord as in the case of the pronephric tubules and the duct is seen
already consisting of radially arranged cells (fig. 58) when it is
cut off from the lateral plate. When established, the duct is the
same in structure in both the somitic and intersomitic spaces; a
cross-section of such a duct in the intersomitic portion is shown
on the right side of fig. 75, while that in the somitic portion
is seen on the left of the same figure.
This condition of the duct is already traceable, in Stage 111,
for no fewer than 10 somites from the hindmost pronephric tubule
backwards, and it forms a direct posterior continuation of the
collecting duct. The duct remains awhile as a solid cord of cells
arranged radially in cross-section, but it soon acquires a lumen
(figs. 75, 86, and 87, sd.). The further development of the duct
goes on more promptly than that of the tubules in the hind part,
and therefore, the embryos at such a stage (Stage 111) have a well
developed duct and more or less primitive tubules (compare fig.
74 with fig. 75).
In the hind region, where yolk-cells are crowded, the pro-
cess is much delayed and more or less modified. Instead of the
differentiation of the cells in situ, it seems to me,. a few cells
are detached from the nephrotome ; a number of cells is produced
by repeated division of these cells (fig. 19, «.sd.) and becomes ar-
364 8. HATTA:
ranged as in the Anlagen in the anterior region. Fig. 89 represents
the section through the twenty-eighth somite in the series of sections
shown in figs. 77-86 ; it is the hindmost section in this series of
sections, in which the cells just spoken of are detected ; there are
found a few cells (a.sd.) of this kind which show no definite struc-
ture, but are scattered. In the next anterior section (fig. 88) the
cells are arranged more or less radially. In the sections lying
further anteriorly to this a perfect tube is formed as seen in fig. 87
(sd.) which shows the frontal section through the seventeenth to
twenty-third somites in the same series as the above two figures.”
In what somite this modified mode of the formation of the duct
begins I can not tell with exactness, but it is certain that the duct
arises by the differentiation of the nephrotomie cells in sıdu more
than 10 segments back of the hindmost pronephric tubule. I have
considered it possible that these cells (a.sd.) might be epiblastic in
origin, but I can not find that the cells composing the epiblast over
this cell-group show any sign of mutiplication; while on the
other hand, the cells on the dorsal edge of the lateral plate
(which corresponds to the nephrotome in the anterior part) are
very active. I see, therefore, no escape from the conclusion that
these cells are mesoblastic in origin.
Also in the anterior part of the body, the epiblast consists
throughout these phases of development always of a single
layer of cubical cells and shows a sharp contour against the
structure inside it, being, in most cases, intervened by a space.
Naturally, mitotic figures are observed at several points, but the
products of these cell-divisions contribute only to the extension
of the ae itself, as may be inferred from the direction of
— — —— a in
1) By the bending of the body-axis, some sections in a series of cross-sections are un-
avoidably cut through frontally.
MORPHOLOGY OF CYCLOSTOMATA. 365
the spindles, the long axes of which are directed always parallel
to the surface of the layer. J have nowhere observed any trace of
either the proliferation or of the casting off of cells from the epiblast
to give rise to the segmental duct.
In the cloacal region, the formation of the segmental duct
goes on a little earlier than in the region next anterior to it.
In spite of much effort, I failed to observe the very beginning
of the formation at the cloacal opening, and I have nothing to
tell of its earliest stage. In the series of sections from which
figs. 77-89, are drawn, I can not yet find in the adjacent part
of the cloaca any trace of the duct; but in the section repre-
sented in fig. 90 which passes through the dorsal lip of the
blastopore of an embryo with about the same number of the
mesoblastic somites (34 or 35) as the one just referred to, the
duct already breaks through into the cloacal cavity (co.sd.)”. Fig.
91 represents the next ventral section which passes through the
dorsal part of the blastopore (Jp.). As seen in these two sections,
immediately inside of the blatlopore (4p.), where the hypoblast
passes over into the epiblast, the walls of the cloacal cavity send out,
right and left, a symmetrical pair of diverticula” (c.dv.), forming
an acute angle, the inner side of which is a part of the enteric wall,
while its outer side is the direct continuation of the epiblast.
The walls of this diverticulum pass over into the segmental duct
(sd.). The communication of the segmental duct with the cloacal
cavity 18 found, therefore, at the point where the epiblastic layer
of the lp is reflected inside and passes over into the hypoblast.
This point of communication is, however, shifted far inside and
RN ne
1) This opening is found in the same vertical plane as the 34th or 35th somite.
2)The right diverticulum only is seen in figs. 90 and 91, the left one being observed in
another section which is unfigured.
u
a}
5
S66 Ss, HATTA:
dorsally when the development proceeds further (fig. 1
and e.dv.).
I have also met with two cases (figs. 90 and 111), i
I have observed some epiblastic cells of the external
walls of the blastopore multiplying actively and having
spindles (z) with axes directed perpendicularly to the plar
epiblast, while the duct comes in firm connection w
point of the epiblast,—the connection is so firm that !
and the epiblast appear to form one and the same tis
this point, thus, there is every appearance of epibla:
partaking in the construction of the segmental duct.
The collecting duct pertaining to the ninth somi
the segmental duct in that segment, having lost the eo
with the tubule.
Up to Stage 1, the duct is represented by the ®
Anlagen in about 8 segments back of the ninth somite ;
1, these Anlagen are converted into the duct in about
terior segments; while in the course of Stage ıv it o
into the cloacal cayity.
From the above account, it is easily conceivable that
lage of the segmental duct and that of the pronephrie tu
perfectly homologous, and that the duct is a continuation oj
of abortive pronephric tubules in the hind region,
Underneath ten and more myotomes lying posterior
the fifteenth somite the proximal portion of the later
whieh corresponds to the nephrotome, contains peculi
cells (figs. 87,88, and 89, ge.) loaded with an enormous qu
yolk-granules; the other mesoblastie cells in this pal
much flattened out, form a thin layer over these cell:
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MORPHOLOGY OF CYCLOSTOMATA. 367
uliar cells are, I think, the equivalent of the primitive
ital cells found in the corresponding part of the Amphibian
Selachian body.
Up to Stage 111, these cells can not be distinguished from
2r mesoblastic cells which are equally rich in yolk-granules.
Stage 1v, they become conspicuous; and in Stage v, again
stinguishable from other constituent cells of this part.
C—The Vascular System in the Pronephros.
In early stages, no trace of the vascular system is perceived
he pronephros. What is recognisable as a fore-runner of the
el is represented by mesenchymatous cells scattered in the
*e between the primary germinal layers (figs. 77 and 82,
.). These free cells are detected, during Stage tv,” in three
ts, viz., beneath the chorda, beneath the ventral wall of the
ric canal and outside the pronephric tubules on either side (figs.
79, 80, 81, and 82, mch.). In Stage v, or at the end of Stage rv,
cells below the hind section of the fore-gut are converted into
endothelium of the heart and of the vessels which are its
et continuations. The cells beneath the chorda are destined
je transformed into the dorsal aorta, and the cells on either
of the pronephros constitute the first indication of the cardinal
is. It is these three vessels—the aorta and the two cardinal
is—which come in relation with the pronephros.
In the embryos in which the degeneration of the tubules is
going on, there is no special vessel supplying the pronephros;
when the process is over, a pair of long blood-spaces (figs.
-104, bs.) is found in communication with the aorta-tract.
) A few of them are observed here and there already in Stage IL
368 8. HATTA:
They are the spaces formed by the slackening and folding of the
median peritoneum which coats the pronephric tubules, as above
stated (p. 355). The fold, i.e. the space, extends throughout
almost the whole length of the pronephros (figs. 100-104) and
contains numerous free-cells. But, as the peritoneum finally
adheres to the median walls of the tubules in each nephrie
segment, the space becomes divided into three pairs: the anterior
pair, which soon disappears, is found between the tubules of
the second and third, the middle is detected between the third
and fourth, and the posterior between the fourth and fifth tubules.
These spaces communicate directly,—medially with the aorta tract
and externally with the tracts of the anterior cardinal veins,
which emerge, in later stages, in the pronephros. They are the
blood-spaces which, I believe, correspond with the intersomitic
arteries demonstrated by Paunt Mayer and others in Selachia.
When the tubules develop further, the arterial portion of
these blood-spaces disappear except the middle portion where it
is sacculated and filled up with the mesenchymatous cells
(figs. 108-109, g/.). This portion is the structure which is called
the glomerulus of the pronephros; it is found one on each side
(see pp. 355 and 358).
Having followed, in the foregoing pages, the successive pro-
cesses which take place in the development of the pronephros
in Petronyzon, step by step, I will give a short resumé of the facts.
1. In the earliest part of Stage 11, the mesoblast consists
simply of the parietal (dorsal) and the visceral (median and
ventral) layers. The proximal portion of the mesoblast is dis-
MORPHOLOGY OF CYCLOSTOMATA. 369
tinguished, first of all, in the histological structure from the
distal portion: the former is composed of columnar cells, and
the latter of irregularly shaped cells. Only the proximal portion
which occupies the largest part of the mesoblast undergoes the
metameric segmentation and gives rise to the scleromyotome and
the nephrotome (in the sense of Ruckert); the distal smaller
portion remains unsegmented and is later converted into the
flattened epithelium of the peritoneum.
2. The earliest traces of the pronephros are noticeable in
exceedingly young stages, that is, in the early part of Stage 11,
in which the embryo has about 16 somites.
3. They are expressed in the form of a diverticulum of the
parietal layer of that section in each mesoblastic somite, which
forms the ventral half of the segmental part of the mesoblast
and is called the nephrotome. This is the Anlage of the pro-
nephric tubule and not of the segmental duct.
4. The pronephric diverticulum or the Anlage is brought
about by the evagination of the parietal layer in each nephrotome,
enclosing a part of the primary coelomic cavity.
5. The nephrotome is separated from the proximal portion
of the segmented mesoblast and forms awhile the proximal portion
of the unsegmented mesoblast or the lateral plate. The separation
begins with an indentation in the anterior and posterior borders
of the mesoblastic somite ; the myocoelome communicates for some
time by a narrow passage with the general coelomic cavity.
6. The Anlage has no histological connection either with
the preceding or the following Anlage or with the other
germinal layers; it is, therefore, segmental in origin and myo-
meric in position.
7. The Anlagen are developed, in Stage 11, in about 12
370 S. HATTA:
segments and are cut off from the scleromyotome in 4 :
In Stage 11, the separation of the Anlage from the
goes as far backwards as the sixteenth or seventeenth
8. The anteriormost Anlage is found in the hind pe
fourth somite and is the first to arise; the second follow
so forth.
9. The Anlagen in each somite are secondarily un
one another by the solid cellular cord which is budde
the anterior and posterior rims of the Anlagen themsel
the collecting duct (Sammelrohr in the sense of RuCKERT)
lished. This process is originally to be looked upc
coming together of the ends of the tubules.
10. The canalization of the collecting duct begins wi
Anlage and proceeds, generally speaking, posteriorly, —
Anlagen in front and back are put in free communica
11. Each Anlage grows dorso-laterally and acquires’
form. The collecting duct is shifted gradually in a dors
direction ; finally it comes to lie between the myot
mesentery, and the chorda dorsalıs.
12. The tubules open in the coelomic cavity at tl
angle of the dorsal corner of that cavity.
13. In the somites posterior to the ninth, the tuk
during Stage 111, cut off also from the lateral plate and
a long duct running, on each side, along the dorsal
the lateral plate where originally the tubules opened.
the segmental duct; the tubules and the collecting du
somites anterior to this constitute the glandular pa
pronephros.
14. The glandular part, or the pronephros proper «
six somites, from the fourth to the ninth. The maximun
MORPHOLOGY OF CYCLOSTOMATA. 371
he pronephic tubules which is attained by the embryo in
e III, is, therefore, six pairs.
15. The tubules ofthe first and second pairs come, in Stage
temporarily in close contact with the epiblast, but do not
ve cells from it; they soon return to their original condition.
16. The anterior extremity of the system shows, from tlıe
‚ degenerating features. The first, second, and sixth of the
les begin, during Stage 111, to decline; and at the end of
e Iv, or the beginning of Stage v, the tubules are reduced
the minimum number, which consists of three pairs from
third to the fifth. These three pairs function as the actual
etory organ for a considerable length of time.
17. Retrogression is first met with in the first pair of tlıe
les, which decline probably without further development,
_after their separation from the myotome is completed ; they
| to atrophy from the free end. The next pair degenerating
ie sixth, which is at first cut off from the collecting duct
remains for a short time, but soon disappears without leaving
ace. The second pair persists for some time seemingly to
tion as the excretory organ, but it atrophies already in
early part of Stage v, the communication with the coelomic
ty being first obliterated ; and in Stage vr, none of the struc-
remains to be recognized.
18. The foremost pair of the persistent tubules comes to
n close contact with the hind wall of the branchial chamber.
two mesoblastic somites which correspond to the first and
nd nephromeres should therefore be looked upon as having
red into the formation of the branchial region.
The stages in which the tubules appear and abort in different
ites are shown in the annexed table.
372 % HATTA:
l#8 | 5 fe le |B le
" | = r - = = _
SNS edit) E lé | à à | §
° = 3 = | £ a | F4
a a a | lé | & 1 8 le
Stage IT | Aut Anz An.) And An 5 Anl.6| Anl.7
[Stage III | | Tubs | Tub.4 Tub.5 | Tab.6| Segm
Stage IV
|
Stage V
Stage VI
19. In older embryos of Stage 111, the visceral la:
nephrotome is folded out, and is called the coelomic
which resembles the coelomic pocket described by 1
Ddellosloma ; however, in Ldellostoma, the fold is der
the parietal and visceral layers of the lateral plate and
wards converted into the Bowman’s capsule, whereas the
projection is the product of only the visceral layer of th
lome ; it gives rise to the radix of the mesentery wh
materials to the mesonephrie tubules and to the gonac
20. The topographical position of the pronephro:
first definite in Stage ıv. It is situated in the chest cavi
lateral to the heart, forward of and dorsal to the liver,
along either side of the chorda. This position is
changed as the development proceeds; the pronephr
in later stages, in front of the liver.
21. During Stage 1v, a structure, which I ha
above the periloncal partition, is observed as an
of the peritoneal wall and disappears during the sa
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MORPHOLOGY OF CYCLOSTOMATA. 373
This horizontal partition is found at three levels. The most
dorsal is well developed, the ventral is a mere trace, and the
middle is intermediate between the above two. I can not state
at present anything definite as regards the significance of this
structure.
22. The convolution of the pronephric tubule takes place in
Stage ıv. With the growth of the myotome, the collecting
duct is prolonged; consequently the connecting points of
the tubules with the duct are farther removed from one another
than before, whilst the nephrostomes retain their original
position ; so that, the two posterior pairs of the tubules are placed
in an oblique direction from dorsal and caudal to ventral and
cranial. Each tubule is, then, convoluted in a cranio-caudal
direction between the heart and the lateral peritoneal wall. In
older stages, the tubules are coiled in all directions, until the
chest cavity becomes filled up with the convolution of the tubules.
23. Up to Stage vi, the nephromeres and the myomeres
exactly coincide one above the other in position. This period is
very long in comparison with other Craniota. As the develop-
ment proceeds further, the pronephic tubules are however shifted
gradually backward, so that, in Ammocoetes 10 mm. long, the
myotomes are already not situated upon the tubules pertaining
to each of them. In later stages, nothing of the relation can be
traced.
24, The segmental duct is looked upon as being brought
about by the union of a series of the abortive pronephric tubules
in about 12 somites lying posterior to the eighth somite”. The
Anlage is laid in the parietal layer of the nephrotome in exactly
the same manner as in the pronephric tubules of the glandular part.
1) In the 9th somite, the aborted tubule actually forms the duct in the segment,
Auen 3:
+
=
.
ve .
— + - mm - een
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ie I enger
a me
wie)
rye net — Ow je
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—
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up
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314 S. HATTA:
The difference is, that the tubules in the posterior region are
soon cut off from the lateral plate and become the duct.
25. Between the epiblast on one hand, and the Anlage or the
duct on the other, there exists always a space, and the duet
has no connection with the epiblast except at its posteriormost
end where the epiblastic cells might, as judged from the mitotic
figures, contribute to the formation of the duct.
26. In the somites posterior to about the twentieth somite,
the Anlage of the duct is represented by a few cells in each
segment probably detached from the dorso-lateral angle of the
nephrotome. These cells multiply and are transformed into
the segmental duct in the posterior part.
27. In Stage ıı, the Anlagen of the segmental duct are cut
off from the mother layer in a few somites; in Stage ıır, the
duct is formed as far as about the eighteenth somite, while in
Stage tv, it breaks out into the cloacal cavity. The cloacal
opening of the segmental duct is found at a point where the
hypoblastic cloacal wall is reflected into the epiblast, these two
layers forming a diverticulum on either side.
28. In stage ıv, the primitive genital cells become apparent
in the nephrotomes of the posterior 10 or more somites; they
can not be discriminated from other mesoblastic cells in the
next advanced Stage.
29. The blood-vessels, which specially supply the pronephros,
acquire definite form in comparatively later stages, viz., at about
Stage v. The dorsal aorta pours out the blood into two pairs of the
blind vessicles, which are formed by the folding of the parietal
peritoneum and are found between the first and second pairs,
and between the second and third pairs, of the persistent tubules
MORPHOLOGY OF CYCLOSTOMATA. 375
ectively. The venous blood is carried away through the an-
or cardinal veins which penetrate the pronephros.
29. These blood-spaces are thus segmental in arrangement
antersomitie in position. The two anterior pairs of them soon
ergo atrophy, but the posteriormost pair persists, becoming
irged and sacculated at the distal extremity. This sacculated
| of the vessel is filled up with free-cells and is called the
nerulus of the pronephros, and, therefore, there is only a pair
‚lomeruli in Petromyzon.
II. Historical Review and Conclusions.
As is well known, Max ScHULTZE (’56) was the first who dis-
red the pronephros in Petromyzon. Having investigated the
æ of P. planeri, the author describes the structure as
rüsenanlage’’ and homologised it with the “ Urnieren (Wolf’sche
per)” of the frog’s larva. His statements on this body are as
ws: “ Nicht lange nach der Bildung dieser Drüse (Thymus)
teht die Anlage einer zweiten, aus dem unter der Chorda
alis angehäuften Blastem über dem Herzen. Aus der durch
mentanlagelungen früh schon sehr undurchsichtig werdenden
se wachsen nämlich nach unten, gegen das Herz zu, 3 oder
urze Fortsätze hervor, welche eine eigenthümliche Wimperung
en’”’ (p. 30).
The stage spoken of probably corresponds to Stage v, or vr,
ny embryo.
Our knowledge on this subject received important additions
the noted investigations of W. Mutter and Max Für-
NGER. MULLER (’75) noticed the first traces of the pro-
hros in a very young embryo, which had yet only four pairs
376 8, HATTA:
of gill-slits. This Anlage gives rise to a much coiled glar
opens into the body-cavity, at first through only on
funnel, but afterwards through four. The gland pa
posteriorly to a pair of ducts, which run along the e
either side and open into the cloaca. Mürter has ho:
the structure with the “ Vorniere” of Myxine and €
duct “ Urnierengang ” (pp. 121-122). He found a pai
meruli projected on the median surface of the gland :
with the peritoneal epithelium.
Max Fursrincer (78) studied the larve of A
planeri, which varied from 4.5 to 180 mm. in length.
ments essentially confirm Murter’s. In his account we
following sentences: “Die auf allen Präparaten au:
Vorniere, die ich im Wesentlichen ganz wie Mounier fai
einen nahmentlich bei den mittleren Stadien volumin
durch 4-5 Myokommata erstreckten Complex von Wi
die vorn durch mehreren Peritonealeanäle (Wimpertr
Bauchhöhle münden und hinten in den Vornierengang ü
Diese auf die 2-3 ersten Myokommata beschränkten
ragen in unregelmässiger Folge bald ventral-medial, bal
lateral in die Bauchhöhle vor nnd wurden (von C.
und mir) meist zu fünf gefunden. Die von rundliche
zellen bekleidete Glomerulus verhielt ganz wie Miner be
(p. 42).
The larve of Ammococles in question seems to @
probably to Stage y, or later stages of my list; in suel
I could not find more than three (or rarely four) pai
tubules, or of the nephrostomes."
1)See the foot-note on p. 358.
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MORPHOLOGY OF CYCLOSTOMATA. 377
The authors who have investigated the development of
tromyzon embryos step by step, are W. Scott, GOETTE, SHIPLEY,
d v. Kuprrer. Their opinions are, however, somewhat
rergent. Scott (’82) derives the pronephric tubules from the
‚mental duct which is, according to him, brought about by
> differentiation im situ of the cells forming the proximal
rgin of the lateral plate. The process takes place in the
ole extent at the same time. At certain points (segmental ?)
the duct thus furmed, evaginations are produced out of it;
se evaginations subsequently open into the body-cavity and
ablish the nephrostomes which are, according to Scorr,
ind from two to three pairs in number. At about the stage
which the funnels are formed, he observed a pair of
meruli.
“In most respects,’ SHIPLEY’s observations (’87) “ confirm
”” (Scorr’s). But “on the origin of the ciliated funnels, the
ults differ from Scorr’s ”” and agree with those of FURBRINGER
mphibian pronephros ?). According to SHIPLEY, “in the region
the heart, where the body-cavity has already appeared, its
gin (2.e., of the segmental duct) seems to be somewhat different.
e lumen of the segmental duct here becomes continuous with a
Dove in the parietal peritoneum, lying near the angle where
: somatopleure and the splanchnopleure diverge. When this
ove closes it leaves four or five openings which persist as the
enings of the ciliated funnels’ (p. 20).
v. KupFrer” (’88) observed, in P. planert, the three pairs of
> tubules arising from three distinct evaginations of the parietal
1)I know this paper only by the abstract in: Jahresbericht ii. die. Fortschr. d. Anat.
>hysiol., Bd. 17. 1889.
we,
3/8 8. HATTA:
layer of the lateral plate; the segmental duct is looked upon as
of the epiblastic origin.
GOETTE (90) worked the development of Petromyzon fluvia-
talis ; his results with respect to the pronephros show some agree-
ment with mine, especially those concerning the later stages. The
author derives also the whole system of the pronephros (including
the segmental duct) solely from the mesoblast. But we diverge in
some important points from each other ; he has found the earliest
traces of the structure at a time when the rudiment of the heart
first becomes apparent (his vi. Periode) (p. 64). From the account °
given in tbe foregoing pages it is clear that this period belongs
to a later stage in which the pronephros has already made a
considerable progress in development; his figures 99, 103, &e.,
which are spoken of as representing the first appearance of the
structure, approximately correspond with my figures 82, 83, &e.,
and with those of even older stages.
The pronephros is, according to GOFTTE, not of a separate
Anlage in its first appearance, but arises in a form of a longi-
tudinal furrow formed, on each side, by an evagination of the
parietal layer of the mesoblast ; the lips of the furrow being fused
at certain points, there remain three openings; these are
converted afterwards into three tubules and ciliated funnels.
The tubules are added by stages until there are usually five, or
more rarely four or six; but how these are multiplied, he can
not say with certainty. The tubules have, it seems to him, no
relation to the metameres of the body; for 3 to 5 tubules are
found in the extent of 2 to 3 metameres (loc. cit, pp. 64-69).
The segmental duct originates, according to GOETTE, in pre-
cisely the same way as the pronephros proper; the only difference
is the complete constriction of it from its mother-layer just as
MORPHOLOGY OF CYCLOSTOMATA. 379
ave made out. From the region of the liver-anlage backward
development of the duct is irregular; he says: “ Auf der
n Seite zeigt sich seine Anlage noch rinnenförmig, während
auf der andern Seite schon" vollkommen röhrenformig ab-
hnürt ist. Endlich wechselt dies Verhalten auch auf derselben
perseite, so dass derselbe Gang, von der Lebergegend rück-
ts verfolgt, bald rinnen-, bald röhrenförmig, geschlossen oder
offener Lichtung sich darstellt ’’ (doco cit., p. 56). The hind
ofthe duct opens in the cloaca (Afterdarm) by the fusion of
r walls and by the communication of the lumen of the duct and
diverticulum of the cloaca. I have not observed in any stage
ny embryos examined the numerous convolutions of the seg-
tal duct demonstrated by GoETTE in the region immediately
ind the “ ursprüngliche Kopfniere.”
GOETTE has made out the three “ peritoneale Scheidewiinde,”’
e calls them : two respectively in the anterior and the posterior
of the pronephros, and the third on either side of the liver.
er, the first contributes, according to him, to the formation of
hind wall of the branchial pouch (Kiementasche) ; the second
converted into “eine Venenbriicke zwischen dem Sinus
osus and der Leibeswand,’ while the third disappears without
ring a trace. They are, according to GoETTE, homologous
a the “ Schlussplatte ’’ of the pronephros in Teleostei; con-
red phylogenetically, nevertheless, they have no intimate
tion to the pronephros in Petromyzon (loco cit., pp. 56-61).
s structure is, as stated on p. 349, doubtless the same as the
ermost peritoneal partition which I have found in my em-
os. I have nothing to communicate on its significance ;
I feel sure that his statement is not accurate when he says
structure appears earlier than the pronephros; for his figs.
380 &, HATTA :
96 and 97, to which his statement refers, represent a sti
siderably later than the first formation of the pronephr:
And the peritoneal partition is not confined to these thre
but is continuous throughout the whole extent of the pror
moreover, beside the “ peritoneale Scheidewinde,” there a
two other partitions of a similar character as above
Also, as to the fate of the structure my results differ ft
I have not been able to observe at all any such cont
to the formation of the hind wall of the branchial cham
of the ‘ Venenbrücke,' as is affirmed by GoETTE.
Rast ('96) says in his recent extensive work on the 8
nephrie organ, that in quite young larvæ of Pelromyzon j
the pronephros also begins in the seventh somite, in wl
first of the four ostia are found, as in Pristiuru.” H
are, however, 501 hours or 20 days and 21 hours ol
larye correspond to my embryos in Stage vı, and upw
which anteriorly two pairs, and posteriorly, one pai
tubules disappeared and only three persistent tubules a
His first nephrostome represents the foremost of the p
nephrostome.
The accounts cited above all agree with the resul
in the present paper in deriving both the pronephros
segmental duct from the mesoblast alone, with the singh
tion of vy. Kurrrer who assumes the epiblastie origin
segmental duct. They differ from the account given in |
going pages in the mode of the formation and in the nu
the tubules formed. The first point of difference is du
1) See the reference tinder Selachia (p, 300).
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MORPHOLOGY OF CYCLOSTOMATA. 381
t that the authors probably overlooked the earliest phases of
mation, which take place, as shown above, in a stage very
ing, but not younger in comparison than that in other Anamnia ;
the formation follows the metameric segmentation of the mesoblast
the anterior region. In later stages, the tubules and the inter-
nitie portion of the collecting duct repeated in sections of a
ies appear, indeed, like the cross-sections of a longitudinal
row or groove of the lateral plate, the lips of which are fused
certain points, as described by SHIPLEY and GOETTE (see my
. 66-74). |
The number of the tubules and nephrostomes varies accord-
to the stages of development. And if some stage or stages
overlooked, it must necessarily lead to an erroneous conclu-
1. This is the probable reason why the statements of the
ters with reference to the number differ.
Indeed, the anterior extremity of the pronephros has already,
m the first appearance, the features of a rudimentary organ ; the
t pair of the tubules can not be observed at the same time
h the following five pairs, except by extremely good luck. In
1e embryos of Stage II, we see occasionally the collecting duct
ne in front of the first tubule, so that we are led to infer
t there were some pairs of tubules in front of the present
t pair, which have degenerated during the course of the
estral history.”
As is seen above, all investigators who have been occupied
h the study of the development of Petromyzon agree in de-
ibing only one pair of glomeruli. SHIPLEY says “there is only
: glomerulus on each side, stretching on each side of the
1)I have stated above that in the earliest part of Stage 111, the anterior extremity of
left collecting duct presents a conical protuberance (see the foot-note on p. 329).
352 S. HATTA :
ılimentary canal extending through about the same space as the
glandular part of the kidnev. Each glomerulus is a diverticulum
of the peritoneum, which generally becomes sacculated ;.........”
(p. 21). The statements by GoETTE confirm SuirLey'’s, and my
results also agree with theirs. However, this is not all of the
vasculur system of the pronephros but represents a posterior
portion of it, the anterior part having disappeared entirely
(see p. 368).
No previous writer on Petromyzon has described such early
stages as given above in the development of the pronephros, nor
lias any one remarked the temporary existence of the pronephric
tubules in the branchial region as well as in the region of the
segmental duct. I will, therefore, extend the comparison over the
allied groups such as Myxinoids and Amphiorus, and higher
Craniota to verify the new facts.
With reference to the development of the nephrie organ in
Myxinoids, there is a great deal of information which we owe
to the unwearied labors of W. MuLLER, Semon, WELDON, and
others”. They had, however, no opportunity to observe the earliest
stage of the embryos. Recently our knowledge on this subject
has been greatly augmented by the new works of Price, Deas,
and Maas.
Price (97) worked out the early development of the prone-
phros observed in a few embryos at different stages of Ddellostoma
slouli. According to him “the first indication of the system
oveurs here in the eleventh segment (of spinal ganglion), and
consists of a simple thickening of the somatic layer of the coel-
1) 1 have not seen the paper by J. MULLER.
MORPHOLOGY OF CYCLOSTOMATA. 383
omic epithelium, which extends through seven sections,............
the thickening has not been caused by a proliferation of cells,
but certain cells having assumed the form of columnar epithe-
lium, while the adjoining cells retained the form of flat epithelium.
are later an evagination will here take place, to form a
segmental tubule ” (p. 209). These evaginations are connected
with one another “by a streak of columnar epithelium, which
in transverse section resembles the first tubule anlage, except that
there is no concavity on the lower surface; this is the segmental
(collecting) duct.” “ The union between the duct and tubules is,’’
in another place he says, “primary and not secondary ”
(loco eit., p. 210).
The pronephros in Bdellostoma comprises, according to PRICE,
69 segments (spinal ganglions). As it begins at the transverse
plane opposite the eleventh spinal ganglion, it is inferred
that the pronephros in Bdellostoma is extended over the whole
length of the branchial region. But “the excretory system
disappears through the greater part of this region before the
gills are formed’ (/oco eit., p. 217).
The segmental duct (in s. s/r.) is, according to the author,
brought about by the rudiments of the hinder 20 degenerated
tubules (in his Stage C); the number of the declining tubules
increases by stages: in Stage A, there are two; in Stage B,
nineteen ; and in Stage C, twenty.
This account is thus in close agreement with that given in
the present work, excepting a slight difference as to the origin
of the collecting duct and as to the number of the tubules. In
Bdellostoma, the collecting duct develops out of the Anlagen
independent from that of the tubules, while in Petromyzon, as
stated in the foregoing description, the Anlage in a mesoblastic
se ve vente on
pes >
384 S. HATTA:
.—
somite develops solely into the tubule, and by the secondary |
union of the tubules’ ends, the collecting duct is brought about.
As regards the number of the tubules, there are, in
—— - Ee
| oy Yee ey ET
- *
Petromyzon, only two pairs in the branchial region instead of
twenty in Bdellstoma. The number is, however, of secondary
od.
| sl end be
importance ; it varies with the stages of embryos and possibly
ig Sc:
af with individuals, and naturally more with the embryos of
at different families. This numerical variation is readily explained
4 by the degenerating tendency of the tubules.
PRICE has made out the segmental evaginations of the dorsal
ee 4 es
corner of the coelomic cavity corresponding to the nephromeres;
they are called by him the “ coelomic pockets.” In Petromyzon,
i I have found a series of solid knobs on the visceral layer of
at the intermediate cell-mass, which are transformed into the |
i segmental folds of epithelium, forming then the direct continua- |
4] tion of the peritoneum. Thus the coelomic pocket in Bdellostoma
i] and the coelomic projection in Pelromyzon are apparently very
a similar structures; the two, however, differ from each other in origin 4
al and in fate. The former (coelomic pocket) is constructed by the
a parietal and visceral layers of the /ateral plate, while the latter
Fait (coelomic projection) is the product of only the visceral layer |
Hi of the nephrotome, the ventral half of the segmented part of the
4 mesoblast. The coelomic pockets become the Malpighian body,
Ha
and the coelomic projections give origin to the radix of the
mesentery, from which the gonad-cells and the mesonephric
.
I Lin 27 A A
tubules are derived. Nevertheless, these two structures are, |
= 2 _ 1°
believe, homologous. PRICE’s statements on the derivation of the
=
+ -
: i ee a a ee as oe... . es
IE TITTEN mes pa 6 Y
n coelomic pocket from the two peritoneal layers, are not as clear as
it is desirable, and its partition from the body-cavity might, it
EA w = | |
Hi seems to me, represent the uppermost peritoneal partition which
t4°
JW
FH
MORPHOLOGY OF CYCLOSTOMATA. 385
n disappears, in Petromyzon, without any definite significance.
any rate, the structure represents “parts of the original
mental coelome, that is, the nephrotome,’’ an unmistakable
t which is denied by Price.
The embryos of Myzine which formed the materials of the
uable works by Maas are too old to be compared with those
Petromyzon used in the present work. But the results ob-
1ed by the author differ from those of Prick in an important
nt, namely, in the derivation of the mesonephros.
The pronephros and mesonephros are, according to PRICE,
erent parts of the same organ. “ Ifthe organ in question could
y be a pronephros alone, or mesonephros alone,” says PRICE
should unhesitatingly pronounce in favour of its being a pro-
yhros’’ (loco cit., p. 120). And he proposes to call “the
ire embryonic kidney holonephros.’’ With Rast, Maas, and
ers, I hesitate to accept Price’s conclusion; for there are,
may be inferred from his statements, great gaps not only
ween the Stages B and C, but also between Stage C and the
lt. The formation of the mesonephros takes place in Petro-
zon only at astage much advanced, in which the processes of
formation and degeneration of the pronephros go on in much
same manner as in Bdellostoma, and it is open to doubt if the
;onephros might not appear in later stages which were lacking
ong Price’s materials.
Up to the oldest embryo observed by Price there were
ther glomeruli nor bloodvessels of a definite form, although there
-e found in the splanchnopleure some vessels whose position
med to suggest their corresponding to the glomeruli of Selachia
| Amphozus; “ but they do not have any relation to the
mings of the tubules, nor have they any direct connection
386 S. HATTA:
with the aorta” (loco cit., p. 213). The glomerulus figured in
his Taf. 17, fig. 12 (gl) corresponds, I think, with a part of the
glomerulus in Petromyzon.”
As is very well known, the independent studies of WEI
(90) and Bovert (’92) on the branchial chamber of Amphiozus
gave a new direction to the morphological investigation of this
field. A number of external openings of ciliated tubes is found
at the dorsal corner of the peribranchial chamber of Amphirus,
Through the morphological study and physiological experiments
this organ-system is demonstrated to be the excretory apparatus, |
or “ Nierencanälchen,”’ as Boveri calls them, of Amphioxus
Bovert counted 91 “ Nierencanälchen’’ in an individual 4 em.
in length and possessing 183 gill-bars on the right side. In the
adult he counted about 180 of the “ Nierencanälchen *’; the number
is, however, by no means constant, but varies within a certain limit.
In the middle region of the branchial chamber, a “ Nieren-
canälchen ” has 3 or 4 “ Seitentrichter,” and 2 “ Endtrichter;’
such is the most complete one. It becomes gradually simplified
both anteriorly and posteriorly, until it is at last represented by 4
short single tubule, as seen in Taf. 33, figs. 9 and 13, given by
Boveri. The tubules in the anterior and posterior part of the
system thus show a sign of degeneration, as in the case of the
pronephros of Cyclostomata.
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1) DEAN has published two papers on the development of the Californian Hag (‘98 and
99); these excellent works contain merely the general account of the course of the develop
ment in surface view. We may expect that the full account will throw much light on the
ontogeny of Craniota. There stand, in the account given by him in these works, the im
portant facts that the “pronephric tubules are apparent in connection with all the m=
blastic somites” (98, p. 274) and that the pronephros is extended far backwards, beyond te
anal region, into the tail (’99, p. 272). It would be highly desirable to observe the pronephric
tubules of the Hag in relation to the wyomere, and not to the spinal ganghia alone, as PRICE
has done.
MORPHOLOGY OF CYCLOSTOMATA. 387
These nephric tubules receive, according to Bovert, the blood
from the aorta, which gives two branchlets to each nephric
segment. These branchlets form in each segment a network in
the neighbourhood of, and winding around, the nephric tubule ;
it is this network that Bovert calls glomerulus.
From the structure, the position, the segmental arrangement,
the physiological function, and the relation of the blood- vascular
system to this system of organs, Boveri regards the latter
as a primitive form of Vertebrate nephric organ and homologised
it particularly with the pronephros of Craniota. The points
of difference which exist between the “ Nierencanälchen ” of Am-
phioxus and the pronephros of Craniota, have been smoothed away
by the author’s masterly arguments. The first of these points
is the want of the segmental duct in Amphiotus ; but this is re-
presented, according to Boveri, by a part of the peribranchial
chamber. The second is the relation of the nephric segments to
3
other systems of organs. The “ Nierencanälchen ’ is branchio-
meric while the pronephros of Craniota is myomeric, in arrange-
ment. But this difference is looked upon by him as only apparent ;
for the number of gill-slits first formed agrees with that of the
muscle-segments in the same region; this is sufficiently demons-
trated by the figure given by Weiss (loco cit., fig. 3).
Thus the author has brought the ‘‘ Nierencanälchen ” of
Amphiozus into perfect harmony with the pronephros of Craniota.
Some additional light is now, I believe, thrown from the side of
Craniota by the facts obtained in Cyclostomata, the lowest
class of Craniota. This harmony will be brought out more in
discussing tle development of the pronephros in Selachia, Teleostei,
and Amphibia, which will be treated further on.
TB oe
_
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A i. a i ah lg eg Be es
-
on *
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CES
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cire Aigle SEL
388 S. HATTA:
Thanks to the labors of many eminent investigators, the
early development of the Selachian pronephros has been, as is well
known, fully studied, so that the facts gathered from this field
are well adapted to be compared with those from other groups,
I have found, in the present investigation, many important points
running parallel with the development of the Selachian pro-
nephros. I may then be allowed to compare my own results ın
Petromyzon with those already arrived at in Selachia. Reference
will, however, be limited to those works which are sufficient to
verify the points I wish to bring out.
Through the excellent work of Ruckert (88) we can best
learn the origin of the pronephros in Selachia. “ Die erste Anlage
der Vorniere”’ is recognised “in Form einer gegen den Ectoblast
gerichteten Vorbuchtung des parietalen Mesoblasts.”’ This Anlage
is first brought about by the thickening of the parietal layer of
the mesoblast, which is found “in den Bereich des segmentirten
Mesoblasts, d.h. Somiten ”’ (p. 209) ; this thickening is called by
the author “‘ Segmentalwulst.”’ The foot-note also runs as follows:
“Der Ursprung des Segmentalwulstes reicht ventral bis zu der
Stelle herab, wo die Somiten in den unsegmentirten Mesoblast
der Peritonealwand übergehen ” (p. 209). The “ Segmentalwulst ”
is so called because it is noticed as the segmental thicken-
ing of the parietal mesoblast of which Ruckert recognised,
in his Stad. 11”, six for Torpedo and four for Pristiurus, stretching
over a corresponding number of the myotomes. The first indication
of the pronephros is expressed, in Selachia also, segmentally ın
the segmental part of the mesoblast at the stage in which the
metameric segmentation of the mesoblast is still going on, and
— ee
ee —
1) The embryos in the stage have 25-27 somites.
MORPHOLOGY OF CYCLOSTOMATA. 389
the myotome is not yet cut off from the lateral plate, just as in
Petromyzon. The foremost of them is found in the hind part of
the third or fourth body-somite. The development of the Anlage
in each segment agrees also with that in Petromyzon; for he says:
“Der Segmentalwulst zeigt in vorliegende Stadium (Stad. 1)
regelmässig die stärkste Entwicklung in seiner mittleren Ab-
schnitt, also ungefähr im Bereich des dritten des ihm gehörigen
Somiten, und verjüngt von da allmählich nach seinem vorderen
und hinteren Ende zu............... ” (loco cit., pp. 210).
The account given by RüCKERT is essentially confirmed by
later investigators such as van WYHE (’89)” Rast (’96), and others,
although they differ from one another in the interpretation of the
facts and in some unimportant points. Rap looks upon the
Anlage of the pronephros (his Vornierenwulst) as the ventral
portion of the somite just as RUCKERT does, while it is, ac-
cording to van WYHE, the product of the lateral plate (his
Hypomer). This is,. as it seems to me, not a contradiction in the
facts, but in the terms used ; for van WYne states: “ Da nun der
Pronephros, wie spiitere Entwicklungsstadien zeigen, ein Produkt
der Seitenplatte ist, wihrend der unmittelbar dorsal davon liegende
Theil des Mesoderms zur Mittelplatte gehört, ist die Segment-
irung des Mesoderms bei Selachiern also nicht auf die Myotom-
platte beschränkt, sondern erstreckt sich auch auf die Mittelplatte
und den dorsalen Theil der Seitenplatte ’’ (/oco cit., pp. 474-475).
The fact is, therefore, no other than that the portion of the meso-
blast dorsal to the ventral limit of the Anlage of the pronephros
undergoes segmentation, and the portion ventral to this point
remains unsegmented, constituting the lateral plate. I will, in
1) The embryo, in which the first traces of the pronephros is seen, is, according to
VAN WYHE, in a stage with 27 somites, whereas RABL has seen in an embryo of Pristiurus
with 23 somites.
u
390 8, HATTA:
this place, not go further, but return in future pages
discussion of this point. It is, however, safe, I believe, t
this portion of the mesoblast as a part of the somite.
Van Wyue found the foremost pronephrie segmen
third body-somite (his Rumpfsegment), and Rasr states
Vornierenwulst begins in the seventh somite form
Gesammtsegment). According to Rann, however, van
third Rumpfsegment corresponds to his seventh Gesammi
To verify this fact Rapr has extended the compari:
Petromyzon, and found that in this case also the pr
begins in the seventh somite; but the pronephrie tubul
somite is, as noticed above (p. 380), not the anteriormo
tubules in his sense, but of the persistent tubules.
Van Wryne noticed five of the pronephrie segn
Raja, and three for Seyllium and Pristiurus ; while Rast
eight Vornierenwülste for Raja, and four for Pristiur:
results in Petromyzon, therefore, best agree with those
by Rückerr in Torpedo.
The authors agree in deriving the collecting duet
lateral extremities of the pronephrie Anlagen, whi
become confluent.
Ruckert has observed, in Pristiurus, as well as in
the secondary connection of the Segmentalwulst with the
which has led him to believe in some contribution of |
cells to the formation of the pronephros, while va:
and Ragr deny this. I have found the same conn
Pelromyzon, but I have found no sign of the cor
of epiblastic cells to the formation of the pronephr
phenomenon is temporary in both Selachia and )
it takes place in Selachia, according to Rückerr, in hi
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MORPHOLOGY OF CYCLOSTOMATA. 391
already in his Stad. 111, a space is seen between these two
ctures. | |
The degeneration of the tubules in Selachia runs a course
lel with that mentioned under Amphiorus and Cyclostomata.
seen above, the Anlagen of the pronephros are developed most
rously in the middle part of the pronephros, as in the case
imphiorus and Cyclostomata ; and degeneration begins at the
ial and caudal extremities as there.
Van WYHE says that the degeneration consists in a confluence
rschmelzung) of the ostia. According to RUCKERT, the
nierenfalte becomes simply flattened out in the cranial part
je pronephros. The reduction in the caudal part is noteworthy:
Anlagen are here constricted off from the mesoblast and con-
ed into the anteriormost section of the segmental duct.
y the middle (the third) diverticulum (in Torpedo) persists
ommunicating with the body-cavity and becomes the ostium
miinale.
In Petromyzon, I have unfortunately failed to observe accu-
ly the manner of degeneration of the tubule in the cranial
It is however probable that it begins either from the
d tip of the tubule (the first tubule), or by obliteration
he nephrostome (the second tubule). In the caudal part, the
cting duct is constricted off from the lateral plate by
eration of the tubule and constitutes the foremost section of
segmental duct, in precisely the same manner as in Selachia.
difference is: in Petromyzon the communication with the
y-cavity is retained by the three middle nephrostomes, while
selachia, it is through only the middle one, that is, the osfium
minale.
The segmental duct becomes apparent in an embryo with
392 Ss. HATTA:
35 (VAN WYHE), or 34 to 35 (Ras) somites. The anterior
small section of the duct is formed, as just stated, in the same
manner in Petromyzon and Selachia. The mode of formation of its
posterior larger portion in Selachia differs from that of Petromyzon.
RückERT (’88) and van Wvue (’88, ’89, ’98) believe that
it is the product of the epiblast”, while Rast maintains its
purely mesoblastic origin. At any rate, the posterior tip of the
duct or the cord is sharply pointed and connected firmly
with the epiblast throughout its growth until it opens into the
cloacal cavity, which is effected, according to van Wvyae and
Rast, in the embryo with 83 to 84 somites. It can be inferred
from van WyHE’s figs 7a and 75, that this communication is found
in a plane vertical to the thirty-eighth Rumpfsegment”. In
Petromyzon, the duct, being formed of a series of abortive pronephric
tubules, has no genetic relation to the epiblast except in the
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cloacal region where the duct seems actually to receive cells from
the epiblast, as fully stated above (p. 366).
The nephric arteries of Selachia which were discovered by
Pıur Mayer without reference to their relation to the pronephros,
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nephric fold, but throw a solid process, the interior of which
consists of round or spindle-shaped cells. This is, according to
RuckertT, the equivalent of the pronephric glomerulus of Au-
phibia described by FüRBRINGER. The development and decline
of these vessels go on parallel with those of the pronephric diver-
1)1 will return to this point again in future pages.
2) According to RABL’s counting, this somite corresponds to his forty-second somite.
MORPHOLOGY OF CYCLOSTOMATA. 393
a. The vessels in the cranial as well as in the caudal part
he pronephros are weaker than those in the middle (the third
fourth) ; only the latter vessels develop further and become the
line artery. Van Wyue confirms Ruckert’s account and
described three vessels in Prestiurus. In addition to these,
WYne has pointed out the very small segmental vessels on
left side, which go not to the intestine, but to the body-
They are not equivalent to the intestinal vessels on the
site side. One of them gives a branchlet to the glomerulus
+h sends out, in its turn, a branchlet to the cardinal vein.
homologous vessels on the right side are to be seen coming out
he root of the vitelline artery. BovErı remarks that the
ls of Pauz MAYER present many points of harmony with
branchial vessels in Amphioxus. Rast agrees essentially with
account given by RÜCKERT and van WYHr, but denies the
ence of a glomerulus. According to Rast, the structure
d the glomerulus by Rucxert does not fulfil the condi-
y of being a glomerulus; he says: ‘‘ Eine einfache Ausbucht-
einer Arterie ist noch keine Gefässschlinge, geschweige
ı ein Glomerulus ” (loco cit., p. 668).
Most of the early investigators, who observed the develop-
t of the Teleostian pronephros, believe it to be mesoblastic
rigin. ‘There are very few writers as RYDER (’87), and Brook
), who derive the segmental duct from the epiblast. According
YELLACHER (’73), GOETTE (’75 and ’88), FÜRBRINGER (’78),
Horrmann (’86), the first Anlage of the pronephros is
ight about by the evagination of the parietal layer of the
blast at the level of the junction of the somite with the
‘al plate, forming thus a longitudinal groove on each side,
394 8. HATTA:
which is subsequently constricted off from the body-cavity.
This takes place at first in the middle region of the body, whence
it proceeds both anteriorly and posteriorly.
OELLACHER observed that the Anlage is converted into a
longitudinal canal or the segmental duct, being completely shut
off from the body-cavity in both the anterior and posterior
parts. The anterior section of the duct is much swollen and
transformed into the pronephric chamber. From the dorsal
aorta, a pair of branches is given off which pushes into the
pronephric chamber, pressing against its median wall and giving
rise to a pair of the glomeruli. This portion of the duct becomes
coiled up and constitutes the pronephros.
GoETTE’s view somewhat differs from the account given
above: the anterior end of the longitudinal groove is not com-
pletely closed from the body-cavity, but leaves awhile the com-
munication with the latter, which is, according to GoETTE, the
morphological equivalent of the nephrostomes of the Amphibian
and Petromyzon pronephros. Opposite this nephrostome, he
says, the glomerulus is formed by evagination of the visceral
peritoneum and projects freely into the body-cavity. This portion
of the peritoneum together with the nephrostome is constricted
from the rest of the peritoneum; the coelomic cavity thus shut
off is converted into the pronephric chamber.
This view is essentially confirmed by subsequent writers such
as FurBRrINGER (’78), Horrmann (’86), and others, although
Horrmann differs in his view of the mode of the formation of
the glomerulus.
According to the results recently arrived at by FELıx” in
———
1)I know his paper only by the abstract in the Jahresberichte über die Fortschritte der
Anatomie und Physiologie, N.F. Bd. III. ’97.
MORPHOLOGY OF CYCLOSTOMATA. 395
the embryos of Salmonidæ, the earliest traces of the pronephros
consist, in embryos -with 11 pairs of the somites, of five solid
proliferations of the lateral plate which is already cut off from
the somite. These proliferations, being coincident with the caudal
half of the third to seventh somites, are strictly metameric
in arrangement and are regarded by the author as the rudimentary
pronephric tubules. These tubules soon become confluent with
one another to form a single outgrowth of the lateral plate, which
is called by the author the “ primäre Vornierenfalte.” The
“ primäre Vornierenfalte,’’ which passes over into the parietal and
visceral layers of the lateral plate, undergoes a longitudinal
constriction (the “sekundäre Vornierenfalte”) by which it is
divided into the dorsal and ventral parts. From the former,
the anterior section of the segmental duct originates, while the
latter is transformed into the pronephric chamber. By stages,
the dorsal part wanders laterally, and the ventral part travels
medianwards. At the same time, these parts are separated from
each other, leaving the communication at only -one point, which
is called the “ Pseudonephrostom.”
This phase of the development of the pronephros observed
by FELIX is, as I believe, undoubtedly earlier than that looked
upon by the previous authors as the earliest indication of the
pronephros.
At the time when the Anlagen of the pronephros are con-
verted into the “ Vornierenfalte,’” the Anlage of the caudal contin-
uation of the segmental duct, becomes apparent in the eighth
to the tenth somite; it is brought about by the division of the
primary lateral plate (lateral plate in the ordinary sense) into (1)
the secondary lateral plate (lateral), (2) the segmental duct (middle),
and (3) the Anlage of the “ Stammvenen ” (median). This pro-
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cess proceeds posteriorly until the duct comes to lie close to the
rectum (Enddarm).
FeLıx thus observed the segmental Anlage of the pronephros
in its glandular part, and derives the rest of the system from
the proximal margin of both the parietal and visceral layers of
the secondary lateral plate ; he has observed neither the posterior
growth, nor the epiblastic origin, of the segmental duct.
The pronephric chamber which results from the confluence of
the five pronephric tubules, is not homologous, according to
FELIX, with that of Amphibia, in which the chamber should be
a constricted part of the body-cavity into which the tubules
open.
Quite recently, Swaen and Bracuet (99) have published a
paper on the early development of the mesoblastic organs in
Salamonide. Although my manuscripts were nearly finished,
when I saw this interesting paper, I must here refer in a few
words to it”.
The authors found the first traces of the pronephros under
the fifth somite, of two embryos, one of which was in the stage of
11 somites, and the other of 13 somites. It is not the product
of the parietal layer of the lateral plate only, but is formed,
as FEezix believes, by the proximal portion of both the parietal
and visceral layers of the secondary lateral plate (l’extremé interne
de la plaque latérale secondaire”). The internal cavity enclosed
by the pronephros is, therefore, not the diverticulum, but a
part of the body-cavity.
ae nT a
1)1 am much indebted to my friend, Dr. A. Oka, who read the paper for me.
2) According to the authors, the “ plaque laterale primitive” is divided into the “ plaque
latérale secondaire” and the “masse intermediaire;” therefore, the “plaque laterale
secondaire ” corresponds to the lateral plate itself of Petromyzon.
MORPHOLOGY OF CYCLOSTOMATA. 397
The Anlage of the pronephros is laid in exactly the same
ner from the fourtli somite to the cloacal region. Under the
rior three somites from the fourth to the sixth, the Anlagen
developed into the pronephric chamber ; the Anlagen posterior
hese are all transformed into the ‘ canal excréteur,’’ as they
the segmental duct, and they have come to the conclusion
9
the ‘‘ canal excréteur ”” of the pronephros has the morpho-
al value of a rudimentary pronephric chamber.
The facts given in the last two papers, are thus in close
rdance with one another as well as with those given by
elf in the foregoing pages. Differences between their
ts and mine are that the authors derive the system from the
al unsegmented mesoblast, and that both the parietal and
ral layers of it partake in the formation of the system. As
been stated in the descriptive part, this derivation is only
rent; a little further study shows that only the parietal layer
3 rise to the system, and this part of the layer belongs to
somite. Indeed, this part appears to form, for some time, the
imal portion of the lateral plate, being early cut off from
rest of the somite. It must be remembered that this separa-
is not the separation of the lateral plate from the somite,
that of the Anlage of the pronephros from the rest of the
te; or, the result of the development of the pronephros. It
erely for a physiological reason that this development or
ration of the pronephric Anlage goes on earlier than, for
ince, in Selachia, it performing in Teleostei the actual ex-
ry function. This will be understood easily, when a com-
son with other groups is made further on.
It has been a well known fact that the development of
398 8. HATTA:
Amphibia shows, in several respects, a parallel course with that
of Petromyzon. Careful observations on the development of
Amphibian pronephros, adduced by recent investigators, have
intensified this similarity with the exception of a few points which
are, however, probably of secondary importance.
Most authors who have worked on the Amphibian develop-
ment agree in deriving the entire system of the pronephros from
the parietal layer of the mesoblast only, and in regarding it as
arising originally as a common pouch, the anterior part of
which is divided secondarily, by a partial closure of the
peritoneal communication, into a number of the pronephric
tubules.
This view has been advanced by earlier authors such as
W. Mürer (’75), GoETTE (’75), FÜRBRINGER (’78), Horrmans
(86), and others. The stage at which the pronephros appears
coincides exactly with that in Petromyzon, as Max FURBRINGER
says in his well known work: “Die erste Entwicklung der
Vorniere und ihres Ausführungsganges findet hier nach der
Scheidung des Mesoderms in Urwirbel und Seitenplatten statt
und folgt unmittelbar der beginnenden Sonderung der ersten in
einzelne Urwirbel amd der Spaltung der letzteren in Haut- und
Darmfaselplatten. Embryonen von Rana temporaria von cirea
2.5 Mm. Länge und von Triton alpestris von ca. 2.0 Mm. L.
entsprechen diesen Stadium ”’ (p. 3)”
Mozrrer (’90) has made out the segmental Anlage of the
Amphibian pronephros, having worked with the embryos of Triton,
1) The .nephrostomes are found, according to the author:
2 in Salamandrina maculala, 3 in Rana temporaria,
2 in Triton alpestris, 3 in Bombinater ignens (GOFTTE) and
2 in Siredon pisciformis, 4 in Cbecilia rostrata (SPENGEL).
MORPHOLOGY OF CYCLOSTOMATA. 399
Bufo, and Rana. His accounts confirm, as a whole, those given
by Rückerr for Selachia above referred to, but differ some-
what from those of most other authors who have worked on
Amphibian pronephros. MOoLLIER states as follows: “ Wir sehen
hier ebenfalls zuerst eine solide, von deın Mesoblast ausgehende
Anlage, deren Structur anfänglich schwer zu erkennen ist und
erst mit dem Hohlwerden, wie bei den Selachiern, klar hervor-
tritt. Dann finden wir, dass hier zwei resp. drei getrennte
Canälchen vorhanden sind, die von den Somiten in conver-
genter Richtung ausgehen und erst nachträglich untereinander
vereinigen zu einem Längscanal, von dem aus die Vornierent-
richter in die Leibeshöhle führen ”” (loco cit., p. 229). The
author derives in this wise the pronephric tubules, exactly as
in the case of Petromyzon, from the segmented part of the
mesoblast only.
MoLLIER’s accounts are for the most part in close accord
with the results given by Frezp (’91, p. 282), who, one year
later independently of MoLLıErR, began with Anura, and
extended the work over Urodele Amphibia. In one point, their
?
results differ widely ; but “the difference is,’ it seems to FreLp,
‘apparent rather than real.” According to Monier, the
nephrostomes communicate with the cavity of the myotome, the
myoccelome of van WYHE; this is denied by Frezn, who believes
that “the pronephric tubules have to do with the ventral seg-
ment of the mesoderm” (loco cit., p. 283). It seems to me
LE
that this ‘‘ ventral segment of the mesoderm ”’ corresponds to the
pronephrotome of van WYHBE in Selachia or to “‘ l’extremé interne
de la plaque latérale secondaire’? of Swaen and BRACHET in
1)It seems that the earliest traces of the pronephros are perceived in an embryo
younger than that with 7 somites.
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Teleostei, and I agree with the view of Ruckert, here represented
by that of MoLLIER.
MoLuiER and FIELD agree with each other in assigning three
pairs of the tubules for Rana and Bufo, extending from the
second to the fourth somite, and two pairs for Trilon (MoLLiıer)
and Amblystoma (Frexp), covering the third and fourth somites."
In ‘addition to these, MOLLIER observed occasional occurrences of
the third tubule in Triton, which is, according to FIELD, not
equivalent, as MOLLIER maintains, to the third tubule in Bufo
and Rana, because the additional third tubule in Z7i/on is found
in the fifth somite, while the third tubule in Rana and Bufo is
under the fourth somite. According to Semon, there are ten
pairs of the tubules on either side of the body in J/chthyophis.
A pair of glomeruli has been made out in Amphibia;
the structure is connected by special vessels with the dorsal
aorta on one hand and with the cardinal vein on the other.
This branch of the aorta is believed by FiELD to correspond
to a part of Mayver’s vessels in Selachia. Beside these, there is
no vessel arranged segmentally or otherwise.
The section of the body-cavity corresponding to the pro-
nephric stretch is gradually exparided, and is shut off temporarily
from the rest of the cavity by a close contact of the parietal
and visceral layers of the ccelome; this part of the cavity is,
according to GOETTE, homologous with the pronephric chamber
in Teleostei and with the homologous structure in Peiromyzon,
which is called by him the “ peritoneale Scheidewiinde.”’
The so-called ventral portion of the Amphibian pronephros
is, according to MoLLIER, brought about by the separation of
ne A — —
1) According to Fre.p, MorrtEr's first body-segment in Triton corresponds to his third
somite in Amblystoma.
MORPHOLOGY OF CYCLOSTOMATA. 401
the ventral portion of the pronephric Anlagen from the dorsal,
which latter is differentiated into the tubules and constitutes the
dorsal portion of the pronephros. The ventral part of the An-
lagen separated from the dorsal retains anteriorly its connection
with the anteriormost tubule and posteriorly with the segmental
duct. It is prolonged and bent out anteriorly in front of the
?
dorsal part. ‘‘ MoLLIER’S description is,” FIELD says, ‘“ sub-
stantially in accord with my own observation,”......... (loco cit.,
p. 286). This feature of the duct shows, it seems to me, a close
resemblance to the anteriormost section of the Teleostean segmental
duct which is, as above referred to, bent in the same fashion.
The segmental duct arises, according to previous writers, as
a longitudinal common furrow oft he parietal peritoneum, which
furrow is later constricted off from the mother-layer and becomes
converted into a long canal. MoLLiER has observed the segmental
duct transformed directly from the mesoblast, just like the
glandular part of the pronephros, in the two somites behind the pro-
nephros. Whether the greater remaining part of the duct is formed
likewise by differentiation of the mesoblast, or by a backward
growth of the hind end of the duct first formed, he could not decide
with certainty ; but the observations of FıeLo elucidate this point.
‘The segmental duct arises,’ FıELD says, ‘‘ throughout its
entire length by a proliferation in situ of the somatopleure ”
(loco cit., p. 223). The author has observed neither its epiblastic
origin nor a free growth of its posterior end, except in the cloacal
region where it “ grows across the cloaca free from adjacent tissue ”
(loco cit., p. 223). In Stage v, the cloacal opening is seen. This
opening is found, in Rana and Bufo, in the vertical plane with
the middle of the twelfth somite, whereas it is below the twentieth
somite in Amblystoma (FIELD).
402 S. HATTA:
The duct is segmental in origin. FıELD says: “TI believe
I am justified in concluding that the segmental duct between
Somites v, and IX, arises in situ from a thickening of the somato-
pleure serially equivalent to that from which in the anterior region
the pronephros is developed ” (doco cit., p. 219). There are no
other Vertebrata which agree more with Peéromyzon with reference
to the development of the segmental duct, than Amphibia. Indeed,
here as there, the segmental duct is of segmental origin and is to
be looked upon, as seen in Petromyzon, as the continuation of
a series of abortive pronephric tubules in the posterior region.
Authors who have observed the epiblastic origin of the
segmental duct in Amphibia are very few. von PERENs1 (87)
has published the results of his study on Rana esculenta, but his
note is unfortunately very short”. This view is opposed, so far
I am aware, by almost all recent observers. After bringing
the results by him into harmony with those by Rucxerr in
Selachia, MOLLIER says : “ Im einen Punkte weichen die Amphibia
von Selachiern ab, dass die Vorniere mit dem Ektoblast in
keine nähere Beziehung tritt. Allerdings heftet sie besonders in
den Stadien, in welchen sie voluminöser erscheint, dem Ektoblast
oft in affallend inniger Weise an. * * = Doch lässt sich
stets eine scharfe Greuze beiderlei Blätter ziehen, wenigstens bei
Bufo, wo die Ektoblastelemente durch ihren Pigmentgehalt deut-
lich gekennzeichnet sind ’’ (loco cit., p. 229).
The historical review undertaken in tlıe foregoing pages
shows the agreement to a large extent of the results arrived at
in several groups of Anamnia. Some points of disagreement are
naturally met with; but these are, I believe, only apparent.
1)I have not seen the paper by Brook.
MORPHOLOGY OF CYCLOSTOMATA. 403
In the groups above referred to, the first indication of the
excretory system becomes apparent at a stage in which some
mesoblastic somites are formed and the metameric segmentation
of the mesoblast is going on. This is the Anlage not of the
segmental duct, but of the pronephros. A single exception is found
in Bdellostoma, in which the early traces of the system become
visible, as we learn from PRICE, at a stage much more advanced
than in other Anamnia, that is, at the stage in which the sclero-
myotome is cut off from the rest of the mesoblast and mesen-
chymatous cells fill up the spaces between organs and organ-systems.
I have endeavoured to reconcile the points, in which the
views of the previous authors diverge from one another, under
the following three headings :—
A.—The Anlage of the Fronephric Tubule is the Product
of the Hesoblastic Somite and not of the Lateral Plate.
The view that derives the pronephros from a single common
groove formed either of only the parietal, or of both the parietal
and visceral layers of the unsegmented mesoblast (the lateral
plate), is advocated by most of the authors who have worked
on the development of Zelromyzon, Teleostei, and Amphibia.
This is due probably to the early separation of the sclero-myotome
from the rest of the mesoblast in these groups. In them thie
Anlagen of the pronephros (or the nephric segments) together
with the lateral plate are cut off from the sclero-myotome and
form, for some time after this separation, the proximal portion
of the lateral plate. It must be borne in mind that this se-
paration is not the separation of the lateral plate, but of the
nephrotome, from the sclero-myotome. This is, therefore, a
404 8. HATTA:
step in the differentiation of the mesoblastic somite, and because
the distal (ventral) portion of the latter happens for a time to be
continuous with the lateral plate, we are not justified in concluding
that it is derived from the lateral plate, which, as we know, never
undergoes segmentation.
It is a significant fact that in Selachia and Amniota, in
which the pronephros does not function as the actual excretory
organ, this separation of the mesoblast into the sclero-myotome
and the nephrotome is not effected so early as in the above
groups, but takes place only at later stages, with the first dif-
ferentiation of the mesonephros. This consideration makes it
reasonable to conclude that the early separation of the mesoblastic
somite into the proximal and distal portions is caused by
physiological necessity and has no morphological significance”.
The case of Lacerta agilis is very instructive. According to
Horrmann (’89), the Anlagen of the pronephros in this animal
are, in the most anterior segment, cut off from the myotome
(sclero-myotome) and remain connected with the lateral plate
just as in Petromyzon, Teleostei, and Amphibia; whilst in all the
following portion, they are the actual diverticula formed segment-
ally in the parietal layer of the lower part of the somite, as in
other Reptilia (pp. 264 and 265). We thus see the two modes
of separation in one and the same animal.
All recent authors agree in thinking that the Anlage of the
pronephros is expressed in itself segmentally and is strictly
myomeric. Now the question arises: How many parts are to be
distinguished in the mesoblast, and to what part of it does the
Anlage of the pronephros belong ?
Van WyueE (’89) has distinguished, in Selachia, three por-
1) This view is grönnded upon the suggestion of Pror. MITSUKURI.
MORPHOLOGY OF CYCLOSTOMATA. 405
tions of the mesoblast which are called by him the “ Epimer,”’
“Mesomer,’’ and ‘“ Hypomer’”’ respectively. The epimere of
VAN WYHE corresponds solely to the myotome; his mesomere
comprises the Anlage of the mesonephros and the sclerotome ;
and the hypomere consists of the Anlage of the pronephros,
the genital gland, and the lateral plate. The epimere, mesomere,
and the dorsal part of the hypomere undergo the metameric
segmentation, while the remaining portion of the hypomere
remains unsegmented. According to van WYHe, the dorsal seg-
mented part of the hypomere is, therefore, the product of the
lateral plate (see p. 389). It seems to me that this division of
the mesoblast does not agree with the facts observed in Petromyzon
and the other Anamnia above referred to; for the mesoblast in
these groups consists, in early stages, of two portions: (1) the
segmented, and (2) the unsegmented, and of nothing more, just as
RucKkERT (’88) and Rast (’88, ’96) have remarked. According to
Rvuckert and Rast, the segmented portion—the somite—com-
prises the myotome and the sclerotome ; the pronephros and the
mesonephros are derived from ita ventral (distal) portion, which is
called by Ruckerr the ‘‘ Nephrotom ” (’88, p. 272).
In Petromyzon, these two portions of the mesoblast, the
segmented and the unsegmented, are, in early stages, clearly dis-
tinguished, being histologically different (see p. 315). The meso-
blast in such an undifferentiated state is almost entirely occupied by
the segmented portion, while the unsegmented portion is very
small, being represented by the loose tissue of a few cells. Such a
mesoblastic segment exactly corresponds to the somite of RuckERT
and Rast. The proximal half of the segmented portion coincides
with the sclero-myotome of those authors. It consists not only
of the myotome, but also includes the sclerotome, And it is the
406 S. HATTA:
distal half of this segmented portion which folds out in each
segment to give rise to the Anlage of the pronephric tubule on
one hand and to the cœlomic projection on the other, and,
of Ruckert.
’
therefore, corresponds to the “ Nephrotom ’
The nephrotome, therefore, constitutes, in both Petromyzon
and Selachia, precisely the same part of the mesoblast, viz. the
distal (ventral) portion of the somite, through which the sclero-
myotome is connected with the lateral plate.
The above early stage in the differentiation of the mesoblast,
in Petromyzon corresponds also to the “ Ursegment ” of Am-
phiozus in HATSCHEK’s sense (’88). By further development of
it the unsegmented mesoblast is brought into light, and we can
then distinguish the “ Urwirbel” and the “Seitenplatte” of
HATSCHEK ('88). And the ventral half of the Urwirbel constitutest
in Petromyzon, the connecting canal between the unsegmented
celomic cavity and the sclero-myotome, that is to say, the
nephrotome. Let us now examine what part of the Ursegment of
Amphiozus represents the nephrotome of the Craniota. -
In his excellent work on “Die Nierencanälchen des Am-
phioxus,” Boveri ventures to solve this important question.
After a discussion he comes to the conclusions:
(1) That the ‘ Gononephrotom ” in Craniota must correspond
to a part of the “ Urwirbel ” of HATSCHEK ;
(2) That the “ Gononephrotom ” of Craniota is homologous
with the genital chambers of the adult Amphiozus.
But these chambers “sind ursprünglich die segmentale
Verbindungscanäle zwischen der unsegmentirten Leibeshöhle und
der Sclero-Myotom gwesen ”” (’92, p. 493). I can, therefore,
ascribe no other significance to the ventral half of the segmented
MORPHOLOGY OF CYCLOSTOMATA. 407
mesoblast in Lelromyzon than that it is the morphological
equivalent of the ‘‘segmentale Verbindungscanäle.”
It thus follows that the distal half of the segmented mesoblast
in Petromyzon undergoes exactly the same fate as that in Am-
phiorus : it is transformed into the pronephros and the celomic
projection or the “ dorsal segmental ccelome,’’ which latter gives
rise, just as Boveri suggests in Amphiorus, to the mesonephric
tubules and, in the hinder region, to the genital gland.
As has been pointed out in the historical review, the rela-
tion of the Anlage of the pronephric tubule to the mesoblastic
somite is the same for Teleostei and Amphibia, as in Petromyzon.
It may, therefore, safely be stated, that the segmented portion
of the mesoblast constitutes in these groups a single integral
structure until the separation of the nephrotome in continuo with
the lateral plate from the sclero-myotome. This separation is, as
above stated, not the separation of the somite from the lateral plate,
but the differentiation of the somite into the sclero-myotome and
the nephrotome, preparatory to the development of the urogenital
system. The reason why the separation takes place earlier in some
groups than in others, rests only on physiological grounds.
B.—The Whole System of the Pronephros of Cyclostomata,
Teleoslei, and Amphibia is Homologous with the
Nierencanälchen of Amphioxus (BovErt) and
not perfectly Homologous with the Selachian
Pronephric System.
I have already stated above (pp. 386 and 387) that Boveri
has brought the pronephric system of Craniota in harmony with
the system of the ‘‘ Nierencanälchen ”’ of Amphiorus, basing his
arguments on the structure, the position, the myomeric arrange-
408 S. HATTA:
ment, the physiological function, and the relation of the vascular
system to the organ. His comparison is, however, almost entirely
limited to Selachia on the side of Craniota, owing perhaps to the
scantiness of the literature at that time. I accept in the main
this homology, and I may perhaps extend this comparison a little
further.
I will begin with the homology of the pronephros of Cyclo-
stomata with the “ Nierencanälchen ” of Amphiorus.
It is well known that the starting point of the hepatie
diverticulum from the enteric canal demarcates, in the Chordata,
the respiratory section of the canal from the nutritious section of
it; and, as GEGENBAUR (’78, pp. 563—581), Barrour”? (’85), and
others affirm, the œsophagus and stomach in the higher forms are a
part of the former section, which is called the fore-gut. And the
homology of the hepatic cecum of Amphiorus with the liver of
the Craniota, has been much strengthened by recent mor-
phological studies and physiological experiments”. The results
of my present study also confirm this view. I will use, therefore,
this fixed point as the landmark of comparison of the two
organ-systems, the pronephros and the “ Nierencanälchen,’ and of
the pronephros in different groups of Craniota.
1) Froin the account of BALFoUR, I will cite the following lines :—
‘In Amphioxus the respiratory region extends close up to the opening of the hepatic
diverticulum, and therefore to a position corresponding with the commencement of the
intestine in higher types. In the craniate Vertebrata the number of the visceral clefts has
become reduced, but from the extension of the visceral clefts in Amphioxus, combined with
the fact that in the higher Vertebrata the vagus nerve, which is essentially the nerve of the
branchial pouches, supplies, in addition the walls of the œsophagus and stomach, it may
reasonably be concluded, as has been pointed out by Gegenbaur, that the true respiratory region
primitively included the region which in the higher types forms the oesophagus and
stomach’ (Vol. 11, p. 758).
BALFOUR has also shown that the solid cord of the œsophagus in Elasmobranchii and
Teleostei, is the remanent of the gill-rudiments in the ancestry (loco eit., pp. 61 and 78).
2) J. A. Hammar, ’99, ’98, and Gcmo SCHNEIDER, ’99.
MORPHOLOGY OF CYCLOSTOMATA. 409
9
The “ Nierencanälchen ” of Amphiozus, according to BovERI
(92), extends over and is limited to, the whole extent of the
branchial region, the posterior larger part of which covers the
hepatic cœcum. The pronephros of Cyclostomata extends
from the anterior body-somite to the cloaca. The anterior section
of the system constitutes afterwards the glandular part represented
by the pronephric tubules and is found in front of, and over,
the Anlage of the liver, or in the region of the fore-gut ; a certain
number (two in Peiromyzon, twenty in Bdellostoma) of the anterior
nephric segments are found in the branchial region, and the
posterior one or two segments of the glandular part (Petromyzon)
cover the liver-Anlage. It follows that the six” to twenty
or more pronephric tubules correspond to as many “ Nieren-
canälchen ”’ in about the middle one third” of the branchial region
of Amphiozus, and that the “ Nierencanälchen ”’ lying posterior to
this point are represented, in Cyclostomata, by a number of the
rudimentary tubules which are converted into the segmental duct.
9
The “ Nierencanälchen ’ are not put in communication with
one another by the collecting duct, as in the pronephros of Cyclo-
stomata, but open to the exterior segmentally. I have stated in
the descriptive part (p. 333) that the free extremities of the
pronephric tubules in Petromyzon are brought into close contact
with the epiblast, so that the latter is pressed out by the enor-
mous growth ofthe tubule and that this is especially the case in
the first and second tubules. This fact throws light upon the
homology of the pronephric tubules in Pelromyzon with the
* Nierencanälchen ”’ of Amphiotus : in other words, the condition
1)The glandular part of the pronephres in Jetromyzon, are represented by the six
pronephric tubules.
2)The branchiomeres in the posterior section of the gill-basket of Amphioius are after-
wards added (see pp. 410-411).
410 8. HATTA:
seen in the ‘‘ Nierencanälchen ” would be brought about, if the
tubules in Petromyzon came to open to the exterior, boring
through the epiblast by the further growth of their free extremity.
This intimate contact of the tubule-end with the epiblast takes
place, as above mentioned, in the middle of Stage 11, where
the tubules have developed a little beyond the mere Anlage.
The pronephric tubules of Petromyzon are, in this stage,
already united with one another by the intersomitic solid cord;
this union is, however, not primary, but secondary. This stage
presents, I think, the phylogenetic stage, in which the “ Nieren-
canälchen ” with separate external segmental openings, and the pro-
nephric tubules with the collecting duct, diverge from each other.
By this assumption, it is not meant that in the ancestry of
Chordata the tubules were closed blindly inside the epiblast ; for
the Anlage of the pronephric tubule might have been, in the
ancestral form too, brought about by the folding of the mesoblast,
to break out finally to the exterior. This perforation would
become unnecessary when the secondary union of the tubules had
been acquired.
Since a certain number of the “ Nierencanälchen ’
of the base of the hepatic cacum, is represented by the pronephric
tubules of the glandular part in Cyclostomata, those lying over
it will be homologous with the pronephric tubules which are
found over and posterior to the hepato-pancreatic Anlage and
converted into the anterior section of the segmental duct, being
secondarily united with one another by the confluence of the free
extremities of the tubules.
There is not to be seen the post-hepatic “ Nierencanilchen ”
in Amphioxus. We learn from LANKESTER (’89) and Wiccey (9)
that in Amphioxus, the new branchial slits are added, by stages,
bf
in front
MORPHOLOGY OF CYCLOSTOMATA. 411
to the posterior end of the pharynx, so that, in later stages, the
coincidence of the number of the slits with that of the myotomes
is lost; and that this addition of the slits continues throughout
life. The nephrotomes in these new slits have been, I think,
originally coincident, in each segment, with the myotomes, from
which they were cut off in early stages and have remained
undeveloped until the new appearance of the added slits. It
seems, therefore, probable that the branchial region of Amphiorus
once extended over the largest portion of the enteric canal,
while a very small section in the posterior part of the canal
performed the nutritious function, as is seen now in the Ascidian”.
The “ Nierencanälchen ”’ in this hinder part may represent the
pronephric tubules in the post-hepatic section of the segmental
duct of Cyclostomata”.
The pronephrie system of Petromyzon comes to have the same
relations with the epiblast as the “‘ Nierencanälchen ” of Amphiozus
at three different points: the free ends of the two anterior
pronephric tubules and the hind end of the segmental duct
(probably the hindmost pronephric tubule). Whilst in the greatest
section of the system the communication with the exterior has
been lost, these three points might have preserved it to a
considerably later phylogenetic stage: the two anterior tubules
playing the same physiological part as the ‘‘ Nierencanälchen ”
of Amphiorus, and the posterior being employed as the only
excretory pore of the system secondarily established by the union
of the tubules.
In main points (with exception of the presence of the
tubules in the branchial region, of the contact or connection of
——
1) Balfour says: “In Ascidians the respiratory sack is homologous with the respiratory
tract of Amphioxus” (loco cit., p. 758.)
2) See p. 108.
412 Ss. HATTA:
them with the epiblast, &c.), the pronephric system of Teleostei
and Amphibia shows, as stated in the historical review, the same
characters as that of Cyclostomata, so that tbe facts established
in Cyclostomata have the same significance for the Teleostei and
Amphibia.
Although such is the case in those Craniota and Amphiozus,
the pronephros of Selachia is quite otherwise: the Anlagen of
the pronephros are here formed in the mesoblastic somites posterior
to the Anlage of the liver (see below), and only one or two
segments of them are converted into the segmental duct (RUCKERT).
These pronephric Anlagen in Selachia are, therefore, the mor-
phological equivalent, not of the glandular portion of the pronephros,
but of those which are converted into the segmental duct in the
Craniota just mentioned.
C—The Segmental Duct in Selachia is not the Morphological
Equivalent of the Duct of the Same Name in Cyclostomata,
Teleostei, and Amphibia.
Contradictory views are met with in the derivation of the
segmental duct. The results arrived at in Cyclostomota, Teleostei,
and Amphibia, well agree in making it of the mesoblastic origin ;
there are a few authors who believe in the epiblastic origin of the
duct in these groups, but their papers are not more than mere
notes. In Selachia, the circumstance is reversed; I am not aware
of any recent author other than Rast, who advocates the meso-
blastic origin of the Selachian segmental duct. The facts given by
RaBz are, however, not the same as those observed in the
groups just referred to. In these, as stated above, the duct is
MORPHOLOGY OF CYCLOSTOMATA. 413
differentiated, so to speak, in situ from the mesoblast in its whole
length, and as recent authors agree, is composed of a series of
the abortive tubules formed in each nephrotome. This is not
the case in Selachia ; here it is brought about, as RaBt states, by
the posterior growth of the collecting duct which is formed by
the confluence of the lateral extremities of the pronephric Anlagen.
It is not easy to bring these two widely divergent modes of
formation into harmony with each other.
A few morphological considerations, however, would, I believe,
enable one to derive one type of the system from the other. I
may be permitted to state here some of these considerations.
I will start with the question: Is the segmental duct of
Selachia the morphological equivalent of that of Petromyzon,
Teleostei, and Amphibia? I believe the question can be answered
safely in the negative, if we consider (1) the position of the
pronephros first formed, and (2) the origin of the duct.
In the first place, the pronephros in Selachia appears, as
we learn from Rast, in the mesoblastic somites lying posterior
to the Anlage of the liver; thus the Anlage of the liver lies
under the fourth and fifth somites, and that of the pancreas
under the sixth, while the pronephros covers the seventh to
tenth somites (°96, p. 667). On the contrary, in Petromyzon” the
pronephros originates in the mesoblastic somites anterior to the
hepato-pancreatic Anlage, only the posterior one or two nephro-
tomes covering the liver. Such being the case, the pronephric
segments in Selachia correspond to the same number of the
abortive tubules in Petromyzon and the other Craniota above
1) As we learn from GoETTE (’75) and Oellacher (’73), the anterior section of the
pronephric system in Amphibia and Teleostei, is also found in the mesoblast opposite to the
posterior section of the fore-gut,
414 Ss. HATTA:
mentioned, which are converted into the anterior section of the
segmental duct in these groups. It follows that the segmental
duct in this part of Petromyzon and the two other groups, is
not -the morphological equivalent of the duct of the same name,
but of the pronephros itself, in Selachia.
In the second place, let us consider the mode of growth
of the segmental duct. Whichever view may be taken of its
origin, whether epiblastic or mesoblastic, this duct in Selachia
does not arise segmentally as in the other Craniota just referred
to. It is a backward growth produced either by delamination
from the epiblast, as RuckertT and van Wy8E affirm, or by
cell-multiplication within the structure of the mesoblastic collect-
ing duct itself, as Ras states. Hence it can not be homologous
with the duct of the same name in Pelromyzon and the two
other groups, which is derived segmentally from the rudimentary
pronephric tubules. The Selachian segmental duct is, in its
whole length, represented, as I believe, by the posterior small
section of the segmental duct in Petromyzon and Amphibia.
In Petromyzon, the hind end of the duct comes into an
intimate connection with both the epiblast and the lateral diver-
ticula of the cloaca, filling up the space between them, and fusing
with both of them. We may suppose that the direct communication
of the duct with the exterior, if such truly existed in the an-
cestral history, may have been at this point of the epiblast.
This fused condition of the duct and the epiblast reminds us of
the early stages of the Selachian duct at the stage when it has
been produced only a little posteriorly from the pronephric region.
I believe that if the duct is to be compared in Petromyzon and
Selachia, a stage such as the above ought to be taken. The
largest part of the Selachian duct is represented by a free hind-
MORPHOLOGY OF CYCLOSTOMATA. 415
ward growth formed after such a condition is passed, and its
homologue can not be found anywhere in Petromyzon. I believe
that the same can be stated of Amphibia which is, to judge from
FIELD’s account, very much like Petromyzon in this respect
(see p. 401).
According to van WyHE and Rast, the duct in Selachia
appears in the seventh to the tenth (in Rast’s sense) somites of
embryos with 34-35 somites and, when it is later connected with the
cloacal wall, the connection is found,—as can be inferred from
fig. 7b of van WyHE,—in the thirty-eighth (forty-second of Rast)
Rumpfsegment (or further backwards) of Pristiurus embryos with
80 (van WYxHE) to 87 (RaBL) mesoblastic somites. There is found
in Selachia, therefore, a number of the mesoblastic somites in
the region back of the pronephros, which do not give rise either
to the pronephric tubules or to the segmental duct, and the duct
grows backwards, free from the mesoblast inside, during the
period in which the somites increase from 34-35 to 80-87, and
for the space reaching from the eleventh or twelfth to the forty-
second (in the sense of Rast) somite. Such a considerable
prolongation of the duct during this period is not observed in
Petromyzon” and in the two other groups of Craniota mentioned.
And furthermore, it is questionable whether, during this
posterior growth, the duct in Selachia receives the constituent cells
from the epiblast along its whole length, as RUCKERT and van
WYHE believe; or only at the point of the epiblast overlying the
hind end of the pronephros, with which the duct is connected,
and posteriorly to this point grows free from both the epiblast
a ee ne rn errr rere rr en en
1) At about this stage (Stage iv), there is no space left behind the pronephric system
segmentally formed; for the embryo of Petromyzon, is retort-shaped and has the anus situated
in the ventral median line of the bulb of the retort.
416 8. HATTA:
and the mesoblast. The latter view seems probable to me. The
figures (’89, figs. 5 a-c) given by van WYHE to illustrate his view
of the epiblastic origin of the duct, are from the vertical plane
of the eighth Rumpfsegment of a Scyllium embryo with 37
somites, which corresponds to the twelfth Gesammtsegment of
Ras. The figures (96, figs. 94, 98, 104, and 108) given by Rast
to the negation of van Wyue’s view, are from the vertical plane
of the twenty-second Gesammtsegment of an embryo with the
G3 somites. ‘These two cases are, I suppose, the two ends of the
same duct in different stages; the anterior end, being the
equivalent of the hind end of the segmental duct in Petromyzon,
actually receives cells out of the epiblast, as the figures by van
WYne show; the other end, which is seen in Rast’s figures, is
the point of mere contact with the epiblast, along which it is
shifting backwards.
If the above comparison be correct, the segmental duct in
Selachia 18, except the anterior very small section which is formed
directly of the abortive tubules, not homologous with the duct of
the same name in Petromyzon, but is a structure secondarily acquired.
From the above account, it may be safely concluded that in
its primary phylogenetic stage, the pronephric system of the
Craniota above referred to consisted of a number of segmentally
arranged tubules, which were directly formed, in each mesoblastic
segment, from the distal (ventral) portion of the mesoblastic
somite, and opened independently to the exterior ; that the lateral
extremities of these tubules were afterwards secondarily united
with one another, thus constructing the collecting and segmental
duct, the hind end of which opened directly to the exterior ; and
that the acquisition of an opening of the duct into the cloaca
MORPHOLOGY OF CYCLOSTOMATA. 417
was the tertiary stage of changes in the system. Such a course
of the phylogenetic development of the system is, however, no
other than that advanced by Rückerr (’88, p. 265).
In the present paper, the historical comparison will be
limited to the groups of Vertebrata stated above; the review of
Amniota and some other theoretical considerations will be reserved
to a future paper, in which I propose to deal with the further
fate of the pronephros and the development of the mesonephros
in Petromyzon.
Having compared the results arrived at in the present work,
with those in different classes of Anamnia, I may be justified
in drawing the following conclusions.
In Petromyzon, the first indications of the pronephros be-
comes apparent at a stage earlier than those hitherto regarded
as the starting point, that is, at a stage in which the mesoblast
in the anterior region has undergone the metameric segmentation
but the lateral plate is not yet cut off from the somite.
The tissue giving rise to the pronephros is the parietal layer
of a small section of the mesoblast, which forms the distal
(ventral) half of the mesoblastic somite. This section of the
?
mesoblast exactly corresponds to the “ Nephrotom ” of RUCKERT
in Selachia.
The Anlage of the pronephros in all the groups of Vertebrata
above referred to is produced by the evagination of the parietal
layer of the nephrotome which theoretically ought to contain a part
of the celomic cavity. In Cyclostomata, such a cavity is
418 S. HATTA:
actually present”: in other groups, the Anlagen are mere thicken-
ings.
As the pronephric tubules are derived, in each segment, from
the distal (ventral) half of each mesoblastie somite, the prone-
phros is, from the first, of a segmental arrangement, being
strictly myomeric. In fact, the separation of the sclero-myotome
from the lateral plate is effected on account of the differentiation
of the Anlage of the pronephros or of the nephrotome.
In Petromyzon, the pronephric tubules which constitute the
glandular part of the system and the anterior section of the
segmental duct, are formed in the region of the fore-gut and some
of them are detected in the region where the gill-pouches are
afterwards formed ; these latter disappear entirely before the gills
come into view.
The segmental Anlagen of the pronephric tubules : are
secondarily connected by the duct formed out of two adjacent
pronephric Anlagen and put in communication with one another.
The degeneration of the pronephric tubules takes place from
both the cranial and caudal extremities of the system. In the
cranial part, the tubules disappear without leaving any trace;
while in the caudal, they are converted into the anterior section
of the segmental duct. The remaining part of the system func-
tions for some time as the excretory organ.
The pronephric Anlagen in the hinder region do not develop
beyond a certain point, but are employed solely to give mise to
the segmental duct just as in the somites having degenerated tubules.
From what has been said, itt is, I venture to think, no rash
conclusion to regard the pronephric tubules in Petromyzon as having
once extended over the body-segments from the branchial region
1) According to SWAEN and BRACHET, the same fact is seen in Teleostei.
MORPHOLOGY OF CYCLOSTOMATA. 419
to the cloacal part, and as having been, in the anterior region,
replaced by gills and, in the posterior, converted into the segmental
duct.
In the two anterior segments which belong to the branchial
region, the free ends of the tubules are brought into close contact
with the epiblast but this germinal layer has, in this region, no
share in the formation of the system. The hind extremity of the
segmental duct, however, strikes against the epiblast and has every
appearance of receiving some cells out of it. These facts allow
us to infer that all the tubules once had each an independent
external opening until they were secondarily united with one
another by the intersomitic duct.
The visceral layer of the nephrotome becomes evaginated
medianwards and forms a series of segmental pouches on either
side of the subchorda ; but this feature is temporary, and the struc-
ture is soon smoothed by their becoming confluent with one another.
This series of pouches is, I believe, the remnant of the primitive
segmental celome, and gives rise to the gonads and the meso-
nephros.
If the accounts given above be correct, the primary mesoblast
is, during early development, divided into two distinct portions : (a)
the larger proximal portion which is segmented, and (6) a small
distal portion which is unsegmented. The former is differentiated
into the sclero-myolome and the nephrotome, and the latter forms
sumply the peritoneal linings.
The pronephric vessels acquire their definitive form in much
later stages; when established, they are intersomitic in position.
The posterior part is transformed into a pair of the glomeruli
of the pronephros.
Petromyzon has for a long time been looked upon as being
420 S. HATTA:
peculiar and standing apart from other Vertebrata in the develop-
ment of the pronephros. But the results brought out in the
present work speak for a complete parallelism between this genus
and the representatives of other classes of Vertebrata.
Biological Laboratory,
The College of Peers, Tokyo.
November, 1899.
Postscript.
By the kindness of Pror. WATASÉ, I have been enabled to
look through Dr. WHEELER’s paper on “ The Development of the
Urogenital Organs of the Lamprey ’”’ which has just been published.
I find a general agreement of his results with mine. The most im-
portant point is the discovery of the earliest traces of the pronephros
as given in the foregoing pages. As to the formation of the
segmental duct, his views are somewhat different from mine; this
and some other points of divergence are, as I believe, due to
gaps in his materials. Thus, his fig. 1, which represents section
through an embryo in his Stage 1, corresponds to my fig. 1, while
the next older stages (Stages 2 and 3) in his series, spoken of as
representing GOETTE’S fig. 9 where the heart is already formed,
coincide with the oldest embryo of my Stage ıv. As seen in the
foregoing description, most of the important processes in the
1) Zool. Jahrbücher, Abtheil. fiir Anat. and Ontog., Bd. xm, ’99.
MORPHOLOGY OF CYCLOSTOMATA. 421
development of the pronephros and the segmental duct take place
in this interval of time, which WHEELER has unfortunately
omitted to study. But in the main, his results confirm mine.
This agreement arrived at independently naturally affords a good
evidence of the correctness of the facts given.
422 8. HATTA:
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Fishes,
(85) — :—A Treatise on Comparative Embryology.
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Anat. Anz., Bd. 11.
(92) Boveri :—Die Nierencanälchen des Amphioxus: Zool. Jahrb., Bd. v.
(88) Brook :—Note on the Epiblastic Origin of the Segmental Duct in
Teleostean Fishes and in Birds: Proceedings of the Royal Society of
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(98) Dean :—On the Development of the Californian Hagfish (Bdellostoma
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(’86) FLemming :—Die ektoblastische Anlage des Urogenitalsystems bei
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(97) :—Contributions to the Morphology of Croate I. On
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(89)
(90)
424 8. HATTA:
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(75) SemPEr :—Das Urogenitalsystem der Plagiostomen und seine Beden-
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m
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MORPHOLOGY OF CYCLOSTOMATA. 495
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PLATE XVIL
Plate XVII.
[The magnification is the same for all figures: Cx 2, with the single
exception of fig. 4 which is Ex 2.]
a.pn.1-6, Anlagen of pronephric | m., median row of mesoblast.
tubules from the first to sixth. | m.p., parietal layer of mesoblast,
a.sd., Anlage of segmental duct. | mes., mesoblast.
cd., collecting duct. mt.I, IL, &c., the first, second, &c.
ch., chorda dorsalis. myotome.
cut., cutis-layer of myotome.
d., dorsal row of mesoblast. .
ep., epiblast. m.v., visceral layer of mesoblast.
hy., hypoblast. n., neural cord or canal.
!.m., lateral plate of mesoblast. | v., ventral row of mesoblast.
mus., muscle-layer of myotome.
Fig. 1. A transverse section through the dorsal region of an embryo inter-
mediate between Stages I and 11.
Fig. 2-7. From a series of transverse sections through a younger embryo
of Stage 11.
Figs, 8-17. From a series of transverse sections through an older embryo
of Stage II.
Figs. 18 and 19. Two sections from a series of cross-sections through a little
more advanced embryo than the last.
Figs. 20-29. From serial cross-sections through the most advanced embryo
of Stage II.
1h ees N° CA 3 (Fee
ut
- i e * . LÉ
oe
we See Sat teat ane
À LT ... Nowe, er .., ee __._d. a N ee ..° = i
ye ® .. se \ as y Le DR En 7, ers a en eme - Vi
ey .? < un... r on . *
se ne fee OR Seg ee 0 tn... OP = “e A N ers e
.° s ~— ps @e e \ “ à ote f 2 + e® ® 5 e
# S Sr in ® . L = le 6 \ . . Pr apr <i. _ Id“: . a fe 2 . .
® ba à re ear” 5 ‘ i \ @, im rn ER “ss °° i ie tas
e. d m 7? ¢ \ . e Lg ° -- wer e is cee”? se te AN
RE 2 -_ 5 Gs, 2 (RE 2 + ® ‘ can : - ..
\ # . CR ‘ ‘ > -
\ ‘ . { « @ % \ {
‘ f. + % \ 4
bla e \ '
LV i t. 3
h m. Ù
y 19. ch.
Fig.6
[9.D.
Fta.5
ch? er nn --
ie mt. x e N
°°" 7. > les e
mel. .* apn.i_ tee ne |
x ge Ver ose |
apn ES ! Im. cr Be SL 4 mtb]
= . ---._ n ASS Erg _ 3 =
ears . . = | vs ss.” ST ~
ee ‘A i Br 27 RE
Im. ve ‘ ; a: F : lm.
.o Is .. x / \ se ®
| a? e a *e Pad . » FL _
CR +> °° ee
e _ . eee? BS gue ‘ = a #@
eran Le --} - ch h ..-"
as ch
-
re
Fig. 10 Fig. 11.
À D) mé.vl.
- e ® =
a® © @e ed * 2 Bee >
met we . Les. las SE
. - % Ban Im. a. T . }
Le e È e ® mt...
cd e > Ss. e N ae * \ 3 Siw int
2 *. 1, ep. age x eo, *e a
. u. oF “ev — mite a ae‘ x . * . [u Zu “. Ve: a .
Um e . Pa - > < ne. Om, pri. ae
“te fe .. A ae FER ont N
. Po be o@ * ° . : .. . e
. A P , o,e? . 2 . .. é
. aay ; f. _ u -„ -_«d 1 res à is
EN, 1 es ds “ey ! Tr. u —
“f te, te Im. ch. os ==
ce e e ‘ r
sch RP RE .
.. *
* +
"tg. 15.
« v° «
| ot*%e Vs D
mi, me ae mt 5. Frg. x... mi SF.
er :
' se arm Pi. arzt dd.
e ‘ (Ae ‘ ere e In. , CYR ..
. se ep cut ote a es
ee > eo 8 à at — =. .
im. ee? pee es ; .’ or De at
. e cae CU MTS > = > 6 Pa
„® . * e e ow ry o
Ped e +e et = ni - .ch. 2 N : €,
Pi a dpn.s. sr Br N ae eee
7 “ 2 eo: Pid / ep.
oS ae oe) . 2
oe 6% S .
. ok Crea \* &
e . = * ee à
u D unies wit . .. .
ea ae ‘ . we aed 7 ch l'ta. Lh.
e CR \ + *
lm “* oe \ a
~~ . , eee®
+ \ .. AE
te 2 ? <
fe *, PULLS mi .kiR. ; aS
o, had e
à : a’
big. 20 N
c/s . op ~ a Va
e,¢. =
‘. we 0; U
na. - n lg. 21. +
à e ¢
I u == ep. mt.V. ER e ”_ eh.
.
r . . er . CE „MN.
, ‘ss . .. 3 Au
‘ - ae .. e 4 So)
ur = ses neo Fe cu. + et es ug ive
" , wo... ge =. 1 e..
7 ey. „ex: .* .. “ eC. _-Cp-
u . , a Ss eA %, e {/ eo
LT Suns : ra A 2% . |
| " Pin . 2 à e .", i @e
+ > ta ” e x
1 _ wi > . ® on te /
e og oe ej
# > e / + o* N 7
im + , cd. EN
f ate ee pf
. ee gt Tha Pat ae jee mv. = pn.‘
+ ® D Me Me SH Fre
* 2 © e 4 x %
bs ee PUIS 7
- . N \
' Hs Im ae ee Im. er
N} eo ee Re eg
, el bs eee |
+ © 7 @*. e
| | he PE ER
NET ey ir
ei at200
Jour, Sc. Coll, Vol, XM, Pl. XVI.
reg we te oe
e ® e@ LL |
= sn Suter — 11. - |
£ Fe + *. |
fy ee. oer wi, -
6 .. .. .% =
. EN ee es © °% Le "ui :
m= pt | % os oe =e ae |
e* 2 Pa . . 2 = 2 ar LEE ss À L
mae .. es 2 soe , e e 2 ee
ER DES eo * “are aa Re ws: 5
= Kr .,. .ı. CA LES
x e . 5 È * © * r . a Te .
» 2 .* e ® ‘ a LEE a |
. e 5 + bd ° bed e . 24 “ + = = !
LA e ' o . of} _ 4 8 . \
e [1 \ e , ® . a pP & |
eo \ . a Ne = = ®
x 4
le PR e #
' \ = = “ ‘
Ye M] » *
: rg. L
ch. * e L ’
4 4 +
ei ' A :
= =. y ‘
. w +
Fig.®. | uni: cm:
roots u 1
u. pe "a, " ”
ait N. ... e, = |
.° --- un -n |
af - « Es } Ye — ae
“ « 4 e >
vy .. one ... a mt IF. |
cé ie & . + TT es Vs
7. Tee =. a n ; . Ÿ Qe Fig.9.
At -* Ye id ee e a *e n.d. |
> - hed 2 - ! ts. Oe PES an
DB, oe 2 o = are t . er en . TT !
' ae ge eh ete ~ xO : 5 . . LA
a Be AR ra @ ae
... ' 0% CP ceased
i | en ge |
ch + ce at? .
ß : fee + Sn. - N.
mLV RES RE
ba: PER à Sig
en, À ; .
Fa | / e x
e a? x + ee... I rid V.
cd... … FRS. 1 ‘al $ es °. t
4 4 ~ + @
‘ we °° .°,, e
pee ee UNE UE { ~f eo. 2
Fig. 13. vo PQ 2 . ° . I / 2 e ae wc cd \
. ~~ Fe Ms e Sie e 2 6 *. +®@
mét.Ÿ. n BEN wee Te (A mtW 3 A 2 H ; FR ae ae
q c ch. we e wre
s
+7 _7 ~ a
ws, Oa: SET im
0 Int. -d o .. bor F 9
bad ep”: eo Im ‘ n > LR CRIER) We ÿ Î
.oe e .. ER : .’ * oe ee À ep.
sn te se on he eee |
oe. 5 e ri PO ay . 7 .
FR "oe - ms Im .. os. ren RS ER A oe tee lg. 15. im
Pew etc on 5% 0 beat u...’ .. ee" : ME | = “se ;
ee y see u
eo” t . | mi.M. .......
~ | \ LE = 2
l I s u .
apn. 4. ae Ps # = ï Eu, n ;
ch. ee syn.
ie 5
~ AR nn SEIEN, f° os = up - - mt VW |
sé RO À a | 24 ge -- -- apr.
ep Te te nett ee we | i a Se
: 5 h u ara - - Un.
Fig. hy. ee -
ch
3 ee tern
A . N,
mt «7° of. fee A
À . ee >’ ich
C2 oe ee > Ÿ
UE LEZ 1 es ie. cee u. 0%, air, - mt V. Fig. 19.
% Pa .* re > .. . „Ag 2 .
„ec PR tee Sur, VA à ., a pn.° Le.
ae: + + re. in Af dete!
Pins .. ls 2. Le ja i site es: vee *, mi ee oe se
7 RUE oo tee re *e tee @ *e, rots bad te 1 ave wet = ae ‘6 f nu
“ ° *.? ° m. E . -
om . rt "ru 16 Le. ir
ea . 3 a “= A e ber; En Pr e a | nn, . Im. IR Ate : “ey (ee PEN vee Br ta wil,
. e , .* mate = . . .
oe 2 a m . is . * e Ÿ s Boi a e ® 5 er “a re = wu . de, Je ewe né Ur
. . : , '
“fs 2 «. e le ; Pi e ” $ bs .. ar en hen _ L 1 j
2 ‘25 > | l : x ; x ! pe ® RC
ge — kg."
. Ay ch sch. i À
CA.
pv >=» ve ‘
Te. - 7 Fe ig.2f. À
D
pees Mes Fig. 25,
a a 7 mil wept Year J
... 76". . e? er __ # Lite
a ; - t ‘
m ae . eh an oh, PR ANNE AR MER SN Te. a ce...
~ %e EP NR s cake 68 ei , et Sa on
= 2. me 1% e oe ee ,” x mt W 3 „’ 2 .
rim 7 .. PER mV. Br .,
ee IE ee a 2
im. wae ~ os 2. « .° See" OR en
~ mt. ten | | ; ed, N Te
e ‘*e ae | u == A > 18 7 @e e
ae ae: . en
Bunt “ bas HU Se As
oe * ‘ PE ge
ee ch. tae À 5
bs oe eal
a xe se
é’ ..
ge‘
% “sy a
Fig. LA. t i ® 29
19.29,
4 “a a Ft.
mt VEL. pete En“ den,
APTE ghee Re Te melt ER ea Belen
-. = te Fire js . . X
. Be Pn. 0. ro DE ee ep. LT Er à Sigg = MN |
#4 - ar . 3 oe* ‘~ ae! . = pn 7 > ‚ie x 2
ee ae Ts .. ° | : peat ce anon FR ; ?
“+ e . 2 x y es
en. PR n 43 | ans 7 Bed ame oe N
o _ Ch - fp » m = e .. Pee ‘ is‘ : N ..,% 7
on re A us ‘ts VE ve, - = ch Im 7e e se. ‘i ie en | =} ch.
Be RE oh Be ee
" = eo’ % Ae fe à x
„2:7 er a 2@ i
; hg ca mp. ER
- ~e a
“. “x
Fe | Hatta del
PLATE XVIII.
Plate XVIII.
a.pn.1-6, Anlagen of pronephric , 2s., mesoblast.
tubules from the first to sixth. | onf.7, 77, &c., the first, second, &c.
a.sd., Anlage of segmental duct. | myotome.
cd., collecting duct. | mus., muscle-layer of myotome.
ch., chorda dorsalis. m.v., visceral layer of mesoblast.
c.p., coelomic projection. n., neural cord or canal.
cut., cutis-layer of myotome.
‚/g., fore-gut. |
EN eriblast pt.1-6, pronephric tubules from the
hy., hypoblast. first to sixth.
!.m., lateral plate of mesoblast. sch., subchorda.
m.p., parietal layer of mesoblast. | sd., segmental duct.
pp.c., pleuroperitoneal cavity.
Figs, 30-31. From the same scries as figs. 20-29 of the last plate.
Figs. 32-50. From a series of transverse sections through a younger embryo
of Stage III.
Figs. 51-58. From a series of transverse sections through an embryo of
Stage Im.
Fig. 59. A section through an older emboyo of Stage 111, the posterior
continuation of which is shown in the next foliowing plate (figs. 60-63).
Digitized by G t
; Ur n. mi.
8 Des
me. À. .° Nr, À Pe n.
.e ..- a Rn Fa e +
. 3 aad, oo >
apa, ar a ¢ Pp. + te” feats N ; .
ae ~ 6 a >. ._, .
u: e-" SEE 5 im . iad Se pare eo. 7
Ore *,° ac: Û es 2. = ci, au
ag res he ° ‘ tA. . u Pare aoe EN . 4
MR Te 2 at "Shae „’ 1 “ ae a OE Bo DE Ve rl.
we Py Par: à ne Ausg
im. ee ke . 2. ep. a BO
a ee = 2 = co
. DE ._
ve es:
1 Ove
pe
Je
ae
is ee a ; oe
a .
: lrg. 36. ._,*
+ “
im. N
ve
4
ee ma. Ls; |
4 ees cl a <i “yy R [ts mi ‘ a“
ss : I .
. TALS, .=Ss a - % N N A
, st ve ni Tea ge : Kasten Ben en.
| ;
cae * | apn. €. get Pate Ya ea > Ve 4
ue oe S Be ee
0 x + 2 ‘ Soe
hog. + i
u A wet ide as oy | 1 rés Zu ch. —
er LA. ıT Ae a eee eres ; RER ee
se ot. gs te Pose e a x NY = à
@ -- .@e™ re * Ad Bo 7
a
- & e 20 A + Ben
eu u ing \ £ J ‘ NM, > u a. Um a ; errr . > *.
; TER \ } fy AY Re La 4 J ee
mia. / À :
pot.
. Lars
.* en
7 D j
= #
e i
a
J *
a 4 =
J u; i
i m
= ä i
= ® % =
= ä
5 r " ä 2
DJs" # ’ =
# a À 4
# De = 7 .
D By # = =
2 aan Far es f
a fs
= » © . | a *
Fr
“ = = = L
a r a
= CS. | . # = 1 à =
m a a æ 5
= r * a - à “ =
Um à . j Sr
“ " = = + LE"
"ls
" " + à = +
a1" à . “
. 3 = i
a
* al . à a i
a - > |
Le : a 1
“ é e Li =
‘
Ib a * m = ‘i
}* 7
| CI
our. Sc. Coll. Vol. XI, Pl. X Vill,
ve ry}
ot % be o-
= = sis i i= 4 1 rd 1}
Ferre, \ Fig, Fig f Mia,
eos,
fe ot ee
N ® ° ° ae
To M
h Fe UE ST EN a. | :
. on | ea i
ees ». | de er = |
A ti ” “ Mi
D RE SE ruf 1 rt
> at Pe ee re Tae mur a]
rt. * Ae 8.5 0 74 ui a a = +
mole fie. Ar i
Te a \ set { |
bd le *: so @ "4 Fire 1
«9 J rE CRE ESA ! Le I 7)
à apa dl Sen 2" \ un ch
Rx P if à \ a a — 5,7 Ep: fier à d d'a Al
. 5 . ec & Pe) : j bowl Cf]
Lu ed ry a! bes. IF} Fr)
et 1 + ss st | Aria
x ep . jee ° es » “ ‘ = ‘ his. le A a
e -% 6 Ir a!
Z “et? 4 (| 1
im ~~ weet! ’ f FIT du? 1
. lt’ ‘et G:- a 14
fa. e eo Ar | ale
. - 2," Luis
Sas oy ie it
Ne " al,
Pes a! |
if #
F dé), i,
‘
je’ : ig
tip. 4 owe
ua .
+
À Li en
rh. hi #7 i
. rufe E y-
L Fat Fee rT
Fm Ne À Le dun
da A hy tu “
Pu. Æ ie » iF "+ .
Fe due inde dala ar dé
at lt \ À 4 L
Im. i ) ae; Tee Ur l'a ar un
AH: mire ur“ re PL EN
Fo tal ‘a 4e [ ees a
PAPE à " r ih. Aa, N= a!
. (u) lm Ay f ne h
LE, ir %
| mn
\
‘
or Pan
ra. IN
Fi _
i: fap. de
né — ee
= — ri CR Te a-
= = = fhe ys a
ete oe nid. NON u pe r=
. [ z Pa ¥ “a i- ra |
F | | Fi ;
“ \ A = = a
— Aly \ né Eur Cala ee, ch
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Matter €
PLATE XIX.
Plate XIX.
ed, collecting duct. mt.I, IT, &c., the first, second, &c.
ch., chorda dorsalis, myotome.
e.p., coelomic projection. mus., muscle-layer of myotome.
cul., cutis-layer of myotome. m.v., visceral layer of mesoblast.
d., dorsal row of mesoblast. n., neural canal.
ep., epiblast. nst.2-3, nephrostome the second
{g., fore-gut. and third.
!., Anlage of liver. pp.1-3, peritoneal partition.
hy., hypoblast. pp.c., pleuroperitoneal cavity.
!.m., lateral plate of mesoblast, pt.1-6, pronephric tubules from the
first to sixth.
| sch., subchorda.
sd., segmental duct.
ms., mesoblast. | sg., spinal ganglion.
m.p., parietal layer of mesoblast. v. ventral row of mesoblast.
m., Median row of mesoblast.
mch., mesenchymatous cells.
Figs. 60-63. From the same series as, and the posterior continuation of,
fig. 59.
Figs. 64-76. From a series of transverse sections through an oldest embryo
of Stage 111.
Figs. 77-81. From a series of transverse sections through an embryo of
Stage 1v; hence the body of embryo in the present stage is twisted,
the sections pass through unavoidably oblique planes,
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. MERE et Fig. 72. |
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Jour. Sc, Colt, Vol. XM. Pl. XIX,
Fi ‚nn. og LE OF
(4 ) ay TR, Fig » /
Ve “Ms + '
A a
ch On Sy n. Bi: Ka Sn
: | À
2, rt IE eg OS DEB) ac Ss Pe: me
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fe ' x ,
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fig : i Ny i = i@ |
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Ko.
PLATE XX.
Plate XX.
a.sd,, Anlage of segmental duct. ' Z.m., lateral plate of mesoblast.
bp., blastopore. mch., mesenchymatous cells.
brg., branchial region. ms., mesoblast.
cc., cloacal cavity. m.p., parietal layer of mesoblast.
cd., collecting duct. mt.I, II, &c., the first, second, &x.
ch., chorda dorsalis. myotome.
c.dv., diverticulum of cloacal m.v., visceral layer of mesoblast.
cavity. n., neural canal.
co.sd., cloacal opening of seg- nst.5, fifth nephrostome.
mental duct. perit., peritoneal membrane.
c.p., ewlomic projection. | PP. 1-3, peritoneal partition.
dl.bp., dorsal lip of blastopore.
fg., fore-gut. ;
ep., epiblast. | pt.1-6, pronephric tubules from
gc. genital cells the first to sixth.
. ge f
hy., hypoblast. | sch., subchorda.
| pp.c., pleuroperitoneal cavity.
int., intestine. | sd., segmental duct.
7., Anlage of liver. " yc., yolk-cells.
Fiss. 82-89. Sections from the same series as fig. 81, lying posterior
tu it. The section shown in fig. 87 passes through somewhat frontally
owing to the bending of the body-axis of the embryo; the neural canal,
which is bent in the same manner as the axis meets with two time
in section.
Figs. 90 and 91. Two sections passing through in the same way as in
fig. 87; in fig. 90 the dorsal lip of the blastopore, and in fig. 91, the
upper (dorsal) portion of it, is cut through.
Figs, 92-96. From a series of transverse sections throngh a little older
embryo than the last; the embryo is twisted in the same way as it.
Fig. 97. Frontal section through an embryo about the same stage as the
last, the body of which has been straightened before cut through.
Digitized by Google
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\
+
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ef). 2
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4
PLATE XXL.
Plate XXI.
«cv, anterior cardinal vein. | !.m., lateral plate of mesoblast.
au, auditery pit. | mch., mesenchymatous cells.
bp, ventral lip of blastopore. | m.p., parietal layer of mesoblast.
ivg., branchial region. perit., peritoneal membrane.
be., blood space. mt.I, If, &., the first, second
vi/., collecting duct. myotome, &c.
el., chorda dorsalis. m.v., visceral layer of mesoblast.
ee, cloacal cavity. n., neural canal.
“dv, diverticulum of cloacal | nst.2-6, nephrostomes from the
cavity. second to sixth.
co,sd., cloacal opening of seg- | pp.c., pleuroperitoneal cavity.
mental duct. pt. 2-5, pronephric tubules from
cw., wall of cloaca. the first to fifth.
up., epiblast. r.m., radix of mesentery.
yl,, glomerulus of pronephros. sch., subchorda.
/., dorsal fin. sd., segmental duct.
/y., fore-gut. t.a, tract of aorta.
/., heart. tac. tract of anterior cardinal
hy., hypoblast. vein.
/., liver, or Anlage of liver. tr.a. truncus arteriosus.
Fiys, 98-106. From a series of transverse sections through an embryo of
Stage V.
lys, 107-110. From a series of transverse sections through an embryo
uf Stage VI.
‘ig. 111. Transverse section through the cloacal region of an older embryo
of Stage v1.
lys, 112-114. A series of sagittal sections through a younger embryo in
Stage Iv.
ig. 115. A frontal section through an embryo a little more advanced
than that of Stage v1.
Digitized by Google
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PATES | | Haun del
Beiträge zur Wachstumsgeschichte der
Bambusgewächse.
Von
K, Shibata, Rigakushi.
Mit Tafeln XXI-XXIV.
I. Einleitung.
Unsere Kenntnisse über Bau und Lebensweise der Bambus-
gewächse waren bisher sehr mangelhaft gewesen, obwohl in neueren
Zeiten einzelne merkwürdige Erscheinungen auf dem Gebiete der
Physiologie dieser eigenartigen Baumgräser durch interessante Beo-
bachtungen') einiger die Tropen besuchender Botaniker zu Tage
gefördert wurden. Bekanntlich gehören die meisten Bambuseen
zu wärmeren Gegenden der alten und neuen Welt, mit Ausnahme
von einigen kälterem Klima angepassten Formen, wie z. B.
Bambusa Kurilensis, die in einer nördlichen Insel Japan’s
bei 46° n. B. gedeiht. Unser Land besitzt eine Reihe von
Bambusformen, welche unserer Pflanzenphysiognomik ein charac-
1)G. Kraus, Physiologisches aus den Tropen. I. Längenwachstum der Bambusrohre.
Ann. d. Jard. Bot. d. Buitenzorg. Vol. XII, p. 196.
H. Molisch, Über das Bluten tropischer Holzgewächse im Zustand völliger Belaubung.
Ann. d. Jard, Bot. d. Buitenzorg. 1898. 2 tes Suppl. p. 23,
428 K. SHIBATA:
teristisches Aussehen verleihen. Einige hochwüchsige Formen aus
der Gattung Phyllostachys sind bei uns überall häufig cultiviert,
hauptsächlich für die mannigfaltigste Verwendbarkeit der Rohre
und auch für ihre Frühjahrsschösslinge, die ein beliebtes Gemüse
darbieten, während andere Arten aus Arundinaria und Bambusa
als Zierpflanzen in unseren Gärten gemein sind.
Nun stellte ich es mir hier als Aufgabe, in erster Linie die
Natur und das Verhalten der wichtigen Baustoffe während der
verschiedenen Vegetationsperioden zu studieren, da mir besonders
die ungemein rasche Entwicklung der Schösslinge etwas interes-
santes in Bezug auf Stoffwanderungs- und Stoffumwandlungsvor-
gänge darzubieten schien, oder in anderen Worten die
Wachstumsgeschichte der Baumgräser mit Berücksichtigung der
Bauverhältnisse näher zu verfolgen.
Die früheren Angaben über die Systematik, Verbreitung und
äussere Lebensweise der Bambusgewächse haben eine Zusammen-
stellung in einem Werk Schröter’s') erfahren, und es schien mir
überflüssig dieselbe hier wiederzugeben. \Vas die Physiologie der
Bambusgewächse anbetrifft, besitzen wir abgesehen von älteren
Beobachtungen über das Wachstum der Schösslinge nur die ein-
gangs erwähnten Arbeiten von Kraus und Molisch. Über die io
Bambuspflanzen vorkommenden Stoffe besitzen wir ebenfalls spär-
liche Angaben. Cohn’) studierte „Tabaschir“ in seiner klassischen
Arbeit. Kozai*) stellte chemische Untersuchungen über die stick-
stoffhaltigen Bestandtheile des Schösslings von Phyllostachys mitis
mr re
1)C. Schröter, Der Bambus und seine Bedeutung als Nutzpflanze. Basel, 1895.
Vergleiche ferner:
E. Hackel, Banıbusacee. Engler’s Die’natürlichen Pflanzenfamilien. IJ, 2. p. 89.
A.et C. Riviére, Les Bambous. Vegetation, culture et multiplication. 1878.
2)F. Cohn, Uber Tabaschir. Beitriige z. Biol. d. Pflanzen. Bd. IV, p. 365.
3)Y. Kozai, On the nitrogenous non-albuminous Constituents of Bamboo shoots.
Bulletin of the College of Agriculture, Vol. I, Nr. 7,
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 429
an, und dabei fand er das Vorkommen von Tyrosin und Asparagin
neben kleineren Mengen der N ucleinbasen. Die Angaben über
die Bauverhältnisse der Bambusgewächse finden wir in Arbeiten
von Schwendener, Haberlandt, Strasburger, Hohenauer,
Güntz, Ross und Magnus. Näheres über die Litteratur wird
noch an geeigneten Stellen Erwähnung finden.
Die vorliegenden Studien wurden im Laufe des academischen
Jahres 1898-1899 im "botanischen Institute der Kaiserlichen
Universität zu Tokio ausgeführt. An dieser Stelle spreche ich
meinem hochverehrten Lehrer Herrn Prof. Dr. Miyoshi meinen
wärmsten Dank für seine vielfache Belehrung und Anregung
aus. Es ist mir auch eine angenehme Pflicht Herrn Prof. Dr.
Matsumura für seine gütige Unterstützung hier meinen herz-
lichsten Dank auszudrücken.
II. Untersuchungsmaterial und Methodisches.
Als Untersuchungsobjecte dienten mir die im botanischen
Garten der Universität cultivierten Bambusarten, insbesondere
Phyllostachys mitis, Riv., die sich in dieser Gegend in voller Ent-
wicklung findet und im hiesigen botanischen Garten auf ziemlich
grossem Grund gepflanzt ist. Die Wachstumsgeschichte der
Schösslinge der obengenannten Art wurde von der ersten Anlage
bis zum mehrere Meter hohen Halmzustand verfolgt. -
Auch die Schösslinge folgender Arten wurden zum Ver-
gleichungszweck untersucht :
im April austreibende— Bambusa palmata, Bambusa Vettchit;
im Mai austreibende—Phyllostachys puberula, Arundinaria
‚japonica ; |
im Juni austreibende—Phyllostachys bambusoides ;
430 K. SHIBATA :
im Juli-August austreibende—Arundinaria Simoni, Arundi-
naria Hindsii ;
im October- November austreibende—Arundinaria Matsu-
muræ, Arundinaria quadrangularis, Arundinaria Hindsu
var gramınea. |
Die Entwicklung der Rhizomspitze wurde bei folgenden
Arten im Herbst untersucht: Phyllostachys mitis, Phyllostachys
bambusoides.
Für andere Arten, die ich in meiner Untersuchung gezogen
habe, verweise ich auf das am Ende dieser Arbeit beigefügte
Artenverzeichniss,
Um die Umwandlung und Wanderung der Stoffe in Reserve-
stoffbehältern und in wachsenden Theilen zu verfolgen,
bediente ich mich unter nöthigen Cautelen der üblichen micro-
chemischen Methoden. Darüber sei hier folgendes bemerkt:
Stärke. Meyer’sche Chloralhydratjodlésung’) wurde mit
Vortheil benutzt.
Glycose. (Reducierender Zucker). Meyer’sche) und
Schimper’sche*®) Methoden wurden neben einander angewandt,
dabei hat sich die letztere zur Nachweisung der kleineren
Menge geeigneter erwiesen. Obwohl diese üblichen Methoden
auch zu unserem Zweck völlig ausreichten, habe ich noch Sicher-
heits wegen eine andere Reaction ausgeführt. Ich habe nämlich
die Wasserauszüge von jungen Halmen, Wurzeln, Rhizomen und
Scheideblättern und auch den Blutungssaft mit essigsaurem
Phenylhydrazin erwärmt, und dabei erhielt ich stets charac-
teristische gelbe Nadelkrystalle von Glucosazon.
1) Vergl. Strasburger, Botanisches Practicum. III. Auflage. p- 277.
2)A. Meyer, Microchemische Reaction zum Nachweis der reducirenden Zuckerarten.
Ber. d. D. B. G. 1885. p. 332.
3) A. Zimmermann, Die botanische Microtechnik. p. 75.
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 431
Rohrzucker. Die Unzuverlässigkeit der bekannten Sachs’-
schen Methode wurde vielfach von Autoren betont. Ich habe
nur ausnahmsweise diese Reaction benutzt, während in den meisten
Fällen ich mich der neulich von Hoffmeister') aufgestellten
Invertinmethode bediente.’)
Gerbstoffe. Die Eisensalzlösungen, insbesondere die ätherische
Lösung des Eisenchlorids, und auch die Kaliumbichromatlösung
wurden angewandt.
Fette. Alkannatinctur und 1-procentige Osmiumsäurelösung
wurden benutzt.
Asparagin. Die bekannte Borodin’sche Methode?) hat sich
zweckenisprechend erwiesen. Zur Erkennung der erhaltenen
Krystalle als Asparagin diente mir hauptsächlich die Winkel-
messung. Vielfach wurde das Borodin’sche Verfahren mit
gesättigter Asparaginlösung angewandt. Ferner diente mir
Diphenylamin-Schwefelsäure zur Unterscheidung von Asparagin
und Salpeter.
Tyrosin. Die nach Borodin’scher Methode behandelten
Schnitte ergaben eine reichliche Ausscheidung von eigenthümlichen
Krystallen, die ohne Schwierigkeit mit Tyrosin identificiert
1)C. Hoffmeister, Über den microchemischen Nachweis von Rohrzucker in pflanz-
lichen Geweben. Jahrb. f. wiss. Bot. Bd. XXXI, p. 688.
2) Die von den ,, Ebisu “-Brauerei bezogene Hefe-Reinkultur wurde zur Darstellung von
Invertin verwendet, dabei habe ich wie folgt verfahren: Die Hefemasse wurde mittelst
Filtration von der Kulturflüssigkeit befreit und nach wiederholtem Auswaschen mit Wasser zum
dicken Brei angerührt. Der Hefebrei kam nach dem Verreiben im Mörser in den Thermostat
bei 50°C, in welchem er für 10-12 Stunden gelassen wurde. Hiernach wurde die abfiltrierte
Flüsigkeit mit 90% Alcohol versetzt, und der dabei entstandene voluminöse Niederschlag
auf Filtrirpapier gesammelt, welcher, nach wiederholtem Auswaschen mit 902 Alcohol und
dann mit absolutem Alcohol auf Schwefelsäure getrocknet wurde. Die wässrige Lösung der
erhaltenen weissen kreideartigen Substanz, die allein niemals Fehling’ sche Lösung reduciert,
zeigte ein energisches Inversionsvermögen. Bei wiederholten Versuchen habe ich
ferner in keinem fall die Beimengung von diastatischen und cellulosespaltenden Enzymen
gefunden. Weitere Verfahren mit den Schnitten genau nach Hoffmeister.
3) A. Zimmermann, Die botanische Microtechnik. p. 80.
432 | K. SHIBATA:
wurden!) (Fig. 58). Belzung’sche Glycerin-Methode*) wurde
auch mit Vortheil benutzt, wobei sich schöne Nadelbüschel in
den Zellen bilden (Fig. 61). Ich habe ferner zur Erkennung der
Vertheilung des Tyrosins in eiweissarmen (Gewebetheilen
Millon’s Reagens benutzt, und dabei wurden die tyrosinreichen
Zellen schnell blutroth gefärbt. Die Färbung bleibt nach dem
Auslaugen der zuvor mit absolutem Alcohol behandelten Schnitte
mit dem warmen Wasser für 10-20 Minuten so gut wie gänzlich
aus. Daher kann diese Rothfärbung niemals von Eiweissstoffen
herrühren.
Eiweiss. Biuretreaction und Raspail’s Reaction wurden
vornehmlich benutzt. Millon’s Reagens kam zur Anwendung
erst nach dem Ausziehen von Tyrosin in oben beschriebener
Weise. | |
Mineralstoffe. Die von Schimper’) empfohlenen Reagentien
wurden verwendet. Die Controllversuche wurden öfters ausge-
1) Dies geschah aus folgenden Gründen:
1. Die Gestalt der Krystalle. Die feine Nadelbüschel in dendritischer Gestalt oder
Doppelpinselform bietet ganz dasselbe Aussehen wie reines Tyrosin.
2. Das optische Verhalten. Im durchfallenden Licht erscheinen die Krystalle briun-
lich und im auffallenden Licht weisslich seidenglänzend. Im polarisierten Licht
zeigen die Krystalle starke Doppelbrechung.
3. Die Löslichkeitsverhältnisse. Die Krystalle sind unlöslich in kaltem Wasser, aber
löslich in kochendem Wasser, Ammoniak und verdünnter Salzsäure. Ferner
sind die Krystalle unlöslich in heissgesättigter Tyrosinlösung.
4. Das Verhalten beim Erhitzen. Wenn man den mit Tyrosinkrystallen besetzten
Objectträger auf der Flamme erhitzt, bis nebenbei vorhandene Asparaginkrystalle
sich zu braunen Schäumen verwandeln (ca. 200° C), so sieht man, dass die
Nadelkrystalle ganz unverändert bleiben.
5. Das Verhalten gegen Millon’s Reagens. Die Krystalle lösen sich im Millon's
Reagens mit einer prachtvoll rothen Färbung der umgebenden Flüssigkeit.
Die oben angeführten Merkmale reichen aus, die Krystalle microchemisch als Tyrosin zu
erkennen.
2) Belzung, Recherches chimique sur la Germination. Ann. d. Sc. nat. Bot. Ser. VII.
T. 15, p. 209.
3) A. F. W. Schimper, Zur Frage der . Assimilation der Mineralsalze durch die
grüne Pflanze. Flora. Bd. 73. 1890. p. 210.
WACHSTUMSGESCHICHTE D. BAMBUESGWAECHSE. 433
führt, um die Reinheit der Reagentien zu prüfen. Die micro-
chemischen Reactionen wurden sowohl an frischen Schnitten als
an auf Deckgläsern geglühten Aschen vorgenommen.
III. Die Bauverhältnisse.
Die Bauverhältnisse der Bambusen sind bisher spärlich
und nur gelegentlich Gegenstand der anatomischen Forschung
geworden. Strasburger') hat den Bau des Gefässbündels von
Bambusa vulgaris kurz geschildert. Das mächtig entwickelte
Bastgewebe in Bambushalmen wurde vielfach von Schwendener)
Haberlandt‘) u. A. erwähnt. Ross‘) bemerkte den anomalen Bau
der Wurzeln. Die Betrachtungen über die Blattstructur finden
wir in den Arbeiten von Güntz’), Magnus‘), Haberlandt’)
und Schwendener’). Übrigens liegen uns noch einige kurze
Angaben von Hohenauer’) und Mébius”) vor. Nun schien
es mir wünschenswerth die Bauverhältnisse der Vegetationsorgane
der Bambusgewächse einem genaueren Studium zu unterwerfen,
damit für die physiologische Forschung dieser interessanten Pflan-
zengruppe eine festere Grundlage geschaffen werde. Meine
1)Strasburger, Über d. Bau u. Verrichtungen d. Leitungsbahnen. 1891. p. 363.
2) Schwendener, Das mechanische Princip in anat. Bau d. Monocotylen. p. 65.
3) Haberlandt, Entwicklungsgeschichte des mech. Gewebesystems d. Pflanzen. p. 23.
4) Ross, Beiträge z. Anatomie d. abnorm. Monocotylenwurzeln. Ber. d. Deutsch. Bot.
Gesells.{ Bd. I, p. 338.
5) Güntz, Onters. üb. d. anat. Structur d. Gramineenblätter. p. 37, 41, 44, 48, 64 etc.
6) Magnus, Einfalzungen d. Zellmembran. (Just’s Jahresber. d. Bot. I, p. 367).
7) Haberlandit,,‚Vergl. Anat. d. assim. Gewebesystems d. Pflanzen. Jahrb. f. wiss. Bot.
Bd. XIII, p. 100.
8) Schwendener, Die Mestomscheide der Gramineenblätter. Ges. Bot. Mitt. Bd. II,
p. 178.
9) Hohenauer, Vergl. anat. Unters. üb. d. Bau d. Startimes. bei d. Gramineen. p, 556,
10) Mébius, Ub. d. eigent. Blühen von Bambusa vulgaris. (Ref. in Bot, Centralbl. 1899.
Nr. 51, p. 479). a eee ad Se; | |
434 | K. SHIBATA :
diesbezügliche Untersuchungen erstreckten sich auf sieben und
zwanzig vorwiegend einheimische Formen, die sich in die vier
Gattungen von Phyllostachys, Arundinaria, Bambusa und
Dendrocalamus vertheilen. Die wesentlichen Ergebnisse will ich
in folgenden Zeilen kurz darzustellen versuchen.
Das BRuızom.
Die Rhizome') von Phyllostachys- und Arundinaria-Arten
sind bekanntlich kurz gegliederte horizontal verlaufende Stengel-
gebilde mit einer rundlichen oder rundlich-ovalen Querschnitt-
form. Die internodiale Markhöhle ist stets stark reduciert, meist
nur einige mm breit und kommt nicht selten zum gänzlichen Ver-
schwinden. Wir wollen zunächst beispielweise ein Rhizominter-
nodium von Phyllostachys mitis ins Auge fassen. _
Nächst unter der Epidermis kommt ein 1-3 schichtiger
Ring von den englumigen langgestreckten sklerotischen Paren-
chymzellen, deren Querwände öfters etwas schief gestellt sind.
Das darinnen liegende 20-25 schichtige bündelfreie Parenchym
wird als die primäre Rinde?) aufgefasst. Es geht ohne scharfe
Grenze zum Grundgewebe des Centralcylinders über, in welchem
in üblicher Weise die collateral gebauten Gefässbündel zerstreut
liegen. Sämmtliche Parenchymzellen sind verholzt und mit
zahlreichen ovalen Tüpfeln versehen. In diesem Gefässbündel
erblickt man ein typisch gebautes Gramineenbündel. Die grosse
Lumenweite der Siebröhren ist dabei höchst auffallend?) ; es wurde
oftmals einen Durchmesser von 0.15 mm erreicht‘), während
1) Vergl. A. et C. Rivière, Les Bambous. p. 68, p. 236.
2) Falkenberg, Vergl. Unters. üb. d. Vegetationsorg. d. Monocotylen. p. 163.
' 8)8Strasburge?, Leitungsbahnen. p. 363.
4) Die Angaben über die Lumenweite der Siebröhren einiger Pflanzen findet man bei
Lecomte (Ann, d, Se. nat, Pot, Ser. VII, T. X, p. 242).
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 435
die grösste Parenchymzelle 0.09 mm und die Geleitzellen meist
nur 0.005 mm weit sind. Die beiden seitlichen Gefässe, deren
maximale Lumenweite 0.2-0.3 mm beträgt, communicieren mit den
einschichtigen Belegzellen durch regelmässig angereihte quer-
gestreckte Tüpfel. Die stark entwickelten Bastbelege um die
Gefässbündel sind seitlich an der Grenze zwischen Hadrom und
Leptom und oft auch unterhalb der seitlichen Gefässe unter-
brochen. Dadurch kommen ein oder zwei Paar Durchlassstellen
zu Stande, die, wie zuerst Schwendener') bemerkt hat, einen
Stoffaustausch zwischen Bündelelementen und Grundgewebe
ermöglichen.
Wenn man die Querschnittbilder der Rhizominternodien der
übrigen Arten in Betracht zieht, so lassen sich unter denselben
folgende drei Typen unterscheiden, nämlich :
Erster Typus. Die äussersten Bündel, welche direct an
die Rinde grenzen, stehen vollkommen isoliert von einander.
Hierher gehören Phyllostachys mitis, Phyllostachys bambusoides,
Phyllostachys puberula und Arundinaria Narihira.
Zweiter Typus. Die Bastbelege der äussersten Bündel
verschmelzen sich häufig unter einander und auch mit den
Baststrängen zu unregelmässigen Bastbändern. Als Beispiele
dienen Arundinaria japonica, Bambusa borealis, Arundinaria
Tootsik, A. Simoni, A. Hindsu etc. In den zwei letztgenannten
Arten befindet sich jedoch oft ein nahezu vollkommener Bastring.
(Fig. 3).
Dritter Typus. Der echte subcorticale Bastring’), an wel-
chen die Mestombündel innen angelehnt sind, befindet sich bei
1)Schwendener, Das mechanische Princip. p. 107.
2) Vergl. Haberlandt, Entwicklungsgeschichte des mechan. Gewebesystems. p. 28.;
Physiologische flanzenanatomie. p. 157.
436 K. SHIBATA:
Bambusa palmata, B. Veitchii, B. paniculata, B. nipponica,
B. ramosa, B. nana, Arundinaria quadrangularıs, A. Matsumure,
A. variabilis, Arundinaria pygmea und ferner Phyllostachys
Kumasasa. (Fig. 2).
Der Bastring der letzterwähnten Arten, welcher je nach
Species verschieden stark ausgebildet ist, geht entwicklungs-
geschichtlich aus einem entsprechend continuirlichen Cambiun-
ring‘) hervor. Schwendener sagt’): „Für Bambuseen ist die
Querschnittform des mechanischen Systems, wie ich sie früher
beschrieben habe, characteristisch genug, um jede nähere Verwandt-
schaft mit den Festucaceen oder irgend einem anderen Tribus
auszuschliessen. Ein Bastring ist nicht vorhanden,......... “ Diese
Bemerkung Schwendener’s passt aber nach obigem Befunde
auf die Rhizome nicht.) Der subepidermale Sklerenchymring ist
nur schwach entwickelt, es ist bei den meisten Arten nur 1-2
Schichten dick. Die Dicke der primären Rinde ist meist un-
ansehnlich und variirt zwischen 4-35 Zellschichten.
Der Bastring wird stets vielfach unterbrochen in den Knoten,
um hier den neu eintretenden Blattspursträngen Platz zu machen.
Ferner ist es als die Regel hervorzuheben, dass innerhalb des
Knotens der Bastbeleg des Mestombündels eine bedeutende Re-
duction erfährt, und sich meist nur auf eine dünne Sichel um
das Leptom beschränkt. Die sämmtlichen Elemente des Bündels
sind hier kurzgliedrig, und die Seitenwände der Siebröhren sind mit
den ausserordentlich zahlreichen Siebtüpfeln versehen. Bekannt-
1)Haberlandt, Entwicklungsgeschichte. p. 28.
2)Schwendener, Die Mestomscheide der Gramineenblätter. p. 188. _
3) Die mittleren Durchmesser der Rhizome dieser drei Typen stehen ungefähr im Ver-
hältnisse 6:3:1. Die Ausbildung der Bastplatte resp. des Bastrings in Rhizomen entspricht
wohl den mit der Dünnheit steigenden Aufforderungen für die Biegungsfestigkeit: Jedenfalls
gehört hier die Anordnung des mechanischen Systems in Rhizomen nicht zum sogenannten
taxonomischen Merkmalen.
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 437
lich findet sich in dem Knoten die Vereinigung der Blattspur-
stränge unter einander und mit den Achselknospenbündeln statt!).
Die hier eintretenden Knospenbündel verbreiten sich in dem
Knotengewebe nach allen Richtungen hin. Die Gefässe der Knos-
penbündel setzen sich in üblicher Weise unter starker Krümmung
an die der Blattspuren an’). Dennoch verdient die Art und
Weise, wie der Übergang des Leptoms erfolgt, eine besondere
Beachtung. Das Leptom des Knospenbündels ist bei der
Ansatzstelle an Blattspuren so stark angeschwollen, dass ihr
ganzer Umriss mit einer Spindel zu vergleichen ist. Figur 4
stellt ein derartiges Gebilde dar. Diese angeschwollene Partie
weicht von dem üblichen Bau des Leptoms in so fern ab, dass
sie die Differenzierung ihrer Elemente in Siebröhren und Geleit-
zellen nicht mehr aufweist, sondern aus lauter gleichartigen
feinen ca. 5-6 w breiten cambiformartigen Elementen?) zusam-
mengesetzt ist (Fig. 4 u. 8), und folglich im Querschnittbild ein
regelmässiges englumiges Maschenwerk darstellt (Fig. 5 u. 7).
Die Anordnung dieser feinen cambiformartigen Elemente bietet
eine grosse Eigenthümlichkeit dar. Die etwas schräg gestellten
Endflächen der seitlich an einander stossenden Elemente scheinen
ungefähr auf derselben Querebene zu liegen, und demgemäss stellt
der ganze spindelförmige Theil im Längsschnitt ein der Länge nach
an einander angereihtes meist 5-7 faches Stockwerk von cambi-
formartigen Elementen dar. Die Elemente in den mittleren 1
1) Vergl. Falkenberg, Vergl. Unters. d. Vegetationsorgane. p. 187.
2) Strasburger, Leitungsbahnen. p. 353 u. p. 365.
3) Selbst bei den relativ weiteren (z. B. bei Arundinaria Simoni) überschreitet die Breite
kaum 9 ew. So weit ich unterrichtet bin, wurde derartige Structur bisher in keiner anderen
Pflanzen beobachtet. Zum Beispiel finden wir keine diesbezügliche Angabe bei verschiedenen
Monocotylen, die von Falkenberg (loc. ci.) und Strasburger (loc. cit.) gründlich
untersucht wurden. Ob sie auch bei anderen Pflanzen vorkommt muss deshalb zur Zeit
dahingestellt bleiben.
438 K. SHIBATA :
oder 2 Etagen dieses Stockwerks sind sehr langgestreckt und
besitzen fein undulierte Seitenwände mit zahlreichen grossen
ovalen tüpfelartig verdünnten Stellen, die zum Beispiel bei
Phyllostachys mitis eine Weite von 5X3 „ erreichen (Fig. 6). In
einem Ende dieses spindelförmigen Theils vermitteln die etwas
breiteren Elemente,—die sich zu 4-5 je einer feinbetüpfelten
Endfläche der Siebröhren und einzeln auch den Geleitzellen
anschlissen,—den Uebergang zum normal gebauten Leptom de
Knospenbündels (Fig. 10). Das andere Ende der Spindel setzt
sich in verschiedener Neigung und oft sogar rechtwinkelig an das
Leptom der Blattspuren an (Fig. 8). Einige Schichten der
Scheideelemente grenzen gewöhnlich den spindelförmigen Theil
vom Grundparenchym ab. Die Zellwandbeschaffenheit der
spindelartigen Theile weicht kaum von der des Leptoms ab; sie
zeigen nämlich ebenso starke Cellulose-Reaction mit Chlorzinkjod
oder Schwefelsäure-Jod, und sie werden auch mit Anilinblau,
Congoroth u.a., im nahezu gleichen Farbenton wie Siebröhren
gefärbt. In späterem Alter tritt jedoch oft eine Spur Holzreaction
in den Elementen der oben erwähnten mittleren Etagen ein. Was
den Plasmagehalt dieser Theile anbetrifft, so scheint es nur auf
einen zarten Wandbeleg beschränkt zu sein, wie es bei Siebröhren
stets der Fall ist. In den ersten Entwicklungszuständen habe ich
constatiert, dass diese Anschwellung aus den entsprechend ver-
mehrten Längstheilungen der procambialen Zellen an der betreffen-
den Stelle hervorgeht. In solchen früheren Stadien zeichnet sich
diese Anschwellung durch besonders reichlichen Plasmagehalt
und auffallend grosse Zellkerne aus, wie es Fig. 9 zeigt.
Derartige spindelförmige Leptomanschwellungen in Knospen-
bündeln habe ich regelmässig in Rhizomknoten sämmtlicher von
mir untersuchter Arten gefunden, aber man findet sie am stärksten
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 439
ausgebildet in den Rhizomknoten der Phyllostachys-Arten, wobei
ihre Querschnittgrösse sogar einem grossen Mestombündel nahe-
kommt. Da die an Rhizombündel sich ansetzenden Achselknos-
penstränge in ihrer Gesammtheit ein physiologisches Analogon des
haustorial Saugorgans darstellen, so ist es nicht unmöglich, dass
dieses Gebilde in dem Sinne ausgebildet ist, dass es eine specifisch
absorbierende Wirkung auf die Siebröhren der Mutterrhizome
auszuüben vermag. Allerdings besitzt es in seinen anatomischen
Merkmalen vieles gemein mit den üblichen Absorptionsgeweben’).
So lange aber die Stofftransportmechanik im Leptome noch nicht
in allen Hinsichten aufgeklärt ist”), möge die nähere Erörterung
der physiologischen Vorgänge, die sich in diesem abweichend
gebauten Leptomtheile abspielen, auf eine künftige Gelegenheit
verschoben werden.
Der Ham.
Die Bambushalme*) sind bekanntlich mit den hohlen Inter-
nodien versehen, die bei Phyllostachys mitis oft eine ansehnliche
Dicke von 20 cm erreichen.
Die primäre Rinde ist stets weit schwächer entwickelt als
in dem Rhizome; diese Verhältnisse, wie sie schon von Falken-
berg‘) und Rothert*) für andere Pflanzen nachgewiesen wurden,
gehen noch aus den folgenden Beispielen deutlich hervor:
1) Haberlandt, Physiologische Pfanzenanatomie. p. 186.
2) Czapek, Uber d. Leitungswege d. organischen Baustoffe in Pflanzenkorper. p. 24;
Lecomte, Etude du Liber des Angiospermez Ann. d. Sc. nat. Ser. VII. T. X, p. 303.
3) Vergl. Riviére, Les Bambous. p. 134.
4) Falkenberg, lc. p. 134.
5) Rothert, Vergl. anat. Unters. üb. d. Differenzen im prim. Bau d. Stengel u. Rhizome.
p- 92.
440 K. SHIBATA :
HALM RHIZOM
Durchmesser| Dicke Durchmesser | Dicke
Central- det Q* Ceutral- der a
cylinders Rinde cylinders Rinde -
Phyllostachys mitis 130.0 | 0.315 | 412] 23.8 | 0.728 | 33
Phyllostachys bambusoides 34.0 | 0.059 | 575) 20.0 | 0.611 | 33
Phyllostachys puberula 56.0 | 0.049 |1142| 22.0 | 0.933 | 23
Phyllostachys Kumasasa 2.7 | 0.023 | 120 5.6 | 0.780 | 7
Arundinaria Simoni 14.0 | 0.049 | 285 8.0 | 0.494 | 16
Arundinaria japonica 17.0 | 0.050 | 340 8.4 | 0.286 | 29
Arundinaria Hindstt 22.0 | 0.059 | 373| 18.0 | 0.364 | 49
Arundinaria quadrangularis| 23.0 | 0.045 | 511 8.0 | 0.260 | 31
Arundinaria Matsumure 2.9 | 0.018 | 139 29 | 0. 20 | 15
Arundinaria pygmea 2.3 | 0.036 64 4.5 | 0.325 | 14
Arundinaria Narthira 14.0 | 0.049 | 285] 21.0 | 0.468 | 45
Bambusa borealis 5.5 | 0.045 | 122 6.5 | 0.212 | 31
Bambusa palmata 10.0 | 0.063 | 159 7.1 | 0.624 | 11
Bambusa Veitchir 4.5 | 0.023 | 200 2.9 | 0.143 | 20
*Q.=Das Verhältniss des ersteren zur letzteren.
Die äussersten 1-2 Schichten Rindenparenchymzellen sind
oft sklerotisch verdickt (Fig. 12) und unterscheiden sich von den
übrigen nur durch eine grössere Länge; folglich haben wir
hierbei keineswegs mit einem Bastring, wie Haberlandt einst
annahm,') zu thun. Die Bastbelege der peripherischen Gefäss-
bündel und die dazwischen liegenden Baststränge verschmelzen
mit einander zu unregelmässigen Bastbändern, besonders häufig
in dünneren Halmen von Arundinaria Matsumuræ, A. pygmea
etc. Dennoch begegnet man hier in keinem Fall dem echten
Bastringe.
1) Haberlandt, Entwicklungsgeschichte. p. 23.
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 441
Die zuerst von Schwendener’) bei einigen Bambusaarlen
entdeckte. eigenthümliche Parenchymlamelle, die quer in dem
innenseitigen Bastbelege inseriert ist, habe ich auch in den
Halmen aller echter Bambusaarten (B. vulgaris, B. nana und LB.
stenostachya), Dendrocalamus latiflorus und bei 2 Arundinaria-
arten (A. Hindsii, A. quadrangularis) aufgefunden. Überdies
habe ich die Fälle beobachtet, dass die Lamelle nur an einer
Seite in das Grundparenchym übergeht, und dass sogar das Paren-
chym in der Mitte des Beleges allseitig von Bastzellen umschlossen
liegt (Fig. 17 u. 18). Nach den letzterwähnten Thatsachen
erscheint es a priori sehr wahrscheinlich, dass dieses Parenchym-
gewebe erst nachträglich aus einem Theil des Procambiums des
Bastbeleges hervorgeht, wie es Haberlandt?) schon vermuthet hat.
In der That konnte ich in einem Procambialstrang in jun-
gen Internodien zuerst nichts von dieser Parenchymlamelle
erkennen. Sie differenziert sich erst später aus einer Strang-
anlage, in welcher alle Formelemente schon fertig angelegt sind,
derart, dass die langgestreckten Procambialzellen in betreffender
Stelle successive Quertheilungen erfahren und zum Epen um-
gewandelt werden (Fig. 21 u. 22). Die feinkörnige Stärke, die
später dem Zucker Platz macht, tritt sogleich in diesem Gewebe
auf (fig. 20 u. 22) und bleibt in demselben während der weiteren
Ausbildung des Stranggewebes. In dieser Weise dient die
Parenchymlamelle den dem Mestom unmittelbar anliegenden
Bastzellen als ein Speicherungsort der nötigen Baustoffe. Die
durch diese Parenchymlamelle vom Mestom abgetrennte Bast-
masse bleibt gewöhnlich in ihrer Ausbildung sehr zurück, wie
das Fig. 19 zeigt. Nun schien es mir berechtigt diese Paren-
1) Schwendener, Das mechanische Princip. p. 66.
2) Haberlandt, le p. 23. ne oy Se
449 K. SHIBATA: :
chymlamelle als eine im Innern des Stranggewebes eingeschobene
„Stärkescheide‘"), die in ihrer physiologischen Rolle der
gewöhnlichen strangumgebenden gleicht, aufzufassen. Derartige
Einrichtungen würden vielleicht zweckentsprechend sein, bei
einem mit so starkem Bastbelege versehenen Bündel, wie es bei
Bambuseen angetroffen wird?).
Die schon erwähnten spindelförmigen Leptomanschwellungen
des Knospenbündels kommen auch in dem Halmknoten vor,
jedoch meist in schwächerer Ausbildung.
Die dünneren Blatttragenden Zweige stimmen in ihrem Bau
mit den dickeren Halmtheilen wesentlich überein. Bei einem
solchen (mit einem Durchmesser kleiner als 1 mm) wird das
Rindenparenchym zu 1-2 Schichten reduciert und mehr oder
minder verdickt. Die aussenseitigen Bastbelege der peripherischen
Bündel stossen oft direct an die Epidermis, so dass eine Art
Bastrippe zu Stande kommt’). Derartige Rippenbildung konnte
ich jedoch bei einigen Arten, wie Arundinaria Matsumura, selbst
in den dünnsten Zweigen (0.7 mm dick) nicht nachweisen.
Was den Gefässbündelverlauf in den Halmen sowie in den
Rhizomen anbetrifft, so gehört er dem Palmentypus‘) an, and
habe ich durch successive Querschnitte und Längsschnitte in
der Spitzenregion constatiert, dass die grossen medianen Blatt-
spurstränge 5-6 Internodien zurücklegen müssen, bevor sie sich
an andere Blattspurstränge ansetzen.
1) Vergl. Heine, Über physiologische Function der Stirkescheide. Ber. d. D. B G.
1885, p. 189.
2) Allerdings wurde die mechanische Bedeutung, die Detlefsen (Ub. d. Biegungselasti-
cität v. Pflanzentheilen. Arb. d. Bot. Inst. Würzburg. Bd. IIL'p. 182.) diesen Parenchym-
lamellen zuzuschreiben versuchte, von Schwendener (Zur Lehre v. d. Festigkeit d. Gewächse.
Ges. Bot. Mitteil. Bd. II. p. 19-20.) genügend widerlegt.
3) Vergl. Schwendener, Die Mestomscheide. p. 183.
4) De Bary, Vergleichende Anatomie d. Vegetationsorgane, p. 271 ff
° WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 443
DER STIEL.
Die vielen untersten Internodien des Schösslings von Phyllo-
stachys milis vereinigen sich sehr früh zu einem verholzten soliden
Gebilde, welches im fertigen Zustande 2-4 cm lang und nur
1.0-1.5 cm dick ist. Das Gebilde, das ich hier ,, Stiel‘) nenne,
lässt keinen Unterschied mehr zwischen Internodien und Nodien
im inneren Bau erkennen.
Die Rinde dieses Theils besteht aus etwa 20 Schichten
parenchymatischer Zellen. Die Bastbelege der äussersten Bündel
verschmelzen sich zu einem vielfach unterbrochenen unregel-
mässigen Band. Nach innen liegen zahlreiche Bündel, welche
die ganze Länge des Stiels hindurch gedrängt verlaufen (Fig. 13)
und dann in Rhizomknoten eintreten, um sich dort an die
Blattspuren anzusetzen. So überwiegen im Querschnitte des
Stiels sehr stark die Bündel, und das dazwischen liegende Paren-
chym ist zu einem 2-4 schichtigen schmalen Gewebe reduciert.
Diese Verhältnisse entsprechen wohl der Function des Stiels als
Leitungswege und nicht als Speicherngsort. Die Querschnitt-
form des Bündels mit Bastbelege ist in den meisten Fällen
rundlich oval und es ist vollkommen von 3-6 schichtigen
Bastzellen umgeben. Daher ist der Stoffaustausch zwischen
leitenden Elementen und Grundparenchym so gut wie gänzlich
ausgeschlossen. Das Leptom nimmt die äussere Hälfte des Bündels
ein und besteht aus einer Anzahl 0.04-0.05 mm breiten Siebröhr-
en und englumigen Geleitzellen. Das Hadrom besteht aus nur
einem (seltener zwei) grossen Gefässe (oft bis 0.15 mm weit),
welches von einigen Schichten kleinzelligen Hadromparenchyms
1) Dieser „Stiel“ stellt also eine einzige Stoflleitungsbahn zwischen dem vom Schösslinge
sch entwickelnden Halme und dem Rhizome dar.
444 K. SHIBATA : .
umgeben ist (Fig. 14). Dazu kommen noch einige Tracheiden
mit netz-, spiral- oder ringförmigen Wandverdickungen. Bei den
übrigen Arten sind die ebenso schmalen Stieltheile in genau
derselben Weise ausgebildet und sie stimmen in ihrer inneren
- Structur mit dem oben beschriebenen gänzlich überein.
Dre WoRZEL.
Die zahlreichen Wurzeln’) befinden, sich radial angeordnet
an den Rhizomknoten und den unterirdischen Halmknoten ; sie
erreichen bei Phyllostachys-Arten eine maximale Länge von 70
cm mit einem Durchmesser von 4 mm. Zunächst will ich hier
den anatomischen Bau der Wurzelrinde von Phyllostachys-, und
Arundinaria-Arten näher betrachten. |
Die äusserste Zellschicht der Rinde lässt sich als die Aus-
senscheide unterscheiden, indem die äusseren und radiaren
Zellwände sehr stark verdickt sind (Fig. 24a u. 25). Damit
spielt sie die Rolle der schützenden Oberhaut anstatt der Epi-
dermis, die sehr früh zerstört und abgeworfen wird. Nach innen
folgt die verholzte Bastschicht (Fig. 24a). Das Rindenparenchym
lässt sich in die äusseren aus unregelmässig polygonalen weit-
lumigen Zellen zusammengesetzten Schichten und die inneren
aus regelmässig in radialen und concentrischen Reihen angeord-
neten Zellen bestehenden Schichten unterscheiden’). Die letzteren
sind von einer Anzahl radialer Lufträume durchzogen. Die
Zellen der Endodermis besitzen eine starke Verdickung von
inneren und radialen Wänden, die in üblicher Weise verkorkt
sind (Fig. 28). Die stark verdickte Wandung ist zierlich ge-
1)A. et C. Riviere, Les Bambous. p. 93.
2) Die Zellschichtenzahl der inneren Rinde ist stets kleiner als die der äusseren.
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 445
schichtet und von feinen verästelten Kanälen förmlich durchsetzt
(Fig. 28). Im Querschnitt stellt demnach diese nach Aussen
zugekehrte C-Scheide ein symmetrisches Bild mit der oben er-
wähnten ebenso C-förmigen Aussenscheide dar'). Die 1 oder 2
innersten unmittelbar der Endodermis anliegenden Rindenschichten
sind bei Phyllostachys Kumasasa, Bambusa borealis und Arundinaria
quadrangularis als Verstärkungsring?) ausgebildet, indem die
Zellen durch innenseitige C-förmig verdickte und stark verholzte
Wände ausgezeichnet sind (Fig. 34).
So weit es den Bau der Wurzelrinde betrifft, zeigen die echten
Bambusa-Arten nämlich B. vulgaris, B. nana, B. stenostachya,
B. arundinacea u. a. ein von dem oben beschriebenen ganz ab-
weichendes Verhalten.‘ Hier weisen die Zellen der subepidermal-
schicht keine Wandverdickung auf und ebenso verhalten sich
die persistenten Epidermiszellen. Darauf folgen 2-3 Schichten
enger Bastelemente, welche nach innen scharf von den weit-
lumigen Rindenparenchymzellen abgesetzt werden (Fig. 26 u. 27).
Die äussere Rinde besteht nur aus einigen Zellschichten, während
die von grossen Lufträumen durchzogenen inneren Schichten viel-
fach dicker sind (Fig. 27). Die Endodermiszellen sind ringsum
verdickt und bilden die sogenannte O-Scheide’). Merkwürdig
ist ferner der Bau des Verstärkungsrings. Die innersten 1-oder
2-schichtigen Rindenparenchymzellen führen an ihren inneren
Wänden eine Anzahl unregelmässig gestalteter aus reiner Cellulose
bestehender Auswüchse, die häufig die äusseren Wände erreichen,
so dass sie im Querschnitt die ganzen Zellen nahezu aus-
1) Dasselbe Verhältnis wurde von Schwendener (Die Schützscheiden und ihre
Verstärkungen. Ges. Bot. Mitt. Bd. II, p. 120, 127.) auch bei einigen Orchideenluftwurzeln
bemerkt.
2)Schwendener, Die Schützscheide und ihre Verstärkungen. Ges. Bot. Mitt. p. 122
3) Vergl. Schwendener, Lc. p. 128, Tabelle. |
446 | K. SHIBATA :
zufüllen scheinen (Fig. 31 u. 32)'). Eine Anzahl einheimischer
Arten, die wegen ihrer 6 Stamen bisher in Bambusa eingereiht
wurden, z. B. B. Veitehii, B. palmata, B. borealis etc., weisen
jedoch im Bau der Wurzelrinde eine vollkommene Übereinstim-
mung mit Arundinaria auf und so schien es mir berechtigt, unter
Berücksichtigung noch anderer Merkmale,— vor allem: Fehlen der
in Bastbelege eingeschobenen Parenchymlamellen, die Gestalt der
Caryopse, die langkriechenden Rhizome u.s.w.—diese Formen-
gruppe von Bambusa loszutrennen und als eine neue Section in
Arundinariee aufzunehmen. Die Aufstellung dieser neuen
Formengruppe bietet uns doppeltes Interesse ; denn einmal erweist
dieselbe, dass die Eintheilung nach der Zahl der Stamen, auf welche
man in der Bambuseensystematik ein grosses Gewicht zu legen
pflegt, nicht immer durchführbar ist. Andererseits kommt diese
Formengruppe’) in ihrer Verbreitung auf Japan?) beschränkt vor.
Wir gehen nun zur Betrachtung des Centralcylinders über.
Die Anordnung der Leitbündel weicht, wie es von Boss‘)
nachgewiesen wurde, vom typischen Bau der Monocotylen ab. Zu
den normalen peripherischen radialen Bündeln, die ausserordent-
lich polyarch sind,’) kommt noch eine Anzahl der inneren iso-
lierten Hadrom- und Leptomstränge hinzu. Im Querschnitte
beliebiger junger Wurzeln bemerkt man innerhalb der dünnwandi-
gen Endodermis ein oder zwei Schichten des ununterbrochenen
Pericambiums (Fig. 23), dessen allgemeine Vorkommniss in Bam-
1) Derartige Structur findet man nicht in Sch wendener’s Aufzählung der verschiedenen
Verstärkungsformen (vergl. l.c. p. 132).
2) Die Anzahl der bis jetzt bekannten hierhergehôrigen Arten ist neun. Vergl. Makino.
Bambusacee Japonicæ. Bot. Mag. XIV, Nr. 156, p. 20.
3) Vielleicht auch in China.
4) Ross, Beiträge zur Anatomie abnorm. Monocotylenwurzel. Ber. d. D. B. G. Bd. J, p.37.
5) Z. B. in einer 4 mm dicken Wurzel von Phyllostachys mitis habe ich mehr als 150
gezihlt.
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 447
buseen um so mehr Beachtung verdient, als bei den meisten
Gramineen die primordialen Gefässe nach van Tieghem’) direct
der Endodermis anzustossen pflegen. Die inneren Hadromstränge
kommen in etwa drei concentrischen Ringen angeordnet vor.
Dazwischen liegen zerstreut die inneren Leptomstränge, welche
jedesmal aus den 1 oder 2 Siebröhren und den englumigen
Geleitzellen bestehen (Fig. 43 etc). Bei echten Bambusa-Arten
besitzt die stets einzeln stehende Siebröhre einen regelmässig
ovalen Umriss (Fig. 38). Die Gesammtanzahl der inneren Leptom-
stränge beträgt in den meisten Fällen, wie folgende Beispiele
lehren, eine Hälfte der peripherischen, aber bei echten Bambusa-
Arten kommen beide fast in gleich grosser Anzahl vor.
Peripherisches
Leptom
Phyllostachys mitis 84
P. bambusoides 83
2 puberula 47
Arundinaria japonica 75
Hindsii
Matsumurcæ 27
variabilis 30
pygmæa 43
palmata 41
Veilchii 29
ramosa 20
paniculata 39
nipponica 33
vulgaris 70
stenostachya 47
À nana 48
Dendrocalamus latiflorus 81
1) Van Tieghem, Les Racine. p. 123; Vergl. Morot, Recherche sur le Pericycle.
Ann. d. Sc. nat. Sèr. VI, T. 20, p. 233 und auch Falkenberg, Vergl. Unters. d. Vege-
tationsorgane. p. 192.
448 | K. SHIBATA :
Die tibrigen Elemente des Centralcylinders werden, abge-
sehen vom centralen Markparenchym, prosenchymatisch zugespitzt
und zugleich stark verdickt. So entsteht hier ein hohlecylindri-
scher mechanischer Ring, in welchem sämmtliche Leitstränge
eingebettet liegen. Hier muss noch eine Frage gelöst werden:
In welcher Weise geschieht die Communication zwischen den
einzelnen leitenden Elementen, die von einander getrennt im
mechanischen Gewebe liegen? Zwar hat Reinhardt!) die in Frage
kommenden Verhältnisse bei den anomal gebauten Wurzeln von
Musaceen, Pandanaceen, Palmeen und Cyclanthaceen ermittelt
und manch interessantes entdeckt. Betreffs der Communication
zwischen einzelnen Leptomsträngen in unserem Fall muss vor
allem bemerkt werden, dass die ausserordentlich stark verdickten
und verholzten Pericambiumzellen als die Leitungswege zwischen
den peripherischen Leptomsträngen kaum in Betracht kommen’).
Wenn man nun die Zahl der in beliebigen zwei Wurzelquersch-
nitten vorkommenden Leptomstränge sorgfältig mit einander
vergleicht, so kann man leicht eine bedeutende Abnahme der-
selben nach dem Wurzelspitze wahrnehmen, wie es aus einigen
beigefügten Beispielen hervorgeht :
Zahl der Leptomstränge in
Proximalende Mitte Distalende
22.5 cm langes Wurzelstück®) 128 — 104
12 is ss - 116 108 102
Der Umstand beruht bloss darauf, dass die inneren Lep-
tomstränge sich unter einander und mit den peripherischen im
weiteren Verlauf allmählig verschmelzen, wie man sich durch
Betrachtung successiver Querschnitte überzeugen kann. Die
1) Reinhardt, Das leitendegewebe einiger anomalgebauten Monocotylenwurzel. Jahrt.
£ wiss. Bot. Bd. XVI, p. 336.
2) Reinhardt, le. p. 361.
3) von Phyllostachys bambusoides.
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 449
Figuren 39 und 40 zeigen einige Fälle der erwähnten Verschmelz-
ung. Dieser Modus des Leptomverkehrs ist nach Reinhardt’-
schen Angaben auch bei anderen anomal gebauten Wurzeln häufig
verwirklicht‘). Der zweite Modus ist aber von mehr wirksamer
und auflälliger Art. Bei jeder Ansatzstelle der zahlreich ent-
springenden Nebenwurzeln an Centralcylinder werden sämmtliche
hier befindliche peripherische sowie verschieden tief liegende
innere Leptomstränge in einem System förmlicher Anastomosen-
bildung zusammengehalten, welche bei den meisten Arten sich
auf die etwa 10 periperischen Leptomstränge hinüberstreckt und
bei Bambusa-Arten sogar die Hälfte des ganzen Umfangs des
Centralcylinders in sich umfasst. Die etwas schematisierten
Figuren 35 und 36 illustrieren das obengesagte. Das hier die
Verbindung zwischen einzelnen Leptomsträngen herstellende
Gewebe besteht aus den plasmareichen parenchymatischen Zellen,
die mit den ansehnlich grossen Zellkernen und den dünnen un-
verholzten Wänden versehen sind (Fig. 43 u. 44). Die Versch-
melzung zweier Gefässe habe ich nur selten gesehen, während
bei der Ansatztelle der Nebenwurzel sämmtliche mechanische
Zellen zufolge reichlicher Tüpfelbildung uud häufiger auftretender
Querwände einen holzparenchymartigen Character annehmen und
demgemäss dem Saftaustausch zwischen den eingebetteten Gefässen
besser angepasst sind. Den directen Anschluss der Leptomelemente
an Holzparenchymzellen, wie es von Reinhardt für Musaceen
und Cyclanthaceen”) nachgewiesen wurde, habe ich auch häufig
bei Bambuswurzeln angetroffen (Fig. 41).
Die Basaltheile des Centralcylinders der Nebenwurzel sind
1) Reinhardt, Le. p. 364, p. 343 ete.
Vergl. Ross, Beitr. z. Anat. abnorm. Monocot. wurzel. p. 334.
2) Reinhardt, Le. p. 343, p. 346 und p. 348,
450 K. SHIBATA :
aus dem stark verdickten porösen rechteckigen parenchymati-
schen Zellen gebildet, durch welche die kurzen Basalglieder jedes
Leptomstrangs abwärts verlaufen, um sich dem oben erwähnten
Leptomanastomosencomplex der Hauptwurzel anzuschliessen
(Fig. 37). Hingegen scheinen die. Gefässe der Nebenwurzel
basalwärts meist blind zu endigen, so dass sie nur selten in
directen Zusammenhang mit denen der Hauptwurzel kommen.
Die Nebenwurzeln zeigen in ihrem Bau alle Merkmale der
bezüglichen Hauptwurzeln, dennoch fehlen ihnen stets die inneren
Hadrom- und Leptomstränge (Fig. 47 u. 45).
Die Ansetzung der Wurzeln an die Stammorgane geschiet
in der bei Monocotylen üblichen Weise). Die Elemente des
mechanischen Rings des Wurzelcentraleylinders breiten sich
scheibenförmig aus und verschmelzen sich mit den äussersten
Bündeln der Stammgebilde. Einzelne losgelöste Wurzelstränge
dringen noch weiter ein und schliessen sich den peripheren
Stammbündeln an, wobei das Leptom der ersteren solch eine
Umgestaltung erfährt, wie sie bei den Knospenbündeln beobachtet
wird?).
Es erübrigt noch einen interessanten Befund kurz zu
erwähnen. Die Rindenparenchymzellen der Nebenwurzeln, mit
Ausnahme von den innersten 2-3 Schichten kleinlumiger Zellen,
sind gewöhnlich von einem Pilz bewohnt, der in jedem Zelllumen
ein ansehnliches Knäuel von dicken verschlungenen Mycelfäden
bildet (Fig. 46 u. 47). Die verpilzten Wurzeln bieten trotzdem
ein ganz normales und gesundes Aussehen dar. Die Mycelfäden
treiben hie und da sogenannte Vesikulen aus und producieren
oft massenhaft gelbe körnige Substanz von nicht genau bekannter
1) Vergl. Falkenberg, Vergl. Unters. p. 196.
2) Vergl. p. 437.
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 451
chemischer Zusammensetzung. Die Stärke verschwindet gewöhn-
lich vom inficierten Gewebe. Es unterliegt also keinem Zweifel,
dass wir in diesem Fall mit einem endotrophischen Myco-
rhiza') zu thun haben. Der Wurzelpilz fehlte in keiner der von
mir untersuchten Arten und ist sowohl in den epidermlosen
Nebenwurzeln von Arundinaria- und Phyllostachys-Arten als
in den mit Epidermis versehenen Bambusa-Nebenwurzeln con-
stant nachweisbar. Die Rindengewebe der Hauptwurzeln habe
ich meist pilzfrei gefunden, abgesehen von den dünneren Wurzeln
von Arundinaria variabilis, Bambusa ramosa, ete. Die Lösung
der Frage nach der physiologischen Rolle’), die dieser Pilzsym-
biont in der Ernährung der Baumgräser spielt, will ich mir
für künftige Studien vorbehalten.
Dir BLATTGEBILDE.
Die in zwei entgegengesetzten Reihen gelegenen, breiten
Scheideblätter*) umhüllen übereinander den ganzen Schössling
und auch die wachsende Spitze des Rhizoms.
In der basalen, zum Knotengewebe übergehenden Region
jedes Scheideblattes weisen die Leitstränge in ihrem Hadrom kein
grosses getüpfeltes Gefäss auf, sondern sie besitzerf nur zahlreiche,
oft mehr als 15 Ring- oder Spiralgefässe, die mit einander man-
nigfach anastomosieren. In dem in der mittleren Partie des
Scheideblattes ausgeführten Querschnitte erblickt man parallel-
1) Frank, Lehrb. d. Botanik. Bd. J, p. 274; Uber neue Mycorlriza-Formen. Ber. d. D.
B. G. Bd. V, p. 400.
2) Es wurde neuerdings vielfach die Ansicht geäussert, dass die) Pflanzen die mit Mycorhiza
ausgerüstet sind, der Assimilation des freien Stickstofls befähigt seien. Vergl. Janse, Ann. d.
Jard. Bot. Buit. Vol. 14, p. 200, und auch Nobbe, Landw. Versuchs-St. Bd. LI, p. 241.
3) Rivière, Les Bambous, p. 76-82, p. 231.
452 K. SHIBATA :
verlaufende, abwechselnd starke und schwache Leitbiindel, die in
ihrem Bau kaum von den dem Stammorgan eigenen abweichen
(Fig. 48). Sie sind mit einander durch die aus einigen Siebröhr-
en und Gefiissen bestehenden Queranastomosen verbunden
(Fig. 53). Die Bastbelege auf der Leptomseite stossen gewöhnlich
unmittelbar an die stark verdickte Epidermis der Aussenfläche an
(Fig. 51), aber bei dicken fleischigen Scheideblättern der Phyllo-
stachys-Arten liegen fast alle Bündel mit ihren Bastbelegen ganz
frei im Parenchym (Fig. 52). Entgegengesetzt den stärkeren
Bündeln liegen die bandförmigen, meist 2-3 schichtigen Bast-
stränge an der Blattinnenseite. Die letzteren kommen bei Arundi-
naria Matsumure sonst auch an der Blattaussenseite hier und da vor
(Fig. 49). Das Scheideblattparenchym besteht aus dünnwandigen,
saftreichen Zellen,') von denen einige subepidermale Schichten bei
unterirdischen, harten Scheideblättern sclerenchymatisch verdickt
sind. Bei den derben oberirdischen Scheideblättern von Arundi-
naria-Arten tragen die an die Intercellularräume angrenzenden
Flächen der Parenchymzellen die eigenthümlichen bald kugel-
förmigen, bald stäbehenförmigen Auswüchse, die starke Holz-
reaction geben (Fig. 54).
Die Spaltöffnungen kommen an der Ober- sowie Unterseite
der Scheideblätter vor.
Die laubblatttragenden Blattscheiden stimmen in ihrem Bau
mit den oben geschilderten Niederblättern wesentlich überein.
Die Laubblätter der Bambuseen sind schon wiederholt von
vielen Forschern anatomisch untersucht worden. So haben
Kareltschikoft’), Magnus und Haberlandt die Armpallisa-
1) Die Parenchymzellen ausge wachsener Scheidebliitter enthalten fast keine Stärke, sondern
viel Glykose.
2) Kareltschikoff, Ub. d. faltenformig: Verdickungen in d. Zellen einiger Gramineen.
p. 180. (Referat).
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 453
dennatur des Assimilationsgewebes erkannt. Bei Güntz finden
wir Angaben über einige allgemeine Characteristik der Bambuseen-
blätter, dabei führte das energische Auftreten der mechanischen
Elemente ihn zur Aufstellung des ,, Bambuseentypus “ der
Gramineenblätter'). Bei dieser Sachlage würde es berechtigt sein,
dass ich mich hier nur auf einige kurze Notizen beschränke.
Um jedes Mestombündel bemerkt man zweierlei Scheiden’),
d.h. eine farblose Parenchymscheide und eine innere verholzte
Bastscheide (Fig. 55 u. 56). Die stets einschichtige Parenchym-
scheide fehlt selbst bei kleinsten Bündeln nicht; bei stärkeren
Bündeln ist sie oft dort stark verdickt und verholzt, wo sie an
subepidermale Bastrippen anschliesst. Die wenigstens um das
Leptom stets vorhandene Bastscheide wurde von Schwendener
als Mestomscheide ausgezeichnet?) und der echten Schutzscheide
zur Seite gestellt. Dieselbe ist um die kleineren Bündel meist
einschichtig, aber bei den stärkeren nicht selten mehr als 4
Schichten dick. Ferner verhalten die Elemente dieser Scheide
sich gegen Schwefelsäure kaum anders als gewöhnliche verholzte
Bastzellen, während sich die unverholzte Parenchymscheide gegen
dieses Reagens sehr widerstandsfähig erweist. So ist die in Rede
stehende Scheide als eine vereinfachte Form der das Mestom
vollkommen umschliessenden Bastscheide, wie ich sie schon bei den
Stielbündeln beschrieben habe, aufzufassen.
IV. Der Entwicklungsvorgang der Schösslinge.
Als Gegenstand der folgenden Darstellung diente mir Phyllo-
stachys mitis.
1) Güntz, Unters. üb. d. anat. Struct. d. Gramineenbl. p. 64.
2)Schwendener, Die Mestomscheiden der Gramineenblitter; Vergl. Strasburger,
Leitungsbahnen. p. 344.
3) Schwendener, lc. p. 178.
454 K. SHIBATA :
Die auf jedem Knoten der wachsenden Rhizomspitze angelegte
Knospe wird erst im nächsten Jahre zu einem kleinen Schossling
mit dem schon differenzierten, verholzten, ca. 1 cm langen Stiel
ausgebildet. Diesen letzteren nenne ich kurzweg den Schösling
des 2ten Stadiums, während die dem Knoten dicht anliegende,
stiellose Knospe als 1stes Stadium von diesem unterschieden
wird. Wenn man einen Querschnitt in der oberen Region dieses
kleinen Schösslings ausführt, so sieht man den Centralcylinder
gesondert in einen peripherischen, schmalen, bündelführenden Ring
und in umfangreiches Markgewebe, welches sich nach unten
allmählig verschmälert. Auf dem Längsschnitt sieht man dicht
unterhalb des Urmeristems vom Vegetationspunkt beginnend eine
grosse Anzahl abwechselnd stärkereiche und stärkearme Zonen,
welche letztere sich in späteren Stadien zu Internodien verlängern.
Der Schössling des 2ten Stadiums nimmt im Laufe des Som-
mers an Grösse zu und wächst im Spätherbst (October-November)
schon zu einem mittelgrossen Schössling des 3ten Stadiums. In
diesem Zustande verharrt er während des Winters.
Anscheinend schon in März tritt eine rasche Zunahme an
Grösse ein und im Anfang April erreichen die Schösslinge unter
der Erde eine ansehnliche Grösse, die ich als 4tes Stadium
kennzeichnete. Der Schössling ist mit zahlreichen geräumigen,
dicken Scheideblättern bedeckt. Der verholzte Stiel ist nun ca.
2 cm lang und 0.9-1.2 cm dick geworden. Die unteren, an den
Stiel sich direct anschliessenden, etwa zehn Internodien, deren
mittlere Höhe 1-2 cm beträgt, sind mit zahlreichen 3-4 mm dicken
und bis etwa 15 cm langen Wurzeln dicht besetzt. Ueber den
inneren Bau ist folgendes zu bemerken. Die Spitze, unterhalb
des Urmeristems, besteht aus abwechselnd stärkereichen und
stärkearmen Zonen, deren Zahl binnen 6 mm 40 beträgt.
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 455
Die Dicke der beiden Zonen nimmt nach unten zu, und erst in
Entfernung von 1.5-2.0 cm vom Vegetationspunkt entstehen im
internodialen Markgewebe sichtbare Querrisse, deren Weite und
Höhe in den nachfolgenden Internodien immer zunehmen. Diese
primordialen Markhöhlen erreichen in den unteren, mit Wurzeln
besetzten Internodien eine maximale Höhe von ca 3 mm, dabei
besitzt das Diaphragm eine Dicke von 2.5 mm. Die Bündel-
anlage in der Spitzenregion besteht aus engen procambialen
Zellen. Oberhalb der Stelle, wo die erste Markhöhle zum.
Vorschein kommt, erfolgt schon die Differenzierung in Elemente
des Biindels.
Indessen tritt die Spitze des Schosslings allmählig auf der
Erdoberfläche hervor; vom Ende April ab erfolgt dann ein
rasches Wachstum desselben. Schon in Mitte Mai erreichen
mehrere Schösslinge eine Höhe von 8-10 Meter, und an den
oberen Nodien findet die Entfaltung von blatttragenden Aesten
statt. Mehrere Internodien auf der Erde sind nur 6-9 cm
lang und nach oben nimmt die internodiale Länge graduell zu.
In mittlerer Höhe der Pflanze erreichen sie die Länge von 20
cm und mehr. Bis auf diese Region haben alle Internodien ihr
Längenwachstum vollendet. Mehrere darauf folgende Internodien
besitzen basale Wachstumszonen. Die von dort nach oben lie-
genden Internodien verjüngen sich allmählig zum Vegetations-
punkt. Die noch in Streckung begriffenen Internodien sind stets
mit Scheideblättern umhüllt. Der Schössling in diesem Zustande
ist im 5Sten Stadium.
Hier lasse ich einige Zahlenangaben folgen:
456 K. SHIBATA :
Stadium II. Stadium III. Stadium IV.
1: 7% oe | | 3 1
Aeussore | 44 (42 | 46 | 202| 198| 160 | ia
Curvatur
| 56.7 48.0| 485
Ana | | 3.4 139 | 184] 120 | | |
Curvatur
Maximalumfang incm.| 5.3
Gewicht in Gr.
Tägliche Zuwachsmessungen
an jedem Mittag vom 24 April
L. II. II. IV. |
Linge | £ | Länge | 2 | Lange | 5 | Länge | 5 |
Datum in 5 in s in a in | e :
cm. Sg m | © cm. Q em. § |
April 24 34.9 37.2 45.7 ge | |
25 38.1 | 321 431 | 59 51.3 | 5.6 310 38
26 442 | 61 50.6 | 7.5| 611 | 98 37.6 | 6.6 |
27 534 | 9.2 62.4 | 11.8 75.0 | 13.9 461 | 85
28 62.5 | 91! 733 | 109! 91.7 | 16.7| 566 105
29 741 | 116| 874 | 141| 1144 | 227 729 | 163
30 868 | 127) 1018 | 130) 1353 209) 868 | 139
Mei 1 106.9 | 20.1) 1239 | 226) 1640 | 287| 1088 | 220)
2 137.9 | 31.0| 1583 | 344] 2058 , 418] 1419 | 33.1.
3 175.9 | 38.0} 2024 | 441] 2533 | 47.5| 1801 ' 382
4 199.1 | 232| 230.7 | 2833| 2835 ' 302) 207.7 | 27.6
Bl Re — | 2771 | 464) 38364 | 52.9] 263.2 | 555!
6 300.5 | 507| 333.7 | 566] 3986 | 622| 3190 55.8
7 325.5 | 25.0} 363.7 | 300! 418.0 | 19.4| 3465 275 |
8 338.6 | 13.1| 375.6 | 119| 447.6 | 29.) 361.2 | 14.7
9 3825 | 43.9] 417.6 | 42.0] 5006 | 53.0] 4094 | 48.2:
10 423.9 | 414| 4645 | 47.0| 5469 | 463] 4526 | 432 |
11 4753 | 514| 515.6 | 511| 605.0 | 581] 5141 | 615
12 651.0 | 75.7| —— = | es — | 5979 | 838
13 | 5801 | 291| 6192 | 518| 7214 | 582| 6362 | 38.3
14 6316 | 515| 6619 | 427| 7864 | &0| 6646 | 284
15 719.6 | 880| 74.7 | 828| 846.1 | 59.7| 7107 | 461)
NB.—* Zuwachs ist Mittel von 2 Tagen.
*{ Nach Beobachtungen des hiesigen meteorologischen Observatoriums.
Anm. 1. Also bei diesen Messungen stieg der maximale Zuwachs nicht selten über 80 cm pro
Anm. 2. Das bisher bekannte stärkste Wachstum der Bambushalme beträgt 91.3 cm pro 24
Anm. 3. Auf die hier beobachteten auffälligen Wachstumsschwankungen und andere interes.
1) Die älteren Angaben über das Wachstum der Bambuspflanzen findet mam\bei Kraus
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 457
Um die erstaunlich grosse Schnelligkeit des Wachstums von
Bambusschösslingen in dem 5ten Stadium zu demonstrieren!),
führe ich hier einige von mir ausgeführte Messungen an Phyllo-
slachys mitis an,
von Phyllostachys-Halmen,
bis 15 Mai ausgeführt.
V VI.
| Linge a Länge à Mittlere M le
in E in 5 Wetterangaben. Tempera- Heidi:
cm. IS cm. 5 tur.** tate
106.6 Kar tleiser Wind 50.2
1212 |146| 526 klar-wenig’ trüb 73.0
144.8 | 23.6 76.3 | 23.7 klar 743
176.8 | 32.0 108.5 | 32.2 klar 69.8
208.3 | 315 | 140.4 | 319 wenig trüb 713
250.4 | 42.1 1844 | 44.0 klar, leiser Wind 43.5
284.8 34.4 216.8 | 32.4 klar, leiser Wind 64.9
34.6 | 39.8 | 260.3 | 43.5 Har, windig 73.0
378.4 | 53.8 322.0 | 61.7 klar, windstill. 66.0
4444 | 66.0| 388.9 | 66.9 Regen 80.0
484.2 | 39.8 429.2 | 40.3 klar, leiser Wind 91.4
551.7 | 67.5 | 4939 | 647 wenig trüb 86.0
633.9 | 82.2 566.7 | 72,8 Regen 82.6
672.4 | 38.5 607.0 | 40.3 Regen 93.3
713.1 | 40.7 625.4 | 18.4 halbklar, windig 86.6
755.5 | 42.4 675.4 | 50.0 halbklar, windstill. 76.1
7705 | 150 | 7389 | 63.5 klar, leiser Wind 81.2
klar, leiser Wind 17°.8 65.0
Regen 18°.3 84.8
klar, leiser Wind. 14°.6 89.6
| klar 18°,5 78.2
klar 20°.7 69.4
24 Stunden ! Riviére fand denselben bei Phyllostachys mitis in Algier 57 cm pro 24 Stunden.
Stunden. (Vergl. Pfeffer, Pflanzenphysiologie. Bd. II, p. 83.)
sante Fragen kann ich’an dieser Stelle nicht weiter eingehen (Vergl. Kraus, Le.)
(Ann. d. Jard. Bot. de. Buit. vol. XII. p. 197.) zusammengestellt.
458 K. SHIBATA :
V. Verhalten der Baustoffe
während der Entwicklung der Schösslinge.
In diesem Kapitel will ich die Umwandlungs- und Wander-
ungsvorgänge verschiedener Baustoffe während der Entwicklung
der Bambusschösslinge in wesentlichen Zügen darzustellen ver-
suchen.
DIE RESERVESTOFFE.
Unter stickstofffreien Reservestoffen, die sich in Bambus-
pflanzen vorfinden, kommt die Stärke in erster Linie in Betracht
Im zeitigen Herbst, wo die Ablagerung der Reservestofie
schon stattgefunden hat, konnte ich sie in oberirdischen und
unterirdischen Theilen aller untersuchten Arten in wechselnden
Mengen auffinden. Die Phyllostachys-Arten, welche mit einem
umfangreichen, unterirdischen Rhizomsystem ausgerüstet sind,
pflegen nur sehr kleine Mengen der Stärke in ihren Halmparen-
chymzellen aufzuspeichern. Bei allen Arten wird die grösste
Menge der Stärke in Rhizomen und Wurzeln deponiert. Die
‘ Parenchymzellen der Knoten sind stets äusserst stärkereich und
die Blattscheiden einiger Arten speichern ebenfalls die Stärke im
Parenchym auf. Die Siebröhren sind dagegen höchst inhaltarn ;
ich habe nur selten winzige Stärkekörner in denselben nach-
gewiesen. Es waren aber kleine Mengen von Glykose und
Rohrzucker stets vorhanden. Beachtenswerth ist die Gestalt der
Stärkekörner ; bei Bumbusa palmata, B. Veilchii und B. pani-
culata sind sie aus zahlreichen kleinen Theilkörnern zusammen-
gesetzt (Fig. 57). Solche polyadelphische Stärkekörner') kommen
nach Nägeli in Stammgebilden nur selten vor.
1) A. Meyer, Untersuchungen über die Stärkekörner. p. 204.
WACHSTUMGSESCHICHTE D. BAMBUSGEWAECHSE. 459
Der reducierende Zucker ist ziemlich reichlich in Halmen
und Rhizomen im Winterzustande nachzuweisen. Die winzigen
Fetttröpfchen sind oft im Halmparenchym von Phyllostachys
mitts, Arundinaria Simoni, Arundinaria Hindsi u.s.w. ange-
troffen, aber sie kommen jedenfalls als Reservestoffe kaum in
Betracht. |
Um eine Vorstellung über die Mengenverhältnisse der auf-
gespeicherten Stärke zu anderweitigen DBestandtheilen der
Reservestoffbehälter zu gewinnen, habe ich einige Analysen der
zweijährigen Rhizome von Phyllostachys mitis ausgeführt‘). Es
ergab folgendes :
% Gehalt der
Trockensubstanz.
Stärke ..... D nok
Reducierender Zucker......eer...... 0.95
Nicht reducierender Zucker...... 4.31
Rohproteinstoffe (N x 6.25) ...... 5.41
Kette en nee DOL
Rohfaser ..........,....,... RE N I)
Asche ....... LINE AR SD e
Unbestimmte Stoffe (Differenz)... 8.65
100.00
1) Das am 25 November gesammelte, kräftige Rhizomstiick von Phyllostachys mitis, dessen
Parenchym sich zuvor bei microscopischer Beobachtung als von Stärke strotzend erwies, wurde
mittelst des Hobels abgeschaubt, schnell bei 70°-80° getrocknet, und zu einem feinen Schrot
gemahlen. Von diesem lufttrockenen Rhizomschrot wurde ein bestimmtes Quantum abge-
wogen und zu jeder Bestimmung verwendet.
Das Trockengewicht des Schrots wurde nach weiterem 4 stündigen Trocknen bei 100°
(zur Gewichtsconstanz) bestimmt.
Die Stärke wurde mittelst der Erhitzung im Soxhlet’schen Autoclave verzuckert.
Die löslichen Kohlehydrate wurden nach 5-6 maligem Ausziehen mit kaltem
Wasser binnen 24 Stunden erschöpft. Der nichtreducierende Zucker wurde nach
Inversion mit verdünnter Schwefelsäure bestimmt. Alle Bestimmungen der Zucker wurden
nach Meissl- Allihn’scher Gewichtsmethode ausgeführt.
Der Gesammtstickstoffwurdenach Kjeldah] und der Eiweisssticks off nach
Stutzer bestimmt.
Die Fasersubstanz wurde durch Weender’sches Verfahren bestimmt.
Das Atherextract wurde ohne weiteres als Oel angenommen.
460 K. SHIBATA :
%
Man sieht also, dass die Stärke wohl als Hauptreservestof
zu betrachten ist, dagegen sind die Proteinstoffe in ver-
hältnissmässig geringer Menge vorhanden.
Nun schien es mir erwünscht zu wissen, eine wie weite Strecke
des Rhizoms zum Auswachsen eines Schösslings dienen sollte, so
habe ich im Anfang Februar von einer Plantation von Phyllostachys
bambusoides eine Anzahl Rhizomstücke ausgegraben und die auf
Knoten vorkommenden Schösslinge (im 3ten Stadium) aufgezählt.
Es ergab folgendes Resultat :
Zahl Gesammtanzahl
der Rhizomstücke | der Internodien Rhizomzweige Schösslinge
69 632 39 15
Aus obigem berechnete ich das Zahlenverhältniss der Rhizom-
internodien zu einem Schössling, wie 42.1:1.
KOHLEHYDRATE.
Die stickstofffreien Reservestoffe in allen untersuchten Arten
bestehen, wie schon erwähnt, hauptsächlich aus der Stärke. Dass
der Stärkegehalt der Rhizome eine bemerkbare Verminderung
während des Winters erleidet, wie es von Rosenberg!) für einige
perennierende Gewächse dargethan wurde, konnte ich nicht in
diesem Fall bestätigen, da ich grosse Anzahl von Rhizomen von
Phyllostachys mitis, Phyllostachys bambusoides, Phyllostachys Kuma-
sasa, Arundinaria Hindsit, Arundinaria Narihira, Arundinarv
quadrangularis, Arundinaria Matsumure, Bambusa palmata,
Bambusa nana u.s.w. im Winter (Anfang Januar—Ende Februar)
1) Rosenberg, Die Stärke im Winter. Pot. Centralbl. Bd. LXVI, p. 837.
WACHSTUMGSESCHICHTE D. BAMBUSGEWAECHSE. 461
untersuchte und dabei keine merkliche Differenz in Bezug auf
Stärkegehalt von den im Herbst beobachteten Exemplaren auf-
finden konnte. Auch die Wurzeln der untersuchten Arten
enthielten in dieser Jahreszeit grosse Mengen von Stärke in
ihrem Rinden- und Markparenchym, so z. B. bei den am 25.
Januar gesammelten Exemplaren :
| Phyllostachys , Phyllostachys | Arundinaria | Arundinaris
! milis | bambusoides Hindsii Narihira
Rindenparenchym 4
Markparenchym 3
Gleiches gilt für die oberirdische Halme von Arundinaria-
und Bambusa-Arten*) Merkwürdigerweise nimmt die Menge des
reducierenden Zuckers während des Winters unverkennbar zu.
Sodann kann man leicht mehr oder minder bedeutende Mengen
desselben im Halm- und Rhizomparenchym obengenannter Arten
nachweisen.)
Aber in Stadium IV, wo die unterirdischen Schösslinge ein
1) Bequemlichkeitshalber habe ich zur Bezeichnung des Stürkegehaltes folgende Ziffern
benutzt:
0—bei gänzlicher Abwesenheit von Stärke;
1—wenn ein Theil des Gewebes stürkefrei ist, wührend der andere wenige Körnchen
in den Zellen führt;
2—wenn alle oder die meisten Zellen wenige Stärkekörner enthalten ;
3— wenn ein Theil der stärkeführenden Zellen wenige stürkekörner enthält, während
der andere recht viel Stärke führt;
4—wenn das Gewebe recht viel Stärke enthält ;
5—wenn alle oder die meisten Zellen strotzend gefüllt sind.
2) Vergl. A. Fischer, Beiträge zur Physiologie der Holzgewächse. Jahrb. f. wiss. Bot.
Bd. XXII, p. 92, p. 112.
3) Dieser Zucker geht aber grösstentheils schon im Anfang Miirz wieder verloren, ohne
duss dabei eme bemerkbare Stärkezunahme stattfand. Auch fielen die Versuche den Zucker
in der abgeschnittenen Halmtheilen durch künstliche Erwärmung (im Treibhaus bei 17°-20°C.)
zur Stärke überzuführen negativ aus.
462 K. SHIBATA :
rasches Wachstum begannen, ist die deutliche Starkezunahme
in mehreren Rhizominternodien in der Nähe von Knoten, an
welchen der wachsende Schössling sitzt, zu beobachten. So
z.B. bei Phyllostachys mitis :
Anfang , Ende Anfang | Anfang ' Mitte
Februar : April
Rindenparenchym
Centralcylinderparen-
chym
Markparenchym
Diese Stärkezunahme mag jedoch darauf beruhen, dass
die von ferneren Theilen des Rhizoms in Form von Zucker
zugeführten Kohlehydrate hier in der Nähe des Schösslings
transitorisch in Stärke umgewandelt werden. Dafür sprechen die
Umstände, dass erstens in entfernteren Rhizominternodien keine
entsprechende Stärkezunahme stattfand, und zweitens schon in
dieser Zeit ein Blutungssaft, der eine wichtige Rolle beim
Zuckertransport spielt, von jeder beliebigen Schnittfläche des
Rhizoms hervorquillt.
Die Stärkezunahme ist vor allem im verholzten Stieltheile
des Schösslings ausgeprägt :
| Ende Anfang Anfang
| December Februar März
Tr Fe el | er on
Rindenparenchym
Centralyclinderparen-
chym
Hadromparenchym
Gleichzeitig wurde die partielle Entleerung der Rhizomknoten,
an welche die Schösslinge sitzen, beobachtet, obgleich die nächst
folgenden Internodien, wie schon bemerkt, noch von Stärke
erfüllt waren.
WACHSTUMGSESCHICHTE D. BAMBUSGEWAECHSE. 463
Von jetzt ab wird die Stärke im Rhizome nach und nach
aufgelöst und schon in dem Stadium V, wo die Schösslinge auf
der Erde 4-6 Meter hoch wuchsen, verschwinden fast sämmtliche
Stärkekörner vom Parenchym der benachbarten Rhizominter-
nodien. So zum Beispiel bei Phyllostachys mitis :
Rhizominternodien—
Rindenparenchym
Centralcylinderparenchym
Markparenchym
N odium—
Subepidermale sclerotische
Schicht
Rindenparenchym
Centralcylinderparench ym
Stiel des Schösslings—
16. April
Subepidermale sclerotische 0 ° 0 0
chic
Rindenparenchym
ex
7
©
©
Centralcylinderparenchym
Hier findet auch in der Wurzel eine entsprechende Stärkeent-
leerung statt:
| 16. April ı 19. Mai
|
Rinden- eh grosszelliges | 5 | 0
parenchyml;nneres kleinzelliges : i) 1-2
Markparenchym
464 K. SHIBATA :
Merkwürdigerweise konnte ich bei so raschem Auflösen der
Stärke eine entsprechende Glykosebildung im Parenchym nicht
beobachten ; bei der Zuckerprobe nach Schimper erhielt ich
nur Spuren von Oxydulkörnern in Parenchymzellen.
Die Kohlehydrate in wachsenden Schösslingen verhalten
sich im Grossen und Ganzen analog mit denjenigen in von
Sachs, H. de Vries u. A. untersuchten Pflanzen.') Indes
ist folgendes noch zu bemerken.
Die Abwesenheit von Stärke im Urmeristem des Vegetations-
punktes habe ich im allgemeinen constatiert. Die feinkörnige
Stärke wird erst an der Stelle, wo die erste Differenzierung der
Bündelanlage und des Grundparenchyms auftritt, nachweisbar
und man kann abwechselnd stärkereiche und stärkearme Zonen
deutlich sehen. Nach unten tritt der Unterschied im Stärke-
gehalt dieser abwechselnden Zonen immer schärfer hervor. Der
reducierende Zucker tritt an der Spitze weiter unten als Stärke
auf und zwar zuerst in dem Marke der internodialen Zone, wo der
erste Anfang der Zellstreckung sich durch Zerreissen von Gewebe
kund thut. Selbst die fertig gestreckten, unteren Internodien des
mehrere Meter hohen Schösslings bleiben noch lange Zeit von der
Glykose erfüllt. Sobald die Streckung eines Internodiums voll-
endet ist, verschwindet die Stärke aus dem Parenchym, abgesehen
von einigen winzigen Körnchen in 2-3 Zellen bei Durchlassstellen
1) Hier seien nur folgende erwähnt:
Sachs, Physiologische Untersuchung üb. d. Keimung von Schminkbohne.—Sachs,
Keimungsgeschichte der Gräser.—De Vries, Wachstumsgeschichte der Zuckerrübe.—
De Vries, Keimungsgeschichte der Kartoflelknollen.—Detmer, Vergl. Physiologie
d. Keimungsprocess der Samen. —Hoflmann, Uber d. Stoffwanderung bei d. Keimang
von Weizen- und Kleesamen —A. F. A. C. Went, Chemisch-physiologische Unter-
suchungen üb. d. Zuckerrohr.
2) Vergl. Sachs, Üb. d. Stoffe welche d. Material z. Wachstum d. Zellhäute liefern.
Jahrb. f. wiss. Bot. Bd. III, p. 207.
WACHSTUMGSESCHICHTE D. BAMBUSGEWAECHSE. 465
der Biindel. Da hierbei sämmtliche Zellwände noch keine
nennenswerthe Verdickung zeigen, so erfolgt die weitere Aus-
bildung der Bündelelemente ohne Gegenwart der umgebenden
Stärkescheide, in welcher nach Heine’) die nöthigen Baustoffe
als Stärke deponiert werden sollen. Die besonders starke Zucker-
ansammlung in einigen Parenchymschichten um die in Ausbildung
begriffenen Bastbelege herum vertritt hier die Stelle der fehlenden
Stärkescheide und daher mag sie als Zuckerscheide’) bezeichnet
werden.
In der wachsenden Wurzel bemerkt man die kleinste Menge
der winzigen Stärkekörner nur in der noch zartwandigen Endo-
dermis. Hingegen ist in der Haube von der ersten Anlage die
Stärke in ihren Zellen festgehalten”) Der reducierende Zucker
kommt in der ganzen Länge der Wurzel, ausser der 4-5 mm
langen Strecke der Spitze und der Haube, reichlich vor. Erst in
der ca. 40 cm lang gewachsenen Wurzel wird die Abnahme und
zuletzt das Verschwinden vom Zucker an der Wurzelbasis
bemerkbar.
Wie schon erwähnt konute ich in Rhizomen und Wurzeln,
wo die Reservestärke in Auflösung begriffen war, gewöhnlich nur
eine Spur von Glykose auffinden. Analoge Fälle sind bereits
bekannt. So z.B. gelangte es Sachs nicht, in Cotyledonen der
keimenden Phaseolus- Samen, in Schildchen von Triticum und
Zea und auch in Funiculus verschiedener Samen die Glykose
nachzuweisen‘), obgleich hier das Vorhandensein der gelösten
1) Heine, Die physiologische Bedeutung der sogenannten Stärkescheide. Landw.
Versuchs-St. 1888. p. 115.
2) H. de Vries hat früher den Ausdruck im analogen Sinne mit „ Leitscheide “
Schimpers angewandt.
3) Vergl. Sachs, Jahrb. f. wiss. Bot. Bd. III, p. 203.
4) Sachs, Über die Stoffe, welche das Material zum Wachstum der Zellhäute liefern,
Jahrb. f. wiss. Botanik. Bd. III, p. 248.
466 K. SHIBATA :
Kohlehydrate von vornherein erwartet werden musste. In
gewissen Fällen dieser Art ist es nicht unwahrscheinlich, dass
die Glykose durch andere lösliche Kohlehydrate ersetzt wird.')
In treibenden Rhizomen von Phyllostachys mitis habe ich
nun den Rohrzucker in stärkehaltigen Parenchymzellen mittelst
der Invertin-Methode nachgewiesen. Ferner in Rhizomen, von
welchen fast alle Stärkekörner schon verschwunden waren
(Stadium V), beobachtete ich noch erhebliche Mengen Rohrzucker.
Uebrigens habe ich in folgenden Arten den Rohrzucker in Rhizo-
men während des Austreibens der Schösslinge beobachtet : PAyllo-
stachys bambusoides, Phyllostachys puberula und Arundinaria
japonica; und in folgenden im Halmparenchym während des
Austreibens der Zweigknospen : Bambusa palmata und Arundi-
naria japonica. Ferner erhielt ich die Rohrzucker-Reaction in
Halmen von Phyllostachys Kumasasa, Arundinaria Simoni, und
Arundinaria Hindsii var. graminea. In den Wurzeln von
Phyllostachys mitis, deren grosskörnige Reservestärke in Auflösung
begriffen war, konnte ich ebenfalls Rohrzucker nachweisen.
Gleiches gilt für verholzte Stieltheile der Schösslinge. In allen
diesen Fällen kommt Rohrzucker hauptsächlich im Parenchym
und viel weiniger in Siebröhren vor. Nun liegt mir der Gedanke
nahe, dass in diesem Falle die Kohlehydrate hauptsächlich in
Form des Rohrzuckers von Zelle zu Zelle wandern’).
Vom Rohrzucker ist noch zu erwähnen, dass ich ihn im
1) So z. B..wurde für Gramineen-Scutellum das Vorhandensein vom Rohrzucker anstatt
Glykose von Grüss auf microchemischem Wege sowie/auf experimentelle Weise sicherge-
stellt. (Vergl. Ber. d. D. B. G. Bd. XVI, p.17.) Puriewitsch (Jahrb. f. wis Bot. Bd.
XXXI, p. 53.) hat bei der ersten Periode der Entleerung der Reservestärke das Auftreten
nichtreducierenden Zuckers beobachtet. Vergl. Leclare du Sablon, Recherche sur les
Reserve Hydrocarbones des Bulbes et des Tubercules. Rev. gen. d. Bot. 1899.
2) Vergl. E. Schulze, Ueber die Verbreitung des Rohrzuckers in den Pflanzen und
über seine physiologische Rolle. Zeitschrift f. physiol. Chemie. Bd. XX, p. 552.
|
|
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 467
jungen Gewebe unterhalb. des Urmeristems, wo schon eine
Differenzierung in nodiale und internodiale Zonen stattgefunden
hat, manchmal, wenn auch nicht immer, durch Invertin-
Methode nachgewiesen habe’). | |
EIWEISS UND AMIDOVERBINDUNGEN.
Im Jahre 1872 hat Pfeffer’) zuerst die hohe Bedeutung des
Asparagins in der Translocation und Bildung von Eiweiss beim
Keimen von Lupinus luteus und einigen anderen Papilionaceen
auf microchemischem Wege nachgewiesen. Er hat nämlich con-
statiert, dass das Asparagin als Auflösungsproduct des Reserve-
proteins in Cotyledonen entsteht und dann wachsenden Theilen
zugeführt wird, und ferner, dass wenn Kohlehydrate bei der
Assimilation sich vermehren, das gebildete Asparagin zum Eiweiss
regeneriert wird’). Die letzterwähnte Thatsache hat er später
durch die Kulturversuche im Dunkeln und in kohlensäurefreier
Luft weiter begründet‘) Seit diesen zum ersten Male exact
ausgeführten Arbeiten Pfeffer’s wurde die Frage nach Eiweiss-
umsetzungen mit immer wachsender Eifrigkeit seitens der
Botaniker und Chemiker verfolgt, was zu zahlreichen Arbeiten
Veranlassung gab. Schulze und seinen Schülern verdanken
wir besonders eine Reihe der Versuche über jene Amidover-
1) Ich habe im Gewebe dieser Region eine schöne rosarothe Färbung mit conc.
Schwefelsäure erzielt. Diese Reaction deutet darauf hin, dass hier ein lösliches Kohlehydrat
neben Eiweiss vorhanden ist. Vergl. Frankfurt, Zur Kenntniss der chemischen Zusam-
mensetznng des ruhenden Keimes von Triticum vulgare. Landw. Versuchs-St, 1896. p. 461.
2) Pfeffer, Untersuchungen über die Proteinkörner und die Bedeutung des Asparagins.
Jahrb. f. wiss. Bot. Bd. VIII, p. 429.
3) Pfeffer, Le. p. 558.
4) Pfeffer, Über die Beziehung des Lichtes zur Regeneration von Eiweissstoffen aus dem
beim Keimungsprocess gebildeten Asparagin. Monatsber. d. Acad. d. Wiss. z. Berlin. Dec.
1873.
468 K. SHIBATA:
bindungen und Hexonbasen, wie Glutamin, Leucin, Tyrosin,
Phenylalanin, Arginin u.s.w., die neben Asparagin beim
Eiweissumsatz auftreten. Namentlich hat Schulze schon in
einer im Jahre 1878 publicierten Arbeit die Ansicht geäussert,
dass in Lupinenkeimlingen andere Nichteiweiss- Verbindungen,
die neben Asparagin auftreten, auch zur Regeneration des
Eiweisses dienen müssen‘). In demselben Jahre hat Borodin
eine allgemeine Verbreitung von Asparagin im Pflanzen-
reiche festgestellt, dabei sprach er aus: „Sobald irgend ein
lebenskräftiger Theil irgend einer Pflanze arm an stickstoff-
freien Substanzen wird, sieht man in ihm Asparagin als
Zersetzungsproduct des Eiweisses auftreten und sich mit der
Zeit immer mehr anhäufen.‘“) Demnächst fand Kellner‘) in
jungen Theilen der Gräser eine bedeutende Menge Amide, und
äusserte zuerst die Ansicht, dass die Amide durch Synthese aus
anorganischen Stickstoffverbindungen entstehen. Auch Horn-
berger‘) meinte, dass die Amide, die in Maiskeimpflanzen auf-
treten, synthetische Producte seien. Suzuki’) hat angegeben, dass
er bei Einführung von anorganischen Salzen wie Ammoniumnitrat
und Natriumnitrat in verschiedenen Pflanzen eine Asparagin-
bildung bewerkstelligen konnte. Emmerling*) hat auch Amido-
säuren als synthetische Producte angesehen, aber es fehlt an
einem experimentellen Beweis. So kann die Bildung des Aspara-
1)E. Schulze, Über Zersetzung und Neubildung der Eiweissstoffe bei der Keimung von
gelber Lupine. (Jahresber. f. Agr. Chem. 1878. p. 211).
2) Borodin, Über die physiologische Rolle und die Verbreitung des Asparagins in
Pflanzenreich. Bot. Zeit. 1878. p. 826.
3) Kellner, Landw. Jahrbücher. Bd. VIII. Suppl. 1879.
4) Hornberger, Chemische Untersuchung über das Wachstum der Maispflanze. Landr.
Jahrb. 1882.
5)Suzuki, On the Formation of Asparagin in Plants under different Conditions. Ball.
of the College of Agriculture. Bd. II, p. 409.
6) Emmerling; Studien über Eiweissbildung in der Pflanze. Landw. Versuchsst. 1887.
p. 7.
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 469
gins und der anderen Amidokörper entweder durch Zerfall des
Eiweisses oder durch geeignete Synthese erfolgen. Ob diese
oder jene geschieht muss von Fall zu Fall bestimmt werden.
Jedenfalls ist es seit Pfeffer’s bahnbrechender Untersuchung
klar, dass die Amide und Amidosäuren, deren Entstehungen in
verschiedenen Fällen verschieden sein können, nachher für
Eiweissregeneration verbraucht werden. Gegenwärtig ist es aber
noch nicht sicher ob verschiedene Amide und Amidosäuren ganz
gleichwerthig für Eiweissregeneration dienen. Zwar hat Hans-
teen!) in seiner interessanten Arbeit gezeigt, dass bei Lemna
minor verschiedene, künstlich eingeführte Amide und Amido-
säuren je nach der Qualität der disponiblen Kohlehydrate sich
für Eiweissbildung verschieden verhalten. So liegt der Gedanke
nahe, dass die in bestimmten Keimpflanzen auftretenden Amido-
körper auch ungleichen Werth für Eiweissbildung besitzen.
Früher war Schulze?) der Meinung, dass das Asparagin schwerer
verwendbar als andere Amidokörper ist und daher in Keim-
pflanzen zur Anhäufung kommt. Aber Loew’) behauptete, dass
das Asparagin dem Eiweiss näher steht als andere Amidokörper,
und vermuthete auch, dass die letzteren weiter zerfallen unter
Bildung von Formaldehyd und Ammoniak, aus denen durch
synthetische Processe Asparagin entsteht. Erst neulich ist
Schulze‘) zu einer ähnlichen Vorstellung gelangt. Er spricht
die Ansicht aus, dass das Asparagin (und auch Glutamin) in den
1) Hansteen, Beiträge zur Kenntniss der Eiweissbildung und die Bedingung der Rea-
lisirung. Ber. d. D. B.G. Bd. XIV, p. 362.
2) Schulze, Uber den Eiweissumsatz im Pfanzenorganismus. 1880. p. 30.
3) O. Loew, The-Energy of living Protoplasm. Bulletin of the College of Agriculture
Bd. II, p. 64.
4) Schulze, Uber den Umsatz der Eiweissstoffe in den lebenden Pflanzen. Zeit. f.
physiol. Chemie. Bd. XXIV, p. 60.
Schulze, Uber die Bildungsweise des Asparagins in den Pflanzen. Landw. Jahrb.
1898. p. 509; p. 513.
470 K. SHIBATA:
Keimpflanzen zum grossen Theil durch Umwandlung der Amido-
säuren, die als directe Eiweisszersetzungsproducte betrachtel
werden können, entstehen, und dass die Amidosäuren einmal zu
leicht verwendbaren Amiden’) übergeführt werden, bevor sie sich
in Eiweiss verwandeln.) Bei dieser Sachlage ist es wünschens-
werth im concreten Falle die Localisation und das Verhalten
von Amiden und Amidosäuren zu verfolgen und damit einiger-
massen Aufschlüsse über die Beziehung zur Eiweissregenera-
tion der beiden verschiedenen Stoffe zu gewinnen.
Bei dem vorliegenden Falle der Entwicklung der Bambus-
schösslinge kommen Tyrosin und Asparagin reichlich vor.
Die genannten Vertreter von beiden Stoffgruppen sind glück-
licherweise leicht auf microchemischem Wege bestimmbar.
Hier lasse ich ältere Angaben über das Vorkommen des
Tyrosins vorangehen. Gorup-Besanetz) fand es zuerst im
Wickenkeimlinge. Schulze und Barbieri‘) fanden es in etwas
grösserer Menge in Kürbiskeimlingen. Auch in Lupinenkeim-
lingen scheint es nicht zu fehlen, da Belzung?) aus Extract der
1) Eine entgegengesetzte Meinung, dass das Asparagin ein für Eiweissregeneration wenig
geeignetes Material sei, wurde neuerdings wieder von Prianischnikow (Landw. Versuchs-
St. 1899. Bd. LII, p. 347 ff.) vertreten.
2) Unter neueren Publicationen über Eiweisssynthese, die nach Vollendung meines
Manuscriptes in meine Hand gelangten, seien nur folgende zu erwähnen:
Prianischnikow, Eiweisszerfall und Athmung in ihren gegenseitigen Verhältnissen.
Landw. Versuchs-8t. Bd. LII, p. 137.
Hansteen, Über Eiweisssynthese in grünen Phanerogamen. Jahrb. f. wiss. Bot.
Bd. XXXIH, p. 417.
Prianischnikow, Die Rückbildung der Eiweissstoffe aus deren Zerfallsproducten.
Landw. Versuchsst. 1899. p. 347.
Schulze, Uber Eiweisszerfall und Eiweissbildung in der Pflanze. Ber. d. B, G. 1900.
Heft. 2. p. 36.
Emmerling, Studien über die Eiweissbildung in der Pflanze. Landw. Versuchs-St
Bd. LIV, p. 215.
3) Gorup-Besanetz, Ber. d. D. C. G. VII, p. 146; p. 569.
4) Landw. Jahrb. Bd. VII, p. 431.
5) Belzung, Recherche sur 1. Germination etc. Ann. d. Sc. nat. Bot. Ser. VI, T. 19.
p- 234.
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 471
Keimlinge von Lupinus luteus Tyrosinkrystalle isolieren konnte,
obwohl ihm der microchemische Nachweis des Tyrosins nicht
gelang. Ferner fand es Schulze‘) in Cotyledonen keimender
Lupinusarten, etiolierten Keimlingen der Zupinus angustifolius,
Endosperm von Ricinus communis und etiolierten Pflanzen von
Tropaeolum majus. In allen diesen Fällen ist die Menge des
gefundenen Tyrosins immer sehr gering, so dass man auf micro-
chemische Verfolgung desselben verzichten muss. Schulze
bemerkte, dass der Grund des geringen Vorkommens von Tyrosin
darin liegt, dass es eine viel regere und schnell verlaufende Um-
wandlung erleidet.”) In unterirdischen Pflanzentheilen ist Tyrosin
öfters auf chemischem Wege gefunden. Schulze und Barbieri
fanden Tyrosin neben Leucin in den Kartoffelknollen und in der
Wurzel von Beta vulgaris’) Auch Planta‘) fand es in den
Knollen von Stachys tuberifera. In der botanischen Litteratur
finden wir nur vereinzelte Angaben. Prantl?) hat Krystalle, die
wie Tyrosin reagierten, aus in Alcohol aufbewahrten Stengeln von
Dahlia variabilis erhalten. Borodin‘) fand in Blättern der
etiolierten Kartoffel, die mit absolutem Alcohol behandelt wurden,
Tyrosinkrystalle. Ferner fand er dergleichen in Vicia sativa,
Tropaeolum majus etc. Aber es ist ‘hier zu bemerken, dass diese
Befunde ausschliesslich von abgeschnittenen und in Wasser weiter
cultivierten Zweigen herrührten und gleichzeitige chemische
1) Schulze, Üb. d. Umsatz d. Eiweissstoffe in d. leb. Pflanze. Zeit. f. physiol. Chemie.
Bd. XXIV, p. 58.
2)Schulze, Le. p. 50.
3) Vergl. Schulze, Über den Eiweissumsatz im Pflanzenorganismus. 1880. p. 24.
4) Pianta, Über die Zusammensetzung der Knollen von Stachys tuberifera. Landw.
Versuchs-St. Bd. 35, p. 473.
Vergl. ferner Schulze, Zeits. f. physiol. Chemie. Bd. XXIV, p. 85.
5)Prantl, Das Inulin. 1870. p. 61.
6) Borodin, Über die physiologische Rolle und die Verbreitung des Asparagins. Bot.
Zeit. 1878. p. 819.
472 2 K. SHIBATA:
Belege fehlten. Erst später hat er’) einmal in normalen, jungen
Dahlia-Blättern Tyrosin aufgefunden. Noch später hat Leitgeb’)
den Gehalt der Dahlia-Knollen an Asparagin und Tyrosin
constatiert.
Bevor ich zur Besprechung meiner Beobachtungen fortsch-
reite, will ich hier die Ergebnisse von chemischen Untersuchun-
gen Kozai’s’) kurz erwähnen. Er hat die Analyse des Schöss-
lings (Stadium IV) von Phyllostachys mitis ausgeführt ; sie ergab
folgendes :
% Gehalt der
Trockensubstanz.
Rohproteinstoffe.........,.....,...…. 25.12
Bette assidu 2.49
Röhfaser.. sea Siehe 11.60
DEATKE an 3.33
(PLY KORG: ana 8.15
Andere N-freie ext. Stoffe...... 30.49
Asche ............, ne ane 9,22
Unbestimmbare Stoffe........ we. 9.60
100.00
Fiir die Vertheilung des Stickstoffs auf Proteinstoffe und
nichtproteinartige Verbindungen ergaben sich folgende Zahlen:
N in Proteinstoffen ........,.............. 1.2296 der Trockensubstanz
N in nichtproteinartigen Stoffen 2.82% ,,
Gesammtstickstoff ...........0:000004.04%
EL
1». ”
So sieht man, dass die Schösslinge grosse Mengen von
stickstoffhaltigen Substanzen enthalten, im auffallenden Gegen-
1) Borodin, Uber einige bei Bearbeitung von Pflanzenschnitte mit Alcohol entstehende
Niederschlag. Bot. Zeit. 1882. p. 589.
2) Leitgeb, Der Gehalt der Dahliaknollen an Asparagin und Tyrosin. Mittheil. a. d.
bot. Inst. z. Graz. 1888. p. 222.
3) Kozai, On the nitrogenous non-albuminous Constituents of Bamboo shoots- Bulletin
of the College of Agriculture. Vol. I. No. 7.
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 473
satz zum Rhizom, und insbesondere kommen die nichteiweissar-
tigen Verbindungen in überwiegender Quantität vor. Kozai hat
nach dem Schulze’schen Quecksilbernitratverfahren die seiden-
glänzenden Nadelkrystalle aus dem Wasserauszug von Schösslingen
erhalten, welche mit Sicherheit mit Tyrosin identificiert wurden.
Ferner hat er auch das Asparagin isoliert und durch verschie-
dene Reactionen und Bestimmung der Stickstoffzahl sicher nach-
gewiesen.
Bei meinen Studien wurden die obengenannten Substanzen,
das Asparagin und das Tyrosin in ihrem Verhalten näher verfolgt.
Beide sind nach Borodin’scher Methode reichlich und sicher
nachweisbar, dabei scheint der vorhandene Zucker kein Hinder-
niss zur Krystallisation darzubieten.
Zunächst will ich das Verhalten von Asparagin und Tyrosin
bei der Entwicklung der Schösslinge von Phyllostachys mitis kurz
angeben.
Ich konnte weder Tyrosin noch Asparagin im urmeristema-
tischen Gewebe der ganz jungen Knospen (Stadium I) finden,
während in deren basalen, von Bündelanlagen durchsetzten Theilen
Tyrosin schon regelmässig vorkommt. Nun die Schösslinge
nehmen sehr langsam an Grösse zu und ihre Stieltheile werden,
wie schon erwähnt, allmählig verholzt. Ich beobachtete, dass das
Tyrosin mit der Zeit im Schösslingskörper erscheint und seine
Menge immer grösser wurde, zugleich auch das Asparagin in
nachstehender Menge. Wenn man einen 4-5 cm langen Schössling
in diesem Stadium (Stadium II) untersucht, so sieht man
folgendes : Der Vegetationspunkt bleibt frei von Amidosubstanzen.
Erst 2-3 mm unten, wo die Bündelanlagen schon differenziert
waren, erscheint die erste Spur von Tyrosin im parenchymatis-
chen Gewebe. Asparagin tritt noch weiter unten ein, wo Zucker
474 K. SHIBATA:
in grösserer Menge vorkommt (etwa in der Mitte von der ganzen
Länge des Schösslings), daneben viel Tyrosin. Zuletzt fand ich
im Stieltheile keine Amide mehr. So coincidiert Asparagin in
seiner Localization fast mit reducierendem Zucker. Pfeffer‘)
bemerkte schon derartiges Zusammentreffen von Traubenzucker
und Asparagin in der ersten Periode der Keimung von Lupinus
luteus. In diesem und auch im folgenden Stadium konnte ich
weder Tyrosin noch Asparagin im Rhizom nachweisen.
Das oben definierte Stadium IIT wird im Laufe des Sommers
erreicht. Während dieser Zeit nimmt die absolute Menge des
Tyrosins sowie des Asparagins immer mehr zu, so dass die
Krystalle des Tyrosins und des Asparagins unter dem Microskop
in grösserer Menge und viel leichter gefunden werden ; die Be-
handlung der Gewebe (die aber eiweissarm sind) mit Millon’s
Reagens bringt überall eine tiefere Färbung als in den vorigen
Stadien. Die Vertheilung des Tyrosins und des Asparagins
stimmt im Wesentlichen mit der des vorigen Stadiums überein.
Dabei ist noch zu bemerken, dass das Tyrosin weniger in Nodien
als in Internodien vorkommt. In diesem Zustande überwin-
tern die Schôsslinge ohne bemerkbare Veränderung bis Ende
Februar. Von jetzt ab erwacht ein regerer Process im Schöss-
linge, und von Anfang— Mitte April wächst es schon zu einer
beträchtlichen Grösse unter der Erde. Die Schösslinge in diesem
Stadium (Stadium IV) werden auf dem Markt als Gemüse feil
geboten. Die oben angegebene analytische Bestimmung Kozai’s
rührt auch von einem solchen Schösslinge her. In diesem Stadium
bemerkte ich folgende Vertheilung: Der Vegetationspunkt ist
frei von Amidokörpern, dagegen reich an Eiweiss, aber in jungen
1) Pfeffer, Über die Proteinkörner und die Bedeutung des Asparagins. Jahrb. f. wis.
Bot. Bd. VIII, p. 539.
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 475
Scheideblättern, die zu dieser Region gehören, lassen sich stets
kleine Mengen Tyrosins nachweisen, daher muss man bei Fest-
stellung der Abwesenheit der Amidosubstanzen in der Spitze sie
thunlichst von Scheideblättern befreien. Das Eiweiss, welches
Biuretreaction giebt, ist in dieser Region besonders reichlich in
Procambialsträngen nachweisbar. Nach Sachs’) wird das Eiweiss
durch diese Gewebe dem Urmeristem zugeführt, und da ich hier in
der Spitze keine Amide auffinden konnte, so kann die Wanderung
des Eiweisses wohl in dem Sachs’schen Sinne geschehen. Das
in dieser Weise von unten zugeführte Eiweiss befindet sich unter-
halb des Urmeristems räumlich getrennt von Stärke in regel-
mässig abwechselnden Zonen von je ca. 0.15 mm Dicke. Diese
Zonen deuten schon zukünftige Internodien und Nodien an, und
befinden sich in den ersteren Eiweiss und in den letzteren Stärke.
Erst 4 mm unter dem Vegetationspunkt tritt die erste Spur von
Tyrosin auf und nach unten nimmt esimmer in den parenchy-
matischen Zellen an Menge zu. Zugleich ist die Abnahme des
Eiweisses in parenchymatischen Zellen leicht constatierbar. Das
Asparagin kommt noch weiter unten (ca. 1-1.5 cm unter dem
Vegetationspunkt) fast gleichzeitig mit reducierendem Zucker zum
Vorschein. Da dicht unter dieser Region die erste Zerreissung
im Markgewebe, die den ersten Anfang der Markhöhle andeutet,
stattfindet, so soll hier die Zellstreckung erst recht ausgiebig
geworden sein. Nach unten nimmt die Menge des Tyrosins und
des Asparagins stetig zu, und dabei übertrifft die Menge des Tyro-
sins bedeutend die des Asparagins. Am reichlichsten findet man
1)Sachs, Über die Leitung der plastischen Stoffe durch verschiedene Gewebeformen.
Flora. 1863.
Es ist bekannt, dass das Eiweiss unter Umständen die Cellulosemembran hindurch
diosmiren kann. Vergl. Puriewitsch, Physiol. Unters. üb. Entleerung der Reserve-
stoffbehälter. Jahrb. f. wiss. Bot. XXXI, p. 68; Pfeffer, Pflanzenphysiologie. Bd. I, p. 613.
476 K. SHIBATA :
Tyrosin an Stellen, wo die Wurzelanlagen zur Bildung kommen,')
so dass Tyrosin beim Schneiden des Gewebes mit dem Messer
sofort im Zelleinneren zu Krystallen erstarrt”) In den Bastzell-
anlagen der Gefässbündel, welche noch keine Wandverdickung
zeigen, kommt das Tyrosin bedeutend reichlicher als im Parenchym
vor. Die Ueberreste des Markparenchyms enthalten nur sehr
wenig Tyrosin. Das Millon’s Reagens bewirkt stark blutrothe
Färbung des Zellsaftes, entsprechend dem hohen Gehalt an
Tyrosin. Jedenfalls hat die absolute Menge des Tyrosins im
Vergleich mit den vorigen Stadien bedeutend zugenommen. In
dieser Region dagegen konnte ich das Asparagin nur mit Sch-
wierigkeit auffinden. Es sei noch hervorzuheben, dass das
Asparagin in der Regel in Knoten und Diaphragmen sich nicht
befindet, dagegen fehlt es hier an Tyrosin nicht.
In den untersten Internodien, wo die Verholzung der Bast-
elemente schon eingetreten ist, verliert sich auch das Tyrosin.
Im Laufe des Aprils durchbrechen die Schösslinge einer nach
dem andern die Erde und wachsen ungemein rasch in die
Länge. Es ist nicht zu bewundern, dass in so schnell wachsenden
Pflanzentheilen ein ausgiebiger Eiweissumsatz vor sich geht.
Tyrosin und Asparagin sind sehr reichlich in den oberen wach-
senden Internodien vorhanden, mit gleicher Vertheilungsweise
wie im vorigen Stadium, d.h. Tyrosin tritt ca. 2 cm unter dem
Vegetationspunkt auf und Asparagin ca. 4 cm unter demselben
gleichzeitig mit reducierendem Zucker. Aber sehr interessant ist
die Vertheilungsweise in halberwachsenen Internodien. Nämlich
in der unteren weichen Wachsthumszone eines solchen Interno-
diums befindet sich das Asparagin ziemlich viel neben reichlichem
1) ca. 10te Internodium von unten.
2) Siehe unten p. 482.
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 477
Tyrosin. Hingegen der obere, schon erwachsene Theil desselben
Internodiums enthält kein Asparagin, aber Tyrosin in einer
nahezu gleich grossen Menge wie im unteren weichen Gewebe.
Hier werden Tyrosinkrystalle in jungen Bastzellen, die eben die
erste Verdickung begonnen haben, reichlich ausgeschieden, aber es
befindet sich viel weniger in Parenchymzellen. Ferner enthalten
die Hadrom- und Leptomelemente niemals Tyrosin. Die Nodien
und Diaphragmen enthalten weniger Tyrosin als in Internodien
und gewöhnlich kein Asparagin. In den eben im Wachstum
vollendeten Internodien kommt Tyrosin noch in fast gleich
grosser Menge vor, aber Asparagin nicht mehr. In weiter unten
liegenden älteren Internodien und Nodien verschwindet allmählig
auch das Tyrosin, und mehrere ganz erwachsene und in Verhol-
zung begriffene Internodien auf der Erdoberfläche sind durch-
gehends von Amidokörpern frei, obwohl sie noch reichlich die
Glykose, wie schon bemerkt, im Parenchym aufspeichern.
Von den Scheideblättern habe ich hier nur zu erwähnen,
dass sich in der basalen, weichen Wachstumszone jedes Blattes
ziemlich viel Asparagin neben reichlichem Tyrosin befindet, und
das erstere verliert sich schon an der Uebergangsstelle zu harten
Theilen, während Tyrosin noch weiter oben im Parenchym der
erwachsenen Spreitentheile reichlich vorkommt. So bemerkt
man hier ein ganz ähnliches Verhältniss wie im Halm.
In einige mm hoher Wurzelanlage, sowohl in Periblem wie
in Plerom, lässt sich fast kein Tyrosin nachweisen, obwohl dicht
darunter liegendes Knotengewebe an demselben reich ist. In
verschieden langen wachsenden Wurzeln ist die Spitze stets
tyrosinfrei, und erst 1.5-2 cm unten ist eine Spur nachweisbar.
Allerdings kommt Tyrosin nur in sehr kleiner Menge im
Wurzelparenchym vor, so dass die microchemische Nachweisung
478 K. SHIBATA :
immer schwierig ausführbar ist. Noch spärlicher komnt
Asparagin in wachsender Region vor. Diese Umstände können
zum Theil dadurch erklärt werden, dass die Wurzeln nur lanr-
sam wachsen und demgemäss hier der ausgiebige Eiweissumsiz
nicht stattfindet.
Die mit dem oben angegebenen ganz übereinstimmende
Vertheilungsweise des Asparagins und des Tyrosins habe ich auch
in Schösslingen folgender Arten constatiert :
Phyllostachys bambusoides,
Phyllostachys puberula,
Bambusa palmata.
In den von mir in dieser Beziehung untersuchten Arundi-
naria-Arten, nämlich:
Arundinaria japonica,
Arundinaria quadrangularis,
Arundinaria Matsumure,
Arundinaria Hindsti,
zeigte Asparagin auch das gleiche Verhalten, während ich
Tyrosin nur schwierig auffinden konnte, in auffallendem Gegen-
satz zu Phyllostachys-Arten. Wie dies zu Stande kommt ist mir
unbekannt.
Ferner ist hier zu bemerken, dass ich in Rhizomen von
Bambusa palmata Asparagin in geringer Menge nachweisen konnte,
während es mir bei Phyllostachys-Arten nicht gelang.
Aus dem oben erörterten Befunde lasse ich folgende vier
Sätze gelten, nämlich :
1. Tyrosin übertrifft Asparagin in Menge.
2. Tyrosin tritt in der Nähe vom Vegetationspunkt auf,
dagegen kommt Asparagin noch weiter unten gleichzeitig
mit Glykose zum Vorschein.- |
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 479
3. Asparagin verschwindet aus Nodien und Internodien
sobald ihre Streckung aufhört, während Tyrosin noch
lange Zeit in denselben erhalten bleibt.
4. Tyrosin verschwindet zuletzt aus ganz erwachsenen und
in Zellwandverdickung begriffenen Internodien und
Nodien.
Nun müssen die einmal vorhandenen und allmählig ver-
schwindenden Amidokörper, wie schon bekannt, zur Eiweissre-
generation, sei es direct oder indirect, verwendet werden, weil
sonst weitere Zersetzungs- oder Oxydationsproducte, wie Am-
moniak, Nitrate u.s.w. in den Schösslingsgeweben angehäuft werden
müssen, was durchaus nicht der Fall ist. Zwar habe ich weder
Ammoniak noch Nitrat mit Nessler’s Reagens resp. Dipheny-
lamin-Schwefelsäure in den betreffenden Geweben nachgewiesen.
Bei der Betheiligung an diesem Eiweissregenerationsprocess
scheint, wie aus den oben erwähnten Thatsachen ersichtlich ist,
Asparagin viel leichter verwendbar zu sein und so kommt es
nur an Stellen, wo regere Eiweissbildungsprocesse stattfinden,
vor. Auch in diesem Sinne lassen sich die folgende Beobach-
tungen erklären. In verkümmerten Schösslingen von Phyllostachys
mitis, die täglich nur einige mm wachsen, während nebenbei
stehende kräftige Exemplare täglichen Zuwachs von mehr als
70 cm zeigten, konnte ich in verschiedenen Internodien Aspara-
gin niemals auffinden, dagegen kam Tyrosin dort reichlich vor.
Ganz ähnlich verhalten sich die Rhizomspitzen von Phyllostachys
mitis und Phyllostachys bambusoides, die im Spätherbst (October)
untersucht wurden, wobei sie äusserst langsam wachsen. In
verschiedenen Internodien derselben konnte ich trotz vielfacher
Bemühungen kein Asparagin mit Sicherheit nachweisen, während
Tyrosin dort reichlich vorkommt. Dass das Vorkommen von
480 K. SHIBATA :
Asparagin stets mit der lebhaften Stoffbildung bei schneller
Streckung verbunden ist, ergiebt sich auch aus folgender Bemer-
kung Pfeffers'): ,, War früher die Wurzel das lebhaftest wach-
sende Organ des Keimpflänzchens, so ist dieses jetzt das Stämm-
chen geworden und dementsprechend wendet sich jetzt der
Hauptstrom von Glykose und Asparagin in dieses.”
Hingegen verhält sich das Tyrosin viel träger in dieser
Beziehung, so dass es in schon erwachsenen Theilen lange Zeit
züruckbleibt. Beim Verschwinden des Tyrosins aus ganz er-
wachsenem Internodium wird ein Theil 2x loco verbraucht, aber
ein anderer Theil wird vielleicht den oberen wachsenden
Internodien zugeführt und dabei müssen die jungen Bastelemente
als Leitungsbahnen benutzt werden, wie besonders reichlicher
Gehalt an Tyrosin es vermuthen lässt,
In jeder Hinsicht sind Asparagin und Tyrosin nicht von
gleichem Werthe. Das erste ist ausgezeichneter Eiweissbaustoff.
während das zweite es nur bis zu einem gewissen Grade ist,
Soweit es leichte Verwendbarkeit des Asparagins anbetrifit, steht
mein Ergebniss mit Hansteen') im Einklang.
Wie entstehen das Asparagin und das Tyrosin ?
Aus der Localisation ergiebt es sich schon, dass das Tyrosin
nur bei der Zersetzung des schon vorhandenen Eiweisses ent-
steht und nicht durch synthetischen Process. In den jungen
Geweben unterhalb des Urmeristems, wo noch kein reducierender
Zucker vorhanden ist, tritt es schon auf und vermehrt sich nach
unten, in dem Masse wie das Eiweiss in den Zellen abnimmt.
Andererseits konnte ich in fungierenden Wurzeln und Rhizomen,
1) Pfeffer, Untersuchungen über die Proteinkörner und die Bedeutung des Asparagins.
Jahrb. f. wiss. Bot. Bd. VIII, p. 548. |
2) Hansteen, Über Eiweisssynthese in grünen Phanerogamen. Jahrb. f. wiss. Bot.
Bd. XXXIII, p. 449, p. 486. |
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 481
die wohl als Bildungsstätte der organischen Stickstoffverbindungen
betrachtet werden dürfen, in allen untersuchten Fällen niemals
Tyrosin nachweisen. Ferner giebt der Blutungssaft, der beim
Stoffabfuhr vom Rhizome eine wichtige Rolle spielt, keine Tyro-
sinreaction. Dafür sprechen noch folgende Versuche, die ich
wiederholt ausgeführt habe. Wenn man irgend eine abgeschnit-
tene Rhizomspitze oder einen Schössling von Phyllostachys milıs,
Phyllostachys bambusoides oder Phyllostachys puber ula in destilliertes
Wasser stellt und am Licht oder im Dunkeln verweilen lässt, so
sieht man nach 2-5 Tagen bedeutende Zunahme von Tyrosin in
beliebigen Internodien. Da in diesem Falle vorher weder Nitrat
noch Ammoniak in den Zellen nachweisbar war, so ist. die
nachträgliche synthetische Bildung von Tyrosin wohl ausgeschlos-.
sen, und die beobachtete Tyrosinzunahme muss allerdings auf
die Eiweisszersetzung zurückgeführt werden.
Mit dem Asparagin ist die Sache schwieriger zu entscheiden.
Obwohl ich in oben erwähnten Versuchen die gleichzeitige
Asparaginbildnng gewöhnlich nicht beobachten konnte, ist es
natürlich nicht ausgeschlossen, dass bei der Eiweisszersetzung
hierbei entstandenes Asparagin schnell zur Eiweissregeneration
verbraucht wurde und demgemäss nicht in gleichem Masse wie
Tyrosin zur Anhäufung kam. Andererseits mag die oben erwähnte
Localisation des Asparaginsin kräftig wachsenden Theilen dadurch
zu Stande gekommen sein, dass das im Rhizome fortwährend
gebildete Asparagin mit dem Blutungssaft den wachsenden
Schösslingen zugeführt und in den betreffenden Theilen ange-
häuft wurde‘). Gegenwärtig haben wir drei Möglichkeiten in
Bezug auf Asparaginbildung: erstens durch directe Eiweiss-
1) Natürlich muss das Material der in wachsenden Schösslingen auf eine so erhebliche
Weise umgesetzten Eiweissstoffe von Rhizomen entstammen.
482 K. SHIBATA :
zersetzung,') zweitens durch Synthese aus Ammoniak’) und
drittens durch Umwandlung von Amidosäuren etc.*) Ob die eine
oder andere von diesen Möglichkeiten in unserem Falle zutrifft
muss vorläufig unentschieden bleiben.
Nun gehe ich zur Besprechung der interessanten Löslichkeits-
verhältnisse des Tyrosins über. Bei der Untersuchung der Schöss-
linge von Phyllostachys mitis im IV und V Stadium habe ich
gefunden, dass alle jungen, noch mit plasmatischem Wandbeleg
versehenen Bastelemente und oft auch parenchymatische Zellen
mit schönen Tyrosin-Nadelbüscheln erfüllt sind (Fig. 59 u. 60).
Dieser Umstand liess mich zuerst vermuthen, dass das Tyrosin
schon in den lebenden Zellen in Krystallform vorkommt. Aber
nach genaueren Untersuchungen lässt es sich bald feststellen, dass
das Tyrosin in den intacten lebenden Zellen ganz gelöst im Zellsaft
vorkommt und nur erst in den beim Schneiden geöffneten Zellen
zu Krystallen erstarrt. Man kann diese Thatsache mit aller
Bestimmtheit in folgender Weise beweisen : ein 3-4-zelllagendicker
Längsschnitt des tyrosinhaltigen Internodialgewebes wird zuerst
durch Herumschwenken in Wasser von den an den Schnittflächen
anhaftenden Tyrosinkrystallen befreit und dann unter dem
Microskop mit feiner Nadel zerzupft, so sieht man bald, dass
in vorher klarem Zellsaft der verletzten Bastelemente und
Parenchymzellen eine Krystallbildung stattfindet, welche nach
wenigen Secunden sich als Tyrosin deutlich erkennen lässt. So
wird hier das äusserst schwerlösliche Tyrosin‘) in so hohem
Masse im Zellsaft in Lösung gehalten, dass es sich schon nach
1) Pfeffer, Pfanzenphysiologie. Bd. I, p. 464.
2)O. Loew, Die chemische Energie der lebenden Zelle. p. 77; p. 78.
3) E. Schulze, Ub. d. Umsatz d. Eiweissstofle in der lebenden Pflanzen. Zeit. f. physiol.
Chemie. Bd. XXIV, p. 63.
4)1 Theil Tyrosin ist löslich in 1900 Theil Wasser bei 16°C.
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 483
blosser mechanischer Verletzung der Protoplasten als Krystalle
abscheiden lässt. Die Tödtung des Gewebes durch Chloroform-
dampf, Osmiumsäuredampf sowie Erhitzung bewirkt ebenfalls die
Abscheidung der Tyrosinkrystalle. Ferner kommt in den durch
3% KNO,-Lösung sehr stark plasmolysierten Bastzellen Tyrosin
nach langer Zeit nicht als Krystalle zum Vorschein‘), aber nach
Verletzung der plasmolysierten Zellen treten die Krystalle bald
hervor. Wie solch eine hohe Löslichheit des Tyrosins zu Stande
kommt ist schwer zu beantworten‘). Allerdings muss hier der
Einfluss von den Saftraum umgebenden, lebenden Protoplasten in
erster Linie in Betracht kommen. Die Ausscheidung von Tyro-
sinkrystallen beim Schneiden des Gewebes lässt sich nicht in
Schösslingen von I, II und III Stadien zeigen. Ebenso verhalten
sich die tyrosinarmen Schösslinge von Arundinaria- und Bambusa-
arten. Die Schösslinge von Phyllostachys puberula und Phyllo-
stachys bambusoides zeigen ganz gleiches Verhalten wie im oben
dargestellten Fall von Phyllostachys mitis.
Ausserdem scheint sich ein Theil des Tyrosins auch in
die Zellwände einzulagern, da die Zellwände der jugendlichen
Bastelemente und später auch des Parenchyms immer stärker
roth durch Millon’s Reagens gefärbt werden, in dem Masse,
dass Tyrosin in den Zellen selbst abnimmt. Bekanntlich haben
früher Correns’) und Fischer‘) die Vermuthung ausgesprochen,
dass die sogenannte Eiweissreaction der Zellwände verschiedener
Pflanzen von in dieselben eingelagertem Tyrosin herrühre. Neuer-
1) Vergl. Pfeffer, Pflanzenphysiologie Bd. I, p. 465.
2) Jedenfalls ist die Ansicht Belzung’s (Recherche chimique sur ]. Germination ete.
p. 219), dass das im Zellsaft gelöste Eiweiss die Krystallisation von Asparagin, Leucin etc
verhindern soll, ganz unzutreffend, denn in unserem Falle wird die Krystallisation schon
durch blosse mechanizche Verletzung des Protoplastes im Zellsaft eingeleitet .
3) Correns, Über die vegetabilische Zellmembran. Jahrb. f. wiss. Bot. Bd. XX VI, p. 616.
4) Fischer, Zur Eiweissreaction der Zellmembran. Ber. d. D. Bot. Gesells. Bd. V, p. 429,
484 K. SHIBATA:
dings vertritt aber Czapek') die Ansicht, dass es sich um die
Reaction eines phenolartigen Körpers handelt, welchen er dem
von ihm in Mooszellwänden aufgefundenen Sphagnol als nahe
verwandt betrachtet.
GERBSTOFFE UND FETTE.
Gerbstoffe und Fette spielen bei Entwicklung der Bambus-
schösslinge nur eine untergeordnete Rolle.
Bei den von mir untersuchten Phyllostachys- und Bambusa-
Arten sind Gerbstoffe in verschiedenen Theilen der Schösslinge
überhaupt nicht in nachweisbarer Menge vorhanden.’) In dieser
Hinsicht bietet Arundinaria quadrangularis ein abweichendes
Verhalten dar. Ein ca. 100 cm langer Schössling zeigte folgende
Vertheilung der Gerbstoffe: Das Urmeristem des Vegetations-
punktes ist frei davon, und dann treten sie plötzlich in reich-
licher Menge im Niveau der 3ten Scheideblattanlage auf, zugleich
mit der ersten nachweisbaren Stärke. Gerbstoffe kommen in
einigen nachfolgenden Internodien in gleich reichlicher Menge
vor und dann nehmen sie nach unten ab. Dabei reagieren am
stärksten die Rindenzellen und die peripherischen Centralcylinder-
parenchymzellen. Selbst in erwachsenen Internodien und Nodien
in der Nähe der Erdoberfläche ist eine schwache Reaction be-
merkbar. Die Wurzelhaube enthält auch kleine Mengen der
eisenbläuenden Gerbstoffe. In der dritten oder vierten Scheide-
blattanlage am Vegetationspunkt treten die Gerbstoffe auf und
nehmen nach unten zu. So sieht man hier eine analoge Verthei-
lungsweise wie in den bisher bekannten Fällen, z. B. bei Vicia,
1) Czapek, Zur Chemie der Zellmembran bei den Laub- und Lebermoosen. Flora. 1899-
Bd. 86, H. 4.
2) Abgesehen von kleinen Mengen in Wurzelhauben, Scheideblättern etc.
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 485
Helianthus'), Zuckerrohr’) u.s.w. Im vorliegenden Falle scheinen
die autochthonen?) Gerbstoffe sich aplastisch zu verhalten, weil
hier Zucker, Stärke u.a. auf ganz gleiche Weise vorkommen wie
bei anderen gerbstofffreien Arten.
Auch kommt den Fetten höchstens nur eine locale Bedeut-
ung zu als Baumaterial der verkorkenden oder verholzenden Zell-
wände, so zum Beispiel verschwinden die kleinen Fetttropfen in
einer subepidermalen Zellschicht der Wurzel schon bei eintreten-
der Verkorkung der Zellwände. In jeder älteren Halmparen-
chymzelle von Arundinaria japonica, A. Hindsu, Bambusa
floribunda etc. befindet sich je ein ölartiger gelber Tropfen, meist
in Verbindung mit Calciumoxalatdrusen. Diese kugeligen
Gebilde unterscheiden sich von echten Fetttropfen dadurch, dass
sie niemals mit Alkannatinktur sich färben. In vielen Punkten
stimmen sie mit dem zuerst von Monteverde‘) in Gramineen
aufgefundenen ,, Harzkorper“ überein.
MINERALSTOFFE.
Ich habe in verschiedenen Jahreszeiten die Vertheilung der
Mineralstoffe in den Reservestoffbehältern und den Schösslingen
verfolgt. Als Untersuchungsmaterial dienten mir hauptsächlich
Phyllostachys mitis und Phyllostachys bambusoides.
Ich konnte in den KRhizomen, die schon beträchtliche
Quantität der Stärke aufgespeichert haben, die Mineralstoffe
leicht auffinden. Sie zeigten folgende Vertheilung sowohl in
Internodien als in Nodien :
1) Vergl. Kutscher, Uber die Verwendung der Gerbsäure im Stoffwechsel der Pflan-
zen. Flora. 1883, p. 33.
2) Went, Chemisch-physiologische Untersuchungen über das Zuckerrohr. Jahrb. f. wiss.
Bot. Bd. XXXI, p. 297.
3) Kraus, Grundlinien zu einer Physiologie des Gerbstoffes. p. 58.
4) Monteverde, Über Ablagerung von Calcium- und Magnesiumoxalat in der Pflanze,
Bot. Centralb. 1890. Bd. XLIII, p. 327.
486 K. SHIBATA:
Phosphor reichlich im Parenchym des Centralcylinders ;
nur wenig in Siebröhren.
Magnesium reichlich vorzugsweise in Siebröhren.
Kalium reichlich im Parenchym.
Calcium nicht nachweisbar in Schnitten ; nur kleine
Mengen in Aschen.
Schwefel nur spurweise in Schnitten.')
Chlor ziemlich reichlich im Parenchym ; aber fast keins in
Siebröhren und anderen Elementen der Gefässbündel.
In den Wurzeln können die obengenannten Stoffe auf über-
einstimmende Weise aufgefunden werden. Die Nitrate’) sind
in Rhizomen nur sehr wenig vorhanden, aber zeitweilig etwas mehr
in der Wurzelrinde. Sie kommen hier also überhaupt nicht zur
nennenswerthen Aufspeicherung. Leitgeb*) hat früher gezeigt,
dass der Phosphor in Dahliaknollen hauptsächlich als Calcium-
phosphat vorkommt. Hier ist aber dies nicht der Fall, da ich
keine lösliche Ca-Verbindung in den Zellen auffinden konnte.
Die Siebröhren der Rhizome und Wurzeln enthalten, wie schon
erwähnt, nur geringe Mengen des Phosphors, dagegen reichlich
Magnesia. Daher scheint Magnesium theils als Phosphat, theils
als lockere organische Verbindung in Siebröhren vorzukom-
men. Schimper‘) und Zacharias?) haben auch im Siebröhr-
ensafte von Cucurbita, Wistaria, Aristolochia und Menispermum
reichliche Mengen des Magnesiums aufgefunden.
1)Schimper (Zur Frage der Assimilation der Mineralsalze. Flora, 1890. p. 223) hat
auch in den meisten Rhizomen keine Sulfatreaction erhalten und es dem Vorhandensein von
Krystallisation verhindernden Substanzen in Zellen zugeschrieben.
2) Vergl. Molisch, Über microchem. Nachweis von Nitraten. Ber. d. D. Bot. G. Bd. I,
p. 154.
3) Leitgeb, Über die durch Alcohol in Dahlia-Knollen hervorgerufenen Krystalle. Bot.
Zeit. 1887. p. 29.
4)Schimper, Zur Frage der Assimilation der Mineralsalze. Flora, 1890, p. 228.
5) Zacharias, Uber d. Inhalt d. Siebröhren von Cucurbita. Bot. Zeit. 1884. p. 71.
WACHSTUMGSESOHICHTE D. BAMBUSGEWAECHSE. 487
Schritt für Schritt mit der Entleerung der Kohlehydrate
verschwinden auch die Mineralstoffe aus dem Rhizomparenchym.
Die Nitrate sind schon im IV Stadium in Rhizomen und
Wurzeln nicht mehr nachweisbar. Ebensowenig konnte ich im
Blutungssaft die Nitratreaction erhalten. Wahrscheinlich werden
die Nitrate hierbei fortwährend zu organischen Verbindungen
verarbeitet.')
An den unteren, mit zahlreichen jungen Wurzeln besetzten
Theilen der Schösslinge besitzen Phosphor und Kalium eine
andere Vertheilungsweise als im Rhizome; sie kommen nun
reichlicher in den Bündeln vor. Das Magnesium befindet
sich hier auch in Siebröhren bevorzugt. Ich konnte niemals
die Nitrate im Schösslingskörper auffinden.’) Da sie auch im
Rhizome fehlen, so ist es höchst wahrscheinlich, dass überhaupt
nur wenig Nitrat als solches in den Schössling eingeführt wird.
Von dieser Region nimmt die direct nachweisbare Menge
von Phosphor, Magnesium, Kalium und Schwefel oben nach dem
Vegetationspunkte hin immer mehr zu, wie man sich durch die
Musterung successiver Querschnitte überzeugen kann. Phosphor
kommt anscheinend am reichlichsten in 2-3 cm Entfernung vom
Vegetationspunkt vor, und von hier nach oben nimmt die direct
nachweisbare Menge desselben wieder ab. Merkwürdigerweise
kommt Phosphor fast ausschliesslich in den eiweissreichen Pro-
cambialsträngen vor.) Im Urmeristem lassen sich die anorgani-
1) Die feinen Nebenwurzeln geben stets mehr oder minder starke Nitratreaction. Ob sich
in irgend einer Weise der hier befindliche Pilzsymbiont an der Stickstoffussimilation
betheiligt, muss zur Zeit dahingestellt bleiben. |
2) Übrigens ist es klar, dass die Nitrate sich nicht Lei so lebhafter Eiweisszersetzung
bilden. (Vergl. Schulze, Über d. Vorkommen von Nitraten in Keimpflanzen. Zeit. f.
physiol. Chemie. Bd, XXII, p. 83).
3) Ich habe auch die Lilienfeld’sche Methode fiir Erkennung der Localisation des
Phosphors mit Erfolg benutzt. (vergl. Strasburger, Das botanische Practicum. III. Aufl.
p. 144).
488 'K. SHIBATA:
schen Phosphorverbindungen nicht mehr nachweisen, sondern
grosse Mengen organischer Verbindungen‘. Magnesium tritt
ebenfalls in der Nähe der Spitze fast ausschliesslich in den
Bündelanlagen auf, aber kleine Mengen sind auch in den
eiweisshaltigen Internodialzonen vorhanden. Beachtet man die
bevorzugten Vorkommnisse des Magnesiums in Siebröhren,
Bündelanlagen, Internodialzonen und auch im Urmeristem, so
darf man wohl annehmen, dass es irgend eine wichtige Rolle
bei Eiweissumsatz oder Eiweisswanderung spielt.”) Kalium lässt
sich in der Asche des Vegetationspunktes reichlich nachweisen.
Hingegen ist die Schwefelsäure nicht direct im Vegetations-
punkt nachweisbar, obgleich sich schon ca. 2 cm weiter unten
eine ziemlich starke Reaction zeigt. Das in minimaler Menge
zugeführte Calcium wird in einiger Entfernung vom Vegetations-
punkte als Kalkoxalat niedergeschlagen. Chlor lässt sich ziemlich
viel in eiweissreichen Internodialzonen nachweisen. Im Urme-
ristem scheint es jedoch gänzlich zu fehlen.
Wenn die Schösslinge über die Erde emporwachsen und die
Internodien nach einander ihre definitive Länge erreichen, so
nimmt die nachweisbare Menge der Mineralstoffe in denselben
stetig ab. In unterirdischen Internodien tritt nun mehr oder
minder starke Nitratreaction ein, da die erwachsenen Wurzeln an
dieser Region schon ihre Thätigkeit entfalteten und begannen
die Bodensalze aufzunehmen.
In der ersten Entwicklungsphase der Wurzel ist eine starke
Ansammlung der Mineralstoffe im meristematischen Gewebe leicht
constatierbar. In etwas länger erstreckten Wurzeln findet eine
ähnliche Ansammlung in der Spitze statt. Phosphor und Mag-
1) Vergl. Schimper, ke. p. 224.
2) Vergl. Hornberger, Chemische Untersuchungen über das Wachstum der Maispflanze.
Landw. Jahrb. 1882. p. 278.
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 489
nesium kommen, wie in Schösslingen, hauptsächlich in jungen
procambialen Elementen vor, die vielleicht ihre Wanderbahn
herstellen. Im Urmeristem fehlen nachweisbare Mengen von
Schwefel und Chlor. Die mehr als 40 cm lang gewachsenen
Wurzeln zeigen ziemlich starke Nitratreaction an der Rinde,
welche von von aussen aufgenommenen Salzen herrührt.
In den jüngsten Scheideblattanlagen in der Umgebung des
Vegetationspunktes lässt sich sehr früh eine Ansammlung von
Phosphor und Magnesium beobachten. Uebrigens bedarf dies
keiner Besprechung mehr.
VII. Ueber die Entleerung der Reservestoffe.
Die Versuche von Hansteen') und Puriewitsch’) haben
die Thatsache festgestellt, dass bei Endospermen, Samenlappen,
Knollen und Rhizomen die Entleerung der deponierten Reserve-
stoffe mehr oder minder selbstthätig stattfinden kann. Nun schien
es mir geboten zu bestimmen, erstens inwieweit die Entleerung des
Rhizoms unabhängig von wachsenden Schösslingen vor sich gehen
kann, und zweitens in welchem Grade die Entwicklung der
Schösslinge durch die totale oder partielle Separierung vom
Rhizomsystem beeinflusst wird. Zu diesem Zweck habe ich
Mitte April eine Anzahl kräftig wachsender, unterirdischer
Schösslinge aufgesucht und verschieden tiefe Einschnitte in ihre
Stieltheile und benachbarte Rhizominternodien gemacht. Die
betreffenden Rhizomtheile waren in diesem Stadium mit Stärke
strotzend erfüllt, wie ich durch die Musterung zahlreicher
Exemplare überzeugt war; es zeigte sich folgendes:
1) Hansteen, Über die Ursache der Entleerung der Reservestoffe aus Samen. Flora.
1894 Bd. 79, p. 419.
2) Puriewitsch, Physiologische Untersuchungen über die Entleerung der Reserve-
stoffbehälter, Jahrb. f. wiss. Bot. Bd. XXXI, p. 1.
Centralcylinderparen-| strotzend erfüllt-
chym recht viel (54) engl
Markparenchym strotzend erfiillt (5)
In der folgenden Tabelle stelle ich die Ergebnisse einiger
Versuche zusammen :
490 K. SHIBATA :
Starke | Glykose | Rohrzucker
Rindenparenchym |strotzend erfüllt JM sehr wenig | ziemlich viel
Opera- Inhalt der Rhizome | Bemer-
tionen | Stärke | Zucker | kungen
Durchschneiden Rinde nd
6sten Inte nh u a, 2
dia rg Centraleylinder- ae a
3ten Int. | parenchym: reer
im Sten Int. /f und _ Mark-
. eine Starke
untersucht: 30. en (0); Mark- | Parenchym: fast
Durchschneiden parenchym : ein Zucker
im Stiel wenig Stirke (2)
operiert :
Rinden-, Cen-
: tralcylinder-
wie oben wie oben und Mark-
untersucht: 30. Mai parenchym:
wenig Zucker
operiert :
Durch en
operiert: A il |im Sten Int. vor : E keine Glykose;
dem Schösslinge; due Hu kein Rohr- |
untersucht: 18. Mai |Durchschneiden
im Stiel
Durchschneiden
au on ne vor
operiert : ; il | dem Schésslinge ; , a keine Glykose;
Br 1.5cm tiefer Ein-| Keine Stärke | ehr chi nig
untersucht: 18. i |schnitt ins 2ten (0) Rohrzucker
Int. hinter dem
Schösslinge
operlert: Durchschneiden fast keine
té cie im Stiel Stärke (0-1)
operiert : ; il | Durchachneiden
im 3ten Int. vor
untersucht: 19. Mai | dem Schösslinge
wenig Stärke fast keine
2 Glykose
ae . | Durchschneiden _
VII BEE j im 2ten Int.| keine Stärke
. : hinter dem (0)
untersucht: Schösslinge
wie oben
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 491
Alle oben angeführten Versuche ergaben übereinstimmend,
dass die Entleerung, zumal die Stärkeauflösung in bestimmten
Rhizompartien unabhängig von Schösslingen vor sich gehen kann.
Puriewitsch hat bei den Versuchen mit den Rhizomen von
Curcuma und Rudbeckia gezeigt, dass in solchen Rhizomen eine
partielle Entleerung selbstthätig stattfand, wenn die continuierliche
Ableitung der Lösungsproducte mittelst Gyps besorgt wurde’).
Nun in meinen Versuchen war die Bedingung für derartige Stärke-
entleerung besonders günstig. Die zahlreichen kräftigen Wurzeln
an Rhizomknoten erzeugten zur Zeit einen ansehnlichen Blutungs-
druck und der zuckerhaltige Blutungssaft wurde immer fort von
den Schnittflichen der Rhizome ausgeschieden. Damit wurde
eine fortwährende Wegführnng der Lösungsproducte erzielt,
welche eine so volkommene Entleerung herbeiführte.
Ferner ist es aus obigen Versuchen ersichtlich, dass die
Entwickelung der Schösslinge durch jeden operativen Eingriff in
benachbarte Rhizominternodien—d.h. durch jede Herabsetzung
des Blutungsdrucks—bald sistiert wird.
Macht man in der Nacht oder frühmorgens ein Bohrloch in
ein beliebiges Internodium des kräftig wachsenden Schösslings, so
quillt bald ein zuckerhaltiger klarer Saft hervor. Am 19. Mai
wurde der Blutungssaft von einem mittleren Internodium eines
ca. 1.5 Meter hohen Schôsslings von Phyllostachys puberula
gesammelt. In 100 cem dieser Flüssigkeit fand ich 0.289 gr
Glykose. Ausserdem enthält der Blutungssaft eine kleine Menge
der Amide, da er nach Beseitigung des Eiweisses’) eine starke
Trübung beim Zusatz von Quecksilberoxydnitrat giebt.
1) Puriewitsch, Le. p. 28.
2) Nach der Stützer’schen Methode.
492 K. SHIBATA :
Schröter!) schrieb: ,, Später, in den ausgewachsenen Glie-
dern, findet sich oft ein klares Wasser, das in manchen trockenen
Gegenden den Reisenden ein höchst willkommener Fund ist.“
Derartige mit Wasser erfüllte Markhöhlen habe ich manchmal bei
jungen Halmen von Phyllostachys mitis gefunden. Das Wasser
ist nichts Anderes als der Blutungssaft, der sich von radialen
Rissen an der Peripherie des Diaphragms ausgeschieden hat. In
einem Falle betrug der Glykosegehalt der Flüssigkeit, die sich in der
unteren Internodialhöhle von Phyllostachys mitis befand, 0.269%.
Wenn man frühmorgens einen Bambusbusch besucht, so wird
man einen förmlichen Regen von Wassertropfen aus den Scheide-
blattspitzen der wachsenden Schösslinge bekommen.) Dies in
bekannter Weise von Blattspitzen ausgeschiedene Wasser enthält
auch Glykose neben einer Spur von Amiden. Eine Zuckerbes-
timmung der am 28. April gesammelten Flüssigkeit ergab
0.0958% Glykose.
Alle diese Thatsachen weisen darauf hin, dass eine erheb-
liche Menge der Kohlehydrate und vielleicht auch Amide mit
dem Blutungssaft den Schösslingen zugeführt werden. Dadurch
werden die Schösslinge mit den Baustoffen genügend rasch ver-
sorgt.) Die oben erörterten Bauverhältnisse der Stieltheile lassen
sich auch nicht anders denken, als dass hier der ausgiebige
Stofftransport nur durch die Bündel und zwar durch die wohl
ausgebildeten Gefässe geschehen kann.
1)Schröter, Der Bambus und seine Bedeutung als Nutzpflanze. p. 14; Cohn, Über
Tabaschir. p. 375.
2)Molisch, Uber das Bluten tropischer Holzgewiichse. Ann. d. Jard. Bot. Buit.
1898. Suppl. II, p. 23.
3) Vergleiche hierzu: Strasburger, Bau und Verrichtungen der Leitungsbahnen.
p. 877; Fischer, Beiträge zur Physiologie der Holzgewächse. Jahrb. f. wiss Bot, Bd. XXII,
p. 75; p. 150.
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 493
VII. Zusammenfassung.
In Obigem habe ich die wesentlichen Züge der Wachs-
tumsgeschichte der Bambusgewächse darzustellen versucht. Die
Hauptresultate werden hier kurz in folgenden Worten zusam-
mengefasst :
1. Die Stärke wird in parenchymatischen Zellen der
Rhizome, Halme und Wurzeln als Hauptreservestoff abgelagert.
Die Verminderung derselben im Winter wurde nicht beobachtet,
während zur Zeit des raschen Austreibens von Schösslingen eine
unverkennbare Stärkezunahme (transitorisch) in benachbarten
Rhizomtheilen constatiert wurde. |
2. Die Glykose dient als Baumaterial in wachsenden Theilen
der Schösslinge und ist in schon fertig gestreckten Internodien
derselben transitorisch reichlich aufgespeichert.
3. Der Rohrzucker tritt als das Lösungsproduct der
Stärke im Parenchym der Rhizome und Halme auf.
4. In schnell wachsenden Schösslingen fand eine ausgiebige
Eiweisszersetzung statt, dabei trat Tyrosin in bedeutender Menge
auf.
Tyrosin und Asparagin zeigen einen weitgehenden
Unterschied in ihrem Verhalten. Tyrosin wird schwerer und
langsamer für Eiweissregeneration verbraucht, so dass es in schon
erwachsenen Theilen eine Zeit lang zurückbleibt. Hingegen ist
Asparagin leicht und rasch dazu verwendet und kommt nur
an Stellen vor, wo eine lebhafte Stoffbildung stattfindet.
5. Gerbstoffe kommen nur in Schösslingen einzelner Arten
vor, und Fette spielen hierbei keine wichtige Rolle sowohl als
Wanderstoffe wie als Reservestoffe.
6. Phosphor, Kalium, Magnesium und Chlor werden
494 K. SHIBATA :
in den Reservestoffbehältern aufgespeichert, dabei kommt Mag-
nesium vorwiegend in Siebröhren vor. Calcium und Schwefel
sind gewöhnlich nicht direct nachweisbar.
7. Die Mineralstoffe wandern bei rascher Entwicklung der
Schösslinge schnell von den Rhizomen aus und werden in den
wachsenden Theilen angesammelt. In der Spitze der Halme,
Rhizome und Wurzeln befinden sich Phosphor und Magnesium
in direct nachweisbarer Form fast ausschliesslich in Procambial-
strangen. Schwefel wird erst im wachsenden Theile der
Schösslinge deutlich nachweisbar.
8. Die vom Boden aufgenommenen Nitrate werden wahr-
scheinlich schon in den Wurzeln und Rhizomen zu organischen
Verbindungen verarbeitet.
9. Die Auflösung der Stärke und die Entleerung der Lö-
sungsproducte aus den Rhizomen können unabhängig von der
Entwickelung der Schösslinge fortgehen.
10. Der ausgiebige und schnelle Stofftransport nach wachsen-
den Schösslingen von den Rhizomen kann in Wasserbahnen
geschehen. Dafür sprechen vor allem die Blutungserscheinungen
der Rhizome und Schösslinge und die Bauverhältnisse der Schöss-
lingsstiele.
Botanisches Institut
Juni 1890. Kaiserl. Universität
zu Tokio.
WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 495
VERZEICHNISS DER UNTERSUCHTEN ÄRTEN.')
Phyllostachys mitis Rivière. (Nom. Jap. Moso-chiku.)
Phyllostachys bambusoides Sieb. et Zucc. (Nom. Jap. Ma-dake.)
Phyllostachys bambusoides Sieb. et Zucc. var. aurea Makino. (Nom. Jap.
Hotei-chiku.)
Phyllostachys puberula Munro. (Nom. Jap. Ha-chiku.)
Phyllostachys puberula Munro. var. nigra. (Nom. Jap. Kuro-chiku.)
Phyllostachys Kumasasa Munro. (Nom. Jap. Okame-sasa.)
Arundinaria japonica Sieb. et Zucc. (Nom. Jap. Ya-dake.)
Arundinaria Simoni Rivière. (Nom. Jap. Me-dake.)
Arundinaria Matsumure Hackel. (Nom. Jap. Kan-chiku.)
Arundinaria quadrangularts Makino. (Nom. Jap. Shikaku-dake.)
Arundinaria Hindsii Munro. (Nom. Jap. Kansan-chiku.)
Arundinaria Hindsit Munro, var. graminea Bean. (Nom. Jap. Taimin-
chiku.)
Arundinaria Fortune: Rivière. (Nom. Jap. Chigo-sasa.)
Arundinaria variabilis Makino. (Nom. Jap. Ne-sasa.)
Arundinaria pygmea Mitf. (Nom. Jap. Oroshima-chiku.)
Arundinaria Narihira Makino. (Nom. Jap. Narihira-dake.)
Arundinaria Tootsik Makino. (Nom. Jap. To-chiku.)
Bambusa borealis* Hackel. (Nom. Jap. Suzu-dake.)
Bambusa palmata* Marliac. (Nom. Jap. Chimaki-sasa.)
Bambusa Veitchit* Carrière. (Nom. Jap. Kuma-sasa.)
Bambusa paniculata* Makino. (Nom. Jap. Nemagari-dake.)
Bambusa nipponica* Makino. (Nom. Jap. Miyako-sasa.)
Bambusa ramosa* Makino. (Nom. Jap. Azuma-sasa.)
1) Die ausführliche Beschreibung der hier angeführten Arten findet man bei Makino,
Bambusaceæ Japonice (The Botanical Magazine, Vol. XIV, Nr. 156, p. 20 ff.), die beige-
fügten japanischen Namen sollen zum Herausfinden der betreffenden Arten in der genannten
Schrift dienen.
Die mit * bezeichneten Arten gehören meiner Ansicht nach nicht eigentlich zu Bambusa,
sondern sie würden vielleicht eine selbständige Gattung bilden. (Vergl. oben p. 446 und auch
Makino, lc. p. 20).
An dieser Stelle spreche ich Herrn Makino für die von ihm gütigst vorgenommene Bestim-
mung einiger Arten und auch Herren Asö und Inami für ihre freundliche Unterstützung
bei einigen analytischen Arbeiten meinen besten Dank aus.
496
K. SHIRATA :
Bumbusa vulgaris Wendl. (Nom. Jap. Daisan-chiku.)
Bambusa nana Roxb. (Nom. Jap. Howo-chiku.)
Bambusa nana var. normalis Makino. (Nom. Jap. Taiho-chiku.)
Bambusa stenostachya Hackel. (Nom. Jap. Shi-chiku.)
Dendrocalamvs latiflorus, Munro. (Nom. Jap. Ma-chiku.)
VII.
INHALT.
Einleitung; ss see see. avs eae ee ea 44 497
Untersuchungsmaterial und Methodisches, ... ... 429
Die Bauverhältnisse. ... ... … see eee nen 0. 433
Der Entwicklungsvorgang der Schösslinge. ... ... 453
Verhalten der Baustoffe während der Entwicklung
der Schôsslinge. ... ... ser … …. nen ne nen 408
Ueber Entleerung der Reservestoffe. ... ... ... 489
Zusammenfassung. ce ee … … tee … .… 493
Erklärung der Tafeln.
Tafel XXIL
Fig. 1. Zwei neben einander stehende Siebrörenglieder mit zahlreichen
Siebtüpfeln (stp) an den Seitenwänden, aus Rhizomknoten von
Phyllostachys mitis. spl Siebplatte, stp Siebtüpfel. Vergr. 360.
Fig. 2. Querschnitt durch das Rhizom von Bambusa nipponica. R Rinde,
Brg subcorticaler Bastring, cent Centralcylinderparenchym. Vergr. 30.
Fig. 3. Querschnitt durch das Rhizom von Arundinaria japonica. B
Bastbinder. Vergr. 30.
Fig. 4. Die spindelförmige Anschwellung des Leptoms eines ende
bei der Ansatzstelle an der Rhizombündel. S Siebröhren, gi Geleit-
zellen, G Gefässe, P Parenchymzellen, chf cambiformartige Elemente
Vergr. 70.
Fig. 5. Querschnitt durch die Anschwellung. B Bastzellen, P u. cdf wie
in Fig. 4. Vergr. 125.
Fig. 6. Theil der langgestreckten cambiformartigen Elemente aus dem
mittleren Theile der Anschwellung, mit Querstreifen auf den Seiten-
wänden. Vergr. 450.
Fig. 7. Derselbe im Querschnitt. Vergr. 450.
Fig. 8. Die Leptomanschwellung in einem früheren Entwicklungsstadium.
Längsschnitt durch den Knoten. $ Siebröhren, B Bastzellen, chf
cambiformartige Elemente. Vergr. 83.
9, Übergangsstelle der cambiformartigen Elemente zum normal gebauten
Leptom, in einem jugendlichen Zustand. Sämmtliche Elemente mit
auffallend grossen Zellkernen und reichlichem Plasmagehalt. Su. chf
wie in Fig. 8. Vergr. 450.
Fig. 10. Dergleichen im fertigen Zustand. tp Tüpfel, cb/ wie oben. Vergr.
450. Figuren 4-10 beziehen sich auf Phyllostachys mitts.
Fig. 11. Ein Gefässbündel aus einem inneren Teil des Rhizoms von Arundi-
naria Hindsii. S Siebröhren, yZ Geleitzellen, G Gefässe, d Durch-
lassstelle. Vergr. 125.
Fig. 12. Eine subepidermale sclerotische Parenchymschicht des Rhizoms
von Bambusa palmata. Längsschnitt. ep Epidermis, sc? sclerotische
Parenchymzellen, 2 Rindenzellen. Vergr. 360.
. 13. Querschnitt durch den Stieltheil. 2 Rinde, B Bastbänder, bs
Bastscheide des Mestombündels, mes Mestom. Vergr. 17.
. 14. Ein Mestombündel im Stieltheile, mit vollkommen umschliessender
Bastscheide. ? Tracheiden, P, bs, S, gl, u. @ wie oben, Vergr. 195.
Fig. 13-14. Phyllostachys mitis.
ig. 15. Querschnitt durch den dünnen Halmzweig von Arundinaria
pygmæa. ep u. & wie oben. Vergr. 83.
. 16. Theil desgleichen von Arundinaria japonica. ep, Bu R wie
oben. Vergr. 360.
ig. 17. Ein Halm-Bündel von Bambusa nana var. normalis, mit Paren-
chymlamelle im innenseitigen Bastbeleg. par. 7 Parenchynlamelk,
S, Gu. B wie oben. Vergr. 200.
. 18. Einige verschiedenartige Vorkommnisse des Parenchymgewebes
im Bastbelege. Arundinaria Hindsii. B Bastbelege, par paren-
chymatisches Gewebe. Vergr. 70.
. 19. Ein Halmbündel von Bambusa nana. Die durch parenchymatische
Zellen vom Mestom abgetrennte Masse des Bastbelegs bleibt unverdickt.
par. l, B wie oben. Vergr. 200.
. 20. Auftreten der Stärkekörner in neu differenzierter Parenchymlamelle.
Vergr. 70.
. 21. Obiges im Längsschnitt. Vergr. 125.
. 22. Die durch successive Quertheilungen von Procambialzellen ent-
standenen Parenchymzellen. pre Procambialzellen, & Kern, st Starke-
körner. Vergr. 360. Figuren 20-22. Arundinaria Hindsii.
Fig.
Fig.
Tafel XXIII.
. 23. Querschnitt durch die junge Wurzel von Bambusa palmata. ı.k
innere Rindenzellen, end Endodermis mit Caspary’schen Streifen,
per Pericambium, p.had peripherische Hadromstränge (primordiale
Netztracheiden), p.lep peripherische Leptomstränge. Vergr. 360.
. 24a. Peripherischer Theil der Wurzelrinde von Phyllostachys mitis.
ep Rest der Epidermis, hyp stark verdickte (an den Aussenwänden)
subepidermale Zellen, scl peripherische sclerotische Elemente, a.R
äussere Rindenzellen. Vergr. 200.
. 246. Desgleichen im jugendlichen Zustand. ep, hyp u. scl wie oben.
. 25. Starkverdickte Subepidermalzellen (Aussenscheide) von Phyllo-
stachys bambusoides var. aurea. Vergr. 360.
. 26. Peripherischer Theil der Wurzelrinde von Bambusa vulgaris. ep
Epidermis, hyp unverdickt gebliebene Subepidermalzellen, 802 peripheris-
che Sclerenchymzellen, a. äussere Rindenzellen. Vergr. 360.
. 27. Wurzelrinde von Bambusa nana. ep, hyp, scl, a.R u.t.R wie oben,
Vergr. 125.
. 28. Endodermis (end) und starkverdickte Pericambiumzellen (per)
von Phyllostachys mitis. Verg. 360.
. 29. Längsschnitt durch die Endodermis von Phyllostachys mitis. Vergr.
360.
. 30. Längsschnitt durch den peripherischen Theil der Wurzelrinde von
Arundinaria Matsumure. hyp u. scl wie in Fig. 24. Vergr. 360.
. 31. Endodermis und angrenzende Rindenzellen von Bambusa steno-
stachya. Längsschnitt. cel Zellstoffauswüchse, end u. per wie oben.
Vergr. 360.
. 32. Dieselben im Querschnitt. end, per u. cel wie oben. Vergr. 360.
. 33. C-förmig verdickte Endodermiszellen von Bambusa palmata.
Vergr. 360.
34. Theil des Wurzelquerschnittes von Phyllostachys Kumasasa. ver
Verstärkungsring, {f Lufträume, end, i.R, per u. p.lep wie oben.
Vergr. 360.
35. Verknüpfung der Leptomstränge durch dünnwandiges Verbindungs-
gewebe. Phyllostachys bambusoides. ver.p. Verbindungsgewebe,
nb.w Nebenwurzel, p.lep, i.lep, G, end wie oben. (schematisiert)
Vergr. 70.
. 36. Dergleichen bei Bambusa vulgaris. Vergr. 45.
. 37. Querschnitt durch die Hauptwurzel an der Ansatzstelle der Neben-
wurzel. Arundinaria Matsumure. n.lep Leptomstrang der Neben-
wurzel, {ep Leptomstränge der Hauptwurzel, end, t.R, per wie oben.
Vergr. 360.
g. 38. Innerer Leptomstrang von Bambusa vulgaris. S Siebrôhre, ch
Cambiformzellen, mz mechanische Zellen. Vergr. 360.
. 39. Verschmelzung des inneren Leptomstrangs mit dem peripherischen.
i.lep innerer Leptomstrang, p.dep peripherischer Leptomstrang, $,
cb wie oben. Vergr. 360.
. 40. Verschmelzung zweier peripherischen Leptomstränge. Vergr. 360.
. 41. Directer Anschluss des Leptomstrangs an Hadromparenchym. ¢
Gefäss, hp Hadromparenchym (Gefässbelegzellen), Zep u. mz wie oben.
Vergr. 360.
. 42. Zusammentreffen zweier Hadromstränge. G, hp u. mz wie oben.
Vergr. 360. |
Figuren 39-42. Phyllostachys bambusoides.
ig. 43. Verbindungsgewebe zwischen inneren und peripherischen Leptom-
strängen. i.lep, p.lep, mz u S wie oben. Vergr. 360.
g. 44, Dasselbe von Bambusa vulgaris im Längsschnitt. $, mz, verp
wie oben. Vergr. 360.
. 45. Querschnitt durch den Basaltheil der Nebenwurzel von Phyllostachys
mitis. lep Leptomstränge, mz mechanische Zellen. Vergr. 360.
. 46. Theil der Rinde der Nebenwurzel von Phyllostachys puberwa, wit
endophytischen Mycelfäden. 2 Rindenzellen, myc Pilzfäden, ves
Vesiculen, kor gelbe körnige Substanz. Vergr. 360.
. 47. Querschnitt durch die Nebenwurzel von Phyllostachys puberula.
hyp Subepidermalzellen, sc? peripherische sclerotische Zellen, 2, myc,
end, lep wie oben. Vergr. 200.
Fig,
Fig.
Fig.
Tafel XXIV.
ig. 48. Ein Scheideblattbündel mit starkem Bastbeleg (B) von Phyllo-
stachys mitis. Vergr. 83.
. 49. Querschnitt durch das Scheideblatt von Arundinaria Malsumure
Vergr. 360.
g. 50. Kleinere Scheideblattbündel von Bambusa stenostachya ; Leptom
(lep) ist stets von einschichtigen verholzten Elementen (h) umgeben.
Vergr. 360.
51. Querschnitt durch das Scheideblatt von Bambusa stenostachya. B
Bastbelege, subep.b subepidermale Bastplatte, P Parenchym, mes
Mestom. Vergr. 83.
. 52. Desgleichen von Phyllostachys mitis. lf Lufträume, B, subep. 6,
P, mes wie oben. Vergr. ca. 10.
. 53. Queranastomose des Scheideblattbündels von Arundinaria Hindsii.
Vergr. 360.
. 54. Ligninauswüchse an Zellwänden. („Zwickel“). Vergr. 360.
. 55. Ein kleines Bündel im Laubblatt von Arundinaria Hindsit. ps
Parenchymscheide, bs Bastscheide, subep.b, Had, Lep wie oben.
Vergr. 360.
ig. 56. Längsschnitt durch einen kleinen Nerv des Laubblattes von Bam-
busa palmata. ep, subep. b, ps, bs wie oben. Bastscheideelemente
sind reichlicher betüpfelt als subepidermale Bastelemente. Vergr. 360.
. 57a. Stärkekôrner aus Rhizom von Phyllostachys Kumasasa. Vergr
830.
. 57b. Polyadelphische Stärkekörner aus dem Halm von Bambusa
palmata. Vergr. 830.
. 58. Einige Tyrosinkrystalle, die ausserhalb des nach Borodin’scher
Methode behandelten Schnittes entstanden sind. Vergr. 360.
. 59. Beim Schneiden sofort in jungen Bastzellen auskrystallisierende
Tyrosinkrystalle. B junge Bastzellen (mit plasmatischem Wand-
belege), P Parenchymzellen, tyr Tyrosinkrystalle. Vergr. 360.
60. Desgleichen in Längsschnittansicht. Vergr. 200.
61. Beim Einlegen vom Schnitt in Glycerin in das Zelllumen ausges-
chiedene Tyrosinkrystalle. tyr Tyrosinkrystalle, st Stärkekörner, P.
Parenchymzellen. Vergr. 360.
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Decomposition of Hydroxyamidosulphates by
Copper Sulphate.
By
Edward Divers, M. D. D. Sc., F. R. S., Emeritus Prof.,
and
Tamemasa Haga, D. &., F. C. S.,
Professor, Tokyo Imperial University.
When copper sulphate is added to a solution of a hydroxy-
amidosulphate and the mixture heated, the acid of the salt is
quickly decomposed into water, sulphur dioxide, sulphuric acid,
amidosulphuric acid and nitrous oxide, with possibly a little.
nitrogen. By itself, a heated solution of an alkali hydroxy-
amidosulphate is in a state of very unstable equilibrium, generally
hydrolysing into a solution of hydroxylamine acid sulphate,
and always doing so in presence of a trace of acid, whilst in
presence of even a trace of alkali it slowly passes into sul-
phite and hyponitrite (this Journ, 3, 219). In the cold with
alkali and copper salt, the hydroxyamidosulphate becomes
oxidised at once to sulphite, sulphate, nitrous oxide, and water
with reduction of the cupric hydroxide (op. cit., 225), and when
heated with cupric chloride it reduces the latter to cuprous
498 DIVERS AND HAGA : DECOMPOSITION OF
chloride, becoming itself converted into sulphur dioxide, sulphate,
nitrous oxide, and water. Mercuric nitrate oxidises hydroxy-
amidosulphate more completely, but ferric chloride seems to act
like copper sulphate, and liberates sulphur dioxide.
An alkali hydroximidosulphate is also decomposed by copper
sulphate, but not so easily, for it can be heated with it at 100°
for a short time without change, and only decomposes (but
‘then suddenly) some degrees above that temperature, yielding
the products which a hydroxyamidosulphate gives, together with
sulphuric acid from its hydrolysis into that salt.
Although the presence of much sulphuric acid prevents the
action of copper sulphate on a hydroxyamidosulphate, the acid
in moderate excess has but little effect.
Sodium hydroximidosulphate, if kept with care, decomposes
only very slowly in a way which has hitherto been obscure
(this Journ, 7, 45), but if considered in connection with the
action of copper sulphate it may be regarded as essentially the
same as that brought about by heating it in solution with that
salt. For, the decomposed hydroximidosulphate contains, besides
acid sulphate and hydroxyamidosulphate, both a little gas
(nitrous oxide or nitrogen) shut up in its pores which escapes
when the mags is dissolved in water, and also a little amido-
sulphate, which can be separated from the other salts by pre-
cipitation with mercuric nitrate (this Journ., 9, 242, also 229,
230).
The decomposition of hydroxyamidosulphates by copper
sulphate is also in evident relation with the gradual decomposi-
tion of impure hydroxylamine hydrochloride, particularly when
ferric chloride is among the impurities, water, nitrous oxide and
HYDROXYAMIDOSULPHATES BY COPPER SULPHATE. 499
ammonia (in place of amidosulphuric acid) being the principal,
if not the sole, products. |
There is a very marked difference in the proportions of the
products of decomposition between a hydroxyamidosulphate and
a hydroximidosulphate, but this seems to be owing merely to
the fact that the temperature of the decomposition is different,
for according as hydroxyamidosulphate is heated slowly or
rapidly the proportions of the products of decomposition deviate
from or approach those which obtain when a hydroximidosulphate
is decomposed, this only taking place at a temperature above 100°.
As little as one-tenth of an equivalent of copper sulphate
hus been found to suffice for the complete decomposition of an
alkali hydroxyamidosulphate, the copper sulphate not being
“ consumed in the change it effects ; this allows of the decomposition
being to a great extent carried out at the boiling temperature,
when again the result approaches that observed where hydrox-
imidosulphate is the sali decomposed. Even much less than the
amount above named will effect an almost complete decomposition
but that the quantity of the catalytic agent cannot be very
greatly reduced seems to be due in part to the simple hydrolysis
of some of the hydroxyamidosulphuric acid set free by the
copper sulphate during the prolonged heating here necessary.
Since the cupric salt suffers no reduction, it will be seen
that one part of the hydroxyamidosulphate becomes reduced to
amidosulphate by yielding oxygen for the oxidation of the other
part to water, sulphate and nitrous oxide. The following equa-
tion shows that the hydroxyamidosulphate may change by
cumulative resolution, half into a reduced product (amidosul phate),
and half into oxidised products together equivalent to the non-
existent dihydroxyamidosulphate :
500 DIVERS AND HAGA: DECOMPOSITION OF
(1). ° 2Cu(H,NSO,),=N,O+ H,0 + H,80, + CuSO,+
Cu(H,NSO;).=Cu(H,NSO;), + Cu(H,;NSO;),.
Such an equation expresses much of what happens in the de-
composition of a hydroxyamidosulphate at a lower temperature,
but even in this case, and much more in the decomposition of
a hydroximidosulphate by copper sulphate, where the tempera-
ture is higher, a third molecule decomposes in another way.
The result is that the free sulphuric acid shown in the above
equation gets neutralised, and the third molecule of hydroxyami-
dosulphuric acid yields neither sulphate nor amidosulphate, all
its sulphur being eliminated as dioxide, its nitrogen as nitrous
oxide, and its hydrogen as water thus reverting to sulphurous
and hyponitrous acids, just as it does under the influence of an
alkali (p 497) adding to equation (1) that of Cu(H,NSO,),=N,0+
2H,0+2S80,+CuO, we get (2), 3Cu(H,NSO,),=2N,0+4H,0+
2S0,+ 2CuSO,+ Cu(H,NSO,)., with products free from acid.
It is possible to express the decomposition of hydroxy-
amidosulphate differently, by making nitrogen one of the pro-
ducts in place of nitrous oxide, thus:
(3) 8Cu(H,NSO,),=2N,+2H,0+ 2H,SO,+ 2CuSO,+ Cu(H,NSO,).;
(4) Cu(H,NSO,).=N,+2H,0+S0,+ CuSO,.
In (8) sulphur dioxide is not a product whilst in (4) it is.
Whether, however, nitrogen is formed, even in small quantity,
is doubtful. Along with the nitrous oxide soluble in alcohol, we
found a little insoluble gas—about 4 per cent. by volume of the
whole gas,—but we are not prepared to assert that this was not
due to air in spite of the precautions we took to expel all air
from the apparatus by carbon dioxide before the decomposition.
It will be seen from the equations that, with nitrous oxide asa
product of the decomposition, the sulphur appearing as sulphate
HYDROXYAMIDOSULPHATES BY COPPER SULPHATE. 501
equals that as amidosulphate, whereas, with nitrogen as a pro-
duct, the sulphur as sulphate is double that as amidosulphate in
(3), whilst in (4) there is none as amidosulphate. Now, in the
observed decompositions of hydroxyamidosulphate the sulphur as
sulphate has been found equal, on the average, to that as
amidosulphate, a result showing that within the limits of accu-
racy of the somewhat ae analytical work, no nitrogen is
generated.
Although, when using copper sulphate or copper hydroxy-
amidosulphate, no change to cuprous salt is observable, the
reduction of cupric chloride to cuprous chloride points clearly
to the activity of the copper salt as a ‘carrier of oxygen’ from
one molecule of the hydroxyamidosulphate to another.
Results and Method of the Quantitative Experiments.
The results of the experiments are given, not in the order
in which they were obtained but in that of the growth in
quantity of the sulphur dioxide produced.
In an experiment in which copper hydroxyamidosulphate
was heated very slowly, so as to carry out the decomposition at
as low a temperature as possible (boiling the solution only at the
end in order to expel the last portions of sulphur dioxide), re-
sults were obtained which agree sufficiently well with those
calculated on the assumption that 3.7 per cent. of the salt gives
all its sulphur as dioxide, its hydrogen as water, and its nitrogen
as nitrous oxide, whilst the rest of the salt Geconiposes accord-
ing to equation (1):
502 DIVERS AND HAGA : DECOMPOSITION OF
Sulphur as dioxide ; as trioxide and amidosulphate.
Found......... 3.5 96.2
Cale. 3.7 96.3
An experiment with sodium hydroxyamidosulphate and its
equivalent of copper sulphate, gave results indicating that about
5.3 per cent yielded all its sulphur as dioxide, the rest of the
salt giving sulphur trioxide (sulphuric acid) and amidosulphate
(equation 1):
Sulphur as dioxide; as trioxide; as amidosulphate ; as acidity.
Found......... 5.9 46.0 48.0 21.6
Calc. 5.3 47.4 47.4 21.0
Copper hydroxyamidosulphate in four experiments gave re-
sults agreeing nearly with the assumption that 13.2 per cent. of
the salt gave sulphur dioxide, the rest decomposing according to
equation (1):
Sulphur as dioxide ; as trioxide ; as amidosulphate ; as acidity.
Found......13.0 43.0 43.6 11.1
ey 13.0 43.3 43.2
es Seems 13.1 86.6
Le es 86.5
Calc. ......13.2 43.4 43.4 15.1
In anothar experiment copper hydroxyamidosulphate gave
the following results, as against calculation for 15.4 per cent. to
decompose so as to yield its sulphur as dioxide:
Sulphur as dioxide ; as trioxide; as amidosulphate ; as acidity.
Found...... 15.1 42.9 41.6 10.7
Cale. cu. 15.4 42.3 42.3 13.5
In one more trial, copper hydroxyamidosulphate decomposed
nearly as if 16.6 per cent. of it yielded all of its sulphur as dioxide :.
HYDROXYAMIDOSULPHATES BY COPPER SULPHATE. 503
Sulphur as dioxide ; as trioxide and amidosulphate ; as acidity.
Found......16.3 83.3 9.3
Cale. ...... 16.6 83.4 12.5
A solution of potassium hydroximidosulphate heated with
very little more than its equivalent of copper sulphate, gave
results showing that 25 per cent. of the salt yielded all the
sulphur of the hydroxyamidosulphate coming from it by hydro-
lysis, as sulphur dioxide: |
Sulphur as dioxide ; as trioxide ; as amidosulphate ; as acidity.
Found...... 25.2 37.3 37.0 5.5
Calc. ......25.0 37.5 37.5 6.25
A solution containing sodium hydroximidosulphate and
copper sulphate decomposed in two experimenis, in such a way
that about 28 per cent. of the hydroxyamidosulphate sulphur
became dioxide:
Sulphur as dioxide ; as trioxide ; as amidosulphate ; as acidity
Found..... 27.6 36.9 35.0 4
spy. ess 28.0 36.2 35.8 5.8
Calc. ...... 28.0 36.0 36.0 4
The numbers in the above table stand for parts per hundred
of the sulphur of the total hydroxyamidosulphate decomposed,
and not of the sulphur of the hydroximidosulphate even when such
a salt has been that experimented with. The ‘acidity’ sulphur
is calculated as if the acidity is due to sulphuric acid, not
amidosulphuric acid. The ‘trioxide’ sulphur is that of the
sulphuric acid and copper sulphate yielded by the decomposition.
The differences between the calculated quantities and those
found must be iargely attributed to imperfect estimation; they
cannot be due to error in theory, because no other explanation
of the change than that adopted is possible. In a copper-salt
504 DIVERS AND HAGA: DECOMPOSITION OF
solution mixed with much barium sulphate, it was not easy to
titrate acid with lacmoid paper as indicator. The separation of
sulphate and amidosulphate is not a simple process, especially
when much sulphate is present derived from sources other than
the reaction to be dealt with.
The salt employed in the experiments was either copper
hydroxyamidosulphate, or sodium hydroxyamidosulphate with
copper sulphate, or one of the alkali hydroximidosulphates with
copper sulphate.
1. A solution of the copper salt, containing only a very
little copper sulphate was prepared from normal barium hydroxy-
amidosulphate and copper sulphate, the barium salt (this Journ.,
3, 213, 216) had to be prepared as wanted, because of the
instability of the hydroxyamidosulphates. The strength of the
solution was determined by a barium estimation (hydrolysis in
sealed tube and weighing of the barium sulphate). Copper sul-
phate in slight excess and carefully weighed was added to the
weighed solution of the barium salt, and the ‘copper hydroxy-
amidosulphate at once used without filtering off the barium
sul phate.
2. Sodium hydroxyamidosulphate solution was prepared just
before use by hydrolysing a centigram-molecule of the hydrox-
imidosulphate by adding to its solution a minute and known
quantity of sulphuric acid (this Journ., 11, 3) and to it was
added after neutralisation with sodium hydroxide half a centigram-
molecule of copper sulphate.
3. Potassium or sodium hydroximidosulphate in the quantity
of a centigram-molecule was dissolved and directly heated with
a half molecule in centigrams of copper sulphate.
The solution (either 1, 2, or 3) being in a small flask
HYDROXYAMIDOSULPHATES BY COPPER SULPHATE. 505
connected with a tube receiver holding bromine water kept
cold, was heated, sometimes quickly, sometimes slowly, either by
a spirit Jamp or in a bath of sulphuric acid, the solution being
finally boiled for some minutes, so as to drive all sulphur dioxide
into the bromine water. Before heating, air was removed from
the apparatus by a current of carbon dioxide. In one experiment
the apparatus was made entirely of glass. The oxidised sulphur,
dioxide was weighed as barium sulphate.
The boiled-out copper solution was titrated with N/10 soda
(free from sulphate), using lacmoid paper as indicator. The
imperfection of this operation was proved beyond doubt on cal-
culating out the nature of the changes which had occurred, but
it was serviceable and the best available under the circumstances.
To the boiling hot solution and precipitate of barium sul-
phate, barium chloride was added in excess, the total precipitate
collected, well washed, and transferred to a pressure tube in
which it was ' sted with hydrochloric acid for three hours at
150°. The arium sulphate was again washed on the filter, then
ignited and weighed. The second filtrate and washings contained
sulphuric acid, the quantity of which was estimated as barium
salt. To make this part of the analytical process intelligible, it
must be explained that barium amidosulphate, although itself
quite soluble in water, is partially precipitated along with barium
sulphate, even in presence of hydrochloric acid (this Journ., 9,
283). At 150°, the precipitated amidosul phate hydrolyses, yielding
barium sulphate and ammonium sulphate in molecular proportions.
The copper filtrate from the crude barium precipitate was
evaporated to a small volume, heated with hydrochloric acid for
some hours at 150°, and mixed with barium chloride. The
precipitated barium sulphate represented the principal quantity
606 DIVERS & HAGA: DECOMPOSITION OF OXYAMIDOSULPHATES,
of amidosulphate sulphur, the full amount of which was ascer-
tained by adding to it twice the quantity of that in the ammo-
nium sulphate extracted by hydrolysis from the crude barium
precipitate. The sulphur from the hydroxyamidosulphate, ob-
tuined as sulphate, was found by subtracting from the total the
sum of the quantities of sulphur present as (a) copper sulphate
taken ; (b) barium sulphate from the hydrolysed barium amido-
sulphate which had been precipitated along with the barium
sulphate by barium chloride; (c) sulphuric acid added for
hydrolysing the hydroximidosulphate, when that salt had been
started with; and (d) in the same case, sulphuric acid resulting
from the hydrolysis of the hydroximidosulphate to hydroxy-
amidosul phate.
Hardly any attempt was made to estimate the amount of
nitrous oxide liberated. To do so would only have been useful
as a check on the accuracy of the determinations of the amido-
sulphate, and for that purpose the two substances would have
had to be estimated in the products of one experiment. This,
it did not seem possible to do. An experiment in which hydroxy-
amidosulphate was decomposed gave 55.3 per cent. of the
nitrogen as nitrous oxide, as against 56.6 calculated from the
equation most in accordance with amidosulphate and other sul-
phur determinations. The method of measuring the nitrous oxide
and nitrogen was to expel air from the apparatus by a current
of carbon dioxide continued for some time, and then heat the
copper salt and boil out the gases which were collected over
mercury and potassium hydroxide and measured. The alkali
was then replaced by absolute alcohol to dissolve the nitrous
oxide, and the residual gas measured.
ORS ITO
SB en eit,
a ee re a a
CONTENTS OF RECENT PARTS.
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On the Fate of the Binntopare, the Relations of the Primitive Streak, and the
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Studien über die Schwefelrasenbildung und die Schwefelbacterien der Thermen
von Yamoto bei Nikkö. Von M. Mivosnr. (Hierzu Tafel XIV).
Die Entwickelung der Gonophoren bel Physnlia maxima. Von S. Goro. (Hierzu.
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Studies of Reproductive Elements. III. Die Entwickelnng der Pollenkörner
von Allium fistalosam L. ein Beitrag zur Chromosomenreduktion ia
Paansenreiche. Von C. IsHIKAWA. Ilierzu Tafeln XVI und XVII).
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Interaction of Nitric Oxide with Silver Nitrate. By E. Divers.
Preparation of Pure Alkali Nitrites. By E. Divers.
The Reduction of an Alkali Nitrite by an Alkali Metal. By E. Divers.
Hyponitrites : their Properties and their Preparation by Sodium or Potassium
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Identification and Constitution of Fremy’s Sulphazotized Salts of Potassium, his
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Vol. XIII, Pt. II.
TAGE,
Contributions to the Morphology of Cyclostomata. II.—The
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JOURNAL
COLLEGE OF SCIENCE, _
IMPERIAL UNIVERSITY OF TOKYO,
JAPAN.
VOL. XIII, PART IV.
—
KR % BK # ED G
PUBLISHED BY THE UNIVERSITY.
TOKYO, JAPAN.
1901.
MEIJI XXXIV.
Publishing Committee.
— ni —
Prof. K. Mitsukuri, Ph. D., Rigakuhakushi, Director of the College
(ex officio).
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Prof. T. Haga, Rigakuhakushi.
Prof. S, Watasé, Ph. D., Rigakuhakushi.
All communications relating to this Journal should be addressed to the
Director of the College of Science. .
Observations on the Development, Structure and
Metamorphosis of Actinotrocha.
By
Iwaji Ikeda, Rigakushi.
With Plates NYV-NXX.
Introductory.
Since the discovery of Actinotroeha by JOHANNES MULLER
in 1846, this peculiar Jarval form and its mother animal,
Phoronis, have been made the subject of investigations by many
distinguished authors such as WAGENER (47), GEGENBAUR (’54),
Kronx (54), Scunemer (62), MeTscHNIKorF (72, '82), FE.
B. Witson (81), and Fortrrincer (82). Among more recent
writers CALDWELL (85), Mc’Ixtos (’88), Benmam (’88),
Roue (’90, ’96), Cort (’9I), and E. Schutze (97) may be
mentioned as having published important contributions; while
MASTERMAN ('97) has made quite an elaborate study of the
animal with the view of establishing its relationship to Palano-
glossus and the Chordata in general. As, however, in spite of all
these works there still existed many gaps and unsatisfactory points
508 I. IKEDA:
in our knowledge of this interesting animal, the investigation,
of which an account is given in the following pages, was under-
taken, and though the results are far from exhaustive, I hope
they will help to advance our knowledge of the subject.
My study was begun in the summer of 1898 during a stay
at the Misaki Marine Biological Station and later was continued
at that Station as well as in the Zoological Institute of the Science
College.
At Aburatsubo, a small inlet close to the Station, is found
a species of Phoronis, which has been named by Dr. OKA (’97)
P. yimar.* Its colonies adhere to the overhanging ledges of rocks
near the shore. As the water at the place is always calm and at
low tides recedes so as to almost expose the ledges, the animals
can be easily collected. During the greater part of the year, eggs
and young embryos, clustered together, in what may conveniently
he called embryonal masses, are found adhering to the lophophoral
crown of the adult, one on each side of the median line. These
furnished materials for the study of fertilization, segmentation and
the early larval stages. The larvie in the Actinotrocha stage are
found swimming in the inlet and are caught with the surface net.
As will later be fully described, there oceur four kinds of the
larvee, which no doubt represent as many species, including the
common Phoronis ijimat.
The ‚specimens, both adult and larval, were killed with the
saturated solution of corrosive sublimate in 1% acetic acid or with
Flemming’s fluid. Of the various colouring methods tried on the
sections, double-staining with eosin or safranin and Delafield’s
haematoxylin gave the most satisfactory results.
ran u a re eee
* For a discussion of the statns of this species, see Supplementary Notes.
ON DEVELOPMENT ETC. OF PHORONIS. 809
Before proceeding further, I beg to tender my sincere thanks
to Professors Muirsukuri and Tsmma for their kind supervision
of my work and for their painstaking revision of my manuscripts.
Contents.
I. The Early Development of the Phoronis Larva,
a. Notes on Fertilization.
b. Notes on Segmentation.
c. Gastrulation and Mesoblast-Formation.
dd. Further Observations on the Development of the Larva, up to
the Period when it becomes free-swimming.
If. The Structure of Actinotrocha.
a. The External Appearance of Actinotrocha.
b. The Internal Structure of Actinotrocha.
1. Body-Divisions and Body-Cavities,
2. Organs of Kctoblastic Origin.
3. Organs of Entoblastic Origin.
4+, Organs of Mesoblastic Origin.
III. Metamorphosis.
IV. Supplementary Notes.
I. The Early Development of the Phoronis Larva.
a. NOTES on FERTILIZATION.
Phoronis is, as is well known, a hermaphrodite, in which doth
the male and female sexual elements mature at nearly the same
time. But few authors seem to have studied the animal during its
breeding season, su that our knowledge of its sexual organs and,
consequently, of its fertilization has remained very imperfect, as
was pointed out by Corr (91). The only existing statement as
to how and where fertilization is accomplished in Phoronis is that
of Kowatewsxy (67). This author thought that fertilization
510 I. IKEDA:
took place in the body-cavities, and accordingly, as Corr remarks,
he must have believed that self-fertilization prevails in Phoro-
nis. Cori considers this as highly improbable, but does not bring
forward any positive facts in contradiction of it, his inference
being drawn solely from facts observed in other marine Metazoa.
Since Kowarewsky’s valuable researches (67), it has
generally been accepted that the nephridia serve also as oviducts.
Thus BENHAM says that he saw an ovum attached to one side
of the nephridial funnel and further mentions that KowaLewsky
observed eggs moving through the nephridial canal towards the
exterior. Unfortunately both observers failed to elucidate what stage
of development these eggs are in.
In Phoronis ijimai mature sexual elements are constantly
discovered throughout about one half of the year (from November
to May or June). By carefully examining a living colony of that
species during this period, it will soon be perceived that some indivi-
duals differ slightly from the rest in the aspect of the foot or
body. We see in them a moniliform series of small white specks
shining through the skin in the uppermost part of the body.
These are the ova ready to escape to the exterior through the
nephridia. It must have been such individuals that were observed
by Kowarewsky and BExuam. The body-cavity, in which the
ova lie, corresponds to the rectal chamber near the anterior end
of the body. I have endeavoured to ascertain whether these ova
are fertilized or not, and have at last succeeded in ascertaining
that they are in a stage prior to the extrusion of polar globules,
—the primary oocytes, in Bovert’s terminology. In the fresh
state, they are spherical or somewhat elliptical in shape and per-
fectly opaque by virtue of the abundant volk-granules contained in
the vitellus. It is characteristic of these ova that the nucleus, which
ON DEVELOPMENT ETC. OF PHORONIS. oll
is situated, not in the centre, but near the periphery, is al-
ways in the meta- or ana-phase of Karyokinesis (fig. 17). In
such an ovum the chromosomes are constantly found to be six in
number, each being dumb-bell shaped with the two ends directed
towards the poles. Fig. 18 represents a portion of the section
passing through the equatorial plane of the nuclear figure. It is
evident that these eggs are in preparation for the extrusion of the
first polar globule. As shown in the above figure, the finely
granular protoplasm of the vitellus contains thickly and uniformly
distributed yolk granules, which have a strong affinity for eosin.
That the eggs in question are mature is further demonstrated
by the fact. that I succeeded in artificially fertilizing them and in
rearing out of them normal embryos which grew to certain ad-
vanced stages of development.
If we now examine the embryonal masses, which, as has
been mentioned, are found attached one on each side of the ten-
tacular crown of the adult Phoronis, we find that the embryos
which are farthest away from the nephridial pores are the most
advanced in development and that they are found in successively
younger and younger stages as we approach the pores, until we
reach such eggs as have just been fertilized or perhaps even such
as have not yet been fertilized at-all. But even the youngest eggs
found in the mass present an appearance very different from
those found in the body-cavities, the former being invariably at
a stage after the expulsion of one or two polar globules, In the
egg taken from the mass and shown in fig. 19, two polar globules
have already been formed ; these are situated close together just
inside the vitelline membrane.
On the other hand, if we examine by means of serial sections
through the posterior region of an adult, where the stomach and
812 I. IKEDA:
the sexual organs lie grouped together, a number of large eggs
are frequently found, floating freely iu the coelomic fluid of the
bodv-cavities. These egus do not differ in any respect from those
in the nephridial region as regards the size, the appearance of
the karyokinetic figure, or the number of chromosomes.
The facts above stated plainly point to the following con-
clusion :— The oogonia fall into the body-cavities by a dehiscence of
the ovarian walls and here develop until they reach the stage of
primary oocytes. These travel gradually upwards to the nephridial
region, retaining meanwhile the nuclear figure formed for the
production of the first polar globule. Reaching that region, the
primary oocytes are destined sooner or later to be carried by way
of the nephridia to the exterior, where they become fertilized by
spermatozoa from other individuals.
Reserving an account of the spermatogenesis and ovogenesis
for a future occasion, I may here refer to a few facts observed
by me relative to the process of fertilization. When the two
sexual elements are artificially brought together, numberless sper-
matozoa svon attach themselves to the surface of the ovum. About
10 minutes afterwards, the first polar globule makes its appear-
ance, followed soon afterwards by the second. Meanwhile a small
clear spot, probably marking the place where the male element has
entered, appears on the surface of the egg; it is however observ-
able for only a very short time. Both figs. 19 and 20 are
sections of ova taken from the embryonal mass. The ovum given
in fig. 19 is fully mature and ready to be fertilized; close to the
polar globules rests the large female pronueleus. The ovum
represented in fig. 20 belongs to a stage of fertilization in which
the two pronuclei stand elosely side by side. The larger female
pronucleus has a nuclear membrane irregular in contour. The
ON DEVELOPMENT ETC. OF PHORONIS. 513
intensely stained chromatin pieces are in both nuelei dispersed
without any apparent order throughout the finely granular nuclear
substance. At one spot outside the male pronucleus, there is
visible a small and clear archoplasmie (?) space surrounded by
a set of exccedinglv fine radial rays. The two polar globules of
this ege were distinctly visible in other sections whieh have
not been figured.
b. NOTES ON SEGMENTATION.
Our knowledge of the mode of segmentation in Phoronis is
far from being satisfactory. METSCHNIKOFF (82) gives no account
of the process, FOETTINGER (82), if one may judge from his
figures, seems to have seen the egg undergoing to taland unequal
sesmentation. According to CALDWELL (’82), the segmentation
‘ proceeds with considerable regularity ”’ (Le., p. 374) ; Roue (’90)
says “Porule fécondé subit une segmentation totale fort requliére...... à
(Le, p.1147). E. Scuuntzr (97) simply says “ /eh sah das Ei
sich total und uniiqual furehen” (Le, p. 6).
My observations of the process were made on eggs found in
the embryonal mass as well as on those artificially fertilized. As
the former showed comparatively rarely the earlier stages of
the segmentation, it was necessary to have recourse to the latter
for filling up the gaps of observation.
Soon after the formation of the second polar globule and the
disappearance of the micropyle-lke spot the first cleavage line
makes its appearance, passing on one side of the polar globules
(fies. 1 and 21). At this stage I can not perceive any difference
in sizé and structure between the two blastomeres. The second
514 I. IKEDA :
cleavage plane passes at right angles to the first (fig. 2 0). It
is a remarkable peculiarity of Phoronis eggs that two sister blas-
tomeres derived by the division of a mother blastomere, never
undergo the next division simultaneously, so that between any two
consecutive stages having an even number of blastomeres there
intervenes an intermediate stage with an odd number of the same.
This phenomenon occurs even at the second cleavage ; thus just
before the egg attains the four-cell stage, there exists a stage of
three cells, such as is seen in fig. 2 a. Among the later stages,
those of 5, 7, 9.0.0.0... cells are of constant occurrence. Consequ-
ently it is scarcely admissible to say that the segmentation pro-
ceeds with considerable regularity.
CALDWELL (’82) has asserted that the first differentation of the
future blastoderm into the ectoblast and entoblast is observable
as early as in the four-cell stage. He says: “ At the stage of 4
segmentation-spheres a division into two smaller clear and
two larger opaque cells indicates the future ectoderm and endo-
derm” (Le, p. 374). At the corresponding stage of Phoronis
ijimat I have not been able to discover any appreciable difference
in the size of its cells (see fig. 2 à). Following the 4-cell stage,
the division of the blastomeres in the equatorial plane puts the egg
on the way to the 8-cell stage. According to my own observations,
the above mentioned difference in size of the blastomeres becomes
first perceptible at this stage. Fig. 3 shows a side view of an egg
with 8 blastomeres ; it will be seen that the upper four blastom-
eres are very slightly smaller than the lower four. TI could not
however, at that period, recognize any difference in the cell-con-
tents of the two classes. ,
The irregularity of division, which, as before mentioned, be-
comes more and more pronounced as segmentation advances, tends
ON DEVELOPMENT ETC. OF PHORONIS. 515
to gradually obscure the orderly arrangement of the cells. At the
16-cell stage the regular arrangement is still, though less distinctly,
maintained, while at the 32-cell stage it is quite disturbed (fig. 4).
From this period on, the polar globules can no longer be detected.
In the earlier stages of segmentation, the blastomeres are
found in close contact with one another, leaving no noticeable
space or segmentation-cavity between them. After they have
increased to about 32 in number, the blastocæle and its opening
to the exterior (fig. 4, dlc.) become recognizable. The embrvo
at the morula stage is somewhat oblong in shape and has
a quite spacious blastocæle, and the blastocælic pore (bl.e.)
is distinet on the ventral side (fig. 5). However, this pore dis-
appears at an advanced morula stage, and apparently the vitelline
membrane also, nearly simultaneously with it. At any rate both
have altogether passed out of sight at the next stage, that of the
blastula. In fig. 23, which represents a median section of a young
morula (the outline of which has undergone mutual compression
by the crowding together of embryos), the pore (d/.c.) is cut through
and appears as a slit-like passage between two of the bounding
blastomeres.
In the blastula (fig. 25) the wall consists of evlindrical cells
and encloses a tolerablv wide blastoccelic cavity, which is now at its
greatest development. In this stage, the bilateral symmetry of
the future larva is already established. It has an oblong plano-
convex form, the flattened face of which corresponds to the future
ventral face; and its ends, one somewhat broader than the other,
indicate respectively the future anterior, and posterior, ends.
The cells of the wall are all cylindrical in form, as shown in fig.
25, those of the ventral side being slightly larger than those on
516 I. IKEDA:
the convex dorsal side. The nuclei in all the blastodermal cells
are always situated in a peripheral position.
Plasmic corpuscle.—A noteworthy fact with regard to the
blastula is that in its older stage a certain number of small and
non-nucleated plasmic spheres is almost constantly met with in the
blastocælic cavity (fig. 25, pl.co.) These have been first described
by FOETTInNGER under the name “corpuscules mésodermiques.”
According to this author, these corpuscles are free nuclei imbedded
in a common protoplasmic mass which is supposed to fill up the
blastocæle, each corpuscle bocoming a mesoblast cell, after appro-
priating to itself a certain portion of the surrounding protoplasm.
This view of FoETTINGER, which certainly can not be accepted
at the present day, was, I believe, partially due to the then
defective technique. His method consisted in pouring dilute acetic
acid over the living embryo, and this, as the author himself was
well aware, is highly detrimental, in that it frequently breaks up
the blastomeres into fragments. The corpuscles deseribed by him
from so early a stage as that with only 8 blastomeres must have
been simply produced by fragmentation, the result of his drastic
treatment. The common protoplasmic mass supposed to be present
in the blastocæle, was probably nothing but a coagulum.
Again the mesodermzellen which MeEtscunikxorr ('82) found
in the blastocælic cavity of the blastula are certainly not true
mesoblast cells but rather certain spheres similar to Fortrix-
GERS “ corpuscules m&sodermiques,” as was rightly pointed out by
CALDWELI. Recently E. ScHuLtze (’97) published a short paper
entitled “ Ueber die Mesodermbildung bei Phoronis,” in which he
writes as follows :—“ Schon auf dem Stadium der rundlichen
Blastula sehen wir einige Mesodermzellen im Blastocel aufsitzten ”
ON DEVELOPMENT ETC. OF PHORONIS. 517
(/.c., p. 6). It seems to me that SCHULZE has fallen into the
sıme mistake as METSCHNIKOFF.
Lastly, CALDWELL (’82) has entertained a view quite different
from those of other writers. According to him, the bodies in
question are not present as such in the blastocæle, but are in
reality only the cut ends of blastoderm cells projecting into the
cavity and as such of course have nothing to do with the true
mesoblast.
In the Phoronis studied by me, the plasmic corpuscles are
present only in the highly advanced blastula (fig. 25, pl.co.).
They are usually round in shape and very much smaller in size
than any of the blastoderm cells, but as to structure, they do not
show any deviation from the latter, except in the important res-
pect that they have no nucleus. Although I tried with them all
the available nuclear stains, the presence of any chromatic sub-
stance in them could in no instance be detected. In an earlier
stage such a free floating sphere has never been met with. In-
stead of it, some unusually elongate blastoderm cells (pl. co., fig.
24), such as were found by CALDWELL, were discovered: protruding
their inner end into the blastocæle. The nucleus of these cells
commonly lies in the periphery as in all other cells of the more
ordinary shape. In my opinion, the proximal ends of the elong-
ate cells break off from the main cell-body and fall into the
blastoceele, where they undergo degeneration, breaking into ever
smaller and smaller spheres. By examining serial sections of an
advanced stage like that of fig. 25, it is easy to convince one’s
self that there exists no connection whatever between the spheres
and the blastoderm cells. The spheres, or the plasmic corpuscles,
are clearly distinct bodies and not mere ends of blastoderm cells
cut off in the process of microtomizing as was supposed by
018 I. IKEDA :
CaLpWeELL. Their small size and the total absence of nuclear
substance make it easy to distinguish them from the true meso-
blast cells.
The corpuscles are still frequently discovered in the blasto-
calie cavity at the beginning of gastrulation, together with a few
mesoblast cells. But in an advanced gastrula they have wholly
disappeared, possibly having been absorbed by the blastoderm cells.
I think the temporary co-existence of the plasmic corpuscles and
of the true mesoblast cells in the blastocæle of the gastrula, has
led some previous authors to confound the two elements. As to
the significance of the corpuscles, I can at present offer no opinion
“unless they be merely an excess of supply of nourishment analo-
gous to food yolk’ as has been suggested by CALDWELL ('82, Le.,
p. 18).
c. GASTRULATION AND MESOBLAST-FORMATION.
In this section, I shall first deseribe what I conceive to be
the true history of gastrulation and mesoblast-formation, and then
pass on to a discussion of the views of other writers. The two
developmental processes are so intimately related to each other,
that it seems best to treat them together.
First as to external changes. The bilateral symmetry of the
plano-convex blastula becomes more clearly marked than before
when the gastral invagination begins on the ventral or tlıe flat-
tened side. The initial depression oceurs over the whole ventral
wall, so that a saucer-shaped embryo is produced. At first it ıs
so shallow as to be perceived with difficulty in the surface view.
Soon it deepens, becoming deepest at a point somewhat nearer to
the broader end than to the narrow end of the embryo. The
ON DEVELOPMENT ETC, OF PHORONIS. 519
deepest portion may conveniently be called the central depression.
Fig. 6 represents the ventral view of an embryo in which the
invagination has become visible from the outside, the central
depression being most deeply shaded in the figure. In a slightly
more advanced stage, as the original wide depression grows decper,
the external opening is gradually drawn together and at a certain
stage (fig. 7) becomes transformed into an almost triangular
blastopore situated at a position slightly anterior to the centre, as
was the case with the central depression. The anterior side of
the triangular blastopore is somewhat rounded by curving uniformly
outwards, while posteriorly the two other sides gradually approach
each other so as to meet at a point which may be called the apex
of the triangle. Leading backwards from this apex, there runs in
the median line the so-called primitive groove. This latter and
also the triangular shape of the blastpore are occasioned, in my opi-
nion, simply by the blastopore, originally broadly open, becoming
narrowed by the special activity. in the lateral posterior parts of
its posterior section. In other words, the cell-multiplication of the
ectoblastic layer is carried on especially in the last mentioned
parts, so that there the pressure, which is exercised by the ccto-
blast towards the invaginated layer, is more marked than in the
anterior and lateral borders of the blastopore. As the result of
the above phenomenon, the definitive blastpore is pushed further
anteriorly, and consequently, the archenteric cavity deepens in the
posterior direction, as shown in fig. 29. The above consideration
is supported by the results of actual measurements of the size of
the embryos concerned. In spite of the fact that the embryo has
developed considerably the body-length does not show any signi-
ficant increase, remaining all the while at about 0.12 mm. on an
520 I. IKEDA:
average. This shows that the growth is lost in the curvature of
the body.
When the growing larva reaches the stage represented in
fig. 8, the blastpore assumes a narrow transversely directed, slit-
like form. That portion of the larval body lying in front of the
blastpore— which is the persistent larval mouth—protrudes more
vr less prominently forwards and ventrally, so as to acquire the
form characteristic of the preoral lobe of Actinotrocha. In such
an advanced gastrula, the primary gut-cavity is well established
and can be plainly traced through the wall in the surface view.
If the larvæ of such an early stage of development be taken out
of the embrvonal mass and set free in water, they will swim about
by means of the well developed cilia, which cover the whole
external surface.
Fig. 9 represents a side view of a larva, in which the pre-
oral lobe has grown to a very considerable size. The nephridial
pit, which is an ectublastic invagination just in front of the
posterior end of the gut, is now distinctly visible from the out-
side. In short, the larva may be said to possess the inceptive
characters of an Actinotrocha.
I will now proceed to describe the internal changes accom-
panying gastrulation. The earliest symptom of this process can
be seen in sections before it can be detected from the surface.
It consists at first in a peculiar disposition of those blastodermic
cells which constitute the ventral portion of the blastula wall.
This portion not only shows a shallow concavity, but also the cells
composing it become, as figs. 26 a and 26 6 show, irregularly
arranged on account of mutual pressure, as a result of which some
of the cells are even forced out of file so as to fall into the
blastocwle. These liberated cells have usually a round shape and
ON DEVELOPMENT ETC. OF PHORONIS. 521
of course contain each a distinct nucleus. Some other cells are
apparently in the process of being pushed out and have a club-like
shape, the narrowed end being still inserted between the cells
of the layer to be invaginated. A further symptom of incipient
invagination consists in the circumstance that the nucleus in most
cells of this portion has no longer a peripheral position, but is
situated in the middle or rather nearer to the inner end of the
cells. Moreover, the nucleus is frequently met with in the form
of the karyokinetie figure which shows that the cells are dividing
and increasing in number in the layer to be invaginated. The
cells pushed out into the blastocele are nothing else than mesoblast
cells, so that it may be stated that the mesoblast-formation begins
simullaneousiy with the gastrulation.
At the beginning of gastrulation we can thus distinguish two
parts in the blastoderm wall, azz., the mesentoblast and the ecto-
hlast. The former corresponds to the whole of the portion to be
invaginated, while the ectoblast forms all the remaining portion of
the embryo. The mesentoblast is composed of large and irregular-
lv arranged cells, while the ectoblast is of taller cylindrical cells
regularly arranged in a single row (figs. 26 a and 26 8). The
mesentoblast, as the name indicates, is destined to give rise to
both the entoblastic and mesoblastic elements. The characteristic
disposition of its constituent cells indicates its being the source of
mesoblast proliferation.
The mesoblast proliferation becomes more and more accen-
tuated in activity as the gastral invagination gradually deepens
(see fig. 27), but the mesoblast cells thus formed do not adhere as
a lining epithelium to the ectoblast, until what are called the ant-
erior diverticula have been formed on both sides of the blastopore.
The blastocele, in which the mesoblast cells are at first loosely
22 I. IKEDA:
seattered about, is henceforth greatly reduced in extent and finally,
as the development of the archenteron progresses, is almost obli-
terated, especially along the dorsal and lateral portions of the
embrvo where the ectoblast and the gut come into direet contact
with each other (see figs. 29 and 302).
Figs. 28 a-c show three cross sections through different parts
of a larva of nearly the same stage as that represented in fig. 6,
in which the invagination has become recognizable in the surface
view. Fig. 28a passes through the central depression which be-
comes gradually shallower posteriorly (figs. 235 and 28e). As
these figures show, the mesoblast. cells are at this stage still being
proliferated uniformly from every part of the mesentoblast and do
not yet form a lining epithelium to the ectoblast. When the blasto-
pore has taken a triangular shape (fig. 7) and the primary archen-
teric cavity has somewhat bent itself towards the hind end, the
posterior border of the blastopore has travelled a certain distance
in an anterior direction. If we examine serial sections of this
region, a narrow and shallow groove is detected running for a
short distance immediately behind, and from, the meeting point of
the lateral blastopore lips. Also at about this stage, a paired in-
vagination, the anterior diverticula of CALDWELL, appears along
the side of the anterior portion of the archenteron. These points
will become clear from a consideration of figs. 30 a-e, which
are drawn from serial transverse sections of an _ embryo
slightly older than that shown in fig. 7. In fig. 30a, showing
the right-hand side of the blastopore, we notice a lateral infolding
(ant. div.) of the archenteric wall a short distance inside the
blastopore lip. Here the component cells are irregularly arranged
and their entire disposition reminds one of the mesentoblastic
laver. Indeed some indubitable mesoblast cells are found pressed
ON DEVELOPMENT ETC. OF PHORONIS. 523
against the tip of the diverticulum. No doubt the mesoblast is
here arising, not by direct cell multiplication, but by the pushing
in of the cells of the diverticulum. This is more clearly illustrated
in fig. 31, which shows a transverse section through the blasto-
pore of a more advanced larva; here the mesoblast cells almost
fill up the blastoccelic cavity on both sides of the blastopore.
In fig. 305, a transverse section just behind the closure of the
blastopore, the most anterior portion of the primitive groove before
mentioned is cut across. Here the wall of the groove underlying
the gut is formed of mutually compressed cells, some of which are
evidently migrating into the blastocæle (on the left-hand side in
the figure). If the sections are followed further posteriorly, the
groove still persists, but no mesoblast cell in the actual immigrating
process can be discovered, although there are those which have
been previously pushed out and are now floating between the two
primary germinal layers at this region. Still more posteriorly the
groove entirely disappears and the entoblastie and ectoblastic
layers are separated from each other by the comparatively wide
blastocelie cavity (fig. 30 +). At this stage, therefore, the greater
part of the archenteric wall has ceased to contribute towards the
mesoblast-formation ; in other words, it has lost its mesentoblastic
nature. The mesoblast is now being produced only from two
limited regions, viz., anterior diverticula and the ventral groove.
In a slightly more advanced larva, the ventral groove is still
present for some distance immediately behind the blastopore, but
the laver which forms the groove has entirely ceased to give rise
to mesoblast cells (fig. 32, which is taken from a transverse sec-
tion very near the blastopore). It appears to me that this groove
is to be regarded as but the posterior portion of the original
mesentoblast, which, owing to the fact that the central depression
De Fi me a En nee A, oad gg A cmt
524 I. IKEDA:
is eccentrically placed nearer to the anterior end, has to traverse
a longer distance before it can be reflected inwards, and thus on
its inward course lags behind the anterior and antero-lateral por-
tions. Eventually all the cells of the wall of the groove that are
left behind after proliferating the mesoblast cells, are without
doubt invaginated and form a part of the entoblast. The groove
then entirely disappears. I could not discover any remnant of it
in any part of the posterior region where, according to Ca.-
WELL, the ectoblast and the entoblast are said to stand in fusion
to give rise afterwards to the anus. In such an advanced stage,
the anterior diverticula have also ceased to give off mesoblast
cells and have become straightened out, their walls acquiring a
normal epithelial character (entoblastic).
From the facts above adduced, it may be concluded that
both the anterior diverticula and the ventral groove, present at a
cerlain developmental stage of the Phoronis embryos, are remnants of
the original mesentoblast which at an earlier stage occupied the
the whole extent of the gastral invagination. They are, therefore,
merely temporary, and destined sooner or later to split into meso-
blastic and entoblastic cells.
As will be seen in figs. 30 a-c, the ectoblast and the archen-
terie walls are brought together into such close contact, especially
along the dorsal and lateral regions that scarcely any interspace
is left between them. In the embryo given in fig. 8, the cavity
of the rudimentary preoral lobe is filled with mesoblast cells pro-
duced from the original mesentoblastic layer. So far as I can make
out, these show no difference whatever from those proliferated
from the anterior diverticula: both are indistinguishably mixed
together. Though most of the mesoblast cells in the preoral lobe
lie loose during the active period of the diverticula, there are
ON DEVELOPMENT ETC. OF PHORONIS. O25
found a few that have already apposed themselves flatty to the
ectoblast (see fig. 29), while the cavity behind the blastopore still
remains without a mesoublastic lining. This last condition persists
till the period when the nephridial invagination makes its appear-
ance. The state of things in question is to be scen in fig. 29,
which represents a median sagittal section through an embryo of
nearly the same stage as fig. 8.
Soon afterwards the anterior diverticula and the ventral
groove entirely disappear and the preoral lobe begins to bend more
distinctly downwards. Meanwhile an unpaired ectoblastic invagi-
nation appears at the posterior end of the larva, on the ventral
side of the blind end of the now greatly elongated gut. It appears
at first as a shallow depression (fig. 33, nep. p.) of purely ectoblastic
nature, having nothing to do with the mesentoblast. It is from
this invagination that the future nephridia of Actinotrocha de-
velop and hence I shall call it the nephridial pit, in preference
to the name “anal pit” of CALDWELL, who for the first time des-
cribed this structure. I have very frequently noticed signs of
vigorous cell-division in the cells of the pit wall, evidently only
for enlargement of the pit itself, since the axis of the karyokinetic
spindle is always placed paratangentially to the wall. I have
moreover often noticed peculiar ectoblastic cells round in shape and
in process of multiplication, situated just outside the edge of the
entrance to the pit (fig. 33).
In larvæ of the stage of fig. 9 the nephridial pit can be well
seen in surface views. This stage further attracts our special at-
tention on account of several important developmental processes
taking place in it. First to be noted is the fact that from the
posterior end of the primary gut a small and short evagination
protrudes itself touching the ectoblast with its blind posterior
026 I. IKEDA :
end. This hollow protuberance is the rudiment of the intestinal
anal of Actinotrocha. In longitudinal section it is shown in fig.
37 (int.).
In fig. 54, representing a slightly oblique frontal section of
a larva of nearly the same stage as that of fig. 9, we see below
the pit-like nephridial sac, which is quite free from the gut. The
ectoblastic wall of the preoral lobe is at this stage somewhat
uniformly lined with flattened mesoblast cells, while in the cavity
behind the blastopore the mesoblast cells are for the most part
freely scattered, though a few have already begun to arrange them-
selves against the ectoblast layer in this region. In fig. 35, a
transverse section through the posterior end of a larva of nearly
the same stage, the nephridial pit appears as a single flattened
sac (nep. p.) lying in front of the intestine (int.) ; the ectoblastic
wall is internally lined with a few isolated and flattened mesoblast
cells. In a slightly more advanced stage, the ectoblast behind
the blastopore, and in a less complete degree the gut wall
also, shows a similar mesoblastic lining, though a few mesoblast
cells still remain free, especially in front of the nephridial
sac.
In order to facilitate comparison with the statements of other
writers, I will here add a few words on the change of form under-
gone by the nephridial pit. When in a larva slightly older than
that of fig. 9, the preoral lobe and the future intestinal portion of
the gut have become considerably elongated, the nephridial pit,
which has meanwhile become deeper than before, begins at i
inner blind end to divide into two lateral branches. Each of the
latter corresponds, as will be fully demonstrated further on, to the
nephridial canal of Actinotrocha. Fig. 38, a frontal section of à
larva at this stage, shows the bifurcation just alluded to. The
ON DEVELOPMENT ETC. OF PHORONIS. 227
relation of the unpaired nephridial sac to the gut will be best
understood from the median sagittal section given in fig. 37.
I may here be allowed to put in a short historical review of
the mesoblast-formation in the Phoronis larva.
KoWwALEWsKY (67) attributed the origin of the mesoblast to
the ectoblast.
METSCHNIKOFF (82), FOETTINGER (’82), and E. SchuLtzE
(97) confounded the plasmic corpuscles with the true mesoblast,
and none of them was aware of the presence of the anterior diver-
ticula.
CALDWELL ('85) made many interesting observations on the
mesoblast-formation. According to his view, there exists no meso-
blast before the closure of the blastupore Hips (lateral), but it arises
later from three distinct sources, viz. 1) the anterior paired
diverticulum (entoblastic), 2) the posterior paired diverticulum
(ectoblastic) and 3) “the primitive streak ” connecting the above
two structures. Further it has been declared by him that the
body-cavities of the larva arise in two different regions. As to
the preoral body-cavity, he writes as follows: “ From the time
when two or three mesoblast cells are budded off from the diverti-
cula on either side, a cavity is present in cach mass thus formed.
These cavities are the two halves of the body-cavity (preoral) ”
(Ze, p. 374). On the other hand with respect to the posterior
body-cavity, he states that “7 28 formed independently in a
paired mass of cells which grow out to the end of the first formed
sacs and remain separated from septum” (l.e., p. 376). Thus he
regards the preoral body-cavity as arising after the enterocælic
type. Lastly the author puts forward in his recapitulation the
opinion that the blastopore gives rise to both the mouth and the
a NUS.
228 I. IKEDA : ,
Route ('90) also distinguished two sorts of mesoblast cells in
view of their different origin and fate : “ Mesenchymes primaires ”
and “ initales mésodermiques.” Both are derived from the “ pro-
toendoderme” which forms the primary archenteric wall. The
latter gives rise to cells grouped together into two compact
‘ bandlettes m&sodermiques,”” which are regarded as homologous with
the mesodermal bands of Annelid larvæ. In reality these bands
are, as have been pointed out by SCHULTZE, nothing else than the
posterior paired diverticula of CaLDWELL.
From the account of the mesoblast-formation given in the
foregoing lines, it is evident that the first stages of that process
are observable from the very beginning of gastrulation (figs. 26 a
& 0), and long before the blastopore takes the small triangular
shape. On this point my observations stand at variance with
CaLDWELL’s. Nor can I agree with that author in the opinion
that the mesoblast produced from the anterior diverticula (even
though consisting of only two or three cells) incloses an enterocwlic
cavity. As already described, the cells in question, after being
budded off, lie loose in the blastocæle together with preexisting
mesoblast cells and without forming a wall to a special cavity of
any sort.
As to the ventral groove, METSCHNIKOFF (’82) was the first
to refer to this structure and wrote as follows: “ Jn passender
Lage des Embryos kann man eine in Verbindung mit dem Blasto-
porus befindlichen Furche (longitudinale) wahrnehmen, welche zum
Ilinterende des Embryos hinzieht und sich nur auf dem Ektoderm
beshränkt. Diese Furche erhöht den bilateralen Bauplan des
Embryos erscheint indessen als eine vergüngliche Bildung, welche
man auf späteren Sladium vergebens suchen würde” (l.c., p. 301).
According to CALDWELL, this groove, which he calls “ the primi-
ON DEVELOPMENT ETC. OF PHORONIS. 929
tive streak,” is produced by a fusion of the blastopore lips; the
cells along the fusion line differentiate after multiplication into the
epiblast, the hypoblast, and the mesoblast. And the rapid growth
of the epiblast in this region soon obliterates the groove, leaving
however its posteriormost portion as the “anal pit.” But such, as
I have tried to show, is not the case, for the so-called primitive
streak entirely disappears leaving no trace whatever, long before
the nephridial or anal pit makes its appearance. Therefore there
exists no direct genctic relation between the primitive streak and
the anal pit.
CALDWELL'S view that the two nephridial pouches give off
the mesoblast, which eventually lines the posterior bodv-cavity,
can not be sustained ; for, according to my own observations, that
body-cavity with its mesoblastie lining wall is already present before
the nephridial pit divides into the two pouches. It is true that
the cells floating in the posterior body-cavity are in some sections
found aggregated at the blind ends of the pouches as shown in
Fig. 38. This is a condition which might mislead one to the
conclusion that mesoblast cells are here in process of proliferation.
But solid cell accumulation in such a section is to be considered
as simply due to the obliteration by compression of the lumen of
the nephridial pouches. Fig. 36 taken from an obliquely cut
sagittal section through a larva of this stage, shows no wander-
ing cells in front of the pouches (of which only the right one is
seen in the figure); in this case there is certainly no doubt about
the matter.
Finally as to the anus, CALDWELL mentions “a solid cord of
cells” which he considers to be the posterior remnant of the
primitive streak. According to him, this acquires a lumen and
forms a fine canal leading from the primary gut cavity to the
530 I. IKEDA :
exterior. However, it seems clear to me that this cord is nothing
else than an early stage of the intestinal outgrewth independently
produced at the posterior end of the gut. Moreover, in Phoronis
yimai, the gut cavity does not come into communication with the
exterior at so early a developmental stage as CALDWELL observed ;
in that species, the anus first opens at a definite stage when the
larva bears two pairs of larval tentacles.
E. SCcHULTZE ('97) rejects CALDWELL’s views in regard to
the anal pit, but regards it as a rudiment of the future ventral
pouch of Actinotrocha, This is, however, certainly not true, since
the ventral pouch is a thing that has a distinct origin and appears
at a much later stage of larval development.
dl. FURTHER OBSERVATIONS ON THE
DEVELOPMENT OF THE LARVA.
Some authors have recorded that the larva swims about
abroad at such a stage of development as is represented in fig. 8.
However in Phoronis tjimai, the larva lies hidden in the lop-
hophoral loops of the mother until it has acquired at least two
pairs of larval tentacles.
In the larva shown in fig. 9, the somewhat prominent preoral
lobe hangs over the larval month. Local eetoblastie thickenings
occur at two places, viz., at the centre of the upper surface of
the preoral lobe and along the mid-ventral line near the posterior
end of the body. The former is the future nerve ganglion; the
latter, the rudiment of the first pair of larval tentacles. The
nephridial invagination at the posterior end is still shallow.
At a little later stage, the tentacular thickening divides into
x
ON DEVELOPMENT ETC. OF PHORONIS. 531
two more prominent ridges running on each side obliquely ant-
eriorly. The preoral lobe grows rapidly so as to hang down on the
ventral side and as a consequence of this an cesophageal canal is
formed (fig. 37, es). The cesophageal wall is, therefore, ecto-
blastic in origin and is composed of strongly ciliated columnar
cells. About this period the nephridial invagination becomes
completely divided into two lobes at the proximal end, as I
have already described (figs. 37 and 38, nep. p.). In more ad-
vanced larvæ, the pit is split throughout its entire length into
two nearly parallel canals, each of which opens independently to
the exterior. Figs. 39a-c show three transverse, though not
consecutive, sections passing through the posterior region of a larva
at such a stage. In the first of these figures, the two cell-masses
(nep. e.) on either side of the stomach represent the uppermost
portion of the nephridial canals. In the second figure, each of the
cell-masses encloses an easily distinguishable lumen. The two
“nals finally open to the exterior each by a small pore (nep. o.),
as seen in the third figure (only one pore is cut through in the
above figure, the section being slightly oblique to the main axis
of the larval body). In the above figure we see an ectoblastic
cell-mass separating the right and the left nephridial canals (nep. e).
How is this partition brought about? I think it is caused by re-
evagination of the distal unpaired portion of the nephridial pit, as
by that process the pit wall forming the above portion is gradually
transferred to the body-surface of the larva.
Meanwhile the cesophagus becomes more and more elongated,
while the paired tentacular thickenings bulge out each into two
perceptible prominences. The latter represent the rudimentary state
of two larval tentacles, each of which has internally a cavity con-
tinuous with the postoral body-cavity. Fig. 10 represents a larva
532 I. IKEDA:
with two pairs of as vet very short larval tentacles; this is the
most advanced developmental stage to be met with in the em-
bryonal masses. Fig. 40 is a median sagittal section of such a
highly advanced larva. Here the wsophagus (æs.) and the in-
testine (in£.), which latter now communicates with the exterior by
the small anus (an.), are highly developed, so that the three parts
of the alimentary tract (esophagus, stomach, and intestine), may
be said to be almost complete. The nerve ganglion (fig. 40, gl.)
is well differentiated from the ectoblast of the preoral lobe, pre-
senting itself in section as a round, well marked mass principally
composed of nerve fibres. I have been unable to ascertain whether
a proctodæum is produced at all, and if so, what part of the post-
gut it gives rise to.
The preoral body-cavity is, at this stage of development, still
very incompletely separated from the postoral cavity by a few
mesoblast cells (fig. 43, mes’... The nephridial canals (fig. 41,
nep. c.) are now distinctly separated and removed from each other,
and are found in a cross section to be situated laterally to the
intestine (ent.). One on the right-hand side of the above figure is
cut through at its external opening, while on the other side the
nephridium is represented by a.thick mass of a few ectoblastic
cells. This lateral shifting of the nephridia becomes more and
more pronounced with the advancement of larval development. A
slightly advanced state of the nephridia is shown in fig. 42, where
the nephridial canals (nep. c.) are now seen tolerably long and have
a wall composed ofa single row of cubical cells. It is often observed
that some mesoblast cells connect the canals with the splanchnic
walls (see the above figure). These cells seem to be the first in-
dication of the future collar-trunk septum. Besides, a certain
number of mesenchymatous cells, which later undoubtedly become
ON DEVELOPMENT ETC. OF PHORONIS. 533
the excretory cells of Actinotrocha, is always found attached to
the blind ends of the nephridial canals. CALDWELL says that he
saw the excretory cells aggregated around the apex of each canal
and that they had numerous plasmic processes, giving them a
strong resemblance to the perforated cells known in Zehiurus. It
seems, however, highly probable to me that this strange appear-
ance of the excretory cells is an artefact, since, as I shall point
out later, the same cells in Actinotrocha are certainly not provided
with any such processes.
I have very frequently detected some gigantic mesoblast cells
fluating freely in the postoral body-cavity of larve with one or
two pairs of tentacles (fig. 44, corp.). They are round and
nucleated and contain numerous large yolk-spheres. After repeated
examination I have come to regard them as mother-cells of blood
corpuscles which are found as corpuscle-massess in the collar cavity
of Actinotrocha. This point will again be treated of in detail in
the proper place in the following section.
II. The Structure of Actinotrocha.
a. EXTERNAL APPEARANCE,
It can scarcely be doubted that each species of the Phoro-
nidee has a characteristic form of Actinotrocha peculiar to it. Some
of the previous observers (e. g., Wınsox and MASTERMAN) have
mentioned two distinct types of larve as occurring in the same
locality. Among the larvee which I observed at Misaki, I was
able to distinguish four different types, each of which had a
characteristic form and a more or less definite topographical
nn ort era aan TE à
934 I. IKEDA:
distribution. I will designate these types by the letters 1, B, C,
and D.
Type À (fig. 13). The larve of this type were principally
collected in Aburatsubo and belong in all probability to the species
Phoronis ijimai, which, as I have said, is found in the same
locality. The body is comparatively short and thick, measuring
about 1.-1.5 wm. in total length. The larval tentacles of a full
grown larva never exceed 16 in number.
Type B (fig. 14). This is a larger form than the preceding
(about 2-2.5 mm. in length). The body and the intestinal canal
are long and slender. The full grown larva has about 28 tenta-
cles which are much more slender than those of Type A. Peculiar
to it is the sensory spot (30.) situated just in front of the ganglion
(g/.). The larvæ were found in greatest abundance near Kitsune-
saki, a point at the mouth of the inlet Moroiso.
Type C. (figs. 15a & b). This type is distinguished from
all the others by several characteristic points. In size of body
it is intermediate between Types A and B (usually 1.5 mm. in
length). The body is relatively short and thick. The number
of tentacles, so far as I know, ranges from 16 to 24. A pair of
flask-shaped glands (g/d.) is found one on either side of the
ganglion (gl). A pair of retractor muscles (reé’.) runs longi-
tudinally through the trunk cavity from the tentacular ring to the
apex of the anal cone. Compared with the first two types this is
much rarer.
Tyre D (figs. 12 and 16). This is a rare form of which I
have obtained only seven specimens in all. It is enormously large
in comparison with the others (4-5. mm. in total length and
1. mm. in width). The preoral lobe is disproportionately small,
while the trunk is long and thick. The tentacles are remarkably
ON DEVELOPMENT ETC. OF PHORONIS. 839
numerous, sometimes reaching 48 in number. In a single living
speeimen, the skin of the trunk was of a light orange colour ; the
subdermal eircular muscles were especially well developed in the
trunk but interrupted at four longitudinal clearly marked zones.
The youngest swimming larva I have ever obtained was of
type A. It was already supplied with four pairs of tentacles
which, however, were still short. The body measured about 0.5
mm. in length. The trunk was short and showed a slight
characteristic curvature, the concavity being turned toward the
dorsal side. The thickly ciliated hood was comparatively large ;
the ganglion and the perianal ciliated belt were already well deve-
loped. In the surface view of this larva during life, I was not
able to detect the ventral pouch nor the corpuscle-masses.
At about the stage with five pairs of tentacles, the trunk
becomes elongated and straightend out. The nephridia may then
be seen in their characteristic bouquet-form, and the ventral
pouch appears as a solid ectoblastic thickening. Neither the
corpuscle-masses nor the retractor muscles are yet to be seen.
As the larva grows, the number of tentacles increases in pairs
proceeding from the ventral side toward the dorsal; hence, the
most dorsally situated tentacles are the youngest and the shortest.
In larve with 12 tentacles and belonging to type A, the ventral
pouch is deep enough to be plainly visible from outside. We
always notice from this stage on a pair of the retractor muscles
which extends between the ganglion (g/.) and the dorsal inner side
of the tentacular circle (rel. in figs. 12, 13, 14, and 15).
The larval organisation of types A and B is nearly com-
pleted in the stages with 14-16 tentacles. Let me next give a
somewhat detailed description of the external appearance of Actino-
trocha in general.
536 I. IKEDA:
The preoral lobe. This is a structure which looks like a
broad hood with its concavity directed downwards. It almost
entirely covers the upper anterior part of the collar,* when not
influenced by external circumstances. MASTERMAN has made the
statement that in its natural attitude the hood has its length
disposed parallel to the principal body-axis. However, if the
larva be examined in the living state, it will at once be discovered
that its normal disposition is horizontal. It becomes turned up
only as the result of preservation. Its whole surface is covered
with cilia, most strongly developed along the free margin which
constitutes the preoral ciliated belt. In the full grown larva, the
vanglion (and also the sensory spot in type B) has also a set of
specially long cilia on the outside. Numerous fine and refractive
nerve fibres are seen radiating from the ganglion (g/.) to the free
margin of the lobe (pre. bel.) (figs. 13 and 14).
MASTERMAN has described and figured two ectoblastic struct-
ures which are said to be situated on the ventral wall of the hood
and which he has named the “oral” and the “ pharyngeal ”
grooves. These he compares, as to their function, to the gill-
slits of the Chordata. I can not but think that that writer has
here fallen into a very grave error, which might have been
avoided, had he examined the structures in question in living
specimens. Among the preserved specimens I have frequently
noticed those in which the lower or oral wall of the hood was
prominently bulged out in front of the mouth. In consequence
of that prominence (fig. 16, prom.), there was produced on either
* ] adopt this name of Masterman’s to denote that portion of the larval body which Lie
in front of the tentacular circle and behind the preoral lobe.
ON DEVELOPMENT ETC. OF PHORONIS. 537
side of the mouth a transyerse groove, which was visible when
viewed from the ventral side. I believe it was to grooves of this
kind that MASTERMAN assigned the above important significance.
In my opinion, they are simply artificial productions due to preser-
vation.
The Collar. The form of the collar as a whole may be
compared to a eylinder obliquely truncated at the posterior end.
Its posterior border is fringed with a regular row of tentacles,
while anteriorly it is joined to the hood by a narrow neck. The
number of tentacles (larval) varies according to the different. stages
of growth and also according to the type to which larve belong.
They are most numerous in type D, most individuals of which
bear 40-48 tentacles (figs. 12 and 16). The rudiments of the
adult tentacles make their appearance as bud-like eetoblastie
thickenings immediately below the base of the larval tentacles.
An exception to this rule is found in the case of larve belonging
to type D, in which the adult tentacles are represented by a local
ventral thickening of the wall of the larval tentacles at their
proximal portion (see fig. 58d, s. &.). It is very probable that
the number of the larval tentacles corresponds to that of
the adult. In type A, at any rate, I have ascertained that the
full grown larva and the worm just metamorphosed bear the same
number of tentacles, namely 16.
The trunk. This portion, which is the shortest of the three
regions in early larval stages (fig. 10), comes with growth to
occupy the largest part of the larval body and assumes a long
evlindrical form. Its anterior boundary is the tentacular circle ;
the posterior end is girdled with the perianal ciliated belt which
serves as the larval locomotory organ.
538 I. IKEDA:
6. Tue INTERNAL STRUCTURE OF ACTINOTROCHA.
1. Body-Divisions and Body-Cavities.
I have endeavored to show in the preceding pages, that the
hody-cavities of Actinotrocha do not arise from the enterie diver-
ticula, as was insisted upon by CALDWELL, but that they are
simply produced by mesoblastic cells applying themselves to and
forming the lining of the ectoblastic, and the entoblastie, wall.
They may, therefore, be classed under the “pseudoccele ” or
“schizocele ” of Hertwic. Moreover, the body-cavities of Acti-
notrocha as a whole do not in their genetic relation correspond to
those of the adult, as I shall attempt to clucidate in the sequel.
During the metamorphosis, the greater part of the former (the
preoral cavity) is almost wholly lost, while the other part (the
collar-cavity) is transformed into a vascular space, so that what is
known by the same name in the adult is of an entirely new origin.
Thus we see that the larval body-cavity of Actinotrocha, 7. e. the
trunk cavity, is the only portion that persists among the body-
cavities of the adult, in which it is known as the foot or infraseptal
cavity. In correlation with this circumstance are observable cer-
tain changes in the position of the nephridia and of the vascular
system. As described by CALDWELL, the nephridia of Actinotrocha,
which are not provided with an internal opening, lie for the most
part in the collar cavity, while after the metamorphosis they are
found wholly in the infraseptal cavity of the worm. Moreover,
the paired corpuscle masses which are found only in the collar
cavity of the larva, are no longer seen in the same cavity of the
adult. These changes to a eertain extent at least establish the
ON DEVELOPMENT ETC. OF PHORONIS. 889
fact that some profound changes in the arrangement of the bodv-
eavities must occur during the metamorphosis. As is ac-
knowledged by all, the supraseptal cavity of Phoronis is greatly
reduced in size as compared with that of the larva, and contains
almost no organ except the blood vessels. The infraseptal cavity
is, on the contrary, very wide, and contains many important
organs, €. g., the alimentary canal, the sexual organs, and the
main part of the vascular system. Thus it becomes necessary to
make distinctions between the body-cavities of Actinotrocha and
those of the adult and to call them respectively by different names.
The former may be termed the larval body-cavities, and the latter,
the adult body-cavities.
Most previous writers have not taken anv particular notice of
the relation which exists between the external body-divisions and
the body-cavities of Actinotrocha, so the words “ hood ” and “ foot ”
do not denote anything but mere external features. The idea of
segment was first introduced by CALDWELL; he considers the larval
body as divided into three parts: (1) the preoral lobe set in front
of the septum, (2) the trunk portion situated behind the septum,
and (3) the foot or invaginated pouch. According to this view,
the body-cavity is divided by the septum into two contiguous parts,
viz., the preoral cavity in front of, and the trunk cavity behind,
the septum. MasrERMAN divides the entire body into three
portions, viz., the preoral lobe, the collar, and the trunk. These
three divisions are not only externally marked by their respective
forms, but also by the presence of two transverse septa or mesen-
teries. Thus we see, the preoral lobe of CALDWELL comprises ‘both
the preoral lobe and the collar of MAsTERMAN.
Whatever may be the value of Masterman’s Diplochorda
hypothesis, I feel inclined to accept with some modifications, his
940 I. IKEDA:
view of the body-divisions. The external appearance of the three
portions I have already described in brief. As to the internal body-
cavities corresponding to these external portions, I can not agree
with MASTERMAN, when he says that they are completely separated
from one another ; for, as I shall soon show, the septum which lies
between the preoral and the collar cavities is always an incom-
plete formation, at least in all the Actinotroche which I have
observed. Besides, I have been unable to detect the first and third
pairs of nephridia, which are said to exist in the preoral, and the
trunk cavities (MASTERMAN). Therefore, I can not regard the body-
divisions of Actinotrocha as “segments ”” in the sense of that author.
The septa or mesenteries are very delicate in structure and
can hardly be recognized in living specimens. I have, therefore,
had to study them mostly in sections. I shall hereafter call the
two septa the preoral, and the postoral, septa.
Larval Preoral Body-Cavity. The larval preoral body-cavity
fills up the interior of the hood, in which there is no entoblastic
organ. Innumerable mesenchymatous fibres traverse the cavity
(figs. 45, 49, 63 a, m.f.). A few blood corpuseles are also
frequently discovered in this cavity (fig. 49, corp.) ; this fact is,
I believe, one of the proofs of the correctness of the view which I
now propose to consider.
I have already spoken of the incomplete formation of the preoral
septum. Whether this is a mere specific difference or not, remains
to me uncertain, as I have had no chance of examining the larve
investigated by MaASTERMAN. In sagittal sections of the larva at
any stage of its growth, the septum can constantly be traced so
long as the œsophagus is contained in the sections. In fige 45
and 63 a, a slender cellular strand (mes’.) behind the ganglion (g/.)
represents the septum in cross section. Jt extends between the
ON DEVELOPMENT ETC. OF PHORONIS. 541
upper and the lower walls of the hood. Thus it will be seen that
the septum completely separates the preoral cavity from the collar
cavity just behind the ganglion. But, when we come to sections
passing through a more lateral region to either side of the gan-
glion or of the esophagus, the upper portion of the septum becomes
abruptly indistinct. In fig. 54, which shows a sagittal section
through the right-hand side of the œsophagus of a larva of 16
tentacles (type A), the septum (mes’.) near its ventral attachment
is indicated by a comparatively thick layer of cells, while the
dorsal portion is divided: into fine protoplasmic branches, of which
some extend to the upper wall of the hood and others stop short
ofit. As the relations of this septum are somewhat complicated,
I will try to make them clear by referring to a series of cross
sections (not continuous) through the hood of a larva of type D
(figs. 59 a-d) and also to the annexed wood-cut. The latter isa
diagrammatic representation of the Acfinotrocha hood and its
neighbouring part, as seen from above, 7. ¢., in horizontal projection.
The dorsal side is above and the ventral, below. Nearly in
the centre is the nerve ganglion. Below it and concealed from
sight, is the mouth, from which the esophagus leads downwards.
The little stellate markings, scattered over the greater part of the
figure, are supposed to represent mesenchymatous cells, which, with
the branched and reticulate fibres arising from them, pervade
the preoral body-cavity, except in a small space immediately in
front of, and below, the ganglion. This free space I shall call the
posterior recess of the preoral cavity. The line abcdefgh,
curved somewhat like the letter M, indicates the position of the
preoral septum. The part shown in full line represents that por-
tion of the septum which is complete in structure and the part in
broken line, that portion of the same which is incomplete. All
42 I. IKEDA!
the space in front of the septum is the preoral cavity, while back
of it lies the collar cavity.
Denser accumulation of fibres
in front of the recess. Preoral cavity
l'rvoral belt,
l’osterior recess of
the preoral cavity.
Filerows Nerve ganglion.
Tui ly Thc =
Il
11 | .” —ı
IV St —E 3 —}]1
Preoral :
nner.
{hs plans,
Retractor muscle
Collar cay ur,
Tentacles.
=
mes of sections (Bes 50 af”), the
pi me clear. Section 59a is the most
ss through the hood at about the
tabove wood-cut, The whole of the
à the fibres of branching mesenchymat-
wentral median part (p.r.). This is the
study
+ +
u
w recess of the preoral cayity, which is
drancous layer consisting of pre
4 is from about the plane of
ganglion (gl) and on each side
1
Uses bare andergone comsiderable disturtame
‘the relation: of the isyer remain umalierel.
Digitized by Goo
ON DEVELOPMENT ETC. OF PHORONIS. 548
the anterior end of the collar cavity (col.c.). Below the ganglion,
the posterior recess (p.r.) is seen to have a complete wall, that is
to say, the posterior septum is fully developed. In the above
figure, the two wide spaces lying on both sides of the posterior
recess correspond to the two anterior horns of the collar cavity
projecting forward (marked 5 and g in the wood-cut). And we
can certainly see that on the dorsal side as well as laterally there
is no distinct partition or continuation of the preoral septum
which, according to MasTERMAN, should entirely divide the preoral
and the collar cavities at every point. As the figure shows, the
dorsal portion of the mesentery (mes’) is decomposed into fine
protoplasmic processes which join with those of the fibrous mesen-
chymes dispersed through the preoral cavity. In section, 59 c,
passing though the middle of the ganglion (the line II-III
of the wood-cut), the collar cavities (col.c.) are much wider and
have become united below the cesophagus (es). The septum (as
the wall of the posterior recess, p.r.) in this region is a little more
definite in form than in the last figure ; the posterior recess (p.r.)
is distinct as before. In section 59d passing through the line
IV-IV of the wood-cut, the posterior wall of the recess (p.r.) is
obliquely cut and appears in the right-hand lower corner as a
membraneous slice, the recess being distinctly bounded by the
septum (mes’.). Outside of it are seen, one on each side, the
sections of the retractor muscles (rei.), of which more will be
said later. The collar cavity (col.c.) is now very spacious, but the
septum laterally remains in the same condition as before.
From the above descriptions, it will be clear that the preoral
septum is complete only in the median portion (indicated by the
full line cde f in the wood-cut), while in the more lateral part
on each side, it is at the best a loose open reticular membrane,
544 1. IKEDA:
through which the cwlomic fluid of the preoral and the collar
cavities 1s put in free circulation.
A questionable structure has been described from the preoral
?
cavity by MASTERMAN under the name “ subneural sinus,” and is
compared to the structure bearing the same name in the Hemi-
chorda. According to him, the subneural sinus is an interstitial
space left between the two laminæ composing the preoral septum,
just under the ganglion and above the so-called “‘ subneural glaud.”
Anteriorly and laterally, it is said to be surrounded by the preoral
cavity, and posteriorly, by the collar-cavity ; its upper and lower
walls are claimed to be directly formed of the ectoblast without
a peritoneal layer. Further it is said, that the sinus communicates
mid-dorsally with the dorsal blood vessel on the œsophagus.
After repeated examinations of the larvæ of the four different types,
I am convinced that Masrerman’s subneural sinus is identical
with what I have called the posterior recess of the preoral cavity.
It has nothing to do with the tissue-space in the preoral septum,
but is clearly a part of the preoral body-cavity, which is free
from the mesenchymatous fibres. Besides, I can not in any way
detect the presence of the dorsal vessel on the œsophagus, a vessel
which connects the subneural sinus with the dorsal vessel on the
stomach. <A view similar to mine as above expressed was given
by HARMER in his paper on Cephalodisus ('97).
MASTERMAN has further given an interesting description of the
situated on each side of the ganglion. They
“ proboscis pores,’
are compared to the proboscis pores of Balanoglossus and are
said to fulfill the same function as the collar nephridium of
Actinotrocha. In the larvæ studied by me, the only things that
bear even a remote resemblance to them, are the flask-shaped glands
which are seen on the upper face of the preoral lobe of the larva
ON DEVELOPMENT ETC. OF PHORONIS. 545
belonging to type C. But the position of these glands in relation
to the ganglion as well as their histological structure at once
reveal their true nature. The internal openings of the organs
were described by MASTERMAN as follows: “Just where the
preoral mesoblastic wall slopes away on either side of the sinus
there are a pair of thickenings, which traced forwards, show
themselves to be the commencement of a pair of internal openings ”’
(l.c., p. 307). The paired thickenings referred to by him are
apperently nothing else than the points of attachment of the
retractor muscles in the collar cavity, as will be seen in fig. 59d
(ret.). Further details respecting these muscles will be given
later.
Larval Collar Cavity. The collar-cavity is a comparatively
wide space extending between the preoral and the postoral septa.
It is produced anteriorly into two horns, embracing between them
the posterior recess of the preoral cavity. It is perfectly separated
by the postoral septum from the trunk cavity. The postoral sep-
tum, or simply the septum, as it is more commonly called, is
stretched obliquely transversely between the splanchnic and the
somatic walls, along a line a little below the tentacular circle (figs.
45, 48, mes.). Its dorsal attachment on the splanchnic layer is, as
represented in fig. 45 (mes.), found at the plane of the junction
of the cesophagus with the stomach, while ventrally the attach-
ment lies much further below. In frontal sections of the larva,
the septum (fig. 48 mes.) is seen on either side of the stomach
and its somatic insertion lies just under the tentacles, so that each
tentacular cavity is continuous with the larval collar cavity (fig. 45).
The adult collar cavity, or the supraseptal cavity, is already
formed in the fully developed larva of every type, as a ring-space
running along the inner side of the tentacular circle and above
546 | I. IKEDA!
the septum (see figs, 58a and d, s.c.c.). This, together with
several other larval organs in the larval collar cavity, had better
be treated at a more suitable place in the sequel.
MASTERMAN has described a dorsal mesentery running along
the mid-dorsal line of the cesophagus, and separating dorsally the
larval collar cavity into two lateral halves. In the Actinotrochae
of all the tvpes observed by me and at every stage of the larval
growth, no such mesentery is present. It is true that the bodv
walls and the cesophageal walls very frequently come close together,
especially in the young larva after preservation, so as to greatly
narrow the collar cavity in this region (figs. 49 and 50a). : But
a mesentery is never to be found. Its absence is quite clear in
the large Actinotrocha belonging to type D, in which the skin
and the cesophagus lie well separated by a considerable space
(figs. 58a and 582).
Trunk cavity. The trunk cavity occupies the interior of the
third body-division—the trunk. It is completely separated by the
postoral septum from the collar cavity, and since the septum is oblique
in position, it extends dorsally nearly to the base of the cesopha-
gus. The ventral mesentery extends along the median ventral line
of the body wall and of the alimentary canal, and is wholly con-
fined to the trunk cavity. In fig. 45, which shows a median
sagittal section of a young larva of type A, a portion of this
mesentery (v.mes.) is represented as a thin cellular membrane
extending between the alimentary canal and the ventral pouch
(p.o.), the latter being still shallow at this stage. The whole
extent of the ventral pouch is stretched by the ventral mesentery
to the skin as well as to the digestive canal. This relation re-
mains the same as the pouch grows in length and finally winds
around the digestive canal. A transverse section through the
ON DEVELOPMENT ETC. OF PHORONIS. 547
trunk of a highly advanced larva of type C, is given in fie. 57 a,
in which the much elongated and convoluted pouch is seen cut
into several sections (po.), connected with one another by the
mesentery (?.mes.).
Very frequently it happens that the peritoneal mesoblastic
epithelium, which lines the perianal ciliated belt, is detached from
the ectoblastic wall. This is a purely artificial appearance caused
by the killing reagent. It seems probable that Masterman has
erroneously considered the space thus formed by splitting to be a
vascular space (the “ perianal sinus”). The same author states,
though with much reserve, that he has discovered a third pair of
nephridia in this trunk cavity, which is considered to be a modi-
fied part of the body-cavity, and also to be rudiments of the adult
nephridia. I can at present say no more than that these are cer-
tainly absent in every type of the Actinotrocha studied by myself.
2. Organs of Ectoblastic Origin.
The epidermis of Actinofrocha is represented by a single
laver of cubical or cylindrical cells, those of the collar wall and
of the upper and the lower walls of the hood being provided with
well developed cilia. Besides, there are three specially ciliated
regions: the preoral belt, the tentacles, and the perianal belt. The
last is the larval locomotory organ; on it the cilia are very long,
thick, and somewhat bristle-like when in active motion. At places,
where cilia are strongly developed, (e.g., the nerve ganglion, the
sensory spot if present, the ciliated belts, efc.) the constituent cells
are cylindrica], the nucleus generally lving near the basal end.
The body wall of the trunk region is very thin and is formed of
greatly attenuated cells (especially slender in the advanced larvæ).
548 I. IKEDA!
Numerous unicellular glands are found in the Actinotrocha not
only all over the two surfaces of the preoral lobe, but also in the
cesophageal wall as well as in the inner ectoblastic wall of the
ventral pouch. They are also, though less abundantly, distributed
over both the collar wall and the tentacular wall. The glandular
cells are all pear-shaped, the nucleus being found always appressed
to the base of the cell (figs. 49 and 64d, mgl.). In their
staining reactions, the secretory contents of the glands agree with
those of mucin. It has been often noticed that living larve
remain adhereing to the objects they have touched with the hood,
and that metamorphosed larvee behave similarly with the tip of
the evaginated pouch.
There exists still another, paired, multicellular gland which
is observed only in the larvæ of type C' (figs. 15, gid. and fig. 15 c).
It is situated on both sides of the median line on the upper sur-
face, and somewhat near, the neck of the preoral lobe. It has the
shape of a round flask with a short neck (fig. 15 c). The appear-
ance of the section through the body of this gland reminds us of
the chorda dorsalis in Vertebrate embryos: it presents to view a
mesh-work of protoplasm, a small number of nuclei being found
here and there closely pressed against the reticular beams or the
nodes of these (figs. 56 a-c).* Each of the meshes corresponds to
one gland cell. In fig. 566, which shows an oblique median
section of the body of the gland, a comparatively wide round
space exists in the centre, surrounded by the gland cells which
are arranged more or less radially. This space, when traced up-
wards, passes into a short and very narrow tubular canal, finally
to lead to the exterior by a small aperture (Fig. 56c). Since
* By an unfortunate oversight, Fig. 56 b has had its number omitted in the plate,
ON DEVELOPMENT ETC. OF PHORONIS. 549
the neck portion of the gland is very short, it is difficult to
prepare a good longitudinal section of it, in which the canal may
be seen opening to the exterior. Fig. 56 c represents the terminal
part of the emptying canal, which, as can be ascertained by regu-
lating the focus, leads to the external pore. Mr. Iızuka tells
me that similar glands of ectoblastic origin are constantly found
on the superior ramus of a parapodium in certain Polychæta.
Ventral Pouch. As the ventral pouch is one of the most
characteristic structures of Actinotrocha, its form and fate have
been fully studied by many previous observers. In the 8-armed
larva of type A, an ectodermal thickening below the tentacular
row represents the origin of the pouch. At the 10-armed stage
the thickening becomes more conspicuous, but no invagination has
as yet taken place. For the first time in the 12-armed stage, the
wall at the thickening begins to sink inwards and backwards
(fig. 45, po.). The invagination is lined with a mesoblastic layer,
and, as before noted, is for its whole length suspended by the
ventral mesentery, joining it to the somatic and the splanchnic
walls. As the growth of the larva advances, the pouch becomes
more elongated and bends on itself around the alimentary canal
)figs. 48 and 57 a, po.). In fully developed larvæ of whatever type,
the inner or ectoblastic wall is thrown into small wavy folds
(beginning at the distal portion near the pouch pore), while the
mesoblastic layer becomes muscular, so that at the end of larval
life, it forms a thick muscular sheath whose constituent cells stand
vertically to the inner wall. As to the form and position of the
pouch pore, I can offer no details in addition to what hay been
observed by METSCHNIKOFF and many other authorities.
Nervous System. The nervous system of Actinotrocha, like
that of Phoronis, is of a very low development, being represented
550 ï. IKEDA:
merely by a local differentiation of the ectoblastic cells into ner-
vous elements. The epidermis over both the ganglion (fig. 14. gl.)
and the sensory spot (so.) is strongly ciliated, so that the organs
are easily recognizable in the living larva. The earliest stage
in which I found the ganglion was a 4-armed larva of type A
(fig. 40 gl.). In it, the ganglion consisted of only a few ganglion
cells and nerve fibres.
Although the ganglion and some nerves directly proceeding
from it can be detected with tolerable distinctness in the living
specimen on account of their peculiar refractivity, the peripheral
nerves are as a general rule so very fine and delicate, that they
can not be satisfactorily made out by means of any ordinary
process. With fair success I have had recourse to vital staining
with ınethyl-blue. Larve of type B have been principally em-
ployed for this purpose. They are left for about 15-20 minutes
in a weak solution of methyl-blue in sea water and immediately
afterwards treated with ammonium molybdate. Sometimes, I have
made supplementary observations on larve lying alive in the
methyl-blue solution under the cover glass, but this can be con-
tinued for only a short time, since a general overstaining of other
tissues soon takes place.
Fig. 60 a* shows the dorsal view of the anterior half of a
larva of type B, which was treated in the above way. ‘The nerves
are shown in blue. The results obtained as to their distribution
differ in many important points from those obtained by Masrer-
MAN. Whether this difference is due to the technique or is
actually existant in the species studied, is difficult to ascertain.
nn — ~—
* As the larva shown in this figure was compressed by the cover-glass, the rim of the hood
which appears like its free margin is in reality the line along which the hood was bent and
reflected over by pressure. The line drawn close to the peripheral dots in blue represents the
true edge of the hood,
ON DEVELOPMENT ETC. OF PHORONIS. dol
MASTERMAN says nothing of the method emploved in his inves-
tigation, and unfortunately there exists no other study than his
with which to compare my results.
As may be gathered from the above-mentioned figure, I can
discover no collar nerve ring, nor dorsal or ventral commissure.
Besides, in spite of repeated efforts, I have always failed to make
out, the presence of the so-called perianal nerve ring. The collar
ring and the dorsal commisure, if they can be so named, are repre-
sented by a small number of parallel fibres, which spring directly
out from the posterior corner of the nerve ganglion. In every
case examined, they could be traced no further than a short dis-
tance from the ganglion. Sometimes, I have been able to discern
in sections the main nerves (commonly 3 in number), which run
close together and parallel to one another along the mid-dorsal
line of the trunk, but they were confined to only a few sections
posterior to the first pair of tentacles. On the other hand, a
very complex and beautiful system of nerve fibres could be seen
on the preoral lobe. The fibres are here exceedingly numerous
und fine, radiating from the ganglion on all sides towards the
free margin of the preoral lobe. In the median line and
anteriorly to the ganglion (gl.), the fibres appear as three longi-
tudinal parallel strands on which the unpaired sensory spot (so.)
is situated not far from the ganglion. After passing through the
sensory spot the strands fray out into fine fibres which continue
their course towards the free margin of the preoral lobe. The
fibres emanating from the ganglion do not all show a regular
radial arrangement, but there are some that arising from the
lateral edge of the ganglion, soon take an anteriorly directed course.
Sometimes there were not wanting, especially near the ganglion,
indications of anastomosis between the fibres. However, it seemed
552 - 4. IKEDA!
to me more probable that these appearances were caused simply
by the juxtaposition of intersecting fibres.
The nerve endings in the preoral ciliated belt deserve special
notice. In fig. 60a, there is shown a row of small dots along
the margin of the band. A portion of the latter more highly
magnified is shown in fig. 605. Here each fibre ends in a small
knob which is devoid of any lateral process. At first sight under
low magnification, the row of knobs appears like a deeply stained
ring, Suspecting that there might exist lateral processes connect-
ing knobs, I have repeatedly made observations and experiments,
but without having ever been able to demonstrate such a connec-
tion between them.
I can not but think it very strange that post-ganglional nerve
fibres, if such really exist in the forms of the collar ring and of
the dorsal and the ventral commissures, should not be revealed by
the method adopted. The negative result may be considered due
to incomplete development of nervous elements in the collar and
in the trunk region; but other anatomical relations prove to a
certainty that the larve investigated were fully grown. As I am
not quite sure that my method was not in some respect imperfect,
I leave the matter undecided for the present.
According to MASTERMAN, there is an ectodermal depression
directed inwards and backwards, just in front of, and under, the
ganglion. He calls it the “neuropore,’’ comparing it to the
neuropore of Amphiorus and even to the medullary canal of
Vertebrates. I must say I was much disappointed in failing to
detect in the Actinotroche studied by me this structure of so much
theoretical interest. As a matter of fact, it happened very fre-
quently, while observing living larve, that the ganglion was
retracted deeply inwards by an active contraction of the two
ON DEVELOPMENT ETC. OF PHORONIS. 553
retractor muscles in the collar cavity (figs. 13, 14, 15, ret.) pro-
ducing at the same time a deep depression just in front of the
ganglion. It is also of almost constant occurrence that the gan-
glion is withdrawn inwards on the application of reagents, so as
to produce a shaflow pit or groove in front of, or below, the
ganglion (figs. 63 a, gl). A quite similar fact is always observed
in Lov£n’s larva. From these circumstances I am much inclined
to regard the “neuropore” of MASTERMAN not as a really exist-
ing structure, but as an artefact.
As to the tentacles, I have at present nothing to add to what
is already known about them.
3. Organs of Entoblastie Origin.
In the fully grown larvæ the alimentary canal is a long and
straight tube; it begins with the mouth which is overhung by the
preoral lobe, and ends at the anus in the centre of the anal cone
surrounded by the perianal belt (figs. 12-16). Of the whole ali-
mentary tract three parts may be distinguished: the Oesophagus,
the Stomach, and the Intestine.
Oesophagus. In the embryological part of this article I have
said that the esophagus of Actinotrocha is of ectoblastic origin,
so that the original gastrula mouth is to be sought at the junc-
ture of the cesophagus with the stomach. The csophagus (Figs.
45, 48, and 49, ws.) is a comparatively short and narrow canal
with a wall composed of densely ciliated cylindrical cells, among
which are scattered numerous unicellular glands (m.gl.). Thus
the wall does not differ in structure from that of the hood or of
the collar.
MASTERMAN has described an unpaired ectodermal invagina-
554 I. IKEDA:
tion situated in front of the mouth and just under the ganglion.
It is called the “subneural gland.” Here again I am not in a
position to confirm his view. In spite of repeated examinations
on living specimens, I have been unable to discover any structure
which has the slightest resemblance to the subneural gland. To
judge from my own observation, the “subneural gland” as well
93
as both the “ oral-” and the “ pharyngeal grooves’ of this author
are products of his fixing method. In preserved specimens, it is
frequently noticed that the lower wall of the hood is bulged out
and downwards in front of the mouth (fig. 16, prom.), and, as a
result of this, there is brought about on the wall behind the
prominence a depression, which appears on sections as a tolerably
deep pit (fig. 63 a). |
Stomach. The stomach forms the largest and widest portion
of the alimentary canal. It is especially long in the larvæ of type
D, in which it extends below nearly to the plane of the perianal
belt. The greater part of the stomach wall is composed of
cylindrical cells with short cilia whose spherical nucleus is usually
situated in the centre of the cell (figs. 45, 48, and 50a, stom.).
But the anterior portion of the wall along the mid-dorsal line and
the posterior portion near the intestine greatly differ in their
constituent cells from the remaining parts. They consist exclu-
sively of tall ciliated cells which contain clongated nuclei, and
are, in a word, of the ssophageal type (fig. 45). In the full
grown larva, the ventro-lateral portions of the stomach wall form
two digestive areas placed in the neighbourhood of the septum.
Here the cell boundaries are indistinct and the nuclei are im-
bedded in a common mass of protoplasm, in which remains of
various unicellular organisms are enclosed.
From the anterior end of the stomach a pretty wide and
ON DEVELOPMENT ETC. OF PHORONIS. 555
unpaired diverticulum protrudes itself forwards (fig. 14, div.). The
position of the organ is wholly ventral to the csophagus (es.),
and the form is like that of a sac compressed in the dorso-ventral
direction (figs. 45, 49, 50 a, and 63 a, div.). The internal cavity
is continuous with the stomach cavity. The roof of the diverticulum
in the fresh state generally shows a reddish brown tint. This
coloration is due to the superposition of the fundamental brown-
ish colour on the hemoglobin of the blood corpuscles which, in
advanced larvæ, overlie the organ in either one, or two masses.
The cells which compose the diverticular wall are tall and slightly
curved, and are ciliated on the free ends (fig. 50 a, div.) In fully
grown larvæ of every type, each cell constantly contains a single
small round vacuole in its distal end (fig. 61). The vacuoles can
not be stained by most of the staining reagents. I have scen
them in the diverticular wall of a highly advanced larva belonging
to type A, which had already evaginated the ventral pouch ; even
in this case, they were found only one in each cell (fig. 61 6).
The whole of the diverticulum is lined externally with the thin
peritoneal layer (sce the above figure).
Many previous observers have noticed this organ and have
called it by various names :—
J. Muzrer ('46)—“ Blinddarme ”’ (paired),
(TEGENBAUR (’54)—“ Haufen der Leberzellen,”’
WAGENER (’47)—“ Leberblinddarme,”’
CLAPAREDE (’63)—‘ A dark mass with globules ”’ (after
MASTERMAN),
METSCHNIKOFF (’71)—“ brown specks,”
Wirtson (A.G.) ('81)—*“ glandular lobes of the stomach,”
MASTERMAN (’97)—“ Notochord ”’ (paired),
Roue (’98)—“ Notochord ” (unpaired).
556 I. IKEDA:
Thus it will be seen that while some authors have apparently
confounded the organ with the overlying corpuscle masses, others
have considered it to be a glandular appendage of the stomach,
and still others have regarded it as a skeletal structure. Accord-
ing to MASTERMAN, who maintains the last mentioned opinion, the
stomach wall is produced, in the antero-lateral region, “ into two
remarkable diverticula which in the fully developed larva he as
a pair of elongated organs, Notochords, laterally to the œsophagus ”
(97, Le., p. 302). The organs are said to soon undergo a remark-
able metamorphosis, 2.e., vacuolization. The vacuoles are produced
successively one after another at the distal ends of the cells and
are arranged alternately in several layers. On account of these
facts MASTERMAN rejects the view that the organ is of a glandular
nature, and holds that it is to be compared in function and structure
with the notochord of the Chordata. In 1898 Rouze published
his third paper on Actinotrocha, in which he denied that the organ
is double in number and lateral in position to the cesophagus,
but admitted the vacuolization in the larva of Phoronis sabateri
(=P. psammophila Cort).
I can not at present decide whether the variations in the
number of the diverticulum and in the degree of vacuolization are
of specific value or not. For the present I must be content with
simply noting that the stomach diverticulum in the larvæ studied
by me is constantly unpaired and undergoes no farther vacuolization
process than the production of one vacuole in each cell.
Intestine. The intestine which leads to the anus is a slender
canal whose wall is composed of a layer of somewhat cylindrical,
ciliated cells with round nuclei (figs. 45 and 48, zné.).
er
ON DEVELOPMENT ETC. OF PHORONIS.
4. Organs of Mesoblastic Oriyin.
As the mesoblastie organs have been but little studied in their
development, so their structure and fate after metamorphosis are
very imperfectly known. Although I have endeavoured to make my
study of the organs as exhaustive as possible, some important
questions remain yet unsolved. The principal organs to be des-
cribed in this place are the muscular elements, the vascular system,
and the nephridia.
Nephridia. I will treat these under the mesoblastic organs,
for, though the nephridial canals are of ectoblastic origin, the
organs as a whole bear intimate relations to the mesoblast. Most
of the earlier observers overlooked the presence of nephridia in
the larva. The first discoverer was WAGENER (’48), whose descrip-
tion is, however, very meagre and gives us no exact idea of the
organ. CALDWELL in his preliminary note (’82-’83) has given a
detailed description of the nephridia. According to his view, the
nephridial canal at no time during larval life, opens into the body-
cavity.
MASTERMAN (’97) has described the excretory system of the
larva in detail and has suggested an hypothesis which seems to
me to be an extraordinary one. Each of the three “ segments ”
of the larval body, he concludes, is provided with a paired organ
which performs the excretory function. The three pairs of organs
are called respectively the “ proboscis pores,” the “collar nephridia,”’
and the “trunk nephridia,”
Of these, however, the presence of
the first and the third is, as I have before pointed out, very
doubtful. The second pair, or the collar nephridia, are the organs
which I consider to be the nephridia. MASTERMAN’S views on the
858 I. IKEDA:
structure of the nephridial canals are in the main similar to those
of CALDWELL, except in one important point, viz., that the canals
are said to open by means of funnels into the collar cavity.
When a larva of any type is examined in the living state,
the proximal ends of the organs are seen, as described by WAGENER,
as two bonquet-shaped masses which are formed by a crowding
together of the excretory cells (fig. 13, neph.). They are placed
symmetrically one on each side of the stomach and in front of
the postoral septum. Each of them consists of two parts, the
nephridial canal and the exeretory cells. The former is com-
posed of a layer of cubical cells, and contains a narrow lumen
which ends blindly at the internal end and distally leads to
the nephridial pore lying on either side of the pouch pore.
The greater part of the nephridial canal, together with the excretory
cells, rests on the upper surface of the postoral septum. Fig. 50 à
shows a cross section of the larva, passing through the left neph-
ridial canal near its internal blind end, where the excretory cells
adhere. In the above figure, a small cell mass (nep.e.) on the right
of the figure, shows the cut end of the nephridial canal which is
attached to the septum (mes.). In the figure, the left canal (nep.c)
containing a small lumen, is found applied to the somatic walls. If
traced a little downwards, these two canals become attached to and
imbedded between the two layers of the somatic walls, and are no
more to be seen in the collar cavity. Such a state is represented
in fig. 50 c, in which the two canals (nep.c.) are wholly imbedded
in the somatic walls on both sides of the stomach (stm.). This
condition is more distinctly shown in figs. 47 (a-c), which are
taken from serial longitudinal sections of a larva of type À with
12 tentacles. These figures show only one portion of the skin,
Where the nephridial canal and the somatic attachment of the
ON DEVELOPMENT ELC. OF PHORONIS. 559
postoral septum (mes.) are situated. In fig. 47 a, one portion of
the peritoneal layer of the stomach wall is also represented. Now
we see in the first two figures of the above series, that the nephri-
dial canal (nep.c.) which is here imbedded in the somatic layers,
lies distinctly below the septum (mes.). So, in the third figure
the nephridial pore (nep.o) is seen as a small pit in the trunk
wall, which is situated considerably below the septum. The in-
fraseptal position of the nephridial pores has also been acknowledged
by Carvwerr. ‘Though Masterman has made no direct state-
ment on this point, it may safely be inferred from his figures,
that he must have regarded the pores as lying in front of the
septum.
Fig. öl « represents a longitudinal section through the middle
of the supraseptal portion of the nephridial canal. Here the canal
appears as a comparatively long tube with a narrow lumen; it is
invested throughout with a thin mesoblastie epithelium. At its
upper extremity where the lumen disappears, a certain number of
spindle-shaped excretory cells is found aggregated together. In
fig. 516, which is taken from the same series as fig. 5la, the
canal has wholly disappeared from the section, leaving only a
bunch of the excretory cells (exc.c.) adhering to the septum (mes.).
All of these spindle-shaped cells have their nuclei in the swollen
ends. I have never found cither among, or in, the neighbourhood
of the cell bunch any perforated excretory cells bearing many
processes, —cells which are said to have been present in the Acéi-
nolrocha studied by CaLDWELL.
MASTERMAN considers that each bunquet of the excretory cells
is composed of a cellular mass traversed by a system of minute
funnels ; and that these funnels communicate with the main canal
of the nephridium as well as with the collar cavity. But I may
560 I. IKEDA:
say with certainty that, at least in the larve studied by me,
there existed no such funnel-system nor any such free communi-
cation between the collar cavity and the nephridial canals. The
same negative result was also reached by me in my examination
of the just metamorphosed larva of type A. Thus in fig. 64/
which is drawn from a section through the tip of the nephridium,
the excretory cells (exc.c.) still remain compactly grouped on the
blind tip of the canal (nep.c.), but are not traversed by any sort
of canal-systems.
Muscular System. The muscular system of Acfinotrocha re-
mains in a low state of development, which may account for the
fact that most previous observers have paid no particular atten-
tion to it. It was therefore of much interest to discover two pairs
of tolerably well developed muscles, which had hitherto remained
apparently unknown. They show a strong resemblance to the
retractor muscles which have been known in many forms of
Trochophora \arve.
Though the longitudinal and circular muscles of the body
wall may be observed with tolerable distinctness in the living larva,
they are usually very poorly preserved after hardening. They are
all subdermal in position and very delicate in structure, so that
as a rule they can not be satisfactorily distinguished from the
underlying peritoneal layer. However, in certain preparations of
the entire larva the circular muscles of the upper and lower
walls of the preoral lobe and of the trunk bodv wall could be
detected as fine deeply stained fibres. In the same way the
longitudinal muscles of the collar wall, especially in the larva
of type D, were fairly traceable. The larva of that type also
exhibited a peculiar arrangement of the circular muscles of the
trunk, in that these formed four, equidistant, loagitudinal series
ON DEVELOPMENT ETC. OF PITORONIS. 561
around the periphery of the trunk (see fig. 12). In the larva
of type € I have always found a comparatively thick layer of
circular muscles. In fig. 576, which shows a portion of the
trunk wall containing the nephridial canal (nep.c.), the muscles
are represented as a thin fibrous layer (cir.m.) intercepted
between the ectoderm and the peritoneal epithelium. The floor
of the mouth just opposite the stomach diverticulum, is always
associated with a particularly well developed muscular sheet. The
mesenchymatous unicellular fibres which traverse the preoral cavity
are to be regarded as a kind of primitive muscles. The most
highly developed parts of the muscular svstem of the somatic, and
of the splanchnic, walls are to be found in the muscular sheaths
of the ventral pouch and of the dorsal wall of the stomach in
the advanced larve of all types. Each of them is formed of
a thick layer of enormously elongated muscular cells which stand
vertically to the ectoblastic, or the entoblastic, wall as the case may
be (figs. 58¢ and 63 e, m.sh.). The sheath of the stomach wall
is thickest along the mid-dorsal line of the stomach ; it is shown
in fig. 8 c and fig. 63e, the former figure being taken from a
cross section and the latter from a longitudinal section through
the dorso-anterior region of the stomach. The muscular sheath
(or the external wall) of the ventral pouch is essentially similar
to that of the stomach.
The Retractor Muscles can be constantly detected in every
type of the larva as two slender threads on both sides of the
cesophagus (figs. 12, 13, 14, 15, ret.). They spring from the hind
lateral corners of the ganglion (g/.) and run divergently downwards
until they insert themselves in the collar walls between the first,
and the second, tentacles. In order to obtain a clear idea of the
position of these muscles, it is necessary to study them in sections ;
562 I. IKEDA:
the larve of types B and D are best suited for this purpose, a
the muscles in these are remarkably large and long. Figs. 63 a-c
are taken from serial sagittal sections through the right side of
the nerve ganglion. In figure a, a median section, we see behind
the ganglion nothing but the preoral septum (mes’.) In the next
figure, d, the septum is found to have shifted to a more anterior
position and its dorsal termination is accompanied by a strong
musele-band (ret.). This band corresponds to that portion of the
retractor muscle which is nearest to its anterior insertion on the
postero-lateral side of the ganglion. The two muscles are shown
in fig. 59d (ret.), a cross section through the posterior recess
(p.r.) of the preoral cavity, where they spring directly from the
septum (mes’.). In fig. 63c, which shows a more lateral region
than fig. 63, the muscle is found to have retreated far backwards,
touching with its posterior portion the cesophageal walls (@s.). The
posterior insertion of the muscles on the somatic walls are best
studied in serial cross sections of the larva. Figs. 58 a and 6 show
two cross sections passing through the mouth (@) and through the
middle of the cesophogus (4). In both figures the muscles (reé.)
are found on both sides of the cesophagus (æs.). A little further
down they soon detach themselves from the œsophagus and begin
to traverse freely the body-cavity (larval collar cavity), and after
that thev again apply themselves to the skin on each side between
the first and second tentacles (¢ and ¢” in fig. 58 6).
There is also present another pair of muscles, which can be
discovered onlv in the larvæ of type C They are so very long
as to equal the entire length of the trunk (fig. 15 5, reé’.). They
arise on each side from the somatic walls just above the nephri-
dial pore and run straight downwards traversing the trunk cavity,
and ending at the terminal portion of the intestine. Fig. 57 a
ON DEVELOPMENT ETC. OF PHORONIS. 363
represents a transverse section through the middle portion of the
trunk where the stomach (sim.) joins the intestine (iné.). There
the muscles appear as two small striated masses (reé.’) lying on
both sides of the intestine. As represented in fig. 57 6, which is
taken from the portion of the body wall containing the nephridial
canal (nep.c.), each of the muscles is in its origin traceable to the
circular muscle layer (eir.m.) which is subdermally interposed bet-
ween the ectoblast and the peritoneal epithelium. When these
two muscles, in their downward course, reach the level of the
terminal portion of the intestine, they fuse together into one on
the dorsal side of the intestine. I think that the above men-
tioned muscles are a kind of retractors serving to contract the
trunk of the larval body. I can not help thinking that the
‘ Afterbänderung ” of WAGENER (’47) is probably simply the post-
erior portion of the muscles in question in the proximity of the anus.
Vascular system. It is a well known fact that the closed
vascular system of Phoronis offers one of the greatest obstacles to
the idea entertained by some naturalists that the animal is of the
Polyzoan type. Many writers are, therefore, much inclined to
attribute the simple body organization of Phoronis to secondary
adaptation, and to erect the animal into a distinct order very closely
related to the Chordata. Putting aside for the present all the-
oretical speculations, it is of great importance in ascertaining the
phylogenetic relation of the animal to note that one portion of the
larval body-cavities is transformed into a blood vessel, and that
the simple and rudimentary vascular system of <Actinotrocha under-
goes a wonderful change and suddenly attains the high organi-
sation seen in the adult during metamorphosis. _
Krons (°50) proved that the “ Leberzellen ” of WAGENER
and GEGENBAUR were reallv blood corpuscles. However, he did
564 I. IKEDA:
not discover any blood vessel in Actinotrocha; he thought that
the blood vessels of the metamorphosed worm arose in the cor-
puscle masses of the larva.
CLAPARÈDE (’63) mentioned a ring-like vascular canal under
the tentacular row of the larva, but did not explain its nature.
SCHNEIDER (’62) discovered two vessels in Actinotrocha, which
ran parallel along the mid-dorsal line of the stomach.
METSCHNIKOFF (’7I) described and figured in a larva of 10
tentacles the “ feinen Häutchen ” situated just above the invagina-
tion pouch, which was said to be the “ Gefiissanlage.’”’ Besides,
it is stated that he saw a ventral “ sinusartigen Schlauch ” which
covered the greater part of the stomach and communicated an-
teriorly with the collar cavity. According to his view, this
Schlauch should give rise to the ring vessel of the adult. But
what are really meant by the “ Schlauch ” and the “ Häutchen ”
is not clear from his text and figures.
Witson (’8I) confirmed the main points of METSCHNIKOFF’S
observations, but disproved the presence of a blood vessel along
the intestine, and also the free communication between the pseudo-
hæmal space and the perisviceral cavitv. According to this author,
there are two sorts of corpuscles: the one kind floats in masses in
the perivisceral cavity, and the other (the pseudohcemal corpuscles)
arise within the cavity of a sinus which is formed in the stomach
walls and form the circular ring vessel of the adult.
CALDWELL (’82-’83) gives us a concise description of the
vascular svstem in Actinotrocha and in its adult form. He says
that the corpuscle masses “arise from the mesoblast cells in front
of the septum,” and that “ The vessels arise as slits in the
splanchnopleure. The adult condition is reached partly by con-
strictions, partly by out-growth from these. Thus we have at the
ON DEVELOPMENT ETC. OF PHORONIS. 565
close of the larval life the blood system in the following condition :
1. Blood corpuscles aggregated in two or more masses, lying
in the body-cavity of the preoral lobe, i. e., in front of the septum.
2. A blood vessel formed on the dorsal wall of the stomach,
a marked structure of the larva. |
3. The splanchnopleure, which in the region of the stomach
forms a loose sac surrounding the gut.
4. Cecal prolongations of this sac.
5. Cecal prolongations into the rudiments of the adult ten-
tacles’’ (’82-’83, l.c., p. 377). Besides, the author insists on the free
communication between the splanchnopleuric sac and the body-
cavity in front of the septum. Thus it may be understood that
CALDWELL detected only one vessel (dorsal) in Actinotrocha and
thought the ring vessel of the adult was produced from the
splanchnopleuric sac around the stomach.
MASTERMAN’S views (°97) of the vascular system differ greatly
from those of all the others above quoted. The subneural sinus
is said to communicate posteriorly by a chink with the dorsal
vessel on the cesophagus. The dorsal vessel runs down till it com-
municates with the ventral vessel at the juncture of the stomach
and the intestine, by means of a small ring siuus. Anteriorly also
the dorsal vessel gives off two branches which, after passing along
the inner side of the two notochords, again meet together in the
mid-ventral line, forming a post oral ring sinus. From that meeting
point originates the ventral vessel which runs down along the
whole length of the gut and opens into a large sinus-ring situated
just within the perianal belt. Further, the author denied the free
communication of the blood vessels with the body-cavity, which
had been maintained by METSCHNIKOFF and CALDWELL.
I have already stated my belief that the mother cells of blood
566 1. IKEDA!
corpuscles appear as the gigantic mesoblast cells in the body-cavity
of the larva with one or two pairs of tentacles (fig. 44, corp.). These
cells may be easily distinguished from other wandering mesoblast
cells in that they have an enormous size and are loaded with an
abundant quantity of large yolk grains. Now in 8-10-armel
larvæ of type 4, not only such peculiar cells but also the so-called
corpuscle masses can not be found in any part of the bodv-
avities. Instead of them the collar cavity and often also the
tentacular cavities contain a few large and isolated mesoblast cells
which closely resemble in size and structure the blood corpuscles
of a highly advanced larva. These cells no longer contain large
yolk grains, but enclose numerous fine, refringent granules. Fig.
46 represents two such corpuscles (corp.) floating in the tentac-
war cavity (é) of a larva of 10 tentacles. It can with pro-
priety, I believe, be admitted, that these corpuscles have arisen
by repeated division from the gigantic mesoblast cells, whose yolk
contents have been gradually used up during the process. If this
be not the case, then how is the presence of those isolated cor-
puscles in the young larvæ to be explained? If, as is imagined
by CALDWELL, they are produced by cell-multiplication taking
place in certain parts of the splanchnopleuric walls and form the
corpuscle masses from the first, why should such freely floating
and isolated corpuscles be actually present in the young larvæ in
which the masses are not yet discernible? So far as I- know, in
the larve of type A, the corpusele masses do not exist until the
animal has so far developed as to possess at least 14 tentacles
(lie. 15, corp.). At the stages of 1-4 and 16 tentacles, these masses
are present commonly in two pairs, the one covering the stomach
diverticulum and the other just in front of the septum and ou
both sides of the stomach (fig. 13, corp.). They appear as pinkish
ON DEVELOPMENT ETC. OF PHORONIS. 567
spheres in fresh specimens. They do not constantly adhere to
the stomach walls as Witson and some others have remarked,
but are very frequently found floating freely in the collar cavity,
showing that there exists no direct connection with the splanch-
nic walls. This fact may be clearly seen in fig. 53 (corp.), where
the mass is located at an appreciable distance from the stomach
walls (sém.}. The same state was also observed in a 14-armed
larva of the same type. Besides, I have noticed at these stages
of growth two sorts of corpuscles in the masses: the one sort 1s
large and somewhat coarsely granular ; the other is much smaller
and finely granular. On sections it was found that the former sort
is imbedded here and there in groups of the latter (see fig. 53,
corp.). I can not exactly see the significance of this fact unless
it be that it shows the developmental process of the blood corpus-
cles, in which the larger ones give rise to the smaller by division
(the karyokinetie figures in the former can be made out with tole-
rable distinctness by staining with cosin methylblue). The larger
cells are essentially identical with the corpuscles of both the
younger (12-armed) and the older larvie (so far as advanced as to
be ready for metamorphosis) of the same type. Thus it seems to
ine probable, though I state this with a certain degree of reserve,
that these smaller corpuscles develop into the normal blood cor-
puscles of the highly advanced larva, for in the latter we no longer
find the smaller forms in the corpuscle masses. Fig. G1 @ represents
four corpuscle cells composing a corpuscle mass of a larva of type
À which has already evaginated the ventral pouch, Of course at
such a stage approaching the end of larval life, the number of the
corpuscles is actually greatly increased as compared with that in
younger stages,
From the above observations it may be concluded that the
568 | I. IKEDA:
blood corpuscles of Actinotrocha do not arise at the expense of the
splanchnic walls, but are produced by a continual division of
certain previously differentiated mesoblast cells.
I will next describe the blood vessels which can be seen
during larval life, with reference to the formation of the adult
collar cavity. I have shown in the foregoing pages that Master-
MAN’S subneural sinus is probably nothing but a posterior recess
of the preoral body-cavity, and that neither the dorsal vessel
nor the dorsal mesentery is present on the œsophagus in any
species of Acfinotrocha I have been able to obtain. I have en-
deavoured to ascertain the presence of the dorsal and the ventral
vessels as well as of the ring-sinuses around the gut, and I am
convinced that at no time during larval life any vessels other than
the dorsal on the stomach and the cecal capillaries are present in
the larve.
In the A type-larva of 14 arms, the dorsal vessel, as figs. 50 0
and 50c will show, is not yet formed and the stomach wall is
uniformly lined with a thin mesoblastic layer. This layer thickens
later and its constituent cells become muscular, beginning first at
the base of the postoral septum and along the mid-dorsal line.
When the larva grows to the stage of 16 tentacles, the dorsal vessel
is Inceptionally formed. It arises as a solid cord of cells inter-
posed between the muscular, and the entoblastie, walls of the
stomach. As shown in fig. 52, the vessel in section is represent-
ed by a loose mass of mesoblast cells distinctly delimited on all
sides from the surrounding parts ; but as yet no lumen is visible
in it. I have not seen the definite lumen establish itself in this
rudiment of the dorsal vessel at any time during the whole larval
life of this type, while, on the other hand, in the advanced larve
of the other three types, it could be readily recognized as such.
ON DEVELOPMENT ETC. OF PHORONIS. 569
Fig. 58 c represents a portion of a transverse section through the
anterior region of the trunk of a larva belonging to type D. The
vessel in question here appears as a small canal (d.v.) running in
the stomach walls (sim. and m.sh.). As will be seen in the figure,
the canal is distinctly lined with an epithelial cell-layer. The
dorsal vessel terminates anteriorly just behind the postoral sep-
tum, so that the whole course of the vessel is confined to the
trunk region. During larval life, the dorsal vessel does not extend
so far posteriorly as to become confluent with the cecal con-
tractile capillaries which are formed at the point of juncture of
the stomach and the intestine. Thus we see in Fig. 554, which
is taken from a transverse section through the lower portion of
the stomach, that the gut is covered with a thin mesoblastic wall
without a trace of the dorsal vessel, but the capillaries (v.c.) are
here already developed at this period. In the above montioned
figure they are found as cell masses protruding into the trunk
cavity from the right side of the splanchnic attachment of the
ventral mesentery to the gut; one capillary is seen in cross sec-
tion. The capillaries shown in that figure are certainly in an
early state of development, and, when fully developed, they appear
like a tuft consisting of tolerably long, blindly ending tubes.
Sometimes I have observed that the capillaries are formed not
only on one side of the ventral mesentery, but on both sides of it;
and that they are not constantly formed on the ‘gut walls, but
sometimes on the ventral mesentery. Thus we see in fig. 57 a on
the ventral mesentery (v.mes.) a rosette-like figure (v.c.), the rudi-
ments of the capillaries seen under a low power. It is represented
highly magnified in fig. 57 (taken from another neighbouring
section of the same series). Here are seen signs of cell-multipli-
cation on either side of the ventral mesentery (e.mes.), in places
510 I. IKEDA:
not in direet contact with, but separated by a considerable distance
from, the gut.
From the facts above stated, we are justified in concluding
that the cæcal capillaries are not produced as out-growths of the
dorsal vessel, but are formed independently bv cell-multiplieation
taking place in certain parts (near the gut) of the ventral mes-
enterv, and, therefore, that the dorsal vessel and the capillaries
have different origins.
Next T will consider the origin of the ring vessel of the adult
animal, Although some early authors have frequently referred to
the so-called ring vessel of Aetinotrocha, vet its origin, form, and
position have never been satisfactorily elucidated.* Nobody has
investigated it by means of sections, and the statements which
have been made about it do not rest, it seems to me, on actual
anatomical studies of Aclinolrocha, but rather are mere inferences
from facts known respecting the metamorphosed larva. Conse-
quently there have been put forth several irreconcible views in
regard to the ring vessel. The structure described under that
name by CLAPAREDE, SCHNEIDER, METSCHNIKOFF, and CALDWELL
does not seem to be even one and the same thing.
In order to make clear the relations of this system of organs,
T must first of all deseribe somewhat minutely the adult collar
cavity. In the fully developed larva of every type, the rudiment
of that cavity is represented by a circular space on the inner
side of the tentacular row just above the septum. The space is
in form not a complete ring, but is interrupted at the median
i
# In passing I should say that MASTERMAN’s ring sinuses have nothing to do with the
vessel under consideration, because of them the first is stated to be priesophageal, the second
peri-intestinal, and the third perianal in position; and thus they must be something entirely
different from the ring vessel of other authors, which is a sinus in the splanchnic walls
around the stomach.
ON DEVELOPMENT ETC. OF PHORONIS. 571
dorsal point. The relative position of it with regard to the sep-
tum and the tentacles can be most conveniently studied on sagittal
sections of the larval body. In fig. 63 a the adult collar cavity
(s.c.c.) is indicated by a vertical club-shaped space just inside the
the body walls and above the postoral septum (mes.). Fig. 63 d
shows a portion of the ventral part of a nearly median sagittal
section similar to Fig. 63a. Here the cavity (s.c.c.) is seen also
as a vertical and comparatively wide space situated inside - the
body walls and below the tentacles (p.é.). It can also be made
out that the wall of the cavity is formed of a single layer of
mesoblastie cells and its ventral wall is in contact with the somatic
walls of the adult tentacle (s.2.), while the posterior wall is super-
posed on the anterior side of the septum (mes.). In fig. 58 @ and
582 the two cellular cireles (s.e.c.) attached to the inner side of
the tentacles (¢’.) represent a somewhat obliquely cut transverse
section of the adult collar cavity. The two tentacles belong to
the first pair, and the farthest dorsal point reached by the cavities
is, therefore, at the bases of these tentacles. When the serial
sections are traced posteriorly, these cavities gradually extend more
and more ventrally along the body walls and at last join with
each other in the median ventral line. In fig. 58c the cavities
appear as two narrow spaces (s.c.c.) appressed against the ectoblast
(ech). In fig. 55a, which is taken from a cross section of an
A-type larva cut nearly parallel with the tentacular row, the
cavity (s.c.c.) appears as a long slit-like space intervening between
the postoral septum (mes.) and the body walls. Here the cavity
is seen entirely free of the septum, because the section passes
through that portion of the collar which lies slightly above the
somatic insertion of the septum (compare fig. 63 d).
In somewhat younger larvæ of all types, the adult collar
212 J. IKEDA:
cavity is not yet extended dorsally as far as in fig. 58. Thus, in
fiz. 63 e it is represented by a mesoblastic cell-mass (s.c.e.) which
is placed just under the second tentacle (¢.”), and encloses no
Jumen in itself as vet. It is evident therefore that the adult
collar eavitv extends itself during development from the ventral
towards the dorsal side of the body, as is also the case with the
tentacles.
As the larva reaches the end of the swimming period, the
adult collar cavity in the adult tentacles becomes wider and wider,
nearly filling up the interior of the latter, while the larval ten-
tacular cavity is henceforth gradually reduced to a narrow space
appressed to the upper roof of the tentacle. Fig. 58 d represents
a median sagittal section of a larval tentacle of a larva of type D.
The portion belonging to the adult tentacle (s.t.) is characterised
by a very thick ectoblastic layer forming the ventral wall of that
tentacle. The adult cavity appears as a remarkably wide space
(s.c.c.) beginning at the somatic insertion of the postoral septum
(mes.) and ending at the tip of the adult tentacle (s.£.) The nar-
row cellular band (p.e.c.) resting upon the adult cavity (s.c.c.)
corresponds to the larval collar cavity of the tentacle. The
larval cavity is clearly seen in cross sections of the tentacle ; in
fig. 58 e it is visible as a small space (p.c.c.) inclosed by the adult
cavity (8.c.c.) except at the median dorsal point. Tracing the
cavity (p.c.c.) in the above figure to the base of the tentacle, we
sce that it communicates by a tiny opening with the larval
collar cavity. That portion of the tentacle, which is thrown off
during the metamorphosis (sce fig. 58 d) is distinctly different from
the persistent portion (the adult tentacle) in that the former has
no trace of the adult tentacular cavity (s.c.c). I have ascertained
after repeated examinations, that the retrogressing larval tentacular
N u."
ON DEVELOPMENT ETC. OF PHORONIS. BYE:
cavity becomes after metamorphosis the tentacular vessel of the
adult, as will soon be further explained.
Concurrently with the growth of the adult collar cavity the
larval cavity reciprocally diminishes in extent, and finally after
metamorphosis it is reduced to a narrow cavity surrounding the
gut in front of the postoral septum: this is the ring vessel of the
adult. In fig. 65, which is taken from a transverse section through
the tentacular region of a larva (type -{) just undergoing meta-
morphosis, a semicircular space (cir.c.) on the left side of the gut
represents the greatly reduced larval collar cavity or the ring
vessel, which is found dorsally attached to the somatic walls and
is laterally quite independent of the latter, though ventrally it is
fused with, and rests on, the postoral septum (mes.). One of the
tentacular vessels which proceeds from the ring vessel (cir.c.), 1s
denoted in the plate by &r. Fig. 64e also shows a transverse
section of the head portion (or a frontal section through the
supraseptal portion) of another partly metamorphosed larva re-
presented in fig. 11. Here a comparatively spacious cavity (cir.c.)
surrounding the gut and filled with the corpuscles, represents the
ring vessel from which the tentacular vessels are given off, though
this is not shown in the figure. In the above two figures the
adult cavity distinctly appears as a narrow space (s.c.c.) outside of
the ring vessel (cir.c.). In these stages of the metamorphosis,
the both dorsal ends of the adult collar cavity beeome continuous
with each other so as to form a completely circular space above the
postoral septum.
I am still uncertain as to how this rudiment of the ring vessel
comes into communication with the dorsal vessel ‘or with other
vascular spaces which make their appearance during metamorphosis,
for it is almost impossible to obtain larvæ of intermediate stages
974 I. IKEDA:
in which this vascular communication is just becoming established.
But from observations on metamorphosing larve I have obtained
certain suggestions respecting this process.
III. Metamorphosis.
As to the external changes of the larval body accompany-
ing metamorphosis, I have searcely anything to add to the exact
and detailed descriptions given by METSCHNIKOFF and WHILson.
I will, therefore, confine myself mainly to some anatomical points
which have been less studied by previous observers. My observa-
tions of the metamorphosis were mostly made with the larve of
types A and D which were most abundant in the neighbourhood
of the Station. Sometimes I have observed under the microscope
the whole course of the phenomenon, the duration of the so-called
critical moment being usually not more than 15-25 minutes.
Among the the material obtained with the surface-net we
often find Jarvee which carry about the partly evaginated pouch,
but these individuals can not be said to be undergoing metamor-
phosis in the strict sense of the term, for they may continue the
free swimming life for several davs after capture. Besides, they
do not show any remarkable change in the internal organs. In
them the corpuscles are still in masses, the nephridia preserve
their original form and position, while the alimentary canal is of
the ordinary form and length.
When the metamorphosis takes place, the partly evaginated
pouch protrudes suddenly outwards to its full extent and the
alimentary canal is thrown into convulsive contractions. Mean-
while the latter, especially its œsophageal and intestinal port-
ON DEVELOPMENT ETC. OF PHORONIS. 375
ions, is drawn out into a long tube, and at the next moment
the junction of the stomach and the intestine is first of all
pushed into the now spacious pouch. Thus, the anus and ten-
tacular region are brought into close approximation on the dorsal
line of the trunk; the digestive canal folds back on itself into an
U-shaped tube, as we find it in the adult. The larval tentacles
and the preoral lobe are cast off and digested in the stomach ; the
perianal ciliated belt atrophies in situ. It is during this moment
that the corpuscular masses break away and their elements are
scattered in the blood vessels, some of which are then being formed.
It is highly interesting to observe the breaking up of the corpus-
cular masses, and the motion of the corpuscles in consequence of
the rhythmical contraction and expansion of the dorsal vessel and
of the cwcal capillaries. The deep pinkish colour of the corpuscles
makes it easy to observe their progress into the vessels. Ordi-
narily after about twenty minutes the vascular system of the
worm 15 completely formed and the circulation characteristic of
Phoronis can be noticed. The skin of the foot which now forms
the principal part of the body, becomes more opaque on account
of the secretory products from the innumerable unicellular glands.
To understand the details relating to the metamorphosis we
must examine the animal in sections. Figs. 64 (a-d) show a dis-
continuous series of cross sections of the larva represented in fig. 11,
which is approximately at the critical moment of metamorphosis.
I will briefly sketch with the aid of these figures the general
internal change during metamorphosis.
Entoblastic Organs. When the ventral pouch is fully evagi-
nated and the stomach is pushed into the pouch, the oesophageal
portion elongates downwards enormously, so that the stomach
diverticulum descends far below the postoral septum, that is to sav,
5/6 t. IKEDA!
into the infraseptal cavity (fig. 64c, div.). The vacuoles in the
cell of the diverticular wall disappear at this period, and
the diverticulum itself is immediately afterwards wholly obliterated,
probably as the result of a histological atrophy. The stomach wall
docs not essentially differ in structure from that of the larva.
Mesoblastic Organs. Among the mesoblastic organs the vas-
cular system undergoes most noteworthy changes. MASTERMAN
has maintained that this forms a completely closed canal system
even in the free swimming stage of the larva. So far as I have
been able to ascertain, the closing up of the vessels into a conti-
nuous system occurs after the critical moment of metamorphosis.
Ifaving already described my own observations respecting the origin
of the ring vessel of the adult, I will now describe other vessels
which arise during metamorphosis.
Fig. 64a shows a cross section of the foot near the posterior
extremity where the alimentary canal is bent upon itself. Here
we see the cut ends of three contractile capillaries (r.c.) and three
sinuses (8.s.) In the stomach walls. A comparatively wide space
(d.v.) is found intercepted between the two limbs of the alimentary
canal. This space corresponds to the most posterior portion of the
dorsal vessel, which, if traced further posteriorly, shows itself to
be continuous with both the capillaries (v.c.) and the sinuses (8.s.).
A short distance more anteriorly, the dorsal vessel divides into
two parts cach of which attaches itself to a limb of the alimentary
canal; still more anteriorly the branch on the intestine disappears.
In that portion of the stomach which lies close to the cesophageal
tract, the dorsal vessel becomes enveloped with a thick muscular
sheath which we have before seen in Actinotrocha (fig. 646, d.v.).
The vascular sinuses gradually tend to unite into one common space
lying on the ventro-lateral side of the gut (fig. 642, ss; fig. 64 c,
ON DEVELOPMENT ETC. OF PHORONIS. 577
v.v.). At the place where the degenerating stomach diverticulum
still persists, the sinuses completely blend together into one large
blood vessel corresponding to the ventral vessel of MASTERMAN
(fig. 64c, v.v.). This vessel acquires a definite form at a more
anterior region of the œsophagus, becoming lined on its sides with
a thin mesoblastie wall (fig. 64d, v.r.). At about this level the
dorsal vessel (d.v.) on the cesophagus becomes a small canal, such
as we know it to be in the adult animal. According to my
observations, the large ventral vessel opens at this stage not by
two branches, as is the case in the adult, but by one directly into
the ring vessel. To my great regret, however, I have not been
able to study microscopically the larvæ in which the communica-
tion between the ring vessel and the dorsal or ventral vessel was
in the process of being established.
Now we see that the dorsal vessel of Actinotrocha corresponds
to the afferent, and the ventral vessel to the efferent vessel of the
adult. The sinuses around the stomach, which have newly
arisen during metamorphosis, develop into the complicated organ
of the adult.
Nephridia. At the stage when the metamorphosis takes
place, the nephridia do not show any important alteration in form
and structure from those of the swimming larva; the excretory
cells (exc.c) are found still attached to the blind end of the
nephridial canal (fig. 64 f, nep.c.) and the external nephridial pores
open behind the septum (fig. 64e, nep. o.). In fig. 64e we
notice only that the nephridia as a whole have shifted to a more
dorsal position than that occupied in the preceding stages (compare
with fig. 50c). This shifting of position becomes more and
more marked as the metamorphosis advances, so that when the
process is nearly finished, the nephridia on both sides come close
518 I. IKEDA:
to the so-called anal ridge on the dorsal side. Thus we see in fig.
66, which shows a transverse section through the nephridial region
of a completely metamorphosed larva of type À, that the two
nephridial canals (nep.c.) are situated very close to the intestine
(int.) which lies in the dorsal median line, and that one of them
(the right in the figure) opens to the exterior. By examining
serial sections it was found that the exeretory cells were entirely
absent on the nephridial canals, and that the latter were of an
inconsiderable length, ending blindly at the inner extremities. It
seems very probable, as was pointed out by CaLpwett, that the
excretory cells of the larval nephridia are thrown off into the body-
avity ; it is also probable that that portion of the nephridial canal,
which lies in the collar cavity of Actinotrocha, is obliterated, since,
as we see in fig. 66, the inner end of the canal (the right in the
figure) lies wholly outside of the septum (mes.), that is to say, in
the trunk walls, as is known to be the case in the adult. Thus
we may assume that the formation of the infraseptal nephridial
funnels of the adult is due to secondary outgrowths of the infras-
eptal portion of the atrophied, larval nephridial canals.
Ectoblastic Organs. As CALDWELL has correctly observed, the
larval tentacles and’the greater part of the preoral lobe are torn
off and are digested in the stomach. I have very often met with
the remains of these organs in the interior of the latter (2, fig.
64a). Although I have said, in agreement with CaLpWE Lt, that
the greater part of the preoral lobe is cast off, yet I can not
agree with the view entertained by several authors, that the epi-
stome of the adult developes from the remnant of the preoral lobe
of the larva. Because, according to my studies, the nerve ganglion
of the larva, which nearly marks the posterior limit of the lobe,
can not at all be discerned in the larva at such a stage as is
ON DEVELOPMENT ETC. OF PHORONIS. 379
represented in fig. 11. It is no doubt thrown off together with
the other parts of the preoral lobe. Besides, it is equally true
that the epistome can not be found in the neighbourhood of the
larval mouth at such a stage, while, on the other hand, in the
worm after metamorphosis the rudimentary organ is seen as a
small bud on the dorsal median margin of the mouth.
As before noted, the ring nerve of the adult is not yet formed
in the swimming larva. This nerve and the so-called brain of the
adult are of new formation; and the complicated nervous system
which had developed only in the preoral lobe, suffers the sume
degeneration as that larval organ. In fig. 64e the ring nerve
(r.n.)” is seen in section, just exterior to the septum (mes.). It
consists of a thick laver of very fine nerve fibres.
IV. Supplementary Notes.
J. Mutter (’46), the first discoverer of <Actinolrocha, des-
eribed the animal as an adult worm under the name Acéinotrocha
branchiata. The ventral pouch was considered by him to be the
sexual organ. Doubts were afterwards thrown on his views by
Kroun ('54); and they were finally refuted ly SCHNEIDER (62),
who maintained that Actinolrocha is the larval form of a certain
Gephyrea. This idea was confirmed by KowaLewsky (°67) who
ascertained that Actinotrocha is the free swimming larva of
Phoronis. Since this renowned discovery of KowaLewsky nu-
merous papers on the anatomy and development of Phoronis lave
been published by many celebrated naturalists. But the singular
fact is that the life history of the animal has not been subjected
* Unhappily the letter r is misprinted in the plate as v.
580 | I. IKEDA:
to a detailed study. The metamorphosis of Actinotrocha is, of
course, one of the most curious phenomena in the animal ontogeny.
But the question which interested me almost to the same extent
was: how do the free swimming larvæ come to establish colo-
nies at such fixed and limited spots as are found in the Abura-
tsubo inlet? Accordingly I made several visits to the Misaki
Station solely for the purpose of obtaining clues to the elucida-
tion of this point. The results obtained are, I think, worth
mentioning.
As I have before stated, the breeding season of Phoronis
ijimai ranges through about one half of the year, say, from
November to June or July, during which months the swimming
larve, though few in number, are constantly found in Aburatsubo.
They, are, however, most abundant from the middle of Julv
to the middle of August. Among these larve some are very
young, having apparently just swum out of their birth-place ; but
the majority of them are fully grown.
On July 16th., 1898, I visited the place where Phoronis
ajimai flourishes, to see if it was still in possession of embryonal
masses. But these could no longer be found in the tentacular coils
of the mother animals which were, however, in the normal state.
On the 22nd. of the same month, I went again to the same
place, and every thing was in the same condition as before.
On August 6th. I visited the place for the third time and
to my astonishment I discovered that the animals had all died
off. Several deformed colonies were brought back to the Laboratory
and were kept in an ærated aquarium. On a close examination of
these colonies after a few hours, nearly all of the chitinous tubes
were found empty, only a few containing the putrefying remains
of the animal body. When I reexamined the colonies in the
ON DEVELOPMENT ETC. OF PHORONIS. 581
aquarium on the next morning, I found that some younger
animals had attached themselves to the tip or inner-side of the
tubes of the departed generation ; no doubt they had been hiding
themselves somewhere in the colonial masses on the previous day.
Most of these young worms measured 1-2 c.m. in length.
I had the same experience in the summer of 1899. On
August 2nd. I visited the place and dived under the ledge of
rock where the colonies had formerly flourished ; but I could ob-
tain nothing but some decaying masses of the tubes which emitted
a disgusting odour.
Judging from these facts, it seems to me not improbable
that Phoronis annually changes its generation.
As to the formation of colonies of Phoronis, it may be sup-
posed that the putrescent remains or a certain fluid secreted by
the adult act on the larvæ as a chemotropie reagent. But this
can scarcely be admitted as taking place in the wide and open
sea. I think, on the contrary, that this phenomenon is not to be
attributed to such a complex cause, but is to be regarded merely
as an accidental matter. The colonies of Phoronis qimat form a
compact and rigid mass together with some Ascidians and Mollus-
an shells, and adhere very tightly to the rocks; so that, when
once the animals form a colony in a suitable place, it may well
be assumed that they become gradually luxuriant. But this is not
really the case in Aburatsubo where the colony has remained al-
most the same in size for several years. TI think, what takes
place must be somewhat as follows : the places where the Phoronis
colonies are established year after year, must naturally be well
adapted to the life conditions of the worm, and when a large
number of larvee 1s metamorphosed, as must be the case, during
the above mentioned months, those larvæ that happen to attach
282 1. IKEDA :
themselves to the tubes of the already formed colonies, flourish and
attain full growth; on the other hand, if the larvæ become
attached at some unfavorable places, they must svon be washed off
by waves and many of them must perish before thev can find other
suitable places. To this wasteful death of the larvæ which have
lost the opportunity of finding suitable localities to grow on, must
be due the fact that they remain comparatively stationary in the
number of colonies and in their distribution.
Specific Position of Phoronis 1jimai OKA. According to
Corr’s table, there are 7 known species of Phoronis. But I can
not refrain from entertaining serious doubts as to the correctness of
the present mode of classifymg the Phoronide in general. Most of
the systematic data have hitherto been taken from the external
characters of the animals, such, for instance, as the colour and size
of the body, the number of tentacles, the general form of the
colony, ete. The question now is whether these external characters
are constant and can be depended upon for systematic use. Is
there not a necessitv for taking internal anatomical points into our
consideration ? According to my observations, Phoronis annually
changes its generation and about one half of the vear belongs to
the growing period. Specimens collected during this growing
season must necessarily differ from the adult in the breeding season
in the number of tentacles, in the length and size of the body,
ele. Wide diserepaneies are therefore found between OKA’s obser-
vation and mine on the same species, viz., Phoronis ıjimar. I
have no doubt that OKA made use of only the younger indivi-
duals as will be obvious from the following comparison :
Rody length. Number of tentacles. Length of tentacles.
Oka 40 mm. 150 (aver.) 2 min.
Ikeda 60-100 ma. 200-210 (aver.) Jmm.
ON DEVELOPMENT ETC. OF PHORONIS. 983
I think it possible that the distinction made between other
species may rest on à similarly unsound basis. |
Moreover there is another no-less important point to be con-
sidered. Soon after the report on Phoronis buski by Mc'Intosit
(88) was issued, BLAXLAND BENHAM published his paper (788) on
the anatomy of Phoronis australis, in which he ascertained and
rectified many important points which had been till then but in-
completely known. Among these the following two are the most
remarkable.
(1) Afferent and efferent blood vessels open respectively into
the recipient and the distributing vessels which run parallel and
form so-called ring vessel, Each tentacular vessel is connected at
its basal end not only with the recipient but also with the distri-
buting vessel.
(2) Each nephridial tube communicates internally with the
infraseptal cavities by means of {wo funnels. One, the smaller,
opens into the lateral chamber, while the second is considerably
larger and opens into the rectal chamber.
It is stated by BENHAM, that “ Mr. CALDWELL dealt only with
the larger of the two funnels, in his ‘preliminary note,’ but he
informed me by letter that he became aware of the existence of
the second funnel, shortly after the publication of his paper.” How
is it now with the ring vessel in P. kowalewsky ? CALDWELL says
nothing about it. Corr, on the contrary, denied the above characters
for P. psammophila.
In Phoronis yimat I have ascertained that these two struc-
tures are the same in every point as in P. australis. Fig. 62 «
shows a dorsal portion of a transverse section through the septum ;
two nephridial canals (nep.c.) are seen one on each side of the
intestine (en) and are partly imbedded in the chondroid tissue of
584 I. IKEDA!
the septum (mes.). These canals open by funnels (f.) into the
bodv-cavity (rectal chamber) which separates the intestine (iné.)
from the cesophagus (@s.). These funnels correspond to the larger
funnels of BENHAM, so that in a longitudinal section they appear
as long ciliated cell-masses running longitudinally along the
lateral mesenteries. If we trace the funnels a little downwards, we
find another kind of funnels on the opposite sides of the lateral
mesenteries, opening into the lateral chambers. They arc re-
presented in fig. 625 (f.), which is taken from the right nephri-
dium, They are short indeed and are apt to be overlooked.
Again as to the ring vessels, they are found always as two
concentric loops (the recipient and the distributing) standing side
by side, and, as was described by BENHAM, each tentacular vessel
receives two small branches respectively from the two vessels.
Besides, in Phoronis hippocrepia* which is known from Ilfracombe,
I lave ascertained that the above cited structures (the nephridia
and the ring vessels) are indubitably present without any modi-
fication.
If these structures can not really be found in Lhoronis psam-
mophila, similar specific anatomical deviations must exist in other
little studied species, for instance, in P. ovalis, LP. gracilis, P. buski,
and so on. From these facts and from the variability of the
external characters, I am at present unable to discover points by
which Phoronis australis, P. hippocrepia, and our Phoronis can be
differentially diagnosed.
Tokvo Imperial University,
Science College.
October, 1899.
ee
re ne ne en ee ee ee om u Gab eee ee ee ee — _
* For the opportunity of investigating this species, Tam much indebted to Prof. Yasuda of
the Second Iligher School who kindly gave me a small portion of a colony.
ON DEVELOPMENT ETC. OF PHORONIS. 585
Postscript.
While the manuscript of the present article was undergoing
revision, I was much pleased to read Roure’s elaborate work on
the development of the Phoronide.® For the sake of brevity I
will not here discuss the author’s theoretical considerations, but
will offer some remarks with regard only to his investigations
relating to developmental facts. Some of his results differ const-
derably from mine and from those of previous observers. And the
differences are such as do not seem to be due merely to different
conditions in the species investigated (Phoronis sabatiéri).
According to Rou e’s observations, the first four planes of
segmentation are all vertical and radial, the fifth being the first
that is horinzontal, thus giving rise to 16 blastomeres; and the
egg is composed of two sorts of larger and smaller cells from the
first cleavage. This differs considerably from the account given
in the preceding pages, which is in agreement with the studies
of FOETTINGER and Masterman (1900). It is, however, verv
difficult to decide which of the two opinions is correct or whether
both are correct. I am at present rather inclined to the latter
view. As to Roure’s belief in the peculiar unequal segmentation,
which is said to return soon after to the equal, I fear that the
eges dealt with by the author may have been somewhat premature.
I have often observed premature eggs undergoing a remarkable
unequal segmentation when mixed in water with spermatozoa.
With respect to the nature of plasmic corpuscles, RoULE’s
view is certainly identical with that of CALDWELL, though he does
* Etnde sur le development embryonaire des Phoronidiens.—Ann. d. Sei. Nat. Zool., T.XT,
No, 1-6, 1900, Ä
586 I. IKEDA:
not refer to the latter author. In examining Rovute’s figures (31
and 32), certain nucleated cells are seen dispersed in the blasto-
ceelic cavity. They are said to be the inner ends of some elong-
ated blastomeres. In the embryos of Phoronis tjimai, similar
processes are indeed discovered protruded from some blastomeres,
but they nerver contain the nucleus in their distal or inner ends:
the nucleus belonging to these elongated blastomeres is situated
also peripherally as in other normal blastomeres. The plasmic
corpuscles which have, as I have described in the present paper,
arisen from subsequent fragmentation of certain elongated blasto-
meres, exist very distinctly as separate bodies dispersed in the
blastocælic cavity.
In his present paper Route reiterates his former views about
the dual origin of the mesenchyme-cells. Upon this point I have
already given my own ideas, and here I have to add only the
following remark :
Though Route, like ScHuLTZE (’97) has regarded, the nephri-
dial pit as the origin of the ventral pouch of Aetinotrocha, the
subsequent development distinctly shows that the two structures are
entirely independent of each other in their origin.
] can not refrain from doubting the correctness of RouLe’s
observation that the larva studied by him possessed at no time
during the swimming life any septal membrane in the body-cavities.
They are structures which in the other forms of Actinotrocha
have been so accurately demonstrated by previous observers. The
technique employed by the author may perhaps be found to be
faulty in this respect. The fine threads denoted by “ brides
mesenteriques ” in his figures (57, 75, and 97) seem to have arisen
from the pieces of the otherwise continuous postral septum broken
by the knife-blade in microtomizing.
ON DEVELOPMENT ETC. OF PHORONIS. 587
Route has described the nephridia of the larva in a some-
what peculiar way. According to him, the organs lie considerably
anteriorly as they are found on both sides of the stomach diverti-
culum, and are said to be constructed of cells forming a synevtial
mass which is attached to the somatic walls. No lumen and no
leading canal have been detected in these masses. But, if judged
from the author’s text and figures, it seems to me highly pro-
bable that his so-called nephridia correspond to the corpuscle
masses of other writers. Having overlooked the postoral septum
and the true corpuscle masses, he seems to have come to mistake
the latter for nephridia. Thus, so far as I can understand his
description, he did not notice the change in position of the
organs with regard to the postoral septum during metamorphosis.
As to the number and the position of the stomach diverti-
culum, the larva of Phoronis sabalicri is described to be in the
same condition as that of P. tjimai. But the vacuolization process
of the organ is in the former species more complex than in the
latter, though it is simpler than in the larvæ studied by MASTER-
MAN. When these three cascs are considered together, it may be
concluded that they indicate specific variations.
Rovute’s “cordon dorsal,’’ which is considered to show the
rudimentary state of the rectum of the adult, has not been des-
cribed by any previous author, and also could not be detected in
any of the types of the larve studied by me. If this “ cordon
dorsal ’’ be reconstructed from Roure’s text and figures, it seems
to me almost without doubt that the structure referred to corres-
ponds to the dorsal vessel on the stomach. When Rovte’s figure
(75) and mine (58c) are compared, both of which show a cross
section through the tentacular region at a similar level, it will
be noticed that the “ cordon dorsal ’’ and the “ vaisseau dorsal ’’ in
588 I. IKEDA:
the former figure correspond respectively to the dorsal vessel and
the trunk cavity (anterior portion) in the latter. Again, in re-
gard to the alimentary canal, Rous states that the end portion
of the intestine atrophies, but according to the observations of
CALDWELL and myself, no portion of the larval alimentary canal,
except the stomach di verticulum, undergoes histolysis during
metamorphosis, the entire tract growing gradually in length.
RovLe’s views as to the origin of the blood vessels greatly
differ from those of CALDWELL, MASTERMAN and myself. Accord-
ing to Roue, the vascular spaces and the body-cavities are on-
togenetically the same thing, and the former is formed from a
coalescence of the irregular lacunal spaces of the latter. Though I
agree with him in considering the ring vessel of the adult as a
derivative of the body-cavity (collar) of Actinotrocha, yet I can
not accept his view attributing other vessels to the same process.
Besides, we see in his figures (82, 83, 86, 87, 88, etc.) that the
same vessel (dorsal) is placed sometimes between the afferent, and
the efferent, branches of the intestine, and sometimes on the somatic
walls, Can such a peculiar disposition of the blood vessels against
the skin be verified in the adult animal?
It may be known from RouLe’s contributions, that the larval
development is more accelerated in P. sabatieri than in other
species, So that the ventral pouch and some other organs are in
that species already well developed even in a larva of 6 tentacles.
But this seems of not much significance, since we know that even
in the same type of the larve the progress of larval organisation
does not always keep pace with the increase in the number of the
tentacles.
_ I es
ON DEVELOPMENT ETC, OF PHORONIS. 889
Finally I can but refer here to MASTERMAN’s new work (1900),*
which I was able to read only a short time ago. His views differ
so radically from those of previous authors and from my own, that
I cannot fully discuss so weighty a matter in a brief postscript.
May, 1901.
me ee ae ee ne u ETS -— —
* On the Diplochorda, III, the early development and anatomy of Phoronis Buskii, AI.
Quat. Jour. Micr. Sci., Vol. 43, 1900.
290 I. IKEDA:
Reference papers.
‘46. J. Müczer.—Bericht über einige neue Thierformen der Nordsee
(Müller’s Arch. f. Anat. u. Phys., 1846).
‘47. R. Wacexner.— Actinotrocha branchiata (ibid., 1847),
‘54. C. GEGEXBAUR.—Bermerkungen über Pilidium, <Actinotrocha, ele.
(Zeitsch. f. Wiss. Zool., Bd. V, 1854).
'54. A. Kroum.— Ueber Pilidium and Actinotrocha (Miiller’s Arch. f. Anat.
u. phys., 1854).
‘62. A. SCHNEIDER.—Üeber die Metamorphose von Actinotrocha branchiata
(ibid., 1862).
‘71, K. Merscunixorr.—Ueber die Metamorphose einiger Seethiere (Zeit-
schr. f. Wiss. Zool., Bd. 21, 1871).
‘80. I. B. Witson.—On Actinotrocha (Amer. Natur., 1880).
‘SL E. B. Wırsox.— The origin and significance of the metamorpliosis of
Actinotrocha (Quart. Journ. of Microsc. Sci, Vol. XXI, 1557).
‘82. E. Mrrscunixorr.—Vergleichende embryologische Studien über die
Gastrula einiger Secthiere (Zeitschr. f. Wiss. Zool., Bd. 27, 1882).
‘89. A. ForrtingEer.—Note sur la formation du mesoderme dans la larve
du Phoronis hippocrepia (Arch. d. Biolog., Tom. III, 1852).
‘82. H. CaLpwELL.—P’reliminary note on the structure, development, and
aftinities of Phoronis (Proc. R. 8, L., Vol. 34, 1882-1853).
‘85. II. Cacpwerr.—Blastopore, Mesoderm, and Metameric Segmentation
(Quart. Jour. Micro. Sci., Vol. 25, 1885).
‘88. Mc’Ixrosu.—Report on Phoronis Buskii (Challenger Report, XX, part
LXIII, 1888).
‘88. B. Bennam.—The anatomy of Phoronis australis (Quart. Jour. Micr.
Sci, Vol. XXX, 1888).
‘90. L. Route.—Sur le développement des feuillets blastodermiques chez
les Gephyriens tubicoles (C. I. Ac. Sci, CX, 1890).
‘96. L. livure.— Sur les metamorphoses Jarvaire du Phoronis sabatieri
(ibid., 1896).
#97, A. Oxa.— Nur une nouvelle espèce japonaise du genre Phoronis
(Auuvt. Zool. Japo., Vol. 1, 1897).
ON DEVELOPMENT ETC. OF PHORONIS. 291
‘97. E Scuuttze.—Ueber Mesodermbildung bei Phoronis (Trav. Soc. Imp.
Sci. Nat. St. Petersburg, Vol. XXVIIT, 1897).
‘97. A. T. Masterman.—Preliminary note on the structure of Phoronis
(Proc. Roy. Soc. Edinb., Vol. XXI, 1897).
‘97, A. T. Masrermax.—On the structure of Actinotrocha considered in
relation to the suggested Chordate affinities of Phoronis (ibid., Vol.
XXI, 1897).
‘97, A. T. MasrermMay.—On the Diplochorda (Quart. Jour. Micr. Sci., Vol.
XL, 1897).
‘98. L. Route.—Sur la place des Phoronidiens dans la classification des
animaux et sur leur relations avec les Vertebrés (C. R. Ac. Sci., 1898).
Postscrivt,
‘99, I. Routr.—Lv structure de la larve Actinotroque des lhoronidieus
(Proc. 4th Intern. Congr. Zoul., 1899).
I. IKEDA :
List of Abbreviations.
an, anus.
ant.div., anterior diverticulum.
a.v., afferent vessel.
bl., blastopore.
bl.c., blastocoelic pore.
cir.c., ring vessel, 2.c., the collar cavity.
col., collar of larva.
col.c., collar cavity.
corp., blood corpuscles and corpuscle
mass.
dig.a., digestive aren.
div., stomach diverticulum.
d.v., dorsal vessel.
ect., ectoblast.
exc.c., excretory cells.
J. and f”. nephridial fuauels.
fn, female pronucleus,
gl., nerve ganglion,
gld., gland.
int., intestine.
n., mouth,
mes., postoral septum.
mes’., preoral septum.
Mes., mesoblast cells.
m.gl., mucous gland.
m.f., mesenchymatous fibres.
m.n., male pronucleus
m.sh., muscular sheath of the stomach.
nep.c., nephridial canal.
neph., nephridium.
nep.o., nephridial pore.
nep.p., nephridial pit.
n.f., nerve fibre.
a@s., oesophagus.
0.po., pouch pore.
p.b., polar globule.
p.c.c., larval collar cavity.
per., peritoneal epithelium.
per.bel., perianal belt.
pl.co., plasmic corpuscle.
po., ventral pouch.
pre.bel., preoral belt.
pre.c., preoral body-cavity.
pre.l., preoral lobe.
p.r., posterior recess of preoral cavity.
p.t., larval tentacle.
ret., retractor muscle in the collar.
rel., retractor muscle in the trunk.
s.c.c., adult collar eavity.
8.0., Sense organ.
s.8., SINUS space.
s.t., adult tentacle.
slin., stomach.
Lo Wg a first, second, third
larval tentacles.
tr,, trunk.
ir.c., trunk cavity.
t.v., tentacular vessel.
v.c., vascular coeca or capillaries.
v.gr., ventral groove.
v.mes. ventral mesentery.
v.v., ventral vessel.
PLATE XX V.
Explanation of Figures.
Plate XXV.
1.—Egg with two blastomeres. x4B (Zeiss).
2.—Eeg with three (a) and four blastomeres (b). x AB.
3.—Ege with eight blastomeres, side view. x 4B.
4.—Esg with thirty two blastomeres, seen from the future ventral
side. x 4B.
5.—Young morula, ventral view. x 2D.
6.—Ventral view of an advanced blastula in which the gastral in-
vagination has become visible from the outside. x 2D.
7.—Ventral view of a gastrula, in which the blastopore has taken a
triangular form. x 2 D.
8.—Side view of an advanced gastrula, in which the blastopore has
become a transverse slit. x 2 D.
9.—Young Actinotrocha in which the nephridial pit is visible from
the outside. x 2D.
ig. 10.—Larva of four tentacles (side view) x 2D.
a. 11.—Metamorphosing larva of type A, sketched from a preserved
specimen, x 4A.
ig, 12.—Larva of type D, sketched from a living specimen. Greatly
magnified.
‘ig, 13.—Larva (of 14 tentacles) of type A, in the living state. Greatly
magnified.
. 14.—Highly advanced larva of type B, bearing 28 tentacles, ventral
view. Greatly magnified.
. 15.—Larva of type C, bearing 20 tentacles. «a represents a dorsal
view, b a ventral view, and c the multicellular gland on the hood.
Greatly magnified.
. 16.—Larva of type D), bearing 48 tentacles, after preservation. Greatly
magnified,
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PLATE XXYVl.
Plate XXVI.
Fig. 17.—Primary oocyte, showing the karyokinetic figure for the first
polar globule, x 2 imm. /s.
Fig. 18.— Section through the equatorial plane of the karyokinetic figure
for the first polar globule; the egg was preserved with sublimate
solution (Winkel oc. 3. x ob. 8).
Fig. 19.—After emission oftwo polar globules (Winkel oc. 3 x ob. 8).
Fig. 20.—Section showing one stage of fertilization, when the two pronuclei
(m.n. and fon.) stand side by side. x 2 imm. '/, (Zeiss).
Fig. 21.—Median section of fertilized egg, in which is found a karyokinetic
figure for the first cleavage. x 2imm. !/..
Fig. 22.—Median section of a young morula. x 2F.
Fig. 23.—Median section of a young morula, showing the blastocælic pore
(dl.c.). x 2F. |
Fig. 24.—Median section of a young blastula, in which one blastoderm cell
is seen giving off plasmic corpuscle (pl.co.). x 2 F.
Fig. 25.—Median sagittal section of an advanced blastula ; two plasmic
corpuscles are detected in the segmentation cavity. x 2F.
Fig. 26 (a, b).—Sagittal sections of a highly developed blastula, in which
the invagination has just begun; «a shows a median section. x 2
imm, '/ıe-
Fig. 27.—Median sagittal section of a gastrula in which the invagination
is deeper. x 2 imm. ‘/.
Fiss. 28 (a-c).—Three transverse sections of an advanced gastrula ; a through
the central depression, ) behind the central depression, and c near
the posterior end of the embryo. x 2F.
ig. 29.—Median sagittal section of an embryo at nearly the same stage
as in Fig. 8. x 2imm. '/..
Figs. 30 (a-c).—Transverse sections of an embryo at nearly the same stage
as the preceding; a shows a portion (left) of a section through the
blastopore, b just behind the blastopore and through the ventral groove
(v.gr.), c near the posterior end. x 2 F, a with the tube drawn out.
ig. 31.—Transverse section through the blastopore of a larva, in which the
anterior diverticula (ant.div.) are well developed. x 2 F.
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PLATE XXYVLI.
Plate XXVII.
Fig. 32.—Transverse section through the middle portion of a larva in which
the ventral groove has ceased to give off mesoblast cells. x 2 F,
with the tube drawn out.
Fig. 33. Slightly oblique sagittal section of a larva in which the nephridial
pit (nep.p.) has made its first appearance. x 2 imm. '/,..
Fig. 34.—Oblique frontal section of a larva nearly at the same stage as in
Fig. 9. x ZF.
Fig, 35.—Transverse section through the nephridial pouch (nep.p.) as yet un-
paired. x 2}.
Fig. 36.—Oblique sagittal section of the nepliridial pouch (ncp.p.) partly
divided. x 2F.
Fig. 37 and Fig. 38.—Show respectively sagittal and frontal sections of
larvæ, in which the proximal end of the nephridial pit is about
to divide into two. x 2F.
Figs. 39 (a-c).—Three transverse sections of the two nephridial canals, cach
of which has respectively an internal opening. x 2}.
Fig. 40.— Sagittal section of a larva at the stage represented in Fig. 10.
x 2F.
Vig. 41.—Frontal section of a larva at the same stage as the preceding, in
which the two nephridial pores have separated widely from each
other. x 2F.
Fig, 42.—Oblique frontal section through the nephridial region of a larva a
little younger than the preceding. x 2F.
Fig. 43.—Transverse section through the upper portion of the wsophagus of
a larva of four tentacles, x 21°.
Fig. 44.—Large mesoblast cells which are found in the body-cavity of the
larva of two or four tentacles. x 2F.
(All the figures from fig. 45 to fiy. 55 ure drawn from larve
of type A.)
Fig. 45.—Median sagittal section of a larva of 12 tentacles. x 2D.
Fig. 46.—Portion of a longitudinal section of a larval tentacle of 10-
armed larva; two blood corpuscles are represented in the tentacular
cavity (corp) x 2F.
Figs. 47 (a-c).—Three longitudinal sections of the uephridium of a larva of
12 tentacles. x 2F.
Fig. 48.—Median frontal section of a larva of 16 tentacles. x 2 D.
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PLATE XX VIII.
Plate XXVIII.
Fig. 49.—Anterior portion of a median sagittal section of a larva of 14
tentacles, x 2F.
Figs. 50 (a-c).—Three transverse sections of a larva of 14 tentacles; a
through the stomach diverticulum (div.), b obliquely through the
postoral septum (ames.), and c above the pouch pore. x 2D.
Figs. 51 (a, b).—Longitudinal sections of the nephridium of a larva of 16
tentacles. x 2F.
Fig. 52.—Transverse section through the junction of the stomach and the
vesophagus of à larva of 16 tentacles, showing the rudiment of the
dorsal vessel (d.v.) x 2imm. '/,.
Figs. 53.—Section of the above larva, through the corpuscular mass which
floats in the collar cavity. x 2F.
Fig. 54.—Sagittal section through the right side of the oesophagus of a larva
of 16 tentacle. x 2F.
Figs. 55 (a and L).—Cross sections of a larva of 16 tentacles. In the figure
a is represented the right half of the section which passes through
the stomach (stm.) and the adult collar cavity (s.c.c.); b through the
junction of the stomach and the intestine, whereto the contractile
cweca (v.c.) are attached. x 2F.
(Lugs. 56-57 are drawn from larve belonging to type C.)
Fiss. 56 (a-c).—Cross sections of the 7. where the multicellular gland is
represented in different planes, 2D. Letter b omitted.
| Bigs. 57 (a-c).—Cross sections through the trunk of a larva of 22 tentacles ;
a under the magnification of 2 x D; b a alo of the trunk
walls containing the nephridial canal, magnified 2 x F with the
tube out; c shows a portion of the ventral mesentery near the gut,
magnified 2 x F.
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PLATE XXIX.
Plate XXIX.
(Figs. 58-60 are drawn from larve belonging to type D).
Fig. 58 (a-e).—These figures show different parts of a larva of 44 tentacles.
Figure a shows a cross section through the mouth, b through the
middle portion of the cesophagus (@s.); magnified x 2B; c a
portion of a cross section of the stomach wall (dorsal) and the
trunk walls, x 2F. d ande show respectively a longitudinal and
a transverse section of a tentacle (the former magnified x 2D,
the latter x 4D).
Figs, 59 (a-d).—Four cross sections taken from a series, not consecutive,
and their respective planes of section are given in the text with
reference to the woodcut (p. 542). Unfortunately in these series,
the tissues have undergone a great disturbance by the killing reagent,
but the relations of the layers remain essentially correct, and those
spaces which have been produced from the mutual splitting of the
layers are denoted by “ artefact.” x 2 D.
Figs, 60 (a and b).—Represent the nervous system of Actinotrocha (of
type D), revealed by vital staining with methyl blue and ammonium
molybdate. In a, as the larva was pressed by the coverglass, the
rim which appears like the free margin of the hood is not that
edge at all, but represents the line along which the hood was
bent by pressure; the line drawn near the peripheral blue dots is
the true edge of the hood. D shows a portion of the free margin of
the hood, where the nerve fibres end. a x 4B, b x 2F.
Figs. 61 (a and b).—Are taken from serial sections of a A-type larva
which bears the evaginated pouch ; a shows four blood corpuscles, 5
one portion of the wall of the stomach diverticulum. x 2 F.
Figs, 62 (a and b).—Taken from serial transverse sections through the
nephridial region of the adult. 6 shows the inner and of the left
nephridial canal where the smaller funnel (/’) is attached.
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PLATE XXX.
Plate XXX.
Figs. 63 (a-e).—Are taken from several parts of serial sagittal sections of
B-type larva of 28 tentacles,
a. median section through the hood and the collar. x 2D.
b. more magnified figure of the nerve ganglion (g/.) and the
posterior recess of the preoral cavity (p.r.) in the preceding figure.
x 2F,
c. taken from a section lateral to the œsophagus and to the right
of that of the figure a. x 2F.
d. ventral portion of the collar-trunk walls, where the septum (mes.)
and the adult collar cavity (s.c.c.) are cut through. x 2 F.
e. portion of a section which passes through the second tentacle
(’) x2F.
Figs. 64 (a-f).—Transverse sections taken from serial sections of a meta-
morphosing larva of type A represented in fig. 11. The respective
explanations of them are introduced in the text (p. 584). From
a to c magnified as x 2D, and from d to fas x 2F.
Fig. 65.—Portion (right side) of a transverse section through the tentacular
region of a metamorphosing larva of type A, slightly younger than
that of fig. 11. x 2F.
Fig. 66.—Transverse section through the nephridial region of wholly meta-
morphosed larva of type 4. x 2F.
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eonr Selsmic and Other
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By F. Omorr. (With
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>um A tes (Selenoaamatear
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Preparation of Hyponitrite from Nitrite through Oxamidosulphonate. By E.
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Absorption of Nitric Oxide in Gas Analysis. By E. Divers.
Interaction of Nitric Oxide with Silver Nitrate. By E. Divers.
Preparation of Pure Alkali Nitrites. By E. Divers.
The Reduction of an Alkali Nitrite by ag Alkali Metal. By E. Divers.
Wyponitrites: their Properties and their Preparation by Sodium or Potassium
By. E. Divers.
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On the Geologic Structure of the Malayan Archipelago. By B. Kotô. (With Plate I.
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Earthquake Measurement at Miyako. By F. Omorı and K. HIRATA. (With Plate)
XVIFXXITI).
Ethyl ammoninmsuiphite. By E. Divers and OGAWA.
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quakes. By F. Omori. (With Plate XXVI-XX VIT).
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Potassium Nitrito-hydroxymidosulphates and the Non-existence of Dihydroxy.
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Identification and Constitntion of Fremy’s Sulphazotized Salts of Potassium, his
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On a Specimen of a Gigantic Hydroid, Branchiocerianthas imperator (Allman).
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Mutual Relations between Torsion and Magnetization in Iron and Nickel Wires.
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CONTENTS.
Vol. XIII, Pt. IV.
PAGE,
Observations on the Development, Structure and Metamor-
phosis of Actinotrocha. By Iwas: Ikepa, Rigakushe.
(With. Plate XXV-XX A) ... on cee one ve ose ve se 907
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