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OBSERVATIONS
ON THE
[ : ‘ANATOMY OF THE CIRCULATORY SYSTEM

= AND THE i
COURSE OF BLOOD FLOW IN DIPLOCARDIA COMNUNIS GARMAN.

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Aete. BARRETT

THESIS PRESENTED OR THE DEGREE OF ©

BACHELOR OF ARTS
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ZOOLOGY

COLLEGE OF SCIENCE

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| This paper is based on the results of some investigation work
carried on in the laboratories of the Zoological Department of the
“University of Illinois as a part of the work preliminary to gradua-
Bion in a specialized course in the college of Science. The subject
‘was chosen at the suggestion of Professor F. Smith who has had im-
“mediate supervision of the work and to whom I have been under obli-
_gations for suggestions as to methods of procedure and for help in
| the matter of literature on the subject.

-~---000----
| INTRODUCTION.
| Diplocardia communis Garman ('88), is a species of large
fliesh-colored earth worm common in the black soil of the prairies
of femoris. The body is made up of from 120 to 170 somites, and

often reaches a length of a foot. It is cylindrical for the great-

,

er mart Of its length and has its greatest diameter in somite VII.
|

The specimens used in this investigation were collected in abune-

H dance on the campus of the University of Illinois. During early

A (morning showers they came to the surface where they remained some
| aatt20 time after the showers hed ceased.

In taking up this investigation the aim was to present the an-
atomy of the circulatory system and the course of blood-flow in
this species, since it belongs to a genus concerning whose circula-

tory System very little is known... Whereas in Lumbricus terrestris

| and in many other species there are seven longitudinal trunks, in

D. communis there are but four, the subneural and the lateral neur-

als being lacking in the latter.

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A comparison of some of the main trunks of D. Communis with

| quite carefully worked out will best prepare the way for a more de-
tailed description. .

"a As the name indicates, Diplocardia communis has a double dore
sal vessel, a condition not entirely lacking in some other worms.

| As Beddard a hive most diverse families exhibit this pe-
Briterity. In Megascolex pearainan (92) the vessel is double only

"eign the anterior part of the body as it also is in Diplocardia eis-
— Michaelsen

) eni,('94); in Octochaetus multiporus and in Acanthodrilus annectens

the vessel is completely double there being two distinctly separate

| tubes running side by side on the dorsal surface of the alimentary

= the two tubes fuse. From Michaelsen's definition of Diploe¢
y fede

_ cardia communis (typica) in Das Tierreich ('00) it is evident that

tract. in D. communis, as in Acanthodrilus novae-zelandiae, there

are two such tubes in the middle region of each somites but at the

| he has the wrong idea concerning this double condition of the dors

|

| vessel. He says that it is double , for instance, in somites

y gle
Vill, X, @nd XII, and single in somites VIL, IX, and Xl. In D.

ie

somite X and the anterior intestino-tegumentary vessels in Megasco-
| lex coeruleus end in a network of capillaries in somite XIII, the

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| daterals of D. communis extend from their origin at the anterior

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oe@sophagus,* leave the same in XIII and pass to the body wall along
which they extend as continuous trunks to XVIII giving off branches

in each of the somites XIII to XVIII inclusive. They have direct

connection with the dorsal by means of the dorso-tegumentaries.

|
| In D. communis as in many Oligochaéta there is in addition to

the dorsal a vessel temned supra-intestinal which extends along the
Rca surface of the oesophageal region where it takes the part in
reference to the intestinal est ea iaeleatenat is elsewhere played by
| the dorsal vessel. This vessel,double in Megascolex and entirely

'absent in Lumbricus,is single in Diplocardia. There is no relation
between the supra-intestinal and the typhlosolar network in D. com-
} munis as has been assumed by some writers to exist in certain other

| forms.

METHODS OF OBSERVATION.

The method of working out the anatomy from sections was depart-

‘ed from to a great extent in working out the anatomy of the circue

| laroty system of D. communis and instead dissected living specimens

_wére used. <A number of points were made out from sections, however,

and as means of checking up those workéd out on the living speci#u

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*The term oesophagus as used by Garman in his description of
Diplocardia communis was applied only to the very narrow region be-
tween the pharynx and gizzard, while the narrow part of the alimen-
tary tract following the gizzard and anterior to somite XVII he |
designated as the first aie of the intestine. As has Bourne
in his description of Megascolex coreuleus, I have used the term
oesophagus to apply to any portion of the alimentary canal anterior

to the large intestine which is not designated by any special name.

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mens they proved quite useful.

The worms used for making the sections were placed in a shal-
low dish of water under a small bell jar in the presence of a small
piece of cloth or cotton saturated with chlorform and allowed to

remain there until completely anesthetized. They were killed and

fixed in picro-sulphuric acid. The sections were cut 10 to 15 mic-

rons thick and stained in Flémming's triple stain. The safranin

gave a beautiful red blood stain and enabled one to follow the cap-

_dllaries into the integument with ease.

The best results were obtained as stated above, by a study of
dissected living specimens. It required some little time to become
proficient and accurate in the dissections but with proper care
very few specimens need be lost in the preliminary work.

the worms were anesthetized by the method stated but were not
allowed to become entirely motionless before dissecting. An anés-
thetized Beene: was placed in a shallow petri dish containing a
4/5% salt solution and a strip of cork weishted down by lead. Af-
ter pinning the patevter end of. the worm to the cork it was opened

either along the ventral or the dorsal median line care being taken

' not to sever any of the blood vessels. After pinning down the edg-

6s Of the worm it was ready for study. If none of the larger vése-
sels were broken the specimen remained alive and in good condition

for more than an hour.
I was quite fortunate in having access to a Zeiss binocular

dissecting microscope belonging to the State Laboratory of Natural

History and made frequent use of it in this investigation. Artifi-

cial light from a 32 candle power incandescent light passed through

a bulls eye condensor and thrown on the specimen enabled one to see

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all vessels with little difficulty.
Very small specimens were examined between slides under a 2/3-
inch objective but aside from the valves and a few points concern-

ing the course of flow nothing of importance was made out from then.

ANATOLY.

| Dorsal Vessel.(Figs. 1, 2, & 3, D. V.).-- The dorsal vessel is the

chief pulsatile organ of the body and extends from the anterior
part of the pharynx to the last somite of the body. It lies ciose
to the dorsal wall of the alimentary tract but is not adherent to
it in any part of its course. It bifurcates in the fourth somite
and each of the two branches into which it divides breaks up into
smaller branches on the pharynx and body wall. It is a single tube
anterior to septum vVI/VII, but posterior to this septum there are
two such tubes in the middle region of each somite, while at the
septa the two tubes fuse as previously stated.

As a general rule in Oligochaeta the dorsal vessel is of nearly
the same diameter througcnout its length but’ in D. communis it is
quite small anterior to somite VI increasing in size to somite XIV
where it reaches its maximum diameter. From somite XIV to XIX it
Meo. nearly a uniform diameter but posterior to somite XIX it is‘a

Tittle smaller and continues with a gradual decrease in size to its

termination at the posterior end of the body.

As stated in the introduction some worms are known which have
the dorsal represented by two distinctly separate tubes while oth-

ers are double only at the anterior end as Megascolex coeruleus and

Diplocardia @iseni. Although the double condition as described

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abovo predominates in D. communis several specimens were found in
which the trunks of the Perre pe separated for as many as seven
somites but this condition was never found anterior to XVII. Among
nearly a hundred’specimens dissected not one- was found in wich the
dorsal vessel was a single tube in any region other than that men-
tioned above.. In Microchaéta-rappi('86) the dorsal vessel is double
but the two halves are closely bound together, and only recognizable
as distinct by the presence of two quite separated blood-clots lying
side by Eke in the apparently single tube. In somite VII the dor-
sal vessel, apparently simple, forms a very wide chamber which is |
divided by a longitudinal septum reaching nearly to the posterior
extremity. One might expect fe find in D. communis, in some cases.
at least, that the single tube of the dorsal at the septa was real-
ly double as in the case mentiondabove, but from sections studied
and all small worms in which such facts could be determined no ine
@ueati On Of the double condition of the dorsal vessel at this point
was found. Posteriorly the dorsal vessel ends abruptly ne shown in
Fig. 6. )

eeterior to somite XIII the dorsal vessel is thickly coated
With chlorogogve cells. <A few may be found in somite XIII but an=
eerior to it they are lacking.
Ventral Vessel. (Figs. 1,2,3 & 4, V.V.).-- This is also known as the
subintestinal or supraneural vessel. It is a non-contractile sin-
gle tube supported from the ventral median line of the alimentary

tract by, a thin mesentery and extends the entire length of the body.

Although it is held somewhat firmly by the septa, in the cavities

of the somites it lies free and may be seen to fold upon itself.

It bifurcates on the pharynx in somite III each branch ending in

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theianterior network. Posteriorly it ends abruptly (Tig. 6).
Laterals.(Figs. 1,2,3 & 4, IV).-- as stated above, the lateral
vessels originate anteriorly in a capillary network on the pharynx
and body wall. mhey are larger than the ventral and lie close to
the latero-ventral surface of the alimentary canal. Each bifur -
cates in somite IV, the anterior end of each branch being connect-
ed with a network of capillaries on the pharyngeal wall. mhis net-
work is connected with. the network into which the anterior branches
of the dorsal and ventral vessels break up (Fig.8). In somite V
the lateral vessels leave the lateral surface of the. gizzard on
either side and assume a more ventral position (Fig.8). The later-
als are of a wuniform caliber from somite VI to IX; from somite IX
to XIII a gradual decrease takes place. In somite XIII they leave
the oesophagus and pass Sslightiy ventrad to the body wall along i
which they extend to XVIII (Figs. A uaone

Supra-intestinal vessel. ( Migs. 1.83, S.1.).-- As stated above, the
supra-intestinal vessel in D. communis is a single tube which ex-
‘tends from somite ix to ZIV inclusive. It reaches its maximum die
ameter in somites X to XII where it equals that of a single portion
of the dorsal in that region. Anterior and posterior to this region
if decreases rather abruptly disappearing in the tissue of the oe-
sophagus. The Je eee vessel is connected with the oe-
sOphagus by several short branches in each somite which are called
by Bourne supra-intestino-intestinal rae fhe supra-intestinal
Bisochas connection with hearts in somites X, XI and XII.
Dorso-tegumentary, Vessels. (Figs. 4&5,D.1T).-- These vessels are

the branches of the dorsal which place it in communication with the

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peripheral nofworks. They are slender, contorted vessels and are
given off from the dorsal vessel just anterior to the septa in all
somites eatari or to XIII. They are entirely free from the intes-
tine but are slightly attached to the septa by a thin mesentery.

| The pairs in genites XIV and XVIII are Woe arco and as Garman

| states, the former without branches reaches the body wall between
tee outer and inner pairs of setae. But contrary to his statement
enet they enter the intesument, they join the clitellar division of
the iaterals directly, as do all the dorso-tegumentaries of the
somites anterior to somite XIX.

The dorso-tegumentaries pass outward and slightly upward from
the dorsal. Theythen follow along the body wall to which they give
three or four branches (vTig.4,b). Posterior to somite XX each dor-
so-tegumentary gives off from one to three branches which enter the
intestinal wall Fig.4, a). As far as I know such a condition has
mot beeén described in any species of earthworm. Bourne describes
them as a of the main intestino-tegumentary vessel a vessel
not present in D. communis. I believe, however, that these vessels
in D. communis serve a similar function to that of the ones referr-
ed to in Megascolex coeruleus. Fach dorso-tegumentary reaches the
body ern between the lateral and ventral rows of setae where it
divides into two branches, the larger extending dorsally along the
body wall and the smaller ventrally.

Ventro-tegumentary Vessels. (Figs. 4 & 5, V.1.).-- There is a pair

of these vessels in each somite posterior to somite XIII. They are
given off from the ventral just anterior to the septa and place it
in communication with the peripheral network. They reach the body

wall at about the same point as do the dorso-tegumentaries. The

9.

*

largest of the ventro-tegumentarios are in somite XIV. They are
larger than the dorso-tegumentaries and are quite short.
Ventro-intestinal Vessels. (wig. .4, V.I).-- These are branches of
the ventral placing it in connection with the betoswine. capillary
networks. In the middle of each somite posterior to somite XIII

the ventral Bives off one of these branches which enters the ventro-
"median perc Of the intestinal wall. In several of the larger spec-
Dtrens two or three of these vessels close together were found in :

~ several of the somites near the anterior end of the large intestine

4

(Tey are not as large as the branches from the dorso-tegumentaries

Be Pe intestinal Wall and are of such a length as to permit a free
ia
_noyonent of the ventral vessel.in the somite.

epee tateetinal Vessels. (Fig . 4, D.I).-- These vessels place the

mA

| orsal vessel in communication with the intestinal capillary net-
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|
|

ere. imere are no such vessels in somites I to XIV inclusive. In
abe .

“each of somites XV and XVI the dorsal gives off one pair of -dorso-

_intestinals. There are two pairs of such vessels in each of the

| x

somites posterior to somite XVI except in the last twenty or twenty-

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five in @ach of which there is but one pair. One pair, the larger,

is given off from the dors«1 just anterior to the middle of the so-

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nite while the second is given off near the posterior septum. Both
pairs ¢ enter the intestinal wall directly. The course of the anter-
or peir in the intestinal wall can be traced easily in a live spec=
n while that of the latter can be seen only in sections. In the
estinal wall they pass as distinct vessels to near the ventral

Pas 0
surface where they become ldst in the intestinal network.

Lateral Branches. (Figs. 1,2,5 & 5).-- The laterals bifurcate in V

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| and anterior to this agite they give off beveral branches to the
pharynx, gizzard and body wall. In somite V and cach succeeding

| somite to Ix inclusive @ach lateral gives off a branch to the body

wall, (latero-tegumontary}, one to the oesophagus and one to the sep-

| on The latter which is quite a large branch spreads out over the |

MRE septa! sending some of its smaller branches to the body wall.

- Corresponding branches of the laterals occur also in somites x, Aly
and XlI. From the septal branches of the laterals in somite XI,
branches extend through septum XI/XII to supply blood to the sperm

peacs in +5 The sperm sacs of somite IX situated on septum Ix/K

are ‘supplied by the septal branches of the laterals of that somite...

The ventral parts of the septa in somites VI, VII and VIII are sup-
plied aith small branches from the latero-tegumentary branches of

‘those somites. In somites V to VIII inclusive each lateral gives o

off a branch to the eeesbaa. thease are quite short and break up
into smaller branches immediately on reaching the oesophageal wall.

In somites IX to XILI inclusive there are in each somite at least

two branches from each lateral to the oesophagus. These branches

| are Larger ies those of the anterior somites. In somite XIII the

| laterals give off two or three branches to the integument just de-

| fore reaching it.

|

eee (Figs. 1,2.& 3).-- There are eight pairs of contractile

1

| hearts connecting the dorsal and ventral vessels. These include

all the contractile branches of the dorsal vessel. The five anter-

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sive are intestinal hearts, i.e. they are connected with both supra-
“intestinal and dorsal vessels.

The dorsal hearts arise from the vessel Just anterior to the
“septa and pass ventrad around the oesophagus to the ventral. In

| ‘the latero-ventral region (Fig .2,¥C), each heart diminishes slight-
1 ly in diameter and then immediately enlarges forming a chamber
“which contains valves Because of the presence of these valves I
have désignated these chambers as valvular chambers. “There are in
| Me gascolex coeruleus such enlarged portions of the hearts which he
calls muscular bulbs. Rach valvular chamber of somite VI in that
species gives off two branches neither of which reach the ventral.
In somites VIE to IX inclusive a branch is given off in each case
“below the, muscular bulb, but in the hearts posterior to IX this
bulb is at the juncture of the hearts and the ventral vessel.
“In.no case in D. communis did I find a a eff from

| the heart at any other place than at one of these valvular chambers
oe bulbs.

From each of these valvular chambers a branch*is given off to
the posterior septum and oesophagus. A short distance from the
“ventral there is a second valvular chamber from which: a branch pass-
bas to the body wall and one Go the ventral vessel entering tt some-

nat wetigrally. In the most mgees a” somites the branch joining

bes

| to the latter: The same condition of the dorsal hearts is repeated
| in each of the somites V to IX inclusive.

The condition of the intestinal hearts is more simple than

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that of the dorsal hearts (Fig. 1, I.H). They are considerably
| larger than the latter and the moniliform character of these ves-
| sels described in other worms is very conspicuous in them. Direct

= |

connection with the oesophagus and body wall.is wanting but whereas

———————— =
at

the dorsal hearts take their origin from the dorsal vessel only,

the intestinal hearts as the name indicates take their origin from
Poth dorsal and supra -intestinal vessels. An examination of Fig.l
Will show that the connections of the hearts with the supra-intes=
tinal vessel are much larger than those with the dorsal vessel.
These intestinal hearts have valvular chambers at the Junction or,
the hearts and the ventral vessel. All three intestinal hearts
have Frélations precisely similar to one another.

Capillary Networks. (Figs. 1,2,3,4,5,7,8 & 9).-- In addition to the

andes —_—s

vessels described, longitudinal trunks and transverse vessels,
there is in D. communis. as in all Oligochaeta, a system of smaller
vessels which form plexuses and may be termed capillaries. These
capillary networks may be divided into two series, the periphersd
and the intestinal.
_ Peripheral Networks.- The network first to be described under
this head is the aaa formed by the anterior branches of the dorsal,
ventral and lateral vessels (Fig. 8, AN). From the figure it will
be seen that this network is located on the body and pharyngeal
Walls in the first somite and the anterior part of the second. This
werk could be made out easily from a small specimen in a con-

pressorium.

The peripheral networks in each of the somites postcrior to

| somite XX are similar to one another. The dorso- and ventro-tegu-
|| ae 2) .

f

Te

ia

e é gee) eter

: 1 ohhh agnaiad .
‘aoe aie eda Forasd
or ee oibstu' eES
nae a) ae ie WANS '

in lyr eed’. eto (sen

if : 2 @4 if

‘ ‘7
d
ine ia satiety am
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LS.
| mentarieés reach the body wall at about the same point and pass dor-
sad along it giving off branches which pass into the muscular lay-
| ers, (Fig. 4, P.N). The ventro-tegumentaries of one somite have
direct connection with those of the following somite.

Anterior to somite Xx and posterior to XII the peripheral ner-
work is none complicated than that just described (Fig. 5). As
stated above, the laterals pass to the body wall in XIII and extend
posteriorly along it near the ventro-median line to somite XVIII.
The dorso-tegumentaries of these somites join these lateral-tegu-
‘mentary vessels from which branches are given off to the body wall
in each somite. -

The ventro-tegumentaries of somite XIV supply the body wall in
somites XII and XIII and in part in a few somites anterior to these.
‘It will be seen in figure 5 that there are no continuous trunks ex-
tending posteriorly along the body wall which the ventro-tegumente
aries join, but that each ventro-tecumentary Ct eee on
reaching the body wall and supplies parts of two somites, the one
in which the ventro-tegumentary is located, and the one posterior
to it. In each somite there are branches given off from the two
SE erich pass to the nephridia and in somites XVIII and xXx .to
| the prostate glands. The septal branches of the laterals in X to
xII inclusive reach the body wall and send branches along it both
dorsally and ventrally. | ;

The anterfor branches of te ventro-tegumentaries of somite
XIV extend anteriorly and fuse with the posterior branches, of the
warts of IX. Branches ere given off from these small trunks to

the body wall in each somite from X to XIV inclusive (Fig. 5). An-

een
iy tee alee

4 ae

(eaeAl eh ¢

> les ee
coe “

ae aE

'~

Alas OE
Pa

—

aw
“a oc c: wee ‘ s (avd!

he de

14.

terior to somite IX the peripheral networks are formed by branches
of the latero-tegumontaries and the tegumentary branches of the
dorsal hearts. These branches divide on reaching the body wall one
large branch extending dorsally and a smaller one ventrally. An-
terior .to somite V there is no particular segmental arrangment of
the network but both the body wall and the pharyngeal wall is sup-
plied with branches from the larger anterior branches of the dor-
sal, ventral and lateral vessesls (Fig.8)._

In @ach specimen in which the clitellum was well developed
branches of the peripheral system in this region are much enlarged
and distended with blood. In those specimens in which the clitel-
lum was not so well developed the branches of the avove system were
‘correspondingly small. Sections showed that the muscular layers
and hypodermis were well supplied with capillaries. I have no
doubt that Eee Gevelopment of the clitellum is correlated with the
well developed condition of the peripheral network in somites XIII
to XVIII.

Hypodermal Capillaries.- By means of sections capillaries were
recognized i the muscular layers and also in the hypodermis of the
body wall. The captilaries of the hypodermis in the:region of the
cliteilum in Lumbricus terrestus have been figured by Claparéde.
Reieerda (°63) first showed that capillaries alsc existed in the
hypodermis of other parts of Megascolex coeruleus and some species
of Perichaeta while Rosa ('87) showed that a similar comdition ex-
isted in the epidermis of Criodrilus.

7 From Figure 9 it will be seen that branches of the peripheral
system pass into the circular muscle layer and follow along it giv-

ing off many small capillaries which pass in among the hypodermal

:
- ates eneig nants ae
. Pad Apne bg hie uot : ah

Lhe @leh ata ail
v ae gi Lakes oct! EES

i. es

3.0 1) 0 eo Oe See ee ;

~

om, uit ~. do ama ae lage

ts ocd suet i uDOUnaree +
cs , TOK YS ‘mgt, Fo aol “tte LS ian, a |
LLY Qa w'gait aut oSahos eee
vue ag ener Spa suengiaaat pt

sik bile sou dats cumnilt to en

silele @tagde bewedd (Fas ae

‘ty | Sag 540 ia)
iw poideiel aad eons. aaa
a. Ee ae _ see “ye l alos. 4

iat se as kengi id oe

4 ‘i
a,

meas

Ae

ets
Le...
e@lis. Corresponding vessels found in the lecch by Lankester were

termed by him intir@-epithelial blood capillaries. Many of the hypo-

) Intestinal Networks. (Figs... 1,7 & 10). - In the region of the
Miesophacus from somite VI. to XVII the epithelial lining is closely
| longitudinally corrugated and especially so in somites IX to XIII
inclusive. This latter region of the oesophagus is enlarged and
well supplied with blood sinuses which are situated between the
-epitheleal lining and the muscular layers (Fig. 1, B.P). Beddard&6)
| describes a peculiar dilatation in the oesophagus of Microchaeta *
rappi which is quite similar to the region deseribed above in D:
communis. Where in M. rappi the network disappeared at the junc-
tion of this dilation with both the preceeding and succeeding parts
of the oesophagus, in sections of D. communis the longitudinal t:
trunks of the plexus could be traced as far as somite XVI. They |
were much smaller than those of the dilatation and more regularly
| arranged. There is also a dilatation at the Posterior end of the
oesophagus in Enchytraeus ventriculosus which seems to be similar
| to that of Microchaeta. Prof. Vejdovsky suggests that this organ

| functions as a liver but Beddard is of the opinion that the oesoph- —
| ageal dilatation of Microchaeta rappi as well as that of fuchyoraes
| us ventriculosus probably correspond to — calciferous glands of
_Lumbricus. I believe that this is the correct view and that the

| dilatation in the oesophagus of D. communis carries on a function

| Similar to that of the structures described above.

The intestine which begins at somite XVII is large and well

as | "att + eG aL ‘Teta: Ys! APL. i) <I ia

yn ak Ee, oe all Mea

aie aera gen
>. od: oda
peewee; a ibsdpotshy Au
} wiuot =} ere ee

B 3

4 S se Be iz
San

2 tbl Ste

g) cot. ie! Lam

ot Lito 6. cele ePeete,

she le 2 y types? ame

tiny Sifts stunt ~ tie “AAD Ty a -

« LYraege beget Sr sidedan | P-

* ates wilt qa beeen IOS ideas

aw be gy af? ct sim Seite. :
oe.
a

: sense)... 0p ne a
_ uae i, Ate er a1 | fi $

a

= “a ae

| or three somites. The typhlosole begins in somite XXI and extends
to the posterior end of the body. A representation of a cross séc-

| tion of the typhlosole is seen in figure 4. fhe typhlosole is ace

companied by an immense development of blood capillaries which are

o part of the intestinal network. This Bs evork is so dense and the

| interstices so small that the typhlosolar network appears to be a

- continuous sinus.

The Eterna network of the intestine is continuous from one

‘somite to another and is formed of nearly parallel longitudinal
| trunks connected by short transverse ones. The meshes are more .re-

a

| gular than are those of Megascolex. Figure 4 is from cross section

/

'of the intestine through a pair of dorso-intestinals showing the

) meshes near the ventral surface of the intestine.

COURSE OF BLOOD FLOW.
| Perrier (*74), according to Harrington (#99) and Bourne ('91),
came to the conelusion that in Lumbricus terrestris the blood flows
Mee orty.in the dorsal vessel, downward in the hearts and poster-
B isray in the ventral vessel, also that the blood flows from the dor-
ett

| sal vessel to the intestine through the dorso+ intestinals and in-

| to the dorsal from the dorso-tegumentaries. He also believed that

!

Ss oT

Hit oe aie meg
oo
haa re Ce ah

: . ~ - >. eid len

Se
4
*

*
| a
bag fide Ks *sonokil
; tpagygceleren santo a ie

\
“uWt ne (ations

Ge ha tery ss

incl ist te78) Oe eee

\ortruoh ,ideane Dearakaiar ae
off , Las2 ev laieeer

Py URE os a. ea an tiewa ng! ond ‘ena

ca» otf] Se ths aeeuset-oaeak, at ott.

ws atest 5 (eS=ord ce dog vteane Ley

Mog 2 +c gheeubog-ontete adete

st seein bree ehh towsd* ier

f ie

fyit oookd Tee aun 4de LOH) rs
7

4 <3 ‘oot Dau? a Beas © sepia!

' rs y POT aes th 2 ‘gots or er U via tf , 7 a >

sy

& probable course of blood flow which is almost directly apposed to
that given by Renta He agrees with Perrier that the dorso-tegz-
‘umentaries and dorso-intestinals play opposite r6éles but asserts
that in each set of vessels the blood flows in the opposite direc-

tion from that asserted by Perrier. Bourne says that the blood

the dorso-tegumentaries blood flows away from it; the ventro-tegu-
| mentaries supply the body wall and nephridia, and the ventral re-
ceives blood through the ventro-intestinals. The blood is carried
from the body wall to the intestine by the intestino-tegumentary
system. The course of the blood in the laterals is toward the

| posterior.

Harrington (*99) studied the course of flow in small trans-

| perent specimens of ee csiee bal: Urochaeta, Allolobophora, and Lum-
| a elheid between watch crystals and with the aid of a dissect-
ing lens. He came to the conclusion that the blood flows outward
from the dorsal vessel boranigh the dorso-tegumentaries ime the

_ sub-neural; into the dorsal from the dorso-intestinals; inward

from the body wall through branches of the dorso-tegumentaries in-
ia

to the sub-neural; either anteriorly or posteriorly in the sub-

_ neural; anteriorly in the laterals outward in the ventro-tegument-
|
_ aries, and into the intestine through the ventro-intestinals.

ite

|
|

terrestris and arrived at some conclusions different from any giv-

Johnston ('02) studied the course of blood flow in Lumbricus

iG
| flows through the dorso-intestinals into the dorsal and that through

fas

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‘
he a a oar

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sere 9 Law eee
d el ‘ay > FP Lend to-%

t ser hon tl We 2 = yet

sy ww i . = ~~ 1OR a
——" t i O° Gg
a
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a
(a
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: 4 \ ary
e 4° mmr
j @ ‘a Anu
i "er "ch. fev vam, seen

nok: sf poy Sao aie
i iy iigegoyvd wigip : he “io
a7 nae io ladedn Apasis (sentiennle
“oo p) Bian -jeregal & Sto nore
bat notre page Wiberg hei 1
wa o* %o e2wits oe bor doe MeDS if
i "49 , eetlent erg: poe. oe nevi
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LP
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18.

P atching the pulsations. He also tried injecting the vessels but
this served no purpose in determining the direction of the blood
_ flow.
Johnston supports Bourne in his view that tie blood flows for-
_ ward in the ventral vessel anterior to the hearts. He believes
that the blood flows into the dorsal vessel through both dorso-in-
e | testinals and dorso-tegumentaries although his experiments on the
datter were not wholly decisive. He also states that the blood
fTiows posterior in the laterals; the ventro-intestinals supply the
intestine, and the blood flows anteriorly throughout the entire
length of the dorsal. |

The results of my observations on the course of blood flow in
Diplocardia commmis given bclow are based on the examination of
_ more than fifty living specimens. The animals were anaesthetized
-and opened as previously described, and by clamping and cutting
‘the vessels, and watching the pulsations the direction of flow was
determined. For the study of the course of flow chloretone proved
| to be a better anaesthetic than chloroforn.
Dorsal Vessel.-- As in worms of other species the blood flows

va teriorly in the dorsal vessel throughout its entire length. The

“pulsations can be traced to the anterior end of the body. They
}

“can be very readily seen anterior to the hearts in quite small
i

|
Ventral Vessel+- By clamping and cutting the ventral vessel
“the course of flow posterior to the hearts was found to be backward

but anterior to the hearts the blood flows forward. The ventral

ae

Os Tin oT EP "od ta
- ecu he.
f 7h an = : p
J+ ee

. m8 Hane 7 es

bey 3
eN as

4
f

i
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i cae J :

19.

appeared in that part of the ventral anterior to the clamp. The

' ventral was cut at the point of the clamp and the blood flowed
‘from the posterior part. In several other specimens the ventral
was cut in a similar position with the same result.

Lateral Vessels.-- After clamping one of the laterals in VI

| the blood collected anterior to the clamp. The same vessel clamped
| farther back gave a similar result. This experiment was performed
repeatedly in different regions of the laterals and in no case did
| the blood collect posterior to the clamp. Quite a number of exper-
ints were made in which the laterals were cut, usually in somite
| XIII: where they pass from the oesophagus to the body wall and in

| almost every case the flow of blood was posteriorly. In one spec-
| imen one of the laterals was punctured in somite VI and clamped al-
ternately anterior and posterior to the puncture. When it was :
‘Clanped posterior to the puncture the flow of blood from the open-
| ing was increased considerably but on clamping it anterior to the

| puncture the flow practically ceased. The blood evidently flows
posteriorly in the laterals.

Dorso-:and Laterd-tegumentaries.-- Owing to the dense covering
(of chlorogogue cells it was quite difficult to see the blood in the

| dorso-teg gumentary vessels of dveancedd living specimens. It could

= resorted to for determining the course of flow was that of

Bevtting the vessels and watching the flow of blood. The dorso-teer-

| umentary vessels were cut at different points particularly near the

| body. wall in some and near the dorsal in others. Except in a few

ott ir likige 605 Sia.
fed

ot) at} YO (athe ae
ave ad yeaa ee
i “~ :

. é Lert Ay
i Olt

ts gan ver:
"18a." eae ‘el
pb, ++ nant apd Siena Lt!
‘fan ite (eto alee ai wi too. ° ye
iL begpeus sh 20- ol PUNEF, yikes
neta spaioed¢ of ty lt then aed,
(pmiifen B Ss abe Beau er oe bai
; 7
eee
RRR:

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‘
sb eg iia een 1 fv Ja¢

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20.

| blood from both ends of the sevord vessel. But while the flow from
the distal end was steady and quite pronounced there was at first
only a slight flow from the proximal end. In two cases in which

| the dorso-tegumentaries of the somites lying just posterior to the
hearts were cut near the dorsal the slight flow of the dorsal end,

| mentioned above, was followed by strong pulses of blood produced by
the pulsations of the dorsal vessel. Such cases by themselves might
Bento indicate that the flow of blood through the dorso-tegumen-
taries is from the dorsal: vessel to the integument but when consid-
ered - connection with the large number of cases in which the flow
mewes Obviously in the @pposite direction, it seems very certain that
the normal course of flow is centripetal.

Quite a number of experiments were made on the branches from
Fane dorso-tegumentaries to the intestine (Fig. 4AM) and in every

| case in which these vessels were cut the blood flowed only from the
(dorso-tegunentary) end, 3 |

| The latero-tegumentary vessels were cut in several different |
| specimens and in the majority of cases a large flow of blood came
from the distal end and only a slight flow from the proximal end.
In one or two cases these vessels were clamped but without giving

| satisfactory evidence as to the course of flow. In all experiments
in which there were any results they indicated a flow of blood from

| the body wall to the laterals. The results of experiments on the

| septal branches of the laterals which in somites X, XI, and XII

Tt Sie Te girly eG 4
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ee every piles yon

* ig §
ee Cn > Peres SAMS 5

a ty : © ¢

yi y hires eae | ie es ait 4
12)

, * B18! ~ ‘. a8 ffl A, lke > :
ov. walt, tpiié a Fiee ‘fag hi
Sy 2.0 Suemle sae @ laser eke Stier

-
é

-
«jie ‘=, pee ee5sea wt ae tue ~ in hh,
ee ts

al Eh paghe tas sect va aa : ORS
“pe Mnan' ‘6 é6)ieswr. ae SE aL § 2) hats

~% ah, nd dena Anji serene

Ri ry

——

Ae
extendto the body wall although not as decisive as in the case of

) the latero-tegumentaries, indicated the same direction of blood
ch

}

ss results than these. The ventro-tegumentaries were cut in
_ many specimens and the blood always flowed from the ventral only.
Experiments were made by cutting a oS ee and then
clamping the ventral alternately posterior and anterior to the sev-
ered vessel: In the former case the blood came from the ventro-
tegumentary more forcibly than when there was no clamp on the ven-
tral, and in the latter case the flow of blood almost ceased. Sim-
| ilar experiments of cutting were tried on the heart-tegumentaries
with a similar result. The experiments show that the blood flows
from the ventral through the ventro-tegumentaries and from the
hearts through the heart-tegumentaries to the body wall. In cons
“nection with the heart-tecumentaries) it may be mentioned that in
several specimens dorsal hearts were cut between the ventral vessel

and the lower valvular chamber of the heart and in each case there

| was a flow of blood from both the proximal and distal ends. Since

| blood from the ventral outward but under different conditions of
| 7
| pressure in the ventral and in the tegumentary system this may per-

Dorso- and Ventro-intestinal Vessels.-- The course of flow in
| the dorso-intestinals was determined by watching the pulsations
which in several specimens were easily seen. Several of these ves-

a
1
I

sels were cut and in every case the blood flowed from the intestine.

arog

” 7
ek bung, cleshah:

22.

These two experiments seem to warrant the conslusion that the pul-
Bations were from the intestine to the dorsal. A number of ventro-
-intestinals were cut in several different specimens and only ina
| few cases was there any flow of blood. The flow of blood was alr

| ways from the ventral. The small size of these vessels may perhaps

| nave permitted a rapid coagulation of the blood by the salt solu-

| tion with which the vessel came in direct contact.

| Supra-intestinal Vessel.-- (Fig. 1, S I). This vessel receives
hall of its blood from that region of the oesophagus over which it
Be462, by several short connecting vessels in each somite. As has
been stated, this blood flows through the intestinal hearts into

the ventral. In one specimen in which the pulsations of the dor-

| sal had ceased, it was empty of blood in the anterior region Bares

| the intestinal hearts were still pulsating and the blood was coming
| from the supra-intestinal vessel. The region of the oesophagus .

supplying the blood to tne supra-intestinal receives its supply of

blood from the laterals.

CONCLUSIONS.
The course of blood flow in Diplocardia communis according to
| the results of the above experiments is as follows:- The ger aes
receives its entire supply of blood from the dorso-intestinals and
| dorso-tegumentaries. This lends support to Johnston's ('02) cone
) clusion in regard to similar vessels in Lumbricus terrestris. The
greater part of the blood passes from the dorsal through the hearts
| toward the ventral while the remainder flows on to the anterior
_ network.

The direction of flow in the ventral vessel was found to agree

Bo

>

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= ra

: ' rc) ie. in If Ge a@

+ wi ef end) BOR
ae ae She ety ail

ae "4
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: ap reren ey *
so te qtea-ae, Sere

-

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<

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g 7

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25.

with that described by Bourne and Johnston, viz; t»nat the blood flows
‘anteriorly in front of the hearts and posteriorly back of them.
gat part of the blood which flows anteriorly enters the anterior

| network (Fig. 8, AN) where it mingles with that from the anterior

| portion of the dorsal.

From the anterior network the blood flows posteriorly through
eine laterals supplying the oesophagus in each. somite. That part

| of the blood in the laterals bat anes not pass into the oesophagus
flows into the clitellar part of the peripheral network thence in-
| to the dorsal vessel through the dorso-tegumentaries. (Fig. 5).

| The body wall is supplied with blood by the ventro- and heart-
Mi beamentarics. This blood after having passed through the periph-
} eral network eippiyine the body wall, nephridia and reproductive
jorgans, is in the anterior region, collected and returned to the
Bieerais by the latero-tegumentaries and septal branches of the
|lateral, and in the poSterior region to the dorsal vessel and in+
|testine by the dorso-tegumentaries. (Fig. 4).

| From figure 5 it will be seen that the body wall of somites

| F
1X =- XV inclusive are supplied with blood by the ventro-tegumentar-

the place of the heart tegumentaries of X, XI and XII. The blood
“supplying the latter somites is returned to the lateral vessels by
\the septal branches of the laterals mentioned above.

The intestine in somites XV - XX inclusive receives blood on-

jto XX it receives blood from both the ventro-intestinals and the

| Lo
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the size of the efferent and the afferent vessels of the intestine,
the amount of blood carried by each is about equal.

Mie direction of flow in the blood vessels of D. communis in
}s0 far as they correspond with those of Lumbricus terrestris agrecs

with that found by Johnston in the latter species.
LITERATURE CITED.

"85. On the Anatomy and Histology of Pleurochaeta Moseleyi,

meee Roy. Soc. Ndin., Vol. XXX, Part 2, pp. 481 - 509,
Pls. XXV - XXVII.

"86. On the Structure and Sretenetile Position of a gigantic
‘Barthworm from the Cape Colony. (Microchaeta rappi). Trans,

Zool. Soc. Lond., XII, pp. 63 = 76, Pl. XIV.

"95. A Monograph of the Order of Oligochaeta. 769 pp, 5 Pls.

91. The Nephridium of Lumbricus and its Blood Supply. Quart.
Suemece Mier. Sci., Vol. XXXII, (n.s.), pp. 293 - 334. Pls.

XXIII - XXV.

191. On Megascolex coeruleus, Templeton from Ceylon; together
: with a theory of.the course of the Blood in Earthworms.
| Quart. Journ. Micr. Sci. (n. s.) XXXII, pp. 49 - 87, Pls.VI-IX.
Garman, H. | |
es. On the Anatomy and Rist olecy of a New Earthworm. (Diploea |
cardia communis, Gen. et sp. Deve) cul). Pll. State. Lab. Nat.

Bigeye (tie art. TV, pp. 45 - 77,+Pls. I - v.

*
&*
?
! ih oe

LGizey (ASO
, ele sity “Saas ytd

mee eee Wy ieee ee

20.

Jarrington, N. R.

"99. The Calciférous glands of the Barthworm, with Appendix
‘fon the Circulation. Jour. of Morph. » Vol. XV, Supplement, pp.
Medes - 162. Pls. VI - Ix.

Johnston, J. B. and Johns on, Sarah W.

| Sea.) course Of The Blood Fiow in Lumbricus. Amer. Nat., Vol.
Mee) REXVI, No. 454, pp. 317 - 327.

ae W.

"94. Die Regenwurm - Fauna von Florida and Georgia. (Zoods
Seue@e Abth. f. System. VIII, Bd.2 Hft. pp. 177 - 194.

"00. Das Tierreich, 10 Lieferung. Oligochaeta. Berlin.
ererrier, E.

74.. Etudes sur ait organisation des Lombriciens terrestris.

:, (Urochaeta). Arch. Zool. Exp., t. III, pp. 331 - 530, Pls.

ot - XVII.
Rosa, D.
187. Sul Criodrilus lecuum. Studio Zoologico ed Anatomico.

_eimounc. Torineg, (2), XXXVIII, pp. 15, One plate.

EXPLANATION OF PLATES.
Anterior network.
Blood capillaries.
Blood plexus of the oesophagus.
Layer of connective tissue.

Circular muscle layer.

Dorso-intestinal vessels.

Dorsal vessel.

ts ie
‘e Df . aiakee

: ,@eé?alpe sl
: ae
‘ t - :
' =

£<¢ aq beoiff
7
}: FT fod gs hd ie | 7 : a

f : oe a wii
re ni Bt - cag ead. :
; . ie >

i : - : _ : :
el tebhy ban tae7Gsee

fr

: . - | pe aoe :
a pe i

Epithelial cells.

Epithelial layer.
Dorsal heart. |
Intestinal heart.
“Int. Cap. Intestinal capillary network. | |
Intestine. |
| Intestinal network.

“LM ei Longitudinal musele layer.
el Latero-tegumentary vessel.
Ly Lateral vessel.
Intestino-tegsumentaries. (2?)

Nn Blood vessels of the nephridia. a

Nerve cord. | ie
Oesophagus. | | ‘sg
Blood vessels of the prostate glands.

Peripheral network.

Supra-intestinal vessel.

: Br. Septal branch.

Typhlosole

Valvular chamber.
Ventro-intestinals.

Ventre vessel.

|
Walves.. : | ;
: : |

Branch from the heart to the oesophagus and septum.

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the somite. X 30.

wig. 2.- A diagramatic representation of a cross section through a
pair of dorsal hearts showing the important vessels of the
somite. <X 35.

Fig. 3. - A diagramatic representation of the vessels of somites
fee mn @na XLIT. X about 17.

Pig. 4.- A diagra atic representation of a cross section of the

intestinal region Dae

| Fig. 5 represents the peripheral network of the clitellar region.
X about 18.

Fig. 6.- A diagramatic sketch of the relation of the dorsal and
ventral vessels at the posterior end.

| Meee, represents a part of a cross section of the intestinal wall
Zz 50C,

Fig. 8.- A diagramatic representation of the relations of the dorsal,
lateral and ventral vessels in the anterior region. X about 8.

eee 2.— From a cross section ane body wall, showing the hypoe::
| - dermal capilaries. % 500.

‘Fig. 10.- From a cross section of the oesophagus of somite XVI. X 40.

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