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Full text of "The Journal of the College of Science, Imperial University of Tokyo, Japan"

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THE 



JOURNAL 



OF THE 



COLLEGE OF SCIENCE, 

IMPERIAL UNIVERSITY OF TOKYO, 
JAPAN. 

YOL. XIII. 



PUBLISHED BY THE UNIVERSITY. 

TOKYO, JAPAN. 

1900—1901. 

MEIJI XXXIII— XXXIV. 



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3 J (o'lo 



CONTENTS. 



Pt. I. Published June 2nd, 1900. 

Notes on the Geology of the Dep^dent Isles of Taiwan. 

By B. Koto, Fh. D. Rigaleuhakvshi, Professor of Grcology, 
Science College, Imperial University, Tokyo. {With Plates 
I-F) 1 

JL ¥ J ••• ••• ••• ••• ••• ••• ••• ••• ••• ••• ••• ••• A 

Change of Volume and of Length in Iron, Steel, and Nickel 
Ovolds by Magnetization. By H. Nagaoka, RigdkuhakmU, 
Professor of Applied Mathematics, and K. Honda, Rigaku8h% 
Post-graduate in Physics. {With Plates VI & VII) 57 

Combined Effect of Longitudinal and Circular Magnetizations 
on the Dimensions of Iron, Steel and Nickel Tubes. 
By K Honda, Bigakushi, Post-graduate in Physics. (With 
Plates VIII & IX) 77 

Studien uber die Anpassangsfahigkeit einiger Infusorien an 
concentrirte Losungen. Von A. Yasuda, Bigahushi, Pro- 
fessor der Naturgeschichte an der Zweiten Hochschule zu 
Sendai. (Hierzu Tafel X-XIl) 101 

Ueber die Wachsthumsbeschleunigung einiger Algen und 
Filze durch chemische Keize. Von N. Ono, Uigakibshi. 
{Hierzu Tafel X-XIII) ... 141 

Pt. II. Published July 2Bth, 1900. 

Ammonium Amidosulphite^ By E. Divers and M. Ogawa. 

Imperial University, Tokyo 187 

Products of heating Ammonium Sulphites, Thiosulphate, 
and Trithionate. By E. Divers and M. Ogawa, Imperial 
University, Tokyo 201 

Fottassium Nitrito-hydroxymidosulphates and the Non- 
existence of Dihydroxylamine Derivatiyes. By K 
Divers, M.O., D. So., F. E. S , Emeritus Prof., and T. Haga, 
D. Sc, F. C. S., Professor, Tokyo Imperial University 211 



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Identification and Constitution of Fremy's Sulphazotised 
Salts of Pottasium, his Sulphazate, Sulphazite, etc. By 
E. Divers, M.D,, D. Sc., F. E. S., Emeritus Prof., and T. Haga, 
D. Sc, F. 0. S., Professor, Tokyo Imperial Uaiversity 225 

On a Specimen of a Gigantic Hydroid, Branchiocerianthns 
imperator Allman)^ found in the Sagami Sea. By M. 
MiYAJiMA, Rigakuahi, Science College, Imperial University, 
Tokyo. {With Plates XIV & XV) 235 

Mutual Relations between Torsion and Magnetization in Iron 
and Nickel Wires. By H. Nagaoka, Bigakuhakmhi, Pro- 
fessor of Applied mathematics, and K. Honda, RigakusMj 
Post-graduate in Physics. {With Plates XVI) 263 

The Interaction between Sulphites and Xitrities. By E. 
Divers, M. D., D. Sc., F. E. S., Emeritus Prof., and T. Haga, 
D. Sc F. C. S., Professor, Tokyo Imperial University 281 

Pt III. Published Dee. 28th, 1900. 

Crontributions to the Morphology of Cyclostomata. IL— The 

Development of Pronephros and Segmental Duct in Petromyzon. 
By S. Hatta, Professor in the College of Peers, Tokyo. {With 
xlcites Jx. V J.J.— /vJxJ. 1 ... ... ••• •.• ..• ••• •.• ••• ... oXX 

Beitrage zur Wachstumsgeschichte der Bambusgewachse. 

Von K. Shibata, Rigahushi. {Mil Tafeln XXILXXIV) ... 427 

Decomposition of Hydroxyamidosulphates by Copper 
Sulphate. By E. Divers, M. D., D. Sc., F. R. S., Emeritus 
Prof., and T. Haga, D. Sc., F. C. S., Professor, Tokyo Imperial 
* LyXiiveFoivy ••• ... •.. ••• .*• ••• ... ••• ••• •«• ••• 7«/ f 

Pt IV. Published Oct. Bth, 1901. 

Observations on the Development, Structure and Metamor- 
phosis of Actinotrocha. Iwaji Ikeda, Rigakushi. {With 
Plates XXV-XXX) 508 



PRINTED at the "TOKYO TSUKIJI TYPE FOUNDRY." 



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Publishing Committee. 
H 

Prof. K. MitSUkuriy Ph. D., Bigakuhakushi, Director of the College 

(eat officio). 
Prof. B. Kotdy Ph. D., RigahuhakusM. 
Prof T. Haga, RigdkuhdkusM. 
Prof. S. Watase, Ph. B., RigahuhakusM. 



All eommnnications relating to this Journal should be addressed to the 
Director of the College of Science. 



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^ ]^ ^ IB :fc '^ >?£ 5- 

if! -h ^ «• ^ :i: 9t 
THE 

JOURNAL 

OF THE 

COLLEGE OE SCIENCE, 

IMPERIAL UNIVERSITY OF TOKYO, 

VOL. XIIL, PART I. 



PUBLISHED BY THE UNIVEESITY. 

TOKYO, JAPAN. 

1900. 

MEIJI XXXIII. 



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Publishing Committee. 
^ 

Prof. K. Yainagaway Ph. B., RigakuhakusUy Director of the College 

{ex officio), 
Pro£ J. Sakuraiy RigdkuhakusM. 
Pro£ B. Kotd, Ph. D., RigakuhakusM. 
Pro£ L Ijima, Ph. D., Rigakuhdkushi, 



AU eommonieatioiiB relating to this Journal should be addressed to the 
Director of the College of Science. 



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Notes on the Geology of the Dependent 
Isles of Taiwan. 



By 



B. Eotd) Ph. D. R!gakiihakiishi, 
Professor of Geology, Science College, Imperial University, Tokyo. 



With Plates LV. 



THE HOKO GBOUP (PE8C ADOBES). 

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 Hoko 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 Hoko Channel, — the only pas- 
sages which allow free communication to the waters of the de- 



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KOTO : NOTES ON THE GEOLOGY 




Fjo. 1.— Index man of Taiwan to show 
the position of the islands descril)ed. 



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 KiirO'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- 
ness of Messrs. Y. Saito and 
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, 
H6ko, Hakusha, and Chd-don, the latter three fuse together, 
especially during low tide, into one mass with the intervening 



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OF THE DEPENDENT ISLES OF TAIWAN. 3 

coral-reefs which stretch from one island to tlie other, making the 
shape very much like Thera. The single island of Ciio-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 Hantorin. 
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 06 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 wMud-protected harbours, being constantly 
exposed to the north-east stiflf 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 



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4 Koto : NOtEs oj^ the geology 

liave a jwasi-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) H6ko island, the largest of the whole group. ' 

2) Haku-sha-t6,'^ lying north to the foregoing. 

3) Impai-sho. 

4) Ch6-sho, the eastern neighbour of Hakusha-to. 

5) Kippai (Bird Island of English Admirality chart), the 

northernmost of the whole group. 

6) Gi-6-t6 (Fisher Island), west of H6ko-t6. 

7) Hatto-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. Saito. 

1) The words *td' and *tho* recurg frequently in the geographical name of Taiwan, the i 

former signifying an island, the lalier an islet or rock. ^i 



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OF THE DEPENDENT ISLES OF TAIWAN. O 

II. Stratigraphical Characteristics. 

HOEO ISLAND. 

Hoko or Tai-san-sho^^ is the largest among the forty-seven 
islands of the Hoko group, having an area of 62.7 square kilo- 
metres. Its general outline is k-shaped, curving in at three 
points in the coves, Fiikibi,^^ Giil-bo-ken,^^ and K6lei.^^ 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. 

LIr. Y. Saito has geologically reconnoitered the principal 
islands of the group during last winter, and has kindly placed at 
ray 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, 

■iiiiiittfilLjnarked by the intervening tufaceous sedimentaries of a 



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4 Koto : NotES oj^ the geology 

liave 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) H6ko island, the largest of the whole group. 

2) Haku-sha-t6,'^ lying north to the foregoing. 

3) Impai-sho. 

4) Ch6-sho, the eastern neighbour of Hakusha-to. 

5) Kippai (Bird Island of English Admirality chart), the 

northernmost of the whole group. 

6) Gi-6-t6 (Fisher Island), west of Hoko-to. 

7) Hatto-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. Saito. 

1) The words *t6' and *«^' recurs frequently in the geographical name of Taiwan, the 
former signifying an island, the lat ter an islet or rock. 



CORRIGENDA. 



Page 4, 12th line, for Gi-6-t6 read Gio^tft. 

' * " for Cholnecky read Cholnoky. 

for there lies read the relics. 

for basite read bastite. 

for crystals read crjstal. 

for Bilking read Bucking 

delete the word * macroecopically.* 

for * leached * read * leached or percolated/ 

for pyrites read pyrite. 

for granual read granular. 

Explanation Pts. I <& II, for octant read quadrant. 



4, 


12th line, 


6» 


the last line, 


18. 


18th line, 


21, 


the last line, 


28, 


9ih line, 


29, 


13th line, 


42, 


2nd line, 


45, 


14th line. 


63, 


24th line, 


57, 


20th line. 



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OF THE DEPENDENT ISLES OF TAIWAN. O 

II. Stratigraphical Characteristics. 

HOEO ISLAND. 

Hoko or Tai-san-sho^^ is the largest among the forty-seven 
islands of the Hoko group, having an area of 62.7 square kilo- 
metres. Its general outline is k-shaped, curving in at three 
points in the coves, FAkibi,^^ Giu-bo-ken,^^ and Kolei.^^ 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. 

Ivlr. Y. Saito has geologically reconuoitered the principal 
islands of the group during last winter, and has kindly placed at 
ray 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 Ihrec different flows, poured out after long intervals, 
are well marked by the intervening tufaceous sedimentaries of a 
considerable thickness. The topmost 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 Fukibi-Jiri^^ 

J) Tai-san-sho (iclUl^), signifying * great mountain islet/ is by no means literally true, 
thongb undoubtedly it is llie largest of the whole Pescadores. 



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6 k:ot6 : notes on the geologV 

tongue, which projects out from Sei-shi-an^^ towards the citadel 
of Bako, thus enclosing within it a safe harbour, — we see the second 
sheet of flow J 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 Area were secured by Sailo from the corresponding bed at 
Run (Lun) point in the Island of Gio-6. 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) *ff«. 

2) 'Vorliiufiger Bericbte uber meioe Forscliuogsreise in CliiQa.' Pdermoam» Mith, 45» 
1899, a 8. 



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OP THE DEPENDENT ISLES OP TAIWAN. 7 

distinguished in Eastern Asia ; the one has served for the well- 
ing out of an 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 Chaiig-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 Eiver 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 V^nukoflf*^ 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 epochy a subaerial condition 
has prevailed over Hoko, 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 Bodies basal tiq ties de la Mongol ie/ BuUetiv de la Scci&i b^ge de GSologie, etc.f 
tome II, p. 441. 

2) < Shantung und seine Eingangspfort Kiautschou/ 1898, S. 66. 



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8 KOTO : 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 deposilSy 
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 Bak6'^ 
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 lies of the young columnar lava, 
being separated from it by a blue rock. The last is a fullers 
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. 



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OP THE DEPENDENT ISLES OP 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-bA-snn, nenr Jiri, showing two upper flowi with 
iutirbedded sedimentnries.* 

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-b6-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 
Bako, 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 

*A11 the figures in the following wood-cuts, not otherwise mentioned, are originally 
sketched bv Y. SaitS. 



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10 



KOTO : XOTES ox THE GEOLOGY 



No. 1. lUisalt. 



^-: 



y, /A No, 2. Basalt. 



' '^\^U '■'■'■J^'?''''^j£. Ba"<l<^l felspar 
sands and clays. 






Felspar wind with 
limouito noJulcs. 



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 
flat and form the ground of 
Pampanga, north of Manila^^ 
After this short digression, I re- 
turn to the former subject. Now, 
a yellowish-brown, loose sandy 
bed, 3 m. thick, comes below the 
middle flow, locally with liraonitic 
nodules (Fig. 3). This is succeed- 
ed by another complex bed, 3 to 
4 m. thick, made up of multi- 
farious alternations of clays and 
sands, all retaining the original 
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 Tertiary ; and this profile 
serves as a type of the stratigraphical order of the island. After 
passing over the second * haul-over ' to the Fiikibi 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 Hoko. 
The adjoining wood-cut shows clearly the geological structure 

1) B. Kolo, * Geologic Structuie of the ^lalayan Archipeljigo.' This Jourval, Vol. XI., 
p. 113. 



'-J No. 3. Basalt. 

Sandy clay witli 
lignite. 

Fin. 3.— Section exposed at the west coast 
of Jiri, irOko. 



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OF tflE DEPENDENT ISLES OF TAIWAI^. ll 

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 




Fio. 4.— A Tiow of tbc Islo of Ko-sei-sho, nu erosion relic of Basaltic m&A, 
as seen from tlie coast 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-au, already referred to, and 

going round the south coast along the points of Kan-on-san^^ 

and Ko-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 Bako, 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 Sait6, 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, i.e. on Front 

Taiwan, there are not wanting evidences tending to prove the 

negative change on the shore. 



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12 tOTO : NOTES ON THfe GEOLOGl" 

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 RL-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 Koto 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 Area zone 
in Gio-6 Island, already referred to. The exposure is meagre and 
soon disappears under the rubbish to be seen no more. This 
mineral combustible is but imperfectly incarbonized, and the 
woody structure is said to be yet well preserved. 

From Sei-kei^^ through Ko-tei*^, and Sha-ko^^ 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-to,^^ or the white sand island is bodily connected 
with Hoko through the intervening islet of ChA-don'^, at the 

1) XJEA 2) 4:« d) HSt 4) Hitt 6) »» 6> ^^ 7} i|i«* 



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OF *HE DfiPEI^DJSNT ISL£S OP tAlWAN. 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 sedimeularics seem to be wanting. 
The other features that strike the eyes of observers are firstly, 
the lowness of its relief, the highest point being K6-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 Hoko, 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 Cho-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 half- 

1) tl«Ul 2) AM 3) A« 4) ^a. 1 . 



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14 K0X6 : NOTtS O^ *HE GtOtOOY 

hardened foraminifer sand {PL Ily 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 w^hich forms 
spur-like marginal appendages, characteristic of Calcarina, and 
its external form and microscopic details agree well with C. 
Spenglerij Linn6'\ dredged for the first time near the coast 
of Amboina at the depth of 1,425 fathoms. This species seems 
to be quite as abundant 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. 

010-6 ISLAND. 

Gio-6, or Fisher Island, lies to the west of Hoko, 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) CkalUrif/er BepOrt, ' Fora^iTejra.' 

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OF THE DEPENDENT ISLES OP TAIWAN. 



lo 







Sandy tuffite. 



No. 2. Pasalt 



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 tulEtes as 
well as the underlying bed are likewise present, and well seen 

especially in that portion that lies 
southwards of Sho-chi-kaku.'^ 
Between the last-mentioned 
locality that is situated in the 
middle of the island and ShA- 
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 muscovUe, plagioclase, 
and Basalt-glass. Concentric 
nodules of hematite are frequent- 
ly found in them. Sait6 is 
fortunate enough to find in this complex bed casts of an Area and 
gasteropod {Turbo) in the matrix of ferruginous felspar sand 
with a little magnetite. Judging from the cast, the shell of the 




Fel par sanl. 



No. 3. Basalt. 



Sandy clay. 



FiQ. 3.— General profile at seen in the aoutbem 
part of Gio-6. 



1) tA^ 2) 9K9- 



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16 KOTO : NOTES ON TEE GEOLOGY 

Area 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-6, it should be remarked that the area 
north of Sho-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 : — Hatto,^^ with the dependent 
isle of Sh6-gun-6^^ ; the Smaller and the Larger Bi6-sho% so 
named cat islands from their appearance as seen from a dis- 
tance ; Tai-sho^^ and Sh6-hei^^ with columnar Basalt ; T6-kitsu^^ 
and Sei-kitsu^^, likewise Basaltic ; all being encircled by coral 
reefs. 



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OP THE DEPENDENT ISLES OP TAIWAN. 17 



ni. Petrography of the EfiUsives. 

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 of the 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 Saito, presents 
the same physiognomy, and consists of the same black rock. The 
specimens, brought back from most 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. Sait6, 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- 



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18 kot6 : 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. Saito recognises, as I 
have already said, three lava-flows in the Northern group, 
viz.y the uppermost or youngest being of columnar^ the middle 
porous and vesicular^ and the lowest also partly vesicularj 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. 



I 



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 Hoko 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, 



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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 Masseniairkunffj 

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 H6ko 

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 L Figs. 1, 
4 and 5) and the kernel {PL L Fig. 6), differs in habit. The 



1) 'Ueber die Natur der Glasbasis, sowie der Krystallisationsvorgange im eniptiyeD 
Magma/ T. M. M. Bd. VIII, 1887. 

2) Ibid. Bd. XVm, 1898. 



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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 
Lawson^^ described a similar mineral and the latter author 
named it iddingsite. In the secondy we fail to find such a 
distinct cleavage, and it seems to me to be the same body which 
Michel- Levy called the mineral rouffe^\ 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. Tbe lattery 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 lamellae begins with the development of a 
fine parting which appears like stripes, and which runs parallel 
to the vertical axis {PI. L 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 difl'erence in the 



1) U. S. Geol. Surv., ' Monograph ' XX., p. 388. Iddings identifies this mineral to 
thermophyllite, a foliated mineral having the composition of serpentine. 

2) *The Geology of (Jarmelo Bay.' Bulletin of the Department of Geology in the 
University of California, Vol. I., p. <M. See also Pirssou's paper, Amer. Journ. Sci., XLV, 
1893, p. 381. 

3) * La Chaine des Puys et le Mont Dore,' BvU, G6oi. Soc. Fmnce^ 3me Serie, XVIU, 
1890. 



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OF THE DEENDENT ISLES OF TAIWAN. 21 

chemical composition of the two alteration-products, yet on the 
whole they must be practically identical. The lamellae 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 L 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 b. Miigge^^ however, says that 
the absorption is stronger in the direction perpendicular to the 
* Ldngsrichtung ' than in that parallel to it. ZirkeP^ 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 b. 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 
piuacoid, 010 or 100, the lamellae 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 lamellae is greater (3l=i) 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, / am rather inclined to consider the iddingsite 
to be a mineral approaching to basite. Prof. Rosen busch^^, 

1) Neues Jahrhuch, 1883, II, S. 205. 

2) * Petrographie,* Bd. I, 1893, S. 353. 

3) * Physiographie,' Bd. I., 1892, S. 469. 

4) 'Physiographic,' 1892, Bd. I., S. 461. 



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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 befordert 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 uie 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 11^ 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. 



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OP THE DEPENDENT ISLES OP TAIWAN. 23 

is the case of glass-enclosures, save that ground-mass which fills 
up the rectangular space between the laraellee, showing as if the 
larger crystals have grown out by the apposition of numerous 
flowing lamellfiB. 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 [PI. /, Figs. 4 and 6., PL 11^ Fig. 5), a characteristic com- 
mon to all the plagioclases of Basalts. It seems more rea- 
Bonable 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 in- 
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- 
lamellse 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) 'Stndien oeber lockere vulcanische Auswurfiinge/ Zeittichr, </. d. geol. Qesdl. Bd. 
XXX, a 101. Taf. v., Pigs. 3, 5, and 7. 



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24 kot6 : notes ox 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 lamellae. Microlithic sections twinned on the albite 
type extinguish at the angles from 0° to 26°, with reference to 
the longer dimension. According to Michel-L^vy, 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. 



AUGITK 

Augite, so says Morozewicz^^ belongs to one of the * ver- 
hangnissvollen minerals. It does not obey Fouqu6 and M.-L6vy'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, i.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/ 
TachenrnJ^s MUth., Bd. XVIII., S. 84. 



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OP THE DEPENDENT ISLES OP 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 oo P «>, oo p(» , oo P, and Pob , 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 violei 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 KiA- 
shiti, and Chiu-goku, in Hondd, 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 titaniferous 
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 wdl-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 
ooPo) 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. 



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26 kot6 : notes on the geology 



HTPERSTHENE. 



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 modem 
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 Hatto, the only exception being 
the one from Sei-kei (West Valley) in Hoko, 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 J 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 //, 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 {PI, 11^ Fi^. 4), with only a few 



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OP 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 
aJBTord 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 basafl 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 Hdko Basalts stands in its chemical composition near to 
that of bronzile. 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 
pinacoidfl by ordinary methods. 



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28 kot6 : NOTES on the GfiotxKJir 

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 Hoko), 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-section. 

ANALCIME AND NATROLITE. 

A cave-rock in the southern Gio-6, presents an anomalous 
habit ; a slide made of it contains a colourless mineral in angular 
or polygonal interspaces between the crystals of plagioclase 
{PL 11^ 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 



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OF THfi DfiPENDfiNT lSt£8 Of tAlWAK. 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 11^ 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 Biiking*^ calls the 
^ Basis zweiter Arty 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 close 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) 'BasalUsche Gesteine, etc./ Jahrh, K, K. preuas. geol* LandeMnstall. 1880} S. 153^ and 
1881, S. 606. 

2) 'Ueber Monchiqaite, ein camptonitisches Ganggestein aus der Gefolgenschaft der 
Eleolithsyenite/ TBchermark's Min. MiUh. XI, 1890, S. 445. 

3) Proc. Cal. Acad. Sci., Vol. Ill, 1390. 

4) Cited in Pirsson's paper. 

5) *Trap Dikes,' Bull. 107, U. S. G. S. 1893. 

6) * On Analdte Diabase from San Luis Obispo County, California,' Bull. Geol. Depart. 
Univ. CaL, Vol. I. p. 27?. 

7) * An Analcite-Basalt from Colorado,' Joum. Geol. Vol. V. p. 684. 

8) *A new Analcite Rock from Lake Superior' Joum. Geol. Vol. VII, 1899, p. 422. 

9) *The Mochiquites or Analcite Group of Igneous Rocks,* Joum. GeoL, Vol. IV. 1896, 
p. 679. 



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30 kot6 : NOTES on the geology 

game 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 11^ 
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 
eflfusive period, as the iron ores were not found enclosed in the 
olivine of the intratelluric crystallisation. Both ores, especially 
the ilmentite, have crystallised later 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 th^ir 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 
linesi and cross several crystals of felspar and augite^ mean- 



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OP THE DEPENDENT ISLES OP 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 coane-cryBtalline type of intersertal, 
or ophitic structure, irrespective of hyperisthene 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 Hattd 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 V^nukoff'\ very abundantly in the Basalts of Mongolia, and 
even transparent lamellse were found by him, just as in the Pes- 
cadores rocks. The ilmenite is fresh and leucoxene not noticed. 

1) * Basalt Ton Bakony/ Zeiischr. d. d geoL Oea., XXIX., 1877, 8. 191. 

2) Bosenboflch, < Mikroekopische Physiographic,,' II., 3te Auflage, S. 1015. 

3) * L^ Roches Basaltiques de la Mongolie,' BulL Soc l^elge de Qido^Ut etc T. 11., p, 443. 



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32 kot6 : notes on the geology 

In my few slides of Basalts, bearing the iddinffsitised 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 isometric 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 H6ko, 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 Hoko 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 y consequently the skeletons inter- 
sect each other at the angles of 60° and 120°, and are said to 
consist of garnetohedrons. They all go into 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 odahedra, arranged rectilinearly 
in the direction of the crystallographic axes with secondary and 

1) ^ Experimeutelle Untersuchungen iiber die Bildang der Minerale im Magma,' 
Jkha-maf^a Mitiheilungm, 18, 1898| S. 90. 



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OF THE DEPENDENT JSLES OF TAIWAN. 33 

tertiary offshoots. This mode of growth, the octahedric dendriley 
so called by Morozewies, is well known in petrographical 
literature, since the publication of Prof. ZirkelV^ 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 Hoko, 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 i?, 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 witliin the rock, and that mineral es- 

1 ) * Die mikroskopische Beschafienheit der Mineralien und Gesteine.' Leipzig, 1873, S. 244. 

2) Collected at R^'6-l0-8an (JJ^Sllj) ; and, according to Mr. Saitft, it appears in I. horizon, 
t. f., the uppermost sheet, consequently the youngest of all the lavas of the H6ko Group 
(il/,/fi?. 1). 



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34 KOTO : NOTES ON THE GEOLOGY 

pfecially accumulates near the margin of the secretionary mass, 
the augite being sometimes arranged along the linear common 
base, with the free ends of the crystals 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, the 
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 ihnenite 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 iron-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 oi' the body by the magmatic 
menstruum proceeds alwa3'S 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 iddingsilizalion {Figs. 1 and 2). 

The ground'ma8$ consists, first of all, of the crystals and 



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OF THE DEPENDENT ISLES OF TAIWAN. 35 

grains of augite, all of a violel-hrovin 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 
glome roporphyri tic phenocryst. 

B. THE TYPE OF THE IDDINGSITE-BEARING BASALT, 
{PL /, Figs. 4, 5 and 6; PI. 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 features of this group are firstly, the pre- 
sence of large phenocrysts of olivine which is more or less iddlngsi- 
tized ; secondly, the majority of the augite is xenomorpliic 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 Hoko, one of 
which was struck off at the locality Tai-san ;^^ according to 
Y. Saito, it forms the uppermost flow there. The same may 
be said of the specimen from Kippai, since the youngest 

1) ^;^pj, ^^ySh iJ^BO- '^^^ J*ock eflervc-ces with acid. The micros'^ope discloses the 
fact that the radially arranged fibres of calcile fill up the jolygonal spaces between other 
components, showing bars which correspond to the position of crossed nicols. 



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36 KOTO : NOTES ON THE GEOLOGY 

lava-flow is the only effusive that can be met with on that 
island. 

The olivine is the sole phenocrysl; it is variable in size (the 
largest one measures even 5 mm.), irreguhir in distribution, and 
multifarious in form, some having partinl crystallographic faces, 
while others have none of them. The iddingssitization 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 seq. By the way, I have only to mention that the name 
iddiiigsite 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 [PL /, Fig. 6), which are prevalent 
in the group under question, the plaffioclases arc all approximately 
of the same size, and surpass the augite both in dimension and 
quantity; while in the augite-rich rocks {PL I, fig* 4), the 
phigioclases 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. 

The 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 phigioclase. The augite is, as usual, of a violet- 
brown colour, but in the specimen from Tai-san, it is almost 
colourless in sections. It is free from foreign inclusions, and 



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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 /, Fig. 6), but comparatively large, lamel- 
lar and flat with glittering bluish lustre on the perfect cleavage- 
surface. It looks rather more like ilmenite than magnetite. Stiff, 
slender a;)<7<i7e-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 plagioclasey which 
resolve at the ends into slightly diverging columns {PL /, Fig. 5), 
and these may be easily mistaken for those of apatite, if needles are 
found detached from the w^aist. Optical properties are not in- 
dependtly shown in them, on account of their extreme thinness. 
Similar bodies are noticed by H. S. 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., May, 1896, p. 3S0. 

2) ' Molecularphysik,' I, 1888, S. 378. 



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38 KOTO : NOTES ON THE GEOLOGY 

needles may possibly of a felspathic nature and such a structure 
is termed xariolitio 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, 
rany 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\ TUE OPHITIG TYPE. 

{PI II, Fig. 2) 

This type is represented by a single specimen from Hoko, 
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 of a 
certain thickness. It is a greyish-black, Anamesitic rock, with the 
brownish, lath-shaped phenocrysts of plagioclase (4 mm. length). 
This is the coarsest type of the Hoko 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- 
talliue, 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 tlie rock (Basalt)." 

Petrology for Students.' 2nd. Edit., p. 191 and 201, Fig. 41 A, 

2) The exact locality being Sho-chi-kaku, ('hft,^) at the middle of the island. 



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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 Mg. 2) 
filled with films of brown hydrous sesquixide of iron, which 
cause the phenocrystic feldspar to appear viacroscopically 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 lamellae of ilmenite 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 inlersertal. Magnetite seems to 
be wanting. Owing to the coarseness of the structure, the rocks 
are often porotiSf and the polygonal, angular spaces are often 
filled up with banded, purplish chalcedony. 

D. THE TYPE OF THE OLIVINE-LESS BASALTS. 
{PI. /, Fig. 3 ; PI. H, Figs^. 2 and 3.) 

The olivine-less, hypersthene-bearing Basalts are represent- 
ed in my collection by two specimens from Wampai'^, and one 
Irom each of the following islands, H6ko^\ Kin-sho^\ and 
Hatto-sho'*^ They are megascopically wet-grey, and fine-granular, 

l)i«M 2) Sei-kei n^ in HOko '3)~^i« 4yA» S^ 



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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 fframi- 
litic. 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 he easily recognised as such in the present group. 

The phenocry Stic plagioclase is narrow-tabular with a few twin- 
ning lamella? (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 07ie lamella^ out of many twin- 
ned parts after the pericline law, is provided with closely 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 augite 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 Hoko, which also lies not very far from the 
present sea-shore. 

Hyperthene occurs exclusively, though insignificant in quan- 
tity, in the form of phenocryst {PI. II, 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, 



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OP THE DEPENDENT ISLES OP 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 fissures. 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 augiie sometimes makes its appearance in 
company with the hypersthene and plagioclase, forming local 
patches of secretional origin, with the hyperilic structure. 

The ffround-masSf 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 graniditic. The 
augites of both generations are of yellowish brown and not violet- 
brown. 

Shingly tridymite 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 /, Fiff. 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. 



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42 KOTd : NOTES ON THE GEOLOGY 

E. THE TYPE OF THE ANALCJME-BASALTS. 
{PL Jl Fig. 5.) 

This to the naked eye is macroscopically deep-greyy and fine- 
granular. Under the microscope it is hypocrystalline and more or 
less porphyritic, either the xenomorphic olivine or the aggregate of 
the autoraorphic 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 geiieral" 
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 fehpar- 
phenocryst. 

The ground-mass consists of laths of plagioclase, crystals 



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OP THE DEPENDENT ISLES OF TAIWAN. 4.S 

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 {FL Ily Fig. 5) produced by the shifting of lamellae 
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 lamellse 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 
lamellse usually in association with the hexahedral iron-ore. The 
lamellae are anisotropic, and distinctly pleochroic, and the 
mineral is conjectured to be Umenite. 

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 nairoUte (as in the left, lower margin in Fig. 5). The 



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44 Kord: notes ox 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 Hoko 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 basicy 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 
Na20'Al203*4 Si02'2H20. 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 H6ko Group, but in the Teschenite 
of the Nemuro promontory in Hokkaido, 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 jyresent mode of occurrence of the analcime 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 a^s those of plagioclase, augite, 

1) * The Monchiquite or Analcite Group of Igneous Kocks.' Journ. Geol., Vol. IV., 1896, 
p. e70. 



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OF THE DEPENDENT ISLES OF TAIWAN. 45 

magnetite, apatite, together with the olivine had then sunk down, 
and formed the heap of O'ystals at the bottoniy meanwhile the un- 
consolidated residuum of the rnagm^ wa^ actually slowly flowing 
through the sieves of crystal-heap, or changed its chemical com- 
positio7i through diffusion, after the manner of liquation a^ 
in a metallurgical p<rocess. 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 Hoko. The hand- 
specimen from Nai-an^^ in Gio-6, is, according to Y. Saito, 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 Hoko Group no doubt belong 
to the same horizon. 



1) Nai-an ftS, mMA- 



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46 kot6 : notes on the geology 



THE ISLAND Olf 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 Islindanao, 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 

i)iBB|. 

2) B. Kotd, * On the Geologic Structure of the Malayan Archipelago.* Tliis Journal, 
vol. XI, pages 111 and 118. Wichmann calls the chain the * North Moluccan bow.* *Der 
Wawani auf Amboina und seine angeblichen Ausbriiche, III.' Tijdachr. r. h. Kon. Nederl 
Gen., Jaargang 1899, S. 32. This bow is now said to start from Batjan, lying to the south 
of Makjan. loe, cit. S. 14. 

3) Kotd, loe, cU, p. 118. 

4) The Japan Mail, August 10th 1897, * Forlorn isles.* 



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OP 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 Qadd and 
Forest Belle, the islands of Sh6-K6t6 (Little Botel-Tobago) and 
Koto (Botel-Tobago), and, lastly, Kasho (Samasana), as the conti- 
nuation of, I conjecture, the Mayon system of volcanoes {Fiff. 1). 

The smaller isle of K6t6 is, geologically speaking, entirely 
unknown, but the Larger K6t6 has been several times visited 
by the Japanese, since the first landing of a staff of the gover- 
norship of Taiwan, in April, 1897. Aniong 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 of the Larger K6t6 {Mg. 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 ri and the breadth IJ n, 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. 



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48 kot6 : 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 are not the reefs of Neocene time, which 
usually attain a considerable height of more than 200 m., as in the 
Apes Hill of Takao, but those of a comparatively recent date, 
possibly representing a Diluvial 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 {Mg. 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 



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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 j^tco/tfe, mixed with the crystals 
and dust of magnetite. 

Plagiocla^ey 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 augile 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 yeUowish-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 ffroufid-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 poly synthetically, 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 ohseryed.hy E. Elich in the Amphibole-pyroxene An- 
desite from the Rio Blanco, West Cordillera, Ecuador. Reiss u. Stiibel, * Reisen in Sud- 
Amerika, Das Hochgebirge der Repnbik Ecuador, I.; Petrographische Untersuchungen, L 
West Cordmere,' S. 163. 



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50 kot6 : 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 is 
evident that the Basalt of the Island of Kdtd does not properly 
belong to the category of normal Basalt with violet augite^ present- 
ing the intersertal structure y which is so common in the rocks of 
the Hoko 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 lelspathic, 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. The 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. 
(PI. Ill, Fig. 5.) 

I have three specimens of rocks in Torii's collection, belong- 
ing to the same category. They (iiffer 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). 



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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, pa^;allel 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 gldss-enclosures with bubbles. 

Augite occurs spoj'odically 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- 



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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 recta ngularity 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 ' orthophyricJ In 
the ? variety, minute felspar-needles make the greater part of 
the ground-mass, exhibiting the typical pilotaxitic structure, 

C. APOANDESITES. 

{PL 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, Amor, joui: Sci,, May, 1898. 



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OP 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 Inrge black spot in Fig, 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 composition 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 21. 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-lamellse, 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- A ndesite, already described, by 
the pneumatolytic process which caused the impregnation of the 
pyrites in the rock-mass. 



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54 kot6 : notes on the geology 

D. AMPHIBOLIZED GABDRO. 

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 lamellae of chlorite, 
calcite-films, and common epidote. The hornblende has been so 
highly altered that the original substance remains but in few 
stripes. The pldgioclase-anhedrsi possess only a few twinned 
lamellae, 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 apaliley full of air-pores, was only 
once observed. 

I conjecture this rock to be a metagdbh^Oj 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. 



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OF THE DEPENDENT ISLES OP TAIWAN. 55 

THE ISLE OF KASHO (SA3IASANA). 

(PL III, Fig. 6.) 

Kasho 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-homblende-AndesUe. 

To the naked eye the rock resembles very closely those of 
H^radak^, 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. The 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 



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56 kot6 : kotes ox the geology 

the magma, so that there remains nothing hot the accumulation 
of magnetite-dust in the place of the hornhlende. 

A hrown 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 lamellae makes -5° to -10° on 010 with P/M, 
indicating the presence of labradorite. The albite-lamallse are 
clear and definite, but the width varies much from one lamella 
to another, and even in the same the widtli 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 raicro-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. 



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OF THE DEPENDENT ISLES OF TAIWAN. 57 



CONTENTS. 



Page* 

The Hdko Group (Pencadores) 1 

I. Introductory 1 

II. Stratigraphical Charjicteristics 5 

H6ko Isiafid 5 

General outlines of the ;;eology of the island 5 

Local detaiU of geology 8 

Uaku'sha Inland 12 

Kippai Island 13 

Gi<h6 Island 14 

III. Petrography of the Effusives 17 

A, Component-minerals of Basalts 18 

Olivine 18 

Plagioclase 22 

Augite 24 

Hypersthene 26 

Analcime and Natrolile 28 

The Iron Ores 30 

B, Special Description of Individual Occurrence of J3asalts 33 

a. The granual type 33 

b. The type of the Iddingsite-bearing Basalts 35 

c. The ophitic type 38 

d. The type of the Oli vine-less Basalts 39 

«. The type of the Analcite-bearing Basnlts 42 

The Island of K6t6 (Rotel-Tobaso) 46 

a. Felspar-Basalts 48 

b. Hornblende-Andesites 50 

c. Apoandesites 52 

d. Amphibolized Gabbro 54 

c» oerpenuue .»• ••• ••• ••• ••• ••• .«• ••• ••• .•• «•• o*x 

The Isle of Kashd (Samasana) 55 



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PLATE I, 

(PUOTOGRAIS.) 



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PLATE I. 



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 granuUtic, B6-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. Hoko 
Island. P. 33. 

Fig. 3. — Olivine-less basalt from Ilatto, Southern Group, 
and it probably belongs to the same type as Figs. 3 and 4 in 
Plate II. 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 Go diame- 
ters. Hoko Island. P. 35. 

Fig. o. — Rock belonging to the same type as the preceding. 
It is also from Hoko Island. Olivine on the left side of the 
figure shows various stages of iddingsitization. 

Fig. G. — 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 Go, as is stated in the 
Plate. Nicols crossed. Kippai Island. 



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KotOf The Hoko Group, etc. 



Jour. Sc. Coll. Vol. XIII. PI. I. 





Fig. L X ^4 



Fig. ^. X ^4 





Fig. 3. X 65 



Fig, 4. X 65 




Fig. 5. X 38 




Fig. 6, x65 + nicols 



Koto pilot. 



,QQPgI^ 



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PLATE II. 

(PHOTOGRAHS.) 



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PLATE II. 



Fig. 1. — Iddingsite-bearing andesite, magnified Go diameters, 
showing the typical intersertal structure. Olivine is here changed 
internally into a red mineral, which the writer believes to he 
iddingsite, as is well seen on the lower right octant in the 
figure (pp. 19 and 3o). Kippai Island. 

Fig. 2.— The slide of ophitic basalt (p. 38). Sho-chi-kaku, 
the Island of Hoko. 

Fig. 3. — Olivine-less hyperstliene- bearing basalt, with two 
large crystals of hyperthene in the centre of the figure. The 
structure is granulitic. The Isle of Wam-pai. P. 31). 

Fig. 4. — The same rock-tyjx) as the preceding, but with 
intersertal structure. Local patches of hypersthene, augite and 
plagioclase, with the hyperitic structure. Sei-kei, the Island of 
Hoko. Pp. 39 and 41. 

Fig. o. — Analcime-basalt from Nai-an, Gio-6. It has granu- 
litic 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 Calcarina Spengleriy besides fragments of 
corals, bivalves and serpula. In natural size. Kippai Island. 
P. 13. 



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Koto, The Hoko Groups etc. 



Jour. Sc. Coll. Vol. XIII. PI. II. 





Fig. 1. xod 



Fig. 2. X 24 





Fig. H. X 24 



Fig. 4. XS8 •\-nicols 




Fig. 5. X 66 




Kotd phot. 



Fig. 0. Nat. size 

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, Tokyo Printing Co., Lm> 



imp. Tokyo Printing < 



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PLATE III. 

(PHOTOGRAMS.) 



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PLATE III. 



The Plate III illustrates the type-rocks from the Isles of 
Koto (Botel-Tobago), and Kasho (Samasana). 

Fig. 1. — Apo or altered andesite, magnified Go diameters, 
showing a phenoerystic 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-crystiils, 
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. G. — Hypersthene-hornblende-andesite from the Isle of 
Kasho, 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. ob). 



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Koto, The Hoko Group, etc. 



Jour. Sc. Coll. Vol. XIII. PI. III. 





Fig, 1. X do 



Fig. '4. y.HH -{-nicols 





Fig, 3. X S8 



Fig. 4. X 38 




Fig. fj nm 




Kot6 phot. 



Fif/,e. XJ3 



m 



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PLATE IV. 

(MAP.) 



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PLATE IV. 



The Plate IV sliows the bathometric condition of the neigh- 
bouring seas of the Pescadores or Hoko 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 tlie east from 
Taiwan by a channel of the same name (p. 3). 



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THE HOKO GROUP. 

Kolojhe Hoko Group, etc. Jour. Sc. Coll. Vol. XI. PI. IV. 



Kiiir 



Xortli (iroij 



\ 



Soiilli (h-oup 




nt-shn J*f /> . *^ ^ S€^i^ ^tikif^l 



\%^i/^ 



^ 



r^z-j^^ 



Formosa Ii€f^iA' 



/ 



\ « 



ttrK.lmmf. 



ZO'B.rmujf, 



J 10-20 [ ? 120-30 1 ^ lan-io t ^ I 

Sralo 1.680,000. 



^MFniMmms 



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PLATE V. 

(GEOLOGICAL MAP.) 



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PLATE V. 



The Plate V is intended to show tlie 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. KSait6, who 
also offered me assistance in colouring the geologic elements 
represented on the map. 



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Koto, The !• 



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Change of Volume and of Length in Iron, Steel, 
and Nickel Ovoids by Magnetization. 

By 

H. Nagaoka, RigakuhakusM, 
Professor of Applied Mathematics. 

AND 

E. Honda, Rigakushi, 
Post-graduate in Physics. 

With Plates VI. & VIL 

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 hundred 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 CoUege of Science, 9, 353, 1898 ; Phil. Mag. 
46, 262, 1898. 



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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, BidwelP^ 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 0.56j2, 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. 63, 487, 1894.; Nagaoka and 
Honda loc. cit. 



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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 experimejits : 



Specimen 
No. 


Metal 


a (cm.) 


c (cm.) 


v(c.cm.) 


P 


N 


1 


Nickel 


0.750 


12.50 


31.50 


8.86 


0.125 


2 


9f 


0.500 


10.00 


10.48 


8.86 


0.0848 


3 


Soft iron 


0.750 


12.50 


31.45 


7.84 


0.125 


4 


7f 


0.500 


10.00 


10.53 


7.83 


0.0848 


5 


Ordinary steel 


0.750 


12.50 


31.60 


7.83 


0.125 


6 


if 


0.500 


10.00 


10.57 


7.81 


0.0848 


7 


Wolfram steel 


0.750 


12.50 


31.82 


7.90 


0.125 


8 


)) 


0.500 


10.00 


10.53 


7.95 


0.0848 



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 
sanie specimens as the ovoids. The prisms were 14.6 cm. long 
^d 0.896 cm. square in cross-section. 



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60 



H. NAGAOEA AND K. HONDA. 



Metal 


E (O.G.S.) 


A(O.G.S.) 


e 


Nickel 

Soft iron 

Steel 

Wolfram steel 


2.OTX10'* 
2.10x10'* 
2.04x10"* 
2.02 X 10'* 


0.771 X 10'* 
0.800x10'* 
0.838x10'* 
0.849x10'* 


1.082 
0.844 
0.384 
0.306 



E gives YouDg's modulus, A' {= n. Thomson and Tait) the 
moduhis of rigidity, and a constant defined by the equation 

2 V l + 3<? /-'*• 
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) 


H 


I 


H 


I 


H 


I 


H 


I 


0.7 


24.2 


1.0 


62 


1.9 


23 


2.7 


18 


1.4 


49.8 


2.5 


160 


2.4 


44 


6.8 


65 


3.0 


138.6 


4.3 


291 


6.9 


183 


12.6 


193 


5.4 


238.0 


9.5 


587 


9.7 


279 


20.2 


498 


10.9 


336.8 


12.7 


750 


13.1 


385 


25.8 


748 


37.8 


395.7 


19.9 


948 


23.3 


651 


44.5 


992 


74.1 


420.0 


37.2 


1111 


32.3 


815 


83.6 


1116 


125.3 


434.5 


99.6 


1255 


50.2 


984 


118.0 


1170 


171.6 


438.7 


155.5 


1309 


116.3 


1196 


191.0 


1224 


240.3 


440.7 


270.3 


1400 


174.4 


1260 


344.6 


1301 


481.4 


443.4 


433.6 


1479 


345.0 


1379 


512.3 


1348 


674.2 


444.5 


584.6 


1520 


520.2 


1440 


666.6 


1373 


914.0 


446.8 


792.8 


1546 


873.7 


1489 


940.3 


1400 


1233.0 


447.7 


992.6 


1562 


1149.8 


1512 


1213.3 


J 423 


1747.0 


448.7 


1585.8 


1607 


1822.6 


1549 


1674.9 


1452 



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CHANGE OF VOLUME AND OP 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 ^ooooo * '^^^ 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 r^ards 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 



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62 H. NAGAOKA. AND K. HONDA. 

at jSrst 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 B. Accad. dei Linoei 6, 487, 1891. 



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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 9^ of iron, besides traces of man- 
ganese and carbon. The ovoids used in the present experiment 

1) Knott, Trans. Boy. Soc. Edinb. 38, 527, 1896; 39, 457, 1898. 



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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 B. Accad. die Llnoei, 6, (l)f 257, 1891. 



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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. 

i? I The observed changes of volume and of length are exhibited 

g: I iu the following table : — 



l)Gmtone, Atti della R. Accad. dei Lincei, 6, (1), 257, 1891. 



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C6 



H. NAGAOKA AND K. HONDA. 



Nickel (1) 
H '." 


Soft 
H 


iron (3) 

At 

r 


Steel (5) 


Wolfram steel (8) j 


i " 




H 


V 


13 


0.09 X lO' 


8 


0.10 xio' 


7 


0.08 X lo' 


19 


0..30xl0 


.30 


0.29 


11 


0.52 


12 


0.47 


42 


l..')2 


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 


958 


5.88 


792 


8.01 


1001 


16.68 


1141 


2.44 


1115 


7.18 


1044 


10.16 


1117 


18.96 


1547 


3.24 


1342 


9.47 


1376 


14.07 


1296 


22.75 


1740 


3..53 


1563 


11.45 


1646 


17.20 


1704 


28.82 


2253 


4.12 


2089 


14.68 


2171 


22.20 


21.53 


32.62 



N 


ickel (2) 

SI 

i 
- I4.1xfd 


Soft iron (4) 
H , 

6 2.5x10 


Steel (6) 


Wolfrara steel (8) 
18 4.1x10 


H 

4 


H 


5/ 


13 


3.1 X 10 


6 


- 64.0 


15 


19.0 


19 


7.1 


25 


12.4 


10 


-118.2 


51 


31.6 


28 


15.1 


39 


21.7 


15 


-16.3.6 


127 


23.8 


54 


22.2 


62 


28.9 


33 


-217.5 


224 


3.1 


96 


23.1 


106 


32.1 


59 


-264.3 


354 


-17.7 


160 


17.8 


170 


32.3 


124 


-317.6 


575 


- 52.6 


225 


8.2 


235 


31.7 


302 


-343.6 


698 


- 62.6 


374 


-11.5 


349 


30.2 


561 


-3.53.8 


883 


- 73.5 


589 


-36.0 


592 


23.1 


839 


-356.0 


1077 


- 78.9 


844 


-49.2 


781 


18.7 


1145 


-360.0 


1180 


- 82.2 


1061 


—5.5,5 


1052 


17.0 


1289 


-360.9 


1324 


- 86.6 


1177 


-59.5 


1188 


15.4 


1483 


-362.2 


1447 


- 89.9 


1361 


-64.5 


1345 


13.9 


1849 


-362.7 


1538 


- 91.6 


1729 


-69.9 


1697 


12.4 


2322 


-36.5.3 


2180 


-102.0 


2172 


-78.1 , 


2235 


10.9 



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CHANGE OF VOLUME AKD OF LEKGTH. 



67 



KirchhofTs Constants ^ and ^ 



jf 



13. 



and 



Starting from the formulfle 
V I 



A7:¥ (J±0\ 

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4i 



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IP 



4 4 5 A'(l + 3<;>' 

which give the change of volume and of length of ferromagnetic 
ovoids in terms of KirchhoS's constants k' and //', we obtain the 
following expressions for these two constants : 

2(l + 3tf) ' 
'3q-p 



y= 



and 7c" = 



2(1 + 3</) ' 



where 
and 



= _±^(l + 3^.+4-r^^+3^-, 



H 



, = -^^t^A + -^(l + ^)+/. 



These constants, as calculated from the change of dimensions 
of ovoids, are given in the following table, and graphically drawn 
in Fig. 3 : 



H - 


Nickel 


W 


ISoft 


iron 


Steel 
k' //' 1 


Wolfram steel 
343 - 1-252 


5 


-229100 


712800 


21900 


-22610 


1017 


-1865 


10 


-188900 


578900 


23520 


-28450 


3840 


-3322 


986 


-1512 


20 


- 71000 


216700 


132S0 


-16420 


4248 


-4615 


3600 


-3983 


30 1 


- 36370 


111200 


7302 


- 8050 


4048 


-5080 


4881 


-5440 


60 I 


- 8163 


34540 


21S9 


— 2222 


1738 


-1864 


1946 


-2217 


80 1 


- 6906 . 


20960 


1207 


- 1102 


1069 


-1004 


1198 


-1385 


100 


- 4053 


14120 


753 


- 550 


701 


- 546 


794 


- 880 


120 


- 3373 


10260 


500 


- 255 


477 


- 279 


557 


- 595 


160 ' 


- 1297 


3968 


239 


18 


240 


- 16 


315 


- 317 


250 ! 


- 843 


2553 


55 


175 


69 


117 


128 


- 109 


300 


- 591 


1790 


25 


175 


38 


124 


88 


- 66 


500 


- 216 


655 


-9 


130 


-1 


99 


28 


- 8 


800 


86 


259 


-9 


70 


-6 


57 


8 


5 


1200 


- 39 


117 


-6 


37 


-4 


31 


3 


5 


1600 


- 22 


66 


-4 


23 


-3 


19 


1 


4 


2000 


14 


42 


-3 


16 


-2 


13 





3 



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68 H. NAGAOKA AKD K. HONDA. 

14. The curves for i' and i" present the same general feature 
in iron and steel. // 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 i' ultimately coincides with the axis of I£. h" 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 h" 
decreases very slowly. The position of maximum for Ic and that of 
minimum for h" 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 k' and //' 
is greater in iron than in steel. In nickel, the values of // and h" 
are far greater than those for iron and steel, and moreover are of 
opposite signs. The maximum of h'\ or the minimum of t', seems 
to lie in a weak field ; the rate of decrease or increase is quite 
rapid and the curves for Ic and h" soon approach the axis of//. 
Compared with the results of former experiments, the absolute 
values of Tc and h" 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 



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CHANGE OF VOLUME AND OF LENGTH. 



69 



8I = H[l^^-3(l^+-^;-)y. 

Putting ^=4.67x 10-«, 4.80x I0-^ 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 : 



1 " ' 


Soft iron. 


Steel. 


Wolfram steel. 


oI 


oY 


61 


10 


0.919 


0.055 


(\044 


20 


1.074 


0.212 


0.170 


30 


0.831 


0.402 


0.350 


GO 


0.399 ■ 


0.254 


0.294 


80 


0.244 


0.1.53 


0.249 


100 


0.127 


0.072 


0.188 


120 


0.039 


0.005 


0.145 


160 


-0.080 


-0.092 


0.094 


200 


-0.164 


-0.146 


0.061 


300 


-0.258 


-0.210 


0.017 


500 


-0.311 


-0.236 


-0.002 


800 


-0.275 


-0.205 


-0.038 


1200 


-0.219 


-0.163 


-0.037 


1600 


-0.183 


-0.134 


-0.033 


1 2OO0 


-0.157 


-0.115 


-0.028 



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, X^ 



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70 



H. NAGAOKA AND K. HONDA. 



4.74x 10-^ wLich corresponds to a pull of 0.1 Kilog. per sq, mm. 



H 


dl 


10 


-24.58 


20 


-18.87 


30 


-14.16 


60 


- 9.09 


80 


- 7.12 


100 


- 5.99 


120 


- 5.22 


160 


- 2.70 


300 


- 2.28 


500 


- 1.39 


800 


- 0.88 


1200 


- 0.60 


1600 


- 0.45 


2000 


- 0.36 



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 dl due to change of volume a by 
hydrostatic pressure is given by 



3l=^ih'^J(pjHa. 



If we calculate the change of magnetization due to contrac- 
tions 4.68 X 10"', 5,38 x 10"', 8,42 x 10^ 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 : 



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CHANGE OP VOLUME AND OP LENGTH. 



71 






H 


Nickel 


Soft iron 


Steel 


Wolfram steel 


SI 


SI 


SI 


SI 





0.000 


0.000 


0.000 


0.000 


10 


0.190 


0.757 


0.230 


0.045 


20 


0.119 


0.840 


0.4.'>7 


0.237 


30 


0.080 


0.713 


0.595 


0.8.58 


60 


0.037 


0.398 


0.565 


0.675 


80 


0.030 


0.362 


0.495 


0.550 


100 


0.024 


0.305 


0.437 


0.467 


200 


0.012 


0.178 


0.268 


0.259 


300 


0.008 


0.135 


0.200 


0.185 


500 


0.005 


0.093 


0.134 


0.118 


800 


0.002 


0.062 


0.088 


0.073 


1200 


0.001 


0.042 


0.061 


0.048 


1600 


0.000 


0.031 


0.043 


0.034 


2000 


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 

tbe 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 

eJ^periment, we found that hydrostatic pressure increases the 

°J«gnetization in nickel, while it decreases it in iron. The 

^^''eetnent between theory and experiment is very close in nickel, 

^^^ there is a wide discrepancy in iron and steel, as we have 

^'ready noticed. 

17. Effect of torsion on longitudinally or circularly magnetiz- 
^^ vnre. There are other important consequences to be drawn 
^^^ the constant h" with regard to the effect of torsion on 



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72 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. 

du - 



dx 
dv 



= -l(or, 



9y 



where (o 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 —Ict/rV^H at a distance r from 
the axis, the mean circular magnetization being ^-coh'^ERy 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 —h"Ii. 

Next suppose that the wire is traversed by an electric current 
of intensity 0. Then the circular magnetizing force at a distance 
r from the axis is 

E^^~ 

By applying similar reasoning, we find that the mean longitudi- 
nal magnetization is equal to - loV C , We therefore conclude 
that twisting the wire carrying the electric current gives rise to 
longitudinal magnetization proportional to —V C . Thus the 
circular magnetization produced by twisting a longitudinally 



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CHANGE OP VOLUME AND OF LENGTH. 73 

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 oeolotropic suscepti- 
bility is similar to what would follow from Kirchhoft'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 
1 from the curves of —h" 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 
I 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 Iheoreti'^hen Phynik, 2, 200, 1896, Leipzig; Drude, Wied. Ann. 
63, 8, 1897. 

2) Ewing, Proe. Bot;, Soc. 36, 1884. 

3) Wiedemann, EledricUatf 3. 

4) Zehnder,\Wied. Ann., 38, 68, 1889. 

6) Nagaoka, Phil. Ma^. [5] 29, 123,1890; Journal of the College of Science, TGkyo, 3, 
335, 1890. 



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74 H. XAGAOKA AVD K. HOXDA. 

these two metals lins 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 statcd.*^^ 

18. Looking at the curves of //'ZT, 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. AYe 
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 spile 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 thati' 



1) Knott, Trans. Roy. Soc. Edinb., 36, 485, 1891. 

2) Matteucci, Annales de Chimie et de Physique, 1858. 

o) Wliile tliis paper was passing t!»rough the press, we found that the direction of tlie 
transient current produced by twisting a magnetized iron wire is reversed in strong magneti- 
zing fields. 



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CHANGE OF VOLUME AND OF LENGTH, 75 



and //' are physically functions of the strain, as is borne out by 
the numerous experiments on the effect of stress on magnetization. 
The present slate of the theory of magnetostriction may perhaps 
be connpared with that stage in the history of the theory of 
magnetization when the intensity of magnetization was supposed 
to be feimply proportional to the magnetizing force* Tii f\iet, the 
theory is still in its infancy, so that there are ample grounds 
for expeciing further developments on further researches. 



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Jour. ScColl. Vol. XIJI. PI. VI. 




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U 



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Joun Sc.Coll. Vol XI 11. PL VIL 



111 1 i ! r 




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I 



III 






5 

.1 



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Combined Effect of Longitudinal and Circular 

Magnetizations on the Dimensions of 

Iron, Steel and Nickel Tubes. 

By 

K. Honda, Rigahushi, 

Post-graduate in Pliysics. 
With Plat€8 VIIL 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 1 J convolutions. 
His experiment was modified by BidwelP^ 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 the external 
change of volume for iron, steel, nickel, and cobalt tubes, Knott^^ 

1) Joule, Scientific papers I, 263. 

2) Bidwell, Proc. Roy. Soc. 56, 94, 1895. 

3) Knott, Trans. Boy. Soc. 89, 457, 1898. 



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78 



K. HONDA. 



II 



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 w^as 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, T is the tube to be tested, jPand 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. F in the lower part of the figure shows 



F^ T F 



R 



p 

=0 



1) Beatson, Archives des. Sc. phys. et nat 2, 113, 1846. 

2) Righi, Mera. 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. 



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CHA^^GJE OF DIMENSIONS BV MAG^J^ETIZATIOX. 79 

the front view of these rings. R and K 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 : 



^ Hil'l'h 



X 



r-ii 



coil 



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80 



K. HONDA. 



material 


length, 
(cm.) 


external 
diam. (cm.) 


internal 
diam. (cm.) 


demagnetizing 
factor. 


nickel 

wolfram steel 

soft iron 


17.02 
20.30 
16.97 


1.328 
1.124 

0.966 


1.252 

1.048 
0.842 


0.0261 
0.0162 
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. 



h 



Sesults of Experiments. 

1. Nickel Tube. 

4. The lube 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 diflference 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 9^. 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. 



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CHANGB OF Dimensions by magn^itization. 



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 
tbe 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- 
tudmal 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=0 


H= 


= 6.9 


H=22.1 


H= 


182.9 


h A X 10' 


h 


^xlO' 


h -^xlO^ 


h 


:^xlO^ 


7.6 4.4 


8.2 


2.7 


8.2 0.5 


8.5 


1.6 


16.1 19.0 


17.5 


40.8 


17.5 18.0 


13.6 


2.2 


26.3 46.2 


31.1 


87.1 


31.2 76.2 


21.3 


4.4 


40.6 75.1 


40.9 


110.4 


40.9 124.0 


31.5 


9.2 


50.5 92.5 


49.9 


130.6 


50.2 157.8 


49.0 


27.2 


64.5 108.8 


57.5 


141.4 


64.2 190.4 


64.2 


59.9 


72.1 117.0 


72.1 


163.2 


70.9 201.3 


69.5 


81.6 


83.2 119.7 


81.7 


168.6 


78.8 217.7 


87.6 


130.6 



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82 K. HONDA. 

Here H and h denote effective longitudinal and circular 
fields respectively, both in C.G.S. units. -V 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 througli 
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 



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CHANGE OF DIMENSIONS BY MAGNETIZATION. 



83 



the observation was finished, the key was reversed, so that the 
circular magnetizing 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. 



11 



1) Na^ka and Honda, Preceding paper. 



Digitized by 



Gi 



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 II. 



h 


=0 


h= 


=10.7 


h = 


= 16.8 


h= 


=22.9 


H 


-?^xlO' 


H 


^xW 


H 


^- X 10' 


H 


-^xlO' 


5.3 


- 1.5 


6.9 


- 10.9 


8.1 


- 17.6 


7.2 


- 13.0 


8.5 


- 25.6 


14.6 


- 61.7 


17.8 


- 73.5 


14.2 


- 49.4 


17.0 


- 71.7 


27.8 


-118.1 


27.4 


-118.1 


22.3 


- 93.9 


29.4 


-107.6 


42.7 


-161.6 


46.7 


-174.9 


37.6 


-149.2 


41.2 


-134.7 


62.8 


-198.6 


71.9 


-223.4 


65.1 


-198.7 


61.0 


-16.3.9 


103.3 


-238.4 






94.0 


-232.7 


101.3 


-202.4 


131.6 


-255.2 


131.9 


-271.0 


129.7 


-269.3 


184.9 


-241.8 


176.8 


-279.2 


177.7 


-291.5 


175.2 


-29.3.4 


274.9 


-261.3 


254.0 


-301.7 


255.4 


-313.7 


255.5 


-317.0 


354.9 


-274.1 


361.0 


-312.9 


363 4 


-329.7 


359.3 


-333.2 


4C8.6 


-279.2 


505.0 


- 324.0 


516.1 


-339.4 


516.1 


-347.4 


709.2 


-289.5 


720.2 


-331.6 


779.0 


-344.5 


725.2 


-351.4 



1) Barrett, Phil. Mag. [4] 47, 51, 1874; Nature 26, 516 586, 1882; Beibl. 7. 201. 



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C?HAKGE OF DIMEJfSIONf? BY MAGNETTZATTON. 



85 



The comparison of Figs. 1 and 4 shows tliat the general 
eliaracter of the change of length is the same in these two 
ms^t except that the sign of the change is opposite. Hence 
similar remarks as in the former hold good in tlie present 
case. • 

In the experiment with nickel and cobalt wires traversed by 
an electric current, Eidwell found that the effect waa inmeasur- 
ably small. The discrepancy in nickel perhaps arises from the 
eSeet of temperature, which he did not take into account ; the 
tliiference 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 Bid well'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 favor 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 
circular magnetising current, a well known phenomenon. Even 
with such a weak current as we obtain from a single DanielFs 
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 



Goo< 



li 



w 



86 



K. HONDA. 



dl 



aH"" 



where a, ^9 and n are constants and H is assumed to be positive. 
The determination of these constants from the experimental 
curve gave the following results: 

«=5.18, /3=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 III. 



H 


-f (cal.) 


-« (exp.) 


10 


- 46x10-' 


- 40x10-' 


20 


- 81 


- 81 


30 


-108 


-109 


50 


-148 


-148 


80 


-185 


-185 


120 


-215 


-214 


150 


-230 


-229 


200 


-247 


-246 


250 


-258 


-258 


320 


-269 


-269 


400 


-278 


-278 


500 


-285 


-284 


600 


-292 


-289 


700 


-293 


-291 



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CHANGE OF Dimensions by magnetization. 



87 



Thus except in weak fields, the coincidence between these 
two is very close; the diflFerence does not amount to l^A. 
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 10' 


h 


-^xlO' 


h 


'/ X 10' 


h 


^ xlO' 


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 


- 2.1 


31.2 


- 0.9 


35.2 -20.2 


41.9 


-32.2 


41.9 


-14.6 


41.6 


- 5.2 


51.8 -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 


-25.8 


78.7 -27.9 


74.7 


-47.3 


747 


-48.1 


75.6 


-30.9 


99.5 -27.9 


88.6 


-49.4 


91.4 


-58.0 


88.6 


-40.0 



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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 Iloncia, loc. cit. 



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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- 
od then decreases very slowly. Barrett^^ did not find the 

the case of iron and cobalt. The upper curve shows 
I 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 

7. The results are reduced to the temperature of 



TABLE, V. 



=0 


h« 


10.8 


h= 


17.7 


h= 


25.8 


« X 10^ 


H 


^xlO" 


H 


-^- X 10' 


H 


-^ixlO' 


0.0 


in.3 


3.7 


14.5 


0.2 


11.3 


0.0 


8.3 


30.5 


18.3 


26.1 


11.9 


21.0 


2.1 


16.2 


37.6 


24.5 


31.9 


18.0 


31.2 


17.2 


27.3 


53.1 


35.8 


50.8 


37.2 


57.7 


. 41.8 


38.8 


84.6 


46.3 


75.7 


50.5 


83.7 


56.8 


42.9 


135.3 


52.4 


105.5 


54.5 


120.3 


67.6 


44.9 


182.5 


54.2 


168.4 


60.1 


165.8 


72.5 


43.3 


233.3 


55.0 


244.5 


63.7 


246.5 


75.2 


42.9 


323.5 


55.6 


350.0 


63.9 


351.5 


73.3 


42.5 


505.0 


54.2 


461.5 


63.6 


459.5 


72.3 


41.3 


70S.8 


52.4 


615.5 


62.8 


671.6 


71.0 



US tlielongitLuliiial magnetization combined with a constant 
one is always to increase the length which is first 




ti 



CiL 



Digitized by 



cm 



90 



K. HOHDA. 



shortened by the circular magoetizatlon. In weak longitudiDal 
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. Soft Iron Tube. 

13. The experiments of the change of length by circulai 
magnetization and of the effect of longitudinal field on the 
change of length led to the following results, which are graphi- 
cally shown in Fig. 8. The observations were taken at th( 
temperature of 18° C. 

TABLE VI. 



H=0 


H= 


=5.7 


H= 


=25.8 


H=67.6 


h *^ xlO' 


h 


^ xlO" 


h 


^ xlO^ 


h -^-xlO" 


5.3 - 7.8 


5.3 


- 4.2 


6.3 


- 0.0 


5.3 - 0.5 


14.0 -13.0 


13.8 


-11.9 


14.0 


- 5.2 


13.8 - 1.0 


21.4 -1.5.6 


20.7 


-16.6 


21.4 


-10.4 


21.0 - 4.2 


37.5 -15.6 


35,7 


-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 



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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 
viih 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 
f length at the temperatures of 18.7°C and 76.1® respectively. 
Vhen the key in the circuit of the circularly magnetizing coil 
Fas 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 
lie 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, 
lie general feature of the change of length is so well known 
liat farther remarks are unnecessary. It is only to be noticed 
liat 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) BidweU, Phil. Mag. 05, 228, 1894. 




Digitized by 






II 



92 



K. HONDA. 



II 

III 

i 



in weak fields and to increase it in strong ones. The field at 
which the temperature produces no eflfect 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 VII. 



h 


=0 


h= 


=5.7 


h= 


=9.2 


h= 


=26.2 


H 


-^xlO' 


H 


*/ xlO' 


H 


« xlO' 


H 


Ax 10' 


5.3 


1.1 


6.9 


4.7 


5.3 


1.1 


5.3 


1.7 


10.3 


12.8 


— 


— 


11.2 


17.2 


10.3 


9.6 


21.5 


17.1 


17.9 


22.4 


22.9 


29.6 


20.5 


22 7 


41.3 


19.2 


37.8 


28.8 


39.4 


37.4 


40.6 


35.6 


70.3 


19.2 


61.8 


32.1 


64.0 


42.3 


61.2 


40.0 


97.9 


17.1 


111.1 


31.0 


97.9 


42.2 


90.9 


41.1 


144.3 


11.1 


142.6 


27.8 


145.0 


38.8 


143.5 


38.2 


223.0 


3.6 


218.0 


21.4 


217.6 


31.0 


217.2 


34.5 


318.5 


- 4.9 


320.3 


12.4 


320.0 


20.2 


312.5 


2.^.3 


481.4 


-20.3 


490.0 


- 4.3 


493.8 


42 


475.0 


8.6 


697.0 


-32.5 


704.0 


-13.5 


684.2 


- 5.7 


647.0 


- 2.6 



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. 



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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 
Bid well 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 eflfect 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 u and v 
represent these two dilatations respectively, the volume change o 
is given by the formula <T=w+2r. 

Assuming the isotropy of our specimens, we find the 
calculation leads to the following results : 

TABLE VIII. 



H 


Nickel 


Wolfram steel 


Soft iron 


Jl 


tv 


Iv 




V 


V 


V 


10 


-18.5xl6' 


0.0 xio' 


- 9.6 xW 


20 


-21.0 


- 7.2 


-13.1 


30 


0.0 


-18.8 


-12.8 


40 


18.0 


-22.2 


-12.2 


60 


46.5 


-21.8 


- 7.7 


80 


.54.0 


-19.2 


- 1.9 


100 




-16.4 


1.5 



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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 10"^ 3.1 x 10"^ 
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 
seolotropy 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 seolotropy of the specimens that in weak 
fields, Bidweirs 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. 



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CHANGE OF DIMENSIONS BY MAGNETIZATION. 



95 



Gonoludiug Bemarks. 

17. From the experiment on the relation between magnetism 
and twistj Knott^^ concluded that the pure strahi 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 hu electric current, the chauge of 
length by magnetization must he greater than when there is no 
longitudinal current. Since the change of length for cobalt is 
scarcely aliected by tension, the same must also be the case for 
longitudinal current. The cousideration is partially verified by 
the experiment of Bid well and also by the present oue. 

The same phenomenon may also be more concisely explained 
ill the following manner- Suppose onr samples to be isotropic 
aiid to have no residual effect. Let ^ and i be two magnetic 
forcts acting longitudinally and circularly in two perpendicular 
diructions* When these two forces act simultaneously, we have 
a resultant force H\ this force occasions the change of dinicn- 
eions in our ferromagnetics- The dilatation in the direction of 
the resultant force, as well as that in the direction perpendicular 
to itj can be expressed by J{II) and F[H) respect! vely^ which 
are even functions of ^, To obtain the dilatation in the longi- 
tudinal direction, we have simply to coustruct 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 



1) Kaottj Triwu, Roy, Soc. EdinU 3fl» pL 11,^ 483. 



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;i 






i 



I 



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 i does not exceed 50 C. G. S. units, the eflfect of 
volume change on the change of length by combined action of 
I and t is negligibly small, for in these strong fields at which the 
change of volume is pronounced, the ratio f/H^ in the above 
expression becomes 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 — ^ becomes, 
in all cases, 



Atl). 



L H- 

Since the material is supposed to be isotropic, fiJS) 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 —j- the ex- 
pression F{1) for the change of length by longitudinal magne- 
tization with a constant circular field i, and/(/) for the reciprocal 
case. 

Assuming for the expression f{I£) a suitable empirical for- 
mula for iron, steel or nickel, a simple analytical discussion of 



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CHANGE OF DIMENSIONS BY MAGNETISATION. 97 

the expression —j-y or numerical calculation of it for different 
values of / and t from the experimental curve of the ordinary 
change of length leads to the conclusion that for iron, steel 
and nickel, all the points, v\rhich 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 —f-- 

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 mngnetizations, 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 
iocrease 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 : 



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.:i|.i 



i 



98 



K- UONDA. 



TABLE IX. 





Nickel Tube, t=10.7 


diSerence 


I 


L' (cal.) i 


L' (exp.) 


10 


- 20x10 


- 25x10 


5 X fo 


2J 


- 74 


- 85 


11 


30 


-112 


-125 


13 


50 


-162 


-175 


13 


80 


-204 


-216 


13 


120 


-237 


-250 


13 


200 


-270 


-285 


15 


300 


-201 


-305 


14 


500 


-309 


-323 


14 


700 


-31S 


-331 


13 



Here X' denotes — ^f- —F{1), 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 oi 
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 teolotropy of the tube. Jf the tube, after it ia 
magnetized l>oth longitudinally and circularly, ia demagnetized 
by reversals with regard to the longitudinal magaetization, 
the circular field remaining constant, as ivas actually the case 
in the present experiment, the elongation due to the circular 
field alone ia usually increased by one or two scale divisions, 



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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 seolotropy 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. 



I 


Wolfram steel, t=17.7 


Soft iron, 


t=26.2 


L' (cal.) 


U (exp.) 


// (cal.) 


L' (exp.) 


id 


2xlo' 


Ixio' 


9xl0' 


8xl0' 


30 


22 


18 


23 


30 


50 


31 


37 


29 


38 


80 


39 


51 


31 


41 


120 


46 


57 


29 


41 


200 


48 


62 


21 


36 


300 


48 


64 


12 


26 


500 


47 


63 


- 5 


7 


700 


45 


62 


-17 


- 5 



For iron and steel, the sensibility of the apparatus was about 
2x10""' and the correction for temperature amounted to 5xl0~^ 

1) K. Honda, Jour. Sc. Coll. XI., 311, 1899^ ' 



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100 



K. HONDA. 



'il 



in the most significant case. I believe that the principal causes 
of discrepancy above enumerated are sufficient to account for 
the diflference 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 verj 
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 againsi 
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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Jour Sc CQiL VoL Xiii. PL Viii 




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Jour. ScColl. Vol. XIII. PI. IX. 



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Studien uber die Anpassungsfahigkeit einiger 
InAisorien an concentrirte Losungen'). 



Von 



Atsushi Yasuda, Rigahusfd, 

Professor der Natargeschicbte an der zweiten Hodiachule zu Sendai. 



Hierzu Tajel X-XII. 



Einleitnng. 

In der Natur finden wir Thiere und Pflanzen stets den 
obwaltenden Bedingungen angepasst. Diese Anpassung an die 
TJmgebung ist in der That erst im Verlaufe langer Generationen 
hervorgebracht worden. Wenn dann aber die Lebensbedin- 
gangen sich andern, wie werden diese Organismen dadurch beein- 
flusst? Unsere bisherigen Erfahrungen lehren uns, dass den 
Organismen im Allgemeinen die Fahigkeit innewohnt, sich diesen 
yeranderten Yerhaltnissen genau anzupassen und so dauernd 
leben zu konnen, nur unter der Bedingung, dass die Veranderung 
nicht plotzlich stattfindet, oder, wenn sie rasch vor sich geht, 
sie doch verhaltnissmassig unbedeutend ist. 

*) Die TorlUufige Mitt hei lung dieser Arbeit erschien in The Botanical Magazine. 
Tokyo 1897. Vol. XI, No. 121. pp. 19-24. und Annotationes Zoologicse Japonenses. 1897. 
VoL I, Part I et 11. pp. 23-29. 



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102 



A. TASUDA : ANPASSUNGSFAEHIGKEir 



Bekanntlich giebt es in der PflaDzenwelt die WasserformeD 
der ampliibischeii Gewachse, wie Polygonum amphibium nnd 
Ranunculus aqucUUiSj die sich in ihrer morphologischen nnd 
aoatomiBchen Beschaffenheit ganz anders als ihre Landformei 
verlialten. Auch sind die Hydrophyten in Bezag auf Gestall 
und Struktur einer grossen Metamorphose unterworfen, die m 
zum Leben im Wasser be^higt. Aehnliche Thatsachen finder 
wir auch ia der Thierwelt. Hierher gehoren z. B, unter der 
Amphibien die Anuren, deren aus den Eiern ausschliipfendi 
Laryeii durch Kiemen athmen, aber im erwachsenen Zustaud< 
dutch Lungen, wahrend die Thiere, welche zu den Perenni 
branchiateu gehoren, fortdauernd Kiemen besitzen, weil si^ 
lebenalaug im Wasser wohnen nnd niemals ein oberirdische 
Lioben fiihren, so dass sie sich jenem Medium voUig accommodir 
habeup 

Es diirfte daher von Interesse sein, wenn wir die Beschaf 
fenheit der in der Natur gefundenen Medien verandern, kunst 
liche Niihrmedien anfertigen und die Anpassungsfi-higkei 
gewisser fiir diesen Zweck ausgewahlter Organismen an diesi 
kiinstlicbeu Medien studiren. Es liegen bereits Untersuchunger 
maucber Forscher iiber derartige Kulturversuche bei niederei 
Organismen vor. Im nachsten Abschnitt fasse ich die Resultat 
der wichtigsten einschlagigen Arbeiten zusammen. 

Vorausschicken will ich noch, dass ich die Aupassung der In 
fusorien an solche kiinstlichen Medien studirt habe, die aus hetero 
genen Ixisungen von hoheren Concentrationen bestanden. Fol- 
gendes siud die Fragen, die ich zu beantworten versucht habe :— 
1) Welche Grade der Concentrationen der Aussenmedien konnei 
die Infusorien ertragen ? 2) Welche relative Widerstandsfahigkeil 
haben sie im Vergleich mit Algen und Pilzen ? 3) Welche 



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EINIGER INFUSORIEN. 103 

Veranderungen ihrer ausseren und inrieren Gestalt werden 
dadurch hervorgebracht, und wie wird ihr Bewegungs- und 
Vermehrungsvermogen durch diese heterogenen Medien beein- 
flosst ? In Folgendem werde ich der Eeihe nach die diesbezug- 
liche Litteratur, die Methodik meiner Versuche, die Beschreibung 
der Experimente und endlich die Zusammenfassung der Eesultate 
mittheilen. 

Die vorliegende Arbeit wurde grossentheils im botanischen 
Institut der kaiserlichen Universitat zu Tokyo unter freundlicher 
Anregung des Herrn Prof. Dr. M. Miyoshi ausgefuhrt, wel- 
chem ich fur seine liebenswiirdige Rathschlage meinen verbind- 
lichsten Dank ausspreche. Herrn Prof. Dr. J. Matsumura 
sage ich auch an dieser Stelle fiir 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 StahP), dass AelhaKum 
iqpticum sich allmahlich an Traubenzuckerlosungen anpasste und 
der Einwirkung einer 29^igen Losung widerstand. Eichter^) 
experimentirte mit Cyanophyceen und fand, dass RivvlaHa 39^ 
ige, Qloeocapsa 69iige, Anc^aena 69iige und Oscillaria 1096ige 
Eochsalzlosung ertragen konnten. Auch gelang es ihm, Dia- 
tomaceen in einer 79iigen Kochsalzlosung ein Jahr und in einer 
10%igen einen Monat lang leben zu lassen. Er zog ausserdem 

») E. Stahl. Zar Biologie der Myxomyoeten. Bot. Ztg. 1884. Nr. 11. p. 166. 
*) A. Bichter. Ueber die Anpaasang der Susswasseralgen an EochflalzldsuDgen. Flora. 
1892. pp. 18^. 



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104 A. YASUDA : ANPAS8UNGSFAEHIGKEIT 

verschiedene andere Algen in den Bereich seiner Untersuchungen, 
darunter Zygnemaj Mougeotia, Spirogyraj (hsmariunij Chlorella, 
Tetrasporay Giaetophoray Vaucheriay Oedogonium, Chara u. s. w.; 
eine gewisse Anzahl von ihnen vermochte sogar in 139iigei 
Losung zu existiren. Femer theilte Klebs^) mit, dass einige ic 
den Losungen organischer Verbindungen kultivirte Siisswasser- 
algen anfanglich eine Plasmolyse zeigten, die aber nach einiger 
Stunden vollstandig ausgeglichen war, worauf sie ohrie Bchaden ir 
den neuen Medien fortlebten. Nach demselben Autor gedieh Zyg^ 
nema in einer 10 bis 209^igen Glycerinlosung eine Woche lang 
Auch 10-509^ ige Rohrzuckerlosungen vermochten dieselbe Alg< 
im Leben halten, aber mit verschiedenen Wirkungen je nael 
der Concentration : 1095ige Losung veranlasste lebhafte Kern- 
theilung, 20-25 9iige Langenwachsthum, SO^^ige Zellhautbildunj 
und 409iige Assimilation und Starkebildung, wahrend in 509^ 
iger Losung die Alge nur wenige Tage lebte. 

Unter den Meeresalgen nahm Janse^) eine ahnliche Er- 
scheinung bei Ohaetoviorpha wahr, und zwar hervorgebracht durcl 
Kalisalpeter- und Kochsalzlosungen. Er fand namlich, dass 
wenn man diese Alge in eine solche Losung legt, in Folge ihrei 
Anpassung an dieselbe nach kurzer Zeit ihre Widerstandsfahig- 
keit bedeutend gesteigert wird. Oltmanns^) machte Experi- 
mente fiber den Einfluss der Concentrationsanderung d« 
Meerwassers auf FucuSj der bei niedriger Concentration sich deno 
neuen Medium ganzlich accommodirte. Eschenhagen*) kulti- 

>) G. Klebs. Beitrage zur Physiologie der Pflanzenzelle. Berichte der deutsch. bot 
GeBellach. 1887. Bd. V, Heft 5. p. 181. 

•) J. M. Janse. Plasmolytische Versuche an Algen. Bot Centralbl. 1887. Bd. XXXIf 
p. 21. 

•) F. Oltmanns. Ueber die Bedeutung der ConcentrationsanderuDg des Meerwassers fui 
das Leben der Algen. Btzb. d. Konigl. preuss. Akad. d. Wissensch. zn Berlin. 1891. p. 193. 

*)F. Eschenhagen. Ueber den £influ88 von Losungen verschiedener Concentratioi 
auf das Wachsthunj von Scbimraelpilzen. Stolp. 1889, 



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EINIGEB INFUSORIEN. 105 

virte AspergiUus niger^ PenicilUum glaucum und Botrytis dnerea 
in verschiedentlich concentrirten Losungen von Traubenzucker, 
Glycerin, Natronsalpeter, Kalisalpeter, Chlornatrium und Chlor- 
kalium, und wies sowohl die Grenzpunkte ihrer Accommoda- 
tion als auch ihr Wachsthumsverhaltniss zu diesen Substraten 
nach. B a c h m a n n^) bewies durch zahlreiche Experimente, dass 
Thammdium elegans durch veranderte aussere Bedingungen 
gezwungen werden kann, diese oder jene Art von Sporangiolen 
zu bilden oder die Bildung derselben ganzlich zu unterdnicken. 
Eay^) sate die Sporen von Sterigmatocystis alba in Medien, 
welche aus Zucker, Starke, Mohren, Kartoffeln, Gelatine und 
mineralischen Salzen bestanden, und erhielt verschiedene aus 
diesen Sporen entwickelte Pilzformen. 

Auch fur das Thierreich fehlt es an diesbezuglichen Unter- 
Buchungen nicht. Als Beispiele seien folgende angefiihrt: — 
Schmankewitsch') beobachtete, dass BrancMpuB stagnalts, der 
immer in Susswasser gefunden wird, sich, wenn man ihn in 
versiisstem Meerwasser zuchtet, der Form von Artemia Milhau- 
seniif einer das Brackwasser bewohnenden Art, nahert, und, 
wenn man das Brackwasser so lange concentrirt, bis dasselbe 
den Salzgehalt des Meerwassers erreicht hat, sich in Artemia 
saKnaf eine Meerwasser- Art, verwandelt. Herbs t*) zuchtete die 
Larven einiger Seeigel in verschiedenen Losungen von Lithium-, 



^) J. Bachmann. Einfluss der ausseren Bedingungen auf die Sporenbildung yon 
Thamnidium ekgana Link. Bot. Ztg. 1895. Abt J. p. 128. 

■) M. J. Ray. Variations des Champignons infiSrieurs sous Tinfluence du milieu. 
Bevne g^n^rale de Botanique. 1897. T. IX. pp. 193-259 et pp. 283-304. 

')W. Schmankewitsch. Zur Kenntniss des Einflusses der ausseren Lebensbeding- 
UDgen auf die Organisation derThiere. Zeitscli. f. wiss. Zool. 1887. Bd. XXIX. p. 429. 

*) C Herbst. Experimen telle Untersuchungen iiber den Eiufluss der veranderten 
chemischen Zusammensetzung des umgebenden Mediums auf die Entwicklung der Tbiere, 
L Theil. Zeitscb. f. wiss. Zool. 1892. Bd. XV. p. 446. 



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106 A. TASUDA : ANPASSUHGSFAEHIGKEIT 

Natrium- nnd Ealiamsalzen, nnd fand, dass die Wirkungsstarke 
anf die EntwicklaDgsstufen derselben in einer Reihe der Salze 
TOD demselben Sadical ihrem Molekulargewicht umgekehrt pro- 
portional ist, d. h. ihre Wirkungsstarke nimmt um so starker 
abp je mehr ihre Molekulargewichte znnehmen. Nach ihm 
wurde in einem Falle die (Jastmlation der Seeigel bedeutend 
irerzogert, in einem anderen Falle wurde die Pluteusorganisation 
entweder mit der runden und gedrungenen Qestalt ohne Fortsatze 
Oder sogar ohne eine Spur des Kalkgeriistes gewonnen. Cohn^) 
bemerkte, dass eine plotzliche Concentrationsanderung des 
Aussenmediums der Infusorien eine schadliche oder todtliche 
Einwirkung ausubt Fabre-Domergue^ beobachtete auch das 
Verhaltnifls der Ernahrung in den Korpem einiger Infusorien, 
und gelangte zu folgendem Schluss : ^^Dans des conditions par- 
fait@3 de nutrition prise dans Tacception la plus large da mot 
11 se produit des aliments de r^erve qui disparaissent quand 
dea conditions deviennent d^favorables k la vie." Weiter 
stodirte Boko my') die Veranderungen der Bewegung, der Qe- 
stalt und der Qrosse der Vacuolen von Paramaecium unter dem 
Einfluss gewisser Basen, wie Coffein, Ammoniak und Kali, deren 
1 promilL oder noch diinnere Losung im AUgemeinen die Bewe- 
gung verlang^amte, die Qestalt abrundete und sowohl Vergrosse- 
rung der Vacuolen als auch das Auftreten von neuen verursachte. 

>) F. Cohn. Entwickelungsgeschichte der microscopischen Algen and Pilze. Nova 
Acta Akad. Caes. Leopold. 1861. Bd. XXIV, Th. 1. p. 132. 

*} M, Fsbre-Domergne. Becherches anatomiques et physiologiqoes sar les infa- 
soiren dl»6i. Aim, d. Sc. nat Zool. 1888. S^r. VII, T. 6. p. 135. 

■) Th. Bokornj. Einige yergleichende Versache iiber das Verhalten yon Pflanzen 
imd nicdorcQ Thieren gegen basische Stoffe. Pfl tiger's Archly. 1895. pp. 557*562. 

Bokoraj gab auch in einer anderen Schrift (Vergleichende Stodien iiber die Qiftwir- 
knng yerscbiedener chemischer Substanzen bei Algen nnd Infhsorien. PflGger's Archiy. 
1896. pp. 202"3(Xi) eine genaue Untersncbung (iber die Giftwirknng yon Basen und Sauren 
nnorgani»chef Nutur, Salzen, Oxydations-Qlften, Phosphor, organischen Sauren, Alkoholen, 
Alkaloiden u, a. m. anf das Leben der Infbsorien und anderer Qrganismen. 



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EINIOEB IKFUSOSIEN. 107 

Endlich mussen noch die Besultate der Untersuchungen von 
Davenport und NeaP) Erwahnung finden ; sie zuchteten Stentor 
2 Tage lang in einer O9OOOO59& Bublimat enthaltenden Kaltur- 
losuDg; die Thiere liessen sich sehr wohl acclimatisiren und 
erwiesen sich gegen eine O^OOl^^ige sonst todtliche Sublimat- 
losung ca. vier Mai langer widerstandsfahig als diejenigen, die 
im Wasser kultivirt worden waren. 

Ueberblickt man die Ergebnisse dieser Untersuchungen, so 
ersieht man, dass sowohl den niederen Thieren als auch den 
niederen Pflanzen die Fahigkeit innewohnt, sich geanderten 
Aussenmedien leicht anzupassen. Da aber diese Fahigkeit bei 
verschiedenartigen Organismen verschieden stark ausgepragt ist 
nod unter Umstanden mannigfaltig auftritt, so muss jeder specielle 
Fall genau erforscht werden. Meine vorliegenden Studien sollen 
in dieser Uinsicht einen kleinen Beitrag bringen. 



Methodisches. 



Als Versuchsmaterial wahlte ich solche Infusorien aus, die 
in Graben und Teichen stets gefunden werden konnen. Da aber 
die in der freien Natur vorkommenden Infusorien nie in reiner 
Eolonie vorhanden sind, so liess ich sie in einem Gefasse 
sich massenhaft entwickeln und unter Vorsichtsmassregeln eine 
langere Zeit fortleben. 

Genau nach den Angaben von Miyoshi^) kultivirte ich 
die Infusorien in einem mit Spirogyra gefullten Gefasse. Sobald 

') G. B. Davenport andH. V. Neal. On the Acclimatization of Organisms to 
Poisonous Chemical Sabetanoes. Archiy fiir Entwicklangsmechanik der Organismen. 1895. 
Bd. II, Heft 4. p. 581. 

*) M. Mijoshi. Physiologische Studien uber Ciliaten. The Botanical Magazine. 
Tokyo 1896. Vol. X, No. 112. p. 43. 



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108 



A. YASUDA : ANPASStrNGSFAEHIGKBIT 



die griiiie Masse der Alge sich allmahlicli zu verfiirben begaan 
tmd die urspniDglich klare Fliissigkeit immer mehr getriibt 
wurde, bemerkte ich die Entwickelung verachiedener Arten von 
Infusorien, die sicb oft mit einer erstaimlichen Sehnelllgkeit 
Yermehrtea uud sicb aiiflallenderweise in bald fadenformigeD, 
bald netzartigen Kolonieri gruppirten* Die mieroseopische 
UntersucbuDg ergab, dass diese Kolouien nur aus wenigen Arten 
bestanden, die im Kampfe ums Daseln den Rest iiberwundea 
liatten, Aus dieser Miscbkultiir isolirte ieb eiuzelne Arten, indem 
icb mittelst eiuer Pipette eiue kleine Menge der Kolonie zusam- 
men mit Wasser berausbolte und in eio ebenfalla mit Brunnea- 
wasser und der Alge gefiilltes Gefas3 versetzte. Bei gewobnlicber 
Ziinmertemperatur zeigten dieseKultureu ca. in einer Wocbe iippige 
Entwickelung ; nacb vier oder fiinf Wochen aber nabm die 
Verniebrung ab, und endUcb iiach secbs Wocbeu konnte nur 
nocb eine ausserordentlicb kleine Anzabl in der Flii^igkeit 
gefunden werden> Um eine und dieselbe Art immer in iippiger 
Kultur zu halten, legte ich deshalb alie drei Wocben neue 
Kulturen an und trng dafilr Sorge, dass sie niebt etwa durch 
Bacterieu inficirt wurden, 

Nacbdem icb die gewunscbten Arten auf solcbe Weiee in 
Kultur batte, warden die Experimente anf zweierlei Weise 
ausgefiihrt: einerseits priifte ieb die Anpassungsfabigkeit der 
Infusorien in dem Zustande, wie sie in der Natur vorkommen, 
d- b. in ibrem Zusammenleben mit Bacterieii ; anderseits wandte 
ich zu demselben Zwecke die Eeinkultur jedes Infusors an, ulso 
frei von Bacterieu. 

Der grdsste Theil meiner Experimente wurde mit unreinen 
Kulturen aasgefiihrt ■ in einigen Fallen wiederholte ich die 
Experimente an Eeiu kulturen, um zu wissen, ob die Gegeuwart 



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EINIGER INFUSORIEN. 109 

der Bacterien etwa das Ergebniss der Experimente modificirt 
hatte. Die Resultate stimmten aber bei beiden Kulturen vollkom- 
men ^berein, wie wir nachher sehen werden. 

Alsaussere Medien verwendete ich Losungen von Rohrzucker, 
Traubenzucker, Milchzucker, Glycerin, Kalisalpeter, Natron- 
fialpeter, Chlorkalium, Chlornatrium und Chlorammonium in 
verschiedenen Concentrationen. Diese Stoffe waren chemisch 
rein und wnrden vor dem Gebrauch voUstandig getrocknet. 
Folgende Infusorien wurden bei meinen Studien ausschliesslich 
verwendet : Euglena viridis, Chilomonas paramaeciumy Mallomonas 
FlossUi, Oolpidium colpoda und Paramaecium caudatum. AUe 
Kulturen, sowohl unreine als reine, wurden bei Zimmertem- 
peraturen von 26'^-30'^ C gehalten und in den Winteimonaten 
in einen Thermostat von etwa 30"^ C gestellt. 

Fiir unreine Kulturen benutzte ich eine grosse Anzahl der 
5 cm hohen und 3 cm weiten cylindrischen Qlasgefasse, deren jedes 
30 ccm der Versuchslosung und etwa 1 Gramm Spiroffyra^Fiidea 
enthielt. Da die Infusorien im Brunnenwasser weit besser gedeihen 
als in destillirtem Wasser, so wendete ich bei der Zubereitung 
der flussigen Yersuchsmedien das letztere als auflosendes Mittel fiir 
verschiedene Substanzen an, wobei die Menge der darin gelost 
Torhandenen 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 KontroUkultur, in welcher nur 
Brunnenwasser und S^rogyra-FSiden angewendet wurden, anzu- 
fertigen und zum Vergleiche dienen zu lassen. 

In Bezug auf die Eeinkultur der Protozoen im AUgemeinen 



Die chemische Analyse des Brunnenwassers zeigte, dass es 0,096^ Eochsalz 
enthielt. 



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110 A. YASUDA : ANPASSUNGSFAEHIGKEIT 

haben viele Forscher^) ia neuerer 2ieit versucht, sie entweder auf 
festen Substratea oder iu fliissigen Medien zu ziichten. Bei der 
Isolirung der Infusorien befolgte ich genau die Methode von 
Ogata^) mit positivem Eesultat. Ich liess mir nach seiner Vor- 
schrift feine Glascapillarrohren anfertigen, deren Durchmesser je 
nach der Grosse des Versuchsobjects variirten. So war z. B. bei 
Chilomonas paramaecium^ dessen Korper 25-30 fi Lange und 
10-12 /^ Breite hat, das Capillarrohr etwa 0.1 mm in innerem 
Durchmesser und 10 cm in Lange, wahrend es bei Calpidium 
colpoda^ dessen Korper 60-70 /^ lang und 25-30 // breit ist, einen 
inneren Durchmesser von 0,15 mm und eine Lange von 10 cm 
hatte. 

Sobald das Capillarrohr nach dem Eintauchen in eine 
sterilisirte Nahrlosung mit der letzteren grossentheils gefiillt 
war, brachte ich das namliche Ende desselben in die Mischkul- 
turfliissigkeit von Bacterien und Infusorien und liess das Rolii 
sich mit der Flussigkeit vollig fiillen. Untersucht man ein 
solches Capillarrohr uuter einem Microscope, so findet man an 
der Capillarrohrmiindung eine grosse Anzahl von Infusorien in 
Bewegung. Einige streben sich ins Innere zuriickzuziehen, bald 
aber kommen sie nach dei* Miindung zuriick. Wegen der starken 
Aerotaxis und schwachen Chemotaxis der Organismen') gelingt 

*) Wahrend Beijerinck (Kalturvereuche mit Amoben anf festen Substraten. Centralbl. 
f. Bak. u. Parasit 1896. £d. XIX, No. 8), Celli (Die Kultar der Amobea anf festen Sub- 
straten. Centralbl. f. Bak. u. Parasit. 1896. Bd. XIX, No. 14/15.), Schardinger (Reinkulturen 
yon Protozoen auf festen Nahrboden. Centralbl. f. Bak. u. Parasit. 1896. Bd. XIX, No. 14/15.), 
Gorini (Die Kultar der Amoben auf festen Substraten. Centralbl. f. Bak. u. Parasit 1896. 
Bd. XIX, No. 20), Tischutkin (Ueber Agar-Agarkulturen einiger Algen und Amoben. 
Centralbl. f. Bak., Parasit. u. Infekt. 1897. Bd. Ill, No. 7/8.) und andere Forscher Amoben 
auf festen Substraten kiiustlich ziichten konnten, ist es Ogata (Ueber die Reinkultur gewis- 
ser Protozoen-Infusorien. Centralbl. f. Bak. u. Parasit. 1893. Bd. XIV, No. 6.) audi gelun- 
gen, Polyioma uveUa in fliissigen Medien rein zu kultiviren. 

•) M. Ogata, he, eU, p. 168. 

») M. Miyoshi. loc. cit. p. 48. 



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EINIGEB INFUSOBIEN. Ill 

es nicht immer, die Infusorien auf diese Weise hervorzulocken, und 
ihrer habhaft zu werden. Nur wena sie zufallig in die sterilisirte 
Flussigkeit tief eindringen, kann man unter dem Microscope das 
Capillarrohr an der betreflfenden Stelle abbrechen und dann das 
Ende des Eohrs zuschmelzen. Sodann impft man den infu- 
soriumhaltigen Capillarrohrinhalt, indem man das Eohr mit 
einem sterilisirten Piucet abbricht und den Inhalt in ein mit 
sterisirter Nahrlosuug gefiilltes Reagensglas hineinblast. 

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 konnten. Bei der Zimmertemperatur 
von 20^ C blieb die geimpfte Nahrfliissigkeit nach zwei oder 
drei Tagen yollkommen klar, und erst nach ungefahr zehn Tagen 
erschienen Hunderte von Individuen, die nahe der Oberflache 
der Flussigkeit als selir kleine weisse Piinktchen bin und her 
schwammen. Die Zahl dieser weissen Piinktchen nahm hemach 
allmahlich zu, und dieselben wareu nicht allein am oberen Theil 
des Eeagensglases, sondem auch am mittlereji und unteren Theil 
desselben zerstreut sichtbar. Eine solche Erscheinung bedeutet, 
dass die Keinkultur gut ausgefallen ist, und dass man in jener 
Nahrlosung nichts anders als die isolirte Art der Infusorien 
findet Wenn aber die Nahrlosung wahrend der Impfung von 
Bacterien inficirt wird, so tritt immer eine starke Triibung zu 
Tage, und man bekommt in diesem Falle selbstverstandlich 
keine Reinkultur der Infusorien. 

Die auf diese Weise hergestellte Reinkultur gedieh 4-6 
Woehen lang, vorausgesetzt dass die Nahrstoffe in der Kultur- 
fliissigkeit nicht voUig erschopft waren. Durch erneuerte Wieder- 



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112 



A. YASUDA : ANPASSUNGSFA£HI6££IT 



ii 



impfuiig konnte icli die Organ ismen in reiuem, gutem Kultur- 
zustande eiue lange Zeit erhalten. 

Die Nahrlosung, die ich fur die Reinkultur gebrauchte, 
stellte ieli nach der Vorschrift von Ogata an, uud zwar war 
ihre Zusammensetzung folgende : 

Fleisebextract 1 g 

Rohrzucker 20 „ 

Conceutrirt gekochte Losung von Porphyra vulgaris. 250 ccm 
DestilHrtes Wasser 729 „ 



Beschreibung der Versuche. 

Wie gesagt, stellte ich die Experimente hauptsachlich mit 
unrein en Kulturen an, in der Absicht, den Einfluss, welchen die 
ausaerea Bedingungen auf die Infusorien in ibrem uatiirlicben 
Vorkommen ausiiben, festzustellen. Dabei versaumte ich aber 
uicht^ Kon troll versuche mit reinen Kulturen zu machen und die 
beideii Resultate zu vergleichen. 

Bei alien Versuchen mit unreineu und reinen Kulturen liess 
ich die Beachaflfenheit des Mediums sich plotzlich andern und 
priiffce die Anpassungsfahigkeit unserer Organismen an das neqe 
Medium. Hatte ich unreine Kulturen, so verglich ich gewohn- 
lich im Verlauf von 1-7 Tagen, zuweilen aber erst nach einem 
Mouat, die Wirkungen der verschiedenen Mediumsconcentrationen 
auf das Reproductionsvermogen und die Gestaltanderungen der 
Versucbsorganismen. Speciell bei den Versuchen mit Rohrzucker 
war es nothig, eine Reinkultur anzuwenden, well bei uureiner 
Kultur der Rohrzucker durch vorhandene Bacterien oder Pilze 
nach und nach invertirt und schliesslich gespalten wurde. Um 



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BINIOEB lOTTJSOBIEN. 118 

za erkenneiiy nach wie vielen Tagen der Eohrzucker zum Trau- 
benzucker invertirt wird, priifte ich mit der Fehling'schen 
Losung und fand, dass in meinen Versuchen nach etwa 4 Tagen 
eine kleine Inversion stattgefanden hatte. Meine unreiuen 
Bohrzuckerkulturen waren daher binnen der ersten drei Tage 
doch noch brauohbar. 

Dass diese Inversion ausser durch Bacterien und Pilze auch 
unter Mitwirkung der Infusorien stattfande, ist schon vou vom 
herein unwahrscbeinlich. Um dies aber experimentell zu con- 
statiren, stellte ich einige Versuche mit Bohrzuckerreinkulturen^) 
an, und gelangte beim Priifen der fraglichen Flussigkeit wie 
erwartet zu negativem Besultat. Die KontroUkultur mit A^fj^^ 
gillus glauQus zeigte eine starke Inversion, 

Bei alien Kulturen mit verschiedenen Stoffen machte ich 
immer KontroUkulturen, und bei den kritischen Versuchen, wie 
z, B, der Bestimmung der Concentrationsgrenze einer Flussigkeit, 
ia welcher die Organismen sich mehr oder minder anpassend 
leben konnen, wurden dieselbe Kulturen einige Male wiederholt 

Ich gehe nun zur Beschreibung der einzelnen Versuche bei 
jeder Art meiner Versuchsorganismen fiber. 

(a) Euglena viridis Yiaihg.^) 
Dieser Organismus hatte in der KontroUkultur folgende 



Da der Bohrzacker bekanntlich bei langem Kochen znm TheU in Trauben- and 
FnichUQcker verwandelt wird, so kann bei den Bohrzuckerreinkalturen die gebrauchliche 
Sterilisirang durch Hitze nicht ohne Vorsichtemassregeln angewendet werden. Ich sterilisirte 
deshalb den Bohrzacker mit abeolutem Alkohol and brachte ibn dann in die vorher 
steriliaierte Nahrloeong ein. 

•) Figuren in Fried rich Bitter v. Stein, Der Organismus der Infusionsthiere. 
Leipag 1878. Abt. Ill, Heft I. Taf. XX., W. Saville Kent, A Manual of the Infusoria. 
1880^1. VoLL PL XX. und O. Butschli, H. G. Bronn's Klaflsen und Ordnungen des 
Thierreiches. 1883-87. Bd. I. Protozoa, Abt. IL Taf. XLVIL 



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114 



A, YASUDA : ANPASSUNGSFAEHIGKEIT 



Merkmale : Gestalt gewohnlich spindelformig, Hinterende scbarfe 
zugespitzt, aber wegen der MetaboHe sich mannigfaltig verau 
dernd. Aus dem Schlunde entspringt eine lange Geissel. Chrc 
matophoren zablreich vorhanden, klein, scheibenforraig und rei 
griin gefarbt. Eine contractile Vacuole nahe dem Vorderend 
gelegen. Dicht bei demselben Ende befindet sich auch ein rotbe 
Augenfleck. 

Versuch 1. Rohrzucher^ Ci2H220ii, — Von einer l9^ige 
Losung anfangend liess ich in anderen Kulturen die Concentra 
tion um je l9^ steigen. Obgleich die Accommodation schwe 
wurde, als die Concentration zunahm, so lebte das Infusor doc 
bis zur 159^igen Losung, welche die Maximalconcentration fii 
den Organismus war. l9^ige, 29^ige und 396ige Kulture 
zeigten keine wesentliche Veranderung am Korper des Organis 
mus. Bei einer 49iigen Losung aber begannen die Chromato 
phoren an Grosse zuzunehmen. Von l^o iger Losung bis z 
79^iger war die spirale Bewegung des Organismus lebhafi 
dagegen iiber 89^ wurde sie allraahlich langsamer, wahrend di 
Chromatophoren selbst sich merklich ausdehnten ; als die Con 
centration des Mediums zunahm, wurde auch die Vermehrunj 
verhindert. Bei 12 9^ iger Losung konnte das Thier nicht meh; 
normal gedeihen, bei 139^ iiberlebte eine kleine Anzahl, di( 
jedoch nach einer Woche alle zu Grunde gingen ; bei 14?^ lebtei 
noch einige Individuen, aber nicht langer als 4 Tage, wahrenc 
sie bei 15?^ kaum einen Tag lebendig blieben. Da der Organis- 
mus metabolisch ist, so konnte keine deutliche Veranderung ar 
seiner ausseren Gestalt beobachtet werden. 

Versuch 2. Traubenzucher^ CeHizOo. — Unser Organisms 
konnte 1-11 9^ ige Concentrationen ertragen. Bei 19^- und 2?^- 
Kultur war noch keine merkliche Veranderung wahrzunehmen, 



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EINIGEH INPUSORIEN. 115 

aber schon bei 39iiger Losung dehnten sich die Chroinatoplioren 
ein wenig aus, und bei der Concentration iiber 39^ wurden sie 
noch etwas grosser. Die Bewegung des Thierkorpers scbien bei 
1-69^-Kulturen normal zu verlaufen ; erst bei 7^A wurde sie 
langsamer mit gleichzeitiger Verminderung der Vermebrungs- 
fahigkeit. Bei einer 99^igen L6sung vermehrten sich die Tbiere 
uberhaupt nicbt mehr, und nach einer Woche war nur noch 
eine kleine Anzahlam Leben. Alle Indiyiduen des Infusoriums 
gingen bei einer 109^- Kultur nach einer Woche, und bei einer 
Tou 11 9^ schon nach einigen Tagen zu Grunde. 

Versuch 3. MilchzuckeVy C12H22O11 + H2O. — Unter den oben 
erwahnten Zuckerarten schien unser Infusor sich an Milchzucker 
am besten anzupassen. Die Maximalconcentration, welcher es 
widerstehen konnte, war eine 179^ige. Von 4?^ an aufwarts 
scbienen die Chromatophoren sich zu vergrossern. Bei 1-1 19^- 
Kulturen nahm die Multiplication rasch zu, aber liber 129^ 
wurde sie etwas verraindert und auch die Bewegung wurde 
einigermassen trage. Eine 179^ige Losung erwies sich als die 
Grenzconcentration fur das Versuchsthier. 

Versuch 4. Glycerin, CsHgOj. — Die Versuche lehrten uns, 
dass sich unser In&sor an Glycerin weit schlechter anpassteals 
an eine der oben erwahnten drei Zuckerarten, denn das Thier 
konnte nur 1-69^ ige Losungen ertragen, Bei einer 29iigen 
Losung erweiterten sich die Chromatophoren ; bei 39^- Kultur 
lebte eine kleine Anzahl noch am funften Tage, und bei 69^ 
blieben nur wenige Individuen noch einige Tage am Leben. 
Die Bewegung wurde bei einer 49^igen Concentration schon 
vielfach retardirt, und bei 69^ horte sie fast ganzlich auf. 
Ferner war in der letzteren Losung eine pathologische Erschei- 
nung wahrzunehmen, indem die Cuticula des Korpers urn die 



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116 A. YASUDA : AWrPASSUNGSPAEHIGKEIT 

Ohromatoplioren etwas einsclirumpftey sodass ilir Umrids im 
optischen Schnitte gesehen zickzackformig aussah^ und die 
Chromatophoren selbst verscbmolzen melir oder weniger mit 
einander. 

Versuch 5. SchwefehaureB Magnesmmj Mg804.— Unter den 
unorganischen Substanzen erwies sich das scbwefelsaure Magnesium 
als dem Leben dcs OrganisiDUS am beaten zusagend. Der 
Organismus konnte die Concentration von 1-6?^ vertragen. Die 
Chromatophoren nahmen in ihrer Grosse fortwahrend zu, als die 
Concentration von 1,5?^ bis auf ihr Maximum stieg. In einer 
8,4^igen Losung zeigte das Thier eine sehr trage Bewegung, 
und schon bei 4-69Sigen Losungen ging es beinahe zum 
Stillstand iiber. Betrug die Concentration nur 1-2,5 96, so gedieh 
unser Thier einen Monat lang vollkommen normal, aber von 
einer 2,69iigen Concentration an auf warts biisste es seine Ver- 
mehrungsfahigkeit ein, und endlich bei 5-69^ blieben nur 
vereinzelte Individuen am Leben. 

Versuch 6. SalpelerBaures Kaliunij KNO3. — Der Organismus 
widerstand einer 2,49iigen Concentration. Von 0,8?^ an fingen 
die Chromatophoren an sich zu erweitem ; fiber 29^ wurde die 
Bewegung sehr langsam. Im Allgemeinen schien das vorliegende 
Salz auf die Mulptiplication des Infusors hemmend zu wirken, 
da selbst bei verdunnten Losungen das Thier eine unbedcutende 
Vermehrung zeigte. 

Versuch 7. Salpetersaures Natrium, NaNOg. — Dieses Salz 
verhielt sich fast wie das vorige. Eine 296ige Losung war das 
Maximum, welches unser Thier ertragen konnte. Die Chromato- 
phoren dehnten sich schon von 0,8 9^ an aus. Die Bew^ung 
war bei 2^A iger Losung nach zwei Tagen sehr trage. 

Versuch 8, ChlorhaUum, KCl. — Nachst dem Magnesium- 



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EINIGER INFUSORIEN. 



117 



wies gich unter den unorganischen Stoffen das Chlor- 
lir die Yermehrung des Organismus am giinstigsten. 
?S]ge Loaung verursacbte soVohl Zahlvermehrung als auch 
lerweiteruog der Chroma tophoren. In 0,2-l9'^igen 
ationen gedieh der Orgauismus noch nach 40 Tagen, 
! Concentration auf 2,89i, welcbes die maximale Grenze 
Organ ism us war, so horte die Bewegung fast ganzlich 
rend die Chromatophoren sich tbeilweise zu grosseren 

verschmolzen. 

3ucli 9. Cklornatriumy NaCl. — 0,2-1,89^ waren die Con- 
nen, bei welchen das Tbier am Leben blieb. Die 
ophoren schienen bei einer 0,8?^ igen Losung an Grosse 
aen, und bei einer l,69^igen Concentration zeigte der 
lus noch eine langsame Bewegung. 
sucli 10. Cklorammonium, NH4CI. — Dieses Salz wirkte 
en oben genannten Stoffen am ungiinstigsten auf das 
^3 Organismus ein, sodass die Anpassungsgrenze hier am 
Bu war. 0,2-0,6?^ Kulturen gedieben noch am Ende der 
l^ocbe, aber liber 1% vermebrte sich das Tbier nicbt mebr, 
1,4?^ lebteo kaum noch einige Individuen. Bei 0,69i 
ung nahm die Grosse der Chromatophoren zu, und bei 
^bmolzen sie sich zu wenigen grosseren Komern. Eine 

I^sung verursacbte immer die Verscbmelzung der 
ophoren und hob gleichzeitig die Bewegung des Orga- 
,uf. 

wiederbolte dieselbe Versucbe zehnmal mit Reinkulturen 
;lich die Resultate mit denjenigen bei den unreinen Kul- 
Uie Ergebnisse stimmten in beiden Fallen vollig uberein. 
eobacbtete ich, dass bei den Versuchen mit Beinkulturen 
elligkeit der Multiplication fur die Losungen verscbiede- 



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118 A. YASUDA : ANPASSUNG8FAEHIGKEIT 

ner Coucentrationen eines und desselben Stoffes nicht allzu 
gleich war, obgleich 8ie gleichzeitig geimpft worden waren 
Ira 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 
Woehen nach der Impfung, die Vermehrung eine starke bei 
296, eine massige bei 4?^, eine sehr unbedeutende bei 69^, eine 
noch sparlichere bei 8^A und keine bei 109^. Ferner war die 
Vermehrung bei deraelben Kultur nach 4 Woehen bei 29^ eine 
sehr starke, bei 4?^ eine starke, bei 69^ eine massige, bei Sfo 
eine unbedeutende und bei 109^ eine hochst sparliche Vermehrung 
zu beobachten, wahrend sich am Ende der sechsten Woche bei 
2-49^ eine sehr starke, bei 69^ eine starke, bei 89^ eine nmssige 
und bei 109^ eine sparliche Vermehrung zeigte. Auch beim 
Milchzucker wurden ahnliche Thatsachen constatirt. So gedieheu 
nach 2 Woehen 2-49^ -Kulturen ausgezeichnet ; 69^ -Kultur zeigte 
eine starke, 89^ eine maBsige, 109^ eine sparliche, 129^ eine noch 
schwachere und 149^ gar keine Multiplication mehr. Nach 4 
Woehen aber vermehrten sich die Organismen bei 2-69^ sehr 
stark, bei 89^ stark, bei 109^ massig, bei 129^ sparlich und bei 
149^ sehr sparlich. Endlich nach 6 Woehen gedieh die Multi- 
plication stark bei 109^, massig bei 129^ und sparlich bei 149^. 
Auch fiir schwefelsaures Magnesium, salpetersaures KSlium, 
Chlornatrium u. s. w. habe ich ahnliche Erscheinungen wahr- 
genommen. 

(b) Chilomonas paramaecium Ehrbg.^) 
Der Organismus in Kontrollkultur hatte folgende Charak- 
teristika : Korper nicht metabolisch, sondern plastisch. Gestalt 

») Figuren in Friedrich Ritter v. Stein, loc. cil. Taf. XIX, W. Sayille Kent 
loc. ciL PI. XXIV und O. Butschli. loc. cit. Taf. XLV. 



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EINIGER IXFUSORIEN. 119 

langlich oval, seitlich comprimirt. Vorderende breiter uud schief 
abgestutzt, Hinterende dagegen rundlich zugespitzt. Zwei 
Geisseln am Vorderende, eine derselben roUte sich, wenn der 
Organismus im Euhezustande war. Zablreicbe spbaroidiscbe 
Amylumkorner dicht unter der Korperoberflache. Eine contractile 
Vacuole am Vordertbeil des Korpers. 

Versuch 1. Rohrzucker. — Mit einer l96igen Losung anfangend 
liess ich die Coucentration um je l9^ steigen, wie es bei Euglena 
viridis der Fall war. Von 29^ an auf warts debnten sicb die 
Korncben fortwabrend aus. Ueber 69^ nabm der Organismus 
an Dicke und Breite zu, nicbt aber an Lange, und sab so 
einfach oval aus. Die Vermebrung wurde scbon bei 4?^ verzogert; 
bei 794 lebten einige Individuen nocb eine Wocbe lang. Die 
Individuen aus der 79^igen Kultur waren so trage, dass sie an 
einem bestimmten Platze still lagen und nur eine zitternde Be- 
wegung zeigten ; vor ibrem Tode biipften sie einige Male 
riickwarts. Die letztere Erscbeinung wurde aucb bei den con- 
centrirten Losungen anderer Stoffe beobacbtet. 

Versucb 2. Traubenzucker. — Der Traubenzucker wirkte starker 
als der Robrzucker. Die bochste Concentration, die der Orga- 
nismus vertragen konnte, war 69^. Bei 49^ -Kultur ging die 
Vermebrung nicbt mebr gut vor sicb, und bei 59^ blieb nur 
eine kleine Anzabl der Individuen am Leben. 29^ige Con- 
centration bewirkte, dass die Korncben sicb vergrosserten, und 
bei bVo wurde die Bewegung sebr langsam. Bei 69^ kam der 
Organismus fast zum Stillstande, und wurde eine Unebenheit des 
Korperumrisses bervorgerufen. 

Versucb 3. Milchzucker. — Der Organismus widerstand 1-89^- 
igen Concentrationen. Ueber 39^ vergrosserten die Korncben ibr 
Volumen, bei 69^ wurde die Multiplication verbindert und end- 



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120 



A. YASUDA : ANPASSUNGSFAEHIGKEIT 




lich bei S^A lebten nur noch einige Individuen eiae Woche lang 
weiter, mit dem erwahnten Unebenwerden der Korperumrisse. 
Merkwiirdig war, dass der Korper an Dicke und Breite zunahm, 
als die CoDcentration stieg. 

Versucb 4. Glycerin. — Eine 49iige Losung war die Maxi- 
mal concentration fiir die Accommodation des Thieres. Bei 294 
erweiteiien sich die Kornchen, bei 39^ horte die Vermehrung auf 
und bei 49^ lebten nur noch einige Individuen, deren Korpei 
uiiregelmassige Umrisse zeigten. 

Versuch 5. Schwefelsaures Magnesium. — In l-39iigen L6- 
suDgen lebte der Organismus fort. Von 0,8 9^ an aufwarts ver- 
gnisserten sich die Kornchen, und in 39iiger Losung wurder 
dieselben auffallend gross. Eine l,49^ige Losung verhinderte di< 
Multiplication. Bei hoheren Concentrationen trat bei einigen In- 
dividuen ein unregelmassiges Aussehen zu Tage. Diese Gestall- 
anderung wurde bei einer 2,59^ -Kultur besonders gut beobachtet 
indem alle Individuen noch 2 Wochen mit einer ungewohnlicher 
Unebenheit ihrer Korpergestalt fortlebten. 

Vei'such 6. Salpelersaurea Kalium. — Eine 29^ige Concentra- 
iiou schien die obere Grenze der Anpassung zu sein. £in( 
0j8?oige Losung veranlasste eine Vergrosserung der Amylum- 
turner, die bei einer l9^igen Losuug nach einer Woche einer 
8€hr grossen Durchmesser zeigten. Die Vermehrung ging nui 
bei iiiederen Concentrationen gut vor sich. 

Versuch 7. Salpetenaures Natrium. — Der Organismus konnte 
iti 0,2-1 ,29^ igen Losungen leben. Die Volumenzunahme det 
Kornchen fand von 0,69^ an statt. Bei hoheren Concentrationen 
getUeh unser Organismus nicht. Im Uebrigen fast dieselben 
Erscheiuungen wie beim vorhergehenden Versuche. 

Versuch 8. ChlorJcalium. — Eine 0,89i-Kultur am Ende des 



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EfiriGEE IirFUaOBIEir. 



121 



Pages nacli der Impfung untersucht zeigte sowohl Ver- 
ag der Kornchen als auch Abrundung des Korpers. Nach 
einer Woche besassen einige Individuen in einer 1^/6 
ang eine fast echeibenformige Gtestalt. Ueber 2^^ konnten 
t mehr leben< 

Buch 9. Chlornatrium. — Eine 0,49^ ige Losung liess die 
fa eich erweitern. Bei 0,8-l9^-Kulturen dehnte sich 
imen bedeutend aus, indessen der Korper des Organismus 
kiirzte und rundlich wurde. Bei l9iiger Concentration 
ausserste Grenze der Accommodation erreicht 
such 1(X Chloramfnonium.—'DeT Organismus konnte eich 
0,6^ ige Concentrationen anpassen. Bei einer 0,29^ 
Bung trat schon Kornchenvergrosserung ein, und bei 
igten einige Individuen unebene Umrisse, mit gleichzei- 
>8clnvachuiig ihrer Bewegung. Wie bei Euglena viridie 
bei CkUonwtim paramaecium iibte der vorliegende Stoff 
Q augewaodtan Chemikalien die starkste Einwirkung aus. 

(c) MaUomonas Plcsslii Perty/) 

' Organismus in der normalen Kultur zeigte folgende 
le; Gestalt oval, am Vorderende etwas schmaler. Die 
Cuticularoberfluche mit langen, biegsamen, borstigen 
u bekleidet; am Hinterende mit einer langen Geissel 
I. Anstatt der Amylumkomer war eine Anzahl von 
n vorhanden. Eine contractile Vacuole be&nd sich nahe 
teren Ende. Das Thier schwamm mit lebbafter Bewegung, 
3 oft plotzlich stillstand. 
rsuch 1. liohrzucker. — Der Organismus vertrug Anpas- 

ureu in W. S* K^iit. loc. cU. PL XXIV. 




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122 A. yasuda: anpassungsfaehigkeit 

sungsconcentrationen von 1-7 9^o. Auch bei hoheren Concentra- 
tionen nahm die Grosse des Korpers mehr oder minder za, 
wahrend die Multiplication und die Lebhaftigkeit der Bewegung 
allmahlich sanken. Ferner wuchs die Zahl und Grosse 
der Vacuolen mit der Conceutrationserhohung bedeutend an; 
diese Erscheinung trat schon bei einer 29^-Kultur ein. Erst 
von 496 an wurde die Multiplication schwacher und bei 79i 
erlitt die Bewegung eine Ketardirung, welche zu der sehr 
schnellen, nornialen Bewegung in grosseni Contrast stand. 

Versuch 2. Traubenzucker. — Eine 69^-Kultur war das Maxi- 
mum der Anpassung des Organismus, Die Vergrosserung des 
Korpers und die Zunahme der Vacuolen bei starkeren Concen- 
trationen waren wie beim Rohrzucker. Eine 29^ige Losung 
erweiterte die Vacuolen einigermassen, und von 39^ an nahm 
die Vermehrung des Organismus ab. Die Bewegungshemmung 
war schon bei 49^ zu beobachten, noch starker bei 59^. 

Versuch 3. MUchzucker. — Das Infusor ertrug l-99^ige 
Losungen. Das allgeuieine Eesultat stimmte mit dem bei den 
anderen Zuckerarten iiberein ; der einzige Untei-schied war der, 
dass der Milchzucker eine schwachere Einwirkung auf den 
Organismus ausiibte als die anderen Zuckerarten. Die Vacuolen 
fingen erst bei einer 39iigen Losung an sich zu vermehren, und 
die Multiplication wurde erst von 79^ an etwas verlangsamt. 

Versuch 4. Glycerin. — Die Grenze der Accommodation war 
eine 49^ige Losung. Bei einer 2?.5- und noch aufifallender bei 
einer 39^-Kultur fand Zunahme der Zahl und Grosse der Vacuolen 
und Anschwellen des Organismus statt. 

Versuch 5. Schwefehaures Magnesium. — Der Orgauismus 
vermochte sich 1-3,49^ igen Losungen anzupassen. Eine 0,89^ 
ige Concentration verursachte sowohl Vermehrung als auch 






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EINIGER INFUSORIEN. 123 

Vergrosserung der Vacuolen, und bei hoher concentrirten L6- 
sungen war Verhiaderung der Multiplication und Eetardation der 
Bewegung wahrzunehmen. 

Versuch 6. Salpetermures Kalium. — Eine 0,79^ige Losung 
yergrosserte die Vacuolen etwas. Die Bewegung begann bei 
l-l,59iigen Losungen sehr langsam zu werden. Bei den Kul- 
turen hoherer Concentrationen pflegte der Organismus sich nicht 
zu vermehren, und im Verlaufe einiger Tage ging der grossere 
Theil der Individuen zu Grande. Eine l,596ige Losung bildete 
die Grenze der Anpassuug, 

Versucb 7. Salpetersaures Natrium. — Fur diesen Stoflf besass 
der Organismus eine besonders grosse Resistenzkraft. Er ver- 
mochte sich sogar einer 2,69iigen Concentration anzupassen, 
wenn auch mit grosser Schwierigkeit. Die Bewegung war bei 
1,5?^ noch lebhaft. 

Versuch 8. Chlorkalium. — ^Bei einer 0,896-Kultur vergrosser- 
ten sich die Vacuolen. Die Grenze der Anpassung des Organis- 
mus war bei 1,49^ iger Losung zu beobachten. Mit der Con- 
centrationssteigerung trat Korperabrundung ein. 

Versuch 9. Chloniatrium. — Die Maximalconcentration war 
1,596. Bei 0,896 -Losung schien der Korper nach 5 Tagen sich 
abzumnden. In einer l96igen Losung konnte das Thier 3 
Wochen lang gedeihen, aber in 1,5?6 starb es schon am Ende 
des vierten Tages. 

Versuch 10. Cfilor ammonium. — Fiir diesen Stoff besass der 
Organismus die kleinste Anpassungsfahigkeit, ganz wie es bei 
den anderen Infusorien der Fall war. Eine 0,896 ige Losung 
war das Maximum. Die Cuticularoberflache des lebendeu 
Organismus zeigte in dieser Losung nach einem Tage einige lougi- 
tudinale Falten. 



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124 A. YASUDA : ANPASSUNGSPAEHIGKEIT 

(d) Golpidium colpoda Ehrbg,*) 

Merkmale des Organismus in der normalen Kultur:— 
Korper mittelgross, nierenfSrmig, Riickenseite massig gewolbt; 
Bauchseite in der Nahe des Mundes etwas eingebuchtet. Vor- 
dereude viel schmaler als das abgerundete Hinterende. Cuti- 
cularwimpern auf der ganzen Oberflache des Korpers reiehlich 
vorhanden und an Grosse alle gleich. Mund in noassiger Ent- 
fernung vom Vorderende, in einer die Bauchseite querenden 
Einbuchtung. Eine contractile Vacuole und einige Nahrungs- 
vacuolen vorhanden. Bewegung lebhaft. 

Versuch 1. Rohrzucher. — Die ConcentrationsdiflTerenz der 
Versuchsserie war 1%. 8^o wurde als das Maximum erkannt. 
Schon bei einer 39^igen Losung begannen die Vacuolen sich 
etwas zu vermehren und zu vergrossern, Diese Erscheinung 
wurde mit der Concentrationserhohung immer mehr merklich. 
Ueber 4^4 sah der Korperumriss rundlich aus und die Grosse 
nahm merkwurdig zu. Multiplicationshemmung schon bei 6^/o. 

Versuch 2. Travhenzucker. — Der Organismus lebte in 1-796 
igen Losungen. Vermehrung und Vergrosserung der Vacuolen 
schon bei 29^ und Abrundung des Korpers bei 39^. Bei einer 
4,59^ -Kultur wurde die Multiplication sehr verzogert, bei Q¥» 
lebte am Ende des funften Tages noch eine kleine Anzahl der 
Thiere ; bei 79^ waren nur noch vereinzelte Individuen am 
Leben, welche schliesslich nach 6 Tagen abstarben. 

Versuch 3, MUchzucher. — ^l-1096ige Losungen wurden ver- 
tragen, Vacuolenvergrosserung von 39^ an aufwarts und Kor- 
perabrundung fiber 49^. Bei einer 796igen Losung wurde die 
Vermehrung verzogert, und bei 109^ konnten nur einige Indi- 



») Figuren in O. Butachli. loc. ciU 1887-89. Abt.111. Taf. LXIL 



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EINIGER INFUSORIEN, 125 

viduen 10 Tage lang leben. Auch hier fand mit der Concentra- 
tionssteigerung Grossenzunahme des Korpers statt. 

Versuch4. Glycerin. — Maximalconcentration 6^/0. Vermeh- 
rung und Vergrosserung der Vacuolen bei 29^ ; Korperabrundung 
bei 396. Bei 59^ lebten nur noch vereinzelte Indi viduen wenige 
Tage lang. 

Versuch 5. Schwefehaures Magnesium. — An passu ngsconcen- 
tration : 1-59^. Vacuolen vergrosserung und Korperabrundung 
begannen bei 2%. Bei schwacberen Coneentrationen gedieh der 
Organismus gut, aber fiber 3% schlecht. 

Versuch 6, Salpetersaures Kalium. — Maximalconcentration 
2%. Das Thier gedieh bei 0,89iiger Losung nicht mehr. 
Zahlzunahme und Vergrosserung der Vacuolen waren wie ge- 
wohnlich. 

Versuch 7. Salpetersaures Natrium. — Anpassungsconcentra- 
tion : 0,2-2?^. Ueber 0,8?^ nahm die Vermehrung stufenweise 
ab. Gestaltanderung u. s. w. waren ahnlich wie in den vorher- 
gehenden Fallen. 

Versuch 8. Chlorkalium. — Anpassungsconcentration : 0,2- 
1,69^. Die hoheren Coneentrationen fiber 0,89^ verursachten 
Multiplicationshemmung. Korperabrundung von 0,69^ an. 
Vacuolenvergrosserung fand auch bei starkeren Losungen statt. 

Versuch 9. Chlornatrium. — Maximalconcentration 1,59^. 
Volumenvergrosserung der Vacuolen wie gewohnlich. 

Versuch 10. ^Chlorammonium. — Der Organismus konnte sich 
nur ausserst verdfinnten Losungen anpassen. Schon bei 0,29^ 
trat Vacuolenvergrosserung ein, bei 0,89^ Bewegungshemmung 
und Unregelmassigwerden der Korperumrisse. Bei 19^, der 
hochsten Concentration, wfelcher das Thier widerstand, waren 
einige Indi viduen nach 2 Tagen noch lebendig. 



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I 



1'^ 



m 



126 A. YASUDA : ANPASSUNGSPAEHIGKEIT 

(e) Paramaedum caudatum Ehrbg.^) 

Merkmale des Organismus in der normalen Kultur : Korper 
yerlaogert, spindelformig, biegsam. Wimpern iiberall an der 
Oberflache des Korpers, dicht und gleichmassig. Trichocysten 
senkrecht zur Oberflache in der unter der Caticula unmittelbar 
befindlichen Rindenschicht gelegen. Mund nahe der Mitte der 
Bauchseite. Schlund ziemlich lang. Zwei contractile Vacuoleu 
am vorderen und hinteren Ende, mit strahligen zufiihrenden 
Kanalen. Nahrungsvocaolen vorhanden. Bewegung lebbaft. 
Verniehrung langsam. 

Versuch. 1. Rohrzucher. — Anpassungsconcentration 1-796. 
Von Wo an auf warts bis 7 96 Zahl- und Durchmesserzunabme 
der Vacuolen. Ueber 49^ Dickwerden des Korpers ; bei 79* 
blieb das Thier noch viele Tage lang lebendig. 

Ver8ucb2. Travhenzucker. — Anpassungsconcentration: 1-59*. 
Vacuolenvergrosserung bei ca. 29*, Korperabrundung bei 39*. 
Sonst wie beim vorhergehenden Versuch. 

Versuch 3. MUchzucher. — Maximalconcentration 89*. Ver- 
mehrung und Vergrosserung der Vacuolen bei 39* u. s. w. In 
hoheren Concentrationen erreichten die Durchmesser der Vacuolen 
bedeutend grossere Dimensionen, und der Korper erhielt ein 
fleischiges Aussehen. 

Versuch 4. Glycerin. — Anpassungsconcentration 1-39*. In 
diesem Medium konnte das Versuchsinfusor nicht lange am Leben 
bleiben. Vacuolenvergrosserung und Korperabrundung wie bei 
den Yorigen Versuchen. 

Versuch 5. Schwefekaures Magnesium. — Maximalconcentration 
2,4?^- Obgleich der Korper bei 0,29* verlangert war, so wurde 

>) Figuren in 0. Butschli. toe. eU, 1887-89, Abt. m. Taf. LXIJI. 



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EnaOER INFUSOEIEN. 



127 




jei 2,4?^ viel fleischiger, wobei sich auch die Vacuolen 
rten. 

uch 6, Saipetersanres Kalium. — ^Anpassungsconcentration 
Die durch dieses Medium hervorgebrachten Greetalt- 
an waren fast dieselben wie die vod Golpidium colpoda 
^Iben Med ill Di. 

uch 7. Salpelersaures Natrium. — Maximalconcentration 
►ickerizuriahme des Korpers von ca. 0,79^ an. In 1,2?^ 
ng lebten nur noch vereinzelte Individuen mit schwacher 

r. 

3 

uch 8, Chlorkalium. — Aupassungsconcentration 0,2-19^. 
I9i-Kultur sterb das Thier nach 3 Tagen ganzlich ab. 
ucli 9. Ghiornairium. — Anpassungseoncentration 0,2-1 9i. 
1 mit Concentradonssteigerung auch Abrundung des 
statt, 

uch 10. ChioT ammonium. — Maximalconcentration 0,59<. 
80r accomraodirte sich an dieses Medium am schwersten ; 
Kultur blieb es lange am Leben. 



AUgemelnes und Schlussbemerkungen. 

den oben angefiihrten Versuchen ergiebt sich, dass mit 

;erung der Concentration unabhangig von der chemischen 

tnheit die Cuticularoberflache der Infusorienkorper ein- 

;t, wenn die Organismen plotzlich in das Medium 

werJen, weil durch con cent rirtere Medien das Wasser 

Tliierkorper herausgezogen wird. Zugleich wird ihre 



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128 



A. YASUDA : ANPASSUXGSFAEHIGKEIT 



Bewegung, die bisher lebhaft gewesen war, immer langsamer, \xn\ 
nach einem kurzdauernden Zittern an einem Platze kommen di 
Thiere endlich zum Stillstande. VVenn aber die Concentratio 
des Mediums nicht zu stark ist, so konnen es die Infusorie 
ohne grossen Schaden ertragen, und die einmal gebildeten longi 
tudinalen Falten der Cuticularoberflache verschwinden nac 
eioiger Zeit wieder. Sogar bei concentrirteren Losungen find< 
man nicht selten einige Individuen, welche mit der contrahirtei 
unebenen Korperoberflache noch einige Tage lang fortlebe 
konnen. 

Je hoher die Concentrationen der Medien sind, desi 
schwerer wird selbstverstandlich die Anpassung, und wenn sic 
schliesslich die Maximumgrenze nahert, so stirbt der grossere The 
der Individuen ab. Im Falle gelungener Anpassung an ei 
gewisses Medium sieht man stets Volumen- sowie Zahlzunaha 
der Chromatophoren, Amylumkorner und Nahrungsvacuolei 
Gleichzeitig nimmt der Korper selbst an Dicke und Breite zi 
dagegen an Lange etwas ab, so dass er ein einigermassen shgf 
rundetes Aussehen erhalt. Zugleich ist ausserdem Grossenzunahn 
des ganzen Korpers wahrnehmbar, wie ich dies ausschliesslic 
mit Zuckerarten bei Golpidium colpoda^ MaUomonas FlossKi un 
Chilomonas paranuiecium nachgewiesen habe. 

Als eine allgemeine Kegel gilt auch, dass die Vermehrungi 
fahigkeit bei hoherer Concentration stark beeintrachtigt wire 
Unsere Versuche mit den Reinkulturen von JEhiglena viridi 
Chilomonas paramaecium und Golpidium colpoda bieten hierfii 
unzweideutige Beweise dar. Zum Vergleich fiihre ich einige dc 
bei Schimmelpilzkulturen gewonnenen Erfahrungen an. Eschei 
hagen^) constatirte, dass das Wachsthum einiger Schimmelarte 

F. Eschenhagen. he, cit, p. 55. 



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IINIGEB INFUSORIEN. 



129 



rch starkere Concentrationen des Substrates stark ver- 
; Klebs*) beobachtete, dass das Auftreten der Konidien- 
und die Perithecienbildung von Eurotium repenSy die 
ig und die Sporaogienbildung von Mvcor rdcemoms durch 
geriing der Coucentration des Mediums retardirt wurden. 
tlich fand iclr) auch, dass Aspergillus nigeVy der in Mag- 
ulfat-Nahrlosungen von verscbiedener Concentration 
it wurde^ nach 4 Tagen verschiedene Grade der Ent- 
Dg zeigte. Der Pilz wuchs in einer 59^-Kultur am besten, 
gut bei IQfo , walirend bei 209^ und 309^ nur noch eine sehr 
le Entwickelung zu beobacbten war. Die weisse Anlage 
nidienfriicbte trat bei 5% und 109^ naeb 4 Tagen, bei 
ich 5 Tagen und bei 309^ erst nach 6 Tagen ein. 
ter den zebn von rair angewendeten Stoflfen — vier organi- 
Lud sechs unorgaiiischen Verbindungen — passten unsere 
ten sich den Zuckerarten am besten an, und wieder unter 
ckeiarten erwies sich der Milchzucker als das beste An- 
;smedium, Ihm folgt der Rohrzucker in seiner Concen- 
hohe, wtihrend der Traubenzucker scbon in weit verdiinn- 
Qsungen auf die Organismen schadlich einwirkt. Glycerin 
s Anpassungamedium den Zuckerarten sehr nach. Unter 
organischen Verbindungen, deren Einwirkung stets viel 
ist al8 die der organischen Substanzen, ist schwefelsaures 
ium zur Verniehrung der Infusorien am geeigneststen, 
1 Chlorammonium fiir ihr Gedeihen das unpassendste 




Kleba. Die Bedmgungen der Fortpflanzung bei einigen Algen und Pilzen. 
pp. 446-535* 

Yaauda. Ueber den Einfluss verscbiedener unorganischer Salze auf die Fort- 
irgime von JspcrtjiUus ni^. The Botanical Magazine. Tokyo 1898. Vol. XII, No. 



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130 



A. YASUDA : ANPASSUNGSFAEHIGKEIT 



Medium ist, und unter den iibrigen Chlorkalium eine mittlere 
Stellung einniinmt. 

Die verschiedenen Infusorien zeigten in Bezug auf ihre 
Anpassungsfiihigkeit grosse Unterschiede, und zwar wohnte unter 
unseren Infusorien Euglena viridis die grosste WiderstandsfiLhig- 
keit inne, wahrend ParaTaaeoium caudatum die kleinste Resi- 
stenz besass. Folgende Tabelle zeigt die Grenze der Concentra- 
tionen, bis zu welcher die Infiisorien am Leben blieben : 



Stoffe 






i 



a; 



I 



I 



I 



d 

5 



i i 

} 



I a 

e .2 






-52 

o 



s 






if 

o s 



Formeln 



a + 






5 

o 



o 



s 






o 






Concentrationen der 
mit 0,1 Aeq. KNO, 
isotoniachen Losungen') 



% 
5,40 



% 
5,13 



% 
2,70 



% 
1,38 



1,80 



1,01 



% 
0,85 



0,75 



% 
0,59 



0,54 



£:§ 

1 * 

p. 2 

p a 

ll 






Euglena 
viridia 



17 



15 



II 



2,4 



2,8 



1,8 



Ckilomonas 
paramae- 



1,2 



McMomoTwa 
Plosslii 



3,4 



1,5 



2,6 



1,4 



1,5 



Oolpidivm 
colpoda 



10 



1,6 



1,5 



Paramae' 

dvm 
eaudcUwn 



2,4 



1,2 



1,4 



0,6 



0,8 



0,5 



>) Hugo de Vries. Eine Methode zur Analyse der TiugorkrafU Jahrb. f. wias. BoL 
1884. Bd. XIV. pp. 536—537. 



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l\ 



flS.*" 



EINIGER INFUSORIEN. 131 

Wie aus der vorstehenden Tabelle ersichtlich ist, muss die 
Wirkung der angewandten Substanzen auf die Infusorien nichl 
allein dem Grade ihrer Concentration zugeschrieben werden^ 
dagegen zeigt sich ein annaherndes Verhaltniss zu den isotoni- 
schen Concentrationen jedes StoflTes. Ich sage ausdriicklieh „ an- 
nahernd," 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*) : 



^) Die bisherigen Untersachungen ergaben in Bezng auf den Zusammenhang zwischen 
isotonischen Concentrationen und Reaktionsgroese durchaus negative Besnltate. Man ver- 
gkiche hieruber B. Stange. loe dL Nr. 22. p. 364, und C. B. Davenport and H. V. 
Neal. loe. eii. p. 579. 



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132 



A. YASUDA: ANPASaUNGSFABHIOKEIT 



StoflFe. 


i 

1 


J 


1 

a 

g 


.2 


if 


ll 

•Jj 


ll 

s 


i 
1 
5 


•£ 

g 
S 

s 
o 


1 

§ 
E 

1 


1 


ii 


Conoentrationen dsr 

Stoffe, die mit l(i% 

Bobrzucker iii^ju>- 

niach aiad 


15 


15,8 


7,9 


4 


5,3 


3 


2,5 


2,2 


1,7 


1,6 


Oefundene Wertlio 
der maximal ^u 

tration 


15 


17 


11 


6 


6 


2,4 


2 


2,8 


1,8 


1,4 


II 


ll 


Conceotrationen d^r 

Stoffe, die mit 7?^ 

Bohrzucker inoiu- 

nisch gind 


7 


7,4 


3,7 


1,9 


2,5 


M 


1,2 


1 


0,8 


0.7 


Gefundene Werttft 

der mazimnlen 

Anpassnagsooiic^ti- 

tration 


7 


8 


6 


4 


3 


2 


1,2 


2 


1 


0,6 


III 


i 


ConoentratioD d^r 

Stoffe, die mit 7% 

Bohrzucker iKoiCH 

niach Bind 


7 


7,4 
9 


3,7 


1,9 


2,5 


1,4 


1,2 


1 


0,8 


0,7 


Gefundene Wcrlhe 

der maximal4?n 

Anpafisungaco Dd^u- 

tration 


7 


6 


4 


3,4 


1,5 


2,6 


1,4 


1,5 


0,8 


IV 


ll 


Concentrationeu dor 

Stoffe, die mit S% 

Bohrzucker iaoto- 

nisoh Bind 


8 


8,4 


4,2 


-2,2 


2,8 


1,6 


1,4 


1,2 
1,6 


0,9 


0,8 


Gefundene Warthe 

der maximiiJfiti 

Anpa8sung80onc€u^ 

tration 


8 


10 


7 


r> 


5 


2 


2 


1,5 


1 


V 


•M 


ConcentrationeiJ tlor 

Stoffe, die mit 7% 

Bohrzucker isjolj- 

niBch Bind 


7 


7,4 


3,7 


1,9 


2,5 


1,4 


1,2 


I 


0,8 


0,7 


Gefundene Wertbo 

der maximal Kii 

AnpaBBungscoiiCfTi- 

tration 


7 
1 


8 


5 


3 


2,4 


1 


1,2 


1 


1 


0,5 



Dieselbe Tabelle zeigt auch zugleich, dass die Anpassongs- 
grenzen unserer Infiisorien an verschiedene Concentrationen im 
Allgemeinen weit niedriger sind als diejenige der niederen Algen 
und Schimmelpilze. So kann Zygnema nach Klebs*) 509^ Eohr- 

>) Q. Elebs. Beitriigc nn Phyaiologie der PflatizenEene. Benchte der deatsch bot. 
Geaellflcli. 1887. Bd. V. p, IB7, 



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EINIGER INFTJSORTEN. 133 

zucker und 209^ Glycerin vertragen. Dass dieselbe Alge sich 
auch einer 696igen Chlornatriuralosung anpassen kann, ist von 
Richter^) erwiesen worden. Was die Schimmelpilze anbelangt, 
80 zeigen sie ebenfalls eine weitaus grossere Widerstandsfahigkeit 
gegeu Starke Concentrationen. Ich gebe hier zura Vergleiche die 
von Eschenhagen^) erhaltenen Ergebnisse wieder. 



Tranbenzucker. 


Glycerin. 


Salpetersaures 
Natrium. 


Chlornatrinm, 


Aspergillus niger 53 9^ 


i3o/„ 


21 9^^ 


179^ 


Penicillium glaucum 55 „ 


43 „ 


21 „ 


18 „ 


Botryiis cinerea 51 „ 


37 „ 


16 „ 


12 „ 



Unsere Infusorien zeigen gegen dieselben Stoffe folgendes Ver- 
halten : 



Traubenzncker. 


Glycerin. 


Salpetersaures 
Natrium. 


Chlomatrium. 


Euglena viridia 


llo/o 


6 o/„ 


2 0/0 


l,8o/„ 


Cdpidium colpoda 


7 „ 


5 „ 


2 „ 


1,5 „ 


Mallomonas Plosslii 


6 „ 


4 „ 


2,6,, 


1,5 „ 


Chilomonas paramaecium 


6 „ 


4 „ 


1,2,, 


I „ 


Paramaecium caudaLum 


5 „ 


3 „ 


1,2,, 


1 „ 



Daraus geht hervor, dass die Resistenzkraft unserer Infusorien 
gegen die angewendeten Stoffe hinter derjenigen der iSchiramel- 
pilze weit zuriicksteht. 



>) A. Richter. loc. cU, p. 24. 

■) F. Eschenhagen. loc. cit. p. 55. 



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134 A. YASUDA : ANPASSUNGSFAEHIGKEIT 



Zusammenfkssung. 

(1) Isotonische Losungen der in Rede stehenden Substanzen 
iiben auf die von mir gepriiften Infusorien nur eine „ anna- 
hern d" gleichartige Wirkung aus. 

(2) Die Grenzen der Concentration, welcher sich diese 
Infusorien unter gewohnlichen Verhaltnissen anpassen konnen, 
liegen im Allgemeinen weit niedriger als die der niederen Algen 
und Schimmelpilze ; selbst das widerstandsfahigste darunter, 
Euglena viridis, vermag nur verhaltnissmassig schwache Concen- 
trationen zu ertragen. 

(3) Wenn die Organismen plotzlich in Losungen hoherer 
Concentrationen gebracht werden, so treten erst an der Cuticular- 
oberflache ihrer Korper longitudinale Falten auf, aber wahrend 
ihre Anpassung an das neue Medium stattfindet, dehnen sich die 
Falten allmahlich aus, bis sie zuletzt ganzlich verschwinden. 

(4) Die hohere Concentration des Mediums verlangsamt die 
Vermehrung der Infusorien. 

(5) Durch Steigerung der Concentration des Mediums wird 
die Bewegung der Organismen vielfach retardirt. 

(6) Bei Zuckerlosungen starkerer Concentration vergrossern 
sich die Korper der Infusorien bis zu einem gevvissen Grade. 

(7) Die Vacuolen, Chromatophoren oder Amylumkorner 
nehmen in dem Masse an Grosse zu, als die Mediumsconcentra- 
tion steigt. 

(8) Je mehr die Concentration des Mediums zunimmt, desto 
mehr runden sich die Korper der Organismen ab, und die Kor- 
perumrisse werden uneben. 

(9) Wenn das Maximum fiir die Accommodation ein nie- 



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\ 



EINIGER INFUSORIEN. 135 

driges ist, so finden die Veranderungen der Korper der Infusorien 
schon bei niederen Concentrationen des Mediums statt. 

(10) Wenn sich die Concentration des Mediums dem Maxi- 
mumpunkt nahert, so versehmelzen die in den Korpern der 
Organismen befindlichen Cbromatophoren oder Amylurakorner 
mehr oder weniger mit einander. 

Tokyo, 30. November 1898. 



Inhaltsiibersicht. 

Einleitung Seite 101. 

Litteratur „ 103. 

Methodisches „ 107. 

Beschreibung der Versuche „ 112. 

Allgemeines und Schlussbemerkungen „ 127. 

ZusammeDfaasuDg », 134. 



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136 A. YASUDA : ANPASSUNGSFAEHIGKEIT 

Erklarung der FigureD. 

Siimmtliche Figuren wurden nach den lebendigen Thieren in 
den unreinen Kultureu unmittelbar skizzirt, well ihre Gestalten 
bed den getodteten Individuen sich mehr oder weniger veranderten. 

TAFEL X. 

Fig. 1-46. Ghilomonas paramaeciuni Ehrl^. Vergr. 420. 
Fig. 1. Individuen aus einer Nahrlosung mit 1 9^ Milchzucker. 



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EINIGBR INFU80RIE». 



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A. YASUDA : ANPA8SUNGSFAEHIGKEIT 



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EINIGER INFUSORIEN. 



139 



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Fig. 22-28. Paramaecium caudatum Ehrbg. Verg. 240. 
Fig. 22. Individuum aus einer Nahrldsung mit 1 ^ Eohrzucker. 





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Ueber die Wachsthumsbeschleunigung einiger 
Algen und Pilze durch chemische Reize. 



VON 



N. 6no, RigakusM. 
Hierzu Tafel XIIL 

I. Einleitung und Litteratur. 

In seiner Arbeit : llfctudes chimiques sur la v^g^tation^), hat 
Raulin schon im Jahre 1869 darauf aufmerksam gemacht, dass 
Zink- und Siliciurasalze in geeigneter Dosis das Wachsthum von 
Aspergillus niger befordern. Auf dieser an sich richtigen Beo- 
bachtung fussend war der genannte Autor mit Unrecht der 
Ansicht geneigt, dass diese Substanzen zur normalen Entwiekelung 
unseres Pilzes nothwendig seien, indem er diese unter „ les 
^l^ments chimiques essentiels" rechnet und eine Nahrlosung von 
recht complicirter Zusammensetzung fiir Pilze vorschreibt. 

Ueber die Mineralstoffbediirfnisse 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. Sc nat. Bot, Ser. V, T. XL, 1869, S. 91. 

2) V. Naegeli, Der Ernahrungschemismus der niederen Pilze. Sitzungsberichte d. Kgl. 
Bayr. Akad. d. Wiss. Math.-phys. a. 1880. 



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142 



N. ONO: WACHSTHUMSBESCHLEUNICtITTIG 



neuerer Zeit wiircle dies Theraa von MoliscV), sowie auch 
Benecke*) wiedGraufgenomraen. Der Miihe dicser Aut 
verdanken \vir unsere heutigen Kenntnisse in dieser Richt 
Nach den iibeitiostimmeuden Angaben der geoanntcn Aut 
Btellt weder Zink noch Silicium einen eigentlichen Niihrstoff 
tind kaiin wohl von Kulturfliissigkeit ausgeschlossen warden 
Daee die wachsthurasbesehleunigende Wirkung geiw 
iretalliodikalo auf einer chemischen Reizung beruht, wurde 
PfefFer^} in seiner im Jahre 1895 erschienenen Arbeit 
ersten Male klar gestellt und spater in allgemeinen Ziigen ii 
2^ '' Aiiflage seiner Pflanzenphysiologie erortert* Er bemerl 
dem letztgenannten Werke, dass anscheinend geringfiigige 
stande in der That nicht selten einen erhebliclien Einfliiss 
Gedeihen nnd Wachsen haben, und sagt : ,,Vermuthlich hai 
uB Bjcli in dieser besehleunigenden Reizwirkuug nm eine 
tnamiigfticben Reaktionen, die darauf abzielen, dureli intensit 
Thatigkeit einen benachtheiligten Eiufluss tbuulicbst entge 
?Aiarbeiten oder Scbadigungen auszugleichen."^) Im vorherge 
genen Jalire wurdy eine Reihe Versuche von Richards'^) anges 
dereu Resultate die Ansicht Pfeffer's bestatigen. Er zog zu sc 
Untersiichiingen verschiedene Schwereumetallsaize wie Zi 
Kobalt-, Nickel-i Eisen-, und Mangansalze und einige at 
giftige Substans?en heran und stellte die Thatsacbe fest, dass 




]) H> Moliscbj Die mineralische Nahrung der niederen Pil^e. SiUungsberi* 
Wit'Dijr Akad., Oct. 1891 

!!) W» Benecke, Irie ziir Erniihnmg der Schimmelpilze nothwenJigen Metalie. P 
Jahrk r wiss, Eot Ed. XXVIII, 1895, S. 487. 

3) Pfefler* Election organischer Niihrstofle. Pringsh. Jahrb. t wii^s. Bot. lil. X] 

4) PflaTi7.cni>liys!ologie. 2 AuQ. Pxl. I. S. 374. 

5) H- M. KicliEirda, Die Beeinflussnng des Wachsthums einjger Pilae darch che 
lUissiJ* PrinpTih. JaUrli. t wiss. Bot. Bd. XXX, 1897, S. 665. 



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EINIGEB ALGEN UND PILZE. 143 

alle gepriiftea Substanzen inehr oder weniger die Pilzernte zu 
vermehren vermochten. 

Auch anderweitige Beispiele fiir Erhohung der Lebens- 
thatigkeiten durch geringe Zusatze giftiger Stoffe findet man in 
der Litteratur. So versuchte Schulz^) zu zeigen, dass die durch 
Saccharomyces verursachte alkoholische Gahrung bei Gegenwart 
von geringeren Quantitaten gewohnlicb als Hefegifte sich verbal- 
tender Substanzen wie Subliraat, Jod, Salicylsaure, Brom, Arse- 
nige Saure u. a., auf langere oder kiirzere Zeit wesentlich gehoben 
werden konnte. Diese Thatsache aber war bereits gewissen 
Gewerben nicht unbekannt gewesen, dass Zufuhrung von sonst 
gahrungshemmenden Stoffe, wie z. B. Kupfervitriol oder Sali- 
cylsaure, unter Umstanden die Hefe zu energischer Thatigkeit 
veranlasse. Derselbe Autor hat friiher eine ahnliclie Erhebung 
der Lebensausserungen bei dem thierischen Organismus beobaehtet 
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 grosseren Intensitat des Reizes^*^). 

Derartigen Verallgenieinerungen begegnen wir auch bei 
Hueppe^). Hauptsachlich auf die auf bakteriologischem Gebiete 
eonstatirten Thatsache sich stiitzend verkiindigt er die Erschein- 
ung als den Ausdruck des allgemein giltigen Gesetzes fiir die 
Wirkung von Chemikalien auf Protoplasma. Er bezeichnet dieses 
als das „ Biologische Grundgesetz,** welches er in folgendem 



1) H. Schulz, Ueber Hef^ifte. Pfluger^s Archiv f. Physiologic. Bd. 42., 1888. a 517. 

2) H. SchalZy Zur Lehre von der Arzneiwirkung. Virchow's Archiv. Bd. 108, 1877. 
a 427. 

3) F. Hueppe, NaturwlBBenschaftliche Einfiihrung in die Bakteriologie. 1896, Wies- 
baden, a 55. 



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144 N. ONO : WACHSTHUMSBESCHLEUNIGUNG 

Satze formulirt : „Jeder Korper, der in bestimmter Concentration 
Protoplasma totet und vernichtet, in geringeren Mengen die Ent- 
wickelungsfahigkeit aufhebt, aber in noch geringeren Mengen, 
jenseits eines IndiflFerenzpunktes, umgekehrt als Reiz wirkt und 
die Lebenseigenschaften erhoht.** 

Zu erwahnen ist noch, dass auch bei hoheren Pflanzen die- 
selbe oder eine wenigstens sehr nahe verwandte Erscheinung 
vorkommt. Bekanntlich pflegt man seit einigen Jahren die 
Weinreben mit Kupferpraparaten, der sogenannten Bordeaux- 
briihe, zur Bekampfung der Pilzkrankheit zu bespritzen. Eine 
derartige Behandlung ruft ausser der indirekten Einwirkung, die 
Schadigung durch parasitische Pilze herabzusetzen, auch eine Schar 
auflfallender Erscheinuugen 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 Chlorophyllbildung und 
daraus resultirende vermehrte Stiirkeproduktion, reichlicherer 
Traubenansatz, Beschleunigung der Reifung u. a. Rumm ist der 
Ansicht, dass die Steigerung der Chlorophyllbildung einem 
chemischen Reiz zuzuschreiben sei. Im darauf folgenden Jahre 
fiihrten Frank und Kriiger^) einige Bespritzungsversuche an 
Kartoflfeln aus, wobei sie auch eine ahnliche Thatsache consta- 
tirten, Ueber das eigentliche Wesen der Wirksamkeit jener 
Stoffe ist vorlaufig nichts weiteres zu sagen. 

Wie aus der oben angefiihrten Skizze ersichtlich ist, bezogen 



1) Bumm, Ueber die Wirkung der Kupferpraparate bei Bekiimpfung der sogenannteii 
Blattkrankheit der Weinrebe. Ber. d. deutsch. Bot. Ges. Bd. XL 1893. S. 709. 

Bumm, Zur Frage nach der Wirkuug der Kupfer-Kalksalze bei Bekampfung der 
Peronoepora vUkcla. ebenda S. 445. 

2) Frank u. Kriiger, Ueber den Beiz, welclien die Behandlung mit Kupfer auf die 
Kartofiel hervorruft. Ber. d. deutsch. Bot. Ges. Bd. XII, 1894. 



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EINIGER ALGEN UND PILZE. 145 

sich, wenn wir von der zuletzt besprocheDen Thatsache absehen, alle 
bisherigen diesbeziiglichen Versuche mit pflanzlichen Organism- 
en ausschliesslich auf chlorophyllose niedere Organismen der 
Pilze. Wenn die erwahnte Erscheinung, wie vielerseits behauptet 
wird, allgemeine Geltung haben wurde, so sollte es a priori zu 
erwarten sein, dass auch ehlorophyllhaltige niedere Organismen 
in gleichem Sinne reagiren. Dies aber benotigt einer experimen- 
tellen Bestatigung. 

Herbeigezogen wurden zu meiner Untersuchung versehiedene 
Schwerenmetallsalze, wie Zink-, Nickel-, Kobaltsulfat u. a., welche 
auf unsere Versuchsalgen gewisse Reizung auszuiiben vermochten. 
Ferner wurden eine Reihe Parallelversuche mit Pilzen angestellt, 
deren Ergebnisse die oben genannten Richards' scbe Unter- 
suchung bestatigt und erweitert. 

Die vorliegende Arbeit wurde auf Veranlassung und unter 
Leitung von Herrn Prof. Dr. Miyoshi im Botanischen Institut 
der Kaiserlichen Universitat zu Tokyo wahrend des Zeitraums 
von August 1898 bis Juni 1899 ausgefiihrt. 

Es ist mir eine angenehme Pflicht, meinem hochverehrten 
Lehrer fur die vielseitige Anregung meinen verbindlichsten Dank 
auszusprechen. 

Herrn Prof. Dr. Matsumura sage ich an dieser Stelle auch 
meinen besten Dank fur die Belehrung und das Interesse, welches 
er meiner Arbeit entgegengebracht hat. 



n. Methodisches. 

Bti unseren Versuchen koraraen stets die Reinkulturen in 
Betracht. Als Kulturgefasse wurden Erlenmeyer'sche Kolben 

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146 N. ONO : WACHSTHUMSBESCHLEUNIGUNG 

von ca 200 cc Inhalt angewendet, und zwar von gleicher Gestalt 
und Qualitat in einer Reihe von Parallelversuchen benutzt. Nach- 
deni die Gefiisse zuerst mit Salzsaure grundlich gewaschen worden 
waren, wurden sie rait Leitungswasser, dann mit destillirtem 
Wasser wiederholt ausgewaschen, getrocknet und gebraucht. 

Wasser. — Zur Zubereitung der Nahrlosungen und zum Auf- 
losen der Reizmittel benutzte ich doppeltdestillirtes Wasser. 
Das auf iibliche Weise gewonnene destillirte Wasser war von 
Glas zu Glas nochmals destillirt worden. 

Chemische Praparate. — Die als Nahrstoffe sowohl als 
auch als Reizstoffe dienenden Chemikalien stammten grossten- 
theils aus Merck's ,,garantirt reinen*' Reagentien. 

Nahrlosungen. — Fiir Algen bediente ich mich der bekann- 
ten Knop'schen Losung^), die ich nach folgeuder Vorschrift 
bereitete : 

(A) MgSO^+TH^O 10.25g (B) Ca(N03)2 20.00g 

KNO3 5.00,, 

KH2PO4 5.00,, 
Wasser 175.00cc Wasser 175.00 cc 

Beim Gebrauch wurden je 10 cc von A und B mit 880 cc 
Wasser verdiinnt. Diese bezeichne ich als Original-Nahrlosung 
fiir Algen. 

Die Original-Losungen fur Pilzkulturen bestanden aus fol- 
genden drei Serien : 



1) Aus keinem besonderen Grande benutzte ich hier Ca-haltige Nahrloeung. Die Ent- 
bebrlichkeit der genanuten Metalle bei Pilzen und niederen Algen ist bekanntlich in 
neuerer Zeit von Moliscli und Bene eke erwiesen worden. 



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EINIGER ALGEN UND PILZE. 147 

(A) 



KH2P04 




0.50 g 


MgSO, 




0.25 „ 


NH4NO3 




1.00 „ 


Eisen 




Spuren 


Rohrzucker 




5.00 g 


Wasser 


(B) 


90.00 CC. 


ie bei A. Asparagin 0.5g 


statt NH4NO3 1.0 g 



(C) 
Wie bei A. Dextrose anstatt Rohrzucker. 

Bei fast alien Kulturreihen wurde A angewendet, wahrend 
B und C nur ausnahmsweise benutzt wurden. 

KulturaDstelliing. — Ichgossin 5Kolbenje 135ccOriginal- 
Losung (Knop'schebezw. Rohrzuckernahrlosung). Sodann setzte 
ich zum ersten Kolben 15 cc destillirtes Wasser hinzu und Hess 
dies als Nahrlosung fur Controlkulturen dienen, zum 2*®°, 3*®°... 
o^°je 15 cc betreffend verdiinnte Losung von Reizstoffen, deren 
Wirkungen versucht werden soil ten. Bei den Algenkulturen 
geschah die Verdiinnung in absteigender geometrischer Reihe im 
Verhaltniss 1:5, bei den Pilzkulturen aber mit 1:2. Alle Kolben 
wurden darauf gut geschiittelt, um die Losungen aufs innigste 
zu mischen, und dann die in jedem Kolben enthaltene Losung in 
drei Kolben gleicbmassig vertheilt, so dass wir 3 Serien von 
je 5 Kolben, deren jede 50 cc Nahrfliissigkeit enthielt, vor 
uns haben. 



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148 N. ONO : WACHSTHUMSBESCHLEUNIGUNG 

Die auf diese Weise zubereitete Kulturfliissigkeit enthielt 
bei der Knop'sche Losungca 2.5X wasserfreie Salze und bei der 
Pilznahrlosung etwa 59^ Kohrzucker^). Dann folgte bei Pilzkul- 
turen die Sterilisation in einem Koch'schen Dampftopf, welche 
72 — 1 Stunde dauerte. 

Bei Pilzen fand die Impfung in iiblicher Weise statt, wah- 
rend ich sie bei Algen in der Weise ausfuhrte, dass ich mittelst 
Platindraht oder Pipette eine moglichst kleine Algenmenge aus 
den zuvor in Nahrlosung von derselben Concentration oder auf 
Agar bereiteten Reinkulturen herausnahm und in VersuchsgeSsse 
brachte. 

Die Kulturen wurden in Zinimertemperatur (ca 15° C im 
Mittel) ansgefuhrt, und in kalteren Jahreszeiten ins Treibhaus 
(16-21° C) gebracht. 

Die Kulturdauer variirte unter Umstanden zumeist zwisehen 
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 Erraittelung des Trockenge- 
wichtes der gebildeten Algen- bezw. Pilzmassen Aufschluss 
gegeben. Die Bestimmung wurde folgendermassen ausgefiihrt. 
Nach Beendigung der Versuche wurde die Kulturfliissigkeit 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- 
stabchens. Dann spiilte ich die Erntemasse mit kaltem destillirtem 
Wasser, um dadurch etwa noch vorhandene Nahrfliissigkeit 
moglichst 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 L<>siing wurde von Pfeffer und Richards vielfach benutzt. 



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EINIGER ALGEN UND PILZE. 149 

Trockengewicht der Emte ist in den angefiihrten Tabellen als 
Ernteertrag notirt. 

Bei den AlgenkuUuren geschah es vielfach, zumal bei 
denjeuigen, welche wahrend der kalteren Jahreszeit angestellt 
worden waren, dass die Vermehrung nur selir langsam vor sich 
ging, und dass nach monatelangem Stehenbleiben eine nur 
schwache Entwickelung sich zeigte. In solchen Fallen musste 
ich mich damit begniigen, durch das Aussehen der Kulturen die 
Starke der Entwickelung zu beurtheilen. 

Was nun die specielle Ausfiihrungsmethode anbelangt, so 
wird sie an geeigneten Stellen beriicksichtigt. 



m. Vorbemerknngen nber Versuchsobjekte. 

Fur Pilzkulturen bediente ich mich der gewohnlichen 
Schimmelpilze Aspergillus niger und Penidllium glaucum. 

Benutzt wurden bei meinen Algenversuchen die folgenden 
Formen : 

Protococcus sp. 
Chroococcum sp. 
Hormidium nitens. 
Stigeoclonium sp. 

Da wir zur Zeit fiber die Lebensbedingungen der Algen 
uberhaupt nuj wenig wissen, so war es mir nicht immer ge- 
lungen, die im Freien rasch wachsenden Algenarten im Labora- 
torium unbeschadigt gedeihen zu lassen. Besonders schwierig war 
die Aufgabe, die grosseren Formen in reinem Zustande langere 
Zeit in einer bestimmten Nahrfliissigkeit zu kultiviren. 

Nach einigen darauf beziiglichen Vorversuchen kam ich 



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160 N. ONO: WACHSTHUMSBESCHLEUNIGUNG 

schliesslich auf die oben genannten niederen Formeu zuruck, die 
ziemlich leicht rein zu erhalten waren, ausserdem sicheres Gedeihen 
zeigten und ferner leichtere KontroUe der Impfmasse gestatteten. 

Unsere kleineren Algen waren zumeist in den fiir grossere 
Algen bezweckten Kulturgefassen spontan aufgetreten, und so 
warden diese durcli wiederholte Uebertraguog rein gezuchtet. Die 
ersteren Algen liessen sich auch auf festera Nahrboden, welcher 
aus 72 Proc. Agar und 2.5 Promille Nahrsalz enthaltender 
Gallerte bestand, gut vegetiren und solcher Plattenkultur bediente 
ich mich bei einigen Beimpfungen. 

Iformidium nitenSy welches sich auf der Oberflache einer 
Vaucheria-KnltuY als eine charakteristische seidenglanzende 
Decke bildete, zerfiel nach dem Uebertragen in unsere Nahr- 
losung in einzelne Zellen und vegetirte als solche weiter. 

Es bleibt noch zu bemerken iibrig, dass die Vermehrung 
bei niederen Algen {Protococcus wurde zunachst untersucht) 
wahrend der kalteren Jahreszeit so gut wie vollstandig herab- 
gesetzt worden ist, wenn auch die Kulturen im Treibhaus bei 
16-20° C sich befanden. Die Ursache wolle man nifcht in man- 
gelndem Licht suchen, da dieselben Kulturen mit noch massigerem 
Lichtgenuss vor einem gegeii Norden gerichteten Fenster eine 
gute Entwickelung bis Anfang November zeigten. Ob es sich 
hier etwa urn eine Vegetationsperiodicitat handelt, beabsichtige ich 
im kommenden Jahre naher zu untersuchen. 



IV. Die Verandernngen in der Wachsthnmsweise 

and die Correlation zwischen Fortpflanzung 

und Wachsthum. 

Wie wird die Wachsthnmsweise einer Pflanze beeinflusst 
werden, wenn ihr Wachsthum bei Gegenwart von Beizstoffen 



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EI^IGEB ALGEN UND PILZE. 151 

fiber 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 Veranderung der Wachsthumsweise beobachten. Die 
Zellengrosse blieb unverandert ; so lag z. B. bei Protococcus sp. 
die Zellengrosse jedenfalls zwischen 7-10 fjt. Sie zeigte ferner 
keinen Unterschied in Amylumeinschliissen. 

Bei Pilzkulturen liess sich aber die Veranderung in der 
Wachsthumsweise raehr oder weniger schon makroskopisch er- 
kennen. So war bei den meisten versetzten Kulturen die 
Beschaflfenheit des Mycels ungewohnlich. Wahrend bei Kontroll- 
kulturen die Hyphe in Nahrlosungen durchsichtig und zart 
waren, bildeten diejenigen der versetzten Kulturen ein dickes, 
weisses, hautartiges Geflecht, welches bei langerem Stehen sich zu 
einer aufgeroUten Masse umgestaltete. 

Auch das Turgorverhaltnis 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- 
thatigkeiten dar. So kommt es nicht selten vor, dass bei einer 
Pflanze, die in iippigem Wachsthum begriffen ist, ihre Fortpflan- 
zungsfahigkeit zeitweilig suspendirt wird ; und wenn hingegen die 



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152 N. ONO : WACHSTHUMSBESCHLEUNIQUNG 

Fortpflanzung herabgesetzt worden ist, so schreitet das Wachsthum 
kraftig fort, 

Es war mir daher nicht ohne Interesse, diese Verhaltnisse 
in unserem Falle kennen zu lernen. Ich konnte jedoch das 
Studium nur bei Pilzen ausfuhren, nicht aber bei Algen, da 
ineine Versuchsalgen hauptsachlich einzellige Formen waren, die 
nur durch Theilimg sich vermehrten. 

Bei Pilzen hingegen war die Wechselbeziehung zwischen der 
Myeelentwickelung und der Sporenbildung deutlich zu erkennen. 

In fast alien Fallen iibten die Versuchsstoffe auf die Pilze 
Sporen- bezw. Conidienbildung verzogernden Einfluss aus. Be- 
sonders ausgepragt trat dies bei Zusatz von ZnSO^ und NaFl 
ein. Ich konnte vielfach constatiren, dass bei normalen, in 
Zimmertemperatur (ca 15^ C im Mittel) geziichteten Aspergillus^ 
Kulturen das Mycelium schon nach 1-2 Tagen sich ausbreitete 
und mit angelegten Sporangientragern versehen war, deren Kopfe 
nach weiteren 2 Tagen durch Reifung der Sporen ganz ge- 
schwarzt worden waren. Bei den Kulturen mit Zusatz von 0.005 
Proc. ZnS04 dagegen habe ich selbst nach Verlauf einer Woche 
vergeblich nach reifen Sporangien gesucht. In den letzteren 
Fallen fand ich nur auf dem ziemlich stark angewachsenen 
hautigen Mycel etwas Sporangientrageranlage mit etwas aus- 
geschwollenen Kopfchen, nebst einer Anzahl von wohl als Luft- 
mycel anzusehenden Gebilden. Erst nach weiterem einwochigen 
Stehen wurden sie mit braunlich-schwarzen Sporen besetzt. 

Die Grosse der Sporen, welche gegen verschiedener Einflusse 
sehr empfindlich ist, blieb in unseren Fallen unverandert. Die 
Beschaffenheit des Myceliums aber war nicht unbedeutend 
beeinflusst. 

Wie erwahnt, fand ich in alien von mir untersuchten Stoffen 



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£INIQEB ALQEN UND PILZE. 153 

mehr oder minder die Tendenz, die Sporenbildung zu verzogern 
oder wenigstens zu verspaten. Dies gilt ohne weiteres auch fur 
PeniciUium sowohl, wie fiir Aspergillus. 

Am ausgepragtesten und am schonsten aber konnte ich 
dieses Verhaltnis bei AsfpergilluS'rK.xxMxxv mit Zusatz von NaFl 
kennzeichnen. 

Ich nehme hier aus meinem ProtokoUe folgendes Beispiel : — 

I. Normal Ganze Oberflache mit schwarzen Sporen bedeckt. 

II. 0.00259^ NaFl Etwa Vs der Decke mit Sporen bedeckt, die tibrigen 
Vs steril. 

III. 0.005 ,, „ Nur sehr sparliche Sporenbildung, weiss. 

IV. 0.0 10 „ „ Steril, hautbildend, weiss, 
V. 0.021 „ „ Ganz steril, weiss. 

(Beobachtet nach 10 Tagen Beit der Aussaatzeit d^r Sporen.) 

AUe NaFl enthaltenden Kulturen zeigten eine uppige vege- 
tative Entwickelung des Myceliums und bildeten hautartige 
Decken. 

Nach weiteren zehn Tagen waren die Kulturen wesentlich 
onverandert im Aussehen. Bei III. sporadisches Auftreten der 
Sporangientrager 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 Eesultat: 

I. n. III. IV. V. 

0.314g 0.336g 0.385g 0.3l6g 0.274g 

(Fiir naheres vgl. Tabelle Pilze H) 

Eine Photographic am Ende dieser Arbeit veranschaulicht 
das eben gesagte Verhaltnis. 

Die Unterdriickung der Sporenbildung in diesem Falle kann 
allem Anschein nach eher dem direkt hemmenden Einfluss des 



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154 



N. ONO : WACHSTHUMSBESCHLEUNIGUNG 



betreffendeu Stoffes zugeschrieben werden, als der durch starkerc 
My celentwickelung hevorgerufenen Correlationserscheinuug, welche 
haufig bei gunstigstem Nahrboden zu Stande kommt. Um diesec 
Pankt aufzuklaren, stellte ich die Versuche derart au, dass id 
ein Stiick Mycelium, welches fiir 19 Tage in 0.0159^ NaFl Losunj 
geziichtet worden war und keine eigentliche Fruktifikation, 
abgeselieu von einigen schon besprochenen haarigen Gebilden 
aufnimnitj nach Bespulung mit Wasser in Normallosung brachte 
Schon uach 2 Tagen kamen die angelegten Trager zur Reifi 
imd schwiirzten, wahrend ich auf der noch in 0.01596 verblei- 
benden Decke vergeblich nach den reifen Sporangien suchte 
Dieses Resultat spricht ohne weiteres fiir die gesagte Ansicht. 

Es fragt sich nun, ob die starkere Entwickelung des vege- 
tativea Organs der Pilze bei der Zugabe einer kleinen Dosii 
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 befdrdert. Es ist schon bestatigt wurden, dass Sporenbil- 
dung der Pilze durch einige Gifte^) , viel empfindlicher beeinflussi 
wird als die vegetative Mycelentwickelung. Fasst man dieses 
Verlialtnis ins Auge, so ist es wohl begreiflich, dass bei einei 
geniigenden Verdiinnung der angewandten Stoffe jene Concentratior 
erreicbt ist, welche an sich fur die Mycelentwickelung unschad- 
lich, aber fiir die Sporenbildung hemmend wirkt, und dass sc 
infolge der Correlation das Wachsthum des vegetativen Theilj 
ungewohnlich gesteigert wird. Dieser indirekten Wirkung dei 
betreffenden Stoffe schreibe ich, ausser dem direkten Eeizeffekt 
die Wachfithumssteigerung zu. 

1) O. Loew, Ein natilrliches System der Giftwirkungcn 1893. 




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EINIGER ALGEN UND PILZE. 155 

V. Einflu8s der Reizstoffe auf die Betriebsstoffwechsel. 

Die Pflanze nimmt durch ihre Lebensthatigkeiten die Nahr- 
stoffe auf und verwendet einen Theil derselben zum Aufbauen ihres 
Korpers, 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 Stoflfumsatze sich, wie bekannt, in 
zwei Kategorien : Bau- und Betriebsstoffwechsel trennen. Um 
die Grosse jedes von diesen Stoffwechseln zu ermitteln, hat man 
einigermassen einen Maasstab. So ist bei der BetriebsstoffiYechsel- 
thatigkeit das Trockengewicht maassgebend, und fiir den Betriebs- 
stoflFwechsel giebt die Ermittelung der Kohlensaureproduktion, die 
Ausscheidung gewisser Stoflftvechselprodukte, einige Aufschliisse. 

Wie aus der in der Einleitung angefiihrten Skizze zu ersehen, 
beziehen sich die bisherigen Untersuchungen fiber die Erhohung 
der Lebensthatigkeiten durch chemische Reize zumeist nur auf 
BaustoflFwechsel. Richards untersuchte in seiner Arbeit nur den 
Emteertrag, berucksichtigte aber nicht den Betriebsstoflfwechsel. 
Schulz beobachtete starkere Entwickelung der Kohlensaure bei 
Hefen. Beim ersten Anblick scheint dies auf nur erhohtem 
Betriebsstoffwechsel zu beruhen, doch blieb hier die Frage immer 
ofifen, ob man es in diesem Falle mit der Erhohung der Gahr- 
thatigkeit einzelner Individuen zu thun hat, oder ob diese 
Erscheinung von der durch die Reizwirkung verursachten Ver- 
mehrung der Gahrungserreger bedingt sei. 

Eb fehlen bisher meines Wissens einschlagige Versuche fiber 
die Beeinflussung des Betriebsstoffwechsels in Gegenwart von 
giftigen Stoflfen. Irgend ein Beitrag in dieser Richtung durfte 
wohl nicht ohne Interesse sein. 



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156 



N, ONO: WACHSTHUMSBESCHLEUinGUNG 



Fiir solchen Zweck bieten die Algen keine geeigneten Objektc 
dar, woH aber die Pilze, welche sich dafiir bequem anwenden lessen, 

Einige Pilze, insbesondere Aspergillus niger^ produciren eine 
nicht unbetrachtliche Menge Oxalsaure als Stoffwechselprodukt, 
wie aug der bekannten Arbeit Wehmer's^) hervorgeht. Diese 
Stoffwechselprodukt bot bei meinen Versuchen einen Angriffspunkt 
und BO warden eine Reihe Bestimmungen fiber die Saurenmeng( 
angefiibrt, Ich muss hier bemerken, dass ich die Saure nur am 
titrimetrisclien Wege bestimmte. Die Methode ist untauglich, weni 
Oxalsaure nicht nur als solche, sondern auch als Salz vorkommt 
Well me r zeigt aber in seiner oben besprochenen Arbeit, dassii 
NH^NOs-haltiger Zuckernahrlosung Oxalsaure stets als freie Sauri 
bei A8peTgiUuB''^vlt\xre£L auftritt und da meine Kulturen haupt" 
saclilicb derartige waren, so war die Titration zuverlassig. Di« 
geschah mit Liquor Alkali Decinormalis und Phenolphthaleii 
als Indikator. 

Nachdem ich von der durch Titration ermittelten gesammtei 
Saure die ursprungliche Aciditat der Nahrlosung subtrahirt hatte. 
rechnete ich diese als Oxalsaure um, welche in der angefuhrter 
Tabelle gegeben ist*). 



1) Wehmer, Entstehung und physiologische Bedeutung der Oxalsaure im Stoflfwechael 
einig^r Pilze, Bot. Ztg. 1890. 

SJ Ilier weide ich einige Versuche, um die titrimetrisch ermittelten Zahlea mit denjenigen, 
die gmvtmetrisc'h bestimmt waren, zu vergleichen, ausgefuhrt angeben: — 
Sanrenmenge in g in 10 oc Nalirfliissigkeit. 
Ztisatz von NiSO^ Titration Gravimetriscli 

0.042 0.048 

0.003^ 0.052 0.060 

0.007,, 0.060 0.068 

0.014,, 0.066 0.075 

0.028,, 0.064 0.060 






0.042 


0.040 


0003,, 


0.047 


0.048 


0.007 „ 


0.052 


0.058 


0.014,, 


0.060 


0.065 


0.028,, 


0.051 


0.054 



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EINIGER ALGEN UND PILZB. 157 

Vergleicht man nun die Saurenmenge bei Gegenwart von 
Reizmitteln mit derjenigen der Kontrollversuche, so findet man in 
unseren Versuchen nur mit der einzigen Ausnahme von NiS04 
stets das Minus im ersteren Falls. Dieses Verhaltnis ist ersicht- 
llch aus der Colonne „Saure pro Ig Pilzsubstanz" in den Tabel- 
len. Steigt der Zusatz von Reizstoffen, so wird die Menge der 
Saure um so kleiner. 

Man kann jedoch nicht annehmen, dass die aufgefiindene 
Saurenmenge die samtliche Menge der ausgeschiedenen Saure 
darstellte, da bekanntlich die Ausscheidung der Saure mit ihrer 
Zersetzung Hand in Hand gehen soUte. Daraus geht hervor, dass 
die Erklarung der besprochenen Verhaltnisse nicht einzig in ihrer 
Art sein kann. Es konnten einige Moglichkeiten, welche fiir 
diese Thatsache sprechen, angegeben werden. 

Erstens, wenngleich die Oxalsaure als normales Stoff- 
wechselprodukt unseres Pilzes auftritt, ist sie doch als ein Pro- 
dukt unvollkommener Oxydation anzusehen, und wenn die Stoff- 
wechselthatigkeit auf einmal gesteigert wird, so wird als das 
Produkt vollkommener Oxydation mehr Kohlensaure entstehen, 
dagegen weniger Oxalsaure. 

Zweitens konnte dieselbe Erscheinung auftreten, falls die 
einmal entstandene Saure durch Wiederverarbeitung seitens der 
Pilze verschwindet. Dabei konnte sie entweder als Baumaterial 
wiederaufgenomraen werden oder, ohne wieder den Pilzen nutzbar 
zu werden, zersetzt werden. Doch, wie schon Wehmer^) betont 
hatte, stellt die Oxalsaure einen nur sehr armen Nahrstoff fur 
Aspergillus dar, so dass es wahrscheinlich ist, dass sie bei 
zureichendem Vorrath von guten Nahrstoffen wie Zucker^) wohl 

1) Wehmer I.e. 

2) Bei meinem Versuche betrag der Zackergehalt nach Beendigung der Yersache wenig- 
Btens 1.5 g in je 50 cc der KultarfliisBigkeit, d. h'. ca 3^. 



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N. ONO: WACH8THUMSBE8CHLEUNIGUNG 



I 



II ! 



intakt geblieben ware. Ferner wurde von Wehmer^) constatirt 
dass weder Licht noch tote Pilzmasse allein die Zersetzung de 
Siure hervorzurufen im Stande sind, wohl aber die Lebenstha 
tigkeiten der Pilze. Es bleibt daher nur die Annahme iibrig 
dass die Saure durch Steigeruog des Betriebsstoffwechsels lebhaft 
erer Zersetzung unterworfen worden sei. 

Die dritte Moglichkeit ist schliesslich die, dass diejenigei 
Stoffe (Kohlenhydrat u. s. w.), welche bei normaler Wachsthums 
energie durch Stoflfwechsel z. Th. als Oxalsaure auftreten, b( 
der infolge der Reizwirkung fiber Norm gesteigerten Wachthumg 
thiitigkeit nicht als jene Form abgesondert werden, sondern sic 
gerade in den integrierenden Theil des Pilzkorpers umwande] 
ten, kurz, dass sie als Baustoff verwendet werden. 

Von einer anderen Seite mussen wir also dieses Problem an 
greifenj um zu entscheiden, ob die eine oder andere von diesen Mog 
lichkeitea fur unseren Fall zutriflft. Die Ermittelung der Kohler 
Siioreausscheidung, des okonomischen CoeflBcienten^) u. a. wird wol 
an diesen Punkt anschliessend oder wenigstens rathgebend seii 

Im Folgenden gebe ich die Resultate meiner Bestimmunge 
okonomischer Coefficienten bei den Kulturen mit Zusatz vo 
ZnSOj, bei denen die Erntezunahme stets am auffallendsten wa: 

Die Bestimmung des Coefficienten fand in folgender Weis 
statt : 

Die Kulturflussigkeit wurde zunachst durch andauernd( 
Koch en mit verdunnter Salzsaure vollkommen invertirt. Dan 
verdihinte ich diese bis zu etwa Ya 9^ Zuckergehalt. EineBureM 
wurde mit der betreffenden Losung gefullt. 

In einem Kolben mischte ich genau 10 cc Fehling^sch 

1) Welimer l.c. 

2) Mim yergleiche hieriiber H. Kunstmann, Ueber das Verbal tniss zwischen Pilsern 
uxiid verbrauchter Nahrung. 1895. (Leipziger DisBertation). 



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EINIGER ALGEN UND PILZE. 



159 



Losung mit etwa 40 cc Wasser und brachte es zum Sieden ; darauf 
fugte ich die obeu genannte Losung hiozu, bis schliesslich durcb 
vollkommene Reduction des Kupfers zu Kupferoxydul die 
Flussigkeit farblos geworden war. 

Da unsere Fehling'sche Losung in 1000 cc 34.64g Kupfer- 
mlfat, 174g Kaliumnatriumtartrat und 120g Natriumhydroxyd 
juthielt, so sollte je 10 cc derselben durch 0.05g Zucker reducirt 
irerden. 

Nun kann man leicht durch die Lesung der Burette die 
iuckermenge in der Kulturfliissigkeit kennen lernen. Die 
)ifferenz zwischen der urspriinglich vorhandenen Zuckermenge 
D der Kulturfliissigkeit und der zuinickbleibenden ergiebt selbsU 
erstandlich die verbraucbte Zuckermenge. 

Kulturen mit Zusatz von ZnSO^. 

AspergUlus niger. 
Kulturdauer 14 Tage. Zimmertemperatur. 



Gehalt an ZnSO. 
(Gew. %) 


Pilzemte in g 


Verbrauchte 
Zuckermenge 


Okonomischer 

Coefficient 
J . Verbrauch 




I 









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 




I] 


[. 







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 



»)l 



* 



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N, OXO : WACMSTHUMSBESCHLEUNIGUNG 



III. 






0.39-2 


1.819 


4.6 


0.0037 


0.910 


2.462 


2.7 


0.0074 


0.908 


2.456 


2.7 


0.0148 


0.844 


2.456 


2.d 


0.0297 


0.827 


2.446 


2.8 



Was sich nun aua cliesem Kesultate beurthdleu liisa 
dass der okouomisclie Coefficient in jedem Falle bei \ 
grosser ist in Kontrolle dJi- in niclit zugegetzter Kulliir 
zugesetzter, Dieses Verhaltnis deiitet also au, dags die Pil 
Auwesenbeit von Zinksnlfat veranlaast wurden, mit einen 
haltnismassig kleineo Verbniueb vou Zucker eine bed* 
grossere Kurpersubstanz aafbauen zu kdnuen- So scheini 
weoigsteiis fiir Zioksulfat^ von den oben besprocbenen 
Mogliclikeiten die dritte die wirklicbe zu sein* 



1 

I 



IV. Spedelle Beaprechuiigeu, 

ZnSOi. 

Unter den von Richards geprtifteu Stoifen iibt diesei 
die Btarkste Wirkung aus, 

Auch bei unseren Versuchen mit Algen wirkte ZnSOi j 
FeSOi sehr gunstig auf das Wachsfchum ein. Scbou bei ! 
von ciner minima len Quantitiit, wie O.OOOCH695, nahm die 
etwas ra, nod dies war noch deutliclier bei 0,000069^ hm 0.0 
Stieg die Concentration auf 0.001096, so litten die Algen 
unerheblich, ohne jedocb das Wacbsthum ganz herabzu 
(cf. Tabelle. Algen A, I-TV). 

Uusere VerBucbe mit Pilzeti Btimmen mit doujeidge] 



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EINIGER ALQEN UND PILZE. 



161 



Richards iiberein. Bei langerem Stehen wurde der Unterschied 
zwischen den Versuchs- und Kontrollekulturen recht iiber- 
raschend (cf. Tabelle. Pilze A. I-III). 

Sehr sonderbar trat einmal bei einer Versuchsreihe mil 
Dextrose es hervor, dass keiu nennenswerther Unterschied in 
der Ernte sowohl, als auch in der Saurequantitat sich erkennen 
liess. Den Qrund davon kann ich aber nicht erklaren (Tabelle. 
Pilze. A. IV.) 

Die gelbliche Farbung von Nahrfliissigkeiten, sowie die 
Bildung der braunlichen Sporen in den Versucbskulturen, welche 
schon von Autoren besprochen wurden, waren hier bemerklich. 

Die Siiurenmenge nacli Beendigung der Versuche war in 
Versucbskulturen viel kleiner als in Kontrollen (Tab. Pilze. 
A. I-III.). 

FeS04. 

Richards giebt an, dass dieses Salz erst bei ziemlich gros- 
sem Gehalte einen schiidigenden Einfluss ausiibt. 

Meine betreflfenden Versuche mit Algen zeigten auch, dass 
iasselbe noch hohere Concentration im Vergleich zu anderen 
Schwerenmetallsalzen ertragt. So lag bei Hormidium das Optimum 
3twa bei 0.0005?^ und sogar bei einer hoheren Concentration 
me 0.01269^ war der Ertrag noch etwas grosser als bei den 
Kontrollen (cf. Tab. Algen. B. I-II.). 

In einer mit Zusatz von FeS04 angestellten Penidllium' 
Kultur trat merkwiirdigerweise das ziegelroth geia,rbte Mycelium 
zu Tage. 

NiS04. 

Bei Algen ruft der Zusatz von NiSOi einen befordernden 
Einfluss hervor. Die optimale Dosis lag etwa zwischen 0.00006 



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N, ONO : WACHSTHDMSBESCHXEUNIGimQ 



und 0.000129^S, wiihreud 0.0028?^ eine beschadigeude Wirki 
ausiibte (Tab. Algen C L II,)- 

Saureprodiiktion bei Pikkultureii war hier iiii GegeusaU 
den lueisten FalleQ grosser rait der Erhuhuug der Zusiitzeproa 
(c£ Tabelle Pilze, C Mil.). 

C0SO4. 

Bei Algen scheiiit dies aiich eineo begiinatigeuden Einf 
auszuiibeu, dock lag der optimale Puukt etwas oiedriger als 
NiSO, ; Optimum etwa bei 0.0001 2?S (cf. Tab, Algen D. L 1 

SaureerzeLiguug war wie gewohnlicli kleiner und zwar £ 
regelmassig in Versuchskultureu, 

CuSOi wnrde von Richards niclit untersucht Im Js 
1897 constatirte Qunther*)^ dass Kupfersalze in grossc 
Mengen das Wachstlium der Pilze retardirten, in geringt 
Mengen dagegeo bessercs Gedeihen mit sich bringen* A 
bei meinen niit Asperffillm und Penicillmni angestellten "^ 
sucben beobachtete icli dieselbe Erscbeinung- Hattori^) f 
auch in seinen Untersuchungen (iber die Giftwirkung der K 
fersalze eine abnliche Thataache. 

Hier werde icb zwei Beispiele angeben ; fiir naheres verw 
ich auf die tabellarische Zusammenstellung (Tab. Pilze. E.). 



Asper^iilus niger. 



Gtlmlt an CtiSO., 





0.00159^ 


O.OOSji 


O.OO696 


0.0): 


Erateertrag in g 


0.273 


0.307 


0.313 


0.324 


0.34. 



1) E. GurilJit'r, BeHmg zur miiniralist^li*;!! Ndmiug der Pilze. ErL-mgen 1897, 

2) H. Hattori, Ueber die Einwirkung doB KupferBulfatea auf cioige Pflj 
AtlinuskriyL 



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EINIGER ALQEN UND PILZE. 



163 



Penidllium glaucum. 



Gehalt an CuSOi 





0.00159^ 


0.0039i 


0.006?^ 


0.0129i 


Ernteeitrag in g 


0.213 


0.320 


0.338 


0.359 


0.410 



Bei Algen konnte ich dagegen keine Wachsthumsbef5rde- 
ruDg nachweisen, wie aus der Tabelle ersichtlich ist. Schon bei 
[).0000l9i steht die Ernte etwas zuruck (Tab. Algen E.). Ob 
bei noch weiterer Verdunnung die wachsthumsbegiinstigende 
CJoncentration erreicht sein koDnte, lasse ich vorlaufig unbestimmt. 

Die Saurenmenge in Pilzkulturen war kleiner in Versuchs- 
iulturen (Tab. Pilze. E.). 

HgCL. 

Es ist von gewissem Interesse, dass dieses heftige Gift in 
jeniigender Verdunnung auch das VVachsthum der Pilze befordert. 
5chulz^) giebt an, dass Kohlensaureentwickelung der Hefe in 
jegenwart 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 
Liitteratur nur die Rede von dem schadigenden Einfluss des 
)etreffenden Stofls. Raulin^) betrachtet z. B. dies mit AgNOg, 
^i^GX^ zusamraen als das giftige Salz fiir Aspergillus. Er gibt 
1/512 000 als die Grenze der Giftwirkung. In seinem Experi- 
uente mit 1/819 200 konnte er jedoch keinen wachsthums- 
)e8chleunigenden Einfluss beobachten. Meines Wissens liegt 
ins zur Zeit kein Versuch vor, welcher die letztgenannte That- 
lache in positivem Sinne zeigt. 

1) H. Scbulz, I.C. 

2) Banlin l.c. p. 134. 




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III 



Meinem Versuclie naeh (ef. Tab, Pilze, F,) tritt echo 
Verdiinnimg von 0*001 7 ?£ oder 1/60 000 ziemlieh giite Entvs 
lung von Aspergilhis eiiK Stieg die Concentration auf 1/3C 
BO kam die EDtwickelnng zum Stillstaod, Die Grenze fiir 
wirkung liegt zwiscben 1/GOOOO und 1/30000, 

Dag Optimum war sowolil bei PeinciUium als auc 
A^jergillus etwas unter OiJOlS?^). 

Hier gebe ich zwei Beispiek (cf. Tab, Pilze. F. I-VI] 

Aspergillus niger. 



Gehalt an HsClj 


' 


0.0003 ?i 


0.000795 


0.0013?^ 


O.OC 


Ernteertrag in g 


0.261 


0.355 


0.354 


0.509 


0.4^ 


Penieillium glaucum. 


Gehalt uii HgUlj 





O.OOOB^i 


0.0007 9^ 


0.0013?^ 


O.OC 


Ernteertrag lii jj 


0.183 


0.24D 


0.213 


0.311 


0.2^ 



Saureprodtiction ist hier wie bei den nidston Fallen klein 
Versuchekultuien als bei Kontrollen ('I ab* Pil^e. F,). 

Auf Algen iibte dieses Stilz keinen beachleynigenden Ei 
au8, sondern wirkte nur giftig ein. Schoii bei 0.00005?^ w? 
scbadigende Effekt deutlicb zu erkennen. Doch weitere 
diinnung durfte icb nicbt ausfiibren, da bei solcben 1 
Verdiinnungen einige Feblerquellen als maa&gebeiid aufi 
(Tab, Algen F.). 

Von Richards wurde Li CI zur Untersucbuog heransge 



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EINIGER ALGEN UND PILZE. 



165 



nd in diesem Salze ein eine betrachtliche Wachsthurassteigerung 
ervorrufender Stoff gefunden. Bei meinem Versuche benatzte 
A LiNOs mit gleichem Resultate (Tab. Pilze G.) 

Saureproduktion war hier auch kleiner in Versuchskulturen 
Is in Kontrollen. 

Algen zeigten auch eine besseres Gedeihen in zugesetzten 
Lulturen (Tab. Algen H.). 

NaFL 

Dieser StoflF iibte auch eine beschleunigende Einwirkiing auf 
Igen aus. Der optimale Punkt liegt etwa bei 0.000039^, stieg 
ie Concentration zu 0.000169^ bis 0.0008 9^, so nahm die Ernte 
was ab, war noch grosser als bei Kontrollen. Erst bei 0.00429^ 
eht der Ertrag im Vergleich zur Kontrolle etwas zuriick 
rab. Algen G.). 

Bei Pilzen beforderte dieser Stoflfdas Wachsthum (cf. Tab. 
ilze H.). Seine Wirkung auf die Sporenbildung wurde schon 
Q vorstehenden Capitel behandelt. 

Die Saurenmenge in der Nahrfliissigkeit war wie gewfihnlich 
leinei in zugesetzten als in Kontrolle-Kulturen. 

Arsen. 

Von Arsenverbindungen ist arsenige Saure giftig, doch ver- 
ragen hohere und niedere Pflanzen viel Arsensaure^). 

Da Arsenigsaureanhydrid nur schwer loslich ist, so bediente 
ch mich des arsenigsauren Kaliums. 

Bei Penicillium'K.ultuT war kein bedeutender Unterschied 
ler Ernte sowohl als auch in der Saurep reduction bemerklich. 

1.) O. Loew, System der Qiftwirkungen. 



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Merkwiirdig war hier eine eigentliche Geruclisentwickeliu 
zugesetzten Kolturen^). 

Auf Algen scheinen die genannten Siilze ctwas Waclistl 
begunstigeod 2:11 wirkeu, (Tab, Alg. I)* 



Tn. Schlussbemerkungen und ZnsammeafasauBg 
dcr Eesultate* 

Aus dem Vorstebenden gebt zunachst bervor, das 
chloropbyllfiihreiKlen niederen Organismen wie Algen in i 
Gedeihen giinstig beeinflusst werden diirch einen geringeii 5 
von einigen Stofien, welche fiir sich nicbt Niihrstofie bu 
sogar giftig wirken. In dieser Reaktion verhalten sicl 
Algen gerade me die Pilze. Nur ist zu bemcrken, das 
opt! male Dosis fur Algen viel kleiner als bei Pilzen ist 
Tbatsache, welcbe viel lei cbt vom oekologiscben Standpunkt 
Ihren Aufschlues haben wird. Von den gepruften Stoffen k 
icb nur bei Quecksilbercblorid und Kupfersulfat die bespro 
Reaktion nicbt constatiren, indem icb bei ibnen, soweit ' 
Versncbe reicbtenj stats Giftwirkung beobachtete. Damns 
aber nicbt geschlossen werden, dass den beiden Stoffe 
naniliche Eigenschaft nicbt zukommt, da man bei ibnen 
Umstauden doch noch jene wacbstburasbegiinstigende Einwii 
wohl erwarten kann. 

Bei Pilzenjlkonnte icb die friiberen Versucbe Ricli^ 
bauptsa-chlich bestatigen, dazu priifte icb rait positiven Resu 
einige bisber nocb nicbt untersuchte Stofle. 

Die Verzogerung oder Verspiitnng der Sporenbildun 
unseren Versucben ist nicbt als infolge eioer iippigen vegeti 

1) Sclion von Gasio (Jabreaber. fiber Giibningsorgaaisinen 1803) erSrlert- 



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EINIGER ALGEN UND PILZE. 



167 



Sntwickelung verursachte Correlationserscheinung, vielmehr als 
lurch Reizstoffe bewirkte Hemmung zu betracLten. 

Was nun die Art und Weise der Reizwirkung anbelangt, 
bemerke ich folgendes : 

Wenn es sich hier zuuachst um zeitliche oder andauernde 
lyperaesthesia handelt, so muss Bau- und BetriebsstofFwecLsel 
leichzeitig gesteigert werden. 

Wenn aber dagegen durch Zusatze der Reizstoffe die Thatig- 
eiten seitens des Organismus so gesteigert werden, dass sie mit 
leinerem Energieaufwand die Nahrstoffe in sich aufnehmen und 
ch bauen, kurz, okonomisch arbeiten konnen, so kann der dyna- 
ische Stoffvvechsel nicht so erheblich beeinflusst bleiben. Um 
iher in dieser Hinsicht eine richtige Auffassung zu gewinnen, ist 
n Einblick in den Betriebsstoffwechsel von Wichtigkeit. Einige 
)n meinen Versuchen in dieser Richtung zeigten andeutungsweise, 
iss Betriebsstoffwechsel nicht parallel mit Baustoffwechsel 
jsteigert werden ; doch sind zur Zeit meine diesbeziiglichen 
ersuche leider unzureichend, um in bezug auf diesen Punkt 
Ilgemeines zu sagen. 

Zum Schluss seien im Folgenden die wichtigsten Resultate 
irz zusammengestellt : — 

1. Das Gedeihen der niederen Algen wird durch Einfiih- 
mg gewisser giftiger Stoffe in hochst verdunnten Zustanden 
3gunstigt. Hierzu gehoren ZnS04, NiS04, FeSO^, C0SO4, NaFl, 
iNOs, K2ASO3. 

2. Die Erntezunahme bei Algen muss auf die vegetative 
ermehrung der Individuenzahl zuriickzufiihren sein, da keine 
ennenswerthe Veriinderung der Korpergrosse bemerkbar war. 

3. Die geeignete Dosis ist bei Algen bedeutend kleiner als 



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N. ONO : WACHSTHUMSBE8CHLEUNIGUNG 



i 



Hi 



bei Pilzcn. Schon der Zusatz von 10"* Gr. Mol. Salz wirkte in 
den allermeisten Fallen scLadigend. 

4. In CUSO4 und HgClg fand ich, soweit unsere StudieD 
ausreichen, keine beschleunigende Wirkung auf Algen, wohl 
aber begunstigend bei Pilzen. 

5. Bei Pilzen tritt durch Zusatze von HgCl2 (Optimum etwa 
bei 0.00139^ und CUSO4 (Optimum etwa bei 0.01294) die Wachs- 
thumsbeschleunigung ein. 

6. Die Saurequantitat in Kulturen rait Zusatz von ZnS04 
C3S|>804, HgClg, NaFl, CUSO4 war stets kleiner als in Kontrol 
kulturen, Nur verhielt NiS04, soweit meine Versuche eii 
Urtbeil gestatten, sich diametral entgegengesetzt. 

7. Die gepriiften Stoffe (speciell ZnS04 und NaFl) neigei 
dazu, die Sporenbildung der Pilze direkt zu hemmen, wenigsten 
das Auftreten der Sporen zu verspaten. 

*S, Die oekonomischen Coefficienten in ZnS04-Kultur sin( 
in der Kontrolle, d. h. in der nicht zugesetzten Kultur, be 
weitem grosser als in der zugesetzten. 



Jtmi, 1899. 



Botanisches Institut 
Kaiserl. Universitat 
zu Tokyo. 



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EIHIGEB ALQEN UND PILZE. 



169 



TABELLARiSGHE ZU8AMMEN8TELLUNG. 



M 



BEMERKUNGEN: 

VcrBuclie mit Algen — Der Entwickelungsgrad ist entweder mit Erntegewicht ocler mil 
lien relativen Werth gebenden Zifiern bezeichnet. 

Versucbe mit Pilzen — In Colonne „Aciditat'' ist die Quantitat des Decinormul AlkuU^ 
in cc gegeben, welche 10 cc der Nahrfliissigkeit neutralisirte (Die ureprungliclxe Afldi*rit 
wrurde naturlich vorher gubtrahirt). Daraus ermittelte ich die Sdurenmenge in je bO vc Niilir- 
Siissigkeit, berechnete sie als Oxalsaure und in Colonne „al8 Oxalsiiure umgerccbnet** nngub. 

In Kolonne „Saure pro Ig Pilzsubstanz" ist das Verhaltnis ^^^^^ gegeben. 

Obwohl bei einigen Peniotfmm-Kulturen die Stiurenmenge g^eben sind, durftc kh ducb 
licht die £^^ ermittein, da bei PenicUlium das Titration unzuverlassig scheini. 



I. Versuche mit Algen. 



A. I. Kulturen mit Zusatz von ZnSO«. 
Ernteertrag in g. Pro(o(X)CCw sp— angestellt 5. Oct 



Zimmer-Tem ] tt* ra t;i r. 



.s 


Gram Mol. 





ixlO-« 


ixlO-' 


i X 10-^ 


Kultur- 
ilaiier 


C5 


Gew. 9^ 





0.000014 


0.00014 


0.0014 


I 
II 

m 


0.010 
0.016 
0.012 


0.018 
0.023 
0.019 


0.018 
0.018 
0.021 


0.002 
0.009 
0.006 


23 Tftge 
26 ., 





A. 

Ernteertrag in g. 


II. Ealtaren mit Zusatz ^ 
Proloeoeeus gp.— angestellt 


ron ZnSO.. 
11. Oct 


Zimnier-Temi lural nr. 




Gram 
Mol. 





^xlO- 


i X 10-» 


10-' 


ixlO-* 


KultH- 
daiier 


G«w. e/g 





0.000012 


0.00006 


0.0003 


0.0014 


I 

II 
m 


0.035 
0.032 
0.030 


0.043 
0.040 
0.040 


0.038 
0.036 
0.037 


0.042 
0.040 
0.042 


0.024 
0.023 
0.020 


44 TagL- 

4G „ 
50 ., 



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170 N. ONO : WACHSTHUMSBESCHLEUNIGUNG 

A. IIL Kulturen mit Zasatz von ZnSO«. 
Ernteertrag in g. aroococcwwi— aDgestellt 24. Dec. j^ Treibhaus 16-20*' C. 






Gram 
Mol. 





ij^xlO-« 


I X 10-" 


10-' 


ixlO- 


Kultur- 
dauer 


1 


Gew. 9i 





0.000012 


0.00006 


0.0003 


0.0014 


I 

II 
III 


0.006 
0.009 
0.010 


0.015 
0.021 
0.024 


0.026 
0.022 
0.026 


0.022 
0.024 
0.026 


0.006 
0.010 
0.009 


71Tage 

ft 



A. IV. Kulturen mit Zuaatz von ZnSO^. 
Ernteertrag in g. Pro^ococcu^-angestellt 5. Feb. i^ Treibhaus 16-20° G. 



.2 1 Gram 

a , Mol. 





^xlO-' 


^xlO-« 


10-" 


^ X 10-^ 


Kultur- 
dauer 


1 Gew. 9^ 





0.000012 


0.00006 


0.0003 


0.0014 


I 
II 

m 


0.008 
0.009 
0.011 


0.017 
0.018 
0.015 


0.019 
0.023 
0.018 


0.019 
0.016 
0.017 


0.007 
0.009 
0.005 


65 Tage 

tf 



£. I. Kulturen mit Zusatz von FeSO«. 
Ernteertrag in g. Hoi-midium ntten/»— angestellt 7. Oct Zimmer-Temperatur. 



a 

C3 


Gram 
Mol. 





^xlO- 


ixlO-^ 


lO-" 


i+io-" 


Kultur- 
dauer 


Gew. 9^ 





0.0001 


0.0005 


0.0025 


0.0126 


I 
II 

III 


0.038 
0.028 
0.031 


0.072 
0.083 
0.070 


0.074 
0.079 
0.082 


0.080 
0.062 
0.074 


0.042 
0.041 
0.039 


65 Tage 

» 



B. n. Kulturen mit Zusatz von FeSO^. 
Ernteertrag in g. Honnidmm niterw-angestellt 10. Nov. j^ Treibhaus 16-20° C. 



.3 

o 


Gram 
Mol. 





^xlO- 


ixlO- 


10-^ 


i X 10-» 


Kultur- 
dauer 


Gew. 0^ 





0.0001 


0.0005 


0.0025 


0.0126 


I 

II 
III 


0.025 
0.023 
0.027 


0.050 
0.067 
0.064 


0.0.52 
0.049 
0.058 


0.054 
0.042 
0.046 


0.041 
0.032 
0.032 


79 Tage 

J) 
V 



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EINIGER ALOEN Uin> PILZE. 



171 



C. I. Eulturen mit Zusatz von NiSO«. 





£niteertrag in g. 


Chroococam 


I— angestellt 2. Oct. 


Zimmer-Temperatur. 


.s 




Gram 
Mol. 





5^x10- 


^ixlO-» 


io-» 


ixlO-« 


Kultnr- 
daner 


Gew. 0^ 





0.000012 


0.00006 


0.00028 


0.0014 


I 

II 
m 


0.012 
0.011 
0.013 


0.012 
0.015 
0.018 


0.025 
0.024 
0.020 


0.021 
0.020 
0.022 


0.004 
0.006 
0.007 


64Tage 




C. n. Kulturen mit Zusatz von NiSO^. 
Ilormidium ntVerw— angestellt 16. Nov. 


in Treibhaus 16-24" C. 


a 

1 

C5 


Gram 
Mol. 





AxlO-» 


I X 10-» 


10-' 


ixlO-* 


Kultur- 
dauer 


Gew. o/o 





0.000012 


0.00006 


0.00028 


0.0014 


I 
II 
III 


4 
4 

3-4 


5 
5 
5 


4-5 

4 

4-5 


4 
4 
4 


1 
1 
1 


70Tage 




N.B. Die ZiSer zeigt den Entwickelangagnd. 

D. I. Eulturen mit Zusatz yon CoSO«. 
HomUdium nitena — angestellt 9. Dec, 


in Treibhaus 16-24° C. 


a 

1 


Gram 
Mol. 





^xlO-» 


JxlO-" 


io-» 


ixlO-" 


Kultur- 
daner 


Gew. Oyi 





0.000012 


0.00006 


0.0003 


0.0014 


I 

n 
III 


4.5 

4 

4 


5 
5 
5 


45 
4.5 
4.5 


4 

3.5 

4 


2.5 

2 

2 


70Tage 


Ernteertrai 


N.B. I>ie ZiScr zeigt den Entwickelungagnd. 

D. II. Eulturen mit Zusatz von CoSO,. 
2 in g B-o/oooeetu— angestellt 19. MaL 


Zimmer-Temperatur. 




Gram 
Mol. 





^xlO-» 


I X 10-» 


10-* 


JxlO-^ 


Kultur- 
dauer 


Gew. 0^ 





0.000012 


0.00006 


0.0003 


0.0014 


I 
n 
in 


0.012 
0.009 

0.010 


0.024 
0.030 
0.024 


0.026 
0.025 
0.020 


0.020 
0.022 
0.018 


0.010 
0.007 
0.006 


32Tage 



Digitized by 




172 



N. ONO : WACHSTHUMSBESCHLEUNIGUNG 



E. I. Ealtnren mit Zusatz yon CaSO«. 
SHgeodonium — angestellt 14. Oct. 



Zimmer-Temperatar. 



.a 

'3 
« 


Gram 
Mol. 





^xlO-^ 


ixl0-» 


10-" 


ixlO-^ 


Kultur- 
dauer 


Gevr. o/t, 





0.00001 


0.00005 


0.00025 


0.0012 


I 

II 
III 


5 
5 
5 


4 
4 

4 


2 
2-3 

2 


1 
1 
1 


1 
1 
1 


27 Tage 



N.B. Die Ziifer zeigt den Entwickelung^grad. 







E. n. Enltoren mit Zosatz von QaSO^. 
CRroococcum— angestellt 13. Dec 


in Tieibhaag 16-2(r C 


.25 


Gram 
Mol. 





2^xl0-» 


\ X lO-'' 


10-" 


JxlO-^ 


Kultur- 


Qew. o/o 





0.00001 


0.00005 


0.00025 


0.0012 


dauer 


I 

II 
III 


5 

4-5 
5 


4 
4-5 

4 


3 
3 
3 


2 
2 

2 







71 Tage 

n 
n 



N.B. Die Zifler zeigt den Eutwickelongsgrad. 

F, I. Eulturen mit Zusatz von HgCl,. 
P7t)tococctM— angestellt 25. Dec. 



in Treibhaus 16-20** C. 



.2 

e) 


Gram 
Mol. 





^xlO-" 


%xlO-» 


10-" 


\ X 10-* 


Kultur- 


Gew. o/a 





0.00001 


0.00005 


0.00025 


0.00124 


dauer 


I 
II 
III 


5 
5 
5 


4 
3 
3 


1 














43 Tage 



N.B. Die Zitfer zei^t den Entwickelungagrad 
G. T. Kultaren mit Zusatz von NaFl. 





Emteertrag in g. 


iVotococcu*— angestellt 24. Dec. 


in Treibhan8l6-2(fa 


C5 


Oram 
Mol. 





ji^xlO-' 


liSxlO-' 


%xlO-' 


io-» 


Kultur- 


Gew. 0/, 





0.00003 


0.00016 


0.0008 


0.0042 


dauer 


I 
II 
III 


0.012 
0.012 
0.010 


0.018 
0.025 
0.027 


0.018 
0.018 
0.015 


0.018 
0.015 
0.015 


0.011 
0.015 
0.011 


76 Tage 



Digitized by 



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i 



EINIOEB ALGEN VKD PILZE. 



173 



Ernteertrng in g. 



H. 1. Kulturen mit Zusatz von liNO,. 

i^ococciM-angestellt 16. April. Zimmer-Tempemt>ir. 



.s 

1 


Gram 
Mol. 


^xlO-* 


*xlO- 


10-^ 


ixlO^ 


Kiiitiir- 
daiu?r. 


Gew. Oyi 





0.00003 


0.00014 


0.0007 


0.0034 


I 

II 
m 


0.010 
0.009 
0.010 


0.020 
0.020 
0.018 


0.017 
0.020 
0.016 


0.012 
0.015 
0.011 


0.009 
0.010 
0.008 


24 Tagy 

J* 


I. I. Kulturen mit Zusatz von K.AsO^. 
Ernteertrng in g. /Vo/ococcitf-angestellt 24. Dec. j^ Treibhaus 16-20^ C, 




Gram 
Mol. 





,ixl0- 


^xlO- 


i^lOr* 


10-^ 


Kiiltur- 
dauer. 


Gew. o/g 





0.00002 


0.0001 


0.0005 


0.0024 


I 
II 

m 


0.011 
0.008 
0.009 


0.015 
0.017 
0.015 


0.012 
0.020 
0.018 


0.011 
0.012 
0.014 


0.008 
0.007 
0.009 


54Tagt! 



Qeemtet 18. Jan. 



n. Versuche mit Pilzen. 



A. I. Eulturen mit Znsatz yon ZnSO«. 
AwpergUha nt^cr— angeetellt 24. Dec. '98. 

Kulturdauer 25 Tage. Temperatur 16-20*" C. 



Gehalt in 


Ernteertrag 
in g. 


SSure 


SSure pro Ig. 
PilzatiliBUiiiz. 


QTBmMoL 


G«v. % 


AddlUt 


als Ozaltfura 
amgeiechnet 



ixl0-» 
JxlO-» 
ixia-» 

io-» 




0.003 
0.007 
0.014 
0.028 


0.216 
0.863 
0.938 
0.944 
0.951 


15.7 
12.9 
13.2 
12.9 
13.0 


0.495 
0.406 
0.416 
0.406 
0.409 


2.245 
0.470 
0.443 
0.430 
0.4.30 



ir.B. A«|Minigin als N-QoeU^. 



Digitized by 



GooQldll 



174 



N. ONO: WACHSTHUMSBESCHLEUNIGUNG 




A. II. Kulturen mit Znsatz von ZnSO^. 

AspergiUw niger, — angestellt 24. Dec. '98. 

Geerntet 20. Jan. '99. Kulturdauer 27 Tage. Temperatur 16-20** C. 



Gehalt in 


Ernteertrag 
in g. 


Siiure 


Sanre pro Ig. 
PilzBnbetanz. 


Gnim MoL 


Gow. * 


Acidit£t 


aU Oxalsfiure 
umgerechDet 




ixlO-» 

ixlO^ 

ixlO^ 

10^ 



0.003 
0.007 
0.014 
0.028 


0.181 
0.868 

0.870 
0.858 
0.821 


14.8 
11.8 
11.1 

12.1 
14.1 


0.467 
0.372 
0.350 
0.381 
0.444 


2.580 

0.428 
0.420 
0.444 
0.541 



N.B. Asparagin als N-Quelle. 



A. in. Knlturen mit Zosatz yon ZnSO«. 

AapergUhu m^— angestellt 24. Dec. '98. 

Geerntet 20. Jan. '99. Kulturdauer 27 Tage. Temperatur 16-20*' C 



Gehalt in 


Ernteertrag 
in g. 


Same 


flaiiie pro Ig. 
PilzBubfitani!. 


Gram Mol. 


Gew. % 


tAcldlUt 


al8 Oxalsfiure 
umgerechDet 



|xlO-» 
JxlO^ 
ixlO^ 

io-» 



0.003 

0.007 
0.014 
0.028 


0.187 

1.017 
1.336 
1.939 
1.204 


17.8 

12.5 
11.8 
12.5 
11.2 


0.561 
0.394 
0.372 
0.394 
0.353 


3.000 
0.387 

0.278 
0.419 
0.293 



Geerntet 11. Jan. 



N.B. Asparagin als N-Qnelle. 



A. IV. Kulturen mit Zusats von ZnSO«. 
AspergiUvs nt^w^angestellt 20. Febr. 

Kulturdauer 20 Tage. Temperatur 16-20** C. 



Uehalt in 


Ernteertrag 
in g. 


SSare 


Saore pro Ig. 
Pilzsubetanz. 


Gram HoL 


Gew. % 


Acidltct 


als Ozals&are 
umgerechnet 




JxlO-» 
JxlO^ 
ixlO-» 

io-» 



0.003 

0.007 
0.014 
0.028 


0.634 
0.641 
0.635 
0.627 
0.585 


7.2 
7.2 
7.2 
7.2 
7.2 







N.B. Dextioee anttatt Bobrxacker. 



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EINIGEB ALGEN UND PILZE. 



175 



B. I. Kulturen mit Zosatz von FeSO«. 
PmieiUium ^toteum —angestellt 11. April. 

Kulturdauer 14 Tage. Temperatur 16-20' C. 



GduJim 


Emteertrag 
in g. 


Ritnre 


Saute iiro Ig. 
Pilisutmtatii. 


Gmn MoL 


0«w. S 


AcldlUt 


all Ozalaiure 



JxlO'' 
^xl0-» 

10- 
2xl0-» 




0.007 
0.014 
0.028 
0.056 


0.191 

0.180 
0.233 
0.201 
0.181 


3.5 
3.3 
3.9 
3.5 
3.4 







KB, HH^NO^ N-41iidl«. In alien eisenhaltigen Kulturen waren die Pilznuusen schdn ziegelruih 
eemrbL Schon bei 0.001% deutUche rothe FSrbang bemerklioh. 



Qme^% Aliir/. 



C, I. Kultaren mit Zusatz von NiSO^. 

AsperffiUtu niger — angestellt 9. Febr. 

Kulturdauer 22 Tage. 



Temperatur. 16-20' 11 



OebsH in 


Ernteertrag 
in g. 


Siiure 


Saure [>ru l^,, 
PilzBiibBtaoii. 


GniB UoL 


<%». % 


AoidiUt 


als Oxalsfiare 




ixlO-» 

JxlO-' 

4 X io-» 
io-» 




0.003 
0.007 
0.014 
0.028 


0.250 

0.297 
0.315 
0.401 
0.295 


11.0 
11.1 

10.7 
14.6 
14.0 


0.346 
0.350 
0.337 
0.460 
0.441 


1.464 
1.179 

1.0G9 
1.147 
1.493 



Geemtet 4. MiinL 



N.B. NH«NO, alfl N-Quelle. 



a II. Kulturen mit Zusatz yon NiSO«. 
Aspergillus ni^r— angestellt 9. Febr. 

Kulturdauer 23 Tage. Temperatur 16-20"" C. 



Geliali in 


Ernteertrag 
in g. 


Saore 


Saore 11 ro Ig. 
Pilau t«tariz. 


Gram MoL 


Ge*. « 


Aoidttit 


als Ozalsfture 
umgerechnet 



I X 10-" 
i X 10-' 
^xlO-* 

10-' 




0.0035 
0.007 • 
0.014 
0.028 


0.288 

0.316 
0.307 
0.387 
0.329 


11.1 

10.7 
11.1 
16.9 
16.7 


0.350 
0.337 
0.349 
0.532 
0.526 


1.216 
1.0G7 
1.130 
1.375 

1.500 



N.a NH«NOs als N-Qaelle. 



"-"-r 




176 



N. ONO : WACHSTHUMSBESCHLEUNIGUNG 



Geerntet 6. Marz. 



C. III. Eulturen mit Zusatz v(hi NiSO«. 
Aspergillus m^vr— angestellt 9. Febr. 

Kulturdauer 35 Tage. Temperatur 16-20° C. 



Gehalt in 


Ernteertrag 
in g. 


Saure 


Saure pro Ig. 
PilzgDbstunx. 


Gram Mol. 


Gew. % 


Additst 


als OialsSure 



I X lO-^" 
J X 10-» 
i X lO-'' 

io-» 




0.003 
0.007 
0.014 
0.028 


0.324 
0.310 
0.329 
0.362 
0.341 


9.8 
10.0 
11.9 

15.2 
17.3 


0.308 
0.315 
0.375 
0.479 
0.544 


0.951 
0.016 

1.140 
1.323 
1.695 



N.B. NH«NOa als N^uelle. 



U 



IP 



1 



Geerntet 4. Mai. 



0. IV. Kulturen mit Zusatz von NiSO^. 
Aspergillus niger — angestellt 22. April. 

Kulturdauer 12 Tage. Temperatur 16-20** C. 



Gehalt in 


Ernteertrag 
in g. 


8iinre 


Saure pro ]g. 
I^lzsubstanz. 


Gram Mol. 


Gew. % 


Acidim 


als OxaliAure 
umgerechnet 




|xlO^ 

ixlO-» 

ixlO-^ 

10-* 




0.003 
0.007 
0.014 
0.028 


0.262 
0.390 
0.404 
0.364 
0.315 











N.B. NH«NOs als N-Quelle. 



Geerntet 4. Mai. 



C. V. Kulturen mit Zusatz von NiSO^. 
Aspergillus ni^rer— angestellt 22. April. 

Kulturdauer 12 Tage. Temperatur 16-20** C. 



Gehalt in 


Ernteertrag 
in g. 


Saure 


Saure pro Ig. 
PilzBubstaDZ. 


Gnm Mol. 


Gew. % 


Acidit&t 


als Ozalsfiare 
umgerechnet 



ixlO-» 
JxlO-^ 

ixlOr^ 

io-» 



0.003 

0.007 
0.014 
0.028 


0.214 
0.311 
0.300 
0.307 
0.296 









N.B. NH«KOs als N-Quello. 



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EIKIOEB ALGEN UND PILZE. 



177 



Qeerotet 4. Mai. 



C. VI. Kulturen mit Zusatz von NiSO^. 
Aspergillus nigtr — angestellt 22. April. 

Kulturdauer 12 Tage. Temperatur 16-20° C. 



Gehalt in 


Ernteertrag 
in g. 


Store 


Saure pro Ig. 
PilzBubetanz. 


Gram Hoi. 


Oev. % 


Acidillt 


ala OxalsSura 
umgerechnet 




ixlO-^ 
JxlO-^ 
ixlO-» 

io-» 




0.003 
0.007 
0.014 
0.028 


0.278 

0.340 
0.325 
0.308 
0.324 









Geerntet 16. Marz. 



N.B. NH«NOa als N.QaeU& 



D. I. Kulturen mit Zusatz von CoSO^. 
AspergUlua m*p«r— angestellt 17. Febr. 

Kulturdauer 27 Tage. Temperatur 16-20° C. 



Gehalt in 


Ernteertrag 
in g" 


Saure 


SScare pro Ig. 
Pilzsubstanz. 


Qiua UoL 


G«w. X 


AcidiUt 


als Ozals&ure 
umgerechnet 




AxlO- 

JxlOr^ 

JxlO^ 




0.0017 
0.0035 
0.007 
0.014 


0.297 
0.439 
0.565 
0.751 

0.872 


9.0 
10.3 
12.3 
11.2 

8.7 


0.283 
0.324 
0.387 
0.353 
0.274 


0.953 
0.738 
0.685 
0.470 
0.314 



Geerntet 20. Marz. 



N.a NH«NOs ab N-QueUe. 



D. II. Kulturen mit Zusatz von GoSO^. 
Aspergillus niger — angestellt 17. Febr. 

Kulturdauer 31 Tage. Temperatur 16-20° C. 



Gehalt in 


Ernteertrag 
in g. 


Saure 


f%nre pro Ig. 


Qnm MoL 


Gew. % 


Aciditst 


als Oxalsfiure 
amgerechnet 




Axio^ 

|xl0-» 
ixlO-* 
ixlO^ 



0.0017 

0.0035 

0.007 

0.014 


0.280 
0.423 
0.582 
0.745 
0.815 


10.0 
12.1 
15.5 
16.7 
8.4 


0.315 
0.381 
0.491 
0.548 
0.265 


1.125 
0.900 
0.844 
0.735 
0.313 



N.B. NH«NO« alfl N-Quelle. 



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Google j 



178 



y. ONO : WACHSTHUMSBESCHLEUHIGUNG 



I 



Geemk't 17. Miir/.. 



D, HX Ealtaren mit Zusatz von CoSO^. 
A^ergiUua niger — angestellt 17. Febr. 

Kulturdauer 28 Tage. Tempenvlur 16-20° C 



<itliall in 


Ernteertrag 
in g. 


Sanre 


Sua re pni 


Gram Mol. 


0«. Jt 


AcidlUt 


ala OiaUlUTa 


Pilzsul>sla 








0.267 


10.6 


0.334 


1.251 


AxlO- 


0.0:J17 


0.393 


13.0 


0.409 


1.041 


ixlO-" 


0.0035 


0.561 


15.5 


0.488 


O.870 


JxlO-» 


0.007 


0.742 


14.8 


0.4G6 


0.62S 


i X 10-^ 


0.014 


0.770 


10.0 


0.315 


0.409 



f Jwrntt't *J0. Miici, 



N.B. NH«NOs als N-QueUe. 



Lh IV. Kulturen mit Zusatz von CoSO,. 
PiTiltiUivm glaueum — angestellt 20. Marz. 

Kulturdauer 10 Tage. Tempefatur 10-20^ C 



(iclinll in 


Ernteertrag 
in g. 


Siiure 


Siure pro 


Grwu H(d. 


G<'W. % 


AciditSt 


al« (Jxalsiiun! 


PilzsufasU 



AxlO- 
JxlO-* 
JxlO^ 
JxlO-' 




0.0017 
0.0035 
0.007 
0.014 


0.108 
0.186 
0.225 
0.317 
0.366 


5.4 

5.0 
5.4 
5.9 
5.9 


0.170 
0.157 

0.170 
0.186 
0.186 





Gevrntet 1, April. 



N.a NH«NO, als N-QueUe. 



D< V. Kulturen mit Zusatz von CoSO«. 
Peindllium glaueum — angestellt 20. MTinu 

Kulturdauer 11 Tage. Temt^emLur 16-20° C 



Gehalt in 


Ernteertrag 
in g. 


Saare 


Siure pro 


Gckm HoL 


a«w. f 


Addltst 


aU Uxalflaure 
uiii|[?€reclniet 


ruzBuiniU] 




Axio-' 

ixlO-» 
JxlO--* 
ixlO-' 




0.0017 
, 0.0035 
0.007 
0.014 


0.242 

0.469 
0.354 
0.482 
0.772 


4.6 
5.7 

5.9 
5.7 
5.9 


0.145 
0.179 
0.186 
0.178 
0.186 





N.B. NH«NOs ala N-Quelle. 



Digitized by 



Google 



I 



EINIGEB ALGEN UND PILZE. 



179 



Qeerntet 13. April. 



D. VL Eulturen mit Zosats von CoSO«. 
Penieilliwn glaueum — augestellt 20. Marz. 

Kulturdauer 23 Tage. Temperatur 16-20** C. 



Gehalt in 


Ernteertrag 
in g. 


eSaie 


Sanre pro Ig. 
PilzBubstanz. 


GitmMoL 


Gew. % 


AcldiUlt 


als Oxalflfiuie 
umgerechnet 




^xlO- 
ixlO-» 
JxlO-» 
ixlO-' 



0.0017 

0.0035 

0.007 

0.014 


0.363 

0.349 
0.520 
0.649 
0.289 


6.4 

6.2 
5.9 
5.7 
6.6 


0.202 
0.195 
0.186 
0.179 
0.207 






N.B. NH«NOs als N.Qaelle. 



Greemtet 31. Marz. 



E. I. Ealturen mit Zusatz von GuSO«. 
AapergUlvs niger — angestellt 21. Marz. 

Kulturdauer 10 Tage. Temperatur 16-20*» C. 



Gehalt in 


Ernteertrag 
in g. 


Sanre 


Sanre pro Ig. 
PiljsubBtanz. 


Gram Mol. 


Qew. % 


Acldltjtt 


als Ozalsftaie 
umgerechnet 




Axio- 

JxlO^ 
JxlO-* 
ixlO-» 




0.0015 
0.003 
0.006 
0.012 


0.307 

0.305 
0.297 
0.311 
0.360 


6.2 
5.2 

5.2 
4.7 
5.1 


0.195 
0.164 
0.164 
0.138 
0.160 


0.635 

0.538 
0.552 
0.444 
0.444 



Geerntet 5. April. 



N.B. NH«NOs als N-Qnelle. 



R n. Enltnren mit Zusatz von GaSO«, 
AaperffUhu ni^— angestellt 21. Marz. 

Kulturdauer 16 Tage. Temperatur IB-SO** C. 



Gehalt in 


Ernteertrag 
in g. 


Sanre 


Siinre pro Ig. 
Pilzsnbstanz. 


GfamHol. 


Gew. % 


Acldltst 


als Oxalsfiure 
umgerechnet 




AxlO- 
JxlO-^ 
JxlO-» 
ixlO-» 




0.0015 
0.003 
0.006 
0.012 


0.273 
0.307 
0.313 
0.324 
0.345 


9.7 
10.0 
10.3 
10.1 

9.4 


0.309 
0.315 
0.324 
0.318 
0.296 


1.121 
1.003 
1.035 
0.967 
0.858 



N.B. NH«NOs als N-Quelle. 



I|i 



Nil 



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180 



N. ONO: WACHSTHUMSBESCHLEUNIGUNG 



Geemtet 6. April. 



E. m. Kultuien mit Zusatz von CoSO^. 
AspergiUu8 m^r— angestellt 21. Marz. 

Kulturdauer 16 Tage. Temperatur 16-20® G 



Gehalt in 


Ernteertrag 
in g. 


Same 


SSure pro Ig. 
RlzRubstenz. 


(inin MoL 


Gew. f 


Acldiut 


als Ozals&ura 
umgerechnet 



^xlO-- 
JxlO^ 
Jxl0-» 
Jxl0-» 




0.0015 
0.003 
0.006 
0.012 


0.218 
0.252 
0.352 
0.358 
0.343 


9.9 
10.1 
9.9 
9.3 
9.6 


0.312 
0.318 
0.312 
0.293 
0.302 


1.431 
1.265 
0.886 
0.818 
0.880 



N.B. NH^NOj N-Quelle. 




Qeeratet 9. Febr. 



F. I. Eulturen mit Zusatz von HgCl,. 
Aspergillus niger — angestellt 1. Febr. '99. 

Kulturdauer 8 Tage. Temperatur 16-20® C. 



Gehalt in 


Ernteertrag 
in g. 


Silaie 


Soare pro Ig. 
PilzsabsUnz. 


Gram HoL 


Gew. % 


AciditJCt 


als OzalsSure 
umgerechnet 




Axio-' 

JxlO-' 
JxlO^ 
JxlO^ 




0.0017 
0.0034 
0.0067 
0.0135 


0.126 
0.180 


12.6 
13.3 


0.397 
0.419 


3.176 
2.328 





N.B, NII^NO, als N-Quelle. QMVJ% gut entwickelt. Sporon braun. 0.0034 ;( fast keine Entwickelnng. 
0.0067 )( u. 0.0185 )( keine Entwickelnng. 



Gt'enitct 9. Febr. 



F. II. Knlturen mit Zusatz von HgCla. 

Aspergillus m^Per— angestellt 1. Febr. '99. 

Kulturdauer 8 Tage. 



Temperatur 16-20* C. 



Gohalt in 


Ernteertrag; 
in g. 


Saare 


Saare pro Ig. 
PilzsabsUnz. 


QnmMoL 


G«w. % 


AcidiUt 


als OxalaSure 




tfexlO- 
JxlO^ 

ixio-» 

Sxio-» 




0.0017 
0.0034 
0.0067 
0.0135 


0.119 
0.188 


10.0 
12.7 


0.315 
0.400 


2.644 

2.128 



N.B. NE«NOa als N-Quelle. Entwickelnng wie vorige. 



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EINIGEB ALGEN UND PILZE. 



181 



G^erntet 9, Febr. 



F. in. Kultaren mit Zosatz von Hgd.. 
AapergiUus m<?er— angestellt 1. Febr. *99. 

Kultordauer 8 Tage. Temperatur IQ^^r C. 



OelialL in 


Emteertrag 
in g. 


Saure 


Siure pro Ig. 
FUzsubeians. 


Brain MoL 


G«w. X 


ActdiUt 


ala Oxal^ure 
umgerechnet 


' 
^xlO- 
JxlO-^ 
^xlO-* 
JxlO^ 




0.0017 
0.0034 
0.0067 
0.0135 


0.153 
0.160 


11.3 
13.7 


0.356 
0.431 


2.366 

2.568 



K.B. NH4NO, als N-Quelle. Entwlckelang wie vorige. 



F. IV. Kulturen mit Zusatz von HgCl,. 
FeniciUium glaucum—AngeeteWt 20. April. 



Gcemtet 1. 


Mai. 


Kulturdauer ] 


11 Tage. 


Temperaf 


111- ]0-2IJ° C. 


Gehalt In 


Emteertrag 
in g. 


Kiure 


Siuro pro Ig. 
Pilzmifastaoz. 


Gram Hoi. 


Oew. % 


AcidlUt 


als OialsSure 
umgerechnet 








0.203 


4.5 


0.142 




Axio-* 


0.0017 


0.243 


4.7 


0.148 




IxlO-' 


0.0034 


0.242 


5.1 


0.159 




JxlO-^ 


0,0067 


0.473 


5.1 


0.159 




^ X lO-" 


0.0135 


0.251 


4.9 


0.154 





N.B. NH4NO8 ala N-QueUe. 



CK'emtet 2. MaL 



F. V. Kulturen mit Zusatz von Hga,. 
PenieiUium p^ctucuni— angestellt 20. April. 

Kulturdauer 12 Tage. Temperatur 16-20^ C. 



Ck'hait in 


Emteertrag 
in g. 


Saure 


Slure pro Ig. 

ril2SUtj8t«P2, 


Gimm Mai. 


G«w. % 


Aoidltftt 


ala Oxals&ure 
umgerechnet 




JxlO-- 

ixlC^ 

ixlO-^ 

10-^ 




0.0003 

! 0.0006 

0.0013 

0.0027 


0.222 

0.282 
0.264 
0.273 
0.301 


46 
4.6 
4.7 
4.6 
4.5 


0.145 
0.145 
0.148 
0.145 
0.142 





N.a NH«NOs als N-Qaelle. 



Digitized by 



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t82 



N. ONO: WACHSTHUMSBESCHLEUNIGUNG 



Geemlet 1. Mai. 



F. VL Kulturen mit Zusatz von HgCl,. 
PeniciUium glaucum — angestellt 20. April. 

Kulturdauer 11 Tage. Tennitrjiltir Ifi-SO** C. 



1 Gefinlt in 


Emteertrag 
in g. 


Saure | 


&iiire pro Ig 


Grflra Mol. 


Gow. % 


AddiUt 


als Oxalsaure 
umgcrectmet 


Pilzsubstani 


n 


0.183 
0.249 
0.213 
0.311 
0.246 


5.0 

5.5 
5.9 
5.5 

5.8 


0.157 
0.173 
0.185 
0.173 
0.1S2 




ixlO- 
1 X 10-' 
J X lO- 

lo-- 


0.0003 
0.0006 
0.0013 
0.0027 







N.B. NH^NOs als N.Quelle. 



F. VII. Kulturen mit Zusatz von HgCl,. 

Aspargillus nigei' — angestellt 30. Marz. 

Geerntet 18. April. Kulturdauer 18 Tage. Ternperatur 16-2{f C. 



Gebalt in 


Ernteertrag 
ing. 


S&ure 


Eittue pro 1| 


Grftui Uol 


Oew. % 


AcidiUt 


als OJcal^UFQ 
umgerechnct 


Pilzsubstani 








0.347 


8.8 


0.277 


0.800 


JxlO-* 


0.0003 


0.517 


9.6 


0.302 


0.586 


4x10-' 


0.0006 


0.513 


9.4 


0.296 


0.555 


ixlO- 


0.0013 


0.552 


10.9 


0.343 


0.621 


io-« 


0.0027 


0.565 


11.3 


0.356 


0.630 



N.B. NH«NO, als N-Quelle. 



F. Vm. Kulturen mit Zusatz von HgCl,. 

Aspergillus ntyrr— angestellt 30. Marz. 

Geerntet 18. April. Kulturdauer 19 Tage. Ternperatur 16-20* C. 



Uebalt in 


Emteertrag 
in g. 


Siare 


Sill re pro 1 


Gjiliu Mol. 


Gew. % 


AcidiUt 


als OxAlsiiire 
umgcrcchact 


PilzEubstan 




JxlO-^ 
JxlO-* 

J X 10-* 
10-^ 




0.0003 
0.0006 
0.0013 
0.0027 


0.341 
0.458 
0.474 
0.630 
0.429 


8.6 

9.4 

9.4 

12.5 

10.3 


0.271 
0.296 
0.296 
0.394 
0.324 


0.793 
0.646 
0.624 
0.625 
0.755 



N.B. NH^NOs als N^uello. 



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EINIQER ALQEN UND PILZE. 



183 



Gwratet 18. April 



F. IX. Kulturen mit Zusatz von HgCl, 
AtperffSluB niger — angestellt 30 Marz. 

Kulturdauer 19 Tage. Teinperatur ie-20® C, 



Gehxlt id 


Emteertrag 


SiUiie 


Sauie pro Ig. 
Pilzsubstaiu. 


Omm Mol. Gtw. f 


in g. 


AciaiWt umgerechnet 


1 

jxio-« : 0.0003 

|xl0- 0.0006 

JxlO-" 0.0013 

10^ 0.0027 


0.261 
0.355 
0.380 
0.509 
0.451 


8.4 ' 0.265 

9.2 0.290 

9.4 0.296 

12.1 0.381 

11.1 0.350 


1.015 
0.816 
0.779 
0.742 
0.776 



N.B. NH4NO, als N-Qoelle. 



Geeratet 18, April- 



G. I. Kulturen mit Zusatz von LiNO,. 
Asperffillus nigei — angestellt 1. April. 

Kulturdauer 17 Tage. Temperatur 16-20** C. 



Gehalt in 


Emteertrag 


Siiure 


SSare pro Jl% 
PilzBubstanii. 


Giuii MoL lier. % 


in g. 


AoidltCt 


als Oxalsfiure 
umgerechnet 




5x10-* 

JxlO-* 
I X 10-'' 



0.004 
0.008 
0.017 
0.034 


0.300 
0.408 

0.428 
0.348 
0.345 


9.0 
9.4 
8.9 

8.7 
8.6 


0.284 
0.296 
0.283 
0.273 
0.271 


0.946 
0.725 
0.661 
0.784 
0.782 



N.B. NH« NO, als N-QueUe. 



Geemtet 21. Febr. 



H. I. Kulturen mit Zusatz yon NaFl. 
AspergUbu ni^— angestellt 7. Febr. '99. 

Kulturdauer 14 Tage. Temperatur 16-20® C. 



Gelntt in 


Emteertrag 
in g- 


Saure 


Saure pro 1^. 
PilMubetany.. 


QnmlloL 


Oew. * 


AcidlUt 


als OxalsSore 
umgerechnet 




J X 10-* 
JxlO-* 
ixlO-« 




0,0025 
0.005 
0.010 
0.021 


0.199 
0.325 
0.312 
0.246 
0.289 


8.8 

9.4 
7.2 
6.1 
6.0 


0.277 
0.296 
0.227 
0.192 
0.189 


1.392 
0.911 
0.727 
0.880 
0.654 



N.B. NH^NOs als N-Qnelle, 



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Googl^ 



184 



N, ONO: WACHSTHUMSBESCHLEUNIGnNG 



Geornlet £3. Febr. 



H. n. Koltaien mit Zosatz von NaFl. 
ABpergSliu m^ey^-angesteUt 7. Febr. '99. 

Kulturdaoer 16 Tage. Tcmperalar 16-2<f a 



Gelialt in 


Ernteertrag 
in g. 


Same 


SUire pro ] 


Gnrai MoT. 1 tiaw. « 


AcIdiUt 




Piligiibstan 



Ax 10-* 

fxlO-' 

4xio-» 

i X 10~* 



0.0025 
0.005 
0.010 
0.021 


0.314 
0.336 
0.385 
0.316 
0.274 


10.0 

10.0 

7.6 

5.9 

5.6 


0.315 
0.315 
0.239 
0.18C 
0.17G 


LOGO 
0.937 
0.621 
0.589 
0.642 



N.B. NH«NO, al8 N-Quelle. 



■i 




Ge'^rntet 25. Febr. 



H. IIL Eulturen mit Zasatz von NaFl* 
Aspergillus niger — angeetellt 7. Febr. '99, 

Kulturdauer 18 Tage. Tcmpcratur lfi'2a** a 



QehAlt in 


Ernteertrag 
in g. 


SSnre 


STiure pro ] 


Qma Uol. 


G«w. % 


AcidlUt 


ala Oxalsaivre 
udgensehDct 


Pilffiubatan 




AX 10-* 1 

JxlO-* 
ixlO-' 
JxlO* 



0.0025 
0.005 
0.010 
0.021 


0.270 
0.280 
0.285 
0.264 
0.265 


8.4 
10.7 

7.7 
6.6 
6.1 


0.265 
0.339 

0.242 
0.208 
0.192 


0.982 
1.207 
0.849 
0.788 
0.725 



N.6. NH«N08 al8 N-Quelle. 



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EINIGEB ALGEN UOT) PILZE. 185 



INHALT. 



I. Elnlcitung and Litteratur. 

IL Methodisch€& 

ni, Vorbfzuerkangeii uber die Veisacbflobjekte. 

ly. Temndernngen in der Wachsthumsweifie und die Correla- 
tion swischen Fortpflanzang und Wachsthmn. 

V. Einlluse der Beizstoflfe auf die Betriebsstofiwechsel. 

VI. Bpecielle Besprechungen. 

YII. Schlossbemerkungen und Zusammenfaasung der Besnltate. 

TABELLAMSCHE ZUSAMMENSTELLUNG. 

I, Venrache mit Algen. 
II* Versuche mit Pilzen. 



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1 



186 



Erklarung der Tafel XIII. 



Kulturen von ABpvrgWm mgtr mil und ohne Zusatx von NaFl. 

(Photographiert 16 Tage nach der S^renaasBaat) 
I. Ohne Zusats ; Kontrollknltur. 
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^ „ 

(Fiir naherea vgL S. 153 and ferner Tabelle Pilz. H.) 



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o 

to 
C 
o 




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1 

'I 



i 

t 



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CONTENTS OP RECENT PARTS. 



Vol. IX., Pt. 1 . . . yen 0.62 (Price in Tokyo). 
On a Certnlii Cliuis of Frmnnbofer** Dlflnractloii*l*beBOineiiii. By H. Naoaoxa. 
Umtm of Equal Intenalty aboni tbe Point of Inieraeetion of Frannliofer** Dif* 

feneUon Banda. By H. Nagaoka. 
BTole on Tinfoil Gratlni^ aa a I>«te«tor for Electric Waves. By T. MizuKO. 
Tke Tbemio-electrlc EflPBCIa of I^onfritndlnal Sireas In Iron. By K. TsuBUTA. 
Tliemio-olectric EflTpcta of I/onirKndlnal Tenaion In Blcrerent Melala. By E. 

TsUBUTA. 
Holes* on tlie Topas from Hlno. Bt T. HiKl. 
Hercnrjr Percblorates. By M. Chikashiqb. 
Potaaalam nltrososnlpliate. By E. DiVERS and T. Haoa. 
Bodlnm nltroaoanlptaate. By £. DiVEBS and T. Haga. 
The Conailtntlon of tlie Hlirososalphatea. By £. D1VBB8 and T. Haga. 

Vol. IX., Pt. 2 . . . yen 1.31 (Prioe in Tokyo). 
The Tinfoil Orailnc Betector for Electric Waves. By T. MiZUNO. 
On the PledmonUte-rhyorlte from Shlnano. By N. Yamasaki. {With Plate VI). 
The Atomic Weiirht of Japanese Tellnrlam. By M. Chikashige. 
Das Johannlsfc&fer-IJcht. Voii H. MufiAOKA. 
On the Prediction of Solar Eclipses. By Shin HirataMA. 
How Mercorons and Mercnric Salts chanipe Into each other. By S. Hada. 
Imldoaalphonates (Second paper). By £. DrvSBs and T. Haga. 
Amldosalphonlc acid. By £. BiVERS and T. Haga. 
Molecnlar Condnctlvlty of Amldosalphonlc Acid. By J. SakuRAI. 
The Physiological Action of Amldosalphonlc Acid. By OSCAB LoEW. 
The Kedactlon of Nltrososnlphates. By £. Divers and T. Haga. 
Economic Preparation of Hydroxylamlne Salphate. By £. D1YBB8 and T. Haga. 
On the Time-lAfp In (he Hai^nedsatlon of Iron. By Y. EATa (With Plates' 

VTI-XV). 

Vol. IX., Pt. 3 . . . yen 0.65 (Price in Tokyo). 
DIAmctlon Phenomena in the Focal Plane of a Telescope with Circular Aper* 

tare dae to a Finite Soarce of I^iflrht. By H. NagAOKA. (With Platee XVI & 

XVII). 
BesMhrchea on Haimetostrlctlon. By H. Nagaoka and K. HoKDA. (With PUOn XVIII 

& XIX). ' 

Vol. X., Pt. 1 . . . yen 1.80 (Price in Tokyo). 
On the Fate of the Blastopore, the Belatlons of the Primitive Streak, and the 
Formation of the Posterior End of the Embryo In Ghelonla, together with 
Remarks on the Mature of Meroblastlc Ova In Tertebrates. (Contributions 
to the Embryology of Reptilia. V.). By E. Mitsukubl (With Plates I-XI). 
Vol. X., Pt. 2 . . . yen 1.20 (Price in Tokyo). 
Veher elae In Hlsakl vorkommende Art von Ephelota and Ober Ihre Sporen* 
bildnnff. Von C Ishikawa. (Hxemi, Tafdn XII und XIII). 



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'Veber <lnii massenliafle Torkommen von Elii^nbaeterleii In den Ttaermen Ton 

Ikno. Von M. MiTOSHi. 
Stadlen fiber dl« Schwefelvasenblldnnir and die Seliwefelbiieterleii der Tbermeii 

von Ynmoto bet Nlkk6. Von M. MiYOSHi. {Hierzu Tafd XIV), 
Die EntwiekeliiniT der Gonophoren bel Physnlla maxima. Von S. GrOTo. {^em 

Taf. XV). 
Stndies of ReprodneCiTe Elements. III. Die Entwickeliini: der Polienk4(roei 

von Alllam fistul<Minm !•., ein Beilraip xnr CbromoeomenrednkUon la 

Pflansenrelehe. Von C. IsHiKAWA. Hierza Tafein XVI und XVII). 
4/on(rlbati»ns to the Horpholoipjr of Cjrclostomatn. 1. On the Formation of thi 

Heart In l>etromyxon. By S. Hatta. ( With Plate XVIII), 
Vol. X., Pt. 3 . . . t/en 1.10 (Price in Tokyo). 
The 9Ietam«»rphosls of Asterlas palllcia, with Special Beference to the Fate ol 

(he B<»djr-eavUleii. By S. Goto. {With Plates XIX-XXIV). 



Vol. XI., Pt. 1 . . . yen 0.34 (Price in Tokyo). 
Preparation of Hyponltrite from Nitrite throoirl* Oxantldosnlpbonate. By E 

Divers and T. Haga. 
Absorption of Nitric Oxide In «as Analysis. By £. DlYEBS. 
Interaction of Nitric Oxide with Silver Nitrate. By E. DiVERS. 
Preparation of Pure Alkali Nitrites. By E. DlVEBS. 

The RedncUon of an Alkali Nitrite by an Alkali Metal. By E. Diyebs. 
Hyponltrltes : their Properties and their Preparation by Sodium or Potasslnu 

By. E. Divers. 

Vol. XI., Pt. 2 . . . yen 0.38 (Price in Tokyo). 

On the Geuloffic Structure of the Malayan Archipelago. By B. Eotd. [With PlaU 1 

Vol. XI., Pt. 3 . . . yen 1.45 (Price in Tokyo). 
Iforlsontal Pendulums for the Mechanical Regrlst ration of Seismic and Othe 

Earth Movements. By F. Omori. {WUh PtaXea II-XTI). 
Note on the Preliminary Tremor of Earthquake Motion. By F. Omobi. {Wh 

Hates Xm-XViy 
Earthquake Measurement at MIyako. By F. OiiORi and K. HiRATA. (WUh Plat 

xvnxxniy 

Ethyl ammonlnmsulphlte. By E. DivERS and Ogawa. 

Ethyl ammonium selenlte and Non-existence of Amidoselenites (Selenosamates 

By E. Divers and T. Hada. 
Notes on the Minerals of Japan. By K. JiNBO. 

Vol. XI., Pt. 4 . . . yen 1,04 (Price in Tokyo). 
On the Mutual Influence between lionipltudlnal and Circular MaffnetlsatioBs li 

Iron and Nickel. By K. HoNDA. {WUh Plates XX J V & XXV). 
The Earthquake Investlfpatlon Committee Catalogue of Japanese Earthqoakei 

By S. Sekiya. 
Notes on the Earthquake Investlflpatlon Committee Catalogue of Japanese Earth 

quakes. By F. Omori. {WUh PhUs XXVI-XXVH). 



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Vol. XII., Pt. 1 . . . yeti 1.20 (Price in Tokjo). 
JnpAiiivclte li««cliii]ie Fulmonaien. AnM. Vni^rsaeh. d. Id Zool. Mnieiini d«r^ 
k. ITnlv. In TOkyi^ eotlialtenen Haterlales, Von A. Jacobi. {Ulenu Taftin 

Vol. XII,, Pt. 2 • . . yen 0.60 (Price in Tokyo). 
£tiiflcA aiir la F^^ondation et l^Embryoff^nle da Ginkgo blloba. Second m^moire. 
Par Sakl'GOKo Hoiase. {Avee Ft. VII-IX). 

Tol, xn., Pt. 3 . . . yen 1.06 (Price in T5ky6). 
ITnt«r«ii«liiiii9«ii ilber ille EntwlekluniT der Geflchlecbtsorsaiie and ileii Torirnnif 

der BerrneUtiiniT bel Cyeas revolata. By S. IeeNO. (With Flaiea jV-XK//)» 
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my 

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CONTENTS. 



Vol. xin., Pt. I. 

PA 

Notes on the Oeolog^ of the Dependent Isles of Taiwan. 

By B. Koto, PA. Z?., Bigakuhakushi ; 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. Naqaoka, Bigaku- 
hakushi, Professor of Applied Mathematics; and K. Honda, 
Rigakushi, Post-graduate in Physics. {With Plates VI&VU). 

Combined Effect of Longitudinal and Circular Magnetiza- 
tions on the Dimensions of Iron, Steel and Nickel 
Tubes. By K. Honda, Rigakmhi ; Post-graduate in Phys- 
ics. {With Plates VIII & IX) ' 

Studien uber die Anpassungsfahigkeit einiger Infusorien an 
concentrirte Losungen. Atsushi Yasuda, Rigakushi ; 
Professor der Naturgeschichte an der zweiten Hochschule zu 
Sendai. {Eierzu Tafel X-XII) 1< 

Ueber die Wachsthumsbeschleunigung einiger Algen und 
Filze durch chemische Reize. Von N. Ono, Rigakushi. 
{Hierzu Tafel XIII) 1^ 



PRINTED AT THE " TOKYO TSUKIJI TYPE FOUNDRY." 



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JOURNAL 



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OF THE * J 



COLLEGE OE SCIENCE, 

IMPERIAL UNIVERSITY OF TOKYO, 

VOL. XIII., PART II. 



Ig, -^ 1^ m :k ^ ^V /ii 

PUBLISHED BY THE UNIVERSITY. 

TOKYO, JAPAN. 

1900. 

MEIJI XXXIII. 



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Publishiiig CkMBunittee. 



Prof. K. Yamagawa, Ph. B., RigakuhakusM, Director of tbe Ooll6( 

Pro£ J. Sakuraif RigakuhakusM. 

Pio£ B. Kotft, Ph. D., RigdkuhdkuOL 

Pro£ I. Ijtfna, Ph. D., RigakuhakusM. 



All cominwnlcattons relating to this Journal shoold be addressed to the 
Director of the College of Science. 



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Anunonitun Amidosulphite. 



By 



Edward Divers ^tuI Masataka Ogawa, 

Imperial UiuTemty, Tf)ky5, 



he ifiterLiction of Biich familiar gases as ammonia and 
ir dioxide ceased to attract with any effect the attention of 
io-ators sixty years ago and more. Yet comparatively 
ig had then been definitely made out about the nature of 
rodiiet, and even the few statements concerning it in some 
i best treatises on chemistry have but little experimental 
Eition, The history of the subject is briefly given on p. 193. 

Nmi'Umon of dry sulphur dioxide and mnmonia* 

liven when comparatively well-dri&d, sulphur dioxide and 
mia unite at once and with great energy when brought 
ler; yet they can remain mixed without combining, pro- 
Bufficient care has been observed to exclude moisture. It 
lOt been necessary, however, in order to demonstrate thii 
og phenomenon, to have resort to the elaborate precautions 



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188 



E, DTVEE8 AFD M, OGAWA : 



adopted by BreretOE Baker, in his famous experiments upon the 
noo-miion of hydrochloric acid and ammonia (/. CL Soc.f 1894, 
fi5i Bll ; 1898, 73, 422), and we have only dried the gaaes 
during their flow through the tubes. As we were able to dry 
sulphur dioxide better than ammonia, because common phoe- 
phorua pentoxide could be used for the purpose, we have had 
success in mixing the gases without their combining only by giving 
this giis 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 ihc in- 
terior of the flask remained clear for some minutes, the mixe^l 
gases only combining on their escape through the mercur 
the air. But the ammonia having, it is presumed, jrn 
brought enough moisture with it through passing more ] 
along the tubes than at first, the walls of the flask becani- 
denly coated with an orange-coloured deposit, while the m 
rose high in the exit tube. «. 

Proportions in wkiak 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 cousiderablLS They vary also according as one 



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AMMONIUM AMIDOSULPHITE. 



189 



ler 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- 
ited product, that may be observed, are results clearly due 
3 secondary changes going on (p. 192). The simple union 
amonia and sulphur dioxide, which can be secured by 
Qg 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 
L of the two gases can appear to take place in other pro- 
ms than the above. It is pretty certain that, by proceeding 
jr enough and using strong cooling agencies, secondary ac- 
30uld be almost entirely prevented and the statement just 

be verijfied, 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 
presence 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 



II 



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190 E. DIVEES 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 an 
water by sodium, was contained in a small flask, fitted wil 
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 poi 
dery. In spite of the external cooling, the heat of combinij 
was sufficient to cause ammonia gas, saturated with ether-vapoi 
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 w( 
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. Tl 
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, ai 
continued for hours, until all the ether adhering to the precif 
tate had been carried away, the flask being all the while sti 



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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 
QOt be replaced by air or hydrogen for drying the salt, 
aid the flask be kept out of the freezing-mixture, so long 
T still moistened the salt, without the latter taking an 
•colour. When dry and in an ammouical atmosphere, 
t is more stable, but cannot long be kept at the ordi- 
temperature 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 
ished into the water, and its very dilute solution distilled 
Ikali 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 

35.09 ; 
34.69 ; 



Sulphur diox. 

64.91 per cent. 
65,31 „ 



Found: 
S0^NH3),: 

he slight excess of ammonia indicated is safely attributable 
means taken to preserve the salt till it was analysed. 




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E. DIVERS AND M. OGAWA : 



Its properties^ constitutioTiy and name. 

The new salt is white and apparently crystalline, am 
Ijears 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 souiu! 
if dissolved by ice or enough ice-cold water, furnislies a sol 
answering all tests for pure ammonium sulphite* In this n 
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 th< 
Bait. When the salt is much decomposed, its sohition 
other reactions besides those of a sulphite. In anhydrous a] 
it dissolves freely, evidently as ethyl ammoniunisulphite ; 
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 u solid part, 
more or less orange- coloured, and into ammonia, the liquii 
undergoing further change into solid matters (p. 197) 

Constitution. — The salt is more probably an amiclo- th 
imido-compound, NH4N(S02NH4)2 (analogue of normal a 
Ilium imidosulphate), because it can be obtained only wh( 
temperature is kept down and the ammonia is in excess- 
still more probably a sulphuryl rather than a thionyl comj 
because of its feeble activity as a reducing agent and of it 
easy passage into ammonium sulphite or ethyl ammoniumsul 
It has accordingly to be formulated as NHg'SO/NHi, an 
NH2SOONH4. 

Name. — Since the salt represents ammonium sul 
NH40'S02*NH4, in which the ammonoxyl is replaced by a 



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AMMONIUM AMIDOSTJLPHITE. 



193 



is properly called ammonium amidosulphite. Berglund's 
of amidosulphonate now in use, for amidosulphate is 
;ly based on a misconception. The name, amidosulphinate, 
[ogy 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 
3, 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. 

istary. — Experiments made earlier than ours on the union 
bur 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 
L837, 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 
jed 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 
I solution, which is at first yellowish but soon, becomes 



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194 E. DIVEKS AlO) M. OGAWA : 

colourless, and gives, when recently prepared, the reactic 
mamly of a mixture of ammonium sulphate and trithionate, 1 
to a small extent those of a sulphite also ; and, lastly, that wl 
the solution is of certain concentration it gives a transient r< 
dish coloration with hydrochloric acid. 

Forchhammer {Compt. rend., 1837, 5, 395) found that, 1 
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 rm 
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 E 
berenier. 

The views advanced as to the nature of the orange b( 
have been, that it is a compound of ammonia with an isoi 
of sulphurous anhydride, which changes at once with wi 
into ammonium sulphate and trithionate, just as ammonium pj 
sulphite slowly changes in hot solution (Rose) ; that it is ami 
gen sulphide, S(NH2)2, mixed with ammonium sulphate (Foi 
hammer) ; that it is, partly, thionamic acid, NH2*S0'0H, par 
ammonium thionamate, both volatile, being its colour due 
impurity (H. Watts) ; and that it is ammonium pyrothionam 
NH2"S204'NH4 (Joergenssen). 

Interaction of the gases. — We have repeated Rose's exp 
ments of measuring over mercury the volumes of the gases wl 



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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 sur^ in water, though mixed 
3ther substances, and quantitative analysis of which shows 
nuch more than three inols. ammonia to two mols. sulphur 
le have gone to its formation. If, instead of examining it 
3e, 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 
ins amidosulphite or gives any sulphite to water, and contains 
luch 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. 

^oducts of the decomposition. — Both Rose and Forchham- 
■Qund ammonium sulphate to be a principal constituent of 
roduct of the interaction of the gases. A suflficiently high 
irature having been reached, this will have been the case ; 
jrmore, the solution of the even less heated product slowly 
les acid and full of sulphate. But when the temperature 
lot been allowed to exceed 30®, or even 40°, the quantity of 
ate in the product is so small that it may almost be dis- 



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196 E, DIVEBS 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 niti 
which might be that of trithionate and, in the case of his p 
duct, gave other reactions of a trithionate. But when the p 
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 hac 
be strongly heated under pressure. Besides this, the abseno 
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 
composition. 

Heating pure ammonium amidosulphite gives the same 
suits 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 i 
receiver kept well cooled, the product is deposited as a s 
waxy, yellow coating on the walls of the vessel and on the 
tubes. Its colour varies in different parts from nearly whit( 
orange-red somewhat irregularly but generally so as to be wh 
near where the ammonia enters, the whiteness not being du( 
moisture in the gases, as Rose assumed. When the product] 
to 30-35°, whether by its own heating or by external heat, i 
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 



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AMMONIUM AMIDOSULPHITE. 



197 



gain 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 
Lon of the gases. With very gradual heating, the tem- 
jr liquid product is much less coloured than in the other 
ts colour being evidently caused by the presence of a 
itter dissolved in it, which gives indications of being 

ds orange-red substance is never formed in more than 
nail quantity. It gives a yellow colour to the aqueous 
1 of the whole product, which, however, slowly fades away. 
1, carbon bisulphide, and other menstrua dissolve it out 
le 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- 
i becomes explosive when heated above 150°, and may 
ave become nitrogen sulphide, but before being heated it 
this substance. 

Lcept the very little sulphate already mentioned, there is 
^et 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 
)08ition that may be expressed by ONHg, 8SO2, assuming 
jsence of 2.5 per cent, moisture. The composition of the 
crude residue does not differ much from this. The alco- 



11 



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198 E. DIVEBS AI^D M, OGAWA : 

holic solution, cooled and charged with ammonia, gives mini 
scaly crystals in small quantity. This substance, dried in 
current of ammonia, has a composition expressed by (NHs)8S2( 
and dried in the sulphuric-acid desiccator, that of (NH8)2S2( 
These three substances all give the silver-nitrate reaction oft 
aqueous solution of the whole residue, and on boiling with dik 
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 deri\ 
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 fij 
heated with hydrochloric acid under pressure. 

From the mother-liquor of the above mentioned S2O3 sail 
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 aqueoi 
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 



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AMMONIUM AMIDOSULPHITE. 



199 



J early stage and the latter in the late stage of the de- 
)sition. This remarkable production of water, though 
5 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 
ilphite. 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- 
i, 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 
ir dioxide. 

Ve hope in a future paper to be able to report the com- 
Q of this investigation. 



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Products of heating Ammonium Sulphites, 
Thiosulphate, and Trithionate. 



By 



Edward Divers and Masataka Ogawa, 

Imperial University, Tskyo. 



^hat has been published upon the effects of heating am- 
n 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 
lulphite. We therefore make known what we have ascer- 



'Tf 



M'l 



Preparation of the salts used. 

mmonium sulphite^ (NH4)2 SOg, OH2. — Statements are con- 
; as to whether this salt can be got from its solution by 
ation (Muspratt, Phil Mag., 1847, iii, 30, 414 ; Marignac, 
&., 1857, 17 ; Forcrand, (hmpt. rend., ,1885, 100, 245 ; 
;, CompL rend., 1887, 104, 1793 ; Roehrig, J. pr. Ch., 1888, 
7), We find that a concentrated solution, charged with 



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E. DIVEE8 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 xn 
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 evaporal 
is to take advantage of the lessened solubility of the salt 
presence of much ammonia. Ammonia solution, sp. gr. Oi 
containing therefore about 28 grm. ammonia in 100 c.c, is tc 
treated in a flask with sulphur dioxide, while it is kept in i 
tion in a mixture of ice and salt, and with the tube convej 
the sulphur dioxide not dipping into the solution. The formal 
of a very little orange- coloured matter in the neck of the fl 
cannot be avoided, but this can be easily removed afterwa: 
When the solution has become thick with crystals, no ra 
sulphur dioxide is to be added, although very much ammc 
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 i 
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 rea 
it evolves ammonia and thus becomes the very deliquesi 
pyrosulphite. 

Anhydrom ammonium sulphite is readily obtained from 
hydrated salt by long enough exposure in the desiccator ; i 
very hygroscopic. 



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LMMONITJM SULPHITES, THIOSULPHATE AND TWTHIONATE. 203 

[mmonium pyrosvlphitey (NH4)28206. — 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 
; mixture, the pyrosulphite crystallises out from it, in quan- 
luivalent to a little over one-fifth of the ammonia taken, 
thrown out of solution by the sulphur dioxide. The salt 
J 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 
Moisture. This salt is also easily obtainable by evapora- 
s aqueous solution, but hardly free from sulphate, and not 
it some decomposition, through loss of sulphur dioxide and 
;h oxidation. It is much more soluble than the normal 
te. 

mmonium thiosulphate. — An old solution of calcium thio- 
te, obtained by boiling lime and sulphur together in water 
laving the solution until much of the pentasulphide had 
)xidised by the air, was decanted from insoluble matters, 

with ammonium carbonate in some excess, filtered, and 
i-eely 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 
«iccator. The well-dried crystals have been found by Lock 
:iuess (J?er., 1889, 22, 3099) to be anhydrous. 
[mmonium trithionate. — ^This salt has apparently not hitherto 
prepared by any one. Being exceedingly soluble in water. 



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204 E. DIVEBS AND M. OGAWA : PRODUCTS OF HEATING 

it cannot be prepared by Plessy's excellent process for the i 
tassium salt {Ann. Gh. Phys.y 1844, iii, ii, 182), or by 
slight modification by Hertlein {Z. phys. Ch.^ 1896, 19, 2i 
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 desiccal 
This very deliquescent and changeable salt cannot be kept 1( 
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, ii 
subliming vessel consisting of a test-tube, 15 cm. long and ab 
15 mm. in internal diameter. The tube was closed by a cao 
chouc stopper, and a very slow current of dried nitrogen throi 
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. 1 
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 



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IMMONITJM SULPHITES, THIOSULPHATE AND TRITHIONATE. 205 

J 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- 
ed only to be referred to the presence of minute quantitities 
purities. It all dissolved readily in carbon bisulphide, and 
Uised out on evaporating the solvent, 
t can hardly be doubted but that ammonium tetrathionate 
oentaihionatej if it can exist) would decompose in the same 
3 trithionate. Ammonium hypomlphate (dithionate) has been 
L by Heeren {Pogg.j 1826, 7, 55), and more definitely by 
3 {Ann.j 1888, 246, 194) to first become anhydrous, if not 
y so when heated, and then to decompose at «bout 130° 
ulphur dioxide and a residue of ammonium sulphate. 
immonium thiosulphate. — Zeise, in 1824 {Ghn. 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 
ig. A much more weighty statement is that made by 
g {Ber.j 1874, 7, 1159), namely, that the dry salt can be 
aed unchanged, intermediate dissociation being admitted, 
lave found it to decompose very slowly at 150°, the main 
cts being a sublimate of anhydrous normal sulphite and 
due of sulphur un fused, as in the case of the trithionate. 
also very small quantities of hydrogen sulphide and am- 
i 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 



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206 



E. DIVERS AND M. OGAWA : PBODUCTS OP 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 t] 
showed the former to be essentially anhydrous normal sulphi 
and the latter unchanged thiosulphate : — 



(NH,),S03 
Sublimate 
(NH,)A03 
Eesidue 



Ammonia 

29.31 
27.54 
22.27 
20.69 



Solphor 

27.59 percent. 
27.55 „ 
43.24 
42.31 



99 



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(N £1.1)28203= 2NH 
8H2+(NH4)2S306. When ammonium thiosulphate is rapidly a 
more strongly heated, ammonia is lost and sulphur sublimes ; tl 
as a matter of course and of no significance, thiosulphate s 
even trithionate are produced on adding water to the mi: 
sublimates. 

Ammonium pyrosulphiie. — We did not get this exceedin 
deliquescent salt into the tube ready for heating before it 1 
condensed some moisture, and to this we attribute part of the resi 
obtained. Change went on slowly in the salt at 130° and soi 
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 sublini 
was pyrosulphite in one experiment ; in another, it was this J 
mixed with a very little anhydrous sulphite. But there wai 



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fMONIUM SULPHITES, THIOSULPHATE AND TRITHIONATE. 207 

rable residue, more than one-third of the weight of the 
en, consisting of sulphate, trithionate, sulphur, and ap- 
f some tetrathionate. There was no sulphite or thiosul- 

The tetrathionate, the sulphur, and the sulphur dioxide 
jry 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- 
i (with no doubt intermediate dissociation), and that the 
e of a little moisture causes it to decompose partly into 
e and trithionate. 

ihydrotLB ammo7iium sulphite volatilises at about 150°, 
; a sublimate of the same salt, or rather, a pseudosub- 

for the salt surely dissociates when heated, 
/drated 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 
3ape 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 272 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 
i grm., and this had now lost one-fifth of its weight, the 
i having the composition expressed by (NH3)io(S02)6(OH2)7, 
ent to a mixture or combination of the three salts, hydrat- 
phite (39.49i), anhydrous sulphite (34.194), and pyrosul- 
[26.69i), dividing equally among themselves the sulphur 



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208 E, DrVEES AND M. OGAWA : PBODUCTS OP 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 


(NH3US0A(0H,), 


25.00 


56.47 percent 


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 oflf, the residue should have been 8472 per < 
of the hydrated normal sulphite, whereas it proved to be 1 
more than 79 per cent., in consequence of volatitisation of 8 
of the (dissociated) salt, made manifest by the production 
little sublimate. 

After renewing the heating in a fresh subliming-tube, all 
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 heal 
sulphur dioxide steadily escaped, but practically ceased to d 
long before sublimation was finished. The residue left ¥ 
sulphur dioxide was no longer coming oif, proved on analysis t 
normal sulphite again, but only half hydrated, 2(NH4)^03, ( 
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 ou 
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 



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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 
Iphate and trithionate, as already described. Heating in 
m tube, and more rapidly, Muspratt's results will be got, 
n weter is more quickly expelled, and some pyrosulphite 
posit as a sublimate. 



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aasimn Nitrito-hydroximidosulphates and the 
«i-e3ristence of Dihydroxylamine Derivatives. 

By 

Edward Divers, 1^1. D., D. Sc., F. R. S., Emeritus Prof., 

and 

Tamemasa Haga, D. Sc, F. C. S., 
Professor, T5ky5 Imperial University. 



ike potasaiimi nitrate (this Journal, 7, 56), potassium 
forms double salts with the potassium hydroximido- 
tes (sulphonBtes), the non-recognition of whose existence 
lowed mistaken notions to arise about the nature and the 
3ts of the sulphonation of nitrous acid. 
hlaMium nitrite and 2/3 normal hydroximidomlphatef KNO2, 
S03K)2 — The sparing solubility of 2/3 normal potassium hy- 
Qidosulphate in water is hardly affected by the presence of 
liim nitrite and when a sufficient quantity of the salt has 
lissolred by heat it crystallises out again almost pure on 
g the hot solution, even though the water has also dissolved 
siamnch aa one-sixth of its weight of the nitrite. When 
lution of the nitrite is stronger than this there crystallises 



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212 E. DIVERS AND T. HAGA : 

out instead of the hydroximidosulphate itself a combination c 
with a molecule of the nitrite. The same double salt is i 
formed in the cold when the hydroximidosulphate is triturs 
and digested with such a solution of the nitrite. Precauti 
being taken against the hydrolysis of the unstable hydroximi 
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 h 
rhombic prisms with 2OH2, its compound with the nitrite 'u 
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 nitrifc 
a strength of 10 per cent, or more nitrite. It is neutral to 
mus and very soluble in water but its solution soon depc 
crystals of the 2/3 normal potassium hydroximidosulphate uu 
it is very dilute. In any case the hydroximidosulphate can 
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 cone 
tratecl 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 niti 
oxide, a result of the hydroximidosulphate being a sulphoni 
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< 



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POTASSIUM NITBITO-HYDBOXIMIBOSULPHATES, 



213 



ively when heated — more so than does the hydroximido- 
tte 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 
rite 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, 
le hygroscopic character of a solution of potassium nitrite, 
aalysis 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 
lost of its sulphur appears as ordinary sulphate, but not 
ill ; so that in estimating the sulphur the solution must be 
[ysed for some hours at 150° under pressure. The results 
lysis were : — 



FouQcl, 
K3HN,SA, 



Potassium 

33.14 
33.10 



Sulphur 

17.95 per cent. 
18.06 „ 



here are other ways in which the potassium nitrito- 
)rraal hydroximidosulphate may be formed all consisting 
ally in producing the hydroximidosulphate by sulphonating 
11 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 



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314 E. DIVERS AND T. HAGA : 

of nitrite be employed. Potassium nitrite, 30 grains ; potass: 
hydroxide, 10 grams; water, 50 to 100 grams are to receiiJ 
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 enU 
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 sola 
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 nc 
first by Raschig but he attached to it a significance unlike 
here presented. Discussion of his views will be found towi 
the end of this paper. 

There is yet another way in which this potassium niti 
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 flufiy or cotton-like lustreless matter: 



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POTASSIUM NITBITO-HYDROXIMIDOSULPHATES. 



215 



Sites. If now to this fluffy matter suspended in its cold 
Br-liquor carefully decanted from every particle of the crys- 
if the 5/6 normal salt a hot solution of this 5/6 normal salt 
' or even 40 per cent, nitrite be poured in, a r^elatively 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 
3n of nitrite containing the 5/6 normal salt dissolved there is 
ble equilibrium between the tendency to yield HON (SO3 K)^, 
(S03K)2,OH2 again and that to form HON(S03K)2, 
0. 

(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 nitrite and normal hydrozimidosulphale KNO2, 
^(S03K)2, 4OH2. — This compound salt is only obtainable 
1 strongly alkaline solution. For when the normal hydrox- 
mlphate 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 



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216 



E. DIVERS AND T. HAGA : 



must be present in some quantity in the solution. The pres( 
of too much alkali causes a little of it to separate with 
normal salt, taking the place apparently of the water of cryg 
lisation of this salt (this Journal 7, o2), and similarly 
separate with the normal salt in its combination with niti 
then also seeming to lessen the capacity of the normal sali 
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 mag 
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 ani 
represent the soft voluminous spheres highly condensed. Probs 
these always begin their growth from a minute granular nuel 
The double salt can only be purified for analysis by pressin; 
on the porous tile, when the soft spheres become a felted lusti 
cake and the hard white granules crumble down like mass© 
wax. Analysis of the two forms has given us the foUoi?^ 
results : — 





Potssm. 


Alk. potssi 


Silky ; found, 


35.21 


9.92 


K;N3SA6,4.40IL, 


35.14 


10.04 


Granular ; found, 


33.99 


9.20 


KvN^SAe, 60H^ 


33.89 


9.68 



Sulphur 

16.23 per cent. 
16.43 „ 
15.90 
15.85 



9f 

J7 



The varying amount of water is only the recurrence of what 



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POTASSIUM KITBITO-HTDROXIMIDOSULPHATBS. 



217 



II 



recorded concerning the normal potassium hydroximido- 
late by itself. The double salt is exceedingly alkaline, its 
iniiy we eatimated 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 
crystals 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 
hydroxiniidosulphate which, simple or combined with 
e, contained less than its K7 to S4. By dissolving the 
0-2/3 normal bydroximidosulphate in a hot concentrated 
on of nitrite containing sufficient alkali the nitrito-normal 
^ximidosulphate can be readily obtained by cooling the 
on- 

Potmmim nitrite and potassium hjG normal hydroximidosuU 
. — We have obtained three compounds of the ^jQ normal 
mi\\ nitrite, one being 7KNO2, 2HK5(NS.A)2, 8OH2. By 
: an almost ^aturtited solution of potassium nitrite con- 
ig a little potassium hydroxide and dissolving in it by heat 
/6 normal hydroximidosulphate there is obtained a compound 
iimte fibrous cryalals 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 
xiches that indicated by the formula given above. For 



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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 (=30H2) a 
then approached in composition Fremy's sulphazite. 





Potasm. 


Sulphur 


Alk. Potssm. 


Original salt, found, 


36.81 


14.35 


480 per cent 


Recrystallised, „ 


36.68 


14.47 


4.51 „ 


K.,H,N„S,0«, 30H„ 


36.87 


14.20 


4.34 „ 



A second double salt, 3KNO2, K5H (NS207)2, OH2, was got 
dissolving one mol. 5/6 normal salt and 1.4 mol. potassium 1 
droxide in a hot 65 per cent, nitrite solution and cooling, 
appearance it resembled the other compound salt. Its analj 
gave : — 





Potssm. 


Sulphur 


Alk. potssm. 


Found, 


36.17 


15.07 


4,51 per cent. 


Gale, 


36.81 


15.06 


4.61 „ 



A third double salt, anhydrous, 7KNO2, 3K6H(NS207)2, ^ 
not prepared synthetically but by treating an almost satura 
solution of the nitrite with alkali and sulphur dioxide, and j 
ding alkali again after the sulphonation, imitating a process 
Fremy's. Then, filtering the heated solution from much cr 
talliue 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 : — 



Found, 


Potssm. 
36.94 


Sulphur 
16.37 


Alk. potstun. 
4.96 per cent 


Gale, 


36.99 


16.51 


5.05 „ 



This salt was quickly resolved by water into nitrite ai 



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POTASSIUM HITBITO-HYDROXIMIDOSULPHATES. 



219 



ala of the very sparingly soluble 0/6 normal hydroximido- 

late, 

rbo varying proportions in which potassium nitrite and the 

iormal hydroxinudusulphate unite would possess but little 

St were it not for the fact that they have evidently been 

ally met with and taken to be salts of specific constitution 

remy and by Rasohig. 

Non-existenee of Dihydroxylaminesulphoyiates. 

^'reray believed in the existence of less sulphonated deriva- 
of potassium nitrite than his sulphazite (see next paper) it- 
jss sulphonated than his sulphazotates (hydroximidosulphates) 
Lttrihuted his failure to find them to the fact of their possess- 
xceediDgly high solubility. Claus held much the same 
and believed that by adding to an aqueous solution of 
miiu nitrite an alcoholic solution of sulphur dioxide in not 
uge a quantity he had obtained an impure crystallisation 
salt, ON SO,K {5er. 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 
jgether 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 formulae : — 






(SO,K)/ \0K 



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220 



E. DIVERS AND T. HAGA 



These he prepared by partial sulphonation of the nitrite 
known ways. They both yielded crystals of a hydroximidosi 
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 1 
cooling an apparently homogeneous crop of crystals of alm< 
the same composition and properties as one or other of Raschij 
salts. Kaschig gave two ways for preparing the salt having t 
second of the formulse just given and in these ways we ha 
obtained the nitrito-2/3 normal hydroximidosulphate already c 
scribed in this paper, but mixed with a little potassium sulphi 
This impurity accounts for the alkaline reaction of Raschij 
preparation and the presence in it of a little more than Kg to I 

He got the other salt (K2 to S) only once and in the fo] 
of white crusts when working unsuccessfully for hydroximidosi 
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 Frei 
— 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 lo renc 
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 



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POTASSIUM NITBITO-HYDBOXIMIDOSULPHATES. 



221 



the next crop of the salt itself as to cause its rejection, 
presence of sulphite in less quantity in the first crop of 
ils will have been masked by the oxidising action of the 
t oxide in becoming nitrous oxide. That sulphite was pre- 
in Raschig's preparation well accords also with the fact that 
slum hydroxide added in excess precipitated potassium sul- 
, for, although hydroximidosulphate is itself decomposed by 
oaost concentrated solutions of potassium hydroxide into 
lite 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 
nee known.* Inspection of Raschig's formulae is of itself 
ient to prevent their getting accepted as in accordance with 
'acts. For from these formulae both salts should be strongly 
ine, while in reality one is neutral. Above all it is hardly 
ble that dissolution in cold water should suflSce to cause 
3sulphonated nitrogen to become disulphonated. 
Raschig held his two salts to be identical with Fremy's 
sium sulphazite and sulphazate respectively ; but the nature 
remy's salts will be found, we believe, more precisely given 
le paper following this. The point we would here insist 
I is that Raschig's preparations, judged by their chemical 
viour, have no claim to be considered as dihydroxylamine 
natives, being in every way indistinguishable from synthe- 

[)f the 3KN0, 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 
iis 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. 



41 



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222 



E. DIVEBS AND T. HAGA : 



b 



tically prepared compounds of nitrite and hydroximidosulphat 
Dihydroxylamine salts have as yet only a hypothetical existei 
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 siuj 
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 uudet 
rained constitution, both of which were most probably a 
nitrito-hydroxiraidosulphates. They may therefore be notic 
here although Raschig did not represent them to be dihydrox 
amine derivatives. Yet they were evidently closely like 1 
other two in properties. One was isomeric with potassium hyj 
nitrososulphate (Pelouze's salt) and also with his (Kg to S) ' dil 
droxylamine ' salt, allowing for different hydration, and the otl 
was isomeric with potassium 5/6 normal hydroximidosulpha 
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, g, 8 
It was got by dissolving nitric oxide in solution of potassii 
sulphite and hydroxide and evaporating to a small volume I 
crusts formed. If we assume that air or nitric peroxide was i 
excluded there were the conditions present for getting a nitril 
hydroximidosulphate, for, as we show in a paper which w 
shortly follow this, nitrous fumes passed into potassium sulph 



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POTASSIUM NITBIT0-HYDR0XIMID0SULPHATE8. 



223 



)ii generate hydroximidosulphate freely together with 

he other salt isomeric with 5/6 normal hydroximidosulphate 
btained in Raschig's attempt to form 2/3 normal salt by 
g surphur dioxide into a solution of potassium nitrite and 
side and letting stand for a day. These, too, are conditions 
3ting nitrito-hydroximidosulphate. Now, both products 
in being decomposed by water in such a way as to yield 
dmidosulphate 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 
Q hot alkaline solution a little nitrous oxide which only 
Kyamidosulphate is known to give. These peculiarities 
ty 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 3KNO2, K5H (NSiOyji, 2OH2, and 
L compound we have described on page 218 ; that for the 
ide of the 5/6 normal hydroximidosulphate treated as being 
te compound is SKNO.,, 6K2HNS2O7, 5K5H(NS207)2, whicli 
er should give crystals of K2HNS2O7, 2OH2. 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. 




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Poti^ntt 


SulpUur 




Kitft>&o isomer, foimd, 


35.72 


14.40 


jwr cent 


Catciiliitedj 


.S6.05 


1475 


IT 


D. & H^^ salt, found, 


.36.17 


15.05 


» 


Oxiraido isomer, „ 


33.04 


21.23 


»I 


Calculntedj 


32.91 


21.54 


1) 



For the present , the existence of isomesides of Peloiize's 
iuhI Fremy^s basic sulphazolate must be regarded as no longer i 
probiible. 



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dentification and Constitution of Premy'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. S., 
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 
im siUphazate. With slightly diminished concentration of 
ution he generally obtained instead the brilliant, often 
rhombic prisms of potassium basic sulphazolate (5/6-normal 
limidosulphate, this journal, 7, 15). But sometimes 
was obtained neither of these salts before the solution 
5 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 
im 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 potassium neutral sulphazotate (2/3-normal hydr- 
osulphate Raschig) and, lastly, with still greater dilution 



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226 



DIVERS A HAGA : IDENTIFICATION AND CONSTITUTION OF 






V 



the minute brilliant needles of his potassium- sulphammoi 
(nitrilosiilphate Berglund). Still other salts he believed tc 
produced in the first stages of the reaction between the nit 
and sulphur dioxide, one of which he named potassium m 
aziie ; 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 F^mall quantity and as crystal 
warty granules — by the action of water upon the ' sulphazi 
whereby this was converted into ' basic sulphazotate ' which 
posited and a solution that on evaporation yielded the ' aul 
azite/ These two salts could together in solution be char 
back into the * metasulphazotate ^ while the ' sulpha zite * 
the ' sulphazate ' could similarly often be changed into the ' m 
sulphazate ' again. These two ^ raeta ' salts he regarded there 
as perhaps merely double salts of the others. The ' sulphaz 
the ' sulphazate/ and the * sulphazotates ' be treated as b 
members of a series of salts in which there were to two aton 
nitrogen from one up to eight atoms of sulphur j — three in- 
" sulphazitOj four in the ' sulphazate', and five in the ' sul 
azotates.' With this conception of the nature of these salts, b; 
on his analyses, it was easy to understand the decompositioj 
the ' sulphazate ' into the ' sulphazite ' and the * sulphazot; 
But this and other of Fremy's interpretations of the facta 
served by him have lost all importance and particular intc 
through the progress of chemistry since his memoir was f 
lisbed and only his account of the facts requires considera 
now. 

Subsequent work by others and ourselves in the same i 
has sliown that Fremy in the account he gave of the prepara 
of his many salts went two little into details as to the coniliti 



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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 
^'s 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. 
I 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- 
in of the other salts. In Fremy's working, this most easily 
i salt came only as the final product of the sulphonation 
berefore gave him no trouble. Claus emphatically displayed 
epticism as to Fremy's results ; yet in nearly every point 
ich he difiered from Fremy as to the facts we find Fremy 
^e been right. When Raschig repeated Fremy 's work — 
ith the modifications in procedure introdticed by Claus — he 
isults similar to, though less unsatisfactory than, those Claus 
btained. He made an approach to Fremy's work in so far as 
le often got very little nitrilosulphate ; nevertheles3 he too 
in his attempts to prepare the * sulphazate ' in Fremy's way. 
n perhaps all essential points we can lay down the method 
)eat Fremy's experimental work successfully. But in some 
a little uncertainty obtains owing to the fact that the very con- 
ited and complex solutions which yield Fremy's salts are apt 
)Osit what is virtually the same salt in diflferent forms as well 
times salts quite distinct from each other under only slight and 
re variations in the circumstances attending their formation. 
Sulphazate. — This is Fremy's first salt directly obtained in 
ilphonation of the nitrite. In getting it he took approxi- 
y 5 mols. potassium nitrite to 2 mols. potassium hydroxide 
I little water and into the solution passed sulphur dioxide 



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228 



DIVERS & HAGA ! IDENTIFICATION AND CONSTITUTION OF 



I' 



until it became almost filled with silky needles very solubL 
water. So far it is easy to follow Fremy with a full measui 
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 fa4:t 
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 * sulpha^! 
was strongly alkaline and very caustic and when decomposer 
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, 19.5Q| 
nitrogen, 4.9. We can learn what his after-treatment was 
reference to other parts of his paper where he speaks of the 
necessary (when sulphonating the nitrite) to maintain the n 
Unity of the solution by adding potassium hydroxide from tin 
time and of dissolving sulphazotised salts for examination in ^^ 
containing this alkali. Certain it is he must have added i 
potassium hydroxide to the solution after getting it to crystal 
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 < 



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FBEMY's 8T7LPHAZOTISED SALTS OP POTASSIUM. 



229 



ion 18 to deprive it of much of its nitrite and to convert the 
lormal into more nearly normal hydroximidosulphate — to 
ice, therefore, potassium nitrite by potassium hydroxide, 
ipting Fremy's mean numbers as accurate, what he analysed 
the composition, 
11K8NS207,0H2; K^HNS^O;, 2OH2; 2(K2HNS207, KNO.). 



Found, 


PotsBin. 

34.9 


Sulphnr 
19.55 


Nitrogen 
4.9 


Alk. potssm. 

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 
flScient to say of his * sulphazate ' that it was the silky 
itus-like nitrito-2/3 normal hydroximidosulphate more or less 
erted into the also silky asbestus-like normal hydroximido- 
late, an account 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 add 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. 
SvJphazite. — What Fremy named potassium svJphazite he only 

obtained, and then not by direct sulphonation of the nitrite, 
le 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 hydroximidosulphate), and left a mother-liquor which 
old evaporation till syrupy yielded the sulphazite. It showed 
i analogy with his sulphazate but was distinguished from it 



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930 DIVERS & HAOA ! IDENTIFICATION AND CONSTITUTION OP 

by having liltle tendency to hydrolyse and by at once evolving 
some nitric oxide when its solution was mixed with a dilute acid. 
Water decomposed the sulphazile^ but into what products was not 
ascertained. 

We have suflSciently realised Freray'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 : 



Fremy's salt, 


Potassm. 
38.16 


Sulphur 
16.27 per cent 


D. & H's salt, 


36.94 


16.37 „ 



and agreed with it in chemical properties, so far as is known, 
At the same time it was indistinguishable from a compound ol 
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 * mammiliated crusts' as Fremy got, which, 
though only in rough agreement in percentage composition with 
hm 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 u double salt. 

MetoBulphazateJ^' — In Fremy's experience it sometimes hap- 
pened, when passing sulphur dioxide into solution of nitrite and 
nlkali of a concentration intermediate to that giving sulphazaU 
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 metasulphazotate in the French original, but not in the Gennan 
tmnslation. 



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FREMY 8 SULPHAZOTISED SALTS OF POTASSIUM. 



231 




sentrated solution of sulphazale and sulphazite ; also by 
; a solution of sulphazate and then cooling it. When 
ly compressed the jelly became a transparent wax-like mass. 
I 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 
therefore have been that of nitrite combined with normal 

normal hydroximidosulphate in such proportions and with 
idditions perhaps of alkali as prevented crystallisation. 
7e have not had Fremy's success in getting this salt in 
3f jelly and wax but have met with just such phenomena 
forming barium sodium hydroximidosulphate, BaNaNS207, 
as will be found described in our paper already frequently 
id 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- 
m 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 
©ion, 3(KN02, 2K3NS2O7, 4OH2); K3NSA; 3OH2. 



)nnd (Fremy), 
„ (D.&H.), 
Uculated, 



Potaem. 
35.10 

35.10 

35.06 



Sulphur 

16.74 
16.68 
16.74 



Nitrogen 
4.81 



5.23 



A Ik. potssm. 
per cent. 

10.47 „ 

10.23 ,. 



Ye got the salt by dissolving the hydroximidosulphate in 



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232 DIVEES & HAGA : IDENTIFICATION AND CONSTITUTION OF 

hot concentrated nitrite solution containing alkali. To 100 cc 
water there were present 4572 grm- nitrite and V/^ grm. potassian 
hydroxide ; for 66 mol. nitrite there were dissolved 10 mol 
anhydrous normal hydroximidosulphate. But for the salt beinj 
in beautiful asbestus-like fibres, there was nothing to distingoisl 
it from the jelly and the wax-like metasulphazate, which, therefore 
we do not hesitate to class as a nitrito-hydroximidosulphate. 

Basic sulphazotale^ which Fremy considers next, has beei 
shown by us already {loc. dt.) to be the 5/6 normal hydroximi 
dosulphate, and not the salt of a distinct acid, the sulphazotic 
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/i 
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 
iral sulphazotates are hopelessly out of accord with its constitutioi 
and properties, though those for the basic sulphazotate are satis 
factory enough. 

SulphazidatCy produced by the hydrolysis of the sulphazotatej i 
hydroxyamidosulphate (Claus). Sulphazilate and meta^ulphazilate 
oxidation products of sulphazotate are ON(S03K)2 and ON(S03K)3 
and have been studied by Claus, Raschig, and Hantzsch. 

Metasulphazotate. — Sometimes Fremy got a salt in the forn 
of spangles {paillettes) j in appearance like minute crystals of icwi 
sulphazotate^ but differing from these in not being hard under pres- 
sure. This salt be named, therefore, metasulphazotate. Accordinj 



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IT 



FREMY S SULPHAZOTISED SALTS OF POTASSIUM. 



233 



a it is also obtainable by mixing (hot) solutions of the (basic) 
izotate and sulphazite. It is very soluble in water, very 
ne and unstable unless the water contains alkali. In pure 
it becomes basic sulpJiazotate and sulphazite again. It 
the greatest analogy with metasulphazate and is distin- 
5d in the same way as this salt from basic sulphazotaie. It 
)e a compound of basic sulphazolate and sulphazite. So far 
f. It will be evident that there is nothing in its history 
)perties to distinguish it, except its occurring in the form 
xkling particles and even that can be met with in the 
mlphazotate suddenly precipitated ; we have also got other 
e sulphazotised salts in what may be called spangles, 
1 not this particular salt. In the preceding paper, page 214, 
\re described an impure form of nitrito-2/3 normal hydrox- 
lulphate obtained by dissolving the 5/6 normal salt in a hot 
itrated 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 metasulphazotate and is 
less alkaline than the metasulphazotate and is much less 
le than the metasulphazate. It may be formulated as 

K3NS2O;; 9(KN02,K2HNS20;,1V20H2). 

PotflBm. Sulphur Nitrogen Alk. potssm. 
und (Fremy), 33.8 18.6 3.5 I>ercent. 

„ (D. &H.), 33.79 18.35 1.09 „ 

Jculated, 33.68 18.37 7.63 1.12 „ 

ulphammonate and sulphamidate are respectively nitrilo- 

ite and imidosulphate (Berglund). 




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On a Specimen of a Gigantic Hydroid, 

Branchiocerianthus imperator (Allman), 

found in the Sagami Sea. 



By 



M. Miyajima, Rigdkushi. 
Science College, Imperial University, Toky5. 



With Plates XIV d: 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 

irmounted a long stalk which evidently fixed the animal 

I sea-bottom. A circle of numerous gi'oceful tentacles hang 

from the margin of the disc, while on its upper surface 

m 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- 




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236 



M. MITAJIMA 1 



meters and the prevailing colour transparent scarlet. It 
agreed on all sides that it was a New Year's gift from Otoht 
and that it Blionld be known in Japanese as Oiohwie 
Hanagam. 

The specimen p 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, piec^ of the tentacles 
the dendritic appendages were fixed in the sublimate aut 
Perenyi's fluid. 

The specimen waa handed over by Prof, Mitsukuri to 
to work out ite finer structure. 

It was evident from the fii'st that the specimen was 
similar to the form only a short time before described by Jl 
('98) as Branchiocerianthtis urceolus. I started, tlierefore, 
an idea that I was dealing with an Actiniun, 

As I proceeded in my investigation, however, it became ] 
that til is idea was not tenable, and the conclusion was fii 
reached that the animal was very closely allied to Coiymoi 
and that it belongs probably to the species obtained l>y the ** C 
lenger '' at about the same locality and named by Allman 
Monocanlits imperalor^ notwithstanding many discrepancies bet 
his description and the specimen. This conclusion was como] 
cated through Prof, Mitsukuri to Dr, Mark and a request was 
sent to him J that during his opportune stay in Europe, he shi 



^Otohunc-' b a befloilful gf}dd<^.^ who h eapposed to have lier palaces al thebol 
the sea. ^Mlanagafia" is Ihe flower-^nn-whatle or amnineiilal fumuwL Thus OttJt\ 
Hanagwfi nie&iis *' the omatiirntQl pnrn^ol of Otuhime/^ 



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BRANCHIOCERIANTHUS IMPERATOR (aLLMAn). 



237 



sible, examine the original specimen of Monocaulvs imperator 
) British Museum. To the results of his examination of the 
lens I shall return in the later part of this paper, 
leanwhile an article was published in the Zoologischer An- 
by O. Carlgren (*99) throwing doubt on Mark's Branchio- 
thus being an Actinian, and contending that it more 
bly is a Gorymorpha or at least a form standing very close 
nfmorpha. 

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- 
v^hich had been communicated to him, as mentioned, by 

and thinks that both his and our specimens belong to the 
genus and that our specimen is probably identical with the 
nuliLS imperator of Allman. 

before going further I wish to express my deepest feeling 
ligation to Prof. Mitsukuri 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 
Dmes, all showing in their arrangement a well marked 



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i 



238 



M. MIYAJIMA : 



bilaterality. The hydrocaulus, which is attached not tc 
center but to the edge of the hydranth, is nearly cylindrica 
iuereases 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 ai 
tentacles, both oral and marginal, were deep scarlet in c 
while the gonosomes possessed light rufous colour. The 1 
caulus was light pink in colour, being quite pale in its na 

The general features and the colours are well sho 
Fig- 1, PI. 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 objecl 

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, butj 
from that of Branchiocerianthus urceolus, in having its si 
diameter less than its transverse^ the two diameters] [being 
pectively 80 and 90 mm. (Woodcut 2). 

At one end of the sagittal diameter is attached the b 
caulas 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 Branchiocerianthus urceolus Mark. 

The edge where the hydrocaulus is attached I shall des 
the lower, and the opposite the higher, edge. 



r 



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Ill 



BBANCHIOCERIANTHUS IMPERATOR (ALLMAN). 



239 



Woodcut 1. 
1. 




mt 



IS showing sagittal (1.) and transverse (11.) sections of the hydranth. 
jrpoetomal region of the disc ; C orifice of the diaphragiu (m) in the liydranth ; (7 orifice 
iphragm (m') in the hjdrucaulus ; eg, central, ig. lateral, globule of the gonophore ; 
', H* lower cavity of the hydranth ; h, hypostome ; ic. intercalated cord ; HO 
lus; ml, marginal, o(, oral, tentacle; P, peduncle of the gonosome; 22. outer 
the disc, provided with the radial canals (r.c). 



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240 M. MIYAJIMA : 

The hypostome (Woodcut 1, I, II, h), the superior pro 
tion of the disc, is slightly conical, diminishing gradua 
its diameter from the base towards the free end whei 
mouth opens. A little below the mouth the hypostome b 
brush-like group of about 180 filiform tentacles {ot) whi( 
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 syj 
glyph which is said to be present in the oral tube of Bra 
ceria7ithus 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 o 
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, corres 
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 si 
stretch as far as the margin of the disc, while laterally and tc 
the higher edge it is distant from the margin 35 mm. and 2! 
respectively. It thus assumes an ovoidal outline, the pointe 
attainuig the lower margin of the disc and passing directly 



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BRANCHIOCERIANTHUS IMPERATOR (aLLMAN). 
Woodcut 2. 



241 





Diagram showing the upper surface of the disc, 
hiatus at the lower edge of the diHC. Other letters as in Woodcut 1. 



of the hydrocaulus. The longer (i. 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. 



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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 an 
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, E.) which run from 
base of the gonosomes to the margin of the disc. This re 
thus assumes the form of a wide horse-shoOj whose two i 
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 (PI. XV, Fig. 1, r.c.) slightly swell oa 
surface of the disc thus giving the latter an undulating apj 
ance. The canals are intercalated by solid cords (PL XV, Fi 
i.e.) which appear on the surface of the disc as opaque Hues. 
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 h 
edge are smaller and shorter than those higher up, until at 
both arm-ends they are practically nil. On the other hand 



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BRANCHIOCERIANTHUS IMPEBATOB (aLLMAn). 



243 



Is on the higher side of the disc are not so long as those 
he lateral 9 but run straight from the base of the gonosomes 
le 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 
le hydranth-cavity where the cavities of the gonosomes stand 
^mmunication 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 (PL 

Fig. 1). 
The name of marginal tentacles {mt) is given to the outer- 

circle of filiform tentacles arranged like a fringe around the 
;in of the disc. The circle is not complete, there being a 
18 (v) at the lower edge of the disc where the surface of the 
)stome 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 
le disc they increase successively in length until about the 

(counting from the lower edge) is reached, of which the 
bh 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 




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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, j 
upper {H) and a lower {S). The superior prolongation of tl 
upper cavity is that of the hypostome, which does not she 
any indication of the septal partition. The lower cavity is mo 
spacious than the upper, and not only occupies the whole loi«? 
part of the hydranth but extends also through the entire leng 
of the hydrocaulus. 

The membrane (m) separating the hydranth-cavity arii 
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 lo^ 
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 spei 
by the numerous intercalated cords {vide supra) to the upi 
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.. 



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BBANCHIOCERIANTHTJS IMPERATOR (aLLMAN). 



245 



>f sections (PI. XV Figs. 4-9) passing through the lines 1-1, 
•3, 4-4, 5-5, 6-6 in Fig. 1, PI. 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 {tb.) 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 
)ond8 to the enlarged end of the solid cord. The cavity 
tentacle is almost filled up by the spongy endoderm which 
ihe 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 
le 2-2, Fig. 1) cut just inside the margin of the disc, the 
canals already assume their characteristic shape in cross- 
i, 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-lamellse, the upper, 
and the vertical. The upper lamella (u.l.) is situated along 
rface of the disc, the basal (i.Z.), in the floor {i.e. in the dia- 
n), and the vertical (v.l.) connects these two lamellae. 

traced inwards, (PL XV Figs. 6, 7) the intercalated cord 
es thinner and thinner, until it no longer shows in cross- 
i the form of a trapezoid, but assumes the shape of a tri- 
formed by two vertical and one basal lamella. Where the 
)me arises (Fig. 8), the vertical lamella does not reach the 
lamella ; hence the radial canals communicate here with one 
3r and form the upper common cavity of the hydranth. 



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M, MIYAJIMA 



Withitt the circle of the gonosomes (Fig, 9) the upper la 
stands entirely separated from the basal, oo which the V€ 
lamella shows itself only an a ridge-like line which in ( 
Bectiou is reeogiiizable as a simple small knob. 

Preserved in formalin, the fine tissues of Uie animal 
unfortunately mostly gone. Luckily, however, the pieces c 
gonosomes and the marginal tentacles, which were preserv 
sublimate, ete,, helped us to ascertain something of the histoli 
character of the animaL 

The wall of the animal-boilyj I need hardly Bay^ consi 
the three layers, ecto-, endo- and mesoderm as in other Cc 
terates* 

The ectoderm, the outermost layer, has been entirely 
olV from the specimen in formalin, but in the pieces fixed 
sublimate was well preserved. This tissue is a single laj 
cyliudrical cells which in their preserved condition are mo 
1^3 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 c 
endoderm. This tissue was well enough preserved ev€ 
formalin so that the structure of the animal could be h 
made out by this layer alone. 

The endoderm, the innermost layer, which lines the i 
cavity of the animal, remained unfortunately only here and 
in the specimen in formalin. From these patches it cou 
made out that the eudoderm lining the hydranth-cavity is st 
cells thick (Figs. 3 & 10)- The cells are irregularly forniei 
contain but a little cytoplasm which is pi-essed towards the 
with the nucleus. Consequently the wall of the cavity gi 



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BRANCHI0CEEIANTHU8 IMPEKATOR (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 
le 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 
itirely filled up with a tissue which reminds one of the verte- 
I notochord. It has the same structure as the spongy en- 
rm of the hydranth-cavity already mentioned. Only near 
3a8e yof 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 
aunicating 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 
be branched tubular stalks, upon which the gonophores 
grouped 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 
dnds (Figs. 13 & 14). The one kind of which there is only 
in each cluster is situated on the top of the terminal branch, 
5 the others take a more lateral position. The former is 
ir than the latter, consisting of the irregularly shaped cells 
ly vacuolised (Fig. 14, e.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, 



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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 tc 
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, Lg.) are mostly spherical 
consist of compactly packed cells rich in cytoplasm. The d 
derm prolonged from the branchlet distinctly separates 
ectoderm from the central cell-mass. After examining b 
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, 1 
is a wide cavity occupying the whole interior of the glol 
which communicates with that of the branchlet. I could 
detect gonophores developed any further than this in our s] 
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 oi 
Hence the dendritic gonosome of this animal is a peculiar o 
which bears on a common stalk the sexual and defensive elerui 



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BBANCHIOCERIANTHUS IMPEKATOR (aLLMAN). 249 

In Other hydroids these two elements are borne on separate stalks, 
as for example in Pennalia. 

HydrocauloB. 

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. Mabk, 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 

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250 



M. MIYAJIMA : 



half of it there are visible from outside 15-20 longitudinal 
bands (Fig. 17). They stand about 2-3 mm. distant froi 
another and run down to about the middle part of the 1 
caulus where they become obscure. From the surface the 
remarkably like the mesenterial filaments of an Anthozoon. 
wavy bands anastomose here and there with one anothe] 
give to the hydrocaulus of our specimen an appearance 
resembling that of Corymorpha. Though the bands are in tl 
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 
Bpecimen, in which the ectodermal and endodermal cells 
mostly lost, I could not ascertain whether the wavy bandi 
the endoderm canals, a structure peculiar to Corymorpha^ c 
I think it, however, very probable that they existed, anc 
rise to these band-like appearances. In the published ac 
of Monocavim imperator of Allman the endoderm canalj 
plainly described and figured. 

The mesoderm is very well developed, especially 
hydrocaulus where it reaches a thickness of about 0.2-0.5 
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, /./.) ai 
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 



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BRANCHIOCEEIANTHTJS 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 (op) 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 (op.), which is seen 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. 



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252 



M. MIYAJIMA 



w 




Summary. 

1. The most striking feature in our specimen is its g 
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 1 
symmetry in a hydriform person as very different from 1 
pressed for instance in the planoblast of Corymorpha and L 
which is but temporary and occurs only at a certain p< 
development, or from the biradial symmetry as express 
few genera like MonobracMum 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 man; 
canals visible even on the surface of the disc. That ren 
structure is not, however, peculiar to our specimei 
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 narrow 
the upper, so that the former forms a slender canal in th( 
of the '' Wulst." Gosta Gronberg ('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. Gronb 
"Mesoderm-wulst." 



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BRANCHIOCEBIAMTHUS IMPEEATOK (aLLMAN). 



253 



ivity 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 
irginal tentacles. 

The tentacles are filiform and arranged in two sets, oral 
and marginal (proximal), as is characteristic of the 
es of TvhviaridcBy OorymorphidcBj and Monocaulidcd. 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 
are, 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. 

ConBiderations on the SyBtematic Position of our 
Specimen. 

hose who would compare the account given above of the 
ire of our specimen with that of Branchiocerianthus urceoluB^ 

('98) will not for a moment doubt that we have in these 
3es essentially similarly constituted animals. It seems almost 
iuous 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 



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254 



M. MrSTAJIHA : 



I 



bilateral symmetry, being interrupted at the lower (or 
posterior) edge where the hydrocaulus is attached. 

That our specimen and Brachiocerianthus urceolus b 

least to the same genus, there can hardly be any doubt. 1 

they belong to the same species is another question. It 

haps premature to decide this point, at present, in as i 

Mark has not yet published his full paper. Judging f 

preliminary notice which deals exclusively with the < 

features, the following are the chief points of diflference. 

a. The general shape. B. urceolus is stated to have an e3 

graceful, symmetrical vase-like figure with flaring Up 

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 produci 

upper end of the animal a sort of eccentric funnel 



depression. 



<t 



a 



A. 



B, 



^ The fancied resembl 
the animal to a little 
which this side view pres 
suggested the specific 
adopted — urceolus.** (M 
p. 148). The pitcher 
shape of the hydranth 
due to two causes : (1) i 
ing in of the disc-marj 
(2) the extreme obliquit; 
axis of the hydranth to 
of the hydrocaulus. In 
nexed woodcut, a repres 
axis of the hydrocaulu 



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BBAKGHIOOEBIAMTHUS IMPEBATOB (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 transverse diameter 
which is the greater of the two. The following measure-r 
ments will make this point clear. 

Sagittal (longit) diam. Trans, diatn. Ratio of trans, 

m mm. in mm. diam. to sagittal. 

B, urceolus. 

Small specimen... 25 15 60% 

Large specimen. ..38 30 nearly 79?^ 

iSSrColC' 80 90 112.5./. 

c. The size: — 

T^*i^K r^p kU^ \.xfA,^ Maximum length Maximum length 

^"J^»'luf if mm"'''" i '^^ r^--' °f *''^ »"'' ♦«"'«=''' 
lemacle m mm. m mm. 

B. urceolus 105-200 125 30-35 

S^^Coile^' 650 300 50-55 

d. The lower end of the hydrocavlus : — Mask 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. 



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256 



M. miyajima: 



e. The radial canal: — Mabe mentions that the radial ca 
B. urceolus run " from the base of the oral tube to th 
of the marginal tenbicles, before reaching which viany 
fork J each of the branches communicating with the luni 
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 diflferences are to be regarded a 
specific or due simply to the diflferences in size, age, i 
must leave for the present an open question. I am ii 
however, to think that B. urceolus and our specimen 
diflferent species. 

References have already been made several times in the 
of the foregoing pages to the resemblance of our speci 
Monocavlus imperator of Allman, a gigantic hydroid i 
by the Challenger oflf 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 as 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 observi 
the fresh object, it is very easy to overlook such a feature as 1: 
symmetry when the disc is lying in the midst of a : 
tentacles. Of course the best thing we could do under I 



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BBANCHI00EBIANTHX7S IMPEBATOE (alLMAN). 257 

cumstances was to appeal to the original specimens. At the 
request of Prof. Mitsukuui, Prof. Mabk, who was opportunely 
staying in Europe at the time, was kind enough to examine the 
type specimens of Monocaulvs 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 eflforts 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 
Branehiocerianthus urceoltts 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. urceohjLS " 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- 
ceoluSf as shown above, the hydrocaulus is attached at one end 
of the sagittal (longitudinal) diameter of the disc. But Prof. 
Mabk adds, '^ the specimens were so much wrinkled and folded 
that I have not much confidence in this conclusion." There is 



* In oor specimen the opening which pnts the hydranth- cavity and that of the hydrocaalus 
in communication is not elongated, but almost circular as already stated. 



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258 



M. MITAJIMA 



one carious point of difference between our specimen and 1 
catUus iwperator. 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 Ali 
there are on the whole strong probabilities in favor ol 
assumption. Those who read carefully Allman's descriptioi 
notice that the points which he brings out distinctly ii 
structure of his species, such as a wide cavity extending thi 
the entire length of the stalk, the presence of the stalk-meso 
in the shape of a fibrillated membrane — a point which Al 
emphasizes as ** the most striking feature in the histology c 
Hydroid " — and so forth, are absolutely similar in our speci 
If we remember in addition that both came from practicall 
same locality, it is, I believe, within the scope of reasoG 
ness to conclude that our specimen belongs to Monocaulus impi 
of Allmann. 

If this is really the case, we must examine other spec 
Monocaulus. The genus includes, besides Monocaulus impe 
two other species ; M. gla^ialtSy (Sars) (for which Allman 
blished the genus) and M. pendula, (Agassiz). These two i 
show, however, a radial symmetry, and now that M. impi 
is shown to have a bilateral symmetry, can not possibly b 
in the same genus with the latter. M. imperator must thei 
be separated from the other two species and placed in a 
genus. According to the rules of nomenclature, this new j 



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BRANCHIOCEBIANTHT7S IMPEBATOR (aLLMAn). 259 

must take the name Branchiocerianthvs^^ first given by Mark, 
and our specimen then ought to be known as 

Branckiocerianthm imperator (Allman). 

P.S. We shall await with interest the full report on the 
specimen of Monocaulus obtained by Prof. Chun in his recent 
deei>-sea expedition (^99). 

December, 1899. Zool. Institute, Science College, 

Imperial University, Tokyo. 



♦Mark ('99) mentions that Bhiaofruma eamea of S. F. Clakke (76) may be a form same 
as, or closely related to, BrandUooeriarUhuB, Clarke's original description is very brief and 
it is impossible to determine whether Mark's suspicion is correct or not. At any rate, 
Clabks makes no mention of any bilateral symmetry in the structure of the animal. 



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260 



M, HFTAJIMA : 



Literature, 

Agassis, L. 

* 62. Contributions to the Natural History of the United Stat 

Americiu VoL IF, (BoBton). 

AllMAN, Gr. J. 

* C4. On the cnnRtniction and limitation of genera among the 

droidea. Ann. and Mag. of N. H.j 3rd series^ No, 77, 
' 7L A monograph of the gymnoblastic or Tnbnlarian Hyd 
(London.) 
* 83-' 88. Report on the Hjdroids dredged by R lif , S. Challenger d 

the years 1873-7C. 
Braubb^ A- 

' 91, Ueber die Entatehung der GeRchlechtsprodnkte nnd Entwick 
von Tnbularia mesemhri/anthemum, Zeit /iir lotss^ 
Bd. 52. 
Carlgrek, 0* 

'99, BranckiorerianilmB urceolus^. L, Mark, eine Hydroid ? 
Aug. Bd, XXIL No. 581. 
Chun, C. 

* 91* Coelenteraten, m Broom's Klass, und Ord, dea Tien^eicl 

* 99, Die deut«;he Tiefaee-Expedition 1898/1899, (Berlin), 
Clarke, K F, 

'7C. Report on thf* Hydroids collected on the ooast of Alaska an 
Aleutian Islands, by W, H, Dall, U,8< Coast-Survey and 1 
from 1871 to 1874, 

DOFLEIN, F. 

^96. Die EihiWnng bei Tubnlaria ; Z. Jur Wififf. Zool. Vol, 62 
Gr5nberg, G, 

' 98, Beitriige znr Kenntnias der Gattang Tnbularia. Zoo!, J 
Bd, IL Heft I, 



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BBANCHIOCEEIANTHUS IMPERATOB (aT,LMAN). 



261 



li 



WN, 0. 

82. Der Oi'ganismus der Hydroid-Polypen. Jen. Zeit, Bd. 8. 
Vol. 8. (Jena). 

:8, Th. 

68. British Hydroid Zoophytes. 1868. 

^OCH, G. 

73. Vorlaufige Mittheilung liber Colenteraten. Jen, Zeit Bd. 7. 

^ E. L. 

98. Preliminary report on Branchiocerianihus urcedns. Bull, of 
the Mm. of Crnip. Zool. Vol. XXXII. No. 8. 

99. *' Branchiocerianthus/' a correction. Zod. Anz. Bd. XXII 
No. 590. 

fANN, A. 

83. Die Entstehung der Sexualzellen bei den Hydronnedusen, 1883. 



Explanation of Plate. 
PI. 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 ; Ect. ectoderm ; End. 
Tm ; iZ^ upper, W lower, cavity of the hydranth ; tc. intercalated cord ; 
nesoderm ; ttU. marginal tentacle ; p. peduncle of the gonosome ; R. 
region of the disc, provided with the radial canals ; re. radial canals ; 
men of the base of the marginal tentacle. 

ig. 1. Surface-view of a portion of the disc with gonosomes (p) and 
lal tentacles {mt.). Nat. size. 

upper wall of the disc, h a part of the diaphragm in the hydranth. 
ig. 2. Longitudinal section through the outermost margin of the disc 
1x4. 



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262 



Fig. 3. Cross-section of the upper wall of the lower cavitj c 
hydranth. Zeiss DDx2. 

Fig. 4-9. Serial sections of the upper part of the disc. Zeis 
Fig. 4. Cross-section through the line 1-1 in Fig. 1^ 



Fig. 5. 






2-2 


Fig. '6. 






3-3 


Fig. 7. 






4-4 


Fig. 8. 






5-5 


Fig. 9. 






6-6 



Fig. 10. Cross-section of the radial canal and intercalated amL 
BBx2. 

Fig. 11. Cross-section of the marginal tentacle. Zeins n^x2. 

Fig. 12. Ijongitudinal section of the same. Zeiss aj^ x 2- 

Fig. 13. Terminal branches of a gonosome, Zeiss a x 2. 

Fig. 14. Longitudinal section of a branchlet of the gonosome. 
Fx2. eg. Central, Lg. lateral, globule. 

Fig. 15. Central globule with nematocyst (n). Zeiss DDx4. 

Fig. 16. Lateral globule in which the bell-nucleus (?jj^) and the 
derm-cup (enc.) are fairly well recognizable. Zeiss Fx2, 

Fig. A part of the wall of the hydrocaulus with the* wavy 
Nat. size. 

Fig. 18. Cross-section of the mesoderm in the hydrocaulus, Z«l^ 
h.l. outer, longitudinally, c.l, inner, circularly, striated layer* a?, spc 
responding to the wavy band. 

Fig. 19. Mesoderm of the hydrocaulus, macerated ^^'lth eaiis?ric ] 
Zeiss a x 2. 

Fig. 20. Surface-view of the root and the sheath. Ks^l, ^ize. ap 
like appendage ; s. sheath. 

Fig. 21. Longitudinal section of the root figured in Fig 20* 
ax 2. 0. outgrowth of the endoderm; other letters as in Fig, 20. 



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Jour. ScLColl. Vol. XIII. PI. Xy. 



Fig. 3 



Fig. 16 



b.n. 








/./...^ iy >f'~^-,jigi,i,edbyi^OO^^ e- 



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I 



lutual Relations between Torsion and Magneti- 
zation in Iron and Nickel Wires. 

By 

H. Nagaoka, RigakuhakusfU, 
Professor of Applied Mathematics, 

and 

E, Honda, RigakusM, 
Post-graduate in Physics. 



With Plate XVI. 



The various effects of stress on the magnetization of ferro- 
^netic metala are of such a complex character that no simple 
itioti seems to exist among them. The strains caused by mag- 
iising the ferromagnetics are of no less complex a nature, so that 

co-ordination of these two classes of complicated phenomena 
up to tlie present, still a matter of doubt. Various isolated 
ts, such as the analogies between the change of magnetization 
longitudinal pull and that of length by magnetization, the 
Ltion 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 
lularly magnetized wire respectively, were long considered as 
rding a clue to the explanation of these phenomena. So far 



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264 



H, HAQAOKA AIS'D K< HONDA : 



as we are aware^ no attempt has yet been made to plac 
these ditterect phenomena oij ti common footing. Some tii 
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 It? 
magnetized wire and the longitudinal magnelization ca 
twisting a circularly magnetized mre. It is our objec 
present paper to show that tliese difl'erent phenomena 
linked together in a common bond. 

§ 1. Twist produced by the intaraetiou of cirouia 
and langitudinal magnetizations. 

Tlie subject was? first studied by G. Wiedemann'^ w 
blished remarkable reciprocal relations with tlie lon| 
magnetization produced by twisting a cii*cularly magnetic 
Di\ Knott^^ fouud that the direction of twist in iron is 
lo that in nickel ; BidwclP* afterwards discovered that tl 
in iron h reversed in high Melds and takes place in t] 
direction as in nickeL Unfortunately some of the exp 
were undertaken with wires which were longer than tha 
coil J so that the magnetization was far from being unifc 
will suffice for qualitative tests, but we can not hope 



l)Nsigniika anM llundn, tliiH JiHim. 13, p. 57, 10(H1; PhiK mag. 49, V* 341, 
2)G. Wietlemaivn, Pugg. Ann. 103, p. 57], 185.^ j 106, p. Jt>l^ 1859; J^iHr, 
:t)KnnW, Tmns. Kt*j, Sue. Kdinij., 32 d), p* lOiS^ 1882/8;jj 35 (*)> P» ^^7,, IS 
p, 4^5, 1B9L 



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TOBSION AND MAGNETIZATION. 



265 



ite quantitative results. The position of maximum twist 
Lckel 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 
ixtremities of an iron or nickel wire 21 cm. long were 

brazed stout brass wires, and a 
light plane mirror was attached 
to the lower one. The end of 
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- 
Qt 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 
int by enclosing it in a wide braes tube with a small win- 
just where the reflecting mirror was attached. The twist 
measured by scale and telescope method, by which the 
3tion of 0.3" i)er. cm. was easily read. The current was 
ured by Kelvin graded amperemeters, whose constants were 




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266 



H. NAOAOKA AND K. HONDA : 



from time to time checked by means of an ampere 
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 longitudini 
magnetizing current. 

2. The longitudinally magnetizing current was kept const 

and the amount of twist measured by varying the circuli 
magnetizing current. 

Before each experiment, care was taken to demagnetize 
wire completely either longitudinally or circularly by passing 
alternate current of gradually diminishing intensity. 

Twist by varying Ihe 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 twh 
reversed increases with the circularly magnetizing field. In nic 
the direction of twist is opposite to that in iron, but the gee 
feature is similar to iron, the only diflFerence being that i 
in strong longitudinal fields, the twist is not reversed. For ? 
of the equal thickness, the amount of twist in nickel is grc 
than that in iron — the maximum twist in iron wire of 1 ] 
diam. by passing 6 amperes through it amounts to about 
per cm., while with nickel wire of 0.83 mm. diam. under sin 
conditions, the maximum twist amounts to about 200/' 



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TOBSION AND MAGNETIZATION. 



267 



Tivist 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, 
lie longitudinal field remains constant. Such is also the 

with nickel in moderate and strong fields. In low longi- 
Qal fields, however, the twist does not continue to increase with 
circular, but we notice a maximum as will be clear in the 
re. There is great experimental difficulty in increasing the 
liar 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- 
m. The twist, however, goes on slowly increasing, till it 
3es 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 
3 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 
jriments 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 
rably high field, while in the present experiment, it occurs 
'ly in the same field as in iron. It may partly be due to the 



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268 



n. NAGAOEA AND K. HONDA! 



tlifference in tlie method of measuring the twist and parti 
the non-uniformity of the field, as was often the case in raoi 
the older experiments. 

The observed angles of twist in iron and nickel are exhil 
in the following tables, where C denotes the longitudinal cut 
per sq, mni< in amperes, H the field strength in C,G.S, i 
and T the angle of twist per cm. expressed in seconds. 

Circular Meld being Qmstant. 
Iron wire: diam.=0.98 mm. 



c= 


= 1.06 


c. 


=3.38 


C = 


= 6.95 


H 


T 


H 


T 


H 


r 


5.6 


18.6" 


5.6 


21.9" 


5.6 


16.4 


12.4 


21.7 


11.3 


35.6 


11.3 


30.0 


26.0 


13.1 


22.6 


32.8 


•22.6 


36.4 


41..3 


7.9 


36.2 


24.7 


49.2 


27.8 


90.5 


2.5 


79.2 


13.7 


78.0 


19.7 


112.0 


1.6 


97.3 


10.7 


96.2 


16.4 


215.0 


- 0.5 


191.0 


6.0 


132.0 


8.2 


492.0 


- 1.9 


442.0 


2.7 


45,10 


2.5 



Nickel wire: diam.=r0.83 mm. 



c= 


=2.45 


C= 


=4.33 


C: 


=6.05 


H 


7 


H 


T 


H 


r 


4.9 


- 941" 


4.9 


- 97.6" 


4.9 


- 88.1 


11.2 


-137.0 


11.2 


-164.4 


11.2 


- 159.i 


24.3 


-12.5.5 


24.3 


-179.4 


243 


-196.^ 


38.8 


-102.1 


38.8 


-156.6 


38.5 


-182.1 


53.6 


- 85.1 


53.4 


-134.1 


53.2 


-159.5 


84.0 


- 62.9 


83.8 


-101.8 


84.0 


- iig.-; 


102.2 


- 54.5 


102.2 


- 88.0 


103.4 


-lOU 


225.0 


- 31.7 


222.0 


- 47.9 


226.0 


- 59.; 


415.0 


- 19.6 


389.0 


- 29.8 


414.0 


- 40.2 



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TOBSION AND MAGNETIZATION. 



269 



Longitudinal Meld being Omstant. 
Iron Wire: diam. = 1.05 mm. 



H= 


=2.67 


U= 


=5.92 


H= 


=19.38 


C 


T 


C 


T 


C 


T 


0.35 


0.3" 


0.58 


0.3" 


0.40 


0.0" 


0.80 


1.1 


1.03 


0.6 


0.89 


0.0 


1.06 


2.0 


1.73 


2.0 


1.36 


0.0 


1.80 


3.7 


2.20 


3.0 


1.92 


0.3 


2.22 


5.7 


2.96 


5.1 


2.66 


0.4 


3.04 


8.8 


3.55 


6.2 


3.48 


0.6 


4.35 


14.2 


478 


9.1 


4.88 


1.4 


6.79 


21.5 


6.73 


13.6 


6,25 


2,5 


7.52 


23.2 


7.34 


14.7 


7.16 


2.8 



Nickel wire: diam.=0.83 mm. 



H= 


=5.7 


H= 


=39.7 


H= 


=96.3 


C 


r 


C 


T 


C 


r 


1.03 


22.1" 


0.95 


22.6" 


0.81 


8.5" 


1.89 


48.1 


2.06 


59.3 


1.76 


19.9 


2.82 


70.2 


3.34 


101.4 


2.82 


32.3 


4.50 


102.0 


4.10 


123.6 


4.29 


50.5 


7.25 


121.2 


5.90 


153.6 


6.22 


73.2 


9.40 


116.4 


7.95 


172.8 


8.65 


99.6 


11.60 


107.4 


11.40 


178.8 


11.50 


1-27.5 


13.38 


99.0 


13.10 


176.4 


13.22 


141.6 



§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, 

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270 



H. NAGAOKA AND K. HONDA ; 



one^^ of us investigated the transient current for iron 
nickel wires. It was then found that the current due to t^^ 
ing was opposite in direction in these 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 sti 
fields. In order to make this point clear and see if any i 
mate relation with the Wiedemann effect could be traced, fresh 
periments were undertaken by the same method as b€fore, 
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 Fij 



Iron 


wire: diam.= 1.33 mm. 


Nickel 


wire: iliiiiu.= 1.09 m 




length =20.90 cm. 




l«?ngth =20.80 en 


0= 


15° 


d= 


50° 


e= 


15° 


6=50° 


H 


Qxio' 


H 


QxlO 


H 


QxlO 


H Qx 


3.2 


25.4 


3.2 


30.7 


1.3 


- 5.8 


1.1 - 


5.3 


29.0 


4.8 


33.6 


3.7 


- 7.1 


3.7 - 


15.6 


24.9 


15.8 


42.0 


16.0 


-17.7 


5.3 -1 


32.6 


16.3 


30.5 


38.4 


33.7 


-21.-2 


10.2 --J 


545 


10.9 


48.6 


30.5 


53.4 


-21.5 


35.6 ~:i 


87.4 


6.4 


86.0 


18.5 


81.2 


-20.1 


44.9 -A 


120.8 


2.5 


139.4 


8.1 


115.4 


-19.2 


67.6 --J 


165.6 


1.5 


183.8 


3.7 


157.0 


-16.4 


107.4 --J 


242.0 


- 0.1 


327.6 


- 3.2 


213.7 


-12.6 


160.2 -3 


495.4 


- 1.7 


447.4 


- 5.0 


319.6 


-10.3 


241.5 -■: 


650.2 


- 2.9 


711.0 


- 5.1 


530.3 


- 6.5 


316.4 ~'2 


908.0 


- 3.3 


959.0 


- 5.2 


703.0 


- 5.4 


699.8 -1 


1298.0 


- 3.1 


1521.0 


- 4.7 


1214.0 


- 3.9 


1150.0 -I 


1790.0 


- 2.5 


1872.0 


- 4.4 


1815.0 


- 2.0 


1S53.0 - 



l)Nagaoka, Journ. Sci. Coll., Tokyo, 4, p. 323, 1891. 



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TORSION AND MAGNETIZATION. 271 

Here 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 a 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 
uruski (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 

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272 



H. KAGAOEA Aim E. 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 maguetL 
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: (liam.= 0.888 mm. 


Nickel 


wire; (Iiara.= 0.965 




length =20.74 


cm. 




length =20.94 c 


d= 


15° 


0= 


50° 


d= 


15° 


fl=50 


C 


Qxio' 


C 


QxlO* 


C 


Qxio' 


G Q: 


0.21 


0.7 


0.21 


16.1 


0.21 


- 45 


0.14 - 


0.85 


3.7 


0.69 


45.3 


0.66 


-HI 


0.23 - 


1.53 


20.5 


1.49 


89.1 


1.56 


-205 


0.90 - 


2.36 


31.4 


2.19 


111.0 


2.40 


-239 


2.05 - 


3.93 


36.5 


3.27 


125.6 


3.35 


-256 


2.87 - 


4.72 


33.6 


4.65 


1242 


4.36 


-2C5 


4.42 - 


6.55 


29.2 


5.91 


115.4 


5.86 


-272 


7.37 - 


7.89 


21.9 


8.29 


109.6 


7.95 


-269 


10.34 - 


12.82 


13.9 


12.48 


86.2 


10.89 


-256 


15.33 - 


19.01 


10.9 


17.08 


65.7 


1407 


-243 


20.85 - 


24.29 


5.8 


24.37 


51.8 


20.18 


-219 


23.04 - 


28.64 


5.8 


29.14 


43.2 


26.46 


-206 


26.13 - 



C denotes the total current through the wire express 
amperes ; 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 i 



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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. S. Shimizu, a post-graduate in physics, to whom 
our best thanks are due. 

§4. Theory. 

As already remarked in our last paper on magnetostriction, 
KirchhoflTs 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 R through 
angle <» amounts to 

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i 



274 H. NAGAOKA AND K. HONDA 



- ^(o V HR. {A) 



in field jET, and that the mean longihidinal magnetizaiion cs 
by twisting a ferromagnetic wire carrying an electric curre 
amounts to 



[; ^\<oV'C. (B) 

I 11 The reciprocal relation between these two phenomena m 

apparent at a glance. We shall next show how the same pi 
mena are reciprocally connected with the torsion produced b 
interaction of the longitudinal and circular magnetizations. 
The stress components in a magnetic medium as giv^e 
KirchhoflF are as follows : 

^'= -(^ + ^ + ^)^+ t(^ + ^ - ^^' ) (-+ ^+ 

Taking the axis ofz in the axial line of the wire, and two 
axes in the plane perpendicular to it, we see that the comp{ 



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TOBSION AKD MAGNETIZATION. 



275 



letic forces in a longitudinally magnetized wire traversed by 
lectric current are 



a= — A s^in 0, /5= h cos d, 

e A denotes circular field given by 

2Gr 



r=u, 



A= 



'W 



ing the current, r the distance of the point from the axis 
le wire, R the radius, and the angle between r and the 
of X. 

The stress components in ferromagnetic medium acted upon 
tie forces above specified are given by 

= -(-^ +*+ -rY'^'^+^i-^ + i-;k') (z/^+/r), 

=jr, =(^^ + 't + -y-)^ Hsind, 
moment about the axis of the wire is given by 
= -ff {-^ + ^ + ^yHrdxay. 



' c/«/<*. 



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276 



H. NAGAOEA AND E. HONDA : 



Since -^- and k are very small compared with k'\ the torai 
couple twisting the wire amounts nearly to 



— -5- CniP= -^ X Cross section, (C) 



Since the amount of torsion of a cylindrical wire by a j 
couple is inversely proportional to the fourth power of its ra 
it is evident that for given longitudinal current and fields 
angle of twist is inversely proportional to the square of 
radius. This inference was approximately verified in the pr 
experiments. 

In deducing the three formulae {A)j {B)^ (C), we 
not, strictly speaking, put k" outside the sigu of integr£ 
because the strain coefficient depends on the tield strength, w 
is not uniform in a wire traversed by electric current, 1: 
in these formulae, we shall have to use a mean value to o 
a close approximation. 

The mutual relations between twist and magnetizatioB 
embodied in the three formulae above given. There we 11 
that the strain coefficient &" determines the nature of the 
different phenomena studied in the above experiments, 
fact that the coefficient k" 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 t 
depending on k\ which depends mostly on the change of vol 

In order to test the consequences of the theory as rej 
the twist produced by the joint action of circular and longitm 
magnetizations, we have calculated the twist by assuming 



jtf 



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TORSION AND MAGNETIZATION, 



277 



3 of lc\ calculated from the changes of volume and of 
b 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 

place in 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 
jrably large in iron. In using the strain coefficients, we 
always bear in mind that these values are widely different 
iing 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. 
\a regards the mutual relations among the three different 
)mena above enumerated, it will suffice to state that several 
Bm have already been noticed by G. Wiedemann in his 
'ches on the relation between torsion and magnetism. He 
tally 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 
\red in the relation between torsion and magnetization in order 
•ing to light his ingenious theory of rotatory molecules. 



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. 






'? 



♦» 



I . 



278 H. NAGAOKA AND K. HOimA : 

Elegant as it at first sight appears to be, Wiedemann's theoi 
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 efiects of varic 
physical agencies. He showed that the existence of a oerb 
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 t 
circular magnetizations, a transient current will be prodn 
simply by twisting a longitudinally magnetized wire anc 
longitudinal magnetization will be developed by twisting a ( 
cularly magnetized wire. 

The peculiar feature of Kirchhoff s theory lies in the sire 
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 sti 
coefficients Ic and ^" which play the r6le& of different mo 
in the theory of elasticity. 

The reciprocal relations between the strain caused by n 
netization, and the elTects 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 



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TORSION AND MAGNETIZATION. 



279 



magnetization, we may conveniently place them under the 
ving parallel statements : 



ins produced by magnetisation. 
—(Experiment and theory). A 
Lidinally magnetized wire is 
i by circular magnetization. 

-(Experiment and theory). A 
rly magnetized wire is twisted 
igitndinal magnetization. 

-(Experiment and theory). Up 
lerate fields, the twist produced 
5 longitudinal and circular mag- 
bions of an iron wire is op- 
to that in nickel. 

—(Experiment and theory). 
wist due to longitudinal mag- 
bion of a circularly magnetized 
r nickel wire reaches a maxi- 
in low fields. 

-(Experiment and theory). In 
fields, the twist due to longi- 
,1 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'j — (Experiment and theoiy). 
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- 

lelmholtzy Crene's Journal 100, p. 137, 1886; Abk, 3, p. 203, 1895. 



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J 

i 



280 



ties, the accelerations and .the forces by p'«, q'», q"s, and Ft, 
the relations are generally expressible by the equations 



dP. 



9P, 



\^J 


9P» 9Pa ' 


(2) 


8P, ap» 

9?* 9?a ' 


(3) 


9P. _ aP* 






It will be easily seen that the relations above cited belong t 
case (2). 

The greatest difficulty that we encounter in establishing tli 
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. If a 
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 1 
detect its various defects, not only from qualitative points of vie\ 
but also in various quantitative details. 



l_ 



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le 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, Tokyo 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 ; 
)yrosulphites are not active to their whole extent upon ni- 
i ; (3) 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 ba^e of the sulphite, that may be used in place 
le 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 
lusions drawn by other workers from their experiments ; the 




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I 

I 

I 

I 



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 ioterpretation 
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 propositiouB, taken along 
what we have already published as to the constitution of Fi^ 
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 ^altB are secoi 
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 hii 
believed. 

I. — a. A Normal Sulphite inactive ufmn a Niinte. 

Dipotassium or dieodium sulphite is quite inactive u 
nitrite. In establishing this fact we have mixed soluti< 
normal sulphite and nitrite in proportions varying in dil 
experiments, and left them in closely corked flasks, almosi 
for days and for weeks. No change has ever happened, 
coloured with rosolic acid, a drop of dilute acid would a 
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. h.). 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^ 



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INTIIHACTION BETWEJEN SULPHltES AND NITJEIITES. 



28S 



—the minute quaotity 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 w,as, 
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. 



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284 



E. DIVEKS AND T. HAGA 



Claus's statement that some potassium sulphite neutml to li 
was as active upou uitrite, after lie 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 marl 
alkaline action upon litmus and yet small enough to leave i 
pyrosulphite unchanged. 



I. — b. Potassium Hydroxide not a Factor in llw Form^tioi 
the Sulphazotised Salts. 

That a normal sulphite, potassium or sodium, remains 
inactive upon nitrite, when alkali hydroxide is added, was i3 
tained by leaving the three substances together in sokition 
closed flask for some time, as in the experimeots ivliei 
hydroxide was present, and then precipitating with barium 
ride after addition of ammonium chloride, and finding no sul 
compound left in the filtrate. (Ammonium chloride pre 
precification of hydroximidosulphate, this Journal 7, 48, i 

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 b( 
neutral to lacmoid paper. In proportion as the hydroxide 
appeared, sulphite became abundant, whilst from the time 
the replacement of hydroxide by sulphite was complett 
quantity of sulphite steadily decreased as the isulphazotised 
formed, sulphazotised sodium salts being very soluble no i 



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INTEllACTION BEtWEJEN SULPHItES AND NITRlT^. 



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 ammonium 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. 



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286 E. DIVEES AND T. HAGA : 



Sulphur dioxide istlOoc. 2nd 10 cc. SrtllOcc. 

as 

Sulphite .0662griii8.=96,69i .r204grm8.=96.59^ .3996grnis.=91.5< 
Sulphazot. .0023 „ = 3Ao/o .0047 „ = 3.5o/o .0365 „ = 8.5 

It will be seen that all but 3.5 per cent, of the sulphi 
dioxide entering the solution in the early stages of the expei 
ment remained in the form of sulphite, and that even up to tl 
time when the last of the hydroxide had been consumed, all b 
8.5 per cent, of the total sulphur dioxide was in the slate of sulphii 
That it must be impossible to prevent all temporary local excess 
sulphur dioxide will be at once admitted, as also that it mi 
be diflScult in the later stages to keep down this local excess 
very narrow limits. Therefore it will seem in the highest degi 
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 sulphnzotised salts and endeavoured, accordingly 
keep some of it always present when passing sulphur dioxi 
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 pots 
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 wl 
the alumina contained in the potassium hydroxide began to pi 
cipitate, since this occurs while the solution is still strong 
alkaline to litmus. Raschig, in attempting to prepare Frem; 



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mlphazaiej 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 sulphazoiised 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 




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288 E. DIVEES AND T. HAGA : 

sulphite must also have been present. The two salts were, then 
fore, together in solution unchanged. Raechig, too, found thi 
sulphite and nitrite are inactive upon each other when in present 
of potassium hydroxide dissolved in only its own weight of wate 



II. — a. Even Pyrosulphite only active upon a Nitrite tiU 
it has become Normal Sulphite. 

Pyrosulphite, neutral to lacmoid paper and containing thei 
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 inacti 
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 tl 
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 :— 2NaN02 + SNa^SsOg + OH2 = 2Na^HNS207 + 2Na2S 
using less pyrosulphite, some nitrite remains at the end alo] 
with normal sulphite. That sodium pyrosulphite is not easi 
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 



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never be enough, with ordinary care, to cause much less than 
one-third of the sulphite to remain inactive. If excess of pyro- 
sulphiteis used the interaction appears to be — NaN02 + 2Na2S20s^ 
NaaNSjOg+NaaSOa, 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, juat 
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 normal 
sulphite as its action as an acid upon the nitrite, and not as the 
yielding up of half of its sulphurous acid for the sulphonatioii 
of the nitrite, the interactions being 2NaN02 + Na2S205 + OHj^ 
2HN02+2Na2SO„ and then 2HNO2 + 2X828205 =2Na2HNS,0; 
(see section III a). 

II. — b. Alkali not produced in the Sulphonation of 
a Nitrite. 

One of the most remarkable things, according to Glaus, is 
the production of potassium hydroxide by the formation of FremyV 
salts through the agency of a sulphite. He explained this pro- 
duction by the equation— KNO2+ 2 K2S03 + 20H2=K2HNS20; + 
3KH0. Such an equation was also published by Bergluiid 

o 

[Lands Univ. Arskr. 1875, 13, 14). Raschig gave the same 
equation for results obtained by himself and, in order to express 



pi)' 



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E. DIVERS AND T. HAGA : 



other results, gave also the equation — NaN02+2NaHE 
Na2HNS207 + NaHO. Finding also, and again in agreement 
Claus, that dipotassiura hydroximidosulphate does not coi 
at once or even at all with potassium hydroxide, lie argnei 
this salt cannot have a similar constitution to that of Fn 
* 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 ; 
or here show in other sections of the present paper, except 
the generation of alkali hydroxide, which we now proec 
deal with. Clauses 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 potasd 
80<lium become hot and strongly alkaline to litmus when 
together, and then contain much hydroximidosulphate and n 
sulphate, both neutral to litmus, which soon crystalliae i 
they are the potassium salts. That is all these chemists h 
evidence for the production of the hydroxide ; let us add to 
facts that addition of excess of barium chloride removes a 
alkalinity. It follows, since pyrosulphites are a little ai 
litmus and normal sulphites are very alkaline to it, thj 
phenomena depended upon offer no grounds whatever fo 
belief that alkali hydroxide is produced. Except by the i 
lime, baryta, or other base, there is, we believe, only one w 



I 



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INTERACTION BETWEEN SULPHITES AND NITRITES. 



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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). 

III. — a. A Pyrosulphite all active upon Nitroics 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 HNO2, 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 



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E. DIVERS AND T. HAG A : 



«olution of pyrosulphite (and of normal sulphite, but of tli 
treat in sect. III. b,) to nitrous fumes which act as nilrou 
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 
Tiitrilot^ulphate, notwithstanding the presence all along of so. 
jjyrosulpliite ; for, as was pointed out by us long ago, in 
eiently cold solutions sulphonation to nitrilosulphate h 
occurs. 

The next five sections (III. b, c, rf, e, f) treat of \i 
mixturcB which, from the acid constitution of one of the 
poneiits, behave like that of nitrite and pyrosulphite, that 
if each contained pyrosulphite and nitrous acid, 

11 L — b, formal Sulphite also all active upon Nitrom Ac 

Replacing the pyrosulphite used in the last experimei 
the normal sulphite, it was found that again but in this 
gradually, 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 yery fa 
wards completion. The reaction is expressed by the equat 
:iHNO,+2K2S03=2KNO,+OH2+K2HNSA, from which 
seen that only one-third of the nitrous acid becomes aulphoii 
the rest being used up simply as an acid. 



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INTERACrriON BETWEEN SULPHITES AND NITRITES. 



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This interaction is what, we believe, Rasehig must inadver- 
tly have got, when seeking to prepare Pelonze's salt (hyponi- 
losulphate) by the use of nitric oxide. The conditions are 
curable to the production of the nitrito-hydroximidosulphate 
s vol., p. 222). 



III. — c. Action of Sxdphur Dioxide upon Normal Sulphite 

and Nitrite. 

It has been shown in this paper (sect. I. b) that the hydrox- 
losulphate which, from the first, accompanies the normal 
)hite as joint product of the action of sulphur dioxide upon 
ili nitrite and hydroxide, keeps steadily to small proportions 
he sulphite until nearly all the hydroxide has been saturated. 
^v that point is passed and when, therefore, sulphur dioxide 
aeeting 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. 
3 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 
lulphonating the nitrite, which it then does by combining 
I it in conjunction with the pyrosulphite, thus: — 

2KN02+K2S205 + 2S02 + OHo=2K2HNSA, the hydroximi- 
ilphate 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 



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E, DIVEBS AND T. HAGA 



It 



with its inhibitory effect upon sulph9nation (sect, II- a), 
this change it still holds true that it is nitrous acid itself m 
is sulphonated, the potassium leaving the nitrite to entei 
sulphonate radical, and being replaced by hydrogen. 

Glaus held that there could be no difference between 
effect of submitting a nitrite to the action of a sulphite and lb 
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. 

Ill, — d. Action of Carbon Dioxide and of an Aeid Carbo] 
upon Normal Sulphite and Nitrite. 

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 eseitin] 
tion in a solution of these salts. Concerning tlie activil 
carbon dioxide there is nothing to add to what was publlsL 
our first paper (J. Ch. Soc, 1887, 51, 661), that the gas is 
slowly absorbed by the mixed salts in solution though m 
either salt alone and at the mean temperature, and that ei 
azotised salts are then produced. Normal carbonates of the al 
are inactive. 

It is known that nitrites are not decomposed by a 
dioxide, and also that alkali carbonates are decomposei 
pyrosulphites as freely at the mean temperature as by sul 
dioxide itself. Accordingly, we have found that potassiu 
sodium acid-carbonate dissolved along with exceas of nc 



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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 -KNO2 + 2K2SO3 + 3KHCO3 =K2HNS A + 3K2CO3 
+ OH2, 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 



{ 



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E. DIVERS AND" T. HAGA : 



sulphite and acid carbonate are intermediate products, the latt< 
of which separates for a time from concentrated solutions. T\ 
linve made furtlier experiments to ascertain the effect of the fir 
portions of the sulphur dioxide in producing hydroximidosu 
pliJite, 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 tii 
instead of one, and of weighing both instead of merely measB 
ing them, then in the one we determined the sodium, as sulphj 
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 wj 
admitting of no refined accuracy, that at a later sampling t 
solution contained at most, as much as 3^/2 per cent, less sodii 
than at an earlier sampling, a difference however hardly lai 
enough to need attention. 

The flask for receiving the portion for the sodium determ: 
at ion 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 necessarj'^ the sodium hydroxide \ 
for fixing the composition of the solution at the time it v 
sampled ; sometimes acid carbonate was deposited by it, sou 
times hardly at all ; sometimes the precipitated acid carbom 
slowly disappeared sometimes not. The solution used contain 



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INTERACTION BETWEEN SULPHITES AND NITRITES. 



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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- 
pbate 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 bad 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, 
02.7 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. 
rhat 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 
;rams had become sulphite. 

Uniform results are here, however, as when hydroxide is 
jtarted with, only obtained by uniform working, of which the 
following experiment is a good example. A solution of sodium 
nitrite and carbonate was divided approximately into one-fifths and 
four-fifths, and both portions were treated, as nearly as could be, 
ilike, their unequal quantities making the only difference. The 
jmaller portion when it had received 20 per cent, of the full amount 
}f sulphur dioxide was found to contain 61.8 per cent, of it in form 
af sulphonate, 38.2 per cent, of it as sulphite. The larger por- 
tion, having received 25 per cent, of the amount necessary for 



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E. DIVERS AND T. HAGA : 



♦* 



4 



its full sulphonation, was found to have only 53.3 per cei 
it as sulphonate and 44.7 per cent, of it as siilpliite, 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 sd 
portion having, in relation to its quantity, received su 
dioxide four times more rapidly than the larger portion hm 
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 sulj 
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 sulpliite 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 sulpliur di 
entering the solution during that time becoming sulpha 
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 experimeni 
the great variability of the point at which acid carbonate 
precipitated, as well as the variability of its quantity. 
quick working acid carbonate precipitated mueli earlier a 



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INTERACTION BETWEEN SULPHITES AND NITBITES. 



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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 experimeut 
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 



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E. DIVERS AND T. HAGA : 



is stopped, owing to sulphoiiation of the nitrite and reconversio 
of acid carbonate to normal carbonate — 

NaNOa + 2Na2S03 + SNaHCOa = Na2HNS207 + 3Na2C03+ HA 
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 mi 
be disregarded as a factor in the process of sulphonating wli( 
sulphur dioxide is also at work. Here we would insert that on. 
to simplify discussion do we speak of normal sulphite and carbc 
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 pyrosulphil 

That in the earlier stages of the process, when mu 
carbonate is present, the normal sulphite plays a very small pj 
in the sulphonation not only follows from the observation of 
rapid increase in quantity at first but is also shown by its tb 
nearly constant quantity for a long time though sulphur dioxi 
is still entering the solution and forming sulphonates. Oi 
later, as the carbonate gets consumed, does the sulphite beco: 
an important factor in the sulphonation by freely becoming pyi 
sulphite, for then its quantity rapidly falls. 

The part played by sulphite in the early stages being tl: 
insignificant, we have to seek in the carbonates the source of t 
early considerable sulphonation of the nitrite. It would be u 
reasonable to assume, with acid carbonate present, that the nom 
carbonate takes part in sulpnonation ; equally so to assume tl 
it remains inactive to sulphur dioxide. We are therefore coi 
pelled to recognise that sulphonation goes on only after conve 



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INTERACTION BETWEEN SULPHITES AND NITRITES. 



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tion of all carbonate locally present to acid carbonate and 
sulphite has been effected. Then the reaction that ensues is — 

NaNO^ + NaHCOa + 280., = Na2H NS-.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. 



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£. 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 1 
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 sulplii 
It does not however depend upon this, for while sulphur diox 
liberates a molecule of carbon dioxide in changing carbonate ii 
sulphite, four mols. of it are needed to liberate one moL 
carbon dioxide in changing carbonate and nitrite into hydr 
imidosulphate. 

An explanation is suggested by a consideration of the i 
that when working the process at a moderate rate, the I 
crystallisation of acid carbonate takes place long after the \k 
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 sulpl 
dioxide, does the acid carbonate continue to separate out in iiii 
such quantity as it could do at the stage of the process read 
The cause in one word is supersaturation. The acid carbou 
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 non 
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 



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INTERACTION BETWEEN SULPHITES AND NITRITES. 



303 



ite in the process, the amount of salt separating is small, and 
3nerally depends then for its existence upon its power to resist 
16 action of acid sulphite in ice-cold solutions. The solution 
hen, potentially at least, it is richest in acid carbonate, was 
lund 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. 

III.—/. 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 
fide, according as one or other of the interacting substances is 
excess. That is the ordinary well-known result, but there are 
ro ways of limiting the extent of the action so as to get either 
^droximidosulphate and nitrous acid or the undoubted products 
' their transformation. By these ways, the interaction of sul- 
lur dioxide and a nitrite is shown to be — 

2KNOo + 2SO2 + OH2 = K2HNS0O7 + HNO2. 

The more important way to thus limit the action is by an 
:periment first tried by Glaus {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 
itrito-hydroximidosulphate which precipitates and ethyl nitrite 
hich boils off* by the heat of the reaction : — 

3KNO2 + 2SO2 + C2H6O = KN02,K,H NS A + C2H5 NO2. 
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 



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304 



E. DIVERS AND T. HAGA 



upon a nitrite was found out by Raschig, when trying to pn 
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 rai 
potassium sulphate had crystallised out, potassium amidosulph 
was finally obtained, as proof that hydroximidosulphate 1 
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 (/. Oh. 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 hydroxyami 
sulphuric acid, but at the time we took it to be hydroxylan 
itself. We have also found that, after adding a dilute solul 
of sodium nitrite to excess of a cooled solution of sulphur Am 
and then blowing out of the solution the residual sulphur di 
ide by a current of air, enough hydroxyamidosulphate (hydrolj 
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 filte 
off) gives sulphurous acid when acidified (this Journal III, 21 

Though less successful than Clauses experiment, Rasch 
method is serviceable for showing that the alcohol used in tl 
plays only a secondary part. While excess of nitrite is suco 
fully used in that experiment, the sulphur dioxide must be 
excess in Raschig's method. To understand this, it has onlj 



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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 Add. 

Fremy believed 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 thai 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. /.) 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 
ise 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- 
)f 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 
irated on the water-bath and further concentrated in the 



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306 



E. DIVERS AND T. HAOA : 



II 



vacuum-desiccator, the amidosulphuric 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 hydrochloi 
acid, into acid ammonium sulphate ; we have also complete 
volatilised the acid by heat thus proving the absence of h 
accidentally derived. 

Nitrosyl sulphate dropped into much excess of cooled sol 
tion of sulphur dioxide also yields the hydroxyamidosulphate : 
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 sulj 
azotised acids could be obtained in this way, because of their 
ability to exist in absence of a base. Moreover, he considei 
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 hydroximidosulphates he could prepa 
indeed, were those of potassium, but from ammonium nitrite 
got the nitrilosulphate, and also obtained evidence that calcui 
strontium, and barium nitrites are convertible into amida 
sulphates. 

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 soi 
bases will doubtless be inferior to others, and those which 



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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, raercurosum, and silver, with the results 
low record. 

Ammonium salts. — 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 
tralent of ammonia-water which had just before been con- 
)d to sulphite by passing sulphur dioxide into it. More 
bur 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. 
3 of the solution was hydrolysed and tested then with copper 
late and potassium hydroxide ; it was thus shown to have 
iined abundance of ammonium hydroximidosulphate. 
ther portion of the solution not hydrolysed gave a large 
ipitation of dipotassium hydroximidosulphate on addition of 
mum chloride. 

Barium salts. — Some barium hydroxide was converted into 
lite 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. 



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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 mucli 
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 salts, — 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 wa?? 
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 wiis 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 hydroximidosulphate 
expected. 

Zinc salts, — Zinc nitrite solution was prepared by precipita- 



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INTERACTION J&ETWEfiN SULPHITES AND NITBITBS. 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. 
MercuroiLS salts and silver salts. — Experiments, already re- 
ferred to in sect. III. /. 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 
efiects the preservation of the products. It is not the salts which 
are sulphonated but nitrous acid itself. 

V. — What Nitrous Acid becomes wlien 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'SOsH or (HO)2NS03H, 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- 

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310 DIVERS & HAGA : INTERACTION BETW. SULPHITES & NITEITES. 

imidosulphate corresponding to the pyrosulphite acting— 
HONO + (S02K)-S03K=HON(S03K)2. The origin of all the other 
salts out of this salt has been traced, partly by others and part- 
ly by ourselves, and need not be gone over again here. 



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CONTENTS OF RECENT PARTS. 



Vol. X., Pt. 1 . . . yen 1.80 (Price in Tokyo). 

• Wmtm of tbe Blantopore, the Relatlonn of the Primitive Streak, and the 
PoriHatlon of the Posterior End of the Emhrjo In Clieioniii, toirether with 
BeBHurks on the Watare of aiertibiastlc Ovn In Vertebrntes. (Contributions 
o the Embryology of Reptilia. V.j. By K. Mitsckuri. {With Plates I-XI). 

Vol. X., Pt. 2 . . . yen 1.20 (Price in Tokyo). 
elno In HIsakI Torkommende Art von Epheiota iind ttber Ihre Sporen* 
^ndus. Von C. ISHIKAWA. iHiisrzii Tafdn XII und Xlfl). 

daa ■taaaenhafte Vorkommen von Elt»enbacterlen In den Thermen von 
Ikne. Von M. MrrosHi. 

n Aher die Schwefeiraaenblidnnir und die .Schnrefeibacterlen der Thermen 
ron Tnmoto bet Nlkkd. Von M. MiYOSHi. {Hierzu Tafel XIV), 
itwlckelnnc der Gonophoren bet Physnlla maxima. Von S. Goto. (Hierzfi 
Vaf. XV). 

m or Keprodnetlve Elements. III. Die Eiitwlckeliiniir der Pollenkdrner 
ron Alllnm fistuloanm Ij., ein Beitray stir €hromo«omenrednktl»n In 
PlinMMnrelehe. Von C. IsHlKAWA. Hierzu Tafeln XVI und XV 11), 
hotioBS to the Horpholon^y of €jrclOflt<»niatn. I. On the Formation of the 
BMurt In Petromyaon. By S. Hatta. ( With Plate XVIII). 

Vol. X., Pt. 3 . . . yen I.IO (Price in TokyG). 
etnanorphoflls or Asterlas pallida, with Special Reference to the Fate of 
IBO Body-cavltlen. By 8. Goifi. {With Plates XIX-XXIV). 



Vol.' XI., Pt. 1 . . . yeti 0.34 (Price in Tokyo). 
mUon Of Hyponltrite from Nitrite throui^h Oxamld<»9nlphonate. By E. 
>iyEB8 and T. Haoa. 
l>Ufl»M Of Nitric Oxide in Gas Analysis. By E. Drs'ERS. 

of Nitric Oxide with Silver Nitrate. By E. DiVEKS. 
of Pure Alkali Nitrites. By E. Divers. 

of an Alkali Nitrite by an Alkali Metal. By E. Drv'ERS. 
iltrltcs: their Properties and their Preparation by jiiociinm or Potassium. 
5y. E. Divers. 

Vol, XI., Pt. 2 . . . yen 0.38 (Price in Tokyo). 
9 G«olOiri« structure of the Malayan Archipolnffo. By B. Kotd. {With Plate J. 

Vol. XI„ Pt. 3 . . . yen 1.45 (Price in TOkyd). 
i»iitnl Pendulums for the Mechanical Registration of Seismic and Other 
Enrtii Movements. By F. Omori. (With Bates II-XTI). 

>m the Preliminary Tremor of Earthquake Motion. By F. Omori. {With 
PkUea Xni'XVI). 



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E«rtliqiiake Heiuiareiiient mt MIyako. By F. Omori and K. HiRATA. {With Pbte) 

XVIIXXIIT). 
Btlijl lunnionAaiiisalplilte. By £. DiTEBS and Ogawa. 
Ethjl lunmonlaiD selenlte and Non-existence of Anildo«elenlte« (Bclen— •!■■>—). 

By R D1VEB8 and T. Hapa- 
Votes on the Minerals of Japan. By K. JiNBO. 

Vol. XI., Pt. 4 . . . fjen 1.04 (Price in Tokyo). 
4^n tlie Hutaal Inllaenee between I^on^tadlnal and Clrenlar Mairnetlsntiont la 

Iron and Nlekel. By K. Honda. (With PtcUes XXIV & XXV), 
Tlie Earthquake InvestJffatlon Committee C^talofcne of Japanese Bartliqaskw- 

By 8. Sekiya. 
Motes on the Earthqnake Investii^Atlon Committee Cataloffoe of Japanese Ssrth- 

quakes. By F. Omori. {Wiih Plates XXVI XXVII). 



Vol. XII., Pt. 1 . . . //c?* 1.20 (Price in Tokyo). 
Japanisehe besehalte l*nlmonaten. Anat. Untersaeh. d. In Zool. Mnseam der 
k. Vnlv. In T6ky6 entiiaitcnen Materiales. Von A. Jaoobi. (HUnu Tafebi 
I- VI), 

Vol. XII., Pt. 2 • . . yen 0.50 (Price in Tokyo). 
£tndes sur la F^oondatlon et I'Embryofr^nie dn Ginkfvo blloba. Second n^moire 
Par Sakcgoro Hirase. {Avec PL VII-IX). 

Vol. XII., Pt. 3 . . . yefi 1.05 (Price in Tokyo). 

Vntersiichungren liber die Entwlcklnns der dteschlechtsoryane nnd den Torsanff 

der Befmchtnnfp bei Cyras revolnta. By S. Ikeko. {With Plcden X'XVII\ 
On a Collection of Batraehlans and Reptiles from Formosa and A<y a c eM t Islands. 

, By L. Stejjjeger. 
Some Points in the netamorphosis of Asterlna nrlbbosa. Bv S. OOTO. {WUk Ptde 
Ilf). 

Vol. XII., Pt. 4 . . . yen 1.86 (Price in Tokyo). 

Further Observations on the Nuclear Division of Jfoctiluca. By C ISHIKAWA* 

( With Plata XIX). 
Hotes on Some Exotic Species of Ecioparasitic Trematodes. By 8. Goto. {Wiik 

Plates XX & XXI). 
Tentamen Florje I«utchuensls. Sectio Prima. Plantie Dicotyledonett P<oljrpetal» 

By T. Ito and J. Matsumura. 



Vol. XIII., Pt. 1 . . . yen 1.65 (Price in Tokyo). 

Hotes on the Oeoloffy of the Dependent Isles of Taiwan. By B. Kot6. ( WUk Pfaief 

I-V). 
Chance of Volume and of I«enfrth in Iron, Steel, and Nickel Ovoids by Xairn^tlaatlsB. 

By H. Naqaoka and K. Honda. {With Plates F/d- VII). 
Oombined Efltect of I^ni^itudlnal and Circular Xaffuetisatlons on the IMaseMtioas of 

Iron, Steel and Nickel Tubes. By K. HoNDA. (WUh Plates VIII ^ IX). 



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fltndlen ttber die AnpnflsaarnnUilKkeU elnlf^er Infasorlen an concentrlrte I«dsan^n. 

Von A. Yasuda. [Hienu Tafd X-XIT). 
Veber cll« WaehatbaiiiBbeschlenDlffaDff elnlffer Allien nnd Pllae dnrch chemliMlie 

Keise. Von N. Ono. (Hiertu Tafd X-XIII). 



Vol. XIII, Pt. 2, published July 28th, 1900. 
Price in Tokyo, .... yen 1.00. 



All parts of this Journal are on sale at 

MARUYA & Co., 

TORI SANCHOME, NIHONBASHl, TOKYO. 

R. FRIEDLANDER & SOHN, 

CARLSTRASSE 11, BERLIN N, W. 



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CONTENTa 



Vol. xin., Pt. n. 



PAGE* 



Ammonium Amidosulpliite. By Edwabd Pivers and Masataka 

Ogawa, Imperial* University, Tokyo 187. 

Products of heating Ammonium Sulphites, Thiosulphate, and 
Trithionate. By Edward Diyers and Masataka Ogawa, 
Imperial University, Tokyo 201. 

Potassium Nitrito-hydrozimidosulphates and the Non- 
existence of Bihydrozylamine Derivatives, By Edward 
Divers, M. D., D. Sc., F, R. S., Emeritus Prof,, and Tame- 
MASA, Haga, D. Sa, F. C. S., Professor, Tokyo Imperial 
University 211. 

Identification and Constitution of Fremy's Sulphazotized 
Salts of Potassium, his Sulphazate, Sulphazite, etc. 
By Edward Divers, M. D., D. So., F. R. S., Emeritus Prof., 
and Tamemasa Haga, D. Sc., F. C- S., Professor, Toky5 
Imperial University 225. 

On a Specimen of a Gigantic Hydroid, Branchiocerianthus 
imperator (AUfnan), found in the Sagami Sea. By 
M. MiYAJi3iA, Eigakushi. Science College, Imperial University, 
T5k}^o. (With Bates XIV dk XF) 235. 

Mutual Relations between Torsion and Magnetization in 
Iron and Wickel Wires. By IL Nagaoka, Eigakuhakushi, 
Professor of Applied Mathematics ; and K. Honda, Eigakushiy 
Post-graduate in Physics. {With Plates XVI) 263. 

The Interaction between Sulphites and Nitrites. By Edward 
Divers, M. D., D. Sc., F. R. S., Emeritus Prof., and Tame3IASa, 
Haga, D. Sc., F. C, S., Professor, Takyo Imperial University. 281. 



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* ^ P a :fc * «B # 

m -^ ^^ 1st m ^ ft 
THE 

JOURNAL 

OF THE 

COLLEGE OE SCIENCE, 

IMPERIAL UNIVERSITY OF TOKYO. 

VOL. Xlir., PART TIT. 



M M 'tf m ic ^ ni Vr 

PUBLISHED BY THp] UNIVERSriT. 

TOKYO, JAPAN. 

1900. 
MEFJi xxxni. 



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Prof. K. Yamagawa, Ph. B., Rigdkuhakushi, Director of the OoUege 

(ex officio). 
Pro£ J. Sakuraif RigahuhakusJu. 
Pro£ B. Kotd, Ph. D., Rigakahakuihi. 
Pro£ I, Ijimay Ph. D., Rigahuhakushi. 



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itributions to the Morphology of Cyclostomata. 

II.— The Development of Pronephros and 
Segmental Duct in Petromyzon.^^ 

By 
S. Hatta, 

Professor in the College of Peers, Tokyo. 



With Plates XVILXXL 



rhe 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). 

)ur 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 MuLLER (75), Scott ('82), Shipley ('87), Goette 
KuPFFER ('90), and others, ten years or more have 

t was my intention to publish this paper shortly after the appearance of ray 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 
0U8 points and tlie results here given are different from those of the preliminary 
1 several important respects. 



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312 



S. HATTA : 



1 



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, oi 
merically arranged, organ-system such as the neural c 
chorda, the pronephros, &c., is rendered peculiarly 
in Petromyzon by the fact that the longitudinal axi 
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 tliis axis is cu 
obliquely, as, for instance, the neural cord shown in fij 
3, PI. 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 toto in this fluid, decoloriz 
proper degree in acid-alcohol, and then washed in 90^ 
In the sections of specimens thus prepared, the histologi 



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MORPHOLOGY OP CY0iX)STOMATA. 



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 
lers, Pkof. Mitsukuri and Prof. Ijima, for much in- 
ible advice and for their kindness in looking through . the 
iscripts 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 1 have to deal 
in the present investigation, is only a little advanced beyond 
llipsoidal gastrula; it is intermediate between Stage I and 
f the list given in my first contribution above referred to 
1, A andB, of that article). The head-fold forms a pointed 
iberance at one pole of the ellipsoid, while the blastopore 
ainly visible at the opposite pole. The prominent neural 
I 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 

8. Hatta, On the form, of the germ. lay. in Petr.: this Journal, vol. V, 1891. 



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314 8. HATTA : 

neural cord is interposed in the median line between the right and 
left halves of the mesoblast, uoderneath 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 cells. 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 ii {loc. city 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, viz. 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. 



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MOBPHOLOQY 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 
3 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 
lose group of somewhat irregularly shaped cells {Im.) 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 Amphioxus. 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 
die of the proximal segmented portion (x). The parietal layer 
il to this notch is composed of a regular cylindrical epithelium 
n.2)i which is slightly arched against the epiblast, so as 
;ause an indentation in the latter, while the visceral layer 
le corresponding portion consists of a more or less disturbed 

of high columnar cells. As the subsequent history teaches, 
proximal half of this extent {rut. V) represents the myotume^^ 



)The somites are reckoned from the anterior end. The first, Le, the foremost lies 

diately behind the auditory vescicle when the vescicle comes into view. 

l)The vertical diameter of the neural cord in tigs. 2 and 3 is shown greater than it 

7 iij, the sections passing obliquely owing to the bending of the longitudinal axis of 

mbryo, as noted in the introduction (j). 312). 

t)This term here means the Sclero-rayotom of German authors. 



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316 S. 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, 1 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, i.e. the sixth (fig. 5), the 
mesoblast shows almost the same condition as that already des- 
cribed ; but in the somite preceding the fifth, i.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 {mt.IV)j while, on the 
right, it is cut through in the middle {mt.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 {mt.IV) and consists of high cylindrical cells 
from the lateral plate formed of a loose mass of cells {Im.). In 
the left half, however, the state of things is quite otherwise: 
the Anlage of the pronephros {a.pn. l) together with the corres- 
ponding visceral layer is entirely constricted off* from the myotome 
{mt.IV) J although it is still connected with the lateral plate {Im.). 
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. 



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MORPHOLOGY OF CYCLOSTOMATA. 317 

j 

Fig. 4 represents a section passing between the two somites 
e mentioned (the fourth and fifth) and is much magnified 
IS, Ey^ 2) to illustrate the finer structure of this portion. The 
jtural cells are all loaded with an enormous quantity of 

I 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 

{ml.V) is the anterior wall of the fifth myotome. Two 

of variously shaped cells {Im.) constitute the lateral plate i; 

ih is histologically quite like that in the somitic portion, being f 

posed of irregularly quadratic cells and tapering towards j 

distal (ventral) extremity (compare with the lateral plate, i 

in figs. 2 and 3). However, in the proximal portion, 
e the Anlage of the pronephros consisting of a regular row 

II columnar cells would be found in the somitic portion, we 
lere a group {x) of a few cells of faint appearance, forming 
proximal edge of the lateral plate. By a comparative study 
iro 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 
ler, I have drawn fig. 7 which represents a section through 
ntersomitic plane between the sixth (6g. 5) and the seventh 
te (fig. 6). In this part the Anlage of the pronephros 
iveloped still more weakly, and the mesoblast remains in a 
I primitive state. In the proximal edge of the lateral plate 
10 special structure is detected, but the edge fades away without 
jtinct 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. 
Froui the fifth somite backwards for 9 or 10 somites, the 



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i 



318 



8. HATTA 



niefiobkBt presents almost the same feature of the Ai 
the iifth somite mentioned above. Fig. 5 represent 
through the sixth somite, next behind thc^ fifth ; when 
with (ig. 2 no marked dilTerence is detected in regi 
stractiire of the niL'Soblagt. But in some segments the d 
of the Anhige ir BOmewhtit weaker than in othera, 
fig, 6, which shows a section throngh the seventh sor] 
in a segment posterior to this somile, we find the Anlaj 
pronounced as in the sixth somite. However^ generally 
the Anlage of the pronephros in an anterior sorail 
furtlier than that in a posterior. It must be remem 
the somite in which the Anlage has already becom< 
does not pass over suddenly into the somite in whic 
of it is to ho seen ; bet its development gradually gro^i 
less distinct from ihe anterior to the posterior part, u 
no trace of it is percoived* 

In the preMni d^tgi, therefnre, ihe Anhge of the 
is detected in more than 4 BomUes !nd u mmplelehj s^ 
the myotome Q?ili/ in one seymenl, viz, the fourth somil 
hai no genetic mnnealion either with the Anla^e % 
following somite or with the rpihhst ; tmd it mud he n 
we find theforemod Anloge not exaelb/ beneath the faurl 
but ahmys tmidernealh its kind Im'der, 

Figfi. 8-17 roprf^Rcnt flections throng! i n Ftill 
hryo of this stage, having al>ont 20 Romites. The cpi 
the neunil cord {n.)^ and tlir. chorda [ch.) are essentiall 
as hefure. Being cut through somewdiat obliquely, the 

l)8tjcli a case is very rure. In mtjst ^iJc^^iiJiery exjuulH<?d, Ibe Aukge 
till? ttiynt4ime is fonnd m mriny sei^maiH ►*'► il*ii* wo can hnony dtvidft m 
the jnei^jimticm takes ^jLice lirsL 



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MORPHOLOGY OF CYCLOSTOMATA. 



319 



be two sides do not exactly correspond. On the right side 
;. 8, the hind border of the fourth myotome {mt.IV) is 
ihrough ; the Anlsge of the pronephros (a.pn.l) presents in 
}n 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- 
Iso distended. The lateral plate (Im.) shows, however, no 
[ed progress. The left side of this figure and the right 
^.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 {mt.VT) 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 
rved 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 {mLVT)^ and the two layers 
lis 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 tlie myotome, their structure is at 
Kisenecl, that is, tlieir component cells become loosely s-et together. Later the cells 
)ly tliemselves, and are again compressed by mutual pressure ; giving a compact structure to 
nlage — probably the same condition observed by van Wyhe in Selachian embryos 
►. 476). 



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320 S. HATTA : 

section through the anterior border of the somite. This phase 
of constriction is donbtless 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, i.e. the seventh. 
The Anlage of the pronephros {a.pn.4) is not yet cut off from the 
myotome {mt. Vlt)^ but the process is beginning as shown by a slight 
constriction and an inclination to arch out, while the suddenly 
weakened lateral plate {Ivi.) 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 facU mentioned ahovej it is easily understood that 
the separation of an Anlage from a myotome 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 offy so that the cavity of the myotome (myocoelome) com- 
municates with the peritoneal cavity ^ during some timCf 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 



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MORPHOLOGY OF CYCLOSTOMATA. 



321 



Ugh the twelfth somite ; we can find no marked difference 
reeu the Anlage in the seventh somite and in this. The 
lents lying still farther backwards are not cut through exact- 
ransversely in this same series of sections, owing to the cause 
id above (pp. 312 and 315), so that we can not trace the dif- 
Qtiation 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, i.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 {Im.) 
ents no longer a weak appearance as in younger embryos 

the statement on p. 317 and figfe. 4 and 7), but is occupied 
31 compact cellular structure (cd.) which suddenly passes over 

the loosely composed lateral plate {Im.). 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- 
)n 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 
bis cord which gives rise to the collecting duct or Sammelrohr 
LucKERT ('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 






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322 s. hatta: 

the structure (cd.) at the proximal portion of the lateral plate is 
almost the same as that on the 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 cells only. 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. In fad, the cord appears after the complete 
separation of the Anlage from the myotome, and when it is first 
established, the nearer the plane of a section to the Anlage either 
anterior or posteiHor, 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 



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MORPHOLOGY OF CYCLOSTOMATA. 



323 



part where the duct has been described above as weakest, 
s also the fact that repeated cell-multiplication takes place 
ihe outer rim of each Anlage of the pronephros. One might 
pose that the product of the cell-division would contribute 
J to the growth of the Anlage itself and has nothing to do 
1 the cord ; but this is not the case : the Anlage does not 
N at the outer (lateral) end, as it might seem, but by cell- 
sion within its own structure. 1 have never observed any 
I 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, Avhen they 
fully established (see the right side of figs. 9 and 10) ; this 
[lection thus is not primary, but secondary. The epiblast has, 
a the first, no share in the formation of the cord, always 
^ing a sharp contour against the mesoblast below. 

Thei^e is tkus no difficulty in accepting the view that the 
lecting cord is formed of the intersomitic cell-outgroioths which 
budded out of the anterior and posterior rims of each Anlage 
he pronephros and are subsequently fused together. The cord 
IhereforCj originally brought about by the conjiuence of the free 
entities 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 ii. The epiblast {e]^) 
aists 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 {mt.IV) is cut, while on the right, the 
-plane of the fifth myotonic is met with. A comparison with 
corresponding parts in the younger stages (ligs. 3 and 8) will 



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324 s. 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.i), 
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 visceral layer of the lateral plate (Im.). 

In a little more advanced embryo, the cross-sections of which 
are represented in figs. 20-31, the neural cord (n.), the myotomes 
(mL), 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 to the same cause as the sections represented in Figs. 2 and 3, tlie vertical 
diameter of the neural cord in Figs. 20-23 is shown somewhat longer than it is in reality. 



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MORPHOLOGY OF CYCLOSTOMATA. 



325 



loosening of the composition of the tissue^^ and assume the 
€ of a scalene triangle (figs. 20-23) ; the median side of the 
igle {vim.) 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.VILX). 

The anteriormost Anlage of the pronephros is found as before 
jr the hind part of the fourth myotome, the section of which 
epresented in fig. 20 {apn.l). 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 
med. 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- 
es 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.pu.3) together with the hind border 
he myotome {mt.Vt). 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 
3ath 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. 



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326 S. 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, cd.)^ 
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 



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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- 
)r or the posterior) is the further developed ; a comparative 
y of other examples shows that the separation in the posterior 
[lent follows that in the anterior. The state of the mesoblast in 
next posterior segment, i.e., the tenth segment (fig. 31), is quite 
irent from that just described ; it is in a more primitive condition 
svelopment. 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, 
ind 16). From this segment backwards, a few segments show 
3st the same condition. Still further posteriorly, the structure 
he mesoblast can not be readily observed, since the planes of 
ions 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 {Im.) 
jists 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 J therefore, the Anlage of the pronephros is com- 
ply separated from the myotome in 4 somites^ i.e., from the 
th to the seventh inclusive ; a,nd these are connected tvith 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 
? posterior somites it is indicated merely by a slight depression 
he parietal layer of the mesoblaM. 



Period 2. 



In the embryos which belong to Stage iit, we observe a 
ded advance in several respects. Figs. 32-50 represent a series 



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328 S. HATTA : 

of cross-sections through one of these embryos whicli 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 {mvs.) 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 (Zm.). 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 celW* 

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.l). 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 fewi myotomes in the anterior somites tend to assame this shape already in the last 
stage (see figs. 20, 21, and 22). 

2) See the foo^note on p. 319. 



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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 
t (Fig. 32, a.pn.i) projecting anteriorly and lying between the 
lal edge of the lateral plate {Im.) and the lower surface of 

fourth myotome {mt.IV). The existence of this conical 
'}^ gives us a strong impression that originally there must have 
I present an Anlage of the pronephros in the anterior segment 
sh was connected by a connecting cord with the Anlage 
nging to the fourth somite, but had disappeared duriug the 
logeny and that this conical tube is the remnant of this 
aecting cord.^^ 

The next posterior Anlage, which is found under the fifth 
itome (figs. 37-39, a,2)n.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, 
1.3) has an outline much like that shown on the left side of 
3. 18 and 24, being in the same phase of development, that is, 
3 of the form of an isosceles triangle whose two basal angles 
jh the myotomes. This Anlage is found under the sixth 

L) The internal cavity of tliis conical tube is not entirely closed, but there is clearly seen 
all canal (z) 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 
ling ; I also can not detect any trace of such a canal in embryos of advanced or younger 
s. 
2) See the description under Period. 4. 



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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) lias 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.d 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, i.e. between the iVnlagen 
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 



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MORPHOLOGY OF CYCLOSTOMATA. 



331 



Anlagen fourth and fifth, and in fig. 49, one between the 
jgen fifth and sixth. 

This inequality in the length of the intersoraitic solid cord 
believe, due to differences in the degree to which th§ canal i- 
m within the Anlage has extended into the connecting cord. 
he anterior section of the pronephros, this process has already 
ceded to some extent into the interior of this cord, while in 

posterior, the cavity is still confined entirely within the 
ige itself. The whole system of the pronephros at the present 
lition may be compared to a bamboo-cane with nodes and 
modes ; 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 
1 entirely disappear later when the collecting duct is fully 
blished. 

From the fact mentioned above, it will be easily seen that the 
less of canalization in the pronephric system of Fetromyzori 
ns in the internal cavity of the pronephric Anlage in each 
lent 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 
Q08t cases, not only the internal cavity in each Anlage is 
lious anteriorly and narrowed posteriorly, but the cavity in 
irior 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 tlie 
inatioD of the planes of sections, referred to above. 



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332 S. 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, tlien, the Anlage of tlie 
pronephros is cut off from the myotome in more than JO segmenU^ 
and the canalization has advanced in the anterior section of the 
system, to a state just ready to put the Anlagen in the succeeding 
somites in communication with one another, although the inter- 
Semitic connecting duct in the posterior part remains still solid. 

Figs. 0I-08 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, u tubular structure and have grown dorsally, beiug 
folded out from the body-cavity ; I will accordingly call them the 
pronephric tubules. 

On the right side of fig. 51, the foremost tubule {pLl) 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 {pt.2) and that on the 
left side of the third section posterior to it (fig. 53, pt.2) are 
respectively the second tubule of the right and left side found 



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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 (cd.) on the right side and 
a slice of the hind wall of the second tubule on the left lpt.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, 
pL2) 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. If the 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 aeries of sections, from which fig. 54 is drawn, I observe mitotic figures at 
that point in several sections. 



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334 



S. HATTA : 



I 



showing no structural alteration. Mitotic figures are r 
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 i 
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.l and 
pL2)j 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 ii 
{pL2). 

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 (ptA) and, on the left, 1 
tubule {pL3) respectively. The latter is not so far d 
as its counterpart on the opposite side (fig. 53, pt3), ^ 
former presents a great progress : it consists of a definite 
ium and contains a distinct cavity of triangular shape, i 
the corresponding tubule on the opposite side (fig. I 
which is found in the third section behind the last, is n 
advanced in development. The fifth tubule, the tubuh 
right side of fig, 56 {pL6), is somewhat more developed t 
which belongs to the anterior somite (the fourth tubuh 
opposite side) ; but it has a feature much resembling th 
tubule on the same side (fig. 55, pL4) and the second 



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MORPHOLOGY OF CYCJLOSTOMATA. 



335 



M)site side (fig, 53, pL2). In short, in this series of sections, 
1 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 
f interior. 

From the tenth somite backwards, the Anlagen are cut oflf 
m both the myotomes and the lateral plate, and constitute the 
mental duct or the posterior continuation of the collecting 
3t, which is distinctly traceable for 7-8 somites. Not infre- 
jntly, however, a somite is met with, in which the segmental 
it 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 
tious, 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 
dty is not so simple as in the last specimen ; besides the 
)nephic tubule, there is seen another structure which projects 
: of the inner angle of the peritoneal cavity (figs. 52, 53, 55, 
i 56, c,p.). This projection is originally a fold of the peritoneal 
U 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,^^ 



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336 S. 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, c,p.)y 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, nccording 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 furtlier discuss this structure, as I intend to do so in a future paper 
in which the development of the mesonephros in Pdramyznn will be delt with. 



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MORPHOLOGY OF CYCLOSTOMATA. 



337 



rithelial structure, forming the continuation of the peritoneum 
id folding out from the peritoneal cavity. Beneath the first 
ibule, 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, arid 56). 

The coelomic projections are not confined to the anterior region 
liere 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 
lelomic projection (cp.) 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- 
jphric tubule and will not partake in the formation of the perito- 
jal membrane. I was at first of this opinion, but a careful 
)8ervation of sections through the embryos in each stage showed 
y errpr. 

To illustrate this point satisfactorily, I have given, in the 
inexed wood-cut (Wood-cut 1), a series of semi-diagramatic 
jures, 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 ; 



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338 



8. HATTA : 



a-b indicates the extent of the 
coelomic projection. When the 




A. 

B. 

a 

D. 
R 
F. 

o. 



WtKxl-cut 1. — Semidiagramatic figures to 
illustrate the succeaiive 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. 5o. 
from the left side of fig. 61. 



called the coelomic projection 
plate of epithelial cells {G^ c.p. 



Anlage ; c^d shows that of the 
myotome is cut off, the point 
of the parietal layer indicated 
by a becomes fused with the 
point a of the visceral layer 
(-B, a^). This piece of the meso- 
blast assumes an ellipsoidal shape 
(C). The component cells of 
this elli{)8oid are multiplied by 
active cell-divisions, and the 
piece almost loses its lumen and 
gets a compact consistence {B). 
Meanwhile the cells in the space 
d'C remain inactive. Conse- 
quently the piece acquires a tri- 
angular form (E), whose upi)er 
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 {F) and produces 
a small knob (jP, 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 
) and assumes then the form of 



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MOEPHOLOGY 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 (G). 

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 ui), of the inner and outer layers which constitute 
resjiectively the Muskelblatt and the Outisblalt of German 
authors (fig. 59 and 60, mu.^, and cut). The cells composing the 
3IuskelblaU {fnus.) are, simply differentiated into a transverse row 
of the muscle-plates. The outer layer (ctit.) undergoes, however, 
subsequently a series of interesting changes : it folds in, just as 
the Sklerablatl or sclerotome described by Hatchek in Amphioxm 
('88) between the Mutikelblail and the chorda and the neural tube.^^ 
As is well known, Rabl ('88) has homologised Hatchek's Sklerab- 
latl with his Sclerotomdiverlikel 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 Ixj treated of in an independent article. 

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340 S. 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 PetromyzoUy which gives rise to the 
pronephic Anlage and the coelomic projection, is doubtless homo- 
logous with the " inlermediale cell-mass '' of Balfouk described by 
him in Selachia and, therefore exactly coincides with the ** Nephro- 
torn " of RucKERT.^^ 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 [ctU.) is still composed of short cubical cells. In the 
anterior region, the true coelome {ppx), becomes conspicuous en- 
closed by the parietal {in.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 form 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 ypite of tlie discussion by RCckert ('8U, pp. lt)-20) on the inexactness of thceipres- 
Mon "intermediate cell-mass," I homologise, with many authors, these two terms with each other. 



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MOKPHOLOGY OF CYCXOSTOMATA. 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.l) 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. 66 representing 
the next posterior section will make the matter clear. On the 
left side of fig. 65, the same remnant structure {pLl) together with 
the collecting duct (ctZ.), 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 {n8t2) 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. 



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342 8. HATTA : 

67 {pL2). The shape, which the tubules of the second pair (fig. 66, 
pL2) 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. G6, j9i.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 {c(L) 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 {pL3) to be of the same structure. 
In these two sections the tubules are cut through lengthwise, and 
the nephrostoraes (nsLS) on the two sides come into view in sym- 
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 
tlirongh the intersomitic plane; but this Ls the piece of it belonging to either the anterior or 
the posterior somite. 



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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 {pt6) ; the condition of the tubules 
and nephrostomes {nst.6) 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 {nst.6) are visible on 
the right side, while the collecting duct {cd.) 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 

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344 S. HATTA : 

region, the neural cord (n.) is still solid ; the mesoblast {7m.) is 
many-cell-layered and its metameric segmentation is still going on. 
On the right side of the figure, the 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 {vit. and Im.) 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 maxhnurn number far Petroinyzon ; this stage ought^ there- 
fai^e^ to be regarded as the highest point of development with reference 
to the pronephros. Even in the jy)'esent stage ^ the foremost tubules 
show a tendency to degenerate. 

Period 3. 

The embryos of Stage iv, 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 {7n.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 



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MORPHOLOGY OP 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 (tnch.) 
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 {pt.2)y together with the 
corresponding nephrostome (nsL'J), 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 {n8L2) 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, mch.) 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 

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346 S. HATTA : 

in other places, for instance, in the space between the lateral 
plate and the epiblast (figs. 79-81, mch.)j 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 {pL2) 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 {pL2) 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 {nsL3) of each side, their respective posterior half being 
found on the left side of fig. 80 and on the right side of fig. 
82 {pt.3 and nstM) ; 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 {pL3) 



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MORPHOLOGY OF CYCLOSTOM ATA . 



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. 
m compared with the tubule of the same pair on the opposite 

represented in figs. 81 and 82 {pL3)y this character of the 
ie will be understood more clearly : the dorsal expansion is seen 
5. 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 
ad and third tubules is found. The left tubule of the fourth 
is shown on the left side of figs. 81 and 82 {pt.4) ; fig. 83 
^s a cross-section through the collecting duct (cd.) between the 
1 and fourth tubules and a slice of the anterior wall of the 
t tubule {pt.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 {pt.6) and on the right 
g. 85 on the other (pL6). It is not developed as much as 
more anterior pairs, but shows considerable prepress as com- 
d with the tubules in fig. 73 which represents the younger 
5 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 
i myotome {niLX). On the right side of the figure, however. 



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I I 



348 8. HATTA : 

besides the cross-section of a duct (cd.)^ there is seen a pronephric 
tubule {pl.6)y 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 of the 
sixth tubule which is in a stage of degeneration, and the duct (cd.) is 
doubtless the collecting duct between the 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 diflference of character between these two 
ducts is noticeable for some time in the younger stages. 

2o sum up tlie results obtained in this stage the tubules of the 
third to the jtfth pairs are vigorously developed^ while the second 
is very weak, the sixth retrograding, and the first ha^ entirely 
disappeared. 

In the present stage, a peculiar structure is observed inside 
the walls of the body-cavity (figs. 77-85, pp.l-d). 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 parti" 
tumJ*^ 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 



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MOEPHOLOGY OP CYCLOSTOMATA. 349 

and cut off a long chamber from the body-cavity along the 
openings of the nephrostomes (figs. 77 -S5y pp. l). 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- 
wdnde '* or ** Peritonealbrucke " described by Goette in Petromyzon 
fluviutilis ('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 {mLII-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 



l)See the historical review under Peironyzon, 

2) The embryo, from which these figures are drawn, is a little younger than that just 
spoken of. 



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350 S. HATTA : 

its caudal continuation {3d.) 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 (fff.) and the com- 
mencement of the raid-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 {ryiLI'l") are noticed ; the cell-maes {au.) seen next anteriorly 
to the first myotome {mLi) is a slice of the wall of the left auditory 
pit. The cross-sections of the pronephric tubules from the second 
to the fourth {pL2'4) follow immediately behind the fifth 
myotome; an oblique section of the fifth tubule (pLd) and the 
nephrostomal part of the sixth tubule (pl-O) are also obvious 
behind the fourth tubule. The nephrostomes of the second 
{n8t.2) and fifth (mLd) 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, nsL3, 6j 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,;?/.^ and nsLG), 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.)y which 



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MORPHOLOGY OF CYCLOSTOMATA. 



351 



Qstitutes the earliest fundament of the heart, is detected. The 
3Sent position of the pronephros y — dorsal to the hearty anterior 
d dorsal to the liver j and along eithe)' 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 
3ule (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 
itrary, the tubule of the next pair (fig. 94, pt.3) has much 
ngated and is bent considerably in dorso-lateral direction, 
that we can no longer observe the nephrostome together with 
i tubule itself on the same section. The following tubule, 
3 fourth (fig. 95, pt.4)y is likewise well developed ; the fifth 
;. 96, pt.S) 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 
jsenterial 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 
B tubules beneath the ninth myotome, where the tubules of the 
Lth pair ought to be found, but only the cross-sections of the 
llecting duct or the anteriormost part of the segmental duct 
3 seen. 



r 



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352 s. hatta: 

Thv^y the tubule!< of the thirds fourthy and' fifth pairs con* 
tinue to growy while the fint pah JiaA disappeared in the early 
part of the present stage {or at the end of the foregoing stage) ; 
the sixth has already commenced retrogression and the second is 
also gromng tveaker ami 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. xviii, 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, F). 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 a 
slender tube {fg.) which is bounded by almost a single layer of 
high cylindrical cells. The parietal layer of the peritoneum as 



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MORPHOLOGY OF CYCLOSTOMATA. 353 

well as the coelomic projectiou {rjfi.) 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 {h. and tr.a.) 
which are suspended by the dorsal and the ventral raensenteries, 
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 {tr,a.) found in the space between this meeting 
point and the ventral wall of the enteric canal, are destined to 

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354 8. HATTA: 

form the anterior continuation of the cardiac tube or the trunciis 
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 >?hich I 
could not at firet 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 {pL2) 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, Ac.). 
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 tuboles 
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 



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MORPHOLCKiY OF CYCLOSTOMATA. 



355 



toneum and the collecting duct ; it is the posterior wall of the 
lie in figs. 99. Fig. 101 represents the section through the axial 
le of the third tubule, the nephrostomes of which are recog- 
d more clearly in the section behind it (fig. 102, nsLS). The 
lies of this pair are comparatively not long. The fourth pair 
he tubules and their nephrostomes are obvious in fig. 104 
4 and nstA) which represents the third section behind that of 
102 ; the tubules much resemble those of the pair in front, 
nug 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 
a 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 
m 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, bs.). This fold is traceable from the anterior part 
le third tubule to the hind part of the fourth (figs. 100-104, bs.). 

space enclosed in this fold communicates freely Avith both the 
t of the dorsal aorta under the chorda and the tract of the 
rior cardinal vein outside of the pronephros and contains a 
iber of mesenchymatous cells which probably wander in from 
tract of the aorta and the anterior cardinal vein. As sub- 
lent history shows, this structure constitutes the beginning of 
glomervltis of the pronephroi^. 

Figs. lOo and 106 represent two contiguous sections immedi- 
j l>osterior to the section shown in lig. 104. In fig. 105 we 



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356 s. hatta: 

observe on either side the cross-section of the collecting duct 
{cd.) together with a part of the fourth tubule {pt.4) ; the longi- 
tudinal section of the fifth tubule {pL6) is seen on the right 
side of iig. 106, standing in wide communication {nst.3) 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 Q. 

In the Stage vi, 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 {fy.) 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 arteriomh 
in which its component cells are of cylindrical or cubical shs^ 



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MORPHOLOGY OF CYCLOSTOMATA. 



357 



J peritoneal raembrane lining the enteric canal immediately 
ind the branchial region is also thicker as compared with 
3r parts (fig. 107), being composed of a single layer of cubical 
B, — a peculiarity observed since the last stage (compare figs. 
d9 with fig. 107). 

The pronephric tubules as well as the collecting duct are 
iposed of a regular epithelium of cylindrical cells ; the former, 
•cover, are much prolonged and, in some parts (fig. 108), much 
ed, so that the peritoneal cavity which was almost a hollow 
30 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 {pt.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 an tero- posterior direction, but are curved laterally and ven- 
ly. The component cells are, in the nephrostomal portion, 
Br 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 {pt.4). 



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358 S. HATTA : 

A pair of the glomeruli (figs. 108 and 109,^/.). 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 been observed already from the forcing 
stage. 

The section represented in fig. 110 fortunately passes sym- 
metrically through a pair of the nephrostomes (nst.S) and of the 
tubules {pt.6) 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. 

T7i^ pronephric tubules in the present stage are, therefore, re- 
duced into the minitnum number, i.e., three pair s^^, all of which are 
retained so long as the organ functions as the excretory apparatus 



l)See p. 355. 

2) We occasionally find the four tubules to persist, and the additional tubule is the sixth. 



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MOBPHOLOGY OP 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 ivith the hind border of the hind wall of the branchial 
chamber whei^e^ in the fcyregoing stage^ the second pair of the tubules 
was founds 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 bMongedy 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- 
mnfzon. 

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 
nephrostonial 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 F). 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 

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360 



S. HATTA 



B 



(nephrostomal) portion of the two following pairs is directed 
backwards. The nephrostoraes retain their first position (B). 

Now the secondary carvatures take 
place ((7). The nephrostomal part 
of the foremost pair is crooked just 
like that of the two hind pairs in 
B ; the middle tubule is bent for- 
wards like the foremost tubule. 
The hindmost tubule makes a 
small forward curvature and a 
large backward bending. In the 
next stage, Z>, the foremost and 
the middle undergo no marked 
change, but the secondary curva- 
tures of the hindmost tubule are 
much more strongly expressed. In 
U 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 
increased degree of the original 
curvatures. It seems that the 
subsequent bondings 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 




Wood-cut 2. — Diagrams 
showing the convolutions 

of the tubules in later stngefl. 
L pronephric tubules. 

»f. segmental duct. 



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MOBPHOLOGfY OF CYCLOSTOMATA. 361 

are caused, therefore, by the 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 Genitcd Cells. 

For the sake of simplicity, the development of the segmental 
duct and of the vpscular system in the pronephros hag been 
entirely put aside in the description given above. 

As already alluded to, the origin of the segmental duct in 
Fetromyzan 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 rell-mass 
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 tha continuity 
with the lateral plate. If fig. 31, which represents the section 
through the tenth somite {ie. 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), is observed for some segments further backwards 
(see fig. 17). 

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362 s. hatta: 

The Anlage thus pronounced in each somite soon assumes a 
characteristic oval form, being completely cut oflF 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 formed 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 



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MORPHOLOGY OP 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 ofl* 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 intei'somitic 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 in, 
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 in) 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 
diflferentiation of the cells in dtuy 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. lU, a.sd.) and becomes ar- 

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364 8. hatta: 

ranged as in the Anlagen in the anterior region. Fig, 89 represente 
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 nephrotoraic cells in sUu 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 epiblast itself, as may be inferred from the direction of 



1) By the bending of the body-axis, some sections in a beries of crosB-sections are un- 
avoidably cut through fronlally. 



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MOBPHOLOGY OP CYCLOSTOMATA. 365 

the spindles, the long axes of which are directed always parallel 
to the surface of the layer. / have nowhere observed any trace of 
eUher 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.yK Fig. 
91 represents the next ventral section which passes through the 
dorsal part of the blastopore {bp.). As seen in these two sections, 
immediately inside of the blatlopore {bp.), 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.)y 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 dxLct with the cloacal 
cavity is founds therefoi^Cj at the point where the epiblastic layer 
of the lip is reflected inside and passes over into the hypoblast. 
This point of communication is, however, shifted far inside and 



l)Thi8 opening is found in the same vertical plane as the 34th or 36th somite. 
2) The right diverticnlara only is seen in 6gs. 90 and 91, the left one being observed in 
another section which is nnfigurcd. 



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II 



sm 



^. HATTA ; 



dorsally when the development proceeds further (tig. 1] 
sod cdv,), 

I have also met with two cases (figs. W and 111), i 
I have observed some epiblastic cells of the external 
walls of the blastopore multiplyiDg actively and having 
spindles {x) with axes directed perpendicnlarly to the plar 
epiblast^ while the duct comes in firm connection w 
point of the epi blast, — the connection is so firm that 1 
and the epihlast appear to iorm one and the same tis 
this point, thus, there is ev^ery appearance of epiblai 
partaking in the construction of the segmental duct* 

Tlie collecting duet peitaining to the nhith somii 
the segmental duct in that segment, having lost the eo 
with the tubule. 

Up to Stage II, the duct is represented by the si 
Anlagen in about 8 segments back of the ninth somite ; 
iiii these Anlagen are converted into the duct in about 
terior segments ; while in the course of Stage iv it o 
into the cloaca! cavity. 

From the above account, it is easily conceivable that 
iffffe of ihr i^egmenifd ifuH nnd Ihaf of the pronepkrie In 
prrfeei/}/ homohgow^^ mvl thai the duct is a eontinualion oj 
of abortive prontphrir tt!hulei< in the hind region. 




Underneath ten and more myotomes lying posterior 

the fifteenth somite the proximal portion of the laten 
which corrGspondn to the nephrotome, contains peculi 
cells (figs. 87 J 8S, and 89, gc\) loaded with an enormous qu 
yolk-granules ; the other mesoblastic cells in this pai 
much flattened out, form a thin layer over these cells 



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MORPHOLOGY OP 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 iir, these cells can not be distinguished from 
3r mesoblastic cells which are equally rich in yolk-granules. 
Stage iv, they become conspicuous ; and in Stage v, again 
Istinguishable 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 iv,^^ in three 
ts, viz., beneath the chorda, beneath the ventral wall of the 
iric 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 iv, 

cells below the hind section of the fore-gut are converted into 

endothelium of the heart and of the vessels which are its 
ct continuations. The cells beneath the chorda are destined 
)e 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 IIJ. 




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368 s. hatta: 

They are the spaces formed by the slackening and folding of the 
median peritoneum which coats the pronephric tubulesp 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 nephric 
segment, the space becomes divided into three pairs : the anterior 
pair, which soon disappears, is found between the tubule 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 Paul Mayer and others in Sclaehia, 
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, gl.). 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 PetronyzoUj step by step, I will give a short resume of the facts- 

1. In the earliest part of Stage ii, the mesoblast consists 
simply of the parietal (dorsal) and the visceral (median and 
ventral) layers. The proximal portion of the mesoblast is dis- 



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M0BI»H0L0GY OP OYCLOSTOMATA. 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 ir, 
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 £he 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 ii, in about 12 

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370 



S. fiATTA : 



segments and are cut oflf from the scleromyotome in 4 i 
In Stage iii, 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 ps 
fourth somite and is the first to arise ; the second folio? 
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 Euckert) 
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 communical 

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 do7*saHs. 

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 tul 
during Stage iii, 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 pai 
pronephros. 

14. The glandular part, or the pronephros proper ( 
six somites, from the fourth to the ninth. The maximun 



-.. 



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MORPHOLOGY OF CYCLOSTOMAtA. 



371 



be pronephic tubules which is attained by the embryo in 
;e III, is, therefore, six pairs. 

15. The tubules of the first and second pairs come, in Stage 
temporarily in close contact with the epiblast, but do not 
ive cells from it ; they soon return to their original condition. 

16. The anterior extremity of the system shows, from the 
, degenerating features. The first, second, and sixth of the 
lies begin, during Stage iir, 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 the 
les, which decline probably without further development, 

after their separation from the myotome is completed ; they 
I to atrophy from the free end. The next pair degenerating 
le 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 vi, 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 diff'ercnt 
ites are shown in the annexed table. 



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372 



». HATTA 




^^ 





J 




1 


> 


>- 
£ 


> 


1 


1 




1 


Slag« II 




AqIA 


Anl.2 


AtiL3 


AnLi 


Anl. 6 


Anl.6 


Anl.T 


Stiigulll 






Tub,! 1 


Tub.2 
Tiib.2 


TuU3 
Titb.3 


Tub.4 


Tiib.5 


TqK6 


Begni 


StiigelV 








Tub,4 


Tiib.r> 


Tiib.6 


SegiE 


stage y 








Tub,2 


Tiib.3 


Tub.4 


Tiib.5 


S€^menU 


Stage V[ 








Tub.3 


Tvib.4 


Tub.5 


Segment 



19. 111 older embryos of Stage iir, the visceral la; 
iieplirotome is folded out, and is called the coelomie 
which resembles the coelomie pocket described by i 
Bdelio$lomu- ; lioweverj in MdeUodoma^ the fold is deri 
the parietal and viscend layers of the luteral plate, and 
wards converted into the Bowman*^ capsule, whereas th( 
projection ii3 the product of only the visceral layer of Ih 
tome ; it gives rise to the radix of the mesentery wL 
materials to the mesonephric tubules and to the gonat 

20. The topographical position of the pronephros 
lirst definite in Stage iv. It is situated in the chest cavi 
lateral to the heiirtj forward of and dorsal to the liver, 
along either side of the chorda. This position is 
changed as the dcvelopmeot proceeds ; the prooephr 
in later stages, in front of the liver, 

21. During Stage iv, a structure, which I ha 
above the peritoneal parlition, is observed as an t 
of the peritoneal wall and disappears during iho m 



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MORPHOLOGY OP 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 IV. 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 nephrostoraes 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. 

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I 



374 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 Aulage or iLe 
duct on the other, there exists always a space, und the duet 
has no connection with the epiblast except at its posterioimost 
end where the epiblastic cells might, as judged from the nutotie 
figures, contribute to the formation of the duct 

26. In the somites posterior to about the twentieth somite, 
tlie Anlage of the duct is represented by a few cells in eacli 
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 ii, the Anlagen of the segmental duct are cut 
off from the mother layer in a few somites; in Stage in^ the 
duct is formed as far as about the eighteenth somite, while in 
Stage IV, it breaks out into the cloacal cavity. The cloacal 
opening of the segmental duct is fouud 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 iv, the primitive genital cells become apparent 
in the nephrotomes of the posterior 10 or more somites ; they 
can not be discriminated from other mesobhistic cells in tlie 



{,!> next advanced Stage 



i<i; 29. The blood-vessels, which specially supply the pronephros, 

'I * acquire definite form in comparatively later stages, viz., at about 

M Stage V. The dorsal aorta pours out the blood into two pairs of the 

:ii blind vessicles, which are formed by the folding of the parietal 

lit'! peritoneum and are found between the first and second pairs, 

^'A and between the second and third pairs, of the persistent tubules 

^1 



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MORPHOLOGY OF OYCLOSTOMATA. 



375 



3ectively. 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 
intersomitic in position. The two anterior pairs of them soon 
lergo atrophy, but the posteriormost pair persists, becoming 
irged and sacculated at the distal extremity. This sacculated 
b of the vessel is filled up with free-cells and is called the 
nerulus of the pronephros, and, therefore, there is only a pair 
glomeruli in Petromyzon. , 



II. Historical Review and Conclusions. 

As is well known, Max Schultze ('56) was the first who dis- 
ired the pronephros in Petromyzon. Having investigated the 
86 of P. planeri, the author describes the structure as 
riisenanlage" and homologised it with the "Urnieren (Wolfsche 
per) " of the frog's larva. His statements on this body are as 
)ws : ** Nicht lange nach der Bildung dieser Driise (Thymus) 
teht die Anlage einer zweiten, aus dem unter der Chorda 
salis angehiiuften Blastem iiber dem Herzen. Aus der durch 
mentanlagelungen friih schon sehr undurchsichtig werdenden 
ise wachsen nSmlich nach unten, gegen das Herz zu, 3 oder 
urze Fortsatze hervor, welche eine eigenthiiraliche 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. Muller and Max Fur- 
jfGER. Muller ('75) noticed the first traces of the pro- 
hros in a very young embryo, which had yet only four pairs 



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8. HATTA 




of gill-Blits. Tlii^ Aiilage gives rise to a much coiled glai 
opens into the body-cavity, at first throiigli only oik 
funnel, but afterwards througli foin\ Tho gland pa* 
posteriorly to a pair oF ductf^, whicli run rilong the c 
either side and open into the cloaca. Mulleb has hoi 
the structure with the *'Vorniere'* of Myxme and c; 
duct '* Urnierengang " (pp* 121-122). He found a pai 
meruli projected on the median surface of the gland s 
with the peritoneal epithelium. 

Max Furbrtnger ('7v8) studied the larvje of A 
planer I J which varied from 4.5 to 180 mm. in length. ] 
ments essentially confirm Muller's* In his account we 
following sentences : '* Die auf alien Praparaten am 
Vorniere, die ich im Wesentliclien ganz w^ie Mtjller fai 
einen nahmenttieh bei den mittleren Stadien volumin 
durch 4-0 Myokommata erst reck ten Complex von Wi 
die vorn durch mehreren Peritouealeaniile (Wimpertri 
Bauchhuhle munden und hinten in den Vornierengang ii 
Diese auf die 2-3 ersten Myokoramatii heschninkten 
rogen in nnregelmassiger Folge bald ventral-medial, bah 
lateral in die Bauchhuhle vor nnd wnrden (von Ci 
und mir) meist zu fiinf gefnnden. Die von rundliehe 
zellen bekleidete Glomerulus verliielt gan^ wie MuLLERbe 
llh 42). 

The larva} of AnnnoenfifH in question seems to C\ 
probably to Stage v, or later stages of my list ; in sucl 
I could not iind more than three (or rarely four) pai 
tubules, or of the nephrostomes.^^ 

l)Se6 the Xoot-note on p. 3j8. 



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MORPHOLOGY OP CYCLOSTOMATA. 



377 



The authors who have investigated the development of 
tromyzon embryos step by step, are W. Scott, Goette, Shipley, 
i v. Kupffer. Their opinions are, however, somewhat 
rergent. Scott ('82) derives the pronephric tubules from the 
;mental duct which is, according to him, brought about by 
5 differentiation in situ of the cells forming the proximal 
irgin of the lateral plate. The process takes place in the 
lOle extent at the same time. At certain points (segmental ?) 
the duct thus formed, evaginations are produced out of it ; 
5se evaginations subsequently open into the body-cavity and 
ablish the nephrostomes which are, according to Scott, 
ind from two to three pairs in number. At about the stage 
which the funnels are formed, he observed a pair of 
imeruli. 

" In most respects," Shipley's observations ('87) *' confirm 
I *' (Scott's). But " on the origin of the ciliated funnels, the 
lults differ from Scott's " and agree with those of Fubbbinger 
mphibian pronephros ?). According to Shipley, " in the region 
the heart, where the body-cavity has already appeared, its 
gin {Le.j of the segmental duct) seems to be somewhat different, 
le lumen of the segmental duct here becomes continuous with a 
)0ve in the parietal peritoneum, lying near the angle where 
5 somatopleure and the splanchnopleure diverge. When this 
)0ve closes it leaves four or five openings which persist as the 
enings of the ciliated funnels" (p. 20). 

V. KuPFFER^^ ('88) observed, in P. planed^ the three pairs of 
5 tubules arising from three distinct evaginations of the parietal 



1)1 know this paper only by the abstract in: Jahresbericht ii. die. Fortschr. d. Anat 
Physiol., Bd. 17. 1889. 



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378 s. UATTA : 

layer of the lateral plate ; the segmental duct is looked upon a3 
of the epiblastic origin. 

GoETTE ('90) worked the development of Pelnmyzon fiuvin* 
tilis ; his results with respect to the pronephros show some agree- 
ment with mine, especially those concerning the later stages, Thu 
author derives also the whole system of the pruuephros (includiiig 
the segmental duct) solely from the mesoblast. But we diverge in 
some important points from each other ; ho 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 aceoiint 
given in the 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, &c., 
which are spoken of as representing the first appearance of the 
structure, approximately correspond with my figures 82, 83, &c., 
and with those of even older stages. 

The pronephros is, according to Goette, 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 evaginatian of the 
parietal layer of the mesoblast ; the lips of the furrow being fused 
at certain points, there remain three openiugn; 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 
j relation to the metameres of the body ; for 3 to o tubules are 

found in the extent of 2 to 3 metameres {he, ciL^ pp. 64-65). 

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 



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MORPHOLOGY OP CYCLOSTOMATA. 



370 






ave made out. From the region of the liver-aiilage backward 
development of the duet is irregular ; he says : " Auf der 
n Seite zeigt sich seine Anlage noch rinnenformig, wiihrend 
auf der andern Seite schon* vollkommen rohrenformig ab- 
hniirt ist. Eudlich wechselt dies Verhalten auch auf derselben 
perseite, so dass derselbe Gang, von der Lebergegend ruck- 
ts verfolgt, bald rinnen-, bald rohrenformig, geschlossen oder 

offener Lichtung sich darstellt '' {loco cit.y p. 56). The hind 

of the 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- 
ital duct demonstrated by Goette in the region immediately 
ind the ** ursprungliche Kopfniere." 

Goette has made out the three ** peritoneale Scheidewiinde," 
le 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 
1 the " Schlussplatte " of the pronephros in Teleostei ; con- 
jred phylogenetically, nevertheless, they have no intimate 
ition to the pronephros in Pelromyzon (loco cit., pp. 56-61). 
s structure is, as stated on p. 349, doubtless the same as the 
)erniost 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. 



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380 



8, ITATTA 



96 and 97, to which his statement refers, represent a bU 
giderably later tlian the fii^st formation of the pronephr* 
And the peritoneal partition is not confined to these threi 
but is continuous throughout the whole extent of the proi 
moreover, beside the '* peritoiieale Sclieidcwande/^ there a 
two other partitions of a similar character as above 
Also, as to the fate of tlie structure my results difier fr 
I have not been able to observe at all any such cout 
to the formation of the hind wall of the branchial ehaiu 
of the ** Venenbriicke/' as is affirmed by Goette, 

Bi.BL ('96) says in his recent extensive work on the S 
nephric organ, that in quite young larvffi of Peiromyzon / 
the pronephros also begins in the seventh soraite, in wl 
first of the four ostia are found, as in Prisiiurus^^ H 
are, however, 501 hour^ or 20 days and 21 hours oL 
larvse correspond to my embryos in Stage vi, and upw 
which anteriorly two pairs, and posteriorly, one paii 
tubules disappeared and only three persistent tubules i\ 
Hia first neplirostonie represents the foremost of the i* 
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 singl* 
tion of \\ KuPFFEB who assumes the epiblastie origin 
segmental duct. They diifer from the account given in i 
going pages in the mode of the formation and in the nu 
the tnbules formed. The first point of difference is du 

l)&k'*; ihe refereuge under Seiavlik (p, oUU), 



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MOEPHOLOGY OF CYCLOSTOMATA. 



381 



t that the authors probably overlooked the earliest phases of 
ination, which take place, as shown above, in a stage very 
inig, but not younger in comparison than tliat in other Anamnia ; 
the formation follows the metameric seginentation of the mesoblast 
the anterior region. In later stages, the tubules and the inter- 
aitic 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- 
a. 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 
18 embryos of Stage in, 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 
testral history.^^ 

As is seen above, all investigators who have been occupied 
h the study of the development of Petromyzon agree in dc- 
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 iir, the anterior extremity of 
left collecting duct presents a conical protuberance (see the foot-note on p. 329). 




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382 S. HATTA : 

alimentary canal extending through about the same space as llie 
glamlular part of the kidney. Each glomerulus is a divert icuhim 

nf ihc [vrritoneum, which generally becomes sacculated; ,../' 

(p. 21), The statements by Goette confirm Shipley's, and my 
I'BHult^ txUf} agree with theirs. However, this is not all of Ihe 
vascular system of the pronephros but represents a posterior 
portion of it, the anterior part having disappeared entirely 
(>=i*e p. 308). 

No previous writer on Petromyzon has described such early 
stiiges as given above in the development of the prone pliros, nor 
\\m 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 
Craniotrt to verify the new facts. 

AVith reference to the development of the nephric organ in 
Myxinoids, there is a great deal of information which we owe 
to the unwearied labors of W. Muller, Semo^% Weldon, and 
others^ \ They had, however, no opportunity to observe the earliest 
stHge of the embryos. Recently our knowledge on this subject 
has been greatly augmented by the new works of Price, Dean, 
and M-4AS. 

Price ('97) worked out the early development of the prone- 
phros observed in a few embryos at different stnges of BdeUmioim 
MmitL According to him ** the first indication of the system 
oceiirs here in the eleventh segment (of spinal ganglion), and 
consists of a simple thickening of the somatic layer of the coel- 

1)1 lure pot seen the paper by J. Muller. 



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MORPHOLOGY OF CYCLOSTOMATA. 383 

omic epithelium, which extends through seven section?, 

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. 

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 eU.y p. 210). 

The pronephros in Bdelhsloma 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 " (loco ciLj p. 217). 

The segmental duct (in s, sfr.) 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 PetromyzoTij as 
stated in the foregoing description, the Anlage in a mesoblastic 

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384 S. HATTA : 



I 



somite develops solely into the tubule, and by tlie secondary 
union of the tubules' ends, the collecting duct is brought about 
As regards the number of the tubules, there are, in 
Petromyzony only two pairs in the branchial region instead of 
twenty in Bdelhtoma. The number is, however, of secondary 
-f importance; it varies with the stages of embryos and possibly 

with individuals, and naturally more with tlie embryoB of 
I'l diflferent families. This numerical variation is readily explained 

by the degenerating tendency of the tubules. 

Pbice has made out the segmental evaginations of the dorsal 
corner of the coelomic cavity corresponding to the nephromeres; 
they are called by him the " coelomic pockets." In PelromyzQn^ 
I I have found a series of solid knobs on the visceral layer of 

the intermediate cell-mass^ which are transformed into tke 
<^ segmental folds of epithelium, forming then the direct continua- 

tion of the peritoneum. Thus the coelomic pocket in Bdelhstomn 
I and the coelomic projection in Petromyzon are apparently very 

similar structures; the two, however, differ from each other m origin 
[riJ I and in fate. The former (coelomic pocket) is constructed bj the 

^;fl parietal and visceral layers of the lateral plate, while the latter 

i^(|f (coelomic projection) is the product of only the visceral layer 

^M of the nephrotorae, the ventral half of the segmenied pai^t oflk 

' ' inesobla^t. The coelomic pockets become the Mulpighian boJy, 






and the coelomic projections give origin to the radix of tbe 
mesentery, from which the gonad-cells and the niesotiepbric 
■;[»^ tubules are derived. Nevertheless, these two structures ai*e, I 

> |t believe, homologous. Price's statements on the derivation of the 

^if coelomic pocket from the two peritoneal layers, are not as clear as 

Lj'j is desirable, and its partition from the body-cavity might, it 

?;l*| seems to me, represent the uppermost periton(^al partition whi^b 



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MORPHOLOGY OF CYCLOSTOMATA. 



385 



n disappears, in Felromyzon, 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 Myxine 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- 
led by the author differ from those of Price 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. " If the organ in question could 
y be a pronephros alone, or mesonephros alone," says Price 

should unhesitatingly pronounce in favour of its being a pro- 
)hros " (loco city p. 120). And he proposes to call *'the 
ire embryonic kidney holone'phrosP With Rabl, 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 
lit. The formation of the mesonephros takes place in Petro^ 
ton only at a stage much advanced, in which the processes of 

formation and degeneration of the pronephros go on in much 

same manner as in Bdelloslcyfiia, and it is open to doubt if the 
30nephros 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 
I AmpMoxtis ; '* but they do not have any relation to the 
inings of the tubules, nor have they any direct connection 



lip 



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386 S. HATTA : 

with the aorta '' (loco ciL, p. 213). The glomerulus figured in 
his Taf. 17, fig. 12 (gl) corresponds, I think, with a part of the 
glomerulus in Peiromyzon^^ 

As is very well known, the independent studies of Weiss 
('90) and Boveri ('92) on the branchial chamber of Amphmm 
gave a new direction to the morphological investigation of this 
field. A number of external openings of ciliated tubes i^ found 
at the dorsal corner of the peribranchial chamber of Ampkmu$, 
Through the morphological study and physiological experimeats 
this organ-system is demonstrated to be the excretory apparatu^j 
or " Nierencanalchen/' as Boveri calls them, of Amphmu^^ 

Boveri counted 91 ^'Nierencaniilchen'' in an individual 4 era. 
in length and possessing 183 gill-bars on the right side. In the 
adult he counted about 180 of the " Nierencaniilchen "; the number 
is, however, by no means constant, but varies within a certain limit. 

In the middle region of the branchial chamber, a '* Nieren- 
caniilchen " has 3 or 4 ** Seitentrichter,'' and 2 ** Eudtriehter;*' 
such is the most complete one. It becomes gradually simplirted 
both anteriorly and posteriorly, until it is at last represented bv it 
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. 



1)Dean has published two papers on the development of the Californian Hag09§in«t 
;u *99); these excellent works contain merely the general account of the con r^ of the dcvdop* 

•;! ment in surface view. We may expect that the full account will throw miicb light m^ 

'J ontogeny of Craniota. There stand, in the account given by him in these works, tbt i^" 

;r portant facts tliat the " proncphric tubules arc apparent in connecliou with all the ujeR^ 

blastic somites" ('03, p. 274) and that the pronephros is extended far buckwartb, heyOfliJ ti** 
anal region, into the tail ('09, p. 272). It would he liighly desirable to observe the ^ronfph^^ 
tubules of the Hag in relation to the myomere, and not to the spinal gangbia aluae, as FKiCt 
has done. 



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MOKPHOLOGY OF CYCLOSTOMATA. 387 

These nephric tubules receive, according to Boveri, 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 Boveri 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 " Nierencanalchen " 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 Amphioxm ; 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 
other systems of organs. The " Nierencanalchen " 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 ciL^ fig. 3). 

Thus the author has brought the '* Nierencanalchen " of 
Amphioxvs 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 the development of the pronephros in Selachia, Teleostei, 
and Amphibia, which will be treated further on. 



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388 S. HATTA : 

Thanks to tbe labors of many eminent mvestigators, tbe 
early development of the Selachian pronephros has been, as is well 
known, fully studied, so that the facts gathered from ihis tield 
are well adapted to be compared with those from other group. 
I have found, in the present investigation, many importaat points 
running parallel with the development of the Selachian pro- 
nephros. I may then be allowed to compare my own results in 
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." Tbis Anlaige 
is first brought about by the thickening of the parietal layer of 
the mesoblast, which is found " in den Bereich des segmeudrten 
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 unsegmeutirten Myoblast 
der Peritonealwand iibergehen '^ (p. 209). The " Segmental wulst '' 
is so called because it is noticed as the segmental thicken- 

IJjK ing of the parietal mesoblast of which Ruckert recogniM 

in his Stad. iv^\ six for Toi-pedo and four for Prislmrm^ stretching 
over a corresponding number of the myotomes. The first indication 
of the pronephros is expressed, in Selachia also, segmentully in 
the segmental part of the mesoblast at the stage in which the 

'jf 1;, metameric segmentation of the mesoblast is still going on, auJ 






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l)The embryos in the stage have 25-27 somites. 



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MOEPHOLOGY OP 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. i) 
regelmassig die starkste Entwicklung in seiner mittleren Ab- 
schnitt, also ungefahr im Bereich des dritten des ihm gehorigen 
Somiten, und verjungt von da allmahlich nach seinem vorderen 

und hinteren Ende zu " (loco cit., pp. 210). 

The account given by ROckert is essentially confirmed by 
later investigators such as van Wyhe ('89)^^ Rabl ('96), and others, 
although^they differ from one another in the interpretation of the 
facts and in some unimportant points. Eabl looks upon the 
Anlage of the pronephros (his Vornierenwulst) as the ventral 
poition 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 Wyhe states : ** Da nun der 
Pronephros, wie spatere Entwicklungsstadien zeigen, ein Produkt 
der Seitenplatte ist, wahrend der unmittelbar dorsal davon liegende 
Theil des Mesoderms zur Mittelplatte gehort, ist die Segment- 
irung des Mesoderms bei Selachiern also nicht auf die Myotom- 
platte beschrankt, sondern erstreckt sich auch auf die Mittelplatte 
und den dorsalen Theil der Seitenplatte '' {loco ciL, 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 tlie lateral plate. I will, in 

l)The embryo, in which the first traces of the pronephros is seen, is, according to 
VAN Wyhe, in a stage with 27 somite?, whereas Babl has seen in an embryo of PriAtiui-us 
with 25 somites. 

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S, HATTA 



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this place, not go furtlier, but return in future page« 
discussion of this point. It is, Iioweveri safe, I believe, t 
thin portion of tlic niesoblast as a part of the somite. 

Va?^ Wyhe found the foremost pronephrie segmen 
third body-somltp (his linnipftegnieiU), and Rabl states 
Voniierenwulst be^:^iiis iji the seveiitli somitt* form 
Gesfinimt.^egment). According to Radl, however, vah 
third Eumptsegment corresponds to his seventh Gemuimt 
To verify this fact Kabl lias extended the comparis 
Pelfomyzon, and fouinl tliat in tins case also the pr 
begins in the seventh somite ; but the pronephrie tubul 
somite is, as noticed above (p. 380), not the ant^^riornio: 
tubules in his sense^ but of the persistent tubules. 

Van Wvoe noticed five of the pronephrie segra 
Rrija^ and three for SvyUium and Prutiurus ; while Rabi 
eight Vornierenwiilste for Hajct^ and four for Pristiurf 
results in Petromyzon, therefore, best «giee -with those 
by RucKEHT in Torpedo, 

The authors agree in deriving the collecting duct 
lateral extremities of the pronephrie Anlagen, wh< 
become eonfiuent. 

RuCKEKT tuis observed, in PnMiiirmy as well as in 
the mcondary connection of the Segmentalwulst with the 
which has led him to believe in some contribution of < 
cells to the formation of the pronephros, while yai 
and Rabl deny this, I have found the same conm 
Peirmnyzon, but 1 have found no sign of the eoc 
of epiblastic cells to the forjnation of the pronephr 
phenomenon is temporary in both Belachia and PeJ 
it takes place in Sehichiap according to Rijckert, in hk 



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MORPHOLOGY OF CYCLOSTOMATA. 



391 



already in his Stad. in, a space is seen between these two 
ctures. 

The degeneration of the tubules in Selachia runs a course 
illel with that mentioned under Amphioxxis and Cyclostomata. 
seen above, the Anlageu of the pronephros are developed most 
rously in the middle part of the pronephros, as in the case 
imphioxuB and Cyclostomata ; and degeneration begins at the 
lial 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 
le 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 PelroviyzoUy 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 
jcting duct is constricted off" from the lateral plate by 
teration of the tubule and constitutes the foremost section of 
segmental duct, in precisely the same manner as in Selachia. 

diflference is : in Fetrwnyzon the communication with the 
jr-cavity is retained by the three middle nephrostomes, while 
Selachia, it is through only the middle one, that is, the ostium 
nninale. 

The segmental duct becomes apparent in an embryo with 



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392 8. HATTA : 

35 (van Wyhe), or 34 to 35 (Rabl) somites. The anterior 
small section of the duet 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 Pelrmiytm^ 
RiJCKERT ('88) and van Wyhe ('88, '89, '98) believe that 
it is the product of the epiblast^^, while Rabl 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 Wyhe and 
Rabl, in the embryo with 83 to 84 somites. It can be inferred 
from VAN Wyhe's figs 7a and 7i, 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 
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 
Paul Mayer without reference to their relation to the pronephros, 
were studied by Ruckert and their true nature was pointed out 
by him. There are six of them in T<yrpedo corresponding to the 
number of the nephric segment ; they are, however, not soraitic 
but intersomitic in position. The vessels not only pass through the 
nephric fold, but throw a solid process, the interior of which 
consists of round or spindle-shaped cells. This is, according to 
Ruckert, the equivalent of the pronephric glomerulus of Am- 
phibia described by Furbringer. The development and decline 
of these vessels go on parallel with those of the pronephric diver- 

1)1 win return to this point again in future pages. 

2) According to Rabl's counting, this somite corresponds to his forty-second somite. 



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MORPHOLOGY OP CYCLOSTOMATA. 



393 



la. The vessels in the cranial as well as in the caudal part 
be pronephros are weaker than those in the middle (the third 
fourth) ; only the latter vessels develop further and become the 
line artery. Van Wyhe confirms Ruckert's account and 
(Jescribed three vessels in Fristiurus. In addition to these, 

Wyhe 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 
ih sends oiit, 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. Boveri remarks that the 
3ls of Paul Mayer present many points of harmony with 
branchial vessels in Amphioxus. Rabl agrees essentially with 
account given by Ruckert and van Wyhe, but denies the 
-ence of a glomerulus. According to Rabl, the structure 
id the glomerulus by Ruckert does not fulfil the condi- 
j of being a glomerulus ; he says : '^ Eine einfache Ausbucht- 

einer Arterie ist noch keine Gefassschlinge, geschweige 
1 ein Glomerulus " {loco city 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 
)ellacher (73), Goette (75 and '88), Furbringer ('78), 

Hoffmann ('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, 



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394 8. HATTA : 

which is subsequently constricted ofif from the body-cavity. 
This takes place at iirst 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 GtOETTE, the 
morphological equivalent of the nephrostonies 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 
11 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 FuBBRiNGER (78), HoFFMANN ('86), and others, although 
Hoffmann differs in his view of the mode of the formation of 
the glomerulus. 

According to the results recently arrived at by Felix*^ in 






1) I know his paper only by the abstract in tbe Jaliresbericbte iiber die FortschriUe der 
Anatomie und Pbysiologie, N.F. Bd. III. '97. 



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MOEPHOLOOY OP CYCLOSTOMATA. 395 

the embryos of Salmonidse, 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 " primare Vornierenfalte.*' The 
" primare Vornierenfalte," which passes over into the parietal and 
visceral layers of the lateral plate, undergoes a longitudinal 
constriction (the " sekundare 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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I 396 8. HATTA : 

ll 

cess proceeds posteriorly until the duct comes to lie close to the 
rectum (Enddarm). 

Felix thus observed the segmental Aulage of the pronepliios 
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 Bracket (*99) have published a 
paper on the early development of the mesoblastic organs in 
Salamonidse. 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 
\l of the parietal layer of the lateral plate only, but is formed, 

i; as Felix believes, by the proximal portion of both the parietal 

j. and visceral layers of the secondary lateral plate (restreme iirterDc 

r de la plaque lat^rale second aire^^). The internal cavity enclosed 

by the pronephros is, therefore, not the diverticulum, but a 
r part of the body-cavity. 



1)1 am much indebted to my fiiend, Dr. A.Oka, who read I be pjiper fur lue. 

2)Accordiug to the authors, the " plaque lat^rale primitive " is divided inio I lie *' pb^ae 
lat^rale secondaire" and the ''masse interm^diaire ;'' therefore, the *' pluf(uc Uterak 
secondaire " corresponds to the lateral plate itself of Petromyzon, 



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MORPHOLOGY OP CfYCLOSTOMATA. 



397 



The Anlage of the pronephros is laid in exactly the same 
ner from the fourth somite to the cloacal region. Under the 
rior three somites from the fourth to the sixth, the Anlageu 
developed into the pronephric chamber ; the Anlagcn posterior 
liese are all transformed into the " canal excr^teur," as they 
the segmental duct, and they have come to the conclusion 
the '* canal excr^teur '' of the pronephros has the morpho- 
jal 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 
5lf in the foregoing pages. Diflferences between their 
Its and mine are that the authors derive the system from the 
al unsegmented mesoblast, and that both the parietal and 
jral layers of it partake in the formation of the system. As 
been stated in the descriptive part, this derivation is only 
irent ; 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 
ite ; or, the result of the development of the pronephros. It 
lerely for a physiological reason that this development or 
ration of the pronephric Anlage goes on earlier than, for 
mce, 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 



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•*. 



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398 8. HATTA : 



Amphibia shows, in several respects, a parallel course with that 
of Petromyzon. Careful observations on the developmeDt of 
Amphibian pronephros, adduced by recent investigators, have 
intensified this similarity with the exception of a few points whicli 
^ 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. MiJLLER (75), GOETTE (76), FiJRBRIKGER (78), HOFFMANS 

('86), and others. The stage at which the pronephros appears 
coincides exactly with that in Petromyzon^ as Max Fubbbingee 

I says in hip well known work : " Die erste Entwicklung der 

Vorniere und ihres Ausfiihrungsganges findet hier nach der 

i I Scheidung des Mesoderms in Urwirbel und Seitenplatlen statt 

und folgt unmittelbar der beginnenden Sonderung der eiBten in 
einzelne Urwirbel amd der Spaltung der letzteren in Haut- und 
Darmfaselplatten. Embryonen von Rana tmup&raria von cirm 

J' 2.5 Mm. Lange und von Triton alpeslris von ea. 2.0 Mm, L. 

•^ entsprechen diesen Stadium" (p. 3)^^ 

I MoLLiER ('90) has made out the segmental Anlage of the 

1^ Amphibian pronephros, having worked with the embryos of Triimt 



\^' l)The.nephro8tomes are found, according to the author: 



2 in Salamandrina maailaia, 3 in Bona tempfyrartn, 

2 in Triton alpesLrisy 3 in Bcmbinaier itpieu* (Goette), Atid 

2 in Sir&lon piscifoimiSj 4 in Ooecilia rostt-ala (SpenqelJ* 



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MORPHOLOGY OF CYCXOSTOMATA. 399 

Bufoj and Rana. His accounts confirm, as a whole, those given 
by RucKERT for Selachia above referred to, but differ some- 
what from those of most other authors who have worked on 
Amphibian pronephros. Mollier states as follows : " Wir sehen 
hier ebenfalls ziierst eine solide, von dem Mesoblast ausgehende 
Anlage, deren Structur anfanglich 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 
Canalclien vorhanden sind, die von den Somiten in conver- 
genter Richtung ausgehen und erst nachtraglich untereinander 
vereinigen zu einera Langscanal, von dem aus die Vornierent- 
richter in die Leibeshohle fuhren " {loco cit., p. 229).*^ The 
author derives in this wise the pronephric tuhules, exactly as 
in the case of Peiromyzorij from the segmented part of the 
mesoblast only. 

Mollier's accounts are for the most part in close accord 
with the results given by Field ('91, p. 282), who, one year 
later independently of Mollier, began with Anura, and 
extended the work over Urodele Amphibia. In one point, their 
results differ widely; but "the difference is,'' it seems to Field, 
" apparent rather than real." According to Mollier, the 
nephrostomes communicate with the cavity of the myotome, the 
myocoelome of van Wyhe ; this is denied by Field, 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 
that this " ventral segmeiU of the mesoderm " corresponds to the 
pronephrotome of van Wyhe in Selachia or to " I'extreme interne 
de la plaque lat^rale secondaire " of Swaen and Bracket in 

l)It seems that the earliest traces of the pronephros are perceived in an embryo 
younger than that with 7 somites. 



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400 S. HATTA : 

Teleostei, and I agree with the view of Ruckert, here represented 
by that of Mollier. 

MoLLiER and Field agree with each other in assigaing three 
pairs of the tubules for Eana and Bufoy extending from the 
second to the fourth somite, and two pairs for TriLon (Molliee) 
and Ambly stoma (Field), covering the third and fourth somites.*- 
In addition to these, Mollier observed occasional occurrences of 
the third tubule in TritoUj which is, according to FielDj not 
equivalent, as Mollier maintains, to the third tubule in Bufo 
and Rana^ because the additional third tubule in Triton 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 lehihyophh. 
A pair of glomeruli has been made out in Amphibia ; 
the structure is connected by special vessels witli the dorsal 
aorta on one hand and with the cardinal vein on the otlier. 
This branch of the aorta is believed by Field to correspond 
to a part of Mayer's vessels in Selachia. Beside these, there is 
,1 no vessel arranged segmen tally or otherwise. 

i' The section of the body-cavity corresponding to the pro- 

;. nephric stretch is gradually expailded, and is shut off teroporarilr 

from the rest of the cavity by a close contact of tlie parietal 
j- and visceral layers of the coelome ; this part of tbe cavity is, 

;.. according to Goette, homologous with the pronephric chaiul)er 

in Teleostei and with the homologous structure in Peiromyzon, 
[^ which is called by him the *' peritoneale Scheidewande," 

[ ': The so-called ventral portion of the Amphibian prouephrcB 

r is, according to Mollier, brought about by the separation of 



M 






l)Acconling to Field, Mollier's first body-segment in Triton correjips^nds ta bis tbtwl 
sonute in Ambfysioma. 



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MORPHOLOGY OP 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 anterior most 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.y 

p. 286). This feature of the duct shows, it seems to me, a close 
resemblance to the an teriormost 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 Field elucidate this point. 

"The segmental duct arises," Field says, ** throughout its 
entire length by a proliferation in situ of the somatopleure " 
{loco ciLj 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 city p. 223). In Stage v, the cloacal opening is seen. This 
opening is found, in Rana and BnfOf in the vertical plane with 
the middle of the twelfth somite, whereas it is below the twentieth 
somite in Amblydoma (Field). 

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402 s. hatta: 

The duct is segmental in origin. Field says : *' I 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 " {loco cit.j p. 219). There are no 
other Vertebrata which agree more with Petromyzon 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 Perenji ('87) 
has published the results of his study on Rana e^culentay 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 Ruckeet in 
Selachia, Mollier says : " Im einen Punkte weiclien die Amphibia 
von Selachiern ab, dass die Vorniere mit dem Ektoblast in 
keino nahere Beziehung tritt. Allerdings heftet sie besonders in 
den Stadien, in welchen sie voluminoser erscheint, dera Ektoblast 
oft in affallend iuniger Weise an. '*' * '•' Doch lasst sicli 
stets eine scharfe Grenze beiderlei Blatter ziehen, wenigsteus bei 
Bufoy wo die Ektoblastelemente durch ihren Pigmentgehalt deut- 
lich gekenuzeichnet sind '' [loco cit.y p. 229). 

The historical review undertaken in the 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)1 have not seen the paper by Brook. 

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MORPHOLOGY OP CYCLOSTOMATA. 403 

la 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 raetameric 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 BdellosUyniay 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 Fronej^hric Tubule is the Product 
of the Mesoblastic 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 Petromyzon, 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 the 
Anlagen of the pronephros (or the nephric segments) together 
with the lateral plate are cut oflF 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 



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404 S. HATTA r 

step in the diflferentiation 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 Lacerla agilis is very instructive. According to 
Hoffmann ('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 Petromyzouy 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 Eeptilia (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 segmen tally 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 pronephi*os belong ? 

Van Wyhe ('89) has distinguished, in Selachia, three por- 

l}Thi8 view is grounded upon tlie suggestion of Pkof. Mitsuxubi. 



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MORPHOLOGY OP 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 raesonephros 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 raetaraeric 
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 
RucKERT ('88) and Eabl ('88, '96) have remarked. According to 
RiJCKERT and Rabl, the segmented portion — the somite — com- 
prises the myotome and the sclerotome ; the pronephros and the 
mesonephros are derived from its ventral (distal) portion, which is 
called by Ruckert the '' Nephrotom " ('88, p. 272). 

In Petroinyzoriy these two portions of the mesoblast, the 
segmented and the unsegmented, are, in early stages, clearly dis- 
tinguished, being histologically diflPerent (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 Rabl. 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 



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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 coelomic projection on the other, and, 
therefore, corresponds to the '* Nephrotom " of Ruckert. 

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 diflferentiation of the mesoblast, 
in Pelromyzon corresponds also to the " Ursegment " of Am- 
phioxus 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 
coelomic cavity and the sclero-myotome, that is to say, the 
nephrotome. Let us now examine what part of the Ursegment of 
Amphioxm represents the nephrotome of the Craniota. 

In his excellent work on ^^ Die Kierencanalchen des Am- 
phioxns," 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 Amphioxus. 

But these chambers *^ sind urspriinglich die segmentale 
Verbindungscanale zwischen der unsegmentirten Leibeshohle und 
der Sclero-Myotom gwesen " ('92, p. 493). I can, therefore, 
ascribe no other significance to the ventral half of the segmented 



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MORPHOLOGY OF CYCLOSTOMATA. 407 

mesoblast in Pelromyzon than that it is the morphological 
equivalent of the " segmentale VerbinduDgscanale." 

It thus follows that the distal half of the segmented mesoblast 
in Petroniyzon undergoes exactly the same fate as that in Am- 
phioxus : it is transformed into the pronephros and the coelomic 
projection or the ** dorsal segmental coelome," which latter gives 
rise, just as Boveri suggests in Amphioxtis, 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 Petromyzan. 

It mayy thereforCy 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^myotonie. This separation is, as 
above stated, not the separation of the somite front the lateral plate, 
but the differentiation of the somite into the sclero-niyotome and 
the nephrotome, preparatory to the development of the urogenital 
system. The reason why the separation takes pla^e earliei' in some 
groups than in others, rests only on physiological grounds. 

B. — The Whole System of the Pronephros of Oyclostoinata, 

Teleostei, and Amphibia is Homologous with the 

Nierencanahhen of Amphioxus (Boveri) 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 " Nierencanalchen '' of Amphioxus, basing his 
arguments on the structure, the position, the myomeric arrange- 

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408 S. HATTA : 

ment, the physiological function, and the relation of the vascnlar 
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 Cycle- 
stomata with the " Nierencanalchen " of Amphioxus. 

It is well known that the starting point of the hepatic 
diverticulum from the enteric canal demarcates, in the Chordata, 
the respiratory section of the canal from the nutritious section of 
it ; and, as Geoenbauk (78, pp. 563 — 581), Balfour^^ ('85), and 
others affirm, the oesophagus 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 ccecum of Amphioxus 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 '^ Nierencanalchen," and of 
the pronephros in different groups of Craniota. 

l)From the account of Balfouk, I will cite the following lines: — 

* Jn 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 cleits 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 e-isentially the nerve of the 
branchial pouches, supplies, in addition the walls of the cesophagus and stomach, it may 
reasonably be concluded, as has been pointed out by Qegenbaur, that tlie true respiratory region 
primitively included the region which in the higher types forms the oesophagus and 
stomach' (Vol. ii, p. 758). 

Bat^four has also shown that the solid cord of the a?sophagU3 in Elasmobranehii and 
Teleostei, is the remanent of the gill-rudiments in the ancestry (loco cU.^ pp. 61 and 78). 

2) J. A. Hammar, '99, '98, and Gmx) Schneider, '99. 



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Morphology of cyclostomata. 401) 

The " Nierencanalchen " of Amphiazus, 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 coecum. 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 Petromyzon, twenty in Bdellosloma) of the anterior 
nephric segments are found in the braachial 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- 
canalchen " in about the middle one third'^^ of the branchial region 
of ArnphioxuSy and that the ** Nierencanalchen " lying posterior to 
this point are represented, in Cyclostomata, by a number of the 
rudimentary tubules which are converted into the segmental duct. 

The '* Nierencanalchen " 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 of the 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 Petromyzon with the 
•' Nierencanalchen " of Am^^hioxus : in other words, the condition 

l)The glandular part of the pronephros in reiromyzoiif are represented by the six 
pronephric tabales. 

2) The branchiomeres in the posterior section of the gill-basket of Amphloius are after- 
wards added (see pp. 410-411). 

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410 S. faCATTA t 

seen in the *' Nieren canal chen " would be brought about, if the 
tubules in Fetroniyzon 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 ir, where 
the tubules have developed a little beyond the mere Anlage. 

The pronephric tubules of Pelromyzon 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- 
canalchen" 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 " Nieren canalchen " in front 
of the base of the hepatic coecuni, 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 ** Nierencanalchen " 
io Amphioxus. We learn from La^jkester ('89) and Willey ('91) 
that in Amphioxus, the new branchial slits are added, by stages, 



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MORPHOLOGY OP 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 

h'fe. 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 Amphtoxus 

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 " Nierencanalchen " in this hinder part may represent the 

pronephric tubules in the post-hepatic section of the segmental 

duct of Cyclostomata-^ 

The pronephric system of Petromyzon comes to have the same 
relations with the epiblast as the ** Nierencanalchen " o( Amphtoxus 
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 ** Nierencanalchen " 
of AmphioxtLSf 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 c'd,^ p. 758.) 

2) See p. 108. 

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412 s. HAttA: 

them with the epiblast, &e.), the pronephric system of Teleostei 
and Amphibia shows, as stated in the historical review, the same 
characters as that of Cyclostomata, so that the facts established 
in Cyclostomata have the same significance for the Teleostei and 
Amphibia. 

Although such is the case in those Craniota and Aviphioxus, 
the pronephros of Selacliia 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 (Ruckebt). 
These pronephric Anlagen in Selachia are^ therefore, the nior- 
phological equivalent, not of the glandular portion of the pronephr<^, 
but of those which are converted into the segmental duct in the 
Graniota just mentioned. 



C. — The Segmental Duct in Selachia is not the Morphological 

Equivalent of the Duel of the Same Name in Cyclostoinaia^ 

Teleosteij 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 Kabl, who advocates the meso- 
blastic origin of the Selachian segmental duct. The facts given by 
Rabl are, however, not the same as those observed in the 
groups just referred to. In these, as stated above, tlie duet is 



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MORPHOTX)GY OF CYCLOSTOMATA. 413 

(liflRerentiated, 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 Rabl 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 PelromyzoUj 
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 Eabl, 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 Petromyzovi}^ the 
pronephros originates in the mesoblastic somites anterior to the 
hepa to- 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 Petroniyzon and the other Craniota above 



l)As we leam from Goette I'Tf)) and Oellacher ('73), the anlerior section of the 
pronephric 6ys?tem in Amphibia and Teleostei, is^ also found in the mesoblast opposite to the 
posterior section of the fore-^it, 



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414 8. 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 Petr(yniyz(m 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 Ruckert and van Wyhe affirm, or by 
cell-multiplication within the structure of the mesoblastic collect- 
ing duct itself, as Babl states. Hence it can not be homologous 
with the duct of the same name in Petrcytnyzon 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 Petrmnyzon and Amphibia. 

In PetroTriyzon^ 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 ix)int of the epiblast 
This fused condition of the duct and the epiblast reminds us of 
the eiirly stages of the Selachian duct at the stage when it has 
been produced only a little posteriorly from the pronephric r^ion. 
I believe that if the duct is to be compared in Petrcmiyzon 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- 



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MORPHOLOGY OP CYCXLOSTOMATA. 415 

ward growth formed after such a condition is passed, and its 
homologue can not be found anywhere in Petramyzoix, I believe 
that the same can be stated of Amphibia which is, to judge from 
Field's account, very much like Petroniyzon in this respect 
(seep. 401). 

According to van Wyhe and Eabl, the duct in Selachia 
appeal's in the seventh to the tenth (in Rabl's sense) somites of 
embryos with 34-3e5 somites and, when it is later connected with the 
cloacal wall, the connection is found, — as can be inferred from 
fig. lb of VAN Wyhe, — in the thirty-eighth (forty-second of Rabl) 
Rumpfsegment (or further backwards) of P^^istiurus embryos with 
80 (van Wyhe) 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 Rabl) somite, Such a considerable 
prolongation of the duct during this period is not observed in 
Petromyzov}^ and in the two other groups of Oaniota 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 



1) At about this stage (Stage iv), there is no space left beliind the pronephric system 
segraentally 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. 



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416 S. 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 
Rabl. The figures ('96, figs. 9a, 9b, 10a, and 10b) given by Rabl 
to the negation of van Wyiie's view, are from the vertical plane 
of the twenty-second Gesammtsegment of an embryo with the 
03 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 Fetromyzon, 
actually receives cells out of the epiblast, as the figures by van 
Wyhe show ; the other end, which is seen in Rabl's figures, is 
the point of mere contact with the epiblast, along which it is 
shifting backwards. 

If the above comj^arison be coi'rect, the segmental duct in 
Selachia is, except the anterior very small section which is fanned 
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 mesoblastie 
segment, from the distal (ventral) portion of the mesoblastie 
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 



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irORPHOIXXiY OF CYCLOSTOMATA. 417 

was the tertiary stage of changes in the system. iSuch a course 
of the phylogenetic development of the system is, however, no 
other than that advanced by Ruckert ('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 PetrmnyzoTiy 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 distill 
(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 coelomic cavity. In Cyclostomata, such a cavity is 

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418 S. UATTA : 

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 mesoblastic 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 PetroDiyzoUy 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 rise to 
the segmental duct just as in the somites having degenerated tubule. 

From what has been said, it is, I venture to think, no rash 
conclusion to regard the pronephric tubules in Pelromyzon as hainng 
once extended over the body -segments from the branchial region 



1) According to Swaen and Bbacuet, the same fact is seen in TeleosteL 

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MORPHOLOGY OF CYCLOSTOMATA. 419 

to the clodcal party and as having been, in the anterior region^ 
replaced by gills andy in the posterior, converted into the segmental 
duel. 

In the two anterior segments which belong to the branchial 
region, the free ends of the tubules are brought into close contact 
with theepiblast 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 
lis to infer that all the tubules once had each an independent 
external opening until they were secondarily united with one 
another by the intcrsomitic 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 coslome, 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 (b) a small 
distal portion which is unsegmented. The former is differentiated 
into the sclero-myotome and the nephrotome, and the latter forms 
simply the peritoneal linings. 

The pronephric vessels acquire their definitive form in much 
later stages ; when established, they are intcrsomitic in position. 
The posterior part is transformed into a pair of the glomeruli 
of the pronephros. 

Petramyzon has for a long time been looked upon as being 

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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 Pkof. Watase, 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 diflTerent 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 iv. As seen in the 
foregoing description, most of the important processes in the 

l)ZooI. Jalirbucher, Abtheil. fiir Anat. and Ontog., Bd. xni, '99. 

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MORPHOLOGY OF CYCLOSTOMATA. 421 

development of the pronephros and the segmental duct take pla^ 
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. 



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422 s. hatta: 



Literature. 



('79) Balfour : — A Monograph on the Development of Elafimobranch 

Fishes. 

('85) : — A Treatise on Comparative Embryologj'. 

('87) Beard: — On the Origin of the Segmental Duct in Elasmobranchs : 

Anat. Anz., Bd. ii. 
('92) BovERi: — Die Nierencaniilchen des Amphioxus: Zool. Jahrb., Bd. v. 
('88) Brook:— Note on the Epiblastic Origin of the Segmental Duct in 

Teleostean Fishes and in Birds : Proceedings of the Koyal Society of 

Edinburgh, Vol. xiv. 
('98) Dean : — On the Development of the Califomian Hagfish (Bdellostoma 

stouti Lockington): Quart. Journ. Micr. Sc., N. S., Vol. xl. 
('99) : — On the Embryology of Bdellostoma stonti : Festschr. z. 

siebenzigsten Geburtst. v. C. v. Kupffer. 
('82-'90) DoHRN : — Studien zur Urgeschichte der Wirbelthirkerpers : Mittheil. 

a. d. Zool. Stat. z. Neapel. 
('91) Field : — ^The Development of the Pronephros and Segmental Duct in 

Amphibia: — Built. Mus. Comp. Zool. Harv. Coll., Vol. xxi. 
('86) Flemming: — Die ektoblastische Anlage des Urogenitalsystems bei 

Kaninchen : Arch. f. Anat. u. Physiol., Anat. Abtheil, 
('97) Felix : — Die PRiCE'sche Arbeit „ Development of the Excretory 

Organs of a Mixynoid (Bdellostoma stouti)'^ und ihre Bedeutung 

fiir (lie Lehre von der Entwickelung des Harnsystems: Anat. Anz., 

Vol. xni. 
('78) Furbringer: — ^Zur Vergleichenden Anatomic und Entwicklungs- 

geschichte der Excretonisorgane der Vertebraten : Morph. Jahrb., 

Bd. IV. 
('78) Gegesbaur: — Grundriss der Vergleichenden Anatomic, 2. Auflage. 
('75) Goette : — ^Die Entwicklungsgeschichte der Unke, Leipzig. 

('88) :— Ueber die Entwickelung von P. planeri : Zool. Anz., Jahi^. vi. 

('90) : — Entwicklungsgeschichte des Flussneunauges (P. fluviatilis), 

Hanburg und Leipzig. 
('97) Gregory : — Origin of the Pronephric Duct in Selachians: Zool. Bullet 

Vol. I. 
('81) Hatschek: — Studien iiber Entwickelung des Amphioxus: Arb. a. d. 

Zool. Inst. d. Univ. Wien., Tom. iv. 



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MORPHOLOGY OF CYCLOSTOMATA. 423 

('98) Hammar: — Zur Kenntniss der Leberentwickeliing bei Amphioxus: 

Anat. Anz,, Bd, xv. 
(»97) : — XJeber einige Hauptztige der ersten embryonalen Leberent- 

wickelung: Id., Bd. xiv. 
('88) Hatschek : — Ueber den Schichtenbaii von Amphioxns : Id., Bd. iii. 
('91) Hatta :— On the Formation of the Germinal Layers in Petromyzon : 

. Jour. Coll. Sc. Imp. Univ., Jap. Vol, v. 
('97) :— Contributions to the Morphology of Cyclostoraata. I. On 

the Formation of the Heart in Petromyzon : Id., Vol. x. 
('97) : — Prelimipary Note on the Development of the Pronephros in 

Petromyzon: Annot. Zool. Jap., Vol. i. 
('90) HERTwiGr—Lehrbuch der Entwicklung^geschichte des Menschen und 

der Wirbelthiere, 3. Anflage. 
('86) Hoffmann : — Zur Entwicklungsgeschichte der Urogenitalorgane bei den 

Anamnien: Zeitschrlft f. wiss. Zool., Bd. XLiv. 
('89) : — ^Zur Entwicklungsgeschichte der Urogenitalorgane bei den 

Eeptilien: Id., Bd. xlviii. 
('88) V. Kuffer: — Ueber die Entwickelung der Neunaugen: SB. d. k. bayer. 

Akad. Wis. zu Miinchen, Bd. I. (Ref. in Jahresb. Fortschr, Anat. 

Phys., '89). 
('90) : — ^Die Entwickelung von P. planeri: Arch. f. mikrosk. Anat,, 

Bd. XXV. 
('89) Lank ester: — Contributions to the Knowledge of Amphioxus laiiceolatus 

Yarrell : Quart. Journ. Micr. Science, N. S., Vol. xxix. 
('90) Lankester & WiLLEY ! — The Development of the Atrial Chamber of 

Amphioxus: Id., xxxi. 

(*97) Maass : — Ueber die Entwicklungsstadien der Vornierc und Umiere bei 
Myxine: Zool. Jahrb., Bd. x. 

(^88) Martin : — Ueber die Anlage der Urniere beim Kaninchen : Arch. f. 

Anatomie and Entwicklungsgeschichte. 
('87) Mayer, (P.): — Ueber die Entwickelung des Herzens und der grossen 

Gefus88tamme bei Selachiern: Mittheil. Zool. Stat. Neapel, Bd. vii. 
('88) MiTSUKURi : — The Ectoblastic Origin of the Wolffian Duct in Cheolonia 

(Prelim. Notice): Zool. Anz. Jahrg. xi. 

('90) MoLLiER : — Ueber die Entstehung des Urogenitalsystems bei Amphi- 
bien: Arch. Anat. Phys., Anat. Abth., Bd. iii. 

(^75) MuLLER (W.) : — Das Urogenitalsystem des Amphioxus und Cyclostomen: 
Jen. Zeitschr, Naturw., Bd. jx. 



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424 8. HATTA : 

('72 & '73) Oeixacher : — Beitrage znr Entwicklnngsgeflchichto der Knochen- 

fisclie: Zeiisch. f. wis. Zool., Bd. xxii u. Bd. xxiir. 
(70) OwsJANNiSKOW : — ^Die Entwickliingsgeschichte der Fhiasnennaiigen : 

Melanges Biologie, Tom. vii. 
('87) V. Peuenji:— Die ectoblastische Anlage des Urogenitalsystems bei Rana 

esculenta und Lacerta viridis : Zool. Anz., Jahrg. x. 
('97) Price : — Development of the Excretory Organs of a Myxinoid (Bdello- 

stoma stonti Lockington): Zool.'^Jahrb. Bd. x. 
('88) Rabl: — ^Ueber die Differenzirung des Mesoderms: Anat. Anz., Bd. iii. 

('87) : — Theorie des Mesoderms: Morph. Jahrb., Bd. xv. 

('96) : — XJeber die Entwickelung des Urogenitalsystems der Sela- 

chier (zweite Fortsetznng der " Theorie des Mesoderms "): Id., Bd. xxiv. 
('88) Ruckert: — Ueber die Entstehung der Excretionsorgane bei Selachiern: 

Arch. Anat. u. Phys., Anat. Abth. 
('89) : — Zur Entwickelung des Excretionssystems der Selachier; eine 

Erwiederung an Herrn van Wyhe : *Zool. Anz., Jahrg. xii. 
('82) Ryder : — A Contribution to the Embryography of Osseous Fishes with 

Special Reference to the Development of the Cod : Report U. S. 

Fishcomraission. 
('99) Schneider, (Guido) :— Einiges iiber Resorption und Excretion bei Am- 

phioxus lauceolatus : Anat. Auz., Bd. xvi. 
('56) ScHULTZE, (Max) : — Die Entwicklung^eschichte von Petromyzon planeri, 

Haarlem. 
('82) Scott : — Beitrage zur Entwicklungpgeschichte der Petromyzonten. 

Morph. Jahrb., Bd. vii. 
('81) Sedgwick : — On the Early Development of the Anterior Part of the 

Wolffian Duct and Body in the Chick: Quart. Journ. Micr. Sc., N.&, 

Vol. XXI. 
('96) Semon : — Das Excretionssystem der Myxinoiden in seinen Bedentuug 

flir die morphologishe AufTassung (Ref. Jahresber ix. d. Fortsch. d. 

Anat. u. Phys., N. F., Bd. ji). 

('97) :— Das Excretionssystem der Myxinoiden : Anat. Anz., Bd. vin. 

('97) : — ^Vorniere und Urniere, Id.,_Bd. xiii. 

('75) Semper: — Das Urogenitalsystem der Plagiostomen und seine Beden- 

tung fiir das der ilbrigen Wirbelthiere : Arbeiten a. d. zool.-zootom. 

Inst, in Wiirzburg, Bd. ii. 

('87) Shipley:— On Some Points in the Development of Petromyzon 
fluviatilis : Quart. Journ. Jdicr. Sc., N.S., Vol. xxvii. 



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MORPHOLOGY OP CYCtiOStOMAtA. 425 

('95) SoBOTA : — ^Die Entwickelung der Vorniere der Salmoniden : Anat 

Anz., Bd. VI. 
(^84) Spee, (Guaf): — Ueber directe Betheiligung des Ektoderms an der Bildung 

der Uroierenanlage des Meerschweinchens : Arch. f. Anat. und Phys., 

Anat. Abth. 
(^99) SwAEN et Brachet : — Entude sur les premieres phases du development 

des organs derives du mesoblaste chez les poissons teleosteeus : Arch. 

de Biologie, Tom. xvi. 
('90) Weiss: — Excretory Tubules in Amphioxus lanceolatus: Quart. Joutd. 

Micr. So., N. S., Vol. xxx. 
('84) Weldon: — On the Head-kiduey of Bdellostoma, with a Suggestion as 

to the Origin of the Suprarenal Bodies : Quart. Journ. Micr. Sc., N.S., 

Vol. XXIV. 
('83) WiEDERsUEiM : — Lehrbuch der vergleichenden Anatomie der Wirbethiere. 
('91) WiLLEY :— The Later Larval Development of Amphioxus: Quart Jour. 

Micr. Sc., N.S., Vol. xxxi. 
('94) :— Amphioxus and the Ancestry of Vertebrates, New York and 

London. 
('86) V. Wyhe:— Die Betheiligung des Ectoderms an der Entwickelung des 

Vernierenganges : 2kK)l. Anz. Bd. ix. 
('89) :— Ueber der Mesodermsegraente des Rumpfes und die Ent- 
wickelung des Excretionssystems bei Selachiern : Arch. f. mikr. Anat., 

Bd. xxxiii. 
('98) :— Ueber die Betheiligung des Ektoderms an der Bildung 

des Vomierengang bei Selachiern: Verhandl. d. Anat. Gesetsch. 

auf. d. zwolften Vorsamml. in Kiel. 
CS8) ZiEGLER: — Der Ursprung der mesenchymatischen GeSvebe bei den 

Selachiern : Arch f. mikr. Anat., Bd. xxxu. 



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PIRATE XVII. 



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Plate XVn. 



[The magnificatioQ is the same for all figures: (7x2, with the single 
exception of fig. 4 which is Ex 2.] 

a.pn.1-6, Anlagen of pronephric 
tubules from the first to sixth. 
a,8d,, Anlage of s^mental duct 
erf., collecting duct. 
cA., chorda dorsivlis. 
cM^., cutis-layer of myotome, 
rf., dorsal row of mesoblast. 
ep.j epiblast. 
hy., hypoblast. 
?.m,, lateral plate of mesoblast. 



m., median row of mesoblast 
m.p., parietal layer of mesoblast. 
mes., mesoblast. 

mt.Ty //, &c., the first, second, &c. 
myotome. 

mus.y muscle-layer of myotome. 

m.v., visceral layer of mesoblast. 

n., neural cord or canal. 

v., ventral row of mesoblast. 



Fig. 



1. A transverse section through the dorsal region of an embryo inter- 
mediate between Stages i and ii. 

Fig. 2-7. From a series of transverse sections through a younger embiyo 
of Stage II. 

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 embiyo 
of Stage II. 



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T>i:.ATE XVIII. 



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Plate XVnL 



a.pnJ-6, Anlagen of pronephric 
tnbules from the first to sixth. 
o.«/., Anlage of segmental duct. 
cd., collecting duct. 
ch., chorda dorsalis. 
o./>., coelomic projection. 
cut, J cutis-layer of myotome. 
fg., fore-gut. 
ep., epiblast. 
hy,, hypoblast. 

l.m,, lateral plate of mesoblast. 
?/?.jo., parietal layer of mesoblast. 



Qns., mesoblast. 

Qntlj 77, &c., the first, second, &c. 

myotome. 
mvs.y muscle-layer of myotome. 
7n,v,, visceml layer of mesoblast. 

n., neural cord or canal. 

pp.c.f pleuroperitoneal cavity. 

ptJ'6, pronephric tubules from the 
first to sixth. 

sck., subchorda. 

sd.y segmental duct. 



Figs. 30-31. 
Figs. 32-50. 



From the same series as figs. 20-29 of the last plate. 

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 in. 

Fig. 59. A section through an older emboyo of Stage in, the posterior 

continuation of which is shown in the next following plate (figs. 60-63). 



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FI.ATE XIX. 



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Plate XTX. 



ciL, collecting diict. 

ch., chorda dorsalw. 

c.p,, coeloraic projection. 

cm/., cutis-layer of myotome. 

f/., dorsal row of mesoblast. 

ep.y epiblast. 

fg., fore-giit. 

/., Anlage of liver. 

hy,, hypoblast. 

Lm.j lateral plate of mesoblaBt. 
7/1., median row of mesoblaBt 
mch., mesenchymatous cells. 
ms.y mesoblast. 
m.p., parietal layer of mesoblast. 



mt,I, IT, &c., the first, second, &c. 

myotome. 
mus., muscle-layer of myotome. 
m,v., visceral layer of mesoblast 
n., neural canal. 
mt.2'3j nephrostome the second 

and third. 
ppJ-S, peritoneal partition. 
pp.c, pleuroperitoneal cavity. 
ptJ^, pronephric tubules from the 

first to sixth. 
8cJu, subchorda. 
8d., segmental duct 
sg., spinal ganglion. 
17. ventral row of mesoblast. 



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 embiTo 

of Stage III. 
Figs. 77-81. From a series of transverse sections through an embiyo of 

Stage IV ; hence the body of embryo in the prasent stage is twisted, 

the sections pass through unavoidably oblique planes. 



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I^LATE XX. 



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Plate 



a,8(l,y Anlage of segmental duct. 

hp,j blastopore. 

hrg., branchial region. 

cc, cloacal cavity. 

cd,y collecting duct, 

ch.j chorda dorsalis. 

c,dt\, diverticulum of cloacal 

cavity. 
co.sd.y cloacal opening of seg- 

montal duct, 
c.p., ccjflomic projection. 
dLbp., dorsal lip of blastopore. 
/(/., fore-gut. 
ep,, epiblast. 
gc, genital cells. 
hy., hypoblast. 
int,y intestine. 
L, Anlage of liver. 



' l.m.j lateral plate of mesoblast. 
I mch,, mesenchymatous cells. 
I ms,f mesoblast. 

m.p., parietal layer of mesoblast. 

mt.l Ily &c., the first, second, &c. 
myotome. 

m.v.f visceral layer of mesoblast. 
r?., neural canal. 
7W»^5, fifth nephrostome. 
per{t,j peritoneal membrane. 
pp.1'3, peritoneal partition. 
pp.c,j pleuroperitoneal cavity. 

ptj'6f pronephric tubules from 
the first to sixth. 

sch,, subchorda. 
sd.j segmental duct. 

yc.f yolk-cells. 



Figs. 82-89. Sections from the same series as fig. 81, lying posteriot 
to it The section shown in fig. 87 passes throtigh 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 axi:^ 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 blasto[)ore, and in fig. 91, the 
upper (dorsal) portion of it, is cut through. 

Fig<. 92-9G. From u series of transverse sections through 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 lx>dy of which has been straightened l>efore cut throiigh. 



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Plate XXL 



fwv.y anterior cardinal vein. 

at*,, auditory pit. 

bp,, ventral Ii[) of blastopore. 

hrg.j branchial' region. 

6^*, blood space. 

rt/,^ collecting duct 

tf^f chorda dorsalis. 

ve,, cloacal cavity. 

t\dv., diverticulum of cloacal 
cavity. 

vo,sfLj cloacal opening of seg- 
mental duct. 

vtt\^ wall of cloaca. 

t^jKf epi blast. 

f/Kf glomerulus of pronephros. 

/., dorsal fin. 

ffj., fore-gut. 

h,f heart. 

hy\f hypoblast. 

/.J liver, or Anlage of liver. 



/.w., lateral plate of mesoblast. 
mch.y mesenchymatous cells. 
77i,p., parietal layer of mesoblast. 
perit.y peritoneal membrane. 
wt.I, Ily &c., the first, second 

myotome, &c. 
m.v., visceral layer of mesoblast. 
n., neural canal. 
n8t2-6, nephrostomes from the 

second to sixth. 
pp.c.j pleuroperitoneal cavity. 
pL 2'5y pronephric tubules from 

the first to fifth. 
r.m,, radix of mesentery. 
scluy subchorda. 
sd.j segmental duct. 
tM, tract of aorta. 
t,ac. tract of anterior cardinal 

vein. 
tr.a, truncus arteriosus. 



¥\^^, 1)8-106. From a series of transverse sections through an embryo of 



Stage V. 



Figs, 107-110. From a series of transverse sections through an embryo 

of Stage VI. 
Pig. 111. Transverse section through the cloacal region of an older embryo 

nf Stage VI. 
Fig8. 112-114. A series of sagittal sections through a younger embryo in 

8tage IV. 
Fig. 1 15. A frontal section through an embryo a little more advanced 

tlum that of Stage vi. 



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i 



Beitrfige zur Wachstumsgeschichte der 
Bambusgewachse. 



Von 



K, Shibatai RigakushL 
MU Tajeln XXILXXIV. 

I. Einleitmig. 

Unsere Kenntnisse iiber Bau und Lebensweise der Bambus- 
gewacbse waren bisher sehr mangelhaft gewesen, obwohl in neueren 
^iten einzelne merkwiirdige Erscheinungen auf dem Gebiete der 
Physiologie dieser eigenartigen Baumgraser durch interessante Beo- 
bachtungen^) einiger die Tropen besuchender Botaniker zu Tage 
gefordert warden. Bekanntlich gehoren die meisten Bambuseen 
zu warmeren Gegenden der alten und neuen Welt, mit Ausnahme 
von einigen kalterem Klima angepassten Formen, wie z. B. 
Bambusa Kurilensis, die in einer nordlichen Insel Japan's 
bei 46^ n. B. gedeiht. Unser Land besitzt eine Keiiie von 
Bambusformen, welche unserer Pflanzenphysiognomik ein charac- 



1) G. Kraus, PIiTsiologisches aas den Tropen. L Langenwachstum der Bambusrohre. 
Ann. d. Jard. Bot. d. Baitenzorg. Vol. KIT, p. 196. 

H. Mol isch, Uber das Bin ten tropischer Holzgewadise im Zastand ToUiger Belaubun^. 
Amu d. Jard Bot. d. Baitenzorg. 1898. 2 tes Sappl. p. 23, 



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428 K. SHIBATA : 

teristisches Ausselien verleihen. Einige hochwiichsige Formen aus 
der Gattnng Phyllostdchys sind bei uns iiberall haufig cultiviert, 
hauptsachlich fiir die raannigfaltigste Verwendbarkeit der Rohre 
und auch fiir ihre Fruhjahrsschosslinge, die ein beliebtes Gemiise 
darbieten, wahrend andere Arten aus Aru7idinaria und Bamiusa 
als Zierpflanzen in unseren Garten gemein sind. 

Nun stellte ich es mir hier als Aufgabe, in erster Linie die 
Natur und das Verhalten der wichtigen BaustoflTe wahrend der 
verschiedenen Vegetationsperioden zu studieren, da mir besonders 
die ungemein rasche Entwicklung der Sehosslinge etwas interes- 
santes in Bezug auf Stoffwanderungs- und Stoffumwandlungsvor- 
gange darzubieten schien, oder in anderen Worten die 
Wachsturasgeseliichte der Baumgraser mit BeriicksiclitiguDg der 
Bauverhaltnisse naher zu verfolgen. 

Die friiheren Angaben fiber die Systematik, Verbreitung und 
aussere Lebensweise der Bambusgewaebse haben eine Zusammen- 
stellung in einem Werk Schroter's^) erfahren, und es schien mir 
uberflussig dieselbe hier wiederzugeben. Was die Physiologie der 
Bambusgewaebse anbetrifft, besitzen wir abgesehen von alteren 
Beobaehlungen fiber das Wachstum der Sehosslinge nur die ein- 
gangs erwiihnten Arbeiten von Kraus und Molisch. Uberdie io 
Bambuspflanzen vorkomraenden Stoffe besitzen wir ebenfalls spar- 
liche Angaben. Cohn^) studierte „Tabaschir** in seiner klassiscben 
Arbeit. Kozai^) stellte chemische Untersuehungen fiber die stick- 
stoflfhaltigen Bestandtheile des Schosslings von Phyllostachys mitts 



1)C. Schroter, Der Bambus und seine Bedeutung als Kutzpflunise. Basel, 1895. 
Yergleiche ferner: 

E. Ilackel, Banibusacese. Engler's Die'naturlichen Pflanzenfamilien. II, 2. p. 89. 

A. et C. Riviere, Les Bambous. Vegetation, culture et multiplication. 1878. 

2)F. Cohn, ijber Tabaschir. Beitrage z. Biol. d. Pflanzen. Bd. IV, p. 365. 

3)Y. Kozai, On the nitrogenous non-albuminous Constituents of Bamboo shoots. 
Pulletin of the College of Agriculture, Vol. I, Nr. 7, 



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WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 429 

an, und dabei fand er das Vorkommen von Tyrosin und Asparagin 
neben kleineren Mengen der Nucleinbasen. Die Angaben iiber 
die Bauverhaltnisse der Bambusgewachse finden wir in Arbeiten 
von Schwendener, Haberlandt, Strasburger, Hohenauer, 
Guntz, Boss und Magnus. Naheres iiber die Litteratur wird 
noch an geeigneten Stellen Erwahnung finden. 

Die vorliegenden Studien wurden im Laufe des academisehen 
Jahres 1898-1899 im botanischen Institute der Kaiserliclien 
Universitat zu Tokio ausgefiihrt. An dieser Stelle spreche icli 
meinem hoehverehrten Lehrer Herrn Prof. Dr. Miyoshi meinen 
warmsten Dank fiir seine vielfache Belehrung und Anregung 
au8, Es ist mir auch eine angenehme Pflicht Herrn Prof. Dr. 
Matsumura fiir seine giitige Unterstiitzung bier meinen herz- 
lichsten Dank auszudriicken. 



n. Untersuchungsmaterial und Methodisches. 

Als Untersuchungsobjecte dienten mir die im botanischen 
Charten der Universitat cultivierten Bambusarten, insbesondere 
Phyllostachys mitisy 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 
Schosslinge der obengenannten Art wurde von der ersten Anlage 
bis zum mehrere Meter hohen Halmzustand verfolgt, 

Auch die Schosslinge folgender Arten wurden zum Ver- 
gleichungszweck untersucht : 

im April austreibende — Banibusa j)almala, Bambicsa Veitchii; 

im Mai austreibende — Fhylloslachys puberula, Arundinaria 
japonica ; 

im Juni austreibende — Phylh^tachys bainbusoides ; 

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430 Js:, sfiiBATA: 

im J u 1 i- A ug u s t austreibende — AruncUnaria Simonif Arundi- 
naria Hindsii; 

im October- November austreibende — Arundinaria Maim- 
murcBj Arundinaria quadran^ularis^ Arundinaria Hindsii 
var graminea. 

Die Entwicklung der Rhizomspitze wurde bei folgenden 
Arten im Herbst untersucht: Phyllostachys mitiSf Fhyllostachys 
barabusoides. 

Fiir andere Arten, die ich in meiner Untersuchung gezogen 
habe, verweise ich auf das am Ende dieser Arbeit beige£agte 
Artenverzeichniss. 

Um die Umwandlung und Wanderung der Stoflfe in Reserve- 
stoffbehaltern und in wachsenden Theilen zu verfolgen, 
bediente ich mich unter nothigen Cautelen der ublichen micro- 
chemischen Methoden. Dariiber sei hier folgendes bemerkt : 

Stfirke. Meyer 'sche Chloralhydratjodlosung^) wurde mit 
Vortheil benutzt. 

Olyoose. (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 ublichen Methoden 
auch zu unserem Zweck vollig ausreichten, habe ich noch Sicher- 
heits wegen eine andere Reaction ausgefuhrt. Ich habe namlich 
die Wasserausziige von jungen Hal men , Wurzeln, Rhizomen und 
Scheideblattern und auch den Blutungssaft mit essigsaurem 
Phenylhydrazin erwarmt, und dabei erhielt ich stets charac- 
teristische gelbe Nadelkrystalle von Glucosazon. 



l)Vergl. Strasburger, Botanisches Practicutn. III. AuBage. p. 277. 

2) A. Meyer, Microchemische Beaction zum Nachweis der reducirenden Zuckerarten. 
Ber. d. D. R G. 1885. p. 332. 

3) A. Zimmermann, Die botanische Microtechnik. p. 75. 



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WACHSTUMSGESCfllCHTE D. BAMBUSGEWAECHSE. 431 

Rohrzaoker. Die Unzuverlassigkeit der bekannten Sachs'- 
schen Methode wurde vielfach von Autoren betont. Ich habe 
nur ausnahmsweise diese Reaction benutzt, Tirahrend in den meisten 
Fallen ich mich der neulich von Hoffmeister^) aufgestellten 
Invertinmethode bediente.^) 

Gerbstoffe. Die Eisensalzlosungen, insbesondere die atherische 
Losung des Eisenchlorids, nnd anch die Kaliumbichromatlosung 
wurden angewandt 

Fette. Alkannatinctur and l-procentige Osmiumsanrelosung 
wurden benutzt. 

Aqparagin. Die bekannte Borodin'sche Methode') hat sich 
zweckenlsprechend erwiesen. Zur Erkennung der erhaltenen 
Krystalle als Asparagin diente niir hauptsachlich die Winkel- 
messung. Vielfach wurde das Borodin'sche Verfahren mit 
gesattigter Asparaginlosung angewandt. Femer diente mir 
Diphenylamin-Schwefelsaure zur Unterscheidung von Asparagin 
und Salpeter. 

Tjnrosin. Die nach Borodin'scher Methode behandelten 
Schnitte ergaben eine reichliche Ausscheidung von eigenthiimlichen 
Krystallen, die ohne Schwierigkeit mit Tyrosin identificiert 



1) G. Hoifmeister, Uber den microchemischen Nachweis von Bohrzucker in pflanz- 
lichen G«weben. Jahrb. f. wish. Bot. Bd. XXXI, p. 688. 

2) Die von den „ Ebisu "-Brauerei bezogene Hefe-Beinkultur wurde zar Darstellnng von 
Invertin yerwendet, dabei habe ich wie folgt verfahren: Die He fe masse wurde mittelst 
Filtration von der Kulturfliissigkeit befreit und nach wiederholtem Auswaschen rait Wasser zam 
dicken Brei angeriihrt. Der Hefebrei kam nach dem Verreiben im Morser in den Thermostat 
bei 50^ C, in welchem er fiir 10-12 Stunden gelassen warde. Hiemach wurde die abfiltrierte 
Fluasigkeit mit 90^ Alcohol versetzt, und der dabei entstandene voluminose Niederschlag 
auf flUrirpapier gesammelt, welcher, nach wiederholtem Auswaschen mit 90^ Alcohol und 
dann mit abeolutem Alcohol auf Schwefelsiinre getrocknet wurde. Die wassrige Losung der 
erhaltenen weiseen kreideartigen Substanz, die allein niemals Fehling' sche Losung reduciert, 
leigte ein energisches Inversionsvermogen. Bei wiederholten Versuchen habe ich 
femer in keinem fall die Beimengung von diastatischen und ccHulosespaltenden Enzymen 
gefunden. Weitere Verfahren mit den Schnitien genau nach Hoffmeister. 

3} A. Zimmermann, Die botanische Microtechnik. p. 80. 



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432 K. SHIBATA : 

warden*) (Fig. 58). Belzung'sche Glycerin-Methode^) wurde 
auch mit Vortheil benutzt, wobei sich schone Nadelbiischel in 
den Zellen bilden (Fig. 61). Ich habe ferner zur Erkennung der 
Vertbeilung des Tyrosine in eiweissarmen Gewebetheilen 
Mil Ion's Reagens benutzt, und dabei warden die tyrosinreichen 
Zellen schnell blutroth gefarbt. Die Farbung bleibt nach dem 
Aaslaugen der zuvor mit absolutem Alcohol behandelten Schnitte 
mit dem warmen Wasser fiir 10-20 Minuten so gut wie ganzlich 
aus. Baher kann diese Bothfarbang niemals von EiweissstoSen 
berruhren. 

Biweiis. Biaretreaction und Raspail's Reaction wurdeu 
vornehmlich benutzt. Mill on 's Reagens kam zur Anwenduog 
erst nach ^eux Ausziehen von Tyrosin in oben beschriebena 
Weise. 

Hineralstoffe. Die von Schimper^) empfohlenen Reagendea 
warden verwendet. Die Controllversuche wurden ofters ausge- 



1) Dies geschah aus folgenden Griinden: 

1. Die Oestall der Krystalle, Die feine Nadelbiischel in dendritischer Gestalt oder 

Doppelpinselform bietet ganz dasaelbe Aaasehen wie reines Tyrcmn. 

2. Das optische Vcrhalien. Im durcbfallenden Licht erscbeinen die Krystalle briian- 

licb und im aufTallenden Licbt weisslich seidenglanzend. Im polarisierten Licht 
zeigen die Krystalle starke Doppelbr^bang. 

3. Die LiJdichkeiUoerhdUnisse. Die Krjstalle sind unUislich in kaltem Wasser, aber 

loslich in kochendem Wasser, Ammoniak and verdunnter Salsaore. Ferner 
sind die Krystalle nnloslich in heissgesattigter Tyrosinlosung. 

4. Das Verkatten beim ErhiUen, Wenn man den mit Tyroeinkrystallen besetiten 

Objecttrager auf der Flamme erhitzt, bis nebenbei vorhandene Asparaginkrysttlle 
sich lu brannen Schaomen verwandeln (ca. 200*^ C), so sieht man, das die 
Nadelkrystalle ganz unverandert bleiben. 

5. Dob VerhaUen gegen MUM$ Beagens, Die Krystalle losen sich im Millon's 

Reagens mit einer prachtvoll rothen Farbung der umgebenden Flussigkeit. 

Die oben angefuhrten Merkmale reichen aus, die Krystalle microchemifich als Tyroein n 
erkennen. 

2)Belzang, Becherches chimique sur la Germination. Ann. d. Sc. nat. Bot.S^. VIL 
T. 16, p. 209. 

3) A. F. W. Schimper, Zur Frage der Assimilation der Minerahalzd dorch die 
grtine Pflanze. Flora. Bd. 73. 1890. p. 210. 



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WACHSTUMSGESCHICHTE P. BAMBUESQWAEGHSE. 433 

fiihrt, urn die Reinheit der Reagentien za priifen. Die raicro- 
chemisehen Reactionen warden sowohl an frischen Schnitten als 
an auf Deckglasern gegliihten Aschen vorgenommen. 



m. Die Bauverbaltnisse. 

Die Bauverbaltnisse der Bambusen sind bisber sparlicb 
und nur gelegentlich Gegenstand der auatomiscben Forschung 
geworden. Strasburger^) bat den Bau des Gefassbiindels von 
Bambusa vulgaris kurz gescbildert. Das macbtig entwickelte 
Bastgewebe in Bambusbalmen wurde vielfacb von Scbwendener^) 
Haberlandt^) u. A. erwabnt. Ross^) bemerkte den anomalen Bau 
der Wurzeln. Die Betracbtungen iiber die Blattstructur finden 
wir in den Arbeiten von Giintz^), Magnus^), Haberlandt^) 
und Schwendener®). Ubrigens liegen uns nocb einige kurze 
Angaben von Hobenauer^) und Mobius^^) vor. Nun scbien 
es mir wiinscbenswertb die Bauverbaltnisse der Vegetationsorgane 
der Bambusgewacbse einem genaueren Studium zu unterwerfen, 
damit fiir die pbysiologiscbe Forscbung dieser interessanten Pflan- 

zengruppe eine festere Grundlage gescbaffen werde. Meine 



1) Strasburger, Uber d. Bau o. Verrichtongen d. Leitungsbahnen. 1891. p. 863. | 

2)Schwendenery Das mechanische Princip in anat. Baa d. Monocotjlen. p. 65. 
3)Haberlandt, Entwicklnngsgeschichte des mech. Q^webesjstems d. Pflanzen. p. 23. 
4) Boss, Beitrage z. Anatomie d. abnorm. Monocotylenwurzeln. Ber. d. Deutsch. Bot. 
Qwells-XBd. I, p. 338. 

5)Guntz, Onters. ub. d. anat. Stnictur d. Gramineenblatter. p. 37, 41, 44, 48, 64 etc. 

6) Magnus, Einfalzungen d. Zellmembran. (Just's Jahresber. d. Bot. I, p. 367). 

7) Haberland|t,;,Verg1. Anat. d.^ssim. Gkwebesystems d. Pflanzen. Jahrb. f. wiss. Bot. \ 
Bd. XIII, p. 100. ' t 

8)Schwendener, Die Mestomscheide der Gramineenblatter. Ges. Bot. Mitt. Bd. 11, | 

p. 178. \ 

9)Hohenauer, Vergl. anat. Unters. ub. d. Bau d. Stamn^ bei d. Gramincen.' p. 556, ^ 

10) Mobius, lib. d. eigent. Bltthen ton BarabUstt tulgaris. (Ref. in Bot Centrftlbl. 1899, } 

Nr. 51, p. 479). - •• ^ 



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434 K. SHIBATA : 

diesbezugliche Untersuchungen erstreckten sich auf sieben und 
zwanzig vorwiegend einheimische Fonnen, die sich in die vier 
Gattungen von Phyllostachys^ Arundinaria^ Bamhusa und 
Dendrocahmus vertheilen. Die wesentlichen Ergebnisse will ich 
in folgenden Zeilen kurz darziistellen yersuchen. 

Das Rhizom. 

Die Rhizome^) von Phyllostachys- und Arundi7Uiria^kr\j^ri 
sind bekanntlich kurz gegliederte horizontal verlaufende Stengel- 
gebilde mit einer rundlichen oder rundlich-ovalen Querschnitt- 
form. Die internodiale Markhohle ist stets stark reduciert, meist 
nur einige mm breit und kommt nicht selten zum ganzlichen Ver- 
scbwinden. Wir wollen zunachst beispielweise ein Rhizominter- 
nodium von Phyllostdchya mitts ins Auge fassen. 

Nachst unter der Epidermis kommt ein 1-3 schichtiger 
Ring von den englumigen langgestreckten sklerotischen Paren- 
chymzellen, deren Querwande ofters etwas schief gestellt sind. 
Das darinnen liegende 20-25 schichtige bundelfreie Parenchym 
wird als die primare Rinde^) aufgefasst, Es geht ohne scharfe 
Greuze zum Grundgewebe des Ceutralcylinders fiber, in welchem 
in ublicher Weise die collateral gebauten Gefassbundel zerstreut 
liegen. Sammtliche Parenchymzellen sind verholzt und mit 
zahlreichen ovalen Ttipfeln versehen. In diesem Gefassbundel 
erblickt man ein typisch gebautes Gramineenbundel, Die grosse 
Lumen weite der Siebrohren ist dabei hochst auffallend*) ; es wurde 
oftmals einen Durchmesser von 0.15 mm erreicht^), wahrend 

1) Vergl. A. et C. Bividre, Les Bambous. p. 68, p. 236. 
2)Falkenberg, Vergl. Unters. iib. d. Vegetationsorg. d. Monoootjlen. p. 163. 
3)St]ra9burge^, I^eitungsbahnen. p. 363. 

4) Die Angaben iiber die Lmnenwetto^ d^ Siebrohren einiger Pflanzen findet man btt 
I^ecomte (Ann, d. |3c. nat, Kptr S$r. VJI, T. X, p. 242). 



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WACHSTUMSGESCHICHTE B. BAMBUSGEWAECHSfi. 435 

die grosste Parenchymzelle 0.09 mm und die Greleitzellen meist 
nur 0.005 mm weit sind. Die beiden seitlichen Gefasse, deren 
maximale Lumenweite 0.2-0.3 mm betragt, communicieren mit den 
einschichtigen Belegzellen durch regelraassig angereihte quer- 
gestreckte TupfeL Die stark entwickelten Bastbelege um die 
Geiassbundel sind seitlich an der Grenze zwischen Hadrom und 
Leptom und oft auch unterhalb der seitlichen Gei^isse unter- 
brochen. Dadurch kommen ein oder zwei Paar DurchladBStellen 
zu Stande^ die, wie zuerst Scliwendener^) bemerkt hat, einen 
Stoffaustausch zwischen Bundelelementen und Grundgewebe 
ermoglichen. 

Wenn man die Querschnittbilder der Khizominternodien der 
ubrigen Arten in Betracht zieht, so lassen sich unter denselben 
folgende drei Typen unterscheiden, namlich : 

Erster Typus. Die aussersten Biindel, welche direct an 
die Rinde grenzen, stehen voUkommen isoliert von einander. 
Hierher gehoren PhyUostachys mitisy Phyllosidchys hamhusoides^ 
Phyllostdchys puherula und Arundinaria Narihira. 

Zweiter Typus. Die Bastbelege der aussersten Biindel 
yerschmelzen sich haufig unter einander und auch mit den 
Baststrangen zu unregelmassigen Bastbandern. Als Beispiele 
dienen Arundinaria japonica^ Bambusa borealis^ Arundinaria 
Tootsikf A. Simoni, A, Hindsii 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 Mestombiindel innen angelehnt sind, befindet sich bei 



l)Schwendener, Das mechanische Princip. p. 107. 

2) Vergl. Haberlandt, Entwicklungsgeschichte des mechan. QewebesTsteros. p. 28.^ 
PhTsiologische HaDzenanatomie. p. 157. 



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436 K. SHIBATA : 

Banibusa palmata, B, Veitchiij B. paniculata, B. nippcnicay 
B. ramosa, B. nana, Arundinaria quadrungularisj A. Matsumum^ 
A. variabilis^ Arundinaria pygmcea und ferner Phyllostachys 
Kumasasa. (Fig. 2). 

Der Bastring der letzterwahnten Arten, welch er je nach 
Species verschieden stark ausgebildet ist, geht entwicklongs- 
geschichtlich aus einem entsprechend continuirlichen Cambium- 
ring*) hervor. Schwendener sagt^) : „Fur Bambuseen ist die 
Querschnittform des mechanischen Systems, wie ich sie friiher 
beschrieben babe, characteristisch genug, um jede nahere 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 primaren 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 Blattspurstrangen Platz za machen. 
Ferner ist es als die Regel hervorzuheben, dass innerhalb des 
Knotens der Bastbeleg des Mestombiindels eine bedeutende Re- 
duction erfahrt, und sich meist nur auf eine diinne Sichel um 
das Leptora beschrankt. Die sammtlichen Elemente des Bundels 
sind hier kurzgliedrig, und die Seitenwande der Siebrohren sind mit 
den ausserordentlich zahlreichen Siebtiipfeln versehen. Bekannt- 

l)Haberlandt, Entwicklangsgeschichte. p. 28. 

2) Schwendener, Die Mestomacheide der Gramineenblatter. p. 183. 

3) Die mittleren Durchmesser der Rhizome dieser drei Typen stehen ungefahr im Ver- 
haltnisse 6:3:1. Die Ausbildung der Bastplatte resp. des Bastrings in Bhizomen entspricht 
wohl den rait der Diinnheit steigenden Aufforderungen fiir die Biegnngsfestigkeit; Jedenfalls 
gehort hier die Anordnung des mechanischen Systems in Jlhizomen nipht zam sogenaimten 
taxonomischen Merkmalen. 



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WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 437 

Kch findet sich in dem Knoten die Vereinigung der Blattspur- 
strange unter einander und mit den Achselknospenbiindeln statt^). 
Die liier eintretenden Knospenbiindel verbreiten sich in dem 
Knotengewebe nach alien Richtuugen bin. Die Gefasse der Knos- 
penbundel setzen sich in iiblicher Weise unter starker Kriimmung 
an die der Blattspuren an^). Dennoch verdient die Art und 
Weise, wie der Ubergang des Leptoms erfolgt, eine besondere 
Beachtung. Das Leptom des Knospenbiindels 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 iiblichen Ban des Leptoms in so fern ab, dass 
sie die Differenzierung ihrer Elemente in Siebrohren und Geleit- 
zellen nicht mehr aufweist, sondern aus lauter gleichartigen 
feinen ca, 5-6 // breiten cambiformartigen Elementen^) zusam- 
mengesetzt ist (Fig. 4 u, 8), und folglich im Querschnittbild ein 
regelmassiges englumiges Maschenwerk darstellt (Fig. 5 u. 7). 
Die Anordnung dieser feinen cambiformartigen Elemente bietet 
eine grosse Eigenthiimlichkeit dar. Die etwas schrag gestellten 
Endflachen der seitlich an einander stossenden Elemente scheinen 
ungej^hr auf derselben Querebene zu liegen, und demgemass stellt 
der ganze spindelformige Theil im Langsschnitt ein der Lange nach 
an einander angereihtes meist5-7 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) iibersclireitet die Breite 
kaam 9 i«" So weit ich unterrichtet bin, wurde derartige Slructur bisher in keiner anderen 
Pflanzen beobachtet Zum Beispiel finden wir keine diesbeziigliche Angabe bei verschiedenen 
Monocotylen, die yon Falkenberg {loe, cit.) and Strasburger {loc» cii.) grundlich 
nntersaclit wurden. Ob sie anch bei anderen Pflanzen vorkommt muss dcshalb zur Zeit 
dahin^estellt bleiben. 



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4S8 i,. SfllBAtA! 

oder 2 Etagen dieses Stockwerks sind sehr langgestreckt und 
besitzen fein undulierte Seitenwaade mit zahlreichen grossen 
ovalen tiipfelartig verdiinnten Stellen, die zum Beispiel bei 
FhyUostachys mitis eiue Weite von 5x3 ;/ erreichen (Fig. 6). In 
einem Eade dieses spindelformigen Theils vermitteln die elwas 
breiteren Elemente, — die sich zu 4-5 je einer feinbetupfelten 
Endflache der Siebrohren und einzeln auch den Geleitzellen 
anschlisseo, — den Uebergang zum normal gebauten Leptom dee 
Knospenbiindels (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 gewohnlich den spindelformigen Theii 
vom Grundparenchym ab. Die ZellwandbeschaflTenheit der 
spindelartigen Theile weicht kaum von der des Leptoms ab ; sie 
zeigen namlich ebenso starke Cellulose-Reaction mit Chlorzinkjod 
oder Schwefelsaure-Jody und sie werden auch mit Anilinblaa, 
Congoroth u.a.y im nahezu gleichen Farbenton wie Siebrohren 
gefarbt. In spaterem Alter tritt jedoch oft eine Spur Holzreaction 
in den Elementen der oben erwahnten mittleren Etagen ein. Was 
den Plasmagehalt dieser Theile anbetriflft, so scheint es nur auf 
einen zarten Wandbeleg beschrankt zu sein, wie es bei Siebrohren 
stets der Fall ist. In den ersten Entwicklungszuslanden habe ich 
constatiert, dass diese Anschwellung aus den entsprechend ver- 
mehrten Langstheilungen der procambialen Zellen an der betreffen- 
den S telle hervorgeht. In solchen friiheren Stadien zeichnet sich 
diese Anschwellung durch besonders reichlichen Plasmagehalt 
und aufiallend grosse Zellkerne aus, wie es Fig. 9 zeigt. 

Derartige spindelforraige Leptomanschwellungen in Knospen- 
biindeln habe ich regelmassig in Rhizomknoten sammtlicher von 
mir untersuchter Arten gefunden, aber man findet sie am starksten 



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WACHSTUMSGESCHlCfiTE D, BAMBUSGEWAECHSE. 430 

ausgebildet in den Rhizomknoten der Phyllostachys-ATteny wobei 

ihre Querschnittgrosse sogar einem grossen Mestombiindel nahe- 

kommt. Da die an Khizombundel sich ansetzenden Achselknos- 

peostrange in ihrer Gesammtheit ein physiologisches Analogon des 

kaustorial Saugorgans darstellen^ so ist es nicht unmoglich^ dass 

dieses Gebilde in dem Sinne ausgebildet ist, dass es eine specifisch 

absorbierende Wirkung auf die Siebrohren der Mutterrhizome 

auszuiiben yermag. Allerdings besitzt es in seinen anatomischen 

Merkmalen vieles gemein mit den iiblichett Absorptionsgeweben^j. 

So laDge aber die Stoflftransportmechanik im Leptome noch nicht 

in alien Hinsichten aufgeklart ist'^), moge die nahere ErorteruDg 

der physiologischen Vorgange, die sich in diesem abweichend 

gebauten Leptomtheile abspielen, auf eine kiinftige Gelegenheit 

verschoben werden. 



Deb Halm. 

Die Bambushalme') sind bekanntlich mit den hohlen Inter- 
nodien versehen, die bei Phylloslachys miUs oft eine ansehnliche 
Dicke von 20 cm erreichen. 

Die primare Rinde ist stets weit schwacher entwickelt als 
in dem Rhizome; diese Verhaltnisse, wie sie schon von Falken- 
berg*) und Rothert^) fur andere Pflanzen nachgewiesen wurden, 
gehen noch aus den folgenden Beispielen deutlich hervor : 



l)HaberIandt, PhjBioIogische Pflanzenanatomie. p. 186. 

2)Czapek, liber d. Leitungswege d. organischen Baostofie in Pflanzenkorper. p. 24; 
Lecomte, Etude da Liber des Angiospermei Ann. d. Sc nat S^r. VII. T. X, p. 303. 

3) Vergl. Bivi^re, Les Bambous. p. 134. 

4) Falkenberg, /.c p. 134. 

5)Rothert, Vergl. anat. Unters. iib. d. Differenzen im prim. Ban d. Stengel u. Bhizome. 
p. »2. 



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440 



K. SBIBATA 





HALM 


BHIZOM 


des 
Central- 
cylinders 


Dlcke 

dor 

Binde 


Q« 


DuTchmeaer 

des 

Cejtral- 

cyllnders 


Dloke 

der 

Blnde 


Q* 


Phyllostachya mitts 


130.0 


0.315 


412 


23.8 


0.728 


33 


PhyUostachya bambusoidee 


34.0 


0.059 


575 


20.0 


0.611 


33 


Phyllostachya puherula 


56.0 


0.049 


1142 


22.0 


0.933 


23 


Phyllostachya Kumasasa 


2.7 


0.023 


120 


5.6 


0.780 


7 


Arundinaria Simoni 


140 


0.049 


285 


8.0 


0.494 


16 


AruTidinaria japonica 


17.0 


0.050 


340 


8.4 


0.286 


29 


Arundinaria Eindsii 


22.0 


0.059 


373 


18.0 


0.364 


49 


Arundinaria quadrangularis 


23.0 


0.045 


511 


8.0 


0.260 


31 


Arundinaria Matsumurce 


2.5 


0.018 


139 


2.9 


0. 20 


15 


Arundinaria pygmcea 


2.3 


0.036 


64 


4.5 


0.325 


14 


Arundinaria Narihira 


14.0 


0.049 


285 


21.0 


0.468 


45 


Bamhuaa borealis 


5.5 


0.045 


122 


6.5 


0.212 


31 


Bambusa palmata 


10.0 


0.063 


159 


7.1 


0.624 


11 


Bambusa Veitchii 


4.5 


0.023 


200 


2.9 


0.143 


20 



*Q.=Da8 Verhaltniss des ersteren zur letzteren. 

Die aussersten 1-2 Schichten Rindenparenchymzellen sind 
oft sklerotisch verdickt (Fig, 12) uud unterscheiden sich von den 
iibrigen nur durch eine grossere Lange ; folglich haben wir 
hierbei keineswegs mit eineni Bastring, wie Haberlandt einst 
annahm/) zu thun. Die Bastbelege der peripherisclien Gefass- 
biindel und die dazwischen liegenden Baststrange verschmelzen 
mit einander zu linregelmassigen Bastbandern, besonders haufig 
in diinneren Halmen von Arundinaria Matsumurce^ A. pygnuza 
etc. Dennoch begegnet man hier in keinem Fall dem echten 
Bastringe. 



1) Haberlandt, Eat wicklung^gesckiclite. p< 23. 



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WACHSTTJMSaESCHICHTE D. BAMBUSGEWAECHSE. 441 

Die zuerst von Schwendener^) bei einigen BambusMvien 
entdeckte. eigenthximliche Parenchymlamelle, die quer in dera 
innenseitigen Bastbelege inseriert ist, habe ich auch in den 
Halmen aller echter Bambusaa^Ti^n {B. vulgaHs, B^ nana und B. 
8teno8tachya)f Dendrocalamus lalifiorus und bei 2 Arwndinaria^ 
arten {A. Hindm^ A. quadrangularis) aufgefunden. Uberdies 
habe ich die Falle beobachtet, dass die Lamelle nur an einer 
Seite in das Grundparenchym iibergeht, und dass sogar das Paren- 
chym in der Mitte des Beleges allseitig von Bastzellen umschlossen 
liegt (Fig. 17 u, 18), Nach den letzterwahnten Thatsachen 
erscheint es a priori sehr wahrscheinlich, dass dieses Parenchym- 
gewebe erst nachtraglich aus einem Theil des Procambiums des 
Bastbeleges hervorgeht, wie es Haberlandt^) schon vermutbet hat. 

In der That konnte ich in einem Procambialstrang in jun- 
gen Internodien zuerst nichts von dieser Parenchymlamelle 
erkennen. Sie differenziert sich erst spater aus einer Strang- 
anlage, in welcher alle Forraelemente schon fertig angelegt sind, 
derart, dass die langgestreckten Procambialzellen in betreflfender 
Stelle successive Quertheilungen erfahren und zum Epen um- 
gewandelt werden (Fig. 21 u. 22). Die feinkornige Starke, die 
spater dem Zucker Platz raacht, tritt sogleich in diesem Gewebe 
auf (fig. 20 u. 22) und bleibt in demselben wahrend der weiteren 
Ausbildung des Stranggewebes. In dieser Weise dient die 
Parenchymlamelle den dem Mestom unmittelbar anliegenden 
Bastzellen als ein Speicherungsort der notigen Baustoflfe. Die 
durch diese Parenchymlamelle vom Mestom abgetrennte Bast- 
masse bleibt gewohnlich in ihrer Ausbildung sehr zuruck, wie 
das Fig. 19 zeigt. Nun schien es mir berechtigt diese Paren- 



l)Sc}iwendener, Das mechanische Princip. p. 65. 
2)Haberlandt, U p. 23. , . 



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442 K. shtbata: 

chymlamelle als eine im Innem des Stranggewebes eingeschobene 
„Starkescheide"^), die in ihrer physiologischen RoUe der 
gewohnlichen strangumgebenden gleicht, aufzufassen. Derartige 
Einrichtungen wiirden vielleicht zweckentsprechend seia, bei 
eiDem mit so starketn Baelbelege versehenen Biindel, wie es bei 
Bambuseea angetroffea wird^). 

Die schon erwahnten spindelformigen Leptomanschwellungen 
des Knospenbiindels kommen auch in dem Halmknoten yori 
jedoch meist in schwacherer Ausbildung. 

Die dunneren Blatttragenden Zweige stimmen in ihrem Baa 
mit den dickereu Halmtheilen wesentlich iiberein. Bei einem 
solchen (mit einem Durchmesser kleiher als 1 mm) wird das 
Rindenparenchym zu 1-2 Schichten reduciert und mehr oder 
minder verdickt. Die aussenseitigen Bastbelege der peripherischen 
Biindel stossen oft direct an die Epidermis, so dass eine Art 
Bastrippe zu Stande kommt^). Derartige Bippenbildung konnte 
ich jedoch bei einigen Arten, wie Amndinaria Matsumurce, selbst 
in den diinnsten Zweigen (0.7 mm dick) nicht nachweisen. 
Was den Gefiassbiindelverlauf in den Halmen sowie in dea 
Rhizomen anbetriflft, so gehort er dem Palmentypus*) an, and 
habe ich durch successive Querschnitte und Langsschnitte in 
der Spitzenregion constatiert, dass die grossen medianen Blatt- 
spurstrange 5-6 Intemodien zuriicklegen miissen, bevor sie sick 
an andere Blattspurstrange ansetzen. 



1) Vergl. Heine, Dber physiologische Function der Starkescheide. Ber. d. D. R G. 
1885, p. 189. 

2) Allerdings wurde die mechanische Bedentung, die Detlefsen (Ub. d. Biegangseitt^* 
citat T. Pflanzentheilea Arb. d. Bot Inst Wurzburg. Bd. Ill,' p. 182.) diesen Parcnchym- 
lamellen zuznschreiben versuchte, von Schwendener (Zur Lehre v. d. Festigkeit d. Gewachae. 
Qes. Bot. Mitteil. Bd. II. p. 19-20.) genugend widerlegt. 

3) Vergl . Schwendener, Die Mestomscheide. p. 183. 

4) P^ Bary, Ver^leicbende Anatomie 4. Vej;etation8organe, p. 271 ff: 



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WACHSTUMSGESCHIOHTE J>. BAMBUSGEWAECHSE. 44S 



Der Stiel. 

Die vielen untersten Internodien des Schosslings von Phyllo- 
stachys mitts vereinigen sich sehr friih 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, 
lasst keinen Unterschied mehr zwischen Internodien und Nodien 
im inneren Bau erkennen. 

Die Kinde dieses Theils besteht aus etwa 20 Schichten 
parenchymatischer Zellen. Die Bastbelege der aussersten Biindel 
verschmelzen sich zu einem vielfach unterbrochenen unregel- 
massigen Band. Nach innen liegen zahlreiche Biindel^ welche 
die ganze Lange des Stiels hindurch gedrangt verlaufen (Fig. 13) 
und dann in Rhizomknoten eintreten, um sich dort an die 
Blattspuren anzusetzen. So iiberwiegen im Querschnitte des 
Stiels sehr stark die Biindel, und das dazwischen liegende Paren- 
chym ist zu einem 2-4 schichtigen schmalen Grewebe reduciert. 
Diese Verhaltnisse entsprechen wohl der Function des Stiels als 
Leitungswege und nicht als Speicherngsort Die Querschnitt- 
form des Bundels mit Bastbelege ist in den meisten Fallen 
loindlich oval und es ist vollkommen von 3-6 schichtigen 
Bastzellen umgeben. Daher ist der Stoffaustausch zwischen 
leitenden Elementen und Grundparenchym so gut wie ganzlich 
ausgeschlossen. Das Leptom nimmt die aussere Halfte des Bundels 
ein und besteht aus einer Anzahl 0.04-0.05 mm breiten Siebrohr- 
en und englumigen Geleitzellen. Das Hadrom besteht aus nur 
einem (seltener zwei) grossen Gefasse (oft bis 0.15 mm weit), 
welches von einigen Schichten kleinzelligen Hadromparenchyms 

1} Dieser y^tieV' stellt also eine einzige StoffleituDgsbahn zwischen dem vom Schosslinge 
t'ch entwickelnden Halme und dem Rhizome dar. 

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444 K. SHrBATA : 

umgeben ist (Fig. 14). Dazu kommen noch einige Tracheiden 
mit netz-, spiral- oder ringformigen WandverdickuDgen. Bei den 
ubrigen Arten sind die ebenso schmalen Stieltheile in genan 
derselben Weise ausgebildet und sie stimmen in ihrer inneren 
Structur mit dem oben beschriebenen ganzlich xiberein. 

Die Wubzel. 

Die zahlreichen Wurzeln^) befinden] sich radial angeordnet 
an den Bhizomknoten und den unterirdischen Halmknoten; sie 
erreichen bei Fhyllostachys-Arten eine maximale Lange von 70 
cm mit einem Durchmesser von 4 mm. Zunachst will ich liier 
den anatomischen Bau der Wurzelrinde von Fhyllostachys-'lmi. 
Jrundinaria- Arten naher betrachten. 

Die ausserdte Zellschicht der Binde laast sich als die Aus- 
senscheide unterscheiden, indem die ausseren und radiaren 
Zellwande selir stark verdickt sind (Fig. 24a u. 25). Damit 
spielt sie die BoUe der schiitzenden Oberhaut anstatt der Epi- 
dermis, die sehr friih zerstort und abgeworfen wird. Nach innen 
folgt die verholzte Bastschicht (Fig. 24a). Das Bindenparenchym 
lasst sich in die ausseren aus unregelmassig poljgonalen weit- 
lumigen Zellen zusammengesetzten Schichten und die inneren 
aus regelmassig in radialen und concentrischen Beihen angeord- 
neten Zellen bestehenden Schichten unterscheiden^). Die letzteren 
sind von einer Anzahl radialer ^Luftraume durchzogen. Die 
Zellen der Endodermis besitzen eine starke Verdickung von 
inneren und radialen Wanden, die in liblicher Weise verkorkt 
sind (Fig. 28). Die stark verdickte Wandung ist zierlich ge- 

1) A. et C. Riviere, Les Bambous. p. 93. 

2) Die Ze)lschichten79lil der iimeren Binde ist stets Ideiner ftls die der ai»ei«D- 



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WAOHdTUMSGEdCHlCHTE D. BAHBUSOEWAECfiSE. 445 

schicbtet und yon feinen verastelten Kanalen formlich durchselzt 
(Fig. 28). Im Querschnitt stellt demnach diese nach Aussen 
zagekehrte C-Scheide ein symmetrisches Bild mit der oben er- 
wahnten ebenso C-formigen Aussenscbeide dar^). Die 1 oder 2 
innersten unmittelbar derEndodermis anliegendenBindenschichten 
Bind bei PhylloUachys Kumasasa, Bambusa borealis und Arundinaria 
quadranfftUaris als Verstarkungsring^) ausgebildet, indem die 
Zellen durch innenseitige C-formig yerdickte und stark verholzte 
Wande ausgezeicbnet sind (Fig. 34). 

So weit es den Bau der Wurzelrinde betriflft, zeigen die echten 
Bambttsa^Avten namlich B. vulgaris^ B. nana, B. sienosia^chya, 
B. arundinacea u. a. ein von dem oben beschriebenen ganz ab- 
weichendes Verhalten. Hier weisen die Zellen der eubepidermal- 
schicht keine Wandverdickung auf und ebenso verhalten sich 
die persbtenten Epidermiszellen. Darauf folgen 2-3 ScLichten 
enger Bastelemente, welche nach innen scharf von deo weit- 
lumigen Rindenparenchymzellen abgesetzt werden (Fig. 26 u. 27). 
Die aussere Rinde besteht nur aus einigen Zellschichten, wabrend 
die von grossen Luftraumen durchzogenen inneren Schichten viel- 
fach dicker sind (Fig. 27). Die Endodermiszellen sind ringfium 
verdickt und bilden die sogenannte 0-Scheide'). Merkwxirdig 
ist femer der Bau des Verstarkungsrings. Die innersten 1-oder 
2-schichtigen Rindenparenchymzellen fuhren an ihren ioneren 
Wanden eine Anzahl unregelmassig gestalteter aus reiner Cellulose 
bestehender Auswuchse, die haufig die ausseren Wande erreichen, 
80 dass sie im Querschnitt die ganzen Zellen nahezu aus- 



l)Das8elbe Verhaltniss wurde von Schwendener (Die Schutzscheiden und ihre 
Verstarkongen. Ges. Hot. Mitt. Bd. 11, p. 120, 127.) auch bei einigen Qrchideenlaftwurzeln 
bemerkt. 

2) Schwendener, Die SchfttzBcheide and ihre Verstarkungen. Gres. Bot. Mitt. p. 182. 

3) VergL Schwendener, Lc p. 128, Tabelle. 



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446 K, SHIBATA : 

zufullen scheinen (Fig. 31 u. 32)^). Eine Anzahl einheimischer 
Arten^ die wegen ihrer 6 Stamen bisher in Bambusa eingereiht 
warden, z. B. B. Veitchiij B. palmataj B. borealis etc., weisen 
jedoch im Bau der Wurzelrinde eine vollkommene Ubereinstim- 
mung mit Arundinaria auf und so schien es mir berechtigt, miter 
Berucksichtigung nocli anderer Merkmale, — vor allem : Fehlen der 
in Bastbelege eingescliobenen Parenchymlamellen, die Gestalt der 
Caryopse, die langkriechenden Rhizome u.8.w. — diese Formen- 
gruppe von Bambusa loszutrennen und als eine neue Section in 
ArundinariecB 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 durchfiihrbar ist. Andererseits kommt diese 
Formengruppe^) in ihrer Verbreitung auf Japan*) beschrankt vor. 
Wir gehen nun zur Betrachtung des Centralcylinders fiber. 
Die Anordnung der Leitbiindel weicht, wie es von Ross*) 
nachgewiesen wurde, vom typischen Bau der Monocotylen ab. Zu 
den normalen peripherischen radialen Biindeln, die ausserordent- 
lich polyarch sind,*^) kommt noch eine Anzahl der inneren iso- 
lierten Hadrom- und Leptomstrange hinzu. Im Querschnitte 
beliebiger junger Wurzeln bemerktman innerhalb der diinnwandi- 
gen Endodermis ein oder zwei Schichten des ununterbrochenen 
Pericambiums (Fig. 23), dessen allgemeine Vorkommniss in Bam- 



1) Derartige Structur findet man nicht inSchwendener's Aufzahlung der verechiedenen 
Verslarkungsformen (vergl. Lc* p. 132). 

2) Die Anzahl der bis jetzt bekannten hierhex^horigen Arten ist neun. Veigl. Makino. 
Bambusaceae Japonic®. Bot. Mag. XIV, Nr. 156, p. 20. 

3) Vielleicht auch in China. 

4) Boss, Beitrage zur Anatomie abnorm. Monocotylen wurzeL Ber. d. D. B. G. Bd. I, p. 337. 

5) Z. B. in einer 4 mm dicken Wurzel von Phyllostadiys mUis habe ich mehr als 150 
gezahlt. 



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WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 



447 



buseen um so mehr Beachtung verdient, als bei den meisten 
Gramineen die primordialen Gefasse nach van Tieghem^) direct 
der Endodermis anzustossen pflegen. Die inneren Hadromstrange 
kommen in etwa drei coneentrischen Ringen angeordnet vor. 
Dazwischen liegen zerstreut die inneren Leptomstrange, welche 
jedesmal aus den 1 oder 2 Siebrohren und den englumigen 
Geleitzellen bestehen (Fig. 43 etc), Bei ecbten Bambusa-Axien 
besitzt die stets einzeln stehende Siebrohre einen regelmassig 
ovalen Umriss (Fig, 38), Die Gesammtanzahl der inneren Leptom- 
strange betragt in den meisten Fallen, wie folgende Beispiele 
lehren, eine Halfte der peripherischen, aber bei echten Bambusa- 
Arten kommen beide fast in gleich grosser Anzahl vor. 





Peripherisches 


Inneres 


, 


Leptom 


Leptom 


Phylloatachys mitts 


84 


42 


P. bambtisoides 


83 


41 


P. puberula 


47 


24 


Arundinaria japonica 


75 


40 


A. Hindsii 


181 


97 


A. Matsumurce 


27 


13 


A. variabilis 


30 


16 


A. pygmcea 


43 


22 


Bambusa palmata 


41 


20 


B. Veiichii 


29 


15 


B. ramosa 


20 


10 


B. paniculata 


39 


20 


B. nipponica 


33 


16 


B. vulgaris 


70 


68 


B. sienosiachya 


47 


41 


B, liana 


48 


45 


Dendrocalamus laiijlorus 


81 


55 



1) Van Tieghem, Les Bacine. p. 123; Vergl. Morot, Recherche but le Pericycle. 
Ann- d. Sc nat. 8^. VI, T. 20, p. 233 und auch Falkenberg, Vergl. Untere. d. Vege- 
tationsorgane. p. 192* 



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448 K. SHIBATA : 

Die xibrigen Elemente des Centralcylinders werden, al^ 
sehen vom centralen Markparenchym, prosenchymaiisch zugespitet 
und zugleich stark verdickt. So entsteht hier ein hohlcjlindri-* 
sober mechanischer Bing, in welchem sammtliche Leitstraoge 
eingebettet liegen. Hier muss noch eine Frage gelost werden: 
In welcher Weise gescbieht die Communication zwischen den 
einzelnen leitenden Elementen, die yon einander getrennt im 
mechanischen Gewebe liegen? Zwar hat Beinhardt^) die in Frage 
kommenden Verbaltnisse bei den anomal gebauten Wurzein von 
Musaceeriy Fandanaceeriy Palmeen und Oyclanthaceen ermittelt 
und manch interessantes entdeckt. Betre£& der Communication 
zwischen einzelnen Leptomstraogen in unserem Fall muss vor 
allem bemerkt werden, dass die ausserordentlich stark verdickten 
und yerholzten Pericambiumzellen als die Leitungswege zwischen 
den peripherischen Leptomstrangen kaum in Betracht koramen^). 
Wenn man nun die Zahl der in beliebigen zwei Wurzelquersch- 
nitten vorkommenden Leptomstrange sorgfaltig mit einander 
vergleicht, so kann man leicht eine bedeutende Abnahme der- 
selben nach dem Wurzelspitze wahrnehmen, wie es aus einigen 
beigefugten Beispielen hervorgeht: 

Zahl der Leptomstrauge in 
Proximalende Mitte Distalende 

22.5 cm langes Wurzelstiick') 128 — 104 

12 „ „ „ 116 108 102 

Der Umstand beruht bloss darauf^ dass die inneren Lep- 
tomstrange sich unter einander und mit den peripherischen im 
weiteren Verlauf allmahlig verschmelzen, wie man sich durch 
Betrachtung successiver Querschnitte iiberzeugen kann. Die 

l)Reinhardt, Das leitendegewebe einiger anoma]gebaaten Monoootylenwurzel. Jahrb. 
t wiflB. Bot. Bd. XVI, p. 336. 
2)Beinhardt, /.c p. 361. 
3j yon FhtfUoHlachya bambu$oide4- 



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WAOHSTUMSGESCHIOHTE J>. BAMBUSGEWAEOflSE. 449 

Figuren 39 und 40 zeigen einige Falle der erwahnten Verschmelz- 
uDg. Dieser Modus des Leptomverkehrs ist nach Reinhardt'- 
schen Angaben auch bei anderen anomal gebauten Wurzeln haufig 
verwirklicht^). Der zweite Modus ist aber von mehr wirksaraer 
und auflfilliger Art. Bei jeder Ansatzstelle der zahlreich ent- 
springenden Nebenwurzeln an Centralcylinder werden sammtliche 
bier befindliche peripheriscbe sowie verscbieden tief liegende 
innere Leptomstrange in einem System formlicher Anastomosen- 
bildung zusammengehalten, welche bei den meisten Arten sich 
auf die etwa 10 periperischen Leptomstrange biniiberstreckt und 
bei Bdmbtbaa-ATten sogar die Halfte des ganzen Umfangs des 
Centralcylinders in sich umfasst Die etwas schematisierten 
Figuren 35 und 36 illustrieren das obengesagte. Das bier die 
Verbindung zwiscben einzelnen Leptomstrangen berstellende 
Gewebe besteht aus den plasmareichen parencbymatischen Zellen, 
die mit den ansehnlich grossen 2iellkernen und den dunnen un« 
verholzten Wanden versehen sind (Fig. 43 u. 44). Die Versch- 
melzung zweier Gefasse babe ich nur selten jgeseheu, wabrend 
bei der Ansatztelle der Nebenwurzel sammtliche mecbanische 
Zellen zufolge reicblicber Tiipfelbildung uud haufiger auftretender 
Querwande einen holzparenchymartigen Character annehmen und 
demgemass dem Saftaustausch zwiscben den eingebetteten Gefassen 
besser angepasst sind. Den directen Anschluss d^r Leptomelemente 
an Holzparenchymzellen, wie es von Eeinhardt fxir Musaceen 
und Oyclanihaceev?) nacbgewiesen wurde, babe ich auch haufig 
bei Bambuswurzeln angetroffen (Fig. 41). 

Die Basaltheile des Centralcylinders der Nebenwurzel sind 



l)Reinhardt, la, p. 364, p. 343 etc. 

Vergl. Ross, Beitr. z. Anat. abnorm. Monocot wttrzel. p. 334. 

2)Reinhardt, Up. 343, p. 346 und p. 348, 



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450 K. 8HIBATA : 

aus dem stark verdickten porosen recbteckigen par6nchymati«> 
schen Zellen gebildet, durch welche die kurzen Basalglieder jedes 
Leptomstrangs abwarts verlaufen, um sich dem oben erwahnten 
Leptomanastomosencomplex der Hauptworzel anzuscbliessen 
(Fig. 37). Hingegen scheinen die Gefasse der Nebenwurzel 
basalwarts meist blind zu endigen, so dass sie nur selten in 
directen ZusammenhaDg mit denen der Hauptwurzel kommen. 

Die Nebenwurzeln zeigea in ihrem Bau alle Merkmale der 
beziiglichen Hauptwurzeln, dennoch fehlen ihnen stets die inneren 
Hadrom- und Leptomstrange (Fig. 47 u. 45). 

Die Ansetzung der Wurzeln an die Stammorgane geschiet 
in der bei Monocotylen xiblichen Weise^). Die Elemente des 
mechaniscben Kings des Wurzelcentralcylinders breiten sich 
sebeibenformig aus und verschmelzen sich mit den aussersten 
Biindeln der Stammgebilde* Einzelne losgeloste Wurzelstrange 
dringen noch weiter ein und schliessen sich den peripheren 
Stammbundeln an, wobei das Leptom der ersteren solch eine 
Umgestaltung erfahrt, wie sie bei den Knospenbiindeln beobachtet 
wird^). 

Es erubrigt noch einen interessanten Befund kurz zu 
erwahnen. Die Rindenparenchymzellen der Nebenwurzeln, mit 
Ausnahme von den innersten 2-3 Schichten kleinlumiger Zellen, 
sind gewohnlich von einem Pilz bewohnt, der in jedem ZelUumen 
ein ansehnliches Knauel von dicken verschlungenen Mycelfaden 
bildet (Fig. 46 u. 47). Die verpilzten Wurzeln bieten trotzdem 
ein ganz normales und gesundes Aussehen dar. Die Mycelfaden 
treiben hie und da sogenannte Vesikulen aus und producieren 
oft massenhaft gelbe kornige Substanz von nicht genau bekannter 

1) Vergl Falkenberg, Veigl. XJnters. p. 196. 

2) Vergl. p. 437. 



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WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 451 

chemischer Zusammensetzung. Die Starke verschwindet gewohn- 
Hch vom inficiertea 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'ATten ale 
in den mit Epidermis versehenen Bambusa-l^ ehen^uvzeln con- 
stant nachweisbar. Die Kindengewebe der Hauptwurzeln habe 
ich meist pilzfrei gefunden, abgesehen von den dunneren Wurzeln 
von Arundinaria vaHabilis^ Bambusa ramosa^ etc. Die Losung 
der Frage nach der physiologischen Rolle^), die dieser Pilzsym- 
biont in der Ernahrung der Baumgraser spielt, will ich inir 
fur kunftige Studien vorbehalten. 



Die Blattgebilde. 

Die in zwei entgegengesetzten Reihen gelegenen, breiten 
Seheideblatter^) umliiillen iibereinander den ganzen Schossling 
und auch die wachsende Spitze des Khizoms. 

In der basalen, zum Knotengewebe libergehenden Region 
jedes Scheideblattes weisen die Leitstrange in ihrem Hadrom kein 
grosses getiipfeltes GeK^s auf, sondern sie besitzerf nur zahlreiche, 
oft mehr als 15 Ring- oder Spiralgefasse, die mit einander man- 
nigfach anastomosieren. In dem in der mittleren Partie des 
Scheideblattes ausgefiihrten Querschnitte erblickt man parallel- 

1) Frank, Lehrb. d. Botanik. Bd. J, p. 274; tber neue Mjcorfaiza-Formen. Ber. d. D. 
B. G. Bd. V, p. 400. 

2) Es wurde neaerdiags yielfach die Ansicht geausBert, dass die^Pflanzen die mit Mjcorhiza 
ansgerustet sind, der Assimilation des freien Stickstoffi befahigt seien. Vergl. Janse, Ann. d. 
Jard. Bot Buit, Vol. 14, p. 200, und auch Nob be, Landw. Versuchs-St. Bd. LI, p. 241. 

3) Rividre, Les Bambous. p. 76-82, p. 231. 

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452 K. SHIBATA : 

verlaufende, abwechselnd starke und schwache Leitbiindel, die in 
ihrem Bau kaum von den dera Stara morgan eigenen abweichen 
(Fig. 48). Sie sind mit einander durch die aus einigen Siebrohr- 
en und Gefiissen bestehenden Queranastomosen verbunden 
(Fig. 53). Die Bastbelege auf der Leptomseite stossen gewohnlich 
unmittelbar an die stark verdickte Epidermis der Aussenflache an 
(Fig. 51), aber bei dicken fleischigen Seheideblattem der PhyUo- 
stachyS'Arten liegen fast alle Biindel mit ihren Bastbelegen ganz 
frei im Parenehyra (Fig. 52). Entgegengesetzt den starkeren 
Bundeln liegen die bandformigen, meist 2-3 schichtigen Bast- 
strange an der Blattinnenseite. Die letzteren kommen bei Arundi- 
naria Matsumurce sonst auch an der Blattaussenseite bier und da vor 
(Fig. 49). Das Scheideblattparencbym besteht aus diinnwandigen, 
saftreichen Zellen,^) von dcuen einige subepidermal Schick ten bei 
unterirdischen, harten Scbeideblattern sclerenchymatisch verdickt 
sind. Bei den derben oberirdischen Scbeideblattern von Arundi- 
narta-ATten tragen die an die In tercel lularraume angrenzenden 
Flachen der Parenchymzellen die eigenthiimlichen bald kugel- 
formigen, bald stabchenformigen Auswiichse, die starke Holz- 
reaction geben (Fig. 54). 

Die SpaltoflFnungen kommen an der Ober- sowie Unterseite 
der Scheideblatter vor. 

Die laubblatttragenden Blattscheiden stimmen in ihrem Bau 
mit den oben geschilderten Niederblattern wesentlich iiberein. 

Die Laubblatter der Bambuseen sind schon wiederholt von 
vielen Forschern anatomisch untersucht worden. So haben 
Kareltschikoff^), Magnus und Haberlandt die Armpallisa- 

1) Die Parenchymzellen ansgewachsener Scheideblatter enthalten fast keine Starke, sondera 
viel Qlykose. 

2) Kareltschikoff, Ub. d. faltenformig? Verdickungen in d. Zellen einiger GramineeiL 
p. 180. (Referat). 



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WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 453 

dennatur des Assimilationsgewebes erkannt. Bei Giintz finden 
wir Angaben iiber einige allgemeine Characteristik der Bambuseeu- 
blatter, dabei fiihrte das energische Auftreten der mechanischeu 
iOemente ihn zur Aufstellung des ,, Bambuseentypus " der 
Gramineenblatter^). Bei dieser Sachlage wiirde es berecbtigt sein, 
dass ich mich hier nur auf einige kurze Notizen beschranke. 

Um jedes Mestombiindel 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 Biindeln nicht; bei starkeren 
Bundeln ist sie oft dort stark verdickt und verholzt, wo sie an 
subepidermale Bastrippen ansehliesst. 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 Biindel meist 
einschichtig, aber bei den starkeren nicht selten mehr als 4 
Schichten dick. Ferner verhalten die Elemente dieser Scheide 
sich gegen Schwefelsaure kaum anders als gewohnliche verholzte 
Bastzellen, wahrend sich die unverholzte Parenchymscheide gegen 
dieses Reagens sehr widerstandsfiihig erweist. So ist die in Rede 
stehende Scheide als eine vereinfachte Form der das Mestom 
voUkommen umschliessenden Bastscheide, \yie ich sie schon bei den 
Stielbundeln beschrieben habe, aufzufassen. 

IV. Der EntwicklungsYorgang der Schosslinge. 

Als Gegenstand der folgenden Darstellung diente mir Fhyllo- 
stachys mitts. 

l)Guntz, Untere. iib. d. anat. Struct, d. Gramineenbl. p. 64. 

2) Schwendener, Die Mestomscheiden der Gramineenbliitter ; Vergi . Strasburger, 
Leitungsbahnen. p. 344. 

3) Schwendener, I.e. p. 178. 



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454 K. SHIBATA : 

Dieauf jedem Knoten der wachsenden Rhizomspitze angelegte 
Knospe wird erst im nachsten Jahre zu einem kleinen SchossliDg 
mit dem schon differenzierten, verholzten, ca. 1 cm langen Stiel 
ausgebildet. Diesen letzteren nenne ich kurzweg den SchossKng 
des 2ten Stadiums, wahrend die dem Knoten dicht anliegeode, 
stiellose Knospe als Istes Stadium von diesem unterschieden 
wird. Wenn man einen Querschnitt in der oberen Region dieses 
kleinen Seho^slings ausfuhrt, so sieht man den Centralcylinder 
gesondertin einen peripherischen, schmalen, biindelfiihrenden King 
und in umfangreicbes Markgewebe, welches sich nach unten 
allmahlig verschraalert. Auf dem Laugsschnitt sieht man dicht 
unterhalb des Urmeristems vom Vegetationspunkt beginnend eine 
grosse Anzahl abwechselnd starkereiche und starkearme Zonen, 
welche letztere sich in spateren Stadien zu Internodien verlangern. 

Der Schossling des 2ten Stadiums nimmt im Laufe des Som- 
mers an Grosse zu und wachst im Spatherbst (October-November) 
schon zu einem mittelgrossen Schossling des 3ten Stadiums. In 
diesem Zustande verharrt er wahrend des Winters. 

Anscheinend schon in Marz tritt eine rasche Zunahme an 
Grosse ein und im Anfang April erreichen die Schosslinge unter 
der Erde eine ansehnliche Grosse, die ich als 4tes Stadium 
kennzeichnete. Der Schossling ist mit zahlreichen geraumigen, 
dicken Scheideblattern 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 Hohe 1-2 cm betragt, 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 starkereichen und 
starkearmen Zonen, deren Zahl binnen 6 mm 40 betragt 



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WACHSTUMSGESCHICHTE I>. BAMBUSGEWAECHSE. 45^ 

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 
Hohe in den nachfolgenden Internodien immer zunehraen. Diese 
primordialen Markhohlen erreichen in den unteren, mit Wurzeln 
besetzten Internodien eine maximale Hohe von ca 3 mm, dabei 
besitzt das Diaphragm eine Dicke von 2.5 mm. Die Biindel- 
aulage in der Spitzenregion besteht aus engen procambialen 
Zellen. Oberhalb der Stelle, wo die erste Markhohle zum 
Vorschein kommt, erfolgt schon die Differenzierung in Elemente 
des Biindels. 

Indessen tritt die Spitze des Schossliugs allmahlig auf der 
Erdoberflache hervor; vom Ende April ab erfolgt dann ein 
rasches Wachstum desselben. Schon in Mitte Mai erreichen 
mehrere Schosslinge eine Hohe 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 Lange graduell zu. 
In mittlerer Hohe der Pflanze erreichen sie die Lange von 20 
cm und mehr. Bis auf diese Region haben alle Internodien ihr 
Langenwachstum vollendet. Mehrere darauf folgende Internodien 
besitzen basale Wachstumszonen. Die von dort nach oben lie- 
genden Internodien verjiingen sich allmahlig zum Vegetations- 
punkt. Die noch in Streckung begriffenen Internodien sind stets 
mit Scheideblattem umhiillt. Der Schossling in diesem Zustande 
ist im 5ten Stadium. 

Hier lasse ich einige Zahlenangaben folgen : 



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456 



K, SHIBATA 







Stadium II. 


Stadium HI. 1 Stadinm IV. 




1 ' 2 3 


1 2 1 3 1 2 1 3 


Lange 
in 
cm. 


Aeunere 
Curratur 


4.4 


4.2 4.6 


20.2 


19.8 


16.0 


1 
56.7 48.0 


4S.5 


lanere 
Currator 


8.7 


3.8 


3.4 


13.9 


13.4 


12.0 


Mftximalnmfang in cm. 


5.3 


4.9 


4.4 


16.6 


14.8 


13.8 


34.0 1 31.5 


315 


Gewicht in Gr. 


4.18 


4.21 


4.11 


173.0 


154.0 1 116.5 


1831.0 1432.0 


2062.0 



Tagliche Zuwachsmessongen 
an jedem Mittag yom 24 April 





L 


n. 


UL 


IV. 


Datam. 


Lange 
in 
cm. 


1 
1 


Lange 
in 
cm. 


1 
1 


Lange 
in 
cm. 


1 


Lange 
in 
cm. 


1 1 


April 24 


34.9 




37.2 




45.7 




27.2 


1 


25 


38.1 


3.2 


43.1 


6.9 


51.3 


5.6 


31.0 


3.8 


26 


44.2 


6.1 


50.6 


7.6 


61.1 


9.8 


37.6 


6.6 


27 


53.4 


9.2 


62.4 


11.8 


75.0 


13.9 


46.1 


8.5 


28 


62.5 


9.1 


73.3 


10.9 


91.7 


16.7 


66.6 


10.6' 


29 


74.1 


11.6 


87.4 


Ul 


1144 


22.7 


72.9 


16.3 


80 


86.8 


12.7 


101.3 


13.9 


135.3 


20.9 


86.8 


13.9 


Mai 1 


106.9 


20.1 


123.9 


22.6 


164.0 


28.7 


108.8 


22.0 1 


2 


137.9 


31.0 


158.3 


34.4 


205.8 


41.8 


141.9 


33.1 ; 


3 


175.9 


38.0 


202.4 


44.1 


253.3 


47.5 


180.1 


38.2 


4 


199.1 


23.2 


230.7 


28.3 


283.6 


30.2 


207.7 


27.6 


5 




— 


277.1 


46.4 


336.4 


62.9 


263.2 


65.5 


6 


300.5 


50.7 


3n3.7 


66.6 


398.6 


62.2 


319.0 


55.8 


7 


825.5 


25.0 


363.7 


30.0 


418.0 


19.4 


346.6 


27.5 


8 


338.6 


13.1 


376.6 


11.9 


447.6 


29.6 


361.2 


14.7 


9 


382.5 


43.9 


417.6 


42.0 


500.6 


53.0 


409.4 


48.2 


10 


423.9 


41.4 


464.5 


47.0 


546.9 


46.3 


452.6 


43.2 


11 


475.3 


51.4 


515.6 


51.1 


605.0 


58.1 


514.1 


61.6 


12 


561.0 


75.7 




— 




— 


597.9 


8M 


13 


580.1 


29.1 


619.2 


51.8 


721.4 


58.2 


636.2 


38.3 


14 


631.6 


51.5 


661.9 


42.7 


786.4 


R).0 


664.6 


28.4 


15 


719.6 


sao 


744.7 


82.8 

1 


846.1 


59.7 


710.7 


46.1 



^J?.— * Znwachs ist Miltel yon 2 Tagen. 

*^ Nach Beobachtungen des hiesigen meteorologischen Observatoriums. 
Anm. 1. Also bei diesen Messungen stieg der mazimale Zuwachs nicht selten uber 80 cm pro 
Anm. 2. Das bisher bekannte starkste Wachstum der Bamboshalrae betragt 91.3 cm pro 24 
Anm. 3. Auf die hier beobachteten auffalligen Wachslumsschwankungen und andere interes. 

1) Die alteren Angaben iiber das Wachstum der Bambuspfl^^gf^^igi^et man bei Kr^s 



WACHSTUM8GESCHI0HTE D. BAMBUSGEWAECHSE. 



457 



Um die erstaunlich grosse Schnelligkeit des Wachstums von 
Bambusschosslingen in dem 5ten Stadium zu demonstrieren^), 
fiihre ich bier einige von mir ausgefiihrte Messungen an Phyllo^ 
staehys mitis ap. 



yon PAy//(wrfac%«-HalmeD, 
bis 15 Mai aasgefuhrt. 



V. 


VI. 


1 


Lange 
in 
cm. 


1 


Lange 

in 
' cm. 


CO 

1 


Wetterangaben. 


Mittlere 

Tempera- 

tnr.** 


Mittlere 
relat. 

Homidi- 
tat** 


106.6 








klar^lfeiser Wind 


14°.4C 


50.2 


121.2 


14.6 


52.6 




klar-wenig'triib 


120.5 


73.0 


144.8 


23.6 


76.3 


23.7 


Mar 


150.4 


74.3 


176.8 


32.0 


108.5 


32.2 


klar 


160.4 


69.8 


208.3 


31.5 


140.4 


31.9 


wenig iriib 


IfPS 


71.3 


250.4 


42.1 


184.4 


44.0 


klar, leiser Wind 


18M 


43.5 


284.8 


34.4 


216.8 


32.4 


klar, leiaer Wind 


14°.5 


64.9 


324.6 


39.8 


260.3 


43.5 


klarf windig 


16*^.7 


73.0 


378.4 


53.8 


322.0 


61.7 


klar, windslill. 


18°.4 


66.0 


444.4 


66.0 


388.9 


66.9 


Begen 


17°.6 


80.0 


484.2 


39.8 


429.2 


40.3 


kUir, leiser Wind 


1.3^8 


91.4 


551.7 


67.5 


403.9 


64.7 


wenig (rub 


17«.5 


86.0 


633.9 


82.2 


566.7 


72.8 


Begen 


17°.7 


82.6 


672.4 


38.5 


607.0 


40.3 


Begen 


1P.9 


93.3 


713.1 


40.7 


625.4 


18.4 


halbldary windig 


11^6 


86.6 


755.5 


42.4 


675.4 


50.0 


halhklarf windstiU, 


15^6 


76.1 


770.5 


15.0 


738.9 


63.5 


klar, leiser Wmd 
Mar, leisei' Wind 

Begen 
klar, leiser Wind. 

klar 

Mar 


16^.6 
17^.8 
18°.3 
14°.6 
18°.5 
20°.7 


81.2 
65.0 
84.8 
89.6 
78.2 
69.4 



24 Stnnden ! Bividre fand denselben bei PkylMathfs miHt in Algier 57 cm pro 24 Stnnden. 

Stunden. (Vergl. Pfeffer, Pflanzenphysiologie. Bd. II, p. 83.) 

sante Fragen kann ich>n dieser Stelle nicbt weiter eingehen (Vergl. Kraus, /.e.), 

(Ann. d. Jard. Bot. de. Bait. vol. XIL p. 197.) zusamraengestellt. Digitized by V^jOOQIC 



458 K. SHIBATA : 

V. Verhalten der Banstoffe 
wahrend der Entwicklung der Schosslinge. 

In diesem Kapitel Will ich die Umwandlungs- und Wander- 
ungsvorgauge verschiedener Baustoffe wahrend der Entwicklung 
der Bambusschosslinge in wesentlichen Zugen darzustellen ver- 
suchen. 

Die Reservestoffe. 

Unter stickstofffreien Reservestoflfen, die sieh in Bambus- 
pflanzen vorfinden, kommt die Starke in erster Linie inBetracht 

Im zeitigen Herbst, wo die Ablagerung der Reservestoffe 
sclion stattgefunden hat, konnte ich sie in oberirdischen und 
unterirdischen Theilen aller untersuchten Arten in weehselnden 
Mengen uuffinden. Die Phylloslachys-Arten, welche mit einem 
umfangreichen, unterirdischen Rhizomsystem ausgeriistet sind, 
pflegen nur sehr kleine Mengen der Starke in ihren Halmparen- 
chymzellen aufzuspeichern. Bei alien Arten wird die grosste 
Menge der Starke in Rhizomen und Wurzeln deponiert. Die 
Parenchymzellen der Knoten sind stets ausserst starkereich und 
die Blattscheiden einiger Arten speichern ebenfalls die Starke im 
Parenchym auf. Die Siebrohren sind dagegen hochst inhaltarm ; 
ich habe nur selten winzige Starkekonier in denselben nach- 
gewiesen. Es waren aber kleine Mengen von Glykose und 
Rohrzucker stets vorhanden. Beach tenswerth ist die Gestalt der 
Starkekorner ; bei Bambitsa palmataj B. Veilchii und B. pant" 
culata sind sie aus zahlreichen kleinen Theilkornern zusammen- 
gesetzt (Fig. 67). Solche polyadelphische Starkekorner') kommen 
nach Nageli in Stamnigebilden nur selten vor. 

1) A. Meyer, Untersnchnngen iiber die Starkekorner. p. 204. 

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WACHSTUMGSESCHICHTE D. BAMBUSGEWAECHSE. 459 

Der rediicierende Zucker ist ziemlich reichlich in Halraen 
und Rhizoraen ini Winterzustande nachzuweisen. Die winzigen 
Fetttropfchen sind oft im Halmparenchym von Phyllostachys 
mitts, Arundinaria Simoni, Arundinaria Hivdsii u.s.w. ange- 
troflfen, aber sie kommen jedenfalls als Reservestoflfe kaum in 
Betracht. 

Um eine Vorstellung fiber die Mengenverhaltnisse der auf- 
gespeicherten Starke zu anderweitigen Bestandtheilen der 
Eeservestoftbehalter zu gewinnen, habe ich einige Analysen der 
zweijahrigen Rhizome von Phyllostachys mitis ausgefiihrt^). Es 
ergab folgendes : 

% Gehalt der 
Trockensubstanz. 

Starke 24.01 

Reducierender Zucker 0.95 

Nicht reducierender Zucker 4.31 

Rohproteinstoflfe (Nx6.25) 5.41 

Fette 0.61 

Rohfaser 47.32 

Asche 8.74 

Unbestimmte StofFe (Differenz)... 8.65 

100.00 

1) Das am 25 November gcsammelte, kriiftige Rhizomstiick von Phyllostachys miltSy deflsen 
Parenchym sich zuvor bei microscopischer Beobachtung als von Starke strotzend erwies, wurde 
mittelst des Hobels abgcschaubt, schnell bei 70®-80° getrocknot, und zn einem feinen Schrot 
geroahlen. Von diesem lafttrockenen Rhizomscbrot wurde ein bestimmtes Quantum abge- 
wogen und zu jeder Bestimmung verwendet. 

Das Trockengewicht des Scbrots wurde nach weiterem 4 stiindigen Trocknen bei 100** 
(zur G^wichtsconstanz) bestimmt. 

Die Starke wurde mittelst der Erhitzung im So xhlet'schen Autoclave verzackert 

Die loslicben Kohlehydrate warden nach 5-6 maligera Ausziehen mit kaltem 
Wasser binnen 24 Sinnden erschopft. Der nicht re ducierende Zucker wurde nach 
Inversion mit verdiinnter Schwefelsaure bestimmt. Alle Bestimmungen der Zucker warden 
nach Meissl-Allih n'scher Gewichtsmethode ausgefiihrt. 

Der Gesammtstickstoff wurde nach Ejeldahl und der Eiweisssticks off nach 
Stutzer bestimmt. 

Die Fasersubstanz wurde durch Weender'sches Verfahren bestimmt. 

Das Atherextract wurde ohne wei teres als Oel angenommen. 

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460 



K. SHIBATA 



Man sieht also, dass die Starke wohl als Hauptreservestoff 
zu betrachten ist, dagegen sind die Proteinstoffe in ver- 
haltnissmassig geringer Menge vorhanden. 

Nun schien es mir erwiinscht zu wissen, eine wie weite Strecke 
des Rhizoms zum Auswachsen eines Schosslings dienen solite, so 
habe ich im Anfang Februar von einer Plantation von Phyllostachyi 
bambusoides eine Anzahl Rhizorastiicke ausgegraben und die auf 
Knoten vorkommenden Schosslinge (im 3ten Stadium) aufgezahlt. 
Es ergab folgendes Resultat : 



Zahl 
der Rhizomstucke 


Gesammtanzahl 
der Internodien 


Halme 


Hhizomzweige 


SchoasUnge 


69 


632 


5 


39 


15 



Aus obigem berechnete ich das Zahlenverhaltniss der Rhizom- 
internodien zu einem Schossling, wie 42.1:1. 

KOHLEHYDRATE. 

Die stickstofffreien Reservestoffe in alien untersuchten Arten 
bestehen, wie schon erwalint, hauptsachlich aus der Starke. Dass 
der Starkegehalt der Rhizome eine bemerkbare Verminderung 
wahrend des Winters erleidet, wie es von Rosenberg^) fiir einige 
perennierende Gewachse dargethan wurde, konnte ich nicht in 
diesem Fall bestatigen, da ich grosse Anzahl von Rhizomen von 
Phyllostachys mitisy Phyllostachys bambusoides, Phyllostackys Kuma- 
sasa, Arundinaria Hindsiij Arundinaria Narihira^ Arundinaria 
quadrangularisy Arundinaria Matsumurce, Bambusa palmata, 
Bambusa nana u.s.w. im Winter (Anfang Januar — Ende Febrnar) 



1) Rosenberg, Die Starke im Winter. Eot. Central bl. Bd. LXVI, p. 337. 



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WACHSTUMGSESCHICHTE D. BAMBUSGEWAECHSE. 



461 



untersuchte Und dabei keine merkliche Dififerenz in Bezug auf 
Starkegehalt von den im Herbst beobachteten Exemplaren auf- 
finden konnte. Auch die Wurzeln der untersuchten Arten 
enthielten in dieser Jahreszeit grasse Mengen von Starke in 
ihrem Einden- und Markparenchyra, so z. B. bei den am 25. 
Januar gesammelten Exemplaren : 



, Fhyllosiaehys , Phylloslachy.H | Ai-undinaria 
miiU bambusoides \ HtncUii 


Arundinaria 
Narihira 


Rindenparenchym i 4^) 4 
Markparenchyrn '3 i 2 


5 
3 


4 
3 



Gleiches gilt fiir die oberirdische Halme von Arundinaria- 
und Bambusa-ArtenJ^) Merkwiirdigerweise nimmt die Menge des 
reducierenden Zuckers wahrend 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 Schosslinge ein 



1) Bequemlichkeitshalber liabe ich zur Bezeichnung des Stilrkegebaltes folgende Zifiern 
benntzt: 

— bei ganzlicher Abwesenheit von Starke; 

1 — wenn ein Thell des G^webes sUirkefrei ist, wtihrend der andere wenige Komchen 

in den Zellen fuhrt; 
2 — wenn alle oder die meisten Zellen wenige Starkekomer enthallen ; 
3 — wenn ein Theil der starkefiihrenden Zellen wenige starkekomer enthalt, wahrend 

der andere recht viel Starke fiihrt; 
4 — wenn das Gewebe recht viel Starke enthalt; 
5 — wenn alle oder die meisten Zellen strotzend gefiillt sind. 

2) Vergl. A. Fischer, Beitriige zur Physiologie der Holzgewachse. Jahrb. f. wiss. Bot. 
Bd. XXII, p. 92, p. 112. 

3) IMeser Zucker geht aber grosslentheils schon im Anfang Miirz wieder verloren, ohne 
duss dabei eiue bemerkbare Starkezunahme stattfand. Auch fielen die Versuche den 2kicker 
in der abgeschnittenen Halmtheilen durch kiinstliche Erwarmung (im Treibhaus bei 17°-20'XIJ.) 
zur Starke iiberzufuhren negativ aus. 



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462 



fe. SHIBATA : 



rasches Wachstum begannen, ist die deutliche Starkezunahme 
in mehreren Rhizominternodien in der Nahe von Knoten, an 
welchen der wachsende Schossllng sitzt, zu beobachten. So 
z.B. bei PhyUostachys mitis : 



1 Anfang , Ende 
: Kovember j December 


Anfang | Anfang i Mitte 
Februar ' Marz . April 


Bindenparencbym < 4 4-3 

Centralcylinderparen- 2 2 
chym 1 

Markparenchym 3 1 4 


3 4 

2 

3-4 


3-4 

2 

3-4 


5 

4-5 

5 



Diese Starkezunahme mag jedoch darauf beruhen, dass 
die von ferneren Theilen des Rhizoms in Form von Zucker 
zugefiihrten Kohleliydrate hier in der Nahe des Schosslings 
transitorisch in Stiirke umgewandelt werden. Dafiir sprechen die 
Umstande, dass erstens in entfernteren Rhizominternodiea keine 
entsprechende Starkezunahme stattfand, und zweitens schon in 
dieser Zeit ein Blutungssaft, der eine wichtige RoUe beim 
Zuckertransport spielt, von jeder beliebigen Schnittflache des 
Rhizoms hervorquillt. 

Die Starkezunahme ist vor allem im verholzten Stieltheile 
des Schosslings ausgepragt : 





Ende An&ng Anfang Mitte 
December Februar Marz April 


Rindenparenchym 

Centralyclinderparen- 
chym 

Hadromparenchym 


2-3 
3 




2 
2 



5-4 
4-3 



5 
5 




Gleichzeitig wurde die partielle Entleerung der Rhizomknoten, 
an welche die Schosslinge sitzen, beobachtet, obgleich die nachst 
folgenden Internodien, wie schon bemerkt, noch von Starke 
erfullt war en. 



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WACHSTUMGSESCHICHTE D. BAMBUSGEWAECHSE. 



463 



Von jetzt ab wird die Starke im Rhizome nach und nach 
aufgelost und sclion in dera Stadium V, wo die Schosslinge auf 
der Erde 4-6 Meter hoch wuchsen, verschwinden fast sammtliche 
Starkekorner vom Parenchym der benachbarten Rhizominter- 
nodien. So zum Beispiel bei Fhyllostachys mitis : 
Rhizominternodien — 





16. April 


9. Mai 


19. Mai 


Kin den parenchym 

Centralcylinderparenchym 
Markparenchym 


5 
5 
5 


1 

1-2 

4-3 




1 


Nodium — 




16. April 


9. Mai 


19. Mai 


Sabepidermale sclerotische 
Schicht 

Kindenparenchym 
Centralcylinderparench ym 


4 
4 
1-2 












Stiel des Schosslings — 




16. April 


9. Mai 


19. Mai 


Subepldermale sclerotische 
Schicht 

Kindenparenchym 
Celntralcylinder parenchym 



5 
5 




1-0 










Hier findet auch in der Wurzel eine entsprechende Starkeent- 
leerung statt : 





16. April 


19. Mai 


Rinden- fausseres grosszelliges , 5 
parenchymiinneres kleinzelliges 5 
Markparenchym l 5-4 




1-2 




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464 K. SHIBATA : 

Merkwiirdigerweise konnte ich bei so raschem Auflosen der 
Starke eine entsprechende Glykosebildung ira Parenchym nicht 
beobachteii ; bei der Zuckerprobe nach Schimper erhielt ich 
nur Spuren von Oxydulkornern in Parenchymzellen. 

Die Kohlehydrate in wachsenden Schosslingen verhalten 
sich im Grossen und Gauzen analog mit denjenigen in von 
Sachs, H. de Vries u. A. untersuchten Pflanzen/) Indess 
ist folgendes noch zu bemerken. 

Die Abwesenheit von Starke im Urmeristem des Vegetations- 
punktes habe ich im allgemeinen constatiert. Die feinkornige 
Starke wird erst an der Stelle, wo die erste Differenzierung der 
Bundelanlage und des Grundparenchyms auftritt, nachweisbar 
und man kann abwechselnd starkereiche und starkearme Zonen 
deutlich sehen. Nach unten tritt der Unterschied im Starke- 
gehalt dieser abwechselnden Zonen immer scharfer hervor. Der 
reducierende Zucker tritt an der Spitze weiter unten als Starke 
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 Schosslings bleiben noch lange Zeit von der 
Glykose erfiillt. Sobald die Streckuug eines Internodiums voU- 
endet ist, verschwindet die Starke aus dem Parenchym, abgesehen 
von einigen winzigen Kornchen in 2-3 Zellen bei Durchlassstellen 



1) Hier seien nur folgende erwahnl : 
Sachs, Physiologiflche Untersuchung ub. d. Keimung von Schminkbohne. — Sachs, 
Keimungsgeschichte der Graser. — De Vries, Wachstumsgeschichte der Zuckerriibe.— 
De Vries, KeimnngBgeschichte der Kartoflelknollen. — Detmer, Vergl. Physiologie 
d. Keimungsprocess der Samen. — H o fim a d n, Uber d. Stoffwanderung bei d. Keimong 
von Weizen- und Kleesamen. — A. F. A. C. Went, Chemisch-pbysiologische Unter* 
suchungen iib. d. Zackerrohr. 
2)Vergl. Sachs, IJb. d. Stoffe welche d. Material z. Wachstum d. ZeUhaute liefem. 
Jahrb. f. wiss. Bot. Bd. Ill, p. 207. 



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WACHSTUMGSESCHICHTB D. BAMBUSGEWAECHSE. 465 

der BiindeL Da hierbei sammtliche Zellwande noch keine 
nennenswerthe Verdickung zeigen, so erfolgt die weitere Aus- 
bildung der Biindelelemente ohne Gegenwart der umgebenden 
Starkescheide, in welcher nach Heine^) die nothigen Baustoffe 
als Starke deponiert werden sollen. Die besonders starke Zucker- 
ansammlung in einigen Parenchymschiehten um die in Ausbildung 
begrifFenen Bastbelege herum vertritt hier die Stelle der feblenden 
Starkescheide uod daber mag sie als Zuckerscheide^) bezeichnet 
werden. 

In der wachsenden Wurzel bemerkt man die kleinste Menge 
der winzigen Starkekorner nur in der noch zartwandigen Endo- 
dermis. Hingegen ist in der Haube von der ersten Anlage die 
Starke in ihren Zellen festgehalten.^) Der reducierende Zucker 
kommt in der ganzen Lange der Wurzel, ausser der 4-5 mm 
langen Slrecke 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 
bem^rkbar. 

Wie schon erwahnt konnte ich in Rhizomen und Wurzeln, 
wo die Reservestarke in Auflosung begriffen war, gewohnlicb nur 
eine Spur von Glykose auffinden. Analoge Falle sind bereits 
bekanut. 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 gelosten 



1) Heine, Die physiologiflche l^deutung der sogenannten Starkescheide. Landw. 
Versucbs-St. 1888. p. 115. 

2)H. de Vries hat friiher den Ausdruck iin analogen Sinne init „ Leitscheide/* 
Schi roper's angewandt. 

3)Vergl. Sachs, Jahrb. f. wiss. Bot. Bd. Ill, p. 203. 

4) Sachs, ijber die Stoffe, welche das Material ziim Wachstura der Zellhaute liefern, 
Jahrb. f. wiss. Botanik. Bd. Ill, p. 248. 



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466 K. SHIBATA : 

Kohlehydrate von vornherein erwartet werden musste. In 
gewissen Fallen dieser Art ist es nicht unwahrscheinlich, dass 
die Glykose durch andere losliche Kohlehydrate ersetzt wird.*) 
In treibenden Rhizomen von Phyllostcichys mitts habe ich 
nun den Rohrzucker in starkehaltigen Parenchymzellen mittelst 
der Invertin-Methode nachgewiesen. Ferner in Rhizomen^ von 
welehen fast alle Starkekorner schon verschwunden waren 
(Stadium V), beobachtete ich noch erhebliche Mengen Rohrzucker. 
Uebrigens habe ich in folgenden Arten den Rohrzucker in Rhizo- 
men wahrend des Austreibens der Schosslinge beobachtet : Phyllo- 
stachys bambusoides, Phyllostachys puberula und Arundinaria 
japontca; und in folgenden im Halmparenchym wahrend des 
Austreibens der Zweigknospen : Bambusa palmata und Arundi- 
naria japoiiica. Ferner erhielt ich die Rohrzucker-Reactiou in 
Halmen von Phyllostachys Kwnasa^a^ Arundinaria Simoniy und 
Arundinaria Hindsii var. graminea. In den Wurzeln von 
Phyllostachys mi lis, deren grosskornige Reservestarke in Auflosung 
begriffen war, konnte ich ebenfalls Rohrzucker nachweisen. 
Gleiches gilt fiir verholzte Stieltheile der Schosslinge. In alien 
diesen Fallen kommt Rohrzucker hauptsachlich im Parenchym 
und viel weiniger in Siebrohren vor. Nun liegt mir der Gedanke 
nahe, dass in diesem Falle die Kohlehydrate hauptsachlich in 
Form des Rohrzuckers von Zelle zu Zelle wandern^). 

Vom Rohrzucker ist noch zu erwiihnen, dass ich ihn im 



1) So z. B. wiirde fiir Gramineen-Scutellum das Vorhandensein vom Rohrzucker anstatt 
Glykose von Griies auf microchemischem Wege sowiejauf experiroentelle Weise sicherge- 
stellt. (Vergl. Ber. d. D. B. G. Bd. XVI, p.']?). Puriewitsch (Jahrb. f. wLss Ifet. Bd. 
XXXI, p. 63.) hat bei der ersten Periode der Entleerung der Reservestarke das Aaftreten 
Dichtreducierenden Zuckers beobachtet. Vergl. Lcclare du Sablon, Recherche sor les 
Reserve Hydrocarbones des Bulbes et des Tubercules. Rev. gen. d. Bot 1899. 

2) Vergl. E. Schulze, Ueber die Verbreitnng des Rohrzuckers in den Pflansen nnd 
fiber seine physiologische Rolle. Zeitschrift f. physiol. Cheniie. Bd. XX, p. 652. 



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WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 467 

juDgen Gewebe unterhalb des Urmeristems, wo schon erne 
Differenzierung in nodiale und internodiale Zonen stattgefunden 
hat, manchmal, wenu auch nicht immer, durch Invertin- 
Methode nachgewiesen habe^). 



EiwEiss UND Amidoverbindungen. 

Im Jahre 1872 hat Pfeffer^) zaerst 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 namlich con- 
statiert, dass das Asparagin als Auflosungsproduct des Eeserve- 
proteins in Cotyledonen entsteht und dann wachsenden Theilen 
zugefiihrt wird, und ferner, dass wenn Kohlehydrate bei der 
Assimilation sich vermehren, das gebildete Asparagin zum Eiweiss 
regeneriert wird^). Die letzterwahnte Thatsache hat er spater 
durch die Kulturversuche im Dunkeln und in kohlensaurefreier 
Luft weiter begriindet.*) Seit diesen zum ersten Male exact 
ausgefuhrten Arbeiten Pfeffer's wurde die Frage nach Eiweiss- 
urasetzungen mit immer wachsender Eifrigkeit seitens der 
Botaniker und Chemiker verfolgt, was zu zahlreichen Arbeiten 
Veranlassung gab. Schulze und seinen Schiilern verdanken 
wir besonders eine Reihe der Versuche liber jene Amidover- 

1) Ich habe im Gewebe dieser Eegion eine schone rosarothe Fiirbung mit cone. 
Scliwefelsaiire erzielt. Diese Reaction deutet darauf hin, daas bier ein losliches Eohlehydrat 
neben Eiweiss vorhandeu ist. Vergl. Frankfurt, Zur Kenntniss der chemiachen Zusam- 
mensetznng des rubenden Eeimes von Triiicum vufgare. Landw. Versuchs-St, 1896. p. 461. 

2)PfefFer, Untereuchungen iiber die Proteinkorner und die Bedeutung des Asparagins. 
Jabrb. f. wis3. Bot. Bd. VIII, p. 429. 

S) Pfeffer, Le. p. 568. 

4) Pfeffer, tjber die Beziehung des Lichtes zur Regeneration von Eiweissstoflfen ans dem 
beim Keimungsprocess gebildeten Asparagin. Monatsber. d. Acad. d. Wiss. z. Berlin. Dec. 
1873. 



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468 K. SHIBATA : 

bindungen und Hexonbasen, wie Glutamin, Leucin, Tyrosin, 
Phenylalanin, Arginin u.s.w., die iiebeii Asparagin beim 
Eiweissumsatz auftreten* Namentlich hat Schulze schon ia 
einer im Jahre 1878 publicierten Arbeit die Ansicht geaussert, 
dass in Lupinenkeimlingen andere Nichteiweiss-Verbindungen, 
die neben Asparagin auftreten, auch zur Regeneration des 
Eiweisses dienen miissen^). In demselben Jahre hat Borodin 
eine allgemeine Verbreitung von Asparagin im Pflanzen- 
reiche festgestellt, dabei sprach er aus : „ Sobald irgend ein 
lebenskraftiger 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 anhaufen/'^) Demnachst fand Kellner^) in 
jungen Theilen der Graser eine bedeutende Menge Amide, und 
ausserte 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 Einfiihrung von anorganischen Salzen wie Ammoniumnitrat 
und Natriumnitrat in verschiedenen Pfianzen eine Asparagin- 
bildung bewerkstelligen konnte. Emmerling^') hat auch Amido- 
sauren als synthetische Producte angesehen, aber es fehlt an 
einem experimentellen Beweis. So kann die Bildung des Aspara- 

1)E. Schulze, Uber Zersetzang und Neubildung der Eiweisastofie bei der Keimuog von 
gelber Lupine. (Jahresber. f. Agr. Chem. 1878. p. 211). 

2) Borodin, tlber die physiologische Rolle und die Verbreitung des Asparagins in 
Pflanzenreich. Bot. Zeit. 1878. p. 826. 

3)Kenner, Landw. Jahrbucher. Bd. VIII. Suppl. 1879. 

4)Hornberger, Chemische Untersnchung uber das Wachstum der Maispflanse. Landw. 
Jahrb. 1882. 

5) Suzuki, On the Formation of Asparagin in Plants under difierent Conditions. Bn!K 
of the College of Agriculture. Bd. IF, p. 409. 

6) Emmerling; Studien uber Eiweissbildung in der Pflanze. Landw. Versnchsst. 1887. 
p. 7. 



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WACHSTUMSGESCHICHTE B. BAMBUSGEWAECHSE. 469 

gins und der anderen Amidokorper entweder durch Zerfall des 
Eiweisses oder durch geeignete Syiithese erfolgeu. Ob diese 
Oder jene geschieht muss von Fall zu Fall bestimmt werden. 
Jeden falls ist es seit Pfeflfer's bahnbrechender Untersuchung 
klar, dass die Amide und Amidosauren, deren Entstehungen in 
verschiedenen Fallen verschieden sein konnen, nachher fiir 
Eiweissregeneration verbraucht werden. Gegeuwartig ist es aber 
nocli nicht sicher ob verschiedene Amide und Amidosauren ganz 
gleichwerthig fiir Eiweissregeneration dienen. Zwar hat Hans- 
teen^) in seiner interessanten Arbeit gezeigt, dass bei Lemna 
minor verschiedene, kunstlich eingefuhrte Amide und Amido- 
sauren je nach der Qualitat der disponiblen Kohlehydrate sich 
fiir Eiweissbildung verschieden verhalten. So liegt der Gedanke 
nahe, dass die in bestimmten Keimpflanzen auftretendeii Amido- 
korper auch ungleichen Werth fiir Eiweissbildung besitzen. 
Friiher war Schulze^) der Meinung, dass das Asparagin schwerer 
verwendbar als andere Amidokorper ist und daher in Keim- 
pflanzen zur Anhaufung kommt. Aber Loew^) behauptete, dass 
das Asparagin dem Eiweiss naher steht als andere Amidokorper, 
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 ahnlichen Vorstellung gelangt. Er spricht 
die Ansicht aus, dass das Asparagin (und auch Glutamin) in den 



l)Han8teen, Beitrage zur Kenntniss der Eiweissbildung und die Bedingung der Rea- 
lisinmg. Ber. d. D. B.G. Bd. XIV, p. 362. 

2)Schulze, t}ber den Eiweissnmsatz im Pflanzenorganismus. 1880. p. 30. 

3) O. Loew, The Energy of living Protoplasm. Bulletin of the College of Agriculture 
Bd. II, p. 64. 

4)SchuIze, IJber den Umsatz der Eiweissstofle in den lebenden Pflanzen. Zeit. f. 
physiol. Chemie. Bd. XXIV, p. 60. 

Schuize, \Jber die Bildungsweise des Asparagins in den Pflanzen. Landw. Jahrb. 
1898. p. 509; p. 513. 



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470 K. SHIBATA : 

Keirapflanzeii zutn grossen Theil durch Umwandlung der Amido- 
sauren, die als directe Eiweisszersetzungsproducte betrachtel 
werdea koniien, entstehen, uQd dass die Amidosauren einmal zu 
leicht verwendbaren Amiden^) iibergefiihrt werden, bevor siesicli 
in Eiweiss verwandeln.^) Bei dieser Sachlage ist es wunschens- 
werth im concreten Falle die Localisation und das Verhalten 
von Amiden und Amidosauren zu verfolgen und damit einiger- 
massen Aufschliisse iiber die Beziehung zur Eiweissregenera- 
tioD der beiden verschiedenen Stoffe zu gewinnen. 

Bei dem vorliegenden Falle der Entwicklung der Bambus- 
schosslinge kommen Ty rosin und Asparagin reicblich vor. 
Die genannten Vertreter von beiden StoflPgruppen siud gluck- 
licherweise leicht auf microchemischem Wege bestimmbar. 

Hier lasse ich altere Angaben iiber das Vorkommen des 
Tyrosins vorangehen. Gorup-Besanetz^) fand es zuerst im 
Wickenkeimlinge* Schulze und Barbieri*) fanden es in etwas 
grosserer Menge in Kiirbiskeimlingen. Auch in Lupinenkeim- 
lingen scheint es niclit zu fehlen, da Belzung^) aus Extract der 

1) Eine entgegcngesetzte Meinung, dass das Asparagin ein fiir Eiweissregeneration wenig 
geeignetes Material aei, wurde neuerdings wieder von Prianischnikow (Landw. Vereuchs- 
St 1899. Bd. Lll, p. 347 ff.) vertreten. 

2) Unter neueren Publicationen iiber Eiweitissynthese, die nacli Vollendung meiues 
Manuscriptes in meine Hand gelangten, seien nur folgende zu erwahnen: 

Prianischnikow, Eiweisszerfall and Athmang in iliren gegenseitigen Verbal tniseen. 

Landw. Versuchs-St. Bd. Lll, p. 137. 
Hansteen, tjber Eiweisssynthese in griinen Pbanerogamen. Jahrb. f. wias. Bot. 

Bd. XXXIII, p. 417. 
Prianischnikow, Die Biickbildung der Eiweissstofie aus deren Zerfallsproducten. 

Landw. Versuchsst. 1899. p. 347. 
Schulze, t)ber Eiweisszerfall and Eiweissbildung in der Pflanze. Ber. d. E G. 1900. 

Hell. 2. p. 36. 
Emmerling, Studien uber die Eiweissbildung in der Pflanze. Landw. VersDchs^ 

Bd. LIV, p. 215. 
3)Gorup-Besanetz, Ber. d. D. C. G. VII, p. 146; p. 609. 

4) Landw. Jahrb. Bd. VII, p. 431. 

5) Belzung, Recherche sur 1. Germination etc. Ann. d. Sc. nat. Bot, Ser. VII, T. 15. 
p. 234. 



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WACHSTUMSGESCHICHTE D. BAMBUSGEWAE0H8E. 471 

Keimlinge von Lvpinus luteiLS Tyrosinkrystalle isolieren konnte, 
obwohl ihm der microchemische Nachweis des Tyrosins nicht 
gelang. Ferner fand es Schulze') in Cotyledonen keimender 
Lupinus^rteUf etiolierten Keimlingen der Lupinus anguslifoliuSj 
Endosperm von Ridnus communis und etiolierten Pflanzen von 
Tr&paeolum majus. In alien diesen Fallen ist die Menge des 
gefundenen Tyrosins iinmer sehr gering, so dass man auf micro- 
chemische Verfolgung d^sselben 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 
ofters 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. PrantP) hat Krystalle, die 
wie Tyrosin reagierten, aus in Alcohol aufbewahrten Stengeln von 
Dahlia variabilis erhalten. Borodin^) fand in Blattern der 
etiolierten Kartoffel, die mit absolutem Alcohol behandelt wurden, 
Tyrosinkrystalle* Ferner fand er dergleichen in Vicia saliva, 
Tropaeolum majus etc. Aber es ist hier zu bemerken, dass diese 
Befunde ausschliesslich von abgeschnittenen und in Wasser weiter 
cultivierten Zweigen herriihrten und gleichzeitige chemische 



l)Scbulze, tJb. d. Umsatz d. Eiweissstoffe in d. leb. Pflanze. Zeit. f. physiol. Chemie. 
Bd. XXIV, p. 58. 

2) Schulze, Iji, p. 50. 

3) Vergl. Schulze, Uber den Eiweissumsatz im Pflanzenorganismus. 1880. p. 24. 

4) Plant a, XJber die Zosammensetzung 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, t)ber die physiologische Bolle und die Verbreitung des Asparagins. Bot. 
Zeit 1878. p. 819. 



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472 - K. SHIBATA : 

Belege fehlten. Erst spater hat er*) einmal in normalen, jungen 
Dahlia-BiatleYn Tyrosin aufgefunden. Noch spater hat Leitgeb^) 
den Gehalt der Dahlia-KuoWen an Asparagin und Tyrosin 
constatiert. 

Bevor ich zur Besprechung meiner Beobachtuugen fortsch- 
reite, will ich hier die Ergebnisse von chemischen Untersuchun- 
gen Kozai's^) kurz erwahnen* Er hat die Analyse des Schoss- 
lings (Stadium IV) von Phylloslachys mitts ausgefiihrt ; sie ergab 
folgendes : 

^ Gehalt der 
Trockensubstanz. 

Eohproteinstoffe 25.12 

Fette 2.49 

Rohfaser 11.60 

Starke 3.33 

Glykose 8.15 

Andere N-freie ext. Stoffe 30.49 

Asche 9.22 

Unbestimmbare Stoife 9.60 

100.00 

Fur die Vertheilung des Stickstoffs auf ProteinstoflFe und 
nichtproteinartige Verbindungen ergaben sich folgende Zahlen: 

N in Proteinstoff'en 1.22^^ der Trockensubstanz. 

N in nichtproteinartigen Stoffen ...2.829^ „ „ 

Gesammtstickstoff 4.049^ „ „ 

So sieht man, dass die Schosslinge grosse Mengen von 
Btickstoffhaltigen Substanzen enthalten, im auffallenden Gegen- 

l)Borodin, Uber einigebei Bearbeitung yon PflanzeDSchaitte mit Alcohol entstehende 
Niederschlag. Bot Zeit. 1882. p. 589. 

2)Leitgeb, Der Grehalt 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* fiulletin 
of the College of Agriculture. Vol. I. No. 7. 



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WACHSTUMSGEfiCHICHTE D. BAMBUSGEWAECHSE. 473 

satz zum Rhizom, und insbesondere kommen die nichteiweissar- 
tigen Verbindungen in uberwiegender Quantitat vor. Kozai hat 
nach dem Schulze'schen Quecksilberuitratverfahren die seideu- 
glanzenden Nadelkrystalle aus dem Wasserauszug von Schosslingen 
erhalten, welche mit Sicherheit mit Ty rosin identificiert warden. 
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 naher verfolgt. 
Beide sind nach Borodin 'scher Methode reichlich und sicher 
nachweisbar, dabei scheint der vorhandene Zucker kein Hinder- 
niss zur Krystallisation darzubieten. 

Zunachst will ich das Verhalten von Asparagin und Tyrosin 
bei der Entwicklung der Schosslinge von Phyllostachys mitts kurz 
angeben. 

Ich konnte weder Tyrosin noch Asparagin im urmeristema- 
tischen Gewebe der ganz jungen Knospen (Stadium I) finden, 
wahrend in deren basalen, von Biindelanlagen durchsetzten Theilen 
Tyrosin schon regelmassig vorkommt. Nun die Schosslinge 
nehmen sehr langsam an Grosse zu und ihre Stieltheile werden, 
wie schon erwahnt, allmahlig verholzt. Ich beobachtete, dass das 
Tyrosin mit der Zeit im Schosslingskorper erscheint und seine 
Menge immer grosser wurde, zugleich auch das Asparagin in 
nachstehender Menge, Wenn man einen 4-5 cm langen Schossling 
in diesem Stadium (Stadium II) untersucht, so sieht man 
folgendes : Der Vegetationspunkt bleibt frei von Amidosubstanzen. 
Erst 2-3 mm unten, wo die Biindelanlagen schon difFerenziert 
waren, erscheint die erste Spur von Tyrosin im parenchymatis- 
chen Gewebe. Asparagin tritt noch weiler unten ein, wo Zucker 

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474 K. SHIBATA : 

in grosserer Menge vorkommt (etwa in der Mitte von der ganzen 
Lange des Schosslings), 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 LupinvM 
luteus. In diesem und auch im folgenden Stadium konnte ich 
weder Tyrosin noch Asparagin im Rhizom nachweisen. 

Das oben definierte Stadium III wird im Laufe des Sommers 
erreicht. Wahrend 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 grosserer Menge und viel leichter gefunden werden ; die Be- 
handlung der Gewebe (die aber eiweissarm sind) mit Millon's 
Reagens bringt uberall eine tiefere Farbung als in den vorigen 
Stadien. Die Vertheilung des Tyrosins und des Asparagins 
stimmt im Wesentlichen mit der des vorigen Stadiums iiberein. 
Dabei ist noch zu bemerken, dass das Tyrosin weniger in Nodien 
als in Internodien vorkommt. In diesem Zustande liberwin- 
tern die Schosslinge ohne bemerkbare Veranderung bis Ende 
Februar. Von jetzt ab erwacht ein regerer Process im Schoss- 
linge, und von Anfang — Mitte April wachst es schon zu einer 
betrachtlichen Grosse unter der Erde. Die Schosslinge in diesem 
Stadium (Stadium IV) werden auf dem Markt als Gemiise feil 
geboten. Die oben angegebene analytische Bestimmung Kozai's 
ruhrt auch von einem solchen Schosslinge her. In diesem Stadium 
bemerkte ich folgende Vertheilung : Der Vegetationspuukt ist 
frei von Amidokorpern, dagegen reich an Eiweiss, aber in jungen 



l)Pfeffer, Uber die Protein korner und die Bedeutung des Asparagins. Jahrb. f. wiss. 
Bot. Bd. VIII, p. 539. 



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WACHSTUMSGESCHICHTE D. BAMBUSQEWAECHSE. 475 

Scheideblattern, die zu dieser Kegion gehoren, lassen sich stets 
kleine Mengen Tyrosins nacbweisen, daher muss man bei Fest- 
stellung der Abwesenheit der Amidosubstanzen in der Spitze sie 
thunlichst von Scheideblattern befreien. Das Eiweiss, welches 
Biuretreaction giebt, ist in dieser Region besonders reichlich in 
Procambialstrangen nachweisbar. Nach Sachs^) wird das Eiweiss 
durch diese Gewebe dem Urmeristem zugefuhrt, 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 zugefiihrte Eiweiss befindet sich unter- 
halb des Urmeristems raumlich getrennt von Starke in regel- 
massig abwechselnden Zonen von je ca. 0.15 mm Dicke. Diese 
2^nen deuten schon zukiinftige Internodien und Nodien an, und 
befinden sich in den ersteren Eiweiss und in den letzteren Starke. 
Erst 4 mm unter dem Vegetationspunkt tritt die erste Spur von 
Tyrosin auf und nach unten nimmt es immer 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 Markhohle andeutet, 
stattfindet, so soil hier die Zellstreckung erst recht ausgiebig 
geworden sein. Nach unten nimmt die Menge des Tyrosins und 
des Asparagins stetig zu, und dabei iibertrifft die Menge des Tyro- 
sins bedeutend die des Asparagins. Am reichlichsten findet man 



1) Sachs, tJber die Leitiing der plastischen Stoffe durch Tcrschiedene Gewebeformen. 
Flora. 1863. 

Es ist bekannt, dass das Eiweiss unter Umstandcn die Cellulosemembran hindurch 
diosmiren kann. Vergl. Puriewitsch, Physiol. Unters. ub. Entleerung der Reserve- 
stoflfbehalter. Jahrb. f. wiss. Bot XXXI, p. 68; Pfeffer, Pflanzenphysiologie. Bd. I, p. 613. 



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476 K. SHIBATA : 

Tyrosin an Stellen, wo die Wurzelanlagen zur Bildung kommeu,^) 
so dass Tyrosin beim Schneiden des Gewebes mit dem Messer 
sofort im Zelleinneren zu Krystallen erstarrt.^) In den Bastzell- 
anlagen der Gefassbiindel, welche noch keine Wandvcrdickung 
zeigen, kommt das Tyrosin bedeutend reichlicher als im Parenchym 
vor. Die Ueberreste des Markparenchyms enthalten nur sehr 
wenig Tyrosin. Das Mi lion's Reagens bewirkt stark blutrothe 
Farbung 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 Schosslinge einer nach 
dem andern die Erde und wachsen ungemein rasch in die 
Lange. 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 halberwachseuen Internodien. Namlich 
in der unteren weichen Wachsthumszone eines solchen Interno- 
diums befindet sich das Asparagin ziemlich viel neben reichlichem 



1) ca. lOte Intemodium von unten. 

2) Siehe unten p. 482. 



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WACHSTUMSQESCHICHTE D. BAMBUSGEWAECHSE. 477 

Tyrosin. Hingegen der obere, schon erwachsene Theil desselben 
Internodiums enthalt kein Asparagin, aber Tyrosin in einer 
naliezu gleich grossen Menge wie im unteren weichen Gewebe. 
Hier werden Tyrosinkrystalle in jungen Bastzellen, die eben die 
erste Verdickung begonnen haben, reichlich ausgeschieden, aberes 
befindet sich viel weniger in Parenchymzellen. Ferner enthalten 
die Hadrom- und Leptomelemente nieraals Tyrosin. Die Nodien 
und Diaphragraen enthalten weniger Tyrosin als in Internodien 
und gewohnlich kein Asparagin. In den eben ini Wachstum 
vollendeten Internodien kommt Tyrosin noch in fast gleich 
grosser Menge vor, aber Asparagin nicht mehr. In weiter uuten 
liegenden alteren Internodien und Nodien verschwindet allmahlig 
auch das Tyrosin, und mehrere ganz erwachsene und in Verhol- 
zuDg begriffene Internodien auf der Erdoberflache sind durch- 
gehends von Amidokorpern frei, obwohl sie noch reichlich die 
Glykose, wie schon bemerkt, im Parenchym aufspeichem. 

Von den Scheideblattem habe ich hier nur zu erwahnen, 
dass sich in der basalen, weichen Wachstumszone jedes Blattes 
ziemlich viel Asparagin neben reichlichem Tyrosin befindet, und 
das erstere verliert sich schon an der Uebergangsslelle zu harten 
Theilen, wahrend Tyrosin noch weiter oben im Parenchym der 
erwachsenen Spreitentheile reichlich vorkommt. So bemerkt 
man hier ein ganz ahnliches Verhaltniss wie im Halm. 

In einige mm hoher Wurzelanlage, sowohl in Periblem wie 
in Plerom, lasst sich fast kein Tyrosin nachweisen, obwohl dicht 
daruuter 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. 
AUerdings kommt Tyrosin nur in sehr kleiner Menge im 
Wurzelparenchym vor, so dass die microchemische Nachweisung 

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478 K. SHIBATA : 

immer scliwierig ausfiihrbar ist. Noch sparlicher komnit 
Asparagin in wacbsender Region vor. Diese Umstande konnen 
zura Theil dadurch erklart werden, dass die Wurzeln nur lang- 
sam wachsen und demgemass hier der ausgiebige Eiweissumsatz 
nicht stattfindet. 

Die mit dem oben angegebenen ganz ubereinstimmende 
Vertheilungsweise des Asparagins und des Tyrosins habe ich auch 
in Schosslingen folgender Arten constatiert : 

Phyllostachys bambusoideSj 

Phyllostachys puberula, 

Bambusa palmata. 
In den von mir in dieser Beziehung untersuchten Arundi' 
Tiana- Arten, namlich : 

Arundinaria japontca, 

Arundinaria quadrangularisy 

Arundinaria Matsumurce, 

Arundinaria Hindsiiy 
zeigte Asparagin auch das gleicbe Verhalten, wahrend ich 
Tyrosin nur schwierig auffinden konnte, in auffallendem Gegen- 
satz zu PhylloslachyS' Arten. Wie dies zu Stande kommt ist mir 
unbekannt. 

Ferner ist hier zu bemerken, dass ich in Rhizoraen von 
Bambusa palmata Asparagin in geringer Menge nachweisen konnte, 
wahrend es mir bei PhyUostachys-Arten nicht gelang* 

Aus dera oben erorterten Befunde lasse ich folgende vier 
Satze gelten, namlich : 

1. Tyrosin ubertrifflt Asparagin in Menge. 

2. Tyrosin tritt in der Nahe vom Vegetationspunkt aui, 
dagegen kommt Asparagin noch weiter unten gleichzeibg 
mit Glykose zum Vorschein. 



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WACHSTDMSGESCHICHTE D. BAMBUSGEWAECHSE. 479 

3. Asparagin verschwindet aus Nodien und Internodien 
sobald ibre Streckung aufhort, wahrend Tyrosin noch 
lange Zeit in denselben erhalten bleibt. 

4. Tyrosin verscbwindet zuletzt aus ganz erwachsenen und 
in Zellwandverdickung begriffenen Internodien und 
Nodien. 

Nun miissen die einmal vorhandenen und allmablig ver- 
schwindenden Amidokorper, 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 Schosslingsgeweben angehauft werden 
miissen, was durchaus nicht der Fall ist. Zwar babe ich weder 
Ammoniak noch Nitrat mit Nessler's Reagens resp. Dipbeny- 
lamin-Scbwefelsaure in den betreffenden Geweben nacbgewiesen. 

Bei der Betbeiligung an diesem Eiweissregenerationsprocess 
scbeint, wie aus den oben erwabnten Tbatsacben ersicbtlich ist, 
Asparagin viel leicbter verwendbar zu sein und so kommt es 
nur an Stellen, wo regere Eiweissbildungsprocesse stattfinden, 
vor. Auch in diesem Sinne lassen sicb die folgende Beobacb- 
tungen erklaren. In verkiimmerten Schosslingen von Phyllostachys 
milisy die taglicb nur einige mm wacbsen, wabrend nebenbei 
stehende kraftige Exemplare taglicben Zuwacbs von mebr als 
70 cm zeigten, konnte ich in verschiedenen Internodien Aspara- 
gin niemals auffinden, dagegen kam Tyrosin dort reichlich vor. 
Ganz ahnlich verbalten sich die Rhizomspitzen von Phylloslachys 
mitis und Phyllostachys bambusoideSj die im Spatherbst (October) 
untersucht wurden, wobei sie ausserst langsam wacbsen. In 
verschiedenen Internodien derselben konnte ich trotz vielfacher 
Bemiihungen kein Asparagin mit Sicberheit nachweisen, wabrend 
Tyrosin dort reichlich vorkommt. Dass das Vorkommen von 



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480 K. SHIBATA : 

Asparagin stets mit der lebhaften Stoffbildung bei echneller 
Streckung verbundeu ist, erglebt sich auch aus folgender Bemer- 
kung Pfeflfers*) : „ War friiher die Wurzel das lebhaftest wach- 
sende Organ des Keimpflanzchens, so ist dieses jetzt das Stamm- 
chen geworden und dementsprechend weudet sich jetzt der 
Hauptstrom von Glykose und Asparagin in dieses.'* 

Hingegen verhalt sich das Tyrosin viel trager in dieser 
Beziehung, so dass es in schon erwachseuen Theilen lange Zeit 
ziiruckbleibt. Beim Verschwinden des Tyrosins aus ganz er- 
wachsenem Internodium wird ein Theil m loco verbraucht, aber 
ein anderer Theil wird vielleicht den oberen wachsenden 
Internodien zugefiihrt und dabei miissen die jungen Bastelemente 
als Leitungsbabnen benutzt werden, wie besonders reichlicher 
Gehalt an Tyrosin es vermuthen lasst. 

In jeder Hinsicht sind Asparagin und Tyrosin nicht von 
gleichem Werthe. Das erste ist ausgezeichneter Eiweissbaustoflf. 
wahrend das zweite es nur bis zu einem gewissen Grade ist, 
Soweit es leichte Verwendbarkeit ^es Asparagins anbetrifll, 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, 

l)Pfe£rer, Untersnchangcn iiber die Proteinkorner und die Bedeutang des Asparagins. 
Jahrb. f. wiss. Bot. Bd. VIII, p. 648. 

2)Han8teen, tjber Eiweisssynthese in griinen Phanerogamen. Jahrb. f. wiss. Bot. 
Bd. XXXm, p. 449, p. 485. 



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WACHSTUMSGE8CHI0HTE D. BAMBUSGEWAECHSE. 481 

die wohl als Bildungsstatte der organischen Stickstoffverbindungen 
betrachtet werden diirfen, in alien untersuehten Fallen niemals 
Tyrosin nachweisen. Ferner giebt der Blutungssaft, der beim 
Stoffabfuhr vom Rhizome eine wichtige RoUe spielt, keine Tyro- 
sinreaction. Dafiir sprechen noch folgende Versuche^ die ich 
wiederholt ausgefuhrt habe. Wenn man irgend eine abgeschnit- 
tene Rhizomspitze oder einen Sehossling von PhylloBtachys miliSf 
Phylloslachys bambusoides oder Phyllostdchys puberula in destilliertes 
Wasser stellt und am Licht oder im Dunkeln verweilen lasst, 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 
nachtragliche synthetische Bildung von Tyrosin wohl ausgeschlos-- 
sen, und die beobachtete Tyrosinzunahme muss allerdings auf 
die Eiweisszersetzung zuriickgefiihrt werden. 

Mit dem Asparagin ist die Sache schwieriger zu entscheiden. 
Obwohl ich in oben erwahnten Versuchen die gleichzeitige 
Asparaginbildnng gewohnlich nicht beobachten konnte, ist es 
naturlich nicht ausgeschlossen, dass bei der Eiweisszersetzung 
hierbei entstandenes Asparagin schnell zur Eiweissregeneration 
verbraucht wurde und demgemass nicht in gleichem Masse wie 
Tyrosin zur Anhaufung kam. Andererseits mag die oben erwahnte 
Localisation des Asparagins in kraftig wachsenden Theilen dadurch 
zu Stande gekommen sein, dass das im Rhizome fortwahrend 
gebildete Asparagin mit dem Blutungssaft den wachsenden 
Schosslingen zugefuhrt und in den betreffenden Theilen ange- 
hauft wurde^). Gegenwartig haben wir drei Moglichkeiten in 
Bezug auf Asparaginbildnng: erstens durch directe Eiweiss- 



1) Naturlich muss das Material der in wachsenden Schosslingen auf eine so erhebliche 
Weise umgesetzten Eiweisssto/fe von Bhizoroen entstammen. 



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482 K. SHIBATA : 

zersetzung/) zweitens durch Synthese aus Amraoniak^) und 
drittens durch Umwandlung von Amidosauren etc.^) Ob die eine 
oder andere von diesen Moglichkeiten in unserem Falle zutrifil 
muss vorlaufig unentschieden bleiben. 

Nun gehe ich zur Besprechung der interessanten Loslichkeits- 
verhaltnisse des Tyrosins iiber* Bei der Untersuchung der Schoss- 
linge von Phyllostachys mitts im IV und V Stadium babe ich 
gefunden, dass alle jungen, noch mit plasmatischem Wandbeleg 
versehenen Bastelemente und oft auch parenchyraatische Zellen 
mit schonen Tyrosin-Nadelbiischeln erfiillt 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 lasst es sich bald feststellen, dass 
das Tyrosin in den intacten lebenden Zellen ganz gelost im Zellsaft 
vorkommt und nur erst in den beim Schneiden geoffneten Zellen 
zu Krystallen erstarrt. Man kann diese Thatsache mit aller 
Bestimmtheit in folgender Weise beweisen : ein 3-4-zelllagendicker 
Langsschnitt des tyrosinhaltigen Internodialgewebes wird zuerst 
durch Herumschwenken in Wasser von den an den Schnittflachen 
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 lasst. So 
wird hier das ausserst schwerlosliche Tyrosin^) in so hohem 
Masse im Zellsaft in Losung gehalten, dass es sich schon nach 



l)Pfeffer, Pflanzenphysiologie. Bd. I, p. 464. 

2)0. Loew, Die chemische Energie der lebenden !Zelle. p. 77; p. 78. 
3)E. Schulze, Ub. d. Umsatz d. Eiweissstoffe in der lebenden Pflanzen. Zeit f. phjsiol. 
Chemie. Bd. XXIV, p. 63. 

4) 1 Theil Tyrosin ist loslich in 1900 Theil Wasser bei 16°C. 



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WACHSTUMSGESCHICHTE D. BAMBUSOEWAECHSE. 483 

blosser mechanischer Verlelzung der Protoplasten als Krystalle 
abscheiden lasst. Die Todtung des Gewebes durch Chloroform- 
dainpf, Osmiumsauredampf sowie Erhitzung bewirkt ebenfalls die 
Abscheidung der Tyrosinkrystalle. Ferner kommt in den durch 
39^ KNOs-Losung sehr stark plasmolysierten Bastzellen Tyrosin 
nach langer Zeit nicht als Krystalle zum Vorschein^), aber nach 
A^erletzung der plasmolysierten Zellen treten die Krystalle bald 
hervor. Wie solcli eine hohe Loslichheit des Tyrosins zu Stande 
kommt ist schwer zu beantworlen^). 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 lasst sich nicht in 
Schosslingen von I, II und III Stadien zeigen. Ebenso verhalten 
sich die tyrosinarmen Schosslinge von Arundinaria- und Bambusa" 
arten. Die Schosslinge von Phyllostachys puberula und Phyllo- 
stocky 8 bambusoides zeigen ganz gleiches Verhalten wie im oben 
dargestellten Fall von Phyllostachys mitis. 

Ausaerdem scheint sich ein Theil des Tyrosins auch in 
die Zellwande einzulagern, da die Zellwande der jugendlichen 
Bastelemente und spater auch des Parenchyms immer starker 
roth durch Millon's Keagens gefarbt werden, in dem Masse, 
dass Tyro3in in den Zellen selbst abnimmt. Bekanntlich haben 
friiher Correns^) und Fischer^) die Vermuthung ausgesprochen, 
dass die sogenannte Eiweissreaction der Zellwande veischiedener 
Pflanzen von in dieselben eingelagertem Tyrosin herriihre. Neuer- 



1) Vergl. Pfeffer, Pflanzenphysiologie Bd. I, p. 465. 

2) JedenfaUs ist die Ansicht Belzung's (Recherche chimique siir 1. Grermination etc. 
p. 219), da83 da« im Zellsaft geloate Eiweiss die Krystallisation von Asparagin, Leucin etc 
verhindern soil, ganz unzutreffend, denn in unserem Falle wird die Krystallisation schon 
durch blosse mechani^che Verletziing des Protoplastes im Zellsaft eingeleitet . 

3) Correns, Uber die vegetabilische Zellmembran. Jahrb. f. wiss. Bot Bd. XXVI, p. 616. 
4j Fischer, Zor Eiweissreaction der Zellmembran. Ber. d. D. Bot. (Jesells. Bd. V, p. 429. 



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484 



K. SHIBATA 



dings vertritt aber Czapek*) die Ansicht, dass es sich um die 
Reaction eines phenolarfigen Korpers handelt, welchen er dem 
von ihm in Mooszellwanden aufgefundeuen Spliagnol als uahe 
verwandt betrachtet. 

Gerbstoffe und Fette. 

Gerbstoffe und Fette spielen bei Entwicklung der Bambus- 
schosslinge nur eine untergeordnete Rolle. 

Bei den von mir untersuchten PhyHoddchys- und Bamhusa- 
Arten sind Gerbstoffe in verschiedenen Theilen der Schosslinge 
uberhaupt nicht in nachweisbarer Menge vorhanden.^) In dleser 
Hinsicbt bietet Arundinaria quadrangvlaris ein abweichendes 
Verhalten dar. Ein ca. 100 cm langer Schossling zeigte folgende 
Vertheilung der Gerbstoffe : Das Urmeristem des Vegetations- 
punktes ist frei davon, und dann treten sie plotzlich in reicb- 
licher Menge im Niveau der 3ten Scheideblattanlage auf, zugleich 
mit der ersten nachweisbaren Starke. Gerbstoffe kommen in 
einigen nachfolgenden Internodien in gleicli reichlicher Menge 
vor und dann nebmen sie nacb unten ab. Dabei reagieren am 
starksten die Rindenzellen uud^die peripherischen Centralcylinder- 
parenchymzellen. Selbst in erwachsenen Internodien und Nodien 
in der Nahe der Erdoberflacbe ist eine schwache Reaction be- 
raerkbar. Die Wurzelhaube enthalt auch kleine Mengen der 
eisenWauenden 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 Fallen, z. B. bei Vicia^ 

1) Czapek, Zur Chemie der Zellmembran bei den Laub- nnd Lebermoosen. Flora. 189d- 
Bd. 86, H. 4. 

2) Abgesehen von 'kleinen Mengen in Wurzelhauben, Scheideblattern etc 



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WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 485 

JSelianthu8^)f Zuckerrohr^) u.s.w, Im vorliegenden Falle scheinen 
die autochthonen^) Gerbstoffe sich .aplastisch zu verhalten, well 
hier Zucker, Starke u.a. auf ganz gleiclie Weise vorkommen wie 
bei anderen gerbstoflffreiea Arten. 

Auch kommt den Fetten hochstens niir eine locale Bedeut- 
UDg zu als Baumaterial der verkorkenden oder verholzendeii Zell- 
wande, so zum Beispiel verschwinden die kleinen Fetttropfen in 
einer subepidermal en Zellsehicht der Wurzel schon bei eintreten- 
der Verkorkung der Zellwande. In jeder alteren Halmparen- 
chymzelle von Arundinaria japonicaj A. Ifindsii, Bambusa 
floribunda etc. befindet sichje ein olartiger gelber Tropfen, meist 
in Verbindung mit Calciumoxalatdrusen, Diese kugeligen 
Grebilde unterscheiden sicli von echten Fetttropfen dadurch, dass 
sie niemals mit Alkannatinktur sich farben. In vielen Punkten 
stimmen sie mit dem zuerst von Monteverde^) in Gramineen 
aufgefundenen „ Harzkorper** iiberein. 

MiNEBALSTOFFE. 

Ich habe in verschiedenen Jahreszeiten die Vertheilung der 
Mineralstoffe in den Reservestoflfbehaltern und den Schosslingen 
v^rfolgt. Als Untersucliungsmaterial dienten mir hauptsachlich 
Phyllostachys mitis und Phyllostdchys bambusoides. 

Ich konnte in den Rhizomen, die schon betrachtliche 
Quantitat der Starke aufgespeichert haben, die Mineralstoffe 
leicht auffinden. Sie zeigten folgende Vertheilung sowohl in 
Intemodien als in Nodien : 



1) Vergl. Kutscher, Uber die Verwendnng der Gerbeaiire im Stoffwechsel der Pflan- 
zen. Flora. 1883, p. 33. 

2) Went, Chemifich-physiologische Unlereuchungen iiber das Zuckerrohr. Jahrb. f. wiss. 
Bot. Bd. XXXI, p. 297. 

3)Krau8, Gmndlinien zu einer Fhysiologie des Gerbstofies. p. 58. 
4}Moateyerde, Uber Ablagerung von Calcium- und Magnesiumoxalat iu der Pflanze. 
Bot. Centralb. 1890. Bd. XLIII, p. 327. 



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486 K, SHIBATA : 

Phosphor reichlich im Parenchym des Centralcylmders ; 
nur wenig in Siebrohren. 

Magnesium reichlich vorzugsweise in Siebrohren. 

Kalium reichlich im Parenchym. 

Calcium nicht nachweisbar in Schnitten ; nur kleine 
Mengen in Aschen. 

Schwefel nur spurweise in Schnitten.^) 

Chi or ziemlich reichlich im Parenchym ; aber fast keins in 
Siebrohren und anderen Elementen der Grefassbundel. 

In den Wurzeln konnen die obengenannten Stoffe auf uber- 
einstimmende Weise aufgefunden werden. Die Nitrate^) sind 
in Bhizomen nur sehr wenig vorhanden, aber zeitweilig etwas mehr 
in der Wurzelrinde. Sie kommen hier also iiberhaupt nicht zur 
nennenswerthen Aufspeicherung. Leitgeb^) hat friiher gezeigt, 
dass der Phosphor in DahliakuoWen hauptsachlich als Calcium- 
phosphat vorkommt. Hier ist aber dies nicht der Fall, da ich 
keine losliche Ca-Verbindung in den Zellen auffinden konnte. 
Die Siebrohren der Rhizome und Wurzeln enthalten, wie schon 
erwahnt, nur geringe Mengen des Phosphors, dagegen reichlich 
Magnesia. Daher scheint Magnesium theils als Phosphat, iheiis 
als lockere organische Verbindung in Siebrohren vorzukom- 
men. Schimper*) und Zacharias'') haben auch im Siebrohr- 
ensafte von Oucurbita^ Wistaria^ Aristolochia und Menispermum 
reichliche Mengen des Magnesiums aufgefunden. 

l)Schimper (Zur Frage der Assimilation der Mineralsalze. Flora, 1890. p, 223j hat 
auch in den meisten Khizomen keine Sulfatreaction erhalten und es dem Yorhandensein von 
Krystallisation verhindernden Substanzen in Zellen zugeschrieben. 

2) Vergl. Mo 1 isc h, Uber microchem. Nachweis von Nitraten. Ber. d. D. Bet G. Bd. I, 
p. 154. 

3) Leitgeb, Uber die durch Alcohol in DoA/ta-Knollen hervorgerufenen Krystalle. Bot 
Zeit. 1887. p. 29. 

4)Schimper, Zur Frage der Assimilation der Mineralsalze. Flora, 1890, p. 228. 
5)Zacharia8, Uber d. luhalt d. Siebrohren von Cucurbita. Bot. Zeit. 1884. p. 71. 



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WACHSTUMGSESOHICHTE D. BAMBUSGEWAECHSE. 487 

Schritt fur Schritt mit der Entleerung der Kohlehydrate 
verschwiuden auch die Mineralstoffe aus dera Rhizompareiichym. 

Die Nitrate sind schon im IV Stadium in Khizomen und 
Wurzeln nicht mehr nachweisbar. Ebensowenig konnte icli im 
Blutungssaft die Nitratreaction erhalten. Wahrscheinlicli werden 
die Nitrate hierbei fortwahrend zu organischen Verbindungen 
verarbeitet.^) 

An den unteren, mit zahlreichen jungen Wurzeln besetzten 
Theilen der Schosslinge besitzen Phosphor und Kalium eine 
andere Vertheilungsweise als im Rhizome ; sie kommen nun 
reichlicher in den Biindeln vor. Das Magnesium befindet 
sich hier auch in Siebrohren bevorzugt. Ich konnte niemals 
die Nitrate im Schosslingskorper auffinden.^) Da sie auch im 
Rhizome fehlen, so ist es hochst wahrscheinlich, dass iiberhaupt 
nur wenig Nitrat als solches in den Schossling eingefiihrt 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 iiberzeugen 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. Merkwiirdigerweise 
komnat Phosphor fast ausschliesslich in den eiweissreichen Pro- 
cambialstrangen 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 Filzsjmbiont an der Stickstoffassimilation 
betheiligt, muss zur Zeit dahingestellt bleiben. 

2) tJbrigens ist es klar, dass die Nitrate sich nicht Lei so lebhafter Eiweisszersetzung 
bilden. (Vergl. Schulze, Uber d. Vorkommen von Nitraten in Keimpflanzen. Zeit. f. 
phjsiol. Chemie. Bd. XXII, p. 83). 

3) Ich habe auch die Lilienfeld'sche Methode fiir Erkennung der Localisation des 
Phosphors mit Erfolg benatzt (vergl. Str as burger, Das botanische Practicum. III. Aufl. 
p. 144). 



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488 K. SHIBATA : 

schen Phosphorverbindungen nicht mehr nachweisen, sondern 
grosse Mengen organischer Verbindungen*). Magnesium tritt 
ebenfalls in der Nahe der Spitze fast ausschliesslich in den 
Biindelanlagen auf, aber kleine Mengen sind auch in den 
eiweisshaltigen Internodialzonen vorhanden. Beachtet man die 
bevorzugten Vorkommnisse des Magnesiums in Siebrohren, 
Biindelanlagen, Internodialzonen und auch im Urmeristem, so 
darf man wohl annehmen, dass es irgend eine wichtige KoUe 
bei Eiweissumsatz oder Eiweisswanderung spielt^) Kalium lasst 
sich in der Asche des Vegetationspunktes reichlich nachweisen. 
Hingegen ist die Schwefelsaure nicht direct im Vegetations- 
punkt nachweisbar, obgleich sich schon ca. 2 cm weiter unten 
eine ziemlich starke Reaction zeigt. Das in minimaler Menge 
zugefiihrte Calcium wird in einiger Entfernung vom Vegetations- 
punkte als Kalkoxalat niedergeschlagen. Chlor lasst sich ziemlich 
viel in eiweissreichen Internodialzonen nachweisen, Im Urme- 
ristem scheint es jedoch ganzlich zu fehlen. 

Wenn die Schosslinge fiber die Erde emporwachsen und die 
Internodien nach einander ihre definitive Laqge 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 Thatigkeit entfalteten und began nen 
die Bodensalze aufzunehmen. 

In der ersten Entwicklungsphase der Wurzel ist eine starke 
Ansammlung der Mineralstoffe im meristematischen Gewebe leicht 
constatierbar. In etwas langer erstreckten Wurzeln findet eine 
ahnliche Ansammlung in der Spitze statt. Phosphor und Mag- 

l)VergL Schimper, Le, p. 224. 

2). VergL Ho r n b erg e r , Chemiiche Untersuchungen iiber das Wachstum der Maispflaoxe. 
Landw. Jahrb. 1882. p. 278. 



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WACHSTUMSGESCHrCHTE D. BAMBUSGEWAECHSE. 489 

nesium kommen, wie in Schosslingen, hauptsachlich 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 aafgenommenen Salzen herriihrt. 

In den jiingsten Scheideblattanlagen in der Umgebung des 
Vegetationspunktes lasst sich sehr friih eine Ansammlung von 
Phosphor und Magnesium beobachten. Uebrigens bedarf dies 
keiner Besprechung mehr. 

Vn. Ueber die Entleenmg der Heservestoffe. 

Die Versuche von Hansteen^) und Puriewitsch^) haben 
die Thatsache festgestellt, dass bei Endospermen, Samenlappen, 
KnoUen und Rhizoraen die Entleerung der deponierten Reserve- 
stoffe mehr oder minder selbstthatig stattfinden kann. Nun schien 
es mir geboten zu bestimmen, erstens inwieweit die Entleerung des 
Rhizoms unabhangig von wachsenden Schosslingen vor sich gehen 
kann, und zweitens in welchem Grade die Entwicklung der 
Schosslinge durch die totale oder partielle Separierung vom 
Rhizomsyslem beeinflusst wird. Zu diesem Zweck habe ich 
Mitte April eine Anzahl kmftig wachsender, unterirdischer 
Schosslinge aufgesucht und verschieden tiefe Einschnitte in ihre 
Stieltheile und benachbarte Rhizominternodien gemacht. Die 
betreffenden Rhizomtheile waren in diesem Stadium mit Starke 
strotzend erfiillt, wie ich durch die Musterung zahlreicher 
Exemplare iiberzeugt war ; es zeigte sich folgendes : 

l)Han8teen, Uber die Ursache der Entleerang der Heservestoffe aus Samen. Flora. 
1894 Bd. 79, p. 419. 

5)Puriewit8ch, Phjsiologische Untersuchungen uber die Entleerung der Reserve- 
stoffbehalter. Jahrb. f. wiss. Bot. Bd. XXXI, p. 1. 



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490 



K. SHIBATA : 





Starke 


Glykose 


Rohrzucker 


Rindenparenchym 


strotzend erfullt (5) 


sehr wenig 


ziemlich viel 


Centralcylinderparen- 
chym 


strotzend erfiillt- 
recht viel (5-4) 


desgl. 


d««l. 


Markparenchym 


strotzend erfullt (5) 


desgl. 


desgl. 



Ill der folgenden Tabelle stelle ich die Ergebnisse einiger 
Versuche zusammen : 





Opera- 
tionen 


Inhalt der Rhizome 


Bemer- 
kuDgen 


Starke 


Zucker 


operiert: 20. April 
untersucht: 30. Mai 


Durchschneiden 
im 68ten Inter- 
nodium vor und 
im 3ten Int. 
hinter dem 
Schosslinge ; 
Durchschneiden 
im Stiel 


lUnden- und 
Centralcylinder- 
parenchym : 
keine Starke 
(0); Mark- 
parenchvm : 
wenig Starke (2) 


Rinden-, Cen- 
tralcylinder- 
und Mark- 
parenchym: fast 
kein Zucker 


Zuwachs 

des 

Schosslings 

-0 


operiert: 20. April 

n. 

untersucht: 30. Mai 


wie oben 


wie oben 


Rinden-, Cen- 
tralcylinder- 
und Mark- 
parenchym : 
wenig Zucker 


Zuwachs 

des 

Schosslings 

=0 


operiert: 16. April 

in. 

untersucht: 18. Mai 


Durchschneiden 
imSten Int. vor 
dem Schosslinge; 
Durchschneiden 
im Stiel 


keine Starke 
(0) 


keine Glykoge; 
kein Bohr- 
zucker 


Zuwachs 

des 

Schosslings 

=0 


operiert: 16. April 
IV. 

untersucht: 18. Mai 


Durchschneiden 
im Sten Int. vor 
dem Schosslinge; 
1.5cmtieferEin- 
sohnitt ias 2ten 
Int. hinter dem 
Schosslinge 


keine Starke 
(0) 


keine Glykose; 
sehr wenig 
Rohrzucker 


Zuwachs 

des 

Schoaslings 

=2.5 cm 


operiert: 20. April 
untersucht: 18. Mai 


Durchschneiden 
im Stiel 


fast keine 
Starke (0-1) 


keine Glykose 


Zuwachs 

des 

SchossluiKS 

=0 


operiert: 16. April 
untersucht: 19. Mai 


Durchschneiden 
im Sten Int vor 
dem School inge 


wenig Starke 


fast keine 
Glykose 


Zuwachs 

des 

Schosslings 

=10 cm 


operiert: 16. April 
VII. 

untersucht: 19. Mai 


Durchschneiden 
im 2ten Int. 
hinter dem 
Schosslinge 


keine Starke 
(0) 


wie oben 


Zuwachs 

des 

Schosslings 

=0 



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WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 491 

AUe oben angefiihrten Versuche ergaben ubereinstimmend, 
dass die Entleerung, zumal die Starkeauflosung in bestimmten 
Ehizorupartien unabhangig von Sehosslingen vor sich gehen kann. 
Puriewitsch hat bei den Versuchen mit den Rhizomen von 
OuTcuma und Rvdbeckia gezeigt, dass in solchen Rhizomen eine 
partielle Entleerung selbstthatig stattfand, wenn die eontinuierliche 
Ableitung der Losungsproducte mittelst Gyps besorgt wurde^). 
Nun in meinen Versuchen war die Bedingung fur derartige Starke- 
entleerung besonders giinstig. Die zahlreiehen kraftigen Wurzeln 
an Rhizomknoten erzeugten zur Zeit einen ansehnlichen Blutungs- 
druck und der zuckerhaltige Blutungssaft wurde immer fort von 
den Schnittflachen der Rhizome ausgeschieden. Damit wurde 
eine fortwahrende Wegfiihrnng der Losungsproducte erzielt, 
welche eine so volkommene Entleerung herbeifiihrte. 

Ferner ist es aus obigen Versuchen ersichtlich, dass die 
Entwickelung der Schosslinge durch jeden operativen Eingriff in 
benachbarte Rhizominternodien — d.h. durch jede Herabsetzung 
des Blutungsdrucks — bald sistiert wird. 

Macht man in der Nacht oder friihmorgens ein Bohrloch in 
ein beliebiges Internodium des kraftig wachsenden Schosslings, so 
quillt bald ein zuckerhaltiger klarer Saft hervor. Am 19. Mai 
wurde der Blutungssaft von einem mittleren Internodium eines 
ca. 1.5 Meter hohen Schosslings von Phyllostachys puberula 
gesammelt. In 100 ccm dieser Fliissigkeit fand ich 0.289 gr 
Glykose. Ausserdem enthalt der Blutungssaft eine kleine Meuge 
der Amide, da er nach Beseitigung des Eiweisses^) eine starke 
Triibung beim Zusatz von Quecksilberoxydnitrat giebt. 



1) Puriewitsch, l.c. p. 28. 

2) Nach der S t ii t z e r *3chen Methode. 



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492 K. SHIBATA : 

Schroter*) schrieb : „ Spater, in den ausgewachsenen Glie- 
dern, findet sich oft ein klares Wasser, das in manchen trockenen 
Gegenden den Reisenden ein hochst willkommener Fund ist." 
Derartige mit Wasser erfiillte Markhohlen 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 Flussigkeit, die sich in der 
unteren Internodialhohle von PhyUostachys mitis befand, 0.269?^. 
Wenn man friihmorgens einen Bambasbusch besucht, so wird 
man einen formlichen Regen von Wassertropfen aus den Scheide- 
blattspitzen der wachsenden Schosslinge bekommen.^) Dies in 
bekannter Weise von Blattspitzen ausgeschiedene Wasser enthalt 
auch Glykose neben einer Spur von Amiden. Eine Zuckerbes- 
timmung der am 28. April gesammelten Flussigkeit ergab 
0.09589^ Glykose. 

Alle diese Thatsachen weisen darauf hin, dass eine erheb- 
liche Menge der Kohlehydrate und vielleicht auch Amide mit 
dem Blutungssaft den Schosslingen zugefiihrt werden. Dadurch 
werden die Schosslinge mit den Baustoffen geniigend rasch ver- 
sorgt.^) Die oben erorterten Bauverhaltnisse der Stieltheile lassen 
sich auch nicht anders denken, als dass hier der ausgiebige 
Stoflftransport nur durcb die Biindel und zwar durch die wohl 
ausgebildeten Gefasse geschehen kann. 



l)8chroter, Der Bambus und seine Bedentung als Nutzpflanze. p. 14; Cohn, Cber 
Tabaschir. p. 376. 

2)Moliscb, th)er das Blaten tropiscber Ilolzgewacbse. Ann. d. Jard. Bot. Bait 
1898. Sappl. II, p. 23. 

3) Vergleiche bierzu: Strasburger, Bau und Verricbtungen der Leitungsbabnen. 
p. 877; Fiscber, Beitrage zur Pbysiologie der Holzgewacbse. Jahrb. f. wiss Bot Bd. XXII, 
p. 76; p. 150. 



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WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 493 

VII. Zusammeufassung. 

In Obigem habe ich die wesentlichen Ziige der Waclis- 
tumsgeschichte der Bambusgewachse darzustellen versucht. Die 
Hauptresultate werden hier kurz in folgenden Worten zusam- 
mengefasst : 

1. Die Starke wird in parenehymatischen Zellen der 
Rhizome, HalmeundWurzelnals Hauptreservestoflfabgelagert. 
Die Vermin derung derselben im Winter wurde nicht beobachtet, 
wahrend zur Zeit des r^schen Austreibens von Schosslingen eine 
unverkennbare Starkezunahme (transitorisch) in benachbarten 
Rhizomtheilen constatiert wurde. 

2. Die Glykose dient als Baumaterial in wachsenden Theilen 
der Schosslinge and ist in schon fertig gestreckten Internodien 
derselben transitorisch reichlich aufgespeichert. 

3. Der Rohrzucker tritt als das Losungsproduct der 
Starke im Parenchym der Rhizome und Halme auf. 

4. In schnell wachsenden Schosslingen fand eine ausgiebige 
Eiweisszersetzung slatt, dabei trat Tyrosin in bedeutender Menge 
auf. 

Tyrosin und Asparagin zeigen einen weitgehenden 
Unterschied in ihrem Verhalten. Tyrosin wird schwerer und 
langsamer fiir Eiweissregeneration verbraucht, so dass es in schon 
erwachsenen Theilen eine Zeit lang zuriickbleibt. Hing^en ist 
Asparagin leicht und rasch dazu verwendet und kommt nur 
an Stellen vor, wo eine lebhafte Stoflfbildung stattfindet. 

5. Gerbstoffe kommen nur in Schosslingen einzelner Arten 
vor, und Fette spielen hierbei keine wichtige RoUe sowohl als 
Wanderstoflfe wie als Reservestoflfe. 

6. Phosphor, Kalium, Magnesium und Chlor werden 



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494 K. SHIBATA : 

in deu ReservestoflFbehalteru aufgespeichert, dabei kommt Mag- 
nesium vorwiegend in Siebrohren vor. Calcium und Schwefel 
sind gewohnlich nicht direct nachweisbar. 

7. Die Mineralstoffe wandern bei rascher Eutwicklung der 
Schosslinge schnell von den Rhizomen aus und werden in den 
waclisenden 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 
Schosslinge deutlich nachweisbar. 

8. Die vom Boden aufgenommenen Nitrate werden wahr- 
scheinlich schon in den Wurzeln und Rhizomen zu organischen 
Verbindungen verarbeitet. 

9. Die Auflosung der Starke und die Entleerung der L6- 
sungsproducte aus den Rhizomen konnen unabhangig von der 
Entwickelung der Schosslinge fortgehen. 

10. Der ausgiebige und schnelle Stoflftransport nach wachsen- 
den Schosslingen von den Rhizomen kann in Wasserbahnen 
geschehen. Dafur sprechen vor allem die Blutungserscheinungen 
der Rhizome und Schosslinge und die Bauverhaltnisse der Schoss- 
lingsstiele. 

Botanisches Institut 
Juni 1890. Kaiserl. Universitat 

zu Tokio. 



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WACHSTUMSGESCHICHTE D. BAMBUSGEWAECHSE. 495 



Verzeichniss deb untersuchten Arten.^) 

Phyllostachya mitts Riviere. (Nom. Jap. Moschchiku,) 
Phyllostaehys bambusoides Sieb. et Zuoc. (Nom. Jap. Ma-dake.) 
Phyllostachys bambusoides Sieb. et Zucc. var. aurea Makino. (Nom. Jap. 

Hotei'ChiJcu.) 
Phyllostachys puberula Mimro. (Nom. Jap. Ha-chiku.) 
Phylloslachys jniberula Munro. var. nigra, (Nom. Jap. Kuro-chiku.) 
Phyllostachys Kuma^asa Munro. (Nom. Jap. Okame-sasa.) 
Arundinaria japonica Sieb. et Zucc. (Nom. Jap. Ya-dake.) 
Arundinaria Simoni Riviere, (Nom. Jap. Me-dake.) 
Arundinaria Matsumurce Hackel. (Nom. Jap. Kan-chiku,) 
Arundinaria quadrangularis Makino. (Nom. Jap, Shikaku-dake.) 
Arundinaria Hindsii Munro. (Nom. Jap. Kansan-chiku,) 
Arundinaria Bindsii Munro. var. graminea Bean. (Nom. Jap. Taimin- 

chiku.) 
Arundinaria Fortunei Riviere. (Nom. Jap. Chigo-sasa.) 
Arundinaria variabilis Makino. (Nom. Jap. Ne-sasa.) 
Arundinaria pygmcea Mitf. (Nom. Jap. Oroshima-chiku.) 
Arundinaria Narihira Makino. (Nom. Jap. Narihira-dake.) 
Arundinaria Toots ik Makino. (Nom. Jap. To-chiku.) 
Barnbusa borealis* Hackel. (Nom. Jap. Suzu-dake.) 
Bambusa palmata* Marliac. (Nom. Jap. Chimaki-sasa.) 
Bambiisa Veitchii* Carriere. (Nom. Jap. Kuma^asa.) 
Bambusa panicvlata^ Makino. (Nom. Jap, Nemagari-dake.) 
Bambusa nipponica^ Makino. (Nom. Jap, Miyako-sasa.) 
Barnbusa ramosa* Makino. (Nom. Jap. Azuma-sasa.) 



l)Die ausfiihrliche Beschreibnng der hier angefuhrten Arten findet man bei Makino, 
Bambnsaoese Japonicse (The Botanical Magazine^ Vol. XIV, Nr. 156, p. 20 ff.), die beige- 
^gten japanischen Namen sollen zam Heransfinden der betrefifenden Arten in der genannten 
Sclirift dienen. 

Die mit * bezeichneten Arten gehoren meiner Ansicht nach nicht eigentlich za BatnbuMf 
sondem sie wiirden vielleicht eine selbetandige Gbittnng bilden. (Vergl. oben p. 446 und auch 
Makino, Ic p. 20). 

An dieser Stelle spreche ich Herm M a k i n o fur die von ihm giitigst vorgenommene Bestim- 
mong einiger Arten and auch Herren A so and In ami fttr ihre freundliche Unterstiitzuog 
bei einigen analytischen Arbeiten meinen beaten Dank aus. 



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496 K. SHIBATA : 

Bumbusa vulgaris Wendl. (Nom, Jap. Daisan-ckiku.) 
Bambusa nana Eoxb. (Nom. Jap. Howo-cliiku.) 
Bamhusa nana var. noi^malis Makino. (Nom. Jap. Taiho^hiku.) 
Bamhma stenostachya Hackel. (Nom. Jap. Ski-chiku.) 
Dendrocalamvs laiiflorus, Munro. (Nom. Jap. Ma-chikM.) 



Inhalt. 

I. Einleitung 427 

II. Untersuchnngsmaterial und Methodisches 429 

III. Die Bauverhaltnisse 433 

IV. Der Entwicklungsvorgang der Schosslinge 453 

V. Verhalten der Baustoffe wahrend der Entwicklung 

der SchOsslinge 458 

VI. Ueber Entleerung der Keservestoffe 489 

VII. Zusammeufassung 493 



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Erklarung der Tafeln. 

Tafel XXIL 

Fig. 1. Zwei neben einander stehende SiebroreDglieder mit zahlreichen 
Siebttlpfelii (sdp) au den Seitenwanden, aus Rhizomtnoten von 
Phyllostachya mitts, spl Siebplatte, sip Siebttipfel, Vergr. 360, 

Fig. 2. Querschnitt durch das Rhizom von Bambusa nipponica. R Rinde, 
Brg subcorticaler Bastring, cerd Centralcylinderparenchym. Vergr. 30. 

Fig. 3. Querschnitt durch das Rhizom von Arundinaria japonica. B 
Bastbander. Veipr. 30. 

Fig. 4. Die spindeliormige Anschwellung des Leptoms eines Knospenbundels 
bei der Ansatzstelle an der Rhizorabftndel. 8 Siebr5hren, gl Qeleit- 
zellen, O Geftlsse, P Parenchymzellen, c6/*carabiformartige Elemente 
Vergr. 70. 

Fig. 5. Querschnitt durch die Anschwellung. B Bastzellen, P u. cbf wie 
in Fig. 4. Vergr. 125. 

Fig, 6, Theil der langgestreckten cambiformartigen Elemente aus dem 
mittleren Theile der Anschwellung, mit Querstreifen auf den Seiten- 
wanden. Vergr. 450. 

Fig. 7. Derselbe im Querschnitt. Vergr. 450. 

Fig. 8. Die Leptomanschwellung in einem friiberen Entwicklungsstadium. 
Langsschnitt durch den Knoten. S Siebrohren, B Bastzellen, cbf 
cambiformartige Elemente. Vergr. 83. 

Fig. 9. tJbergangsstelle der cambiformartigen Elemente zum normal gebauten 
Leptom, in einem jugendlichen Zustand. Sammtliche Elemente mit 
auffallend grossen Zellkernen und reichlichem Plasmagehalt. S u. cbf 
wie in Fig. 8. Vergr. 450. 

Fig. 10. Dergleichen im fertigen Zustand. ip Tilpfel, c6/wie oben. Vergr. 
450. Figuren 4-10 beziehen sich auf Phyllostachys mitis. 

Fig. 11. Ein Gefassbtindel aus einem inneren Teil des Rhizoms von Arundi- 
naria Hindsii. S Siebrohren, gl Geleitzellen, G Gefasse, d Durch- 
lassstelle. Vergr. 125. 

Fig. 12. Eine subepidermale sclerotische Parenchymschicht des Rhizoms 
von Bambusa palTnaia. Langsschnitt. ep Epidermis, scl sclerotische 
Parenchymzellen, R Rindenzellen, Vergr. 360. 



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Fig. 13. Querschnitt durch den Stieltheil. B Rinde, B Bastbander, k 

Bastscheide des Mestombtindels, mes Mestom. Veigr. 17. 
Fig. 14. Ein Mestombundel im Stieltheile, mit vollkommen umschliessender 

Bastscheide. i Tracheiden, P, bs, S, gl, u. G wie oben. Veigr. 125. 

Fig. 13-14. Phyllostachys mitts. 
Fig. 15. Querschnitt durch den dtinnen Halmzweig von Arundimria 

pygmcea. ep u. B wie oben. Veip:. 83. 
Fig. 16. Theil desgleichen von Arundinaria japonica, ep, B n R wie 

oben. Vergr. 360. 
Fig. 17. Ein Halm-Btindel von Bambusa nana var. normalis, mit Paren- 

chymlamelle im innenseitigen Bastbeleg. par. I Parenchymlamelle, 

S, O 11. B wie oben. Vergr. 200. 
Fig. 18. Einige verschiedenartige Vorkommnisse des Parenchymgewebes 

im Bastbelege. Arundinaria Hindsii. B Bastbelege, par paren- 

chymatisches Gewebe. Vergr. 70. 
Fig. 19. Ein Halmbundel von Bambusa nana. Die durch parenchymatische 

Zellen vom Mestom abgetrennte Masse des Bastbelegs bleibt unverdickt. 

par. Z, B wie oben. Vergr. 200. 
Fig. 20. Auftreten der Starkekorner in neu diflferenzierter Parenchymlamelle. 

Vergr. 70. 
Fig. 21. Obiges im Langsschnitt. Vergr. 125. 
Fig. 22. Die durch successive Quertheilungen von Procambialzellen ent- 

standenen Parenchymzellen. pre Procambialzellen, k Kern, st Stfirke- 

kdrner. Vergr. 360. Figuren 20-22. Arundinaria Hindsii. 



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Tafel S:llIL 

Fig. 23. Querschnitt durch die jimge Wurzel von Bambnsa palmata, i.H 
innere Rindenzellen, end Endodermis mit Caspary'schen Streifen, 
per Pericambium, pjiad peripherische Hadromstrange (primordiale 
Netztracheiden), p.lep peripherische Leptomstrange. Vergr. 360. 

Fig. 24a. Peripherischer Theil der Wurzelrinde von Phyllostachys mitts, 
ep Best der Epidermis, hyp stark verdickte (an den Aussenwanden) 
subepidermale Zellen, scl peripherische sclerotische Elemente, a.R 
aussere Kindenzellen. Vergr. 200. 

Fig. 246. Desgleichen im jugendlichen Zustand. ep, hyp u. scl wie oben. 

Fig. 25. Starkverdickte Subepidermalzellen (Aussenscheide) von Phyllo- 
stachys bambusoides var. aurea. Vergr. 360. 

Fig. 26. Peripherischer Theil der Wurzelrinde von Bambusa vtUgaris, ep 
Epidermis, hyp unverdickt gebliebene Subepidermalzellen, sol peripheris- 
che Sderenchymzellen, a.R aussere Rindenzellen. Veigr. 360. 

Fig. 27. Wurzelrinde von Bambusa nana, ep, hyp, scl, a.R u, t. R wie oben. 
Vergr. 125. 

Fig. 28. Endodermis (end) und starkverdickte Pericambiumzellen (per) 
von Fhyllostachys mitis. Veig. 360. 

Fig. 29. Langsschnitt durch die Endodermis von Fhyllostachys mitis. Vergr. 
360. 

Fig. 30. Langsschnitt durch den peripherischen Theil der Wurzelrinde von 
Arundinaria Maisumurot. hyp u. scl wie in Fig. 24. Vergr. 360. 

Fig. 31. Endodermis und angrenzende Rindenzellen von Bambusa steno- 
stachya. Langsschnitt. eel ZellstoflSiuswuchse, end u. per wie obeu. 
Veigr. 360. 

Fig. 32. Dieselbon im Querschnitt end, per u. eel wie oben. Vergr. 360. 

Fig. 33. C-fbrmig verdickte Endodermiszellen von Bambusa palmata. 
Veigr. 360. 

Fig. 34. Theil des Wurzelquerschnittes von Fhyllostachys Kumasasa. ver 
Verstarkungsring, 1/ Luftrftume, end, i.R, per u. p.lep wie oben. 
Veigr. 360. 

Fig. 35. Verknupfung der Leptomstrange durch dunnwandiges Verbindungs- 
gewebe. Fhyllostachys bambusoides. ver.p. Verbindungsgewebe, 
nb.w Nebenwurzel, p.lep, i.lep, G, end wie oben. (schematisiert) 
Veigr. 70. 



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Fig. 36. Dergleichen bei Bambusa vulgaris. Veigr. 45. 

Fig. 37. QuerschDitt durch die Hauptwurzel an der Ansatzstelle der Neben- 

wurzel. Arundinaria MatsumurcB. n.lep Leptomstrang der Neben- 

wurzel, lep Leptomstrange der Hauptwurzel, end, i.R, permd oben. 

Vergr. 360. 
Fig. 38. Innerer Leptomstrang von Bambiisa vtdgaris. S Siebrohre, cb 

Cambiformzellen, rnz mechaniscbe Zellen. Vergr. 360. 
Fig. 39. Verschmelzung des inneren Leptomstrangs mit dem peripherischen. 

iJep innerer Leptomstrang, p.lep peripherischer Leptomstrang, S, 

cb wie oben. Vergr. 360. 
Fig. 40. Verschmelzung zweier peripberiscben Leptomstrange. Vergr. 360. 
Fig. 41. Director Anscbluss des Leptomstrangs an Hadromparenchj-m. 6 

Gefass, hp Hadromparencbym (Gefassbelegzellen), lep u. mz wie oben. 

Vergr. 360. 
Fig. 42. Zusammentreften zweier Hadromstrange. G, hp u. mz wie oben. 

Vergr. 360. 

Figuren 39-42. Fhyllostachya bambv^oides. 
Fig. 43. Verbindungsgewebe zwiechen inneren und peripherischen Leptom- 

strangeu. iJep, p.lep, mz u S wie oben. Vergi\ 360. 
Fig. 44. Dasselbe von Bambusa vulgaris im Ijangsschnitt, S, mz, wrf 

wie oben. Vergr. 360. 
Fig. 45. Querschnitt durch den Basaltheil der Nebenwurzel von Phyllostachys 

mitis. lep Leptomstrange, mz mechaniscbe Zellen. Vergr. 360. 
Fig. 46. Tlieil der Rinde der Nebenwurzel von Phyllostachys pubervlaj mit 

endophytischen Mycelfaden. R Rindenzellen, onyc Pilzfaden, ves 

Vesiculen, kor gelbe kornige Substanz. Vergr. 360. 
Fig. 47. Querscbnitt durch die Nebenwurzel von Phyllostachys pubenda, 

hyp Subepidermalzellen, scl peripherische sclerotische Zellen, B, fnyc, 

end, lep wie oben. Vergr. 200. 



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Fig. 48. Ein Scheideblattbiindel mit starkem Bastbeleg (B) von Phyllo* 

stachys mitis. Vergr. 83. 
Fig. 49. Querschnitt durch das Scheideblatt von Arundinaria Maisumurce 

VeTgjr. 360. 
Fig. 50. Kleinere Scheideblattbundel von Bambusa stenostachya ; Leptom 

{lep) ist stets von eioschichtigen verholzten Elementen (h) umgeben. 

Veigr. 360. 
Fig. 51. Querschnitt durch das Scheideblatt von Bambusa sienostachya. B 

Bastbelege, avbepj) subepidermale Bastplatte^ P Parenchym, mea 

Mestom. Vergr. 83. 
Fig. 52. Desgleichen von Phyllostachya mitis. If Luftr&ume, B, stibep. 6, 

P, mes wie oben. Vergr. ca. 10. 
Fig. 53. Queranastomose des ScheideblattbGndels von Arundinaria Hindsii. 

Vergr. 360. 
Fig. 54. Ligninauswuchse an Zellwanden. („Zwickel"). Vergr. 360. 
Fig. 55. Ein kleines Biindel im Laubblatt von Arundinaria Hindsii. ps 

Parenchymscheide, bs Bastscheide, svbepj), Had, Lep wie oben. 

Vergr. 360. 
Fig. 56. Langsschnitt durch einen kleinen Nerv des Laubblattes von Bam- 
busa palmata. ep, subep, b, ps, bs wie oben. Bastscheideelemente 

sind reichlicher betiipfelt als subepidermale Bastelemente. Vergr. 360. 
Fig. 57a. Starkekomer aus Rhizom von Phyllostachys Kumasasa, Vergr 

830. 
Fig. 576. Polyadelphische Starkekomer aus dem Halm von Bambusa 

paimala. Vergr. 830. 
Fig. 58. Einige Tyrosinkrystalle, die ausserhalb des nach Borodin'scher 

Methode behandelten Schnittes entstanden sind. Vergr. 360. 
Fig. 59. Beim Schneiden sofort in jungen Bastzellen auskiystallisierende 

Tyrosinkrystalle. B junge Bastzellen (mit plasmatischem Wand- 

bel^), P Parenchymzellen, tyr Tyrosinkrystalle. Vergr. 360. 
Fig. 60. Desgleichen in Langsschnittansicht. Vei^r. 200. 
Fig. 61. Beim Einlegen vom Schnitt in Glycerin in das Zelllumen ausges- 

chiedene Tyrosinkrystalle. tyr Tyrosinkrystalle, st Starkekomer, P. 

Parenchymzellen. Vergr. 360. 



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Jour. So. Coll. Vol. XIII. PI. XXII. 




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Decomposition of Hydroxyamidosulphates by 
Copper Sulphate. 



By 



Edward Divers, M. D., D. Sc., F. K. S., Emeritus Prof., 

and 

Tamemasa Haga, D. Sc., F. C. S., 
Professor, Tokj^5 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 Joum, 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.j 225), and when 
heated with cupric chloride it reduces the latter to cuprous 



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498 DIYEBS AND HAOA : DECX)MPOSinON OP 

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 mass 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 



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HYDR0XYAMID08ULPHATES BY COPPER SULPHATE. 499 

ammooia (in place of amidosulphuric acid) being the principal, 
if not the sole, products. 

There is a very marked diflference 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 
has 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 sail 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 heatiupr 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 (amidosulphate), 
and half into oxidised products together equivalent to the non- 
existent dihydroxyamidosulphate : 



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500 DIVEBS AND HAGA : DECOMPOSITION OF 

(1). 2Cu(H2NS04)2=N20+H20+H280,H-CuS04+ 
Cu(H2NS03)2=Cu(H2NS05)2+Cu(H2NS03)2. 
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 (I) that of Cu(H2NSO4)2=N20+ 
2H2O+2SO2+CUO, we get (2), 3Cu(H2NSO,)2=2N2O+4H20+ 
2S02+2CuS04+Cu(H2NS03)2, with producls 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) 3Cu(H2NS04)2=2N2+ 2H2O+ 2H28O4+ 2CU8O4+ Cu(H2NS03)2; 

(4) Cu(H2NS04)2=N2 + 2H2O + SO2+ CUSO4. 

In (3) 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 as a 
product of the decomposition, the sulphur appearing as sulphate 



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HYDROXYAMIDOSULPHATES BY CX)PPEB 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 complex 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. 



jResiUls and Method of the Qiumtitative 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 8.7 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 decomposes accord- 
ing to equation (1) : 



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502 PITEBS AND HAGA : DECOMPOSITION OP 

Sulphur as dioxide; as trioxide and amidosulphate. 

Found 3.5 96.2 

Calc 3J 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 5.5 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 

„ 13.0 43.3 43.2 

„ 13.1 86.6 

„ 13.3 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 

Calc 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 :. 



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HYDBOXYAMIDOBULPHATES BY COPPBB STTLPHATE. 603 

Sulphur as dioxide ; as trioxide and amidosniphate ; as acidity. 

Found 16.3 83.3 9.3 

Calc 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- 
lysisy 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 

, 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 diflferences between the calculated quantities and those 
found must be largely 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 

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604 DIVEBS AND HAOA : DEOOMPOSITION OP 

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 oflf the barium 
sulphate. 

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., ii, 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 



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HYDBOXYAMIDOSULPHATES BY COPPER SULPHATE. 505 

connected with a tube receiver holding bromine water kept 
cold, was heated, sometimes quickly, sometimes slowly, either by 
a spirit lamp 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. lu 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 ^ .^ted with hydrochloric acid for three hours at 
150°. Thf arium sulphate was again washed on the filtery 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 160°, the precipitated amidosulphate 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 

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606 DIVEBS & HAGA ! DECOMPOSITION OP OXYAMIBOSULPHATEfi. 

of amidosulphate sulphur, the full amount of which was ascer- 
tained by adding to it twice the quantity of that in the ammo- 
Bium sulphate extracted by hydrolysis from the crude barium 
precipitate. The sulphur from the hydroxyamidosulphate, ob- 
tained as sulphate, was found by subtracting from the total tbe 
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- 
amidosulphate. 

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 
^nd nitrogen was to expel air from the apparatus by a current 
of carbon dioxide continued for some lime, 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. 



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CONTENTS OP RECENT PARTS. 



Tol. X., Pt. 1 . . . yen 1.80 (Price in Tokyo). 

On tlie Fate of the Bln«t»p«re, tta« Relatlonii of the Primitive Streak, and the 
Formation of the Posterior En4 of the Emhrjo In Chelonia, toc^ther with 
Bomarha on the Natnre of Bleroblaatle Ova In Vertebrates. (Contributions 
to the Embryology of Reptilia. V.). By K. Mitsukuri. {WWi PUttet I-Xf). 

Vol. X., Pt. 2 . . . i/en 1.20 (Price in Tokyo). 
t/eber elne In Mlsakl irorkoniniende Art iron Ephelota iind fiber Ihre Sporen- 

blldnnic. Von C ISHIKAWA. (Hier&i Tafdn XII und Xllf), 
Ceber das mnssenhafte Torkommen %'on Elsenbaeterlen In den Thermen Ton 

Ikao. Von M. MiTOSHi. 
Stndlen Hber die Schwerelrasenblidnnir nnd die Aehwerelbaeterien der Thermen 

▼on Tamoto bel Nlkk6. Von M. MiYOSHi. {ERerm Tafd XIV), 
Die Entwickelunc der CU^noplioren bel Plijsaiia maxima. Von S. GrOTu. (Hienu^ 

Taf. XVy 
Stndles of Reprodnetlve Elements. III. Die Entwlckelnnir der Polienk4rner 

von Aillnm fistnlosnm I«.» ein Beltrac *■■* €hromosomenrednktl«>n la 

Pflansenreiehe. Von C. Ishikawa. ITiersa Tafefn XVI und XVII), 
t*ontrlbDtions to the Morphoiogpy of Cjrelostomata. I. On the Formation of I he 

Heart In Petromyson. By S. Hatta. ( WUh Piate XV I 11), 

Vol. X., Pt. 3 . . . yen 1.10 (Price in Tokyo). 
The Metamorphosis of Asterias pallida, with Special Reference to the Fate of 
the Bodj .cavities. By 8. GoTo. (With PlaUs XIX-XXIV), 



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Preparation of Hyponltrite from Nitrite throagrh Oxamldosnlphonate. By E. 

Divers and T. Haga. • 

Absorption of Kltrie Oxide In Gas Anal.Tsls. By E. Dn'ERS. 
Interactfon of Nltrle Oxide with Silver Nitrate. By E. Dhebs. 
Preparation of Pare Alkali Nitrites. By E. Divers. 

The Redaction of an Alkali Nitrite by an Alkali Metal. By E. DlVFRS. 
Hjrponltrltes : their Properties and their Preparation by Sodlnm or Potassium. 

By. E. Divers. 

Vol. XI., Pt. 2 . . . yen 0.38 (Price in Tokyo). 
On the Geoloffic Stmctare of the Halayan Avchlpela(ro. By B. Kot6. {With Plate L 

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Horlsontal Pendulums for the Bfechanleal Re§rlstratlon of Seismic and Other 

Earth Movements. By F. Omori. {With Plates IlXfl), 
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Plates xm-xviy 



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Enrthqnake Meamiremeot at Mlyako. By F. Omori and K. HiBATA. {With Ptaie) 

xvnxxniy 

Eibyl animonlunMiilplilte. By E. DivERS and Ogawa. 

Etliyl ammoninni selenlte and Non-existence of Aniifloselenlle» (.Selenosa mates). 

By E. Divers and T. Hada. 
Notes on the Minerals of Japan. By K. JiNBO. 

Vol. XI., Pt. 4 . . . yen 1.04 (Price in Tokyo). 

On the Wntiial Inlliienee between Ijonsritudlnal and CirciilHr aiagrnetlxatlon!! In 
Iron and Nickel. By K. HoNDA. {WUh PI^e» XX J V cfc XXV). 

The Earthquake Investigation Committee (Tatalogrue of Japanese EArthqnakes* 
By S. Sekiya. 

Notefl on the Earth qin.^ke Investiipatlon Committee C.italojrae of Japanese Earth- 
quakes.' By F. 0.v:oRi. {WUh Flales XXVLXXVIf). 



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JTapanlsche besclialte Fulmonaten. Anat. ITntersneh. d. in Zool. 91useiim dcr 
k. IJnIv. In T6kyd enUialtcnen Slateriales. Yon A. Jacobi. (Uienu Tafein 
I- VI), 

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der Berrnchtunsr bel Cyras revoiuta. By S. iKf^'O. ( With Plaieft X-XVIl). 

On a Collection of Batraehluns and Reptiles from Formosa and A<Uaeent Islands. 
By L. Sticjneger. 

Some Points in the Metaniorphtwis of Asterlna grlbbosa. By S. Goto. {WUh PhU 
TH). 

Vol. XII., Pt. 4 . . . yen 1.86 (Price in Tokyo). 

Further Observations on the Nnclear Division of Noctilnea. By C. IsHIKAWA. 

( With Ptates XIX), 
Notes on Home Exutic Species of Ectoparasitic Trematodes. By S. Goto. ( WitJk 

PicUe8 XX & XXI). 
Teutamen Flora; Ijulchuensls. Sectio Prima. Plantie Dlcotjrledoneje PoljrpetalaB 

By T. Ito and J. Matsumura. 



Vol. XIII., Pt. 1 . . . yen 1.66 (Price in Tokyo). 

Notes ou the Geology of the Dependent Isles of Taiwan. By B. Kotd. {WUh Plates 

I-V). 
Ckani^ of Tolamc and of I^enirtli In Iron, Steel, and Nickel Ovolfis bj Macnetlxation. 

By H. Nagaoka and K. Honda. {With Plates VI & VII). 
Combined Effect of I«ong:itDdlnal and Circular Mairn^tl>Atlons on tke Dlmenalons of 

Iron, Steel and Nickel Tubes. By K. HONDA. {With Plates VIII d IX). 



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-StDdlcn ttber die AnpamnnvMnihiirkeU elnl^r^r lafhMorlen an coiiceii(rlrt« I^tenngren. 

Von A. Yasuda. (ITiazu Tafd X-XII), 
Ueber die WaclmUiainsbeflelileunlKrniii: elnlffer Alffen mid Pilse dnreh ehemtache 
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Ammoiiiuni Amldosulptaite. By E. Divers and M. OoAWA. 

Prodncls of beatlog: Ammonlnm SalpliUes, Thlosnlphafe, and Trithionate. By 

E. DiVEBS and M. Ogawa. 
Potassium Nltrlto-iiydroxymidosulpiiateii and the STon^exUtenee of Diiiydroxy- 

lamlne DerlvallireM. By K. Djveks nnd T. IIaga. 
Ideotiflcatton and Conntltnllon of Fremy'^s !<nlpIiaxo(lBed Sniis of Potasslnm, lil» 

Salphaaate, Snlphaxlie, etc By E. DiVEKS and T. IIaga. 
On a Specimen of a Glgrantic Ifydrold, Branehlocerlanltaiis Imperator (Allman), 

fonnd In the Sa«anii Sea. By M. Miyajima. ( With Plates XIV & XV), 
Xntnal Relations between Torsion and 9Ia)ic>aetlaEation In Iron and Nickel Wlre.«. 

By H. Naoaoka. (With Plates XVI). 
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CONTENTS. 



Vol. xin., Pt m. 

PAGE, 

Contributions to the Morphology of Cyclostomata. II. — ^The 
Development of Pronephros and Segmental Duct in Pctro- 
myzon. By S. H.\tta, Prof. (With Plates XVII-XXI) ... 311. 

Beitrage zur Wachstumsgeschichte der Bambusge^w^adise. 

Vou K. Shibata, Itigakushi. {rait Tafehi XKILXXIV) ... 427. 
Decomposition of Hydroxyamidosulphates by Copper Std- 
phate. By Edward Divers, M. D., D. So., F. R. S., Emeritus 
Prof., and Tamemas/^ Haga, D. Sc., F. C. S., Prof. 497. 



riUNTP:!) AT THE ♦'TOKYO TbUKIJI TYPE FOVNDEY." 



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3R ^ 5(9 S :fc *¥ /?5 S* 

^ "^ $!^ «r ^ fs 9i 

THE 

JOURNAL 

OF THE 

COLLEGE OF SCIENCE, 

IMPERIAL UNIVERSITY OF TOKYO, 
JA.FAN. 

VOL. XIIL, PART IV. 



M M '^ m ± m f9 n 

PUBLISHED BY THE UNIVERSITY. 

TOKYO, JAPAN. 

1901. 

MEIJI XXXIV, 



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Publishing Committee. 
^ 

Prof. K. MitSUkuri, Ph. D,, Rigakuhahushi, Director of the College 

{ex officio). 
Prof. B. Kotdt Ph. /)., Rigakuhdkushi. 
Prof. T. Hagat RigakuhdkushL 
Pro£ S. WataS6f Ph. D., Rigahihakushi. 



AU commTmleations relating to this Journal shonid be addressed to the 
Director of the College of Science. . 



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1 



Observations on the Development, Structure and 
Metamorphosis of Actinotrocha. 

By 
Iwaji Ikeda, RigahishU 



With Plates XXr-XXX. 



Introductory. 

Since the discovery of Actinotrocha by Johannes Muller 
in 184G, this peculiar larval form and its mother animal, 
Phoi'onis, have been made the subject of investigations by many 
distinguished authors such as WagenePw ('47), Gegenbaur ('54), 
Krohn ('54), Schneider ('62), Metschnikoff ('72, '82), E. 
B. Wilson ('81), and Foettinger ('82). Among more recent 
writers Caldwell ('85), ]\rc'lNTOSH ('88), Benham ('88), 
RouLE ('90, '96), CoRt ('91), and E. Schultze ('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 Balano^ 
(/losms and the Chordatii in general. As, however, in spite of all 
these works there still existed many gaps and unsatisfactory points 



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e508 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 resuUs are far from exhaustive, I ho])e 
they will help to advance our knowledge of the subject. 

My study was begun in the summer of 18i)8 during a stay 
at the Misaki Marine Biological Staticm 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 sjK^cies of Phoronis, which has been named by Dr. Oka ('97) 
P. IjbnaL^ 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 animnls 
c^an be easily collected. During the greater part of the year, eggs 
and young embryos, clustered together, in what may conveniently 
be called embryonal 7na8seSy are found adhering to the lophophoral 
crown of the adult, cme on each side of the median line. These 
furnished materials for the study of fertilization, segmentation and 
the early larval stjiges. The larvjc in the Actinotrocha stage are 
found swimming in the inlet and are caught with the surface net. 
As will later be fully described, there occur four kinds of the 
larvie, which no doubt represent as many species, including the 
common Ph(yroni8 ijimai. 

The .specimens, both adult and larval, were killed with the 
saturated solution of corrosive sublimate in l^o 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 
lurmatoxylin gave the most satisfoctory results. 



*For .1 discussion of the stains of this species, see Sufyplementwij NnfeJi, 

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ON DEVELOPMENT ETC. OF PHORONIS. 509 

Before proceeding further, I beg to tender my sincere thanks 
to Professors jMitsukuri and Ijima for their kind supervision 
of my work and for their painstaking revision of my manuscripts. 

Contents. 

I. The Early Develoimient of the Phuronis Larvu. 

a. Notes on Fertilization. 

b. Notes on Segmentation. 

c. Gastrulation and Mesoblast-Formation. 

iL Further Observations on the Development of the Lirva, up to 
the Period when it becomes free-swimming. 
IL 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 Ectoblastic Origin. 

3. Organs of Entoblastic Origin. 

4. Organs of Mesoblastic Origin. 

III. Metamorphosis. 

IV. Sui)p]ementary Notes. 



I. The Early Development of the Fhoronis Larva. 

a. Notes on Fertilization. 

Fhoronis i«, as is well known, a liermaplirodite, in which both 
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, so that our knowledge of its sexual organs and, 
consequently, of its fertilization has remained very imperfect, as 
was pohited out by Cori ('91). The only existing statement as 
to liow and where fertilization is accomplished in Fhoronis is that 
of KowALEWSKY ('67). This author thought that fertilization 



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510 I. IKEDA : 

took place in the body-cavities, and accordingly, as Cori remarks, 
l^e must have believed that self-fertilization prevails in Phoro- 
nis. Cori considers this as highly improbal)le, but does not bring 
forward any positive facts in contradiction of it, his inference 
being drawn solely from facts ol)served in other marine Metazoa. 

Since Kowalewsky's valuable researches ('67), it lias 
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 Fhoronis 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 fool 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 observeil 
by Kowalewsky and Benham. 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 Boveri's terminology. In the fresh 
state, they are spherical or somewhat elliptical in shape and per- 
fectly opaque by virtue of the abundant yolk-granules cout^iined in 
the vitellus. It is characteristic of these ova that the nucleus, which 



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ON DEVELOPMENT ETC, OF PHORONIS. 511 

is situated, not in the centre, but near the periphery, is al- 
ways in the meta- or ana-phase of Karyokinesis (fijz;. 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 prei)aration 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 fiict 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 Pharonis, 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 yomiger stages as we approach the pores, until we 
reach such eggs as have just been fertilized or i)erhaps even such 
as have not yet been fertilized at all. But even the youngest eggs 
foimd in the mass present an appearance very different from 
those found in the bodv-cavities, the former being invariablv 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 
liave already been formed ; these arc situated close together just 
inside the vitelline membrane. 

On the other hand, if we examine by means of serial sections 
through the ])osterior region of an adult, where the stomach and 

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512 I. IKEDA : 

the sexual oii::ans lie jiirouped toj^ether, a numl>er of Vav'^e ejjus 
ari' frequently found, Hoatinii; freely iu the c(x4oinic fluid of the 
hody-eavities. These ej::i;s do not ditfer in any respeet from tho^^^ 
in the nephridial rei^ion as rei::ards the size, the apjK^aranee of 
the karyokinetic lij^ure, or the number of ehromosomes. 

The facts ahove stated plainly |K>int to the followint:: con- 
clusion : — The oor/onia fall into the body-cavitlcs by a dehiscence of 
the orarUin tvalls and here develop utitll 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 
})rimary oocytes arc destined sooner or later to be carried by waij 
of the nephridia to the exterior^ tvhere they become fertilized by 
spermatozoa from other individuals. 

Reserving an account of the si)ermatogenesis and ovogenesis 
for a future occasion, I may here refer to a few facts observeil 
by nie relative to the proceas of fertilization. AVheii the two 
sexual elements are artificially brought together, numberless sjK^r- 
matozoa soon attiich themselves to the surface of the ovum. About 
10 minutes afterwards, the first polar globule makes its a])|x»ar- 
ance, followed soon afterwards by the second. Meanwhile a snudl 
clear spot, probably marking the place where the male element has 
entered, apj^ears on the surface of the egg ; it is however observ- 
able for only a very short time. Both figs, li) and 20 are 
sections of ova tiiken from the embryonal mass. The ovum given 
in fig. 19 is fully mature and ready to be fertilized; close to the 
l)olar globules rests the large female pronucleus. The ovum 
represented in fig. 20 belongs to a stage of fertilization in which 
the two pronuclei stand closely side by side. The larger female 
pronucleus has a nuclear membrane irregular in contour. The 



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ON DEVELOPMENT ETC. OF PHORONIS. 51Z 

intoiisely stained clironiatin pieces are in both nuclei disj)erse(l 
without any apparent order throuiihout the finely <»:raindar nuclear 
substance. At one s])ot outside the male ])ronucleus, there is 
visible a small and clear archophismic (?) spnce surrounded by 
a s(4 of exceedingly fine radial rays. The two polar j»:lobules of 
this egj;' were distinctly visible in other sections which have 
not been fiuured. 



/;. Notes on Segmentation. 

Our knowledge of the mode of segmentation in Phoronh is 
far from being satisfactory. IMETsriiNiKOFF ('82) gives no account 
of the process. Foettinger ('82), if one may judge from his 
figures, seems to have seen the e\^^^ undergoing to taland unequal 
segmentation. According to Caldwell ('82), the segmentation 
''proceeds with consider a f)le regvlarily " {Lc, p. 374) ; "Roule ('90) 

sjiys '' r ovule fecondc i^uhif line .segmentation totahforl regidiere " 

{I.e., p. 1147). E. ScHULTZE ('97) simply says '* Ich mh das Ei 
sirh total und unihjual furrhen^ {I.e., p. G). 

My observations of tlu^ process were made on eggs found in 
the embryonal mass as well as on those artificially fertilized. As 
the former showed comparatively rarely the e^irlier 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-like spot the first cleavage line 
makes its appearance, passing on one side of the polar globules 
(figs. 1 and 21). At this stage I can not perceive any differoncc 
in size and structure between the two blastomeres. The second 



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/)14 I. IKED A : 

(ileavage piano j^assos at riglit angles to the fii'st (fig. 2 b). It 
is a reniiirkable |x^culiarity of Phoronis eggs that two sister blas- 
tonieres derived by the division of a mother blastomere, never 
undergo the next division simultaneously, so that between any two 
eonsecutive 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 o, 7, y 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 obsen'able 
as early as in the four- cell stage. He says: ^^ At the stage of 4 
i^egmentation-spheres a division into two smaller clear and 
two larger opaque celh indicates the future ectoderm and endo- 
derm " (/.r., ]>. 374). At the corresponding stage of Phoronis 
ijimai I have not been able to discover any appreciable differen(*e 
in the size of its cells (see fig. 2 b). 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 |X3rceptible at this stage. Fig. 3 shows a side view of an egg 
with 8 blastomeres ; it will be seen that the upjxir four blastom- 
eres are very slightly smaller than the lower four. I couW 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, Ix?- 
comes more and more pronounced as segmentation advances, tends 



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ON DEVELOPMENT ETC. OP PHORONIS. /)15 

to gradually ohseuro the ordorly arrangomont of the colls. At the 
16-cell stage the regular arrangement is still, though loss (listiuctly, 
maintained, while at the 32-cell stage it is quite disturhed (fig. 4). 
From this period on, the polar globules can no longer be detected. 
In the earlier stages of segmentation, the blastomores 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 blastocfole and its opening 
to the exterior (fig. 4, bLc) become recognizable. The embryo 
at the morula stage is somewhat oblong in shape and has 
a quite spacious blastocoele, and the blastocoolic pore {bl.c.) 
is distinct on the ventral side (fig. 5). However, this pore dis- 
appears at an advanced morula st^ige, 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 eml)ryos), the pore [bLc.) is cut through 
and appears as a slit-like passage between two of the bounding 
blastomeres. 

In the blastula (fig. 2/)) the wall consists of cylindrical colls 
and encloses a tolerably wide blastoca»lic cavity, which is now at its 
greatest development. In this stage, the bilateral symmetry of 
the future larva is already est^iblished. 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 



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510 I. ikeda: 

tlie convex dorsal side. The nuclei in all the blastodermal cells 
are always situated in a peripheral position. 

Plasmic corpuscles. — A noteworthy fact ^vith 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 
blast(X*oelic cavity (fig. 25, pLco.) These have been first described 
by FoETTiNGER Under the name "corpuscules mesodermiques." 
According to this author, these corpuscles are free nuclei imbedded 
in a common protoplasmic mass which is supposed to fill up tlio 
l)lastoc(elc, (»ach 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 technicpie. His method consisted in pouring dilute acetic 
acid over the living embryo, and this, lus the author himself was 
well aware, is highly detrimental, in that it frequently breaks up 
the blastomeres into fragments. The corpuscles described l)y him 
from so early a stage as that with only 8 blastomeres must have 
been simply produced by fragmentiition, the result of his drastic 
treatment. The common protoplasmic mass supposed to be present 
in the blastocoele, was probably nothing but a coagulum. 

Again the mesodernizellen which Metschnikoff ('82) found 
in the blastocoelic cavity of the blastula are certainly not true 
mesoblast cells but rather certain spheres similar to Foettin- 
gee's '' corpuscules m6=<odermiques," as was rightly pointed out by 
Caldwell. Recently E. Schultze ('97) published a short piper 
entitled " Ueber die Mcsodennbildunff bei PhoroniSy" in which he 
writes as follows : — " Schon auf dera Stadium der rundlichen 
Blastula sehen wvr einige ilesodermzellen im Blastoccel aufsitzlen " 



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ON DEVELOPMENT ETC. OF PHORONIS. 517 

{Lc.y p. 6). It seems to me that Schulze has fallen into the 
siime mistake as Metschnikoff. 

Lastly, Caldwell ('82) has enteitoined a view (^uite difierent 
from those of other wTiters. According to him, the bodies in 
(juestion are not present as such in the blastocoele, l>ut are in 
reality only the cut ends of blastoderm cells projecting hito the 
cavity and as such of course have nothing to do with the true 
mesoblast. 

In the Fhoronis studied by me, the plasmic corpuscles are 
present only in the highly advanced blastula (fig. 25, pLco.). 
They are usually round in shai)e 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 sfeiins, 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 blastocoele. The nucleus of these cells 
conmionly lies in the periphery as in all other cells of the more 
ordinary shape. In my opinion, the i)roximal ends of the elong- 
ate cells break off from the main cell-body and fall into the 
blastocade, 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 l)odies and not mere ends of blastoderm cells 
cut off in the process of microtomizing as Wcis supposed by 



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518 I. IKEDA : 

Caldwell. Their sruall size and the tottil 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- 
ccelic cavity at the beginning of giistrulation, 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 blastocoele 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, l.c.y 
p. 18). 

c. Gastrulation and Mesobl AST-Formation. 

In this section, I shall first describe 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 Ijlastula becomes more clearly marked than before 
when the gastral invagination begins on the ventral or the flat- 
tened side. The initial depression occurs over the whole ventral 
wall, so that a saucer-shaj)ed embryo is produced. At first it is 
so shallow as to be perceived with difficulty in the surfece view. 
kSoou it deepens, becoming deepest at a point somewluit nearer to 
the Ijroader end than to the narrow end of the embryo. The 



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ox DEVELOPMENT ETC, OF PHORONIS. 519 

deepest i>ortion may conveniently be called the central depression. 
Fig. 6 represents the ventral view of an embryo in which the 
invagination has become visible fi'om the outside, the central 
depression being most deeply shaded in the figure. In a slightly 
more advanced stage, as the original wide depression grows deei>er, 
the external ojKining is gradually drawn together and at a certain 
stage (lig. 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 romided by curving uniformly 
outwards, while posteriorly the two other sides gradually approach 
ejich other so as to meet at a point w^hich may be called the ajx^x 
of the triangle. Leading backwards from this apex, there runs in 
the median line the so-cidled primitive groove. This latter and 
also the triangular shape of the blastpore are occasioned, in my opi- 
nion, simply by the blastoix)re, 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 esi)ecia]ly in the last mentioned 
}xirt«, so that there the pressure, which is exercised by the ecto- 
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 l)laHtpore 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 fiict that the embryo has 
developed considerably the body-length does not show any signi- 
ficant hicreasc, remaining all the while at al)out 0.12 mm. on an 



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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 
or 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 am be plainly traced through the wall in the surface view. 
If the larvae of such an early stage of development be taken out 
of the embryonal 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 ectoblastic 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 bhistodermic 
cells which constitute the ventral portion of the blastula wall. 
This i)ortion not only shows a shallow concavity, but also the cells 
composmg it become, as figs. 26 a and 26 b 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 
blastoca^le. These liberated cells have usually a round shape and 



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J 



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 cluh-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 karyokinetic figure which shows that the cells are dividing 
and increasing in number in the layer to be invaginated. The 
cells pushed out into the blastocoele are nothing else than mesoblast 
cells, so that it may be stated that the mesoblast-form.ation begins 
simultaneously with the gastrulation. 

At the beginning of gastrulation we can thus distinguish two 
pai'ts in the blastoderm wall, viz,, the mesentoblast and the ecto^ 
h1<tsL The former corresponds to the wdiole 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- 
ly arranged cells, while the ectoblast is of taller cylindrical cells 
regularly arranged in a single row (figs. 2() a and 2G i). The 
mesentoblast, as the name indicates, is destined to give rise to 
both the entoblastic and mesoblastic elements. The characteristic 
clis]"K)sition of its constituent cells indicates its being the source of 
raeBoblast proliferation. 

The mesoblast proliferation becomes more and more accen- 
tuated in activity as the gastral invagination gradually deepens 
(see^g. 27), but the mesoblast cells thus formed do not adhere as 
a lining epithelium to the ectoblast, until wliat are called the ant- 
erior diverticula have been formed on })oth sides of the blastopore. 
Tlie blastocoele, in which the mesoblast cells are at first loosely 



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''>22 T. IKEDA : 

scnttorod {ibout, is honceforth greatly reduced in extent and finally, 
as the development of the arehenteron progresses, is almost obli- 
terated, esj)eeially along the dorsad and lateral portions of the 
embryo where the eetoblast and the gut come into direct contact 
with each other (see figs. 29 and 30 i). 

Figs. 28 a-c show three cross sections through different parte 
of a larva of nearly the sjime stage as that represented in fig. 6, 
in which the invagination has become recognizable in the surface 
vicnv. Fig. 28 a passes through the central depression which bc- 
(HHues gradually shallower posteriorly (figs. 28 b and 28 c). 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 ectol)last. When the blasto- 
j>ore has t^iken a triangular shape (fig. 7) and the primary archen- 
teric cavity has somewhat l)ent 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 arehenteron. These points 
will become clear from a consideration of figs. 30 a-^, which 
are drawn from serial transverse sections of an embrj^o 
slightly older than that shown in fig. 7. In fig. 30 a, shoeing 
the right-hand side of the blastopore, we notice a lateral infolding 
{anl. <Uv.) of the archenteric wall a short distance inside the 
blastoix)re lip. Here the component cells are irregidarly arranged 
and their entire disposition reminds one of the mesentoblastic 
layer. Indeed some indubitable mesoblast cells are found pressed 



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ON DEVELOPMENT ETC. OP PIIORONIS. 523 

against the tip of tlie diverticulum. No doubt the mesoblast is 
hero 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 blastocoelic cavity on both sides of the blastopore. 
In fig. 30 by 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 blastoco^le (on the left-hand side in 
the figure). If the sections are followed further posteriorly, the 
groove still i)ersists, 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 entoblastic and ectoblastic 
layers are separated from each other by the comparatively wide 
blastocijelic cavity (fig. 30 c). 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 layer 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 api)ears 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 

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524 T. ikeda: 

is eccentrically placed nearer to the anterior end, lias to traverse 
a longer distance before it can be reflected inwards, and tluis 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 
dou])t 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 Cald- 
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 
certain developviental stage of the Phoronis e^nbryoSy are remnanls of 
the original mesentoblast which at an earlier stage occupied the 
the whole extent of the gaMraJ invagination. They are, therefore, 
merely temporary y and destined sooner or later to split into ineao- 
blastic and entoblastic cells. 

As will be seen in figs. 30 a-c, the ectoblast and the archen- 
teric 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 



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ON DEVELOPMENT ETC. OF PHORONIS. 525 

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 mesoblastic 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 seen in fig. 29, 
which represents a median sagittal section through an embryo of 
nearly the same stiige 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 api>ears at the posterior end of the larva, on the ventral 
side of the bhnd end of the now greatly elongated gut. It appears 
at first as a shallow depression (fig. 33, 7iep. 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 
sj^indle is always placed paratangentially to the wall. I have 
luoreover 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 larvae of the stage of fig. 9 the nephridial pit am 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 
2>rotrudes itself touching the ectoblast with its blind posterior 



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526 I. iKEDA : 

end. This hollow protuberance is the rudiment of the intejstiual 
Ciinal of Actinotrocha. In longitudinal s>ection it is shown in fig. 
37 (InL). 

In fig. 34, 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 nej)hridial sac, which is quite free from the gut. The 
eetoblastic 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 abeady 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 siime stage, the nephridial pit appears as a single flattened 
sac {nep. p.) lying in front of the intestine {hit.) ; the eetoblastic 
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 
waiters, I m\\ 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 ix)rtion of 
the gut have become considerably elongated, the nephridial pit? 
which has meanwhile become deeper than before, begins at its 
inner blind end to divide into two lateral branches. Each of the 
latter corresponds, as will be fully demonstrated fiirther on, to the 
nephridial canal of Actinotrocha. Fig. 38, a frontal section of a 
larva at this stage, shows the bifurcation just alluded to. The 



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ON DEVELOPMENT ETC. OF PHORONIS. 527 

relation of the unimired nephridial sac to the gut will be best 
understood from the median sagittal section given in lig. 37. 

I may here be allowed to put in a short historical review of 
the mesoblast-formation in the Phoronis larva. 

KowALEWSKY ('67) attributed the origin of the mesoblast to 
the ectoblast. 

MeTSCHNIKOFF ('82), FOETTINGER ('82), and E. SCHULTZE 

C97) 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 blastojK)re lips (lateral), but it arises 
hiter 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 each mass thus formed. 
These cavities are the two halves of the body-cavity {preoral) '^ 
{I.e. J p. 374). On the other hand with respect to the ]M>sterior 
body-cavity, he states that " it is formed independently in a 
paired mass of cells which grow out to the end of the first farmed 
sacs and remain separated from septum " {I.e., \). 37G). Thus he 
regards the preoral body-cavity as arising after the enterocoelic 
tjpe. Lastly the author puts forward in his recapitulation the 
ojnnion that the blastoi)orc gives rise to both the mouth and the 
amis. 



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528 I. iKEDA : 

llouLE (*90) also distinguished two sorts of mesoblast cells in 
view of tlieii* different origin and fate : *' Mesencliymes primaires " 
and ** initales mesoderm iques." Both are derivcM.1 from the " pro- 
t(X4id(Hlerme" which forms the primary archcnteric wall. The 
latter gives rise to cells gi'ouj)ed together into two com|)aet 
** bandlettes mesodcTiuiques/' which are regjirded as homologous with 
the mescxlermal bands of Annelid larvie. In reality these bauds 
are, as have been pointed out by Schultze, nothing else than the 
j)0sterior piired diverticula of Caldwell. 

F'rom 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. 20 a 
& b)j and long before the blastopore tjikes the small triangular 
shajxi. 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 enteroca^lic 
cavity. As already described, the cells in question, after being 
budded off, lie loose in the blastocoele together with preexisting 
mesoblast cells and without forming a wall to a special cavity of 
any sort. 

As to the ventral groove, Metschxikoff ('82) was the first 
to refer to this structiu-e and wrote as follows : " /n passender 
Lagc des Embryos hann man cine in Verbindung mil dem JBlasto- 
pones bejindlichcn Farche {longitudinale) wahrnehmerij icelcJte zum 
Hinlcrende des Embryos hinzieht und sich niir auf dem Ekloderm 
beshrllnkL Diese Farche erhohi den bilateralen Bauplan des 
Embryos erschelnt indessen als eine vergiingliche Bildung^ irelche 
man auf spltUren Stadium vergebens suchen wurde*^ {I.e., p. 301). 
According to Caldwell, this groove, which he adls ** the primi- 



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OK DEVELOPMENT ETC. OP PHORONIS. »529 

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 ciise, for the so-called primitive 
streak entirely disappears leaving no trace whatever, long before 
the nephridial or anal pit makes its appearance. Tlierefore there 
exists no direct genetic relation between the primitive streak and 
the anal pit. 

Caldwell's view that the tw^o nephridial pouches give off 
the mesoblast, which eventually lines the posterior bmly-cavity, 
am not be sustained ; for, according to my own observations, that 
body-cavity with its mesoldastic lining wall is already present before 
the nephridial pit divides into the two pouches. It is tme that 
the cells floating in the posterior body-(*avity are in some sections 
found aggregated at the blind ends of the pouches as shown in 
Fig. ^58. This is a condition which might mislead one to tlie 
conclusion that mesol)last cells are here in process of proliferation. 
But solid cell accumulation in such a section is to be considered 
as simply due to the ol)literation by compression of the lumen of 
the nephridial pouches. Fig. 3G t^iken from an obliquely cut 
sagittal section through a larva of this stage, show^s no w^ander- 
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 tlie ]x)sterior 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 



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O30 I. IKEDA : 

exterior. However, it seems elear to me that this eord is nothing 
else than an early st^ige of the intestinal ontgro^th independently 
produced at the posterior end of the gut. Moreover, in Phoronu 
ijimai, the gut cavity does not come into communication with the 
exterior at so e^irly 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. ScHULTZE ('97) rejects Caldwell's \'iews in regard to 
the anal pit, l)ut regards it as a rudiment of the future ventral 
])0uch of Actinotro(»ha. 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. 



d. Further Observations on the 
Devfxopment of the Larva. 

Some authors have recorded that the larva swims about 
abrofid at such a stage of development as is represeut^nl in fig. 8. 
However in Phoronis ijimaij 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 ectoblastic 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 gjmglion? the 
latter, the rudiment of the first pair of larval tentiicles. The 
nephridial invagination at the posterior end is still shallow. 

At a little later stage, the tentacular thickening divides into 



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OK DEVELOPMENT tlTC. OP PHOROXLS. 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 oesophageal canal is 
formed (fig. 37, aes). The oesophageal wall is, therefore, ecto- 
bhistic in origin and is composed of strongly ciliated cohmmar 
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 larvae, the pit is split throughout its entire length into 
two nearly parallel canals, each of which opens independently to 
the exterior. Figs. 39 a-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. c.) on either side of the stomach represent the uppermost 
portion of the nephridial canals. In the second figure, e^ich of the 
cell-masses encloses an easily distinguishable lumen. The two 
canals 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 ne])hridial canals {nep. c). 
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 oesophagus 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 



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532 T. IKEDA : 

with two pairs of as yot very short larval tontacles ; this is the 
most advanced developmental stage to be met with in the em- 
bryonal masses. Fig. 40 is a median sjigittal section of such a 
highly advanced larva. Here the a\sophagns {(f^s.) and the in- 
testine (inL), which latter now communicates wth the exterior by 
the small anus (an,), are highlj^ developed, so that the three pirts 
of the alimentary tract (oesophagus, stomach, and intestine), may 
be said to be almost complete. The nerve ganglion (fig. 40, ffl.) 
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 ascertiun whether 
a proctodoeum 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, 7tics\), The nephridial canals (fig. 41, 
nep. c.) are now distinctly separated and removed from e^ich other, 
and are found in a cross section to be situated laterally to the 
intestine {int,). 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 ectoblastio 
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 of a 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 



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ON DEVELOPMENT ETC. OF PHORONIS. 533 

the excretory cells of Actinotrocha, is always found attaclied to 
the blind ends of the nephridial canals. Caldwell siiys that he 
sjiw 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 Echiuvus. 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 cert^iinly not provided 
with any such processes. 

I have very frequently detected some gigantic mesoljlast cells 
floiiting freely in the postoral body-cavity of larvae with one or 
two pairs of tentacles (fig. 44, corp,). They are round and 
nucle^ited 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 corj)uscle-massess in the collar cavity 
of Actinotrocha. This point will again be treated of in detiiil in 
the i)roi)er place in the following section. 



n. The Structure of Actinotrocha. 

a. EXTEKNAL ApPEAKANCE. 

It can scarcely be doubted that eiich species of the Plioro- 
nidcB has a characteristic form of Actinotrocha i)eculiar to it. Some 
of the previous observers [e. g., Wilson and Masterman) have 
mentioned two distinct types of larvai as occurring in the sinne 
locality. Among the larvte which I ol)served at Misaki, I wns 
able to distinguish four dittcrent typs, each of which had a 
characteristic form and a more or less definite topographical 



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534 I. IKEDA : 

distribution. I Nvill dcHigiiate these tyjxis by the letters Ay B, C, 
and D. 

Type A (fig. 13). The larvie of this type were principally 
collected in Aburatsubo and belong in all probability to the sjK^'ies 
Phoronis ijwial, which, as I have said, is found in the same 
locality. The body is comparatively short and thick, measuring 
about l.-l.o mm. in total length. The larval tentacles of a full 
grown larva never exceed IG in number. 

Type B (fig. 14). This is a larger form than the preceding 
(about 2-2.6 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 {ho.) situated just in front of the ganglion 
{gL). The larvse were found in greatest abundance near Kitsune- 
saki, a point at the mouth of the inlet Moroiso. 

Type C. (figs. 15 a & 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 l.bmm. 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 {gld.) is found one on either side of the 
ganglion {gl). A jmir of retractor muscles {ret\) runs longi- 
tudinally through the trunk cavity from the tentiicular ring to the 
apex of the anal cone. Comimred with the first two types this is 
much rarer. 

Type D (figs. 12 and IG). 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 lon^ and thick. The tentacles are remarkably 



'o 



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ON DEVELOPMENT ETC. OF PHOBONIS. 535 

numerous, sometimes reaching 48 in number. In a single living 
specimen, the skin of the trunk was of a light orange coloiu* ; the 
subdermal circular muscles were especially well developed in the 
trunk but interrupted at four longitudinal clearly marked zon(*s. 

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 
7nm. 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 larvae 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 
^vhich extends between the ganglion {(/L) and the dorsal iinier side 
of the tentacular circle {rel. in figs. 12, 13, 14, and 15). 

The larval organisiition of types A and B is nearly com- 
pleted in the stages with 14-10 tentacles. I^t me next give a 
somewhat detiiiled description of the external apix^arance of Actino- 
trocha in general. 



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536 I. ikeda: 

The prcoral lobe. This is a structure which looks like a 
broad hood with its couaivity directed downwards. It almost 
entirely covers the up})er anterior part of the collar,* when not 
influenced l)y external circumsUinces. Mastermax has made the 
statement that in its natural attitude the hood has its length 
disposed parallel to the principd body-axis. However, if the 
larva I)e examined in the living sUite, it will at once l^e discovered 
that its normal disj)osition is horizontal. It becomes turned up 
only as the result of invservation. Its whole surfiice is covered 
with cilia, nuxst strongly developed along the free margin which 
constitutes the prcoral ciliated belt. In the full grown larva, the 
}:anglion (and also the sensory spot in type B) has also a set of 
s])ecially long cilia on the outside. Numerous fine and refractive 
nerve fibres are seen radiating from the ganglion {gl.) to the free 
margin of the lobe {pre. hel.) (figs. 13 and 14). 

Masterman has described and figured two ectoblastic struct- 
ures which are siiid to be situated on the ventral wall of the hood 
and which he has named the " oral " and the *' j)haryngc*al '' 
grooves. These he compares, as to their function, to the jicill- 
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 
sjK'cimens. Among the preserved sixicimens I have frequently 
noticed those in which tlie lower or oral wall of the hood wu> 
prominently bulged out in front of the mouth. In consequence 
of that prominence (fig. 1(5, p)wn.), there was produced on either 



*I adopt this name of Masternian's iv denote that portion of tiie lan'al body wliicli lies 
in front of liie tentacular circle and behind the preoral lobe. 



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ox DEVETX)PMENT ETC. OF PHORONTS. 537 

side of the mouth a transverse groove, which was visi1)le when 
viewed from the ventral side. I believe it was to grooves of this 
kind that Masterman assigned the above import^int 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 cylinder 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 l)y a narrow neck. The 
numl)er of tentacles (larval) varies according to the different stages 
of growth and also according to the type to which larv«3 belong. 
They are most numerous in type 2), most individuals of which 
bear 40-48 tentacles (figs. 12 and 1(5). The rudiments of the 
adult tentacles make their appearance as Inul-like ectol)lastic 
thickenings immediately l)elow the base of the larval tentacles. 
An exception to this rule is found in the case of larvte belonging 
to tyjx^ Dj 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. 58 d^ s. L). It is very probal)le that 
the number of the larval tentiicles 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 
re«;ions in early larval stiiges (fig. 10), comes with growth to 
oe<*upy the largest part of the larval body and assumes a long 
cylindrical form. Its anterior boundary is the tenbicular circle ; 
the posterior end is girdled with the perianal ciliated belt which 
serves as the larval locomotory organ. 



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'538 T. IKEDA : 

b. The Internal Structure of Actinotrocha. 

1. Body-Divisions and Body-Cavities. 

I have endeavored to show in the preeeding pagers, that the 
body-eavities of Actinotrocha do not arise from the enteric diver- 
ticula, as was insisted upon by Caldwell, but that they are 
simply produced by mesoblastic cells applpng themselves to and 
forming the lining of the ectoblastic, and the entoblastic, wall. 
They may, therefore, be classed under the ** pseudocoele " or 
** schizocoele " of Hertwtg. ^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 elucidate 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-ca\nty) 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, i. e. the 
trunk cavity, is the only portion that persists among the body- 
cavities of the adult, in which it is kno\Mi as the foot or infraseptal 
C4ivity. In correlation with this circumstance are observable cer- 
\i\\\\ 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 ca\nty 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 chan«:es to a certain extent at least establish the 



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ON DEVELOPMENT ETC. OF PHORONIS. 539 

feet that some profound changes in the arrangement of the body- 
eavities must occiu* 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, e. 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 Aclhiotrocha and 
those of the adult and to call them respectively by diflferent names. 
The former may be termed the larv^al body-cavities, and the latter, 
the adult body-cavities. 

Most previous WTiters have not taken any 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 
sefftnent was first introduced by Caldwell ; he considei's 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,y the preoral cavity in front of, and the trunk cavity behind, 
the septum. Masterman divides the entire body into three 
portions, m., 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 inodifications, his 



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540 I- IKEDA : 

view of the body-divLsions. 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 lie« 
between the preoral and the collar cavities is always an incom- 
plete formation, at least in all the Aclinoirochcd 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 (Mastebman). Therefore, I can not regard the body- 
divisions of Actinoirocha 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./.). A few blood corpuscles are also 
frequently discovered in this ca\aty (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 larv® 
investigated by Masterman. In sagittal sections of the larva at 
any stage of its growth, the septum can constantly be traced so 
long as the oesophagus is contained in the sections. In figs. 45 
and 63 a, a slender cellular strand {me8\) behind the gangUon (ffl.) 
represents the septum in cross section. Jt extends between the 



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O^ DEVELOPAtENT ETC. OP PHORONlS. o4l 

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 oesophagus, the upper portion of the septum becomes 
abruptly indistinct. In fig. 54, which shows a sagittal section 
through the right-hand side of the oesophagus of a larva of 16 
tentacles (type A), the septum {fnes'.) 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 
of it. 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 is a 
diagrammatic representation of the Aclmotrocha hood and its 
neighbouring part, as seen from above, i. e., in horizontal projection. 
Tlie 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 oesophagus 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, i)ervade 
the preoral body-cavity, except in a small si)ace immediately in 
front of, and below, the ganglion. This free space I shall call the 
posterior recess of the preoral cavity. The line abcdefghj 
curved somewhat like the letter M^ indicates the position of the 
preoral septum. The part shown in fiill 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 



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542 



1. IKEDA : 



tli*^* sjHici* ill front of the ^*ptiiin is tlir [»ivonil anity, wliilf back 
of it tit's till* nillar aivitv. 



iVnscr aarumtibtiftn »f fibres 
in front of i\w rece^«* 



Prcuial cavtlj 




ri«»terii*r retx^ of 
Xervc gasigliuii. 



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it^ f> |»t|^^^^^^MiK^ dvsT. i^^tiuD 50 a k tlie most 

^s through the hood at about Uie 

alH>rt^ wo(xlHruu The whole of tbe 

\h die fibres of branclunj raeaenchyiEiat- 

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ON DEVELOPMENT ETC. OF PHORONIS. 543 

tlie anterior end of the collar cavity (coLc). 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, tlie two wide spaces lying on both sides of the posterior 
recess correspond to the two anterior horns of the collar cavity 
projecting forwai'd (marked b 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 Mastebman, should entirely divide the preoral 
and the collar cavities at every point. As the figure shows, the 
dorsal portion of tlie mesentery {r)ie8^) is decomposed into fine 
protoplasmic processes which join with those of the fibrous mesen- 
chymes dispersed through the preoral cavity. In section, 59 c?, 
|xissing though the middle of the ganglion (the line III-III 
of the wood-cut), the collar Ciivities (coLc.) are much wider and 
have become united below the oesophagus {oss). The septum (as 
the wall of the posterior recess, pj\) 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 59 d passing through the line 
IV-IV of the wood-cut, the posterior wall of the recess (p.r.) is 
obliquely cut and appears m the right-hand lower corner as a 
membraneous slice, the recess being distinctly bounded by the 
septum {fne8\). Outside of it are seen, one on each side, the 
sections of the retractor muscles (re/.), of which more will be 
said later. The collar cavity (coLc.) 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 ix)rtion (indicated by the 
fvill line c d ef in the wood-cut), while in the more lateral part 
on each side, it is at the l^est a loose open reticular membrane, 



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544 t. ikeda: 

through which the coelomic fluid of the preoral and the collar 
cavities is put in free circulation. 

A questionable structuie 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 liim, the subneural sinus is an interstitial 
space left between the two laminse 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 oesophagus. 
After repeated examinations of the larvse of the four different types, 
I am convinced that Masterman'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 oesophagus, 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 Habmeb in his paper on Cephalodisus ('97). 

Masteeman has forther given an interesting description of the 
" proboscis pores/' situated on each side of the ganglion. They 
are compared to the proboscis pores of Bodanoglo^sus and are 
said to fiilfill the same function as the collar nephridium of 
Actinotrocha, In the larvue 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 



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ON DEVELOPMENT ETC. OP PHORONIS. 545 

belonging to typo 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 : " JuBt where the 
prearal mesoblastic wall slopes away on eUhei' side of the sinus 
there are a pair of thickenings^ which traced forwards^ show 
themselves to he the commencement of a pair of internal openings ^^ 
(Lc.9 p. 307). The paired thickenings referred to by him are 
apparently nothing else than the points, of attachment of the 
retractor muscles in the collar cavity, as will be seen in fig. 59 d 
(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- 
tuniy 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.)y found at the plane of the junction 
of the oesophagus with the stomach, while ventrally the attach- 
ment lies much further below. In frontal sections of the larva, 
tlie 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 canty, is already 
formed in the fully developed larva of every type, as a ring-space 
yunning along the inner side of the tentacular circle and above 



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546 I. iKEDA : 

the septum (see figs. 08 a 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 oesophagus, and separating dorsally the 
larval collar cavity into two lateral halves. In the Actinolroehae 
of all the types observed by me and at every stage of the larval 
growth, no such mesentery is present. It is true that the body 
walls and the oesophageal 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 50 a). 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 oesophagus lie well separated by a considerable space 
(figs. 58 a and 58 b). 

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 oesopha- 
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 -4, 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.)y 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 



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ON DEVETX)PMENT ETC. OP PITORONKS. 547 

trunk of a highly advanced larva of type C, is given in fig. Tu a^ 
in which the much elongated and convoluted pouch is seen cut 
into several sections (;>o.), connected with one another by the 
mesentery {v.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 Aclinotrocha studied by myself. 

2. Organs of Ectohlastic Origin. 

The epidermis of Actinolrocha is represented by a single 
layer 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, etc.) the constituent cells 
are cylindrical, the nucleus generally lying near the basal end. 
The body wall of the trunk region is very thin and is formed of 
greatly attenufited cells (espcially slender in the advanced larvfc). 



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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 
oesophageal 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 64 rf, m.gl). In their 
staining reactions, the secretory contents of the glands agree with 
those of mucin. It has been often noticed that lixang larvae 
remain adhereing to the objects they have touched with the hood, 
and that metamorphosed larvre behave similarly with the tip of 
the evaginated pouch. 

There exists still another, paired, multicellular gland which 
is observed only in the larvse of type (figs. 15, ffld. 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. 56 A, 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. 56 c). Since 

* By an unfortunate oversight, Fig. 56 b has had its number omitted in the plate. 

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ON DEVELOPMENT ETC. OP 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. 66 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. Iizuka tells 
me that similar glands of ectoblastic origin are constantly found 
on the superior ramus of a parapodium in certain Polychceta. 

.Ventral Pouch. As the ventral pouch is one of the most 

clianicteristic structures of Actinolrocha, 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 larvce of whatever type, 

the inner or ectoblaatic 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 has been 

observed by Metschnikoff and many other authorities. 

Nervous System. The nervous system of Actinotrocha, like 
tliat of PhoroniSy is of a very low development, being represented 



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550 t. tkEDAi 

merely by a local differentiation of the ectoblastic cells into ner- 
vous elements. The epidermis over both the ganglion (fig. 14. ffl) 
and the sensory spot {so.) is strongly ciliated, so that the organs 
are easily recognizable in the living larva. The ciirliest stage 
in which I found the ganglion was a 4-armed larva of type A 
(fig. 40^/.). In it, the ganglion consisted of only a few ganglion 
cells and nerve fibres. 

Although the ganglion and some nerves directly j^roceeding 
from it can be detected with tolerable distinctness in the living 
sjxicimen on account of their peculiar refi'activity, the peripheral 
nerves are as a general rule so very fine and delicate, that tliey 
can not be satisfactorily made out by means of any ordinary 
process. With feir success I have had recourse to vital staining 
with methyl-bhie. Larvse of type £ liave 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 larvae 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 tyj^e 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 Master- 
man. Whether this difference is due to the technique or is 
actually existant in the si)ecies studied, is difficult to ascertain. 

* 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 tlie 
true edge of the huod. 



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O^ DEVELOPMENt ETC. OF PIIORONIS. Ool 

Masterman siiys nothing of the method employed in his inves- 
tigation, and nnfortunately there exists no other study than his 
with whieh 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 eflforts, I have always failed to make 
out the presence of the so-called perianal nerv^e ring. The collar 
ring and the dorsal commisure, if they can be so named, are repre- 
sented by a small number of imrallel fibres, which spring directly 
out from the ix)sterior 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 beantifiil system of nerve fibres conld be seen 
on the preoral lobe. The fibres are here exceedingly numerous 
and 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 {ffl)y the fibres appear as three longi- 
tudinal parallel strands on which the unpaired sensory sjx^t (so.) 
is situated not fer 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 



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5o2 t IKEDA 1 

to me more probable that these appearances were caused simply 
by the juxtaposition of intersectmg fibres. 

The nerve endings in the preoral ciliated belt deserve special 
notice. In fig. 60 a, there is shown a row of small dots along 
the margin of the band. A portion of the latter more liiglily 
magnified is shown in fig. 60 b. 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 experimente, 
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 relatimis prove to a 
certainty that the larvae investigated w^ere fiiUy 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 Amphioxics and even to the medullary canal of 
Vertebrates. I nuist say I was much disiippointed in failing to 
detect in the Actinotrochxe studied by me this structure of so much 
theoretical interest. As a matter of fact, it happened very fre- 
quently, while observing living larvae, that the gangUon was 
i-etracted deeply inwards by an active contraction of the two 



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ON DEVELOPMENT ETC. OF PHORONTS. 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 reagent^i, so as 
to produce a shallow 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 artefiict. 

As to the tentacles, I have at present nothing to add to what 
is already known about them. 

3. Organs of Entoblmtic Origin. 

In the fully grown larvae 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-lG). 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 oesophagus of Actinotrocha is of ectoblastic origin, 
so that the original gastrula mouth is to be sought at the junc- 
ture of the oesophagus with the stomach. The oesophagus (Figs. 
45, 48, and 49, ces.) is a comparatively short and narrow canal 
¥rith 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. 

Mastibi^MAN has described an unpaired ectodermal inviigina- 



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554 T. IKEDA : 

tion situated in front of the mouth and just under the gangHoii. 
It is called the "subneural gland." Here again I am not hi a 
position to confirm his view. In spite of repeated examinations 
on li\ang 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 
as both the " oral-" and the " pharyngeal grooves " of this author 
are products of his fixing methoil. 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. 03 a). 

Stomach. The stomach forms the largest and widest portion 
of the alimentary canal. It is especially long in the larvae of type 
Dy 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 50 a, slam.). 
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 elongated nuclei, and 
are, in a word, of the [esophageal tvpe (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 



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ON DEVELOPMENT ETC. OF PHORONIS. 555 

unpaired diverticulum protrudes itself forwards (fig. 14, div.). The 
position of the organ is wholly ventral to the oesophagus {ces.), 
and the form is like that of a sac compressed in the dorso-ventral 
direction (figs. 45, 49, 50 a, and G3 a, dii\). 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 h?emoglobin of the blood corpuscles which, in 
advanced larvas, 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 larvae of every type, each cell constantly contains a single 
small round vacuole in its distal end (fig. 01 h). The vacuoles can 
not be stained by most of the staining reagents. I have seen 
them in the diverticular wall of a highly advanced larva belonging 
to type Ay which had already evaginated the ventral pouch ; even 
in this ciise, they were found only one in each cell (fig. 61 b). 
The whole of the diverticulum is lined externally with the thin 
peritoneal layer (see the above figure). 

Many previous observers have noticed this organ and have 
called it by various names : — 

J. MuLLER ('46)—" Blinddarme " (paired), 

Gegenbaur ('54) — '* Haufen der Leberzellen," 

Wagener ('47)—** Leberblinddarme," 

Claparede ('63) — " A dark mass with globules " (after 
Masterman), 

Metschnikoff ('71) — " brown specks," 

W1L8ON (A.G.) ('81)— *' glandular lobes of the s