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Sr En cl TS 


044 106 339 5 























Digitized by Google 


Rin oh A ie BS 
mo # 


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. 








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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 























LES 
Sue 
EU, 

a 
4 
GaP | 
t 
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ins 
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oe 
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G, 
fe Si in Y Æ rn o '¢, 
a £ FL ' 
f rn # en | 
u ug } phi 
+ nn 


be 
= 
9 } 
à 
= 


& 4 rar ae 
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à Ge R = mir 
en LES Le 9 
3 


” je: 
té “dg 


be: 
tr 


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A 
an 
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16 





15 


14 








30 


29 
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28 
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21 
27 
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20 
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19 
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PLAT, 


Jour, Sc. Coll. Vol. XM PT. 


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16 


15 


14 


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11 








23 


22 


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15 


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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 


Digitized by Google 





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. 





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Potassium nitrososulphate. By E. Divers and T. Haca. 

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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. 

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Hyponitrites: their Properties and their Preparation by Sodium or Potassium 
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TE a a —— 





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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, 
Science College, Imperial University, Tokyo. (With Plates I-V). 

Change of Volume and of Length in Iron, Steel, and Nickel 
Ovoids by Magnetization. By H. Nacaora, Rigaku- 
hakushi, Professor of Applied Mathematics; and K. Honpa, 
Rigakusht, Post-graduate in Physics. (With Plates VI&VII). 

Combined Effect of Longitudinal and Circular Magnetiza- 
tions on the Dimensions of Iron, Steel and Nickel 
Tubes. By K. Honpa, Zigakushi ; Post-graduate in Phys- 
ics. (With Plates VIII & IX) Er cabin do 

Studien über die Anpassungsfähigkeit einiger Infusorien an 
concentrirte Lösungen. Arsusuı Yasupa, Rigakushi ; 
Professor der Naturgeschichte an der zweiten Hochschule zu 
Sendai. (Hierzu Tafel X-XIT) sis 

Ueber die Wachstbumsbeschleunigung einiger Algen wa 
Pilze durch chemische Reize. Von N. Ono, Rigakushi. 
(Hierzu Tafel XIIT)... 


PRINTED AT THE “ TOKYO TSUKIJI TYPE FOUNDRY.” 


PA 


RR BABE 
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re mn K 
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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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 


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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 





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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, 





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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, 


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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. 





CONTENTS OF RECENT PARTS. 


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e Fate of the Bilastopore, the Relations of the Primitive Streak, and the 
Formation of the Posterior End of the Embryo in Chelonin, together with 
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eine in Misaki vorkommende Art von Ephelotn und über thre Sporen- 
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m über die Schwefelrasenbildung und die Schwefelbacterien der Thermen 
jem Yumoto bei Nikko. Von M. MryosHi. (Hierzu Tafel XIV). 
atwiekelunmg der Gonophoren bei Physalia maxima. Von 8. Goto. (Hierzu 
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s of Reproductive Elements. III. Die Entwickelung der Pollenkörner 
rom Allium fistulosum L., ein Beitrag zur Chromosomenreduktion in 
Pfiaämzenreiche. Von C. IsHIKAWA. Hierzu Tafeln XVI und XVII). 
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ration of Hyponitrite from Nitrite through Oxamidosulphonate. By E. 
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stiom ef Nitric Oxide in Gas Analysis. By E. Divers. 
ttiom of Nitric Oxide with Silver Nitrate. By E. Divers. 
‘ation of Pure Alkali Nitrites. By E. Divers. 
eduction of an Alkali Nitrite by an Alkali Metal. By E. Drivers. 
ätrites: their Properties and their Preparation by Sodium or Potassium. 
3y. E. Divers. 


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Ethyl ammoniumsulphite. By E. Divers and OcAwa. 

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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, 
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JAPAN. 


VOL. XIII, PART III. 





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1900. 


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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 


| 


ie I enger 
a me 


wie) 
rye net — Ow je 
RE ewe 


u 


-- ah ges 
— 
aot: Oe = 


ss urn 
2X. 
- we 


>= 
ee | ee 
= wy eee + - te ln ee + a nf U” end . 


up 
Ser 


PR on, AU 


mary quae. Le mia 


1-2 Fi 
VE 


„me 


yn 


h 
| 









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. 





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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 


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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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396 Ss. HATTA: 


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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MORPHOLOGY OF CYCLOSTOMATA. " 423 


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424 8. HATTA: 


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m 


by: so! 


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 
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we See Sat teat ane 
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Jour, Sc. Coll, Vol, XM, Pl. XVI. 


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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). 





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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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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. 











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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. 





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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 


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Thee Interaction between Sulphites and Nitrites. By E. Divers and T. Haca. 


Vol. XIII, Pt. 8, published Dec. 28th, 1900. 
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CONTENTS. 


Vol. XIII, Pt. II. 


TAGE, 


Contributions to the Morphology of Cyclostomata. II.—The 


Development of Pronephros und Segmental Duct in Petro- 
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Beiträge zur Wachstumsgeschichte der Bambusgewachse. 
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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 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 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. 


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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Vol. XL, Pt. 1... yen 0.34 (Price in Tokyô). 

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. 


Vol, XL, Pt. 2... yen 0.38 (Price in Tökyö). 


On the Geologic Structure of the Malayan Archipelago. By B. Kotô. (With Plate I. 


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Horizontal Pendulums for the Mechanical Registration of Seismic and Other 
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Note on the Preliminary Tremor of Earthquake Motion. By F. Omori. (With 
Plates XITI-X VI). 

Earthquake Measurement at Miyako. By F. Omorı and K. HIRATA. (With Plate) 
XVIFXXITI). 

Ethyl ammoninmsuiphite. By E. Divers and OGAWA. 

Ethyl ammonium selenite and Non-existence of Amidoselenites (Selenosamatesf 
By E. Divers and T. Hapa. 

Notes on the Minerals of Japan. By K. JINBo. 


Vol, XL, Pt. 4... yen 1.04 (Price in Tökyö). 
On the Matual Influence between Longitudinal and Circular Magnetizations in 
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The Earthquake Investigation Committee Catalogue of Japanese Earthquakes, 


By S. SEKIYA. 
Notes on the Earthquake Investigation Committee Catalogue of Japanese Earth- 
quakes. By F. Omori. (With Plate XXVI-XX VIT). 


—— [on 


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Further Observations on the Nuclear Division of Noctiluen. By C. IsHikawa. 
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Tentamen Flore Lutchnensis. Sectio Prima. Plantre Dicotyledones Polypetal:e 
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Ammonium Amidosulphite. By E. Divers and M. Ogawa. 

Products of heating Ammoninm Sulphites, Thiosulphate, and Trithionate. By 
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Potassium Nitrito-hydroxymidosulphates and the Non-existence of Dihydroxy. 
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Untersuchungen über die Schrampfkrankheit („Ishikubyö) des Maulbeerbaumen. 
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Vol. XIII, Pt. 4, published Oct. 8th, 1901. 
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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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