a?

nm
cf

TROD UC ad

ANATOMY,
h,

aoe
J ees oF 2

hae ee,
2 oy “72S 2) ail
oad

Pae
 ul
rn '
ia ia
ee)
yuh U
Rly
a) Les
A ,

pve ae j a).
S Oawe ie
ee au,
Lig
Aesiny

¢
aye a Pa iegle Ue
és Poyia
catty
1 i isd i
at
atvl eave PGP ee: itary yt§ rai
ses ai ani
ra ae ny ie cet week prvi | if Mid
secre iia we PapRotei'Ne, Ovegelags" piel |

\ ee iat sacha:
 . ue
“9i fe :
ioe pa gy. Ts hi

* 4

ae
"|
ale \ ty
aK
EAIM3L7
1394 bi ae eis OG
Kerr AN
|
INTRODUCTION TO

ANATOMY, HISTOLOGY, AND

EMBRYOLOGY.

BY

A, MILNES MARSHALL, M.D, D.Sc, M.A, F.R.S,

3 Py

LATE FELLOW OF ST. JOHN’S COLLEGE, CAMBRIDGE 5

PROFESSOR IN THE VICTORIA UNIVERSITY 5

BEYER PROFESSOR OF ZOOLOGY IN OWENS COLLEGE.

P aM Fitth Loition.
i me
VA \ REVISED AND ILLUSTRATED.

‘> se Nn
1 =~ iene i!
hon > =
j- ws =|
4 ee 0 |
‘oe Pe J MANCHESTER:
( ]./E. CORNISH, 16, ST. ANN’S SQUARE
a>
‘aay LONDON:
a,,
..
ITH ELDER, & CO., WATERLOO PLACE.
ff a
1894.
 PREPACE TO THE FIRST EDITION.

THE Owens College Course of Elementary Biology, which forms

part of the scheme of study prescribed by the Victoria Univer-
sity, is of a rather more extended and comprehensive nature
than the courses held elsewhere under the same name; and
experience has shown me that there is want of a book that will
guide and direct the student through the practical part of his
work, the whole ground of which is covered by no one of the
existing manuals. It is to meet this want that the present
little work has been prepared.
This first instalment of the work consists of an Introduction,
containing practical instruction in the methods employed in
biological investigation ; followed by the application of these
methods to the examination, both anatomtical and _histological,
of an actual animal. For this purpose, the frog has been
selected as being easy to obtain, convenient to dissect, and a
fairly typical example of the great group of Vertebrate animals.
Where, from its small size or for other reason, the frog has
proved unsuitable, other animals have been substituted for it.
For convenience of reference, and in order to definitely stamp
the practical character of the work, directions for dissection,
etc., have throughout been printed in italics.
It is not expected that the student should do the whole of
 iv ;

the work here given the first time he goes over it. The dis-
section of the muscles and of the cranial nerves should only be
attempted if time remain after the other work is completed.
In preparing this first part I have received very valuable
assistance from Dr. Hartog, Demonstrator of Biology in the
College, and from my friend and pupil Mr. C. H. Hurst. I am
also much indebted to Prof. Gamgee and to Mr. Waters for the
important help they have given me in the histological portions.

OwENS COLLEGE,
August, 1882.
 PREPACE TO THE THIRD EDITION.

SincE the appearance of the former editions, the publication of

the Junior Course of Practical Zoology by Dr. Hurst and
myself has completed, so far as Animal Morphology is concerned,
the Owens College Course of Elementary Biology.
I have retained The Frog in its original form, as the mode of
treatment is more elementary, and in some ways different from
that of the larger book. The present edition has been carefully
revised, and an account of the development of the Frog has
been added. In preparing this account, I have had the
advantage of examining some excellent series of sections
mounted by my friends and former pupils, Mr. H. Sidebotham,
and Mr. A. E. Giles.

OWENS COLLEGE,
July, 1888.
 vi

PREFACE TO THE FIFSH EDITION:

THE principal changes in the present edition are in the chapter

on Embryology, in which some new figures have been intro-
duced and some mistakes corrected.
In revising the book I have received valuable aid from my
friends Dr. Hurst and Mr. Gamble, Assistant Lecturers in
Zoology in the College.

OweENS COLLEGE,
December, 1898.

NO iE:

THE preparation of the present (fifth) edition for the printer

was Professor Marshall’s last professional act, and was com-
pleted only a week before his death.
In correcting the proofs I have made no change of any
importance.
C. Herpert Hurst.
OwENs COLLEGE,
Apru, 1894.
 oO NENtS:

INTRODUCTION. PAGE
Laboratory Rules. Apparatus required. Dissection. Drawing.
Use of the Microscope. eee of Microscopical
Objects. Section: Cubmn es. 2200) i. oncnsssnssavieere: neancees, Aobe

CHAP. I.
GENERAL ANATOMY OF THE FROG.
External Characters. Buccal Cavity. Abdominal eae
Peritoneum. Digestive Organs............. 13-21

CHAP. II.
THE VASCULAR SYSTEM OF THE FROG.
The Heart. The Veins. The Arteries. The Structure of the
Heart. Microscopical Examination of Blood .. 22-37

CHAP. IIT.
ELEMENTARY Histonoey.
Epithelium. Sieeciis Muscle. Connective Tissue. Cartilage.
TB 5 parerAh emo at heelSt MOCHA gee NY pra ER

CHAP. IV.
Tue SKELETON OF THE FROG.
The Axial Skeleton. The Appendicular Skeleton .................. 50-64

CHAP. V.
THe Muscutar System oF THE F Roc.
Muscles of Trunk. Muscles of Head. Muscles of Hind-limb... 65-77

CHAP. VI.
THe Nervous System or THE FRrRoa.
The Central Nervous System. The Peripheral Nervous
System. Histology of Nerves .0..i: ges sncsoe-cocspe-cernses 78-96
 Vili

CHAP. VIL.

THE EyE AND Har.

The Eye ofthe Frog. The Eye of the oe ores “ee loee
of the Eye. The Ear of the Frog... <g 7-105

CHAP. VIII.
Tue REPRODUCTIVE ORGANS AND THE CLOACA.
The Male Frog. The. Memale Prog. ..ci.¢...gecnc: cap scnssens 200 Scene e.d OG ans

CHAP. IX.
DEVELOPMENT OF THE FROG.
General Account. Formation of the Egg. Maturation of the
Keg. Fertilisation. Segmentation. Formation of the
Germinal Layers. Development of the Nervous
System. Development of the Sense Organs. Develop-
ment of the Alimentary Canal, The Gill Clefts md
Arches. The Vascular System. The Muscular System
and the Celom. Development of the Skeleton.
Development of the Urinary System... ... 109-155
 INTRODUCTION.

I.—_LABORATORY RULES.
1. The Laboratory is open to members of the Biology Class
from 10-30 to 5 on Tuesdays and Thursdays. Students are
required to attend at least three hours on each of these days.
2. Each student has a definite seat assigned him in the
Laboratory, which he is not allowed to change without per-
mission.
3. Each student has the free use of the drawer and locker
belonging to his seat: the key may be obtained on payment of
a deposit of half a crown, which will be returned if the key is
given up before the end of the term, but otherwise will be
forfeited.
4, All necessary reagents and specimens for dissection are
provided by the Laboratory, but each student is required to
furnish himself with dissecting instruments, note book, and
pencil, as explained in the next section. Microscopes are pro-
vided by the College at a charge of five shillings per term.

II—LIST OF APPARATUS REQUIRED.

Each student is required to provide himself with the fol-
lowing :—
1, Two or three scalpels or dissecting knives of different sizes.
2. 'Two pairs of forceps, one large and one small. Both pairs
should be straight, and should have the tips roughened in order
to secure a firmer hold.
3. Two pairs of scissors; one pair large and strong, for cut-
ting bone and other hard tissues; the other pair small, for
fine dissections. The latter pair should have the blades bent
at an angle (elbow scissors), In selecting scissors be careful te
see that they cut quite up to the points of the blades.
4, A pair of stout needles, firmly mounted in handles.
B
bo ON DISSECTION.

5. A pair of the finest sewing needles, mounted in handles:

only about a quarter of an inch of the needle should project.
They are used for teasing histological preparations.
6. A seeker, z.¢., a blunt needle mounted in a handle, and
bent at an angle half an inch from the end.
7. A metal blow pipe: and a glass cannula with india-rubber
cap.
8. A pocket lens, containing two or three lenses mounted in
a handle, and giving when combined a magnifying power of at
least six diameters.
9. Slides and coverslips, for mounting microscopical speci-
mens. The coverslips should be the thinnest sold (No. 1).
10. A blank note book, for drawing in; an HB pencil, and a
piece of india-rubber.
11. A cheap pair of compasses, for measuring the dissections.

III—ON DISSECTION.
The object of dissection is to separate the several parts and
organs from one another, so as to define their boundaries and
display clearly their mutual relations. Dissection consists
mainly in removing the “connective tissue” which binds the
several parts together.
The following rules should be carefully attended to :—
1. Pin down the animal firmly to the dissecting board.
Never attempt to dissect a specimen that is not so fixed.
2. In pinning out a dissection stick the pins in, not vertically,
but obliquely, so that their heads do not get in the way or
obscure the dissection. )
3. Never cut away anything until you are quite certain what
it is you are removing.
4. Put the part you are dissecting slightly on the stretch;
e.g., When dissecting the bloodvessels or nerves of the throat,
distend it by passing a small roll of paper or the handle of a
seeker down the esophagus ; or when dissecting the muscles of
the leg, pin out the leg in such a position as to stretch the
muscles you are cleaning.
5. In cleaning bloodvessels or nerves always dissect along
them and not across them ; and avoid laying hold of them with
the forceps. Similarly when cleaning muscles, dissect along
their fibres and not across them,
 ON DRAWING. 3

6. Fine dissections should be done under water, which sup-

ports the parts and greatly facilitates the operation. A stream’
of water allowed to play gently on the dissection from time to
time is often a valuable aid. |
7. The dissection of muscles, and still more of nerves, is
greatly aided by placing the specimens in spirit for a day
before dissecting.
8. Keep your instruments clean and sharp. Be careful not
to blunt your fine scissors or scalpel by using them for cutting
hard parts.
9. If you get in a muddle, stop and wash the dissection
thoroughly under the tap before proceeding further.

IV.—ON DRAWING.
It is absolutely essential to draw your dissections, and this
must on no account be omitted. Keep a separate book for your
drawings, and draw every dissection you make. Do not be
discouraged if you find it difficult at first : you will never regret
time spent on it.
The following rules will be useful to those who have not
learnt drawing systematically :—
1. Make your drawing to scale, z.e., either the exact size of
the natural object, or half or double or treble that size, as the
case may be.
2. In commencing a drawing, first determine by careful
measurement the positions of the principal points, and sketch
in lightly the whole outline before finishing any one part.
3. If the object you are drawing is bilaterally symmetrical,
draw a faint line down the middle of your paper, and sketch in
the left hand half first; by measuring from your median line
it will be very easy to make the two halves symmetrical.
4. Name on your drawing the several parts shown, and
mark also the scale adopted. If your drawing be of the
natural size mark it thus— x 1; if it be double the size of the
object mark it x 2; if half the size, x 4, and so on.
5. Draw on one side of the page only : and write an explana-
tion of your drawing on the opposite page.
6. Coloured pencils are very useful, and water-colour paints
still better. Keep certain colours for particular organs or
tissues ; e.g., when drawing the skeleton colour the cartilage
4 THE MICROSCOPE.

blue, the cartilage bones yellow, and the membrane bones either
red or white ; when drawing the bloodvessels colour the arteries
red and the veins blue.

V.—THE USE OF THE MICROSCOPE.

The microscope consists essentially of a stand and a body, the
latter of which bears at its ends the lenses by which the magni-
fying power is obtained.
The stand is an upright pillar, the lower end of which is
attached to a heavy foot to ensure steadiness. A little way
above the foot the stand supports a horizontal plate—the stage—
on which the object to be examined is placed. The stage is
perforated in the middle by a hole, the size of which can be
varied by means of diaphragms. Through this hole light is
directed on the object to be examined by means of a mirror
attached to the stand below the stage. Above the stage the
stand supports a vertical tube, in which the body of the micro-
scope slides up and down.
The body is a tube, in the upper end of which is placed a
combination of lenses, known as the eyepiece, while to the lower
end is screwed another combination of lenses—the objective.
A microscope is usually provided with a couple of eyepieces
and a couple of objectives of different magnifying power. An
objective magnifying only a small number of times is called a
low power; one magnifying many times (200 diameters or more),
a high power. Similarly eyepieces are spoken of as high or low
according to their magnifying power.
In order that an object may be seen clearly the objective
must be at a certain definite distance from the object, this
distance varying with different objectives, and to a slight extent
with different observers. The higher the power employed the
closer must the objective be brought to the object. As the
position of the object on the stage of the microscope is fixed,
this distance is regulated by moving the body of the microscope
up and down in the tube in which it slides.
This process of focussing is effected in two ways:
(1) By simply sliding the body up and down by hand: this
is known as the coarse adjustment ; it should be performed with
a slight screwing motion, and can only be used when low
powers are being employed.
 THE USE OF THE MICROSCOPE. 5

(2) With high powers the objective has to be brought so

close to the object that a more delicate method of adjustment
is necessary. This fine adjustment is effected by a screw with a
milled head placed at the top of the vertical pillar forming the
stand. By turning the head from right to left, in the direction
of the hands of a watch, the body of the microscope is lowered
and the objective brought nearer to the object: by turning in
the reverse direction the objective is raised.
In using the microscope attend to the following rules:—
I. Always examine an object first with the low power. Having
adjusted the eyepiece and objective, direct the light up the tube
of the microscope by means of the mirror, and then place the
object on the stage. Twist down the body until the objective
is about a quarter of an inch from the cover-glass; look down
the microscope, and gradually twist the body up until the
object becomes visible. Focus accurately by means of the fine
adjustment screw.
2. When using a high power begin with the objective close to
the cover-glass, and then focus with the fine adjustment. It
will facilitate the process if, while focussing with the right hand,
you move the object about slightly with the left hand.
3. Take extreme care never to let the objective touch the
cover-glass; and never to touch or allow any dirt to get on the
face of the objective. The face of an objective cannot be cleaned
without doing harm to it.
4. Should by any chance a drop of glycerine get on the face
of the objective, wash it carefully with water from a wash-
bottle, and wipe it gently with a silk handkerchief or piece of
chamois leather. Should Canada balsam be allowed to get on
the objective, do not attempt to clean it yourself, but hand it at
once to the assistant.
5. See that the body of the microscope slides smoothly in its
tube. If it does not, remove it, and clean it by rubbing with a
few drops of olive oil: wipe off the oil before replacing the body
in the tube.
6. Keep both eyes open when looking through the microscope:
a very little practice will enable you to do this, and it will save
you much fatigue. Also get into the habit of using either eye.
7. With a high power, use a small diaphragm: the amount
of light will be somewhat diminished, but the clearness and
definition of the object much increased.
 6 THE PREPARATION OF MICROSCOPICAL OBJECTS.

8. When examining an object, keep one hand on the fine

adjustment, and keep screwing it up and down slightly the whole
time: in this way parts of the object at different depths are
brought into focus successively, and a clearer idea of the object
is obtained.
9. If the object appears dim or dirty, find out where the
fault lies in this way :—
While looking down the microscope, turn round the eyepiece
with your right hand: If the dirt turns round too, remove and
clean the eyepiece. If the fault is not in the eyepiece, move
the slide about gently; if the dirt moves with the slide, remove
the slide and clean it. If the dirt does not move with either
the eyepiece or the slide the fault is almost certainly in the
objective, which should be removed and examined; if dirty, it
must be cleaned very carefully with a piece of silk or chamois
leather.

VI—THE PREPARATION OF MICROSCOPICAL OBJECTS.

In mounting microscopical objects be careful that your slides
and coverslips are thoroughly clean. Slides should be labelled
as soon as they are prepared, and should be kept in a box or
cabinet in which they lie flat.
A. Methods of Mounting.
There are various media in which objects may be mounted.
The method of procedure is much the same with all. Put a
small drop of the fluid in the middle of the slide, place: the
object in the middle of the drop, and arrange it with needles in
the position desired. Then place the cover glass carefully on
the top, letting it rest by one edge on the slide and supporting
the opposite edge by a needle: withdraw the needle gradually
so as to let the cover-glass down slowly, and drive out any air-
bubbles there may be in the fluid. If any air-bubbles still
remain, leave them alone, as they will probably work out by
themselves. Be careful not to use too large a drop of your
mounting medium, and above all things be careful not to let
any of it get on the top of the cover-glass ; should this happen,
the cover-glass must be removed at once and the specimen
mounted afresh with a clean one.
The most important mounting media are the following,
 THE PREPARATION OF MICROSCOPICAL OBJECTS. ~I

1, Normal Salt Solution: a 0°75 per cent. solution of com-

mon salt in water. This is very useful in the examination of
fresh specimens of animal tissues, as, unlike water, it has
practically no action on them. It-cannot be used, however,
for making permanent preparations.
2. Glycerine can be used either pure or diluted with its own
bulk of water. If the preparations are intended to be per-
manent, @ narrow ring of cement must be painted round the
edge of the cover-glass to fix it to the slide.
3. Canada Balsam is the most generally useful, requiring no
cement. Specimens that are to be mounted in balsam must
first be deprived of all water they may contain by placing for
an hour or so in absolute alcohol, and should then, before
mounting, be soaked for a few minutes in oil of cloves or tur-
pentine in order to clear them, i.e., render them permeable by
the balsam. Canada balsam, if too thick, may be diluted with
chloroform, turpentine, or benzole.

B. Teasing.
The object of teasing is to separate the several parts of a
tissue or organ from one another in order to show their minute
structure.
The fragment to be teased should be placed on a slide in a
drop of the medium in which it is to be mounted, and then
torn up into shreds by means of a couple of needles held one
in each hand. The process is often greatly facilitated by
placing the slide on a piece of black paper, which renders the
particles easier to see. When torn up as finely as possible, a
cover-glass is placed on as before. The two rules to be borne
in mind in teasing are the following.
1, Take a very small fragment to commence with.
2. Tease it as finely as you can. Your object is to separate
the component parts from one another.

C. Maceration.
The process of teasing is in many cases facilitated by pre-
' viously macerating the specimen, @.e., soaking it in some fluid,
which, while preserving the individual cells, tends to loosen
them from one another. The most important macerating fluids
are as follows,
8 THE PREPARATION OF MICROSCOPICAL OBJECTS.

1. Ranvier’s Alcohol: a mixture of one part of strong spirit

with two parts of water. The specimen should be put fresh into
the mixture and allowed to remain twenty-four hours or more.
2. Miuller’s Fluid: a solution of bichromate of potash with
a little sodic sulphate in water.
D. Staining.
Various re-agents are employed for the purpose of staining
preparations ;some of these merely colour the whole prepara-
tion more or less uniformly, but the most useful ones are those
which stain certain parts of the cells only, or at any rate stain
these much more strongly than the other parts. The most im-
portant are the following.
1. Hematoxylin. There are various preparations of hemato-
xylin, or logwood, used in microscopical work: the best is that
proposed by Kleinenberg and called by his name. It is pre-
pared thus :—
(a) Make a saturated solution of crystallised calcium chloride
in 70 per cent. alcohol, and add alum to saturation.
(6) Make a saturated solution of alum in 70 per cent. alcohol,
and add (a) to (>) in the proportion of 1 to 8.
(c) To the mixture of (a) and (b) add a few drops of a
saturated solution of hematoxylin in absolute alcohol.
The specimens, which must be perfectly free from all trace
of acid, should be left in the hematoxylin in a covered vessel
or stoppered bottle for from one to twenty-four hours, according
to the size of the specimen and the depth of staining desired,
and then placed in strong spirit for some hours before mounting.
Heematoxylin stains the nuclei of cells much more strongly than
the other parts.
2. Borax-Carmine. This, which is perhaps the most generally
useful of all the stains in ordinary use, is prepared as follows.
Dissolve 2 parts of carmine and 4 parts of borax in 100 parts of
water: add an equal volume of 70 per cent. alcohol; let the
mixture stand for a couple of days, and then filter.
Specimens may be left in borax-carmine for from one to
twenty-four hours, or even for two or three days: on removal
they should be placed in acid-alcohol—+.ec., 70 per cent. alcohol
to which a few drops of hydrochloric acid have been added—
until they become a bright scarlet colour, when they should be
 THE PREPARATION OF MICROSCOPICAL OBJECTS. a

transferred to 70, and then to 90 per cent. alcohol, in which

latter they may be left till required. The time of immersion in
acid alcohol will vary, according to the nature and size of the
specimen, from a quarter of an hour up to a day or more.
3. Picro-carmine is a very useful, and to a certain extent a
differential stain, as it colours the several tissues different
tints. It may be prepared thus. Dissolve 1 gramme of carmine
in 4 cc. of liquid ammonia and 200 ce. of distilled water. Add
5 grammes of picric acid; shake the mixture well for some
minutes, and then decant from the excess of acid. Leave the
decanted liquid for some days, stirring it occasionally : then
evaporate it to dryness, and to every 2 grammes of the dried
residue add 100 ce. of distilled water.
Picro-carmine answers best with specimens preserved in 70 per
cent. alcohol. They should be left in the stain for a day, and
then placed in 70, and afterwards in 90 per cent. alcohol. Some
specimens give better results when washed freely with water
on removal from the picro-carmine, and then placed in | per
cent. acetic acid for an hour before transferring to alcohol.
4, Magenta stains very rapidly but diffusely: the colour
also is not permanent.
5. Silver Nitrate. A} per cent. solution in water stains the
intercellular substance, which binds together the several cells of
a tissue, much more strongly than the cells themselves, and is
therefore chiefly used when we wish to render prominent the
outlines of the individual cells. The specimens should be placed
fresh in the silver solution for from two minutes to a quarter
of an hour, then washed thoroughly with distilled water, and
exposed to the light until stained sufficiently deeply, when they
may be mounted in glycerine. Such preparations are rarely
permanent, as the reduction of the silver, to which the staining
is due, continues until the specimens ultimately become too
dark to be of any use.
6. Osmic Acid. A 1 per cent. solution of osmic acid in water
forms an extremely useful staining reagent. It is especially use-
ful for the detection of fat, which is stained by it a dark brown
or black colour. Specimens, which must be quite fresh, should
only be left in it a few minutes, and may then be mounted in
glycerine, or else washed, dehydrated, and mounted in balsam,
7. Acetic Acid. Although not strictly a staining agent
10 ON SECTION CUTTING.

inasmuch as it does not colour the specimens, acetic acid may

conveniently be mentioned here as it is used for the same pur-
pose as the true stains, ze. for the sake of rendering certain
parts of the cells especially distinct. Acetic acid, of which a
1 per cent. solution is employed, causes the protoplasm of cells
to swell up and become transparent, and brings the nuclei into
special prominence. It is used with fresh specimens.
VII—ON SECTION CUTTING.
Many tissues and organs can only be studied satisfactorily by
cutting them into thin sections, and this method of investiga-
tion is of such importance as to require special notice. There
are three chief stages: Hardening, Imbedding, and Cutting,
which will be noticed in succession.
A. Hardening.
Before the object can be cut into sections it is necessary to
harden it; this may be effected by freezing, but the more usual
plan is by means of reagents. The general action of these
hardening reagents is to coagulate the protoplasm of the tissues ;
and the objects to be attained are to effect this coagulation
quickly, before the tissues can undergo any alteration ; and
thoroughly, 2.e., throughout the whole thickness of the object to
be hardened. ‘To ensure the latter result it is always advisable
to use small pieces of the substance to be cut.
The hardening reagents in most common use are as follows.
1. Alcohol. Specimens may be placed at once in 70 per
cent. alcohol ; and thence transferred after a couple of days to
90 per cent., in which they may be left till required.
2. Osmic Acid, For this purpose a 1 per cent. solution in
water is used : it acts almost instantaneously, and so allows no
change to occur in the tissues ; it has also the merit of staining
the tissues as well as hardening them. It can, however, only
be employed when the specimens are very small, as it hardens
the surface layers so rapidly that it is unable to penetrate
beyond a very slight depth. A few minutes immersion is
usually sufficient.
3. Corrosive Sublimate. A saturated solution in water is
employed, in which the object is placed for half an hour or more.
After removal it is thoroughly washed with water or weak alcohol,
and then transferred to 70 per cent. alcohol before staining.
 ON SECTION CUTTING. Dt

4, Chromic Acid. A 0°25 to 0°5 per cent. solution of chromic

acid in water is a useful hardening reagent; it acts much more
slowly than osmic acid, but penetrates to greater depths.
Specimens should usually be left in the solution for one or more
days. |
5. A Mixture of chromic acid with a few drops of osmic acid
is often very useful, as it combines the advantages of both
reagents.
6. Picric Acid is a very valuable hardening reagent, of which
the best preparation is Kleinenberg’s. Specimens should be left
in it from 12 to 24 hours. It is prepared thus: with 100 ce.
of water make a cold saturated solution of picric acid: add 2
ec. of concentrated sulphuric acid: filter, and add to the filtrate
three times its volume of water.
B. Dehydration.
Specimens that have been hardened in any of the preceding
reagents, except alcohol, should, on removal, be placed for a few
hours in 30 per cent. alcohol, and then transferred to 50 per
cent. alcohol: on the following day they should be transferred
to 70 per cent. alcohol, which should be changed daily until the
Specimens are free from the hardening reagent: they may then
be left in 90 per cent. alcohol until required.
C. Staining.
The hardened specimens, if not too large, may now be stained
with either hematoxylin, borax-carmine, or picro-carmine ; they
should then be replaced in 90 per cent. alcohol. If the
specimen is too large to stain whole, the sections must be stained
atter they are cut.
: D. Imbedding.
The preparation of sections is greatly facilitated by imbedding
the specimen in some waxy substance. For this purpose various
materials have been employed, but by far the most useful is
parafiin, which is used in the following manner:
The stained specimen is placed in absolute alcohol for an
hour or two in order to completely dehydrate it. It is then
transferred to turpentine, in which it is left for half an hour or
more until completely saturated. From the turpentine it is
transferred to melted paraffin, which is kept by means of a
water bath at a temperature just above its melting point. In
12 ON SECTION CUTTING.

this it is left for several hours, or even for a whole day, in

order that it may be thoroughly permeated. It is then placed
in a small box of paper, or other material, filled with melted
paraffin. By means of hot needles it can readily be arranged
in any desired position ; and the paraffin should then be cooled
quickly.
E. Section-Cutting.
When thoroughly set the block is removed from the box, and
the paraffin pared away with a knife until the specimen just
comes into view.
The block is then placed in a microtome, and cut into thin
sections. ‘These may be transferred one by one to a slide, but
a great saving of time is effected by the method of cutting
continuous ribbons, devised by Mr. Caldwell.
This depends on the fact that if the paraffin is of proper
consistency the successive sections, as they are cut, will stick
together at their edges, so as to form a ribbon. To ensure this
the edge of the razor should be placed at right angles to the
direction of stroke, and the edges of the block of parafin cut
parallel to one another, and to the edge of the razor. If for
any reason it is desirable to imbed the specimens in a paraffin
too hard to form ribbons, the block should before cutting, be
coated with a layer of soft parafin, by dipping it for a moment
in a dish of melted soft paraffin. This outer coating should be
left on the sides of the block parallel to the edge of the razor,
but cut away from the sides at right angles to it.
The razor should be used dry: and the sections, when cut,
placed on slides painted, just before they are used, with a thin
layer of a mixture of collodion and oil of cloves in equal parts.
The slide is then heated by a water-bath to a temperature not
exceeding 55°—60° C., so as to melt the paraffin and evaporate
the oil of cloves. The melted paraffin should then be washed
off by turpentine, when the sections will remain fixed to the
slide by the collodion, and may be mounted in balsam in the
usual manner.
Instead of the mixture of collodion and oil of cloves, a solution
of shellac in absolute alcohol may be used: this should be
spread over the slide in a thin layer by means of a glass rod,
and allowed to dry. Immediately before being used the slide
should be brushed over with oil of cloves.
 CHAPTER I.

GENERAL ANATOMY OF THE FROG.

\ rd. \
=e VA
= SS O's Y \

IeS\ a \

Fig. 1. The Common Frog (Rana temporaria) : (from Ecker.)

A. External Characters.
Lay the frog on a board before you ; note, and make drawings
showing the following points :
1. The division into head, trunk, and limbs ; and the absence.
of neck and tail.
2. The two great surfaces.
a. The dorsal surface, or back, is directed upwards when
the frog is in the natural position.
b. The ventral surface, or belly, is directed downwards
towards the ground.
 14 GENERAL ANATOMY OF THE FROG.

3. The skin is moist and smooth ; and devoid of hairs, scales,

and claws. The colour of the skin is variable in different specimens
and at different times: it is mottled on the dorsal surface, paler
on the ventral.
4. The head is flat and triangular, with a blunt apex directed
forwards.
At the sides of the head are the eyes, which are large and
prominent. Each eye has two eyelids, of which the upper is
thick, pigmented, and almost immoveable, while the lower is
semi-transparent and freely moveable.
Behind the eye on either side is an obliquely placed elongated
patch of a dark colour, in the middle of which is a circular
area—the tympanic membrane—supported by a firm marginal
ring. :
5. The limbs. There are two pairs of limbs, fore and hind ;
each limb being composed of three segments.
a, The Fore-limb presents the following divisions :
i, Arm.
ii. Forearm.
iii, Hand, with four digits, corresponding to the four
fingers of man; the thumb being very small and
inconspicuous. In the male frog there is a thicken-
ing along the inner edge of the first digit, specially
developed at the breeding season.
b. The Hind-limb is much longer than the fore limb, and
divided into the following parts:
i. Thigh.
ii. Leg.
iii. Foot, with five toes webbed together. The shortest
toe corresponds to the big toe of man, and the
longest to his fourth toe.
6. External apertures: or openings on the surface of the body.
a. Median apertures.
i. The Mouth is a wide horizontal slit.
ii. The Cloacal aperture is a small hole at the posterior
end of the body, between the legs: it lies slightly
on the dorsal surface, just behind the bony pro-
jection formed by the posterior end of the urostyle.
 THE BUCCAL CAVITY. 15

b. Paired apertures.
i. The Nostrils or anterior nares are two small open-
ings on the dorsal surface of the head, close to its
anterior end.
B. The Buccal Cavity.
Open the mouth to tts full extent : note the wide buccal or mouth
cavity, of which the hinder part or pharynx is continued back into
the esophagus. Note also the following structures:
1. On the Roof of the Mouth.
a. The Teeth.
i. The maxillary teeth are a row of fine teeth,
attached round the edge of the upper jaw.
ii. The vomerine teeth are two small patches of sharp
teeth in the fore part of the roof of the mouth
and near the middle line.
b. The posterior nares are two small holes lying to the outer
sides of and slightly in front of the two patches of
vomerine teeth.
Pass bristles through the nostrils, and see that they
come out through the posterior nares into the buccal cavity.
c. The Eustachian tubes or recesses are a pair of much
larger holes, at the sides of the posterior part of the
buccal cavity. Each hole opens into a slightly dilated
chamber—the tympanic cavity—which is closed
externally by the tympanic membrane already seen
on the surface of the head.
Perforate the tympanic membrane on one side with a
needle, and pass a bristle or seeker through the hole and
down the K'ustachian tube into the mouth.
d. Two rounded prominences at the sides of the roof of
the mouth are caused by the eyeballs.
Press down one of the eyes with your finger, and note
that it can be made to project very considerably into the
buccal cavity.
2. On the Floor of the Mouth.
a, The lower jaw, which is devoid of teeth, forms a bony
margin to the floor of the mouth: the rest of the floor
is soft and fleshy, but is slightly stiffened by a car-
tilaginous plate—the body of the hyoid.
16 GENERAL ANATOMY OF THE FROG.

b. The tongue, which is thin and fleshy, is attached to

the front part of the floor of the mouth, and has its
free bilobed end turned backwards towards the throat.
Turn the tongue forwards with the forceps.
c. The glottis, or aperture of the larynx, is a longitudinal
slit in the floor of the posterior part of the mouth
stiffened laterally by the arytenoid cartilages.
Pass bristles through the glottis into the lungs. If
: any difficulty 1s experienced in finding the glottis
snip through the angles of the mouth with scissors,
so as to allow the mouth to be opened more widely.

C. The Abdominal Viscera.

Lay the frog on its back under water, and fasten it down to the
dissecting board by pins through the limbs. Cut through the skin,
along the middle line, the whole length of the ventral surface.
Separate the skin from the underlying parts, noticing its very loose
attachment to these parts, and the large space—lymph cavity—
beneath it. Turn the flaps of skin outwards and pin them back.
Notice -—
a. The muscles of the body wall.
b. The pectoral or shoulder girdle: a bony arch running
across the body, opposite the fore-limbs.
Pinch up with forceps the muscular body wall, and cut through
ut into the body cavity with scissors a little to one side of the
median plane, being careful not to injure the anterior abdominal
ven which runs along the inner surface of the body wall in the
middle line.
Continue the cut backwards, to the hinder end of the body, and
forwards to the jaw, cutting through the pectoral girdle with strong
scissors, and taking care not to injure the parts beneath.
Note on the inner surface of the larger flap the anterior abdominal
ven, and carefully dissect this from the flap. Pull the two flaps
apart, cutting through them transversely at their posterior ends to
facilitate the process, and turn them back so as to display the viscera.
Inflate the lungs with a blow-pipe through the glottis, and
inflate the bladder through the cloacal aperture.
Note and draw the general arrangement of the viscera, showing
the following structures :
 ABDOMINAL VISCERA. bi

1. The heart, enclosed in the pericardium, is situated in the

middle line in front, and in the natural condition of the parts
is covered by the pectoral girdle and the sternum.
2. The liver, is a large reddish-brown bilobed organ, behind
and at the sides of the heart.
3. The lungs are two thin-walled elastic sacs at the sides of the
heart: they lie dorsal to the liver, and are often hidden by it.
Note the bristles already passed into the lunys through the glottis.
4, The corpora adiposa, or fat bodies, are two bright yellow
tufts of flattened processes attached to the dorsal wall of the
body cavity ; they vary much in size, and usually come to the
surface just behind the liver.
5. The small intestine is a light coloured convoluted tube;
in the middle line behind is the much wider large intestine.
6. The bladder is a thin-walled bilobed sac at the posterior
end of the body cavity.

s\ S55. ts, Sa:

\ SSSJ aa,
NSS
Z a

nS
=a
Wea
WU
YMA

lgoaam
PT)
=
i,Y,tka
Sa
i”

Fig. 2. A diagrammatic transverse section across the posterior part of

the body of a female frog.
a, urostyle: 6, muscles of body wall: d, large intestine: d.a, dorsal
aorta : 7, ilium: /, lymph space between the skin and the muscular body
wall: 7, spinal nerves: o, kidney: ov, oviduct: p, peritoneum: s, skin:
t, fold of skin at groin: w, ureter: v, posterior vena cava.

7. In the female froy note, in addition to the above parts,

a. The ovaries: two large bodies of irregular shape, each
consisting of a mass of spherical black and white
egos, like small shot.
18 GENERAL ANATOMY OF THE FROG.

b. The oviducts: two long, very much convoluted tubes

with thick white walls, lying at the sides of the body
cavity.
8. In the male frog note,
a. The testes: a pair of ovoid bodies of a pale yellow
colour, attached to the dorsal wall of the body cavity.

D. The Peritoneum.
Notice the thin pigmented membrane—the peritoneum—
which lines the body cavity. Trace this to the mid-dorsal line
where it is reflected downwards as a double layer—the mesen-
tery—which embraces at its edge the alimentary canal, and
binds its several coils together. (See Fig. 2.)
Notice also that all the abdominal viscera are really outside
the peritoneum, which forms a closed sac into which the viscera
are as it were pushed from without.
E. The Digestive Organs.

Fig. 3. General view of the viscera of the male frog, from the right side.
a, stomach: b, bladder: c, small intestine : c/, cloacal aperture :
d, large intestine : e, liver : f, bile duct : g, gall bladder : h, spleen : 7, lung :
k, larynx : J, fat body: m, testis: n, ureter: 0, kidney: p, pancreas:
r, pelvic symphysis: 8, cerebral hemisphere : sp, spinal cord : ¢, tongue :
u, auricle: wr, urostyle: v, ventricle : v.s, vesicula seminalis: w, optic
lobe : x, cerebellum : y, Eustachian recess : z, nasal sac.
 DIGESTIVE ORGANS. 19

Turn the liver forwards, and note the stomach lying beneath its
left lobe. Pass the handle of a seeker through the mouth and
down the esophagus into the stomach. *
[Lf the specimen be a female, remove the ovaries and oviducts
completely, taking care not to damage the alimentary canal.|
1. The Alimentary Canal.
a. The esophagus is a short. wide tube leading from the
buccal cavity to the stomach.
b. The stomach is a wide tubular sac about an inch and a
half in length: it is narrower behind, and separated
from the duodenum by a distinct pyloric constriction.
Cut open the stomach longitudinally along its left side, and
wash out ws contents: note the hindle of the seeker already in-
serted through the mouth ; also the longitudinal folds of the
mucous membrane lining the stomach, which increase the extent of
ats surface.
c. The duodenum is the first part of the intestine, rather
more than an inch in length: beyond the pylorus it is
bent back so as to lie parallel to the stomach. At its
further end it is continuous with the small intestine.
d. The small intestine is a slender convoluted tube about
four and a half inches long, opening at its distal end
by a small orifice into the large intestine.
e. The large intestine is a short straight tube about an
inch and a quarter long: it is very much wider than
the small intestine, and opens behind to the exterior
at the cloacal aperture.
f, The cloaca in the frog is the last half inch of the large
intestine, into which open the renal and genital ducts
as well as the bladder: it will be described more
fully when considering the urinary and reproductive
organs (see chapter VIII).
2. The Liver.
The liver is a large reddish-brown organ, divided into right
and left lobes, connected together by a narrow bridge of liver-
tissue. Of the two lobes the left one is much the larger, and is
again subdivided into two.
20 GENERAL ANATOMY OF THE FROG.

a. The gall-bladder is a small spherical sac lying between

the right and left lobes of the liver.
b. The bile duct is a slender tube leading from the liver
and gall-bladder to the duodenum, into which it opens
about half an inch beyond the pylorus, and on the
inner or concave side of the loop formed by the
duodenum and stomach. The distal half of the bile
duct traverses the pancreas: it has rather thick white
walls and is easy to see; the upper half is more
slender and more difficult to trace.
To trace the bile duct turn the liver forwards so that the point
ofattachment of the gall-bladder is clearly seen ; and slightly stretch
the duodenum by a pin passed through the pylorus. Determine the
position of the two ends of the bile duct from the description given
above, and dissect with a scalpel along the line thus indicated.
To see the opening of the bile duct, slit up the first three quarters
of an inch of the duodenum along its convex border and wash out
its contents: squeeze the gall-bladder so as to drive the bile along
the duct into the duodenum : note the point at which it enters, and
insert a bristle through the opening into the duct. Notice also
the strong wavy transverse folds of the mucous membrane of the
duodenum.
3. The Pancreas.
The pancreas is a whitish irregularly lobed mass lying in the
loop between the stomach and duodenum, and best seen by
turning the whole loop forwards. The pancreatic ducts are
numerous and open into the bile duct, which passes through
the pancreas to reach the duodenum.
Cut through the mesentery along its attachment to the intestine -
uncoil the intestine, leaving it attached at both ends, and spread it
out on your dissecting board: measure the lengths of the several
portions and draw them to scale.
F, Other Abdominal Viscera.
1. The Kidneys are two flat elongated oval bodies of a red
colour attached to the dorsal body wall, close to the middle line,
one on each side of the back-bone or vertebral column. They
lie in the large lymph space behind the peritoneum. (See Fig. 2,
p. 47.) |
 ABDOMINAL VISCERA. 21
a. The ureters, or ducts of the kidneys, area pair of white
tubes arising from the outer edges of the kidneys at
about a quarter of their length from their hinder ends,
and running back to open into the dorsal wall of the
cloaca, opposite the opening of the bladder.
In the male frog a pouch-like dilatation, the
vesicula seminalis, is present on the outer side of
each ureter, close to its opening into the cloaca.
3. The Spleen is a small round dark-red body lying in the
mesentery, opposite the commencement of the large intestine.
 CHAPTER IL.
THE VASCULAR SYSTEM OF THE FROG.

The vascular system is a closed system of tubes or vessels

filled with blood, and ramifying through all parts of the body:
its main parts are: (1) the heart, which by its contractions is
continually driving the blood round and round the system of
vessels: (2) the arteries, which are the vessels taking the
blood from the heart to all parts of the body: (3) the veins,
which carry the blood from those parts back to the heart : and
(4) the capillaries, a system of very small vessels connecting
the arteries and veins together.
A. The Heart.
Pin down the frog on its back under water and open the body
cavity as before, taking special care to preserve the anterior abdomi-
nal ven. Dissect this vein carefully from the body wall. In
freeing the pectoral girdle from the underlying muscles take care
not to injure the neighbouring bloodvessels.
Open the pericardial cavity and dissect the pericardium from
the heart and the roots of the great vessels, examine and draw the
heart in situ, showing its several divisions.
1. The divisions of the heart.
i. The auricles form the anterior and dorsal division
of the heart: they are thin-walled and appear
dark in colour owing to the blood being seen
through their walls. On close examination the
division into right and left auricles can be seen.
ii. The ventricle is posterior to the auricles: it is paler
in colour owing to the greater thickness of its
walls; and is conical in shape, with the apex
pointing backwards.
iii, The truncus arteriosus is a cylindrical body arising
from the right anterior border of the ventricle,
and running obliquely forwards across the auricles.
 THE VEINS. 23

Lift up the ventricle and turn it forwards so as to expose the

SUNUS VENOSUS.
iv. The sinus venosus is a thin-walled sac, lying dorsal
to the ventricle and behind the auricles iG
receives the three large vene cave.
2. The pulsation of the heart.
a. Note that the contractions of the heart continue some
time after the frog has been killed, or even after the
heart is completely removed from the body.
b. Note the character of the heart’s pulsations : a regularly
alternating series of contractions and dilatations.
¢. Note further that in each contraction or systole of the
heart all four divisions of the heart contract, but not
simultaneously. The sinus venosus ‘contracts first,
then the two auricles, then the ventricle, and finally
the truncus arteriosus.

B. The Veins.
my, by. sy. jy.

Fig 4. Diagrammatic figure of the venous system of the frog, from

the
right side.
a, Stomach : a.v, anterior abdominal vein : }, bladder: b.v, brachial
vein : c.l, cloacal aperture : ¢c.v, cardiac vein : d, large intestine
: e, liver:
e.v, external jugular vein: f.v, femoral vein: g, gall bladder : h, spleen :
2.€, posterior vena cava: 7.0, innominate vein : j.v, internal jugular
vein
L.p, left pelvic vein : m.v, musculo-cutaneous vein: o, kidney: p.v, hepatic :
portal vein: 7.p, right pelvic vein : r.v, right renal portal vein : s,
venosus : 8.c, sciatic vein: s.v, subclavian vein : ¢, tongue : ¢.a,
sinus
truncus
arteriosus : uw, right auricle : v, ventricle : v.v, Vesical veins.
24 THE VASCULAR SYSTEM OF THE FROG.

The veins should be dissected before the arteries, because, as

a rule, they lie nearer the surface and are therefore met with
first. The veins are further distinguished from the arteries by
their larger size, thinner walls, and darker colour, due to the
blood being seen through their walls.
Dissect from the ventral surface. In cleaning a vein take hold
with the forceps, not of the vein itself but of the tissue surrounding
it ; and take especial care not to prick the vein, as by doing so you
allow the blood to escape and obscure the dissection, and also render
the vein itself difficult to see owing to the loss of colour. Always
dissect along and not across a bloodvessel, and pin out the parts so
as to stretch it slightly.
I. Veins opening into the Sinus Venosus.
a. The right anterior vena cava is a large vein opening
into the right side of the sinus venosus, and returning
to it the blood from the right side of the head and
body, and from the right fore-limb. It is formed by
the union of three veins.
1. The external jugular vein is formed by
i. The lingual vein, from the floor of the mouth
and the tongue.
ii. The mandibular vein, from the margin of the
lower jaw.
In close connexion with the ventral surface of
the external jugular vein is a small round
vascular body, the thyroid gland.
2. The innominate vein is formed by
i. The internal jugular vein, returning blood
from the interior of the skull, which it leaves
by an aperture at the posterior border of
the orbit.
il. The subscapular vein, a small vein from the
back of the arm and shoulder.
3. The subclavian vein, the largest of the three,
is formed by
i. The brachial vein, from the fore-limb.
ii. The musculo-cutaneous vein: a very large
vein returning blood from the skin and
 THE VEINS. 25

muscles of the side and back of the body,

and of the head as far forwards as the nose.
b. The left anterior vena cava corresponds in its course
and branches to the right one.
c. The posterior vena cava is a median vein which, com-
mencing between the kidneys, runs forward to open
into the posterior end of the sinus venosus. It returns
to the heart the blood from the liver and from the
kidneys, and indirectly from other viscera and from
the hind limbs. It receives the following veins :—
i. The right and left hepatic veins, from the liver:
these open into the posterior vena cava just before
it joins the sinus venosus.
ii. The renal veins, from the kidneys: of these there
are four or five on each side which open into, or
rather form by their union, the posterior vena
cava. The most anterior of these receive the
veins from the fat bodies.
; iii. The ovarian veins (in the female), or spermatic
veins (in the male); returning blood from the
ovaries or testes. They are usually four or five
in number on each side, and open into the
posterior vena cava between the renal veins.

II. Vein opening into the Left Auricle.

a. The pulmonary vein is formed by the union of the
right and left pulmonary veins, returning to the heart
the blood from the right and left lungs respectively.
Each pulmonary vein runs along the inner side of its
lung.

III. The Portal Systems.

A portal vein is one which, returning blood from the capil-
laries of some part, breaks up before reaching the heart into a
second set of capillaries within some other organ; these again
unite to form a vein which carries the blood to the heart. In
the frog there are two portal systems, one supplying the kidneys,
and the other the liver.
26 THE VASCULAR SYSTEM OF THE FROG.

a. The renal portal system.

Trace back the anterior abdominal vein to the hinder end of the
body, where it will be seen to be formed by the union of the two
pelvic veins. /ollow back the pelvic vein of one side to the base
of the hind limb ; here tt will be seen to be one of two branches into
which the femoral vein, the large vein returning blood from the
hind limb, divides. The other branch of the femoral vein is the
renal portal vein, which is to be followed to the outer side of the
kidney.
1. The right renal portal vein is the dorsal branch of
the right femoral vein: it runs forwards along the
outer side of the kidney and ends in numerous
branches in its substance. It receives the follow-
ing branches.
i. The right sciatic vein, from the muscles and
skin of the back of the thigh, joins the
renal portal vein close to its commencement,
before it reaches the kidney.
’ ii. The right dorso-lumbar veins are small veins
from the dorsal wall of the body, and, in the
female, from the oviduct: they join the
renal portal vein opposite the kidney.
2. The left renal portal vein corresponds in its course
and branches to the right vein.
b. The hepatic portal system.
This is formed partly by the anterior abdominal vein, which
brings to the liver blood from the hind limbs; and partly by
veins returning blood from the alimentary canal.
1. The anterior abdominal vein is a median vein
formed by the union of the two pelvic veins, the
ventral branches of the femoral veins. . It runs
forwards along the middle line of the ventral
body-wall to the level of the liver, where it leaves
the body-wall and divides into right and _ left
branches, which enter the right and left lobes of
the liver respectively. During its course it
receives the following veins.
i. Vesical veins, from the bladder
ii. Parietal veins, from the ventral body-wall.
 THE ARTERIES. 27

iii. A cardiac vein, from a network of vessels on

the truncus arteriosus.

2. The hepatic portal vein is a wide vein which runs

in the mesentery and joins the anterior abdo-
minal vein at its point of division into right
and left branches ; giving off, before doing so,
a branch to the left lobe of the liver. It
carries to the liver the blood from the walls of
the alimentary canal, and is formed by the union
of the following veins.
i. The gastric vein, from the stomach.
ii. Intestinal veins, from the whole length of the
intestine, both small and large.
iii. The splenic vein, from the spleen: this usually
joins one of the intestinal veins.

C. The Arteries. .

= <a i Ch \
dg

eoltie.. 5a Diagrammatic figure of the arterial system of

the male frog,
from the right side.
a, stomach : 6, nostril: c, small intestine: c.a, carotid
carotid gland: ¢.m, cceliaco-mesenteric artery: c.m, cutaneoartery: c.g,
d, large intestine: d.a, dorsal aorta: J, femur: h, spleen: us artery:
artery: 7, right lung: /.a lingual artery: m, testis: o, h.a, hepatic
occipito-vertebral artery : p.a, pulmonary artery: 7, kidney: o.a,
pelvic girdle ;
8, sternum :.s.a, subclavian artery: S.c, sciatic artery:
i.a, truncus arteriosus : w.a, urinogenital arteries : v, ventricl 7¢, tongue:
arch : 2, systemic arch : 3, pulmo-cutaneous arch. e: 1, carotid
28 THE VASCULAR SYSTEM OF THE FROG.

Dissect as for the veins. Pass a roll of paper down the

cesophagus, so as to distend it and stretch the aortic arches.
Clean carefully the aortic arches, commencing at the truncus
arteriosus ; and follow the several arteries to their distribution,
removing the veins and other structures which overlie them. Note
the dwision of the truncus arteriosus in front into right and left
branches, each of which again divides into three aortic arches—the
carotid arch, the systemic arch, and the pulmo-cutaneous arch.
1, The Carotid Arch is the most anterior of the three arches :
it runs round the side of the cesophagus, and is connected dorsally
with the second or systemic arch: its chief branches are as
follows.
1. The lingual artery is a small artery supplying the
tongue. Immediately beyond the origin of the lingual
artery the carotid arch presents a small spongy
swelling, the carotid gland.
2. The carotid artery runs round the side of the cesopha-
gus to its dorsal surface: it is connected with the
systemic arch by a short branch, the ductus Botalli,
which in the adult frog is usually impervious ; and
then turns forwards beneath the base of the skull,
dividing in front into the two following vessels :
i. The external carotid artery, supplying the roof
and sides of the buccal cavity, and the orbit.
ii. The internal carotid artery, which enters the skull
and supplies the brain.
II. The Systemic Arch, the middle arch of the three, runs
somewhat obliquely round the cesophagus to the dorsal surface,
and unites with its fellow of the opposite side about the level of
the anterior ends. of the kidneys to form the dorsal aorta: near
the level of the posterior ends of the kidneys the aorta divides
into the two iliac arteries. The branches of the systemic arch
are as follows:
a. Branches given off before the union of the two arches.
1. The laryngeal artery is a small branch arising from the
inner side of the systemic arch near its origin from
the truncus arteriosus, and supplying the larynx.
2. The esophageal arteries are one or two branches arising
from the upper part of the arch and entering the
dorsal wall of the cesophagus.
 THE ARTERIES. 29

3. The occipito-vertebral artery is a short branch arising

from the dorsal part of the arch: it runs upwards
immediately in front of the transverse process of the
second vertebra, and divides into two :—
i. The occipital artery: which runs forwards, supply-
ing the side of the head and jaws.
ii, The vertebral artery: a large artery which runs
back alongside of and above the vertebral column,
and gives branches to the muscles of the body-
wall and to the spinal cord.
4. The subclavian artery: arises from the arch immedi-
ately behind the occipito-vertebral artery, and runs
outwards, supplying the shoulder and fore-limb.
b. Branches given off after the union of the two arches to
form the dorsal aorta.
1. The ceeliaco-mesenteric artery is a large median artery
arising immediately beyond the point of union of the
two arches, or sometimes from the left arch just
before the union, and supplying the stomach and
intestines. Its branches are as follows:
i. The celiac artery: which divides into
a. The gastric artery, supplying the stomach.
(. The hepatic artery, supplying the liver and
gall-bladder.
ii The mesenteric artery: which divides into
a. The anterior mesenteric artery, supplying
the proximal part of the intestine.
(. The posterior mesenteric artery, supplying
the distal part of the intestine.
y. The splenic artery, supplying the spleen.
2. The urinogenital arteries are four to six small arteries
which arise from the ventral surface of the aorta
between the kidneys, and immediately divide into
right and left branches, supplying the kidneys, the
reproductive organs and ducts, and the fat bodies.
3. The lumbar arteries are small paired lateral branches
supplying the body-walls.
30 THE VASCULAR SYSTEM OF THE FROG.

4. The hemorrhoidal artery is a small median artery

arising from the hinder end of the aorta, and supplying
the large intestine.
c. Branches formed by the division of the aorta.
1. The iliac arteries are the two large arteries formed by
the division of the aorta, and supplying the hind-limbs.
Each gives off a hypogastric artery, which supplies
the bladder, giving epigastric branches to the ventral
body-wall, and then continues as the sciatic artery
down the leg, giving off branches to the muscles and
skin of the thigh, and dividing at the knee into
peroneal and tibial arteries supplying the leg and
foot.
III. The Pulmo-cutaneous Arch is the hindmost of the three
aortic arches: it divides about the level of the carotid gland
into the following branches.
1. The cutaneous artery is a large artery which at first
runs forwards and upwards and then turns backwards,
supplying the skin of the back along the whole length
of the body, and sending smaller branches to the sides
of the head and to the skin of the ventral surface.
2. The pulmonary artery runs with somewhat sinuous
course along the outer side of the whole length of
the lung, giving off branches into its substance.

D. The Structure of the Heart.

Having completed the dissection of the blood-vessels, cut them
across, about half an inch from the heart ; remove the heart com-
pletely, and dissect it carefully under water. It is well to cut the
vessels of unequal lengths on the two sides, as this will facilitate the
recognition of the sides of the heart during the dissection.
Place the heart at first with the dorsal surface upwards.
1. The Sinus Venosus (Fig. 4, p. 23) is a thin-walled sac on
the dorsal surface of the heart ; it is triangular in shape, with
the apex directed backwards. Into its anterior angles the right
and left anterior venz cavee open, and into its posterior angle
or apex the posterior vena Cava.
Cut away with scissors the dorsal wall of the sinus venosus so as
to expose its cavity: wash out any contained blood.
 THE HEART. fl

The sinu-auricular aperture (Fig. 6, S.V.) leading from the

sinous venosus to the right auricle, is a transversely oval opening,
guarded by imperfect anterior and posterior valves, in the
ventral wall of the sinus venosus, close to its anterior end, and
very nearly in the median plane.
eSp
LLL, am

Fig. 6. The frog’s heart seen from the ventral surface, and dissected
so as to show its structure. The ventral walls of the truncus arteriosus,
and of the auricles and ventricle have been removed. (From a drawing
by Dr. Hurst.)
A, auriculo-ventricular aperture and one of its valves: B, aperture
leading from ventricle to truncus arteriosus, with one of its valves :
C, left carotid arch: C’, style passed down right carotid arch into the
truncus arteriosus: LA, left auricle: P, left pulmo-cutaneous arch:
P’P’. style, passed down right pulmo-cutaneous arch into the truncus
arteriosus : PV, opening of pulmonary vein into left auricle: RA, right
auricle : S, left systemic arch: sg’, style passed down right systemic
arch into the truncus arteriosus: SV, opening from sinus venosus into
right auricle : Vv, ventricle.
32 THE VASCULAR SYSTEM OF THE FROG.

2. The Auricles. Zurn the heart over, with its ventral surface
upwards. Cut away the ventral wall of both auricles wrth fine
scissors, tuking care not to damage the truncus arteriosus which
lies across the right auricle. Wash out the blood from the auricles.
b. The right auricle (Fig. 6 RA) is the larger of the two.
It has thin walls, thickened by muscular strands
which form interlacing reticular ridges on its inner
surface. In the dorsal wall of the auricle, very near
the median plane of the heart, is the aperture from
the sinus venosus already described (Fig. 6 SY).
b. The left auricle (Fig. 6 LA) is smaller, sometimes
much smaller, than the right auricle, which it resem-
bles in the structure of its walls. In its dorsal wall,
very close to the sinu-auricular aperture, is the opening
of the pulmonary vein (Fig. 6 PY).
c. The interauricular septum is the thin partition between
the right and left auricles. It is much thinner than
the walls of the auricles, and is placed somewhat
obliquely, the left auricle lying rather more dorsally
than the right. The septum ends with a free posterior
edge, opposite the auriculo-ventricular aperture.
Cut away with scissors the ventral wall of the ventricle, taking
care not to damage the truncus arteriosus.
3. The Ventricle (Fig. 6 V) is conical in shape with the
apex backwards, and has a small central cavity, with thick
spongy walls. The spongy character is due to great develop-
ment of a reticulum of interlacing muscular strands similar to
those of the auricles: the true outer wall of the ventricle is no
thicker than that of the auricles, and the meshes of the sponge-
work are really part of the cavity of the ventricle, and are filled
with blood.
The auriculo-ventricular aperture lies at the base of the
ventricle, and rather to the left side. It is guarded by valves
(Fig. 6, A) which hang into the ventricle, and are tied down at
their edges by fine tendinous threads ; and it is divided by the
free lower edge of the interauricular septum into right and
left divisions, admitting blood from the right and left auricles
respectively.
Cut away carefully, with fine scissors, the ventral wall of the
truncus arteriosus so as to expose its cavity and the contained valves.
 THE HEART. 33

4, The Truncus Arteriosus consists of two parts ; a proximal

part or pylangium, which is a single vessel arising from the
ventricle ; and a distal part or synangium, which consists of the
basal parts of the aortic arches closely united together.
a. The pylangium (Fig. 6) is a short tube arising from
the right hand ventral corner of the anterior end of
the ventricle: it has thick muscular walls and is
widest about the middle of its length.
The opening from the ventricle to the pylangium
(Fig. 6 B) is guarded by three semilunar pocket valves.
The opening from the pylangium to the synangium
is also guarded by three semilunar valves which are
of very unequal size, a large right one, a small left
one, and a still smaller dorsal valve.
The spiral valve is a longitudinal ridge, projecting
into the cavity of the pylangium: it commences at
the left side of the ventricular aperture and runs
forwards somewhat spirally along the dorsal wall of
the pylangium to its anterior end, where it fuses
with the large right valve of the three between the
pylangium and the synangium. The ventral edge of
the spiral valve is free and rounded, and the valve ig
much wider at its anterior than at its posterior end.
b. The synangium is the distal part of the truncus
arteriosus. In its dorsal wall, immediately beyond
the valves separating it from the pylangium, is an
aperture (Fig. 6 P’) leading to the right and left
pulmo-cutaneous arches (Fig. 6 P,P’). Beyond this
the synangium contains a wide cavity continued right
and left into the two systemic arches—S,S’. The
cavity is partially divided by a vertical tongue-like
projection from its dorsal wall: on the ventral
surface of this tongue are two small openings, very
close together, which lead into the right and left
carotid arches, C,C’.
Cut across the aortic arches, just beyond the division of the
truncus into right and left branches, and note that though each
branch is apparently a single vessel its cavity is really divided
unto three vessels corresponding to the three aortic arches. Pass
bristles down these aortic arches, and note the points at which they
severally open into the truncus arteriosus.
D
34 THE VASCULAR SYSTEM OF THE FROG.

E. The Lymphatic System.

The lymphatic system forms an accessory part of the vascular
system. Its main divisions are as follows :
1. The lymphatic vessels are a series of thin-walled tubes,
very variable in diameter and irregular in shape, which
traverse all the parts and organs of the body and are
in free communication with the veins. They are of
small size, and can only be recognised with the
microscope.
2. The lymph sacs are large irregular spaces communi-
cating with the lymphatic vessels. The most
important are the following :
a. The subcutaneous lymph sacs are the large
cavities between the skin and the muscles,
which have already been seen when removing
the skin. They are separated from one
another by narrow septa of connective tissue,
which bind the skin to the underlying body-
wall.
b. The abdominal lymph sacs are the large
spaces along the dorsal surface of the body-
cavity, ventral to the kidneys, and between
the peritoneum and the body-walls. (See
Fig. 2, p. 17.) The body cavity itself also
communicates with the lymphatic system
through small openings or stomata in the
peritoneum.
3. The lymph hearts are two pairs of small globular con-
tractile sacs placed at points where the lymphatic
vessels communicate with the veins.
a. The anterior lymph hearts lie immediately
behind the transverse processes of the third
vertebra, and beneath the shoulder girdle:
they open into the subscapular veins.
b. The posterior lymph hearts lie at the sides of
the urostyle, close to its hinder end. They
communicate by short vessels with the
femoral veins. Their pulsations can easily
be seen in a living frog.
 BLOOD. oo

F. Microscopic Examination of Blood.

I. Frog’s Blood.
1. Normal. |
Place on a slide a small drop of blood from the heart of a frog;
dilute vt with a drop of normal salt solution (0°75 per cent) : put
on a thin cover-glass, and run a ring of oil round the edge to pre-
vent evaporation : examine with the high power.
Blood consists of a colourless fluid, the liquor sanguinis
or plasma, in which float the blood corpuscles. These
corpuscles are of two kinds.
i. Red corpuscles. These are very numerous, pale
red or yellowish red in colour, and of a flattened
oval shape, with rounded edges and a central
bulging, the nucleus. The flattened shape is best
seen when a corpuscle turns edgeways. They
measure 0°0235 mm. in length by 0°0145 mm. in
width ; or about 3759 x rs'o0 Of an inch.
ii. White corpuscles. These are much fewer in number
and of smaller size: they are colourless, granular,
subspherical in shape, and exhibit “amoeboid”
movements. Sketch one half-a-dozen times at
intervals of half a minute.
2. Action of acetic acid on blood.
Place a fresh drop of blood on a clean slide: add a drop of
acetic acid. cover, and examine with the high power. note the
changes produced.
i, Red corpuscles: the nuclei become much more
apparent than before, and the red colour dis-
appears.
ii, White corpuscles: become clearer, and show nuclei,
sometimes more than one in a single corpuscle.
II. Human Blood.
1. Normal.
Prick the tip of your finger, and place a small drop of the
blood on a slide: add a drop of normal salt solution, cover, and
examine as before. Note the following points :
i, Red corpuscles. These, which are much smaller
36 THE VASCULAR SYSTEM OF THE FROG.

than in frog’s blood, are in the form of circular

biconcave discs with rounded edges, but no nuclei.
They have a tendency to run together into
rouleaux, like piles of coins. Their average
diameter is 0°008 mm., or about 355, of an inch.
ii. White corpuscles. These are very similar to those
of the frog: they are slightly larger than the
red corpuscles, averaging about 0°01 mm., or
zso0 Of an inch in diameter: their amceboid
movements are not well seen unless the slide is
warmed.
2. Action of acetic acid.
Treat with acetic acid as before: note that, unlike the frog’s
blood, no nuclei are visible in the red corpuscles.

G. Circulation of the Blood in the Web of a Frog’s Foot.

The web uniting the toes of the frog’s foot is so thin and
transparent, that with the microscope the blood in it can readily
be seen coursing along the capillaries.
Examine a frog prepared to show the circulation in the web of
the foot. Note the following pornts :
1. With a low power.
a, The irregularly branched pigment cells to which the
colour of the frog’s skin is due.
b. The fine meshwork of bloodvessels along which the
blood can be seen flowing. These bloodvessels are
of three kinds. ,
i. The arteries, carrying blood to the web, are dis-
tinguished by the fact that when they divide,
the direction of flow of the blood is from the
larger trunk to its branches.
ii. The capillaries form a close network of very small,
very thin-walled vessels, along which the blood
flows from the arteries to the veins.
iii. The veins, carrying the blood away from the web
back towards the heart, are distinguished from
the arteries by the fact that the blood in them
flows from smaller to larger vessels.
 CIRCULATION OF BLOOD. ot

2. With a high power: note the following points:

a. The walls of the arteries and veins are much thicker than
those of the capillaries, which latter are often difficult
to see.
b. The white corpuscles have a marked tendency to creep
along the sides of the vessels, while the red corpuscles
rush far more rapidly along the middle of the stream:
this is seen best in the small arteries.
c. The variations in calibre of the small arteries and
capillaries: whilst under observation an artery or
capillary may be seen to change its size to a consider-
able extent.
d. The indefinite character of the capillary circulation.
Owing to changes of size in adjacent vessels, the
direction of flow of the blood in a given capillary may
become reversed.
e. The elasticity of the red corpuscles: seen best when
they are turning the corners of the capillary network.
f. The tendency of the white corpuscles to migrate through
the walls of the capillaries into the tissues outside.
This is much increased by the application of some
irritant substance, as a drop of weak acid, to the web.
38

CHAPTER III.

ELEMENTARY HISTOLOGY.

When examined under the microscope, all the different tissues

and organs of the body are found to consist of elementary bodies
called cells and of an intercellular substance, connecting the
several cells together ; in much the same way as a wall is built
of bricks cemented together with mortar. These cells, of which a
white blood corpuscle is a typical example, vary much in shape,
size, and structure in different tissues, but are to be considered as
fundamentally equivalent to one another. The intercellular
substance varies very much in quantity; it may be almost
absent, so that the several cells are practically in contact with
one another; or it may be so abundant as to separate them
widely : it is to be viewed as formed by the cells, and, therefore,
as secondary in importance to these.
When drawing histological preparations, it is well to look out
for, and draw, a few red blood corpuscles, to the same scale as the
rest of the drawing. The blood corpuscles form most useful
standards of measurement, as their dimensions are already known
(p. 35).
A. Epithelium.
Epithelium consists of cells placed side by side so as to form
layers, which form the surface covering, or epidermis, of the
body, and line the alimentary canal, the blood vessels, and the
various internal cavities of the body. At the external apertures
of the body, the epidermis is directly continuous with the
epithelial lining of the internal cavities.
The layers may be one or more cells in thickness; in the
former case the epithelium is said to be simple, in the latter
stratified.
Epithelium is of different kinds, according to the shape and
structure of its component cells.
I. Squamous Epithelium. In this the component cells are
flattened parallel to the surface they cover: if the epithelium is
stratified, the flattening is most marked in the superficial cells.
 EPITHELIUM. 39

a. Isolated Cells.
Serape gently the inside of your cheek with the handle of a
scalpel, and put the scrapings on a slide; cover, and examine with
(a high power ; draw, showing the following points:
i. The cells are large, flattened and scale-like in shape,
often slightly curled up at their edges.
ii, The nucleus is oval and granular, and lies near the
middle of the cell: it may be rendered more dis-
tinct by acetic acid or magenta.
b. Cells in situ: cast skin of newt.
Take a small piece of the prepared specimen, which has been
stained in hematoxylin, and then, after treatment with alcohol,
cleared with oil of cloves. Mount the specimen in balsam 3
cover, and examine with the high power.
i, The cells are flattened, and fitted together at their
edges, like a mosaic, to form a continuous layer.
Each cell has a large nucleus near its centre.
II. Columnar Epithelium. This consists of elongated rod-like
cells, placed vertically to the surface on which they rest. Ifthe
epithelium is stratified the columnar character is most marked
in the superficial cells.
a. Isolated cells: from the small intestine of the frog:
isolated by maceration for 24 hours in Ranvier’s
alcohol, and stained with picro-carmine.
Mount a drop of prepared specimen in glycerine + paint a
ring of cement round the cover-glass; and examine with the high
power.
1, The cells, which often remain side by side in little
groups, are columnar in shape, with nuclei near
their inner or deeper ends.
b. Cells in situ.
Take a prepared section of dog’s stomach which has been stained,
and then cleared in oil of cloves. Mount in balsam, and examine
with the high power.
i, The superficial layer consists of long narrow colum-
nar cells, packed together side by side, with nuclei
at their inner or deeper ends.
40 ELEMENTARY HISTOLOGY.

III. Ciliated Epithelium. In this the cells, which are usually

columnar, bear at their free ends tufts of exceedingly fine hair-
like processes—cilia—which, when living, exhibit active lashing
movements.
a. Isolated cells. From trachea of rabbit: isolated by
maceration for 24 hours in Ranvier’s alcohol; stained
with picro-carmine, and scraped into glycerine.
Mount a small drop of the prepared specimen tn glycerine:
paint a ring of cement round the cover-glass; examine with the
high power, and note :-—
i. The shape of the cells: their nuclei; and the tuft
of cilia at one end of each cell.
b. Cells in situ: ciliary movement.
Snip off a small piece of epithelium from the roof of the
mouth of a freshly killed frog, near the eyeball : mount in normal
salt solution, and adda small drop of gamboge water to render
the movements more clearly visible: examine with the high power ;
note :—
i. The currents due to the ciliary motion.
i. The movements of the individual cilia: best seen
when the specimen is beginning to die, and the
movements to slacken in speed.

IV. Stratified Epithelium. This is characterised by the

epithelium being several cells in thickness.
Take a prepared section of esophagus of rabbit, or of conjunctiva
of rabbit or pig, which has been hardened in chromic acid, stained,
and cleared in oil of cloves. Mount in balsam, examine with the
high power, and note -—
i. The stratification of the epithelium.
ii. The transition from the deeper spherical or columnar
cells to the superficial squamous cells.
B. Glands.
A gland consists essentially of a layer of epithelial cells
secreting some special fluid. The epithelial surface may be flat,
but is more usually folded or pitted, often in a very complicated
manner, so as to increase the extent of the secreting surface.
 GLANDS. 4]

I. Simple Glands. In simple glands the epithelial surface is

increased by simple pit-like depressions, whose mouths serve
to discharge the secretion on the free surface.
Take a prepared section of large intestine of rabbit which has
been hardened in chromic acid, stained, and cleared in oil of
cloves. Mownt in balsam, and examine first with the low power,
then with the high. Note the following points:
i. The glands are simple tubular depressions of the
surface.
ii. The glandular epithelium lining the pits is a single
layer of short columnar granular cells, many of
which are swollen to form goblet cells.

II. Compound Glands. In compound glands each depression

instead of being a simple pit is itself subdivided or branched,
often in a very complicated manner. There are two chief
varieties: (1) tubular glands, in which the several sub-
divisions are tubular, and of tolerably uniform diameter
throughout : and (2) racemose glands, in which the blind ends
of the pits are dilated into globular chambers or alveoli, to which
the special glandular epithelium is usually confined.
a. Compound tubular glands. Take a prepared section of
kidney of frog: mount in balsam, and examine with both
low and high powers.
i, The tubular gland-cavities are cut at various angles.
If cut transversely a tube appears as a circular
ring: if cut obliquely, as a more or less elongated
elliptical ring: if cut longitudinally, as two
parallel rows of epithelial cells.
ii. The gland cells form a single layer of cubical granular
cells, lining the tubes.
iii, The Malpighian bodies are spherical dilatations on
the tubes, into which project little knots of capillary
bloodvessels. Their structure is most readily made
out in specimens in which the bloodvessels have
been injected with a coloured substance to make
them more distinct.
42 ELEMENTARY HISTOLOGY.

NZ '
GM AWWEA! RBA
2
RA
NZ
\\ hy

e S|

| WAN Ht
: S|RS|
ey) =|
| A gS

(VRS
me |) SS } \\
PAS
a) |
VE
Hy =>

A\e.
Ae
ot

cc

co

GB

MC
P ML
Fig. 7. Section through mucous membrane of the cardiac end of a dog’s
stomach. ~ 140.
B, bloodvessel :CC, cubic or peptic cells: CM, columnar cells: CO,
ovoid cells : GB, fundus or bottom of gland cavity : GC, gland cavity cut
across: GM, mouth of gland: MC, circular muscle fibres :ML, longitudinal
muscle fibres : P, connective tissue layer between the mucous membrane
and the outer muscular walls of the stomach.
 GLANDS OF STOMACH. 43

III. Gastric Glands. The glands of the stomach are well

adapted for a more minute examination of the histology of
glands.
Examine again with a high power the section of the cardiac end
of the dog’s stomach already used for columnar ephithelium.
1. Characters of the glands. The gastric glands are good
examples of simple or slightly branched tubular glands.
They are deep, but very narrow, cylindrical pits,
imbedded vertically in the wall of the stomach, with
their open mouths discharging into its cavity. The
glands are lined by epithelial cells, and are set very
close together side by side. In the microscopical
sections, some of the glands may be seen cut along
their entire length ; but in most cases, owing to the
glands being not quite straight, or the plane of section
being oblique to the surface of the stomach, the tubes
will be cut more or less obliquely, or even trans-
versely.
2. Characters of the gland cells. There are three distinct
kinds of epithelial cells found at different parts of
the length of the gland.
i, Columnar cells, arranged in a somewhat radiate
manner round the mouths of the glands, and
extending a short way down the tubes.
ii. Cubic cells, or peptic cells, lining the deeper parts
of the glands and the greater part of their length:
these are cubical granular cells with centrally
placed nuclei.
iii, Ovoid cells: large oval cells with large nuclei:
these are less numerous than the other two forms,
and occur most abundantly a short way below the
mouths of the glands. They lie along the sides of
each gland, outside the cubical cells, and are said
to secrete the acid of the gastric juice.
C. Muscle.
In muscular tissue the component cells are much elongated
and, in the higher forms, very highly specialised. Muscular
tissue is of two kinds: (1) striated, or voluntary; of which all
44 ELEMENTARY HISTOLOGY.

muscles that are under the control of the will consist : and (2)
non-striated or involuntary forming those muscles over whose
contractions the will has no direct control. The muscular tissue
of the heart, which though involuntary is striated, forms the
chief exception to this rule,
I. Striated, or Voluntary Muscle.
a. Crab’s muscle. ease in glycerine a small piece of crab’s
muscle that has been hardened in alcohol ; cover, and
examine with both low and high powers « note :—
i. The elongated fibres of which the muscle consists.
Each fibre is a single cell, and is enclosed in a
delicate elastic sheath—the sarcolemma—which
will be visible in but few cases ; it is most readily
seen at places where the fibre has been torn
ACYOSS.
ii. The alternate light and dark bands with which the
muscle fibres are marked transversely, and from
which the name, striated muscle, is derived.
iii. The readiness with which the fibres split up longi-
tudinally into fibrils.
b. Frog’s muscle. Z'ease gently a piece of fresh frog’s muscle
in normal salt solution : cover, and examine with the
high power : note :—
i. The transverse striations.
ii. The sarcolemma: best seen by slightly crushing
the specimen.
ili. The nuclei in the fibres: seen on addition of acetic
acid.
II. Non-striated, or Involuntary Muscle.
Take a prepared specimen of frog’s bladder which has been
macerated in Ranvier’s alcohol for 24 hours ; pencilled with a fine
brush to remove the epithelium of the inner surface ; stained, and
cleared with oil of cloves. Mount in balsam, and examine with
low and high powers : note :—
i. The bands of muscular fibre. :
ii, The formation of each band by a number of
elongated, fusiform, nucleated muscle-cells.
iii. The absence of transverse striation in the muscle.
 MUSCLE : CONNECTIVE TISSUE. 45,

D. Connective Tissues.
Under the name ‘connective tissue” are included various
tissues whose functions are mainly passive, and which serve to
support, strengthen and bind together the various organs and
parts of the body. Histologically the connective tissues consist
of elements of four kinds, united together in very varying pro-
portions in different situations: (1) white fibrous tissue: (2)
yellow elastic tissue ; (3) connective tissue corpuscles, which are
comparatively slightly altered cells, usually branched ; and (4)
ground substance, or intercellular substance.
I. White Fibrous Tissue. This consists of a number of fine
transparent fibres of a more or less cylindrical shape, and
with a very characteristic wavy outline: between the
fibres are connective tissue cells, usually in small num-
bers. The fibres are arranged side by side in bundles,
and each fibre presents a number of longitudinal fibrillar
striations. The cellular origin of white fibrous tissue is
difficult to determine. The fibres are believed to be
formed by modification of the intercellular matrix rather
than from the bodies of the cells themselves.
a. Tendon of rat’s tail. Pull out a small piece of tendon
Jrom the tail of a rat: place it on a slide in a drop of
normal salt solution: spread it out with needles, cover
and examine with low and high powers ; note :—
i. The fibres, with wavy outlines.
ii, The fibrille, indicated by longitudinal wavy stria-
tions within the fibres.
Add a drop of acetic acid to the preparation: note that
iii, The fibres swell up and become transparent.
iv. Longitudinal rows of tendon cells, with nuclei,
become visible between the fibres.

II. Yellow Elastic Tissue. This consists of fine branching

homogeneous fibres, with great power of resisting
chemical reagents: the fibres are formed from an inter-
cellular matrix, and not from cells directly.
a. Ligamentum nuche of ox. Tease finely a small shred
in water ; examine with low and high powers ; note :—
46 ELEMENTARY HISTOLOGY.

i. The branching fibres, with very sharp outlines.

ii. The tendency of the branches to anastomose with
one another and so form networks.
iii. The tendency of the fibres and branches to curl up
at their broken ends.
Add a drop of acetic acid ; note that
iv. No alteration whatever is produced in the fibres.
v. No nuclei appear.
III. Areolar tissue. This is a meshwork composed of both
white fibrous and elastic tissues.
a. Subcutaneous tissue of mammal. Take a freshly killed
rat, and snip off a small piece of the loose fibrous tissue
which connects the skin with the subjacent parts: spread
it on a slide: add a drop of normal salt solution : cover,
and examine with low and high powers ; note :—
i. The meshwork, composed of white fibrous tissue
with wavy outlines, mingled with which are
branched elastic fibres.
Add acetic acid ; note that
ii. The white fibrous tissue swells up and becomes
transparent.
iii, The elastic tissue is unaltered.
iv. Connective tissue corpuscles, with nuclei, become
visible.
IV. Adipose tissue. This consists of a fine network of
vascnlar connective tissue, in the meshes of which are
fat cells, i.e, connective tissue corpuscles in which large
quantities of fatty or oily matter have accumulated.
a. Omentum of rabbit or kitten. Mount a small piece of
Sresh omentum in normal salt solution ; protect it from
the pressure of the cover glass ; examine with low and
high powers ; note :—
i. The vascular connective tissue meshwork, in which
lie the fat cells.
ii, The fat cells: large, spherical, or from mutual pres-
sure polyhedral, cells ;distended with fatty matter,
and with their nuclei near the surface.
 CARTILAGE. 47

b. Osmic Acid specimen.

Note the reduction of the osmic acid by the fat, which
becomes stained a dark brown or black colour.

E. Cartilage.
In cartilage or gristle the intercellular substance, which in
most other tissues is only present in small quantity, is greatly
increased so as to far exceed in bulk the cells which it connects
together. The intercellular substance forms a dense translucent
matrix resembling an extremely stiff jelly, in which are
imbedded the cartilage cells, either singly or in groups. In
young cartilage the intercellular substance is much less
abundant, and the cells consequently closer together than in
older or more mature specimens.
Cartilage when free from other tissue is called hyaline
cartilage from the clear or glassy appearance of the matrix, in
contra-distinction to fibro-cartilage, in which the matrix is fibrous
from admixture with white fibrous or elastic tissues.

I. Hyaline cartilage.
a. Cartilage of newt. Zake a small piece of cartilage from
the shoulder girdle of a newt: scrape away gently any
muscle or other tissue that may adhere to it ; mount in
normal salt solution, and examine with low and high
powers.
i, The intercellular matrix is either hyaline or faintly
granular.
ii, The cartilage cells are imbedded in the matrix;
each cell is nucleated, and occupies a cavity or
lacuna in the matrix. In places the cells are in
groups of twos or fours owing to recent division.
Wash the specimen thoroughly in water: stain with carmine,
and mount as a permanent preparation in glycerine; examine
with the high power, and note that
iii, The cell nuclei are stained deeply, and the matrix
very slightly: the layer of matrix immediately
surrounding each cell—the capsule—stains more
deeply than the other parts.
48 ELEMENTARY HISTOLOGY.

b. Articular cartilage. This forms caps covering the

ends of those bones which fit together to form move-
able joints: the caps act as elastic cushions to break
the force of shocks.
Mount in balsam a prepared section of articular cartilage from
the head of the femur, the section being made perpendicular to the
articular surface: examine with low and high powers.
i. The matrix is hyaline or faintly granular.
ii. The cartilage cells. Towards the free surface the
cells and cell groups become gradually flattened,
and arranged paralled to the surface.

F. Bone.
Bone consists of a dense fibrillar intercellular matrix, in
which are imbedded cells which lie in cavities connected with
one another by fine branching canals. The matrix is richly
impregnated with inorganic salts, chiefly phosphate and car-
bonate of lime, which form about two-thirds by weight of the
substance of the bone, and give it its great hardness and strength.
The matrix, with its contained bone-cells, is arranged in con-
centric layers or lamelle, around tubular passages, the Haversian
Canals, in which lie the bloodvessels, which penetrate the bone
in great numbers. A Haversian canal with its contained
bloodvessels, and its surrounding layers of matrix and cells, are
together spoken of as a Haversian system. .
1. Examine with both low and high powers prepared trans-
verse sections of a long bone.
i. The Haversian systems form the greater part of
the bone, and are readily recognised by the con-
centric arrangement of the lamelle, and the
central canals.
ii. The interstitial lamelle fill up the spaces between
the Haversian systems. They form parts of circies
which are in many cases of much larger radius
than the circles of the Haversian systems.
iii. The lacune are the spaces in the matrix in which
the bone-cells lie. In sections of dried bone the
lacunee appear black, through being filled either
with air or with dirt.
 BONE. 49

iv. The canaliculi are very fine branching canals con-

necting the lacune together: they are occupied
while the bone is living by branching processes
of the bone-cells. At the outer part of each
Haversian system, some of the canaliculi are
looped, opening at both ends of the loop into
the same lacuna.
v. The large central medullary cavity of the bone is
occupied during life by the marrow, which
consists of adipose tissue, with very numerous
bloodvessels and large nucleated reddish marrow
cells.
vi. The peripheral or circumferential lamelle are a
series of concentric lamelle parallel to the surface
of the bone, and forming its most superficial
layer.
vii. The perimedullary lamelle are a series of concen-
tric lamellee lining the central medullary cavity
of the bone.
50

CHAPTER IV.

THE SKELETON OF THE FROG.

The skeleton, which forms the hard internal parts of the frog,
is composed partly of cartilage and partly of bone. It forms a
framework giving definite shape to the body, and precision to
the movements ; and serves also to protect from injury some of
the more important and delicate organs, notably the central
nervous system, the sense organs and the heart. In the early
stages of its development the skeleton consists entirely of car-
tilage: in the adult this primary cartilaginous skeleton is replaced
to a greater or less extent by bone. Bone may also be developed
in places where there was no pre-existing cartilage, and is then
called membrane-bone, in contradistinction to the former kind,
or cartilage-bone, which replaces cartilage. Membrane bones
arise in the first instance as ossifications in the dermis or deeper
layer of the skin: in many fish they retain this primitive
position, but in the frog and most higher vertebrates they sink
below the skin and graft themselves on to the more deeply placed
cartilaginous skeleton. Cartilage may also become calcified,
2.e. have calcareous salts deposited in its matrix, without in any
way taking on the character of true bone.
The skeleton may conveniently be divided into (1) the axial
portion, including the skull and the vertebral column: and
(2) the appendicular portion, including the limbs, and the
limb-girdles which attach them to the body.
Examine the prepared skeletons, and make careful drawings to
scale of the several parts. In your drawings colour the cartilage
blue, the cartilage bones yellow, and the membrane bones white or
red, Prepare skeletons for yourself by soaking the parts in hot
water, and carefully brushing away the soft tissues until the
skeleton is clean,
 THE SKELETON OF THE FROG. 51

ra pls
igh

&

Fig. 8. The skeleton of the frog, seen from the dorsal surface
Suprascapula and scapula have been removed. ; the left

a, astragalus : c, caleaneum : d, suprascapula: €, exoccip

Jp, frontoparietal : g, metarcarpals : h, humerus: 7, ilium: ital :f, femur:
4, carpus: m, maxilla: n, nasal: 0, pro-otic: p, pterygo4, metatarsals:
id: pm, pre-
maxilla: g, quadratojugal: 7, radio-uln a: 8, Squamosal: se, sphene
8.v, sacral vertebra : ¢, tibio-fibula : u, urostyle. thmoid:
52 THE SKELETON OF THE FROG.

A. The Axial Skeleton.

I, The Vertebral Column or “ back bone.”
This is a bony tube which surrounds and protects the spinal
cord ; it consists of an anterior part which is divided trans-
versely into nine rings or vertebre, and a posterior unsegmented
portion of about equal length—-the urostyle. At the sides of
the tube, between the successive vertebree, are the intervertebral
foramina through which the nerves pass out from the spinal
cord to the various parts of the body.
a. Structure of a vertebra. Haamine one of the vertebra,
say the third, more closely ; draw it, showing the follow-
ing points :
i. The vertebra is a bony ring ; the spinal cord lying
during life in the central neural canal.
ii. The centrum or body is the thickened ventral por-
tion of the ring: it articulates with the centra of
the vertebree in front of and behind it ; and forms
the floor of the neural canal.
ili. The neural arch consists of the lateral and dorsal
portions of the ring; and forms the sides and
roof of the neural canal.
iv. The spinous process or neural spine is a small blunt
median process, projecting upwards and _back-
wards from the top of the neural arch.
v. The transverse processes are a pair of large processes
projecting horizontally outwards from the sides
of the neural arch,
vi. The articular processes or zygapophyses, on the
anterior and posterior borders of the neural arch,
articulate with corresponding processes on the
vertebree in front and behind, and so serve to
link the vertebrze together.
a. The anterior articular processes, or prezyg-
apophyses, face upwards and slightly in-
wards.
GB. The posterior articular processes, or post-
zygapophyses, face downwards and slightly
outwards.
 THE VERTEBRAL COLUMN, 5S

b. Special vertebra.
i. The atlas or first vertebra articulates in front
with the posterior end of the skull: it has no
transverse processes. Note the large gap on the
dorsal surface between the skull and the neural
arch of the atlas: through this gap, which is
closed by the strong occipito-atlantal membrane,
the central nervous system is divided and des-
troyed in the operation of pithing a frog.
ii. The sacrum, or ninth vertebra, has very stout
backwardly directed transverse processes which
support at their outer ends the pelvic arch.
c. The urostyle is the unsegmented posterior portion of
the vertebral column. It articulates in front with the
body of the sacral vertebra by two surfaces. Along
its dorsal surface runs a prominent vertical ridge,
highest in front and gradually diminishing posteriorly:
the neural canal is continued down the anterior part
of this ridge. At the sides of the urostyle, and about
the length of a vertebra from its anterior end, are a
pair of small holes through which nerves pass out,
and which therefore correspond to intervertebral
foramina,

II. The Skull.

The skull consists of, (1) an axial portion, the cranium,
enclosing the brain and forming an anterior continuation of the
vertebral column: (2) the olfactory capsules and the auditory
capsules, which are fused with the anterior and posterior ends
of the cranium respectively : (3) the bony framework of the
jaws: and (4) the hyoid apparatus.
In the skull the original cartilage, or chondrocranium, is not
so largely replaced by bone as in the vertebral column, large
tracts of unossified cartilage persisting in the adult. Besides
the cartilage-bones the skull is further strengthened by the
addition of numerous membrane-bones.
1, The Cranium is an unsegmented cartilaginous tube, whose
cavity forms the anterior part of the neural canal,
and lodges the brain. The roof of the tube is imper-
54 THE SKELETON OF THE FROG.

fect, there being one large. anterior fontanelle, and

two smaller posterior fontanelles, which are closed by
membrane only. In the cartilage are developed carti-
lage-bones, and around it membrane-bones.
To study the cranium satisfactorily, the membrane-bones should
be stripped from one of the skulls you have prepared for yourself.
a. Cartilage-bones of cranium.
i. The exoccipitals are two irregular bony masses at
the sides of the posterior end of the skull. They
almost completely surround the foramen magnum
or entrance to the cranial cavity ; and bear on
their posterior surfaces the occipital condyles,
two oval convex processes which articulate with
the first vertebra or atlas.

Fig. 9. The frog’s skull, from the ventral surface.

a, parasphenoid : c, columella: e, exoccipital: fp, frontoparietal:
m, maxilla: nm, vomer: o, pro-otic: p, pterygoid: pa, palatine:
pm, premaxilla : g, quadratojugal : se, sphenethmoid.
ii. The sphenethmoid or girdle-bone is a bony tube
which encircles the anterior end of the cranial
cavity, and extends forwards into the olfactory
region: in front it is divided by a vertical partition
into right and left cavities, in which lie the
olfactory sacs.
b. Membrane-bones of cranium.
i. The fronto-parietals are two long flat bones on the
top of the brain-case, covering the fontanelles,
 THE SKULL. 55

and overlapping the hinder end of the spheneth-

moid.
iu. The parasphenoid is a L shaped bone on the ventral
surface of the cranium; its lateral processes
underlying the auditory capsules,
2. The Sense Capsules are cartilaginous and bony capsules
which surround and protect the olfactory and auditory
organs ; they are fused with the cranium so as to form
parts of the skull.
a. The auditory capsules are fused with the sides of the
posterior end of the cranium, to which they form
wing-like projections: they consist largely of cartilage.
i, Cartilage-bone of auditory capsules.
The pro-otics are a pair of irregular shaped bones
in the anterior walls of the capsules, and forming
also. parts of their roof and floor.
b. The olfactory capsules are fused with the anterior end
of the cranium, and also with each other. They
consist very largely of cartilage, which is produced in
front into the rhinal processes.
i, Cartilage-bone of olfactory capsules. The spheneth-
moid, as already noticed, extends forwards so as to
invade the olfactory region, but does not properly
belong to the olfactory capsules.
li, Membrane-bones of olfactory capsules.
a, The nasals are two triangular bones on the dorsal
surface of the anterior end of the head: the bases
of the triangles are turned towards the middle
line and meet each other in front, while their
posterior ends diverge and enclose,. with the
anterior ends of the frontoparietals, a diamond
shaped patch in which the sphenethmoid is visible
on the dorsal surface of the skull.
B. The vomers are two triradiate bones on the ventral
surface of the fore part of the skull: each vomer
bears in its inner and posterior angle a small
group of pointed teeth, and forms the inner bound-
ary of the posterior narial opening of its side.
56 THE SKELETON OF THE FROG.

3. The Jaws consist of two cartilaginous arches on each side,

maxillary and mandibular, in connection with which
cartilage-bones and membrane-bones are developed.
Each arch meets its fellow in the middle line in front;
and the maxilliary arches, forming the upper jaw, are
firmly connected with the cranium by anterior and
posterior bony struts.
a. The Maxillary Arch. All the bones of the upper jaw
are membrane-bones except the quadratojugal. In the
case of the pterygoid and palatine bones ossification
may extend into the underlying cartilage.

Fig. 10. The frog’s skull from the right side,

A, parasphenoid: AS, angulosplenial: B, anterior cornu of hyoid:
C, columella: D, dentary: E, exoccipital : F, nostril: FP, frontoparietal:
‘H, body of hyoid: L, aperture for exit of optic nerve: M, maxilla:
MM, mentomeckelian: M’, aperture for exit of fifth and seventh nerves :
N, nasal: O, pro-otic : P, pterygoid :PM, premaxilla: Q, quadratojugal :
R, aperture for exit of ninth and tenth nerves: S, squamosal: SE, sphen-
ethmoid : T, posterior cornu of hyoid.

i. The pterygoid is a large triradiate bone, the inner

limb of which is connected with the auditory
capsule ; while the posterior limb runs back to
the angle of the mouth, and the anterior limb
forwards along the upper jaw to the palatine
bone.
ii. The palatine is a slender transverse bone, connecting
the upper jaw with the anterior end of the
sphenethmoid.
 THE SKULL. 57

ui. The quadratojugal is a short bone forming the pos-

terior part of the margin of the upper jaw.
iv. The maxilla is a long thin bone forming the greater
part of the margin of the upper jaw: it bears
teeth along its whole length which are anchylosed
with the bone. It is connected behind with the
quadratojugal ; about the middle of its length
with the anterior limb of the pterygoid and with
the palatine ; and in front with the premaxilla.
v. The premazilla is a small bone which meets its
fellow in the middle line in front, and so com-
pletes the margin of the upper jaw: like the
maxilla it bears teeth. It gives off on its dorsal
surface a backwardly projecting process which
forms part of the inner boundary of the nostril.
b. The Mandibular Arch. The upper part of the arch
remains unossified as the quadrate cartilage, which
forms the suspensorium, 7.¢., serves to connect the lower
jaw with the skull: this is a rod of cartilage which is
fused above with the auditory capsule, and runs down-
wards and backwards to the angle of the mouth, where
it is connected with the hinder end of the quadratojugal
bone. In the adult frog, the quadrate cartilage lies
between the squamosal and pterygoid bones, and is
almost completely concealed by these. The lower part of
the arch persists in part unossified as Meckel’s cartilage,
which forms the basis of the lower jaw, and is ensheathed
by cartilage-bones and membrane-bones.
i, The squamosal is a T shaped bone, the stem of
which is closely applied to the outer surface of the
quadrate cartilage. The posterior limb of the T
is attached to the outer surface of the auditory
capsule, and with the body of the squamosal
helps to support the annulus tympanicus.
ii, The angulosplenial ensheaths the inner and lower
surfaces of Meckel’s cartilage along the greater
part of its length: near its hinder end it is
produced upwards into the coronary process.
58 THE SKELETON OF THE FROG.

ili, The dentary is a flat bone covering the outer sur-

face of the distal half of Meckel’s cartilage, as far
forward as the mentomeckelian bone,
iv. The mentomeckelian is a small ossification in
Meckel’s ‘cartilage at the symphysis, ée, the
union of the arches of the two sides at the chin.
Fpl
+ Kolm

Fig. 11. A transverse section across the posterior part of the frog’s
head, to show the position and relations of the auditory organs,
Eustachian tubes, and hyoid apparatus. On the right side the section
passes through the tympanic cavity and the columella; on the left side
through the anterior cornu of the hyoid. The cartilage is dotted, and
the bones, except the columella, are represented black.
A, parasphenoid : As, angulosplenial : B, buccal cavity : C, columella:
D, tympanic membrane: E, Eustachian tube: F, anterior cornu of the
hyoid : FP, frontoparietal :G, glottis: H, arytenoid cartilage: 1, posterior
cornu of hyoid : K, auditory nerve: L, vestibule: M, anterior vertical
semicircular canal: N, horizontal semicircular canal: O, pro-otic :
P, pterygoid: Q, quadrate cartilage : R, quadratojugal: S, squamosal :
T, annulus tympanicus: V, vocal cord: X, mid-brain.

4, The Hyoid Apparatus (Fig. 10). This consists of the

hyoidean arch and the remains of the branchial arches
of the two sides, together with a median ventral
plate, the body of the hyoid, which unites their
lower ends together, and lies in the floor of the mouth.
The hyoid apparatus consists almost entirely of
cartilage.
a. The hyoid arch.
i. The columella (Figs. 10 and 11) is formed from the
top of the hyoid arch: it is a small rod, partly
 THE APPENDICULAR SKELETON. 59

bone and partly cartilage, the inner end of which

is inserted into the fenestra ovalis, an aperture
in the outer wall of the auditory capsule ; while
the outer end is attached to the tympanic mem-
brane rather above its middle.
ii, The anterior cornu of the hyoid (Fig. 10) is a long
slender curved rod of cartilage, attached above to
the auditory capsule just below the fenestra ovalis,
and curving at first backwards and then forwards
and downwards to be attached to the anterior
outer angle of the body of the hyoid.
b. The body of the hyoid is a flat squarish plate of cartilage,
formed by the fused ventral ends of the hyoid and
branchial arches, and lying in the floor of the mouth:
short processes are given off from its angles.
c. The posterior cornua of the hyoid are a pair of stout
bony processes, diverging from the hinder border of
the body of the hyoid.

B. The Appendicular Skeleton.

This comprises the limbs and the limb-girdles, As in the case
of the axial skeleton it consists at first entirely of cartilage, which
becomes afterwards replaced to a greater or less extent by carti-
lage-bone. Membrane-bones, 7.¢., bones developed independently
of cartilage, are very rare, the clavicles being the only examples
met with in the frog. (Fig. 8, p. 51).
1. The Pectoral Girdle.
This consists originally of two half rings of cartilage, one on
each side of the body, which they encircle a short way behind
the head: the dorsal ends of the half rings are attached by
ligaments and muscles to the vertebral column, while the ven-
tral ends are united together in the median plane by the sternum
or “breast bone.”
Kach half ring bears in the middle of its hinder surface a cup-
shaped cavity, which, with the first bone of the fore-limb, forms
the shoulder joint. The part of the arch above the joint is the
scapular portion: and the part below, which is divided into
anterior and posterior divisions, the coracoid portion.
60 THE SKELETON OF THE FROG.

a. The scapular portion is divided into two parts.

i. The suprascapula, the upper portion, is a thin
expanded plate of cartilage overlapping the first
four vertebra: it is partly calcified and partly
ossified, though very imperfectly.
ii. The scapula is an oblong bony plate, constricted in
the middle, and forming the upper half of the
glenoid cavity, or cavity of the shoulder joint,
b. The coracoid portion forms the lower half of the glenoid
cavity: it is divided into anterior and posterior portions,
separated by the coracoid foramen.
i. The precoracoid is a slender horizontal bar of car-
tilage connecting the anterior edge of the scapula
with the sternum.
ii. The clavicle is a slender bone, closely applied to the
anterior border of the precoracoid ; its outer or
scapular end is bent forwards almost at a right
angle.
iii. The coracoid is a stout bone, wider at its inner
than its outer end: it connects the posterior
edge of the scapula with the sternum.
c. The sternum lies in the mid-ventral line: it consists
originally of two lateral halves which fuse com-
pletely in front and behind, but remain distinct
in the median portion. It presents from before
backwards the following parts :
i. The episternum is a flat circular plate of cartilage.
ii. The omosternum is a slender bony rod projecting
forwards in front of the clavicles.
iii. The epicoracoids are a pair of narrow strips of
cartilage closely applied to each other, and lying
between the ventral ends of the precoracoids and
coracoids.
iv. The sternum proper is a rod of cartilage, ensheathed
in bone, projecting backwards behind the coracoids.
v. The xiphisternum is a broad expanded plate of
cartilage at the hinder end of the sternum.
 THE APPENDICULAR SKELETON. 61

2. The Fore-Limb.
The bones of the fore-limb are all cartilage-bones. With the
exception of the small bones of the wrist, they are elongated,
with enlarged ends capped with articular cartilage. The
enlarged ends or epiphyses ossify independently of the shaft of
the bone, with which they do not unite until late in life. The
end of a bone which, when the limb is extended, is nearer to
the body, is called its proximal end, the opposite extremity the
distal end
a. The arm. In the arm there is only a single bone.
i. The humerus. The proximal end or head is en-
larged, and articulates with the glenoid cavity
of the pectoral girdle: below the head is the
strong deltoid ridge extending along the proximal
half of the anterior surface. At the distal end
is a spheroidal articular surface for the bone of
the forearm: and at either side of this a
prominent condylar ridge, the inner or postaxial
one being the larger of the two, especially in the
male frog.
b. The fore-arm.
i. The radio-ulna corresponds to two bones, radius.
and ulna, in other animals: it is single at its
proximal end, but in its distal half is imperfectly
divided by a groove into anterior or radial, and
posterior or ulnar portions. Its proximal end is
hollowed out to articulate with the lower end of
the humerus, and so form the elbow joint, be-
hind which it projects backwards as the olecranon
process.
c. The wrist consists of six small carpal bones arranged in
two rows, proximal and distal, each row having three
bones.
d. The hand has four complete digits and a rudimentary
pollex or thumb. Each digit consists of a proximal
metacarpal bone, beyond which are a variable number
of phalanges.
i. The pollex, the anterior or preaxial digit, consists
simply of a small metacarpal bone.
62 THE SKELETON OF THE FROG.

ii. The first complete digit, corresponding to the fore

finger of man, consists of a metacarpal and two
phalanges.
iii, The second digit, corresponding to the middle
finger of man, consists of a metacarpal and two
phalanges.
vi. The third, corresponding to the ring finger of man,
consists of a metacarpal and three phalanges.
v. The postaxial digit, corresponding to the little finger
of man, consists of a metacarpal and three phalanges.

3. The Pelvic Girdle.

This consists primitively, like the pectoral girdle, of a couple
of half-rings of cartilage, fused together below and attached
above to the tips of the transverse processes of the sacrum. In
the adult frog the girdle is placed very obliquely so as to be
nearly parallel with the vertebral column instead of at right
angles to it.
Each half presents on its outer aspect a cup-shaped cavity—
the acetabulum—forming, with the thigh bone, the hip-joint : we
accordingly distinguish an iliac portion above the acetabulum
and an ischio-pubic portion below it, corresponding respectively
to the scapular and coracoid divisions of the pectoral girdle.
i. The ilium forms the anterior and upper half of
the acetabulum, and extends forwards as an
elongated laterally-compressed bar, which is
attached in front to the transverse process of the
sacrum and bears along its dorsal surface a
prominent vertical ridge of bone, the iliac crest,
ending behind in an abrupt vertical border.
Posteriorly the two ilia meet each other and are
united together in the median plane to form the
iliac symphysis.
ii, The pubes consists entirely of cartilage: it forms
the anterior portion of the ventral division of the
girdle, and therefore corresponds to the pre-
coracoid in the pectoral girdle. The two pubes
are completely fused together in the median
plane, and form only a very small portion, about
one-sixth, of the acetabular cavities.
 THE APENDICULAR SKELETON. 63

iii. The ischium is the posterior portion of the ventral

division and corresponds therefore to the coracoid
in the shoulder girdle. It forms the posterior
third of the acetabulum. ‘The two ischia are
completely fused together in the median plane.

4, The Hind-Limb.
The bones have the same general characters as those of the
fore-limb, to which they correspond very closely.
a. The thigh.
i. The femur is a long slender bone, expanded at both
ends, and curved slightly in a sigmoid manner.
The proximal end or head is spheroidal, and fits
into the acetabulum to form the hip joint: the
distal end is somewhat expanded laterally.
b. The leg.
i. The os cruris or tibio-fibula isa single bone, rather
longer than the femur, slightly curved, and
expanded laterally at both ends. It presents
along the greater part of its length a groove
indicating its correspondence with two bones,
tibia and fibula, which in man and many other
animals remain distinct from each other.
¢. The ankle, corresponding to the wrist in the fore-limb,
consists of two rows of tarsal bones.
a. The proximal row of tarsal bones consists of two
elongated bones united together at both ends, but
widely separated in the middle.
i. The astragalus is on the preaxial or tibial side.
ii. The caleaneum is on the postaxial or fibular
side, and is the larger of the two bones.
B. The distal row of tarsal bones consists of two very
small bones.
d. The foot has five complete digits, and a supernumerary
toe as well. Each digit consists of a proximal
metatarsal bone, beyond which are a variable
number of phalanges.
64 THE SKELETON OF THE FROG.

i. The hallux or preaxial digit, corresponding to the

great toe of man, is the smallest of the series.
It consists of a metatarsal and two phalanges.
On the inner side of the hallux is the calcar,
supposed to be an additional or supernumerary
toe: it may have one or two joints in addition to
a short metatarsal.
il. The second toe consists of a metatarsal and two
phalanges.
ii. The third consists of a metatarsal and three
phalanges.
iv. The fourth, the longest of the five, consists of a
metatarsal and four phalanges.
v. The postaxial digit, corresponding to the little toe
in man, consists of a metatarsal and three
phalanges.
 65

CHAPTER YV.

THE MUSCULAR SYSTEM OF THE FROG.

The muscles, or flesh, are the direct means by which the

various movements of the body and of its several parts are
brought about. A muscle consists of a fleshy belly, which is
usually attached at each end by means of tendons to some, hard
part, very commonly to bone. Motion is effected by the muscle
contracting, 7.c., shortening, and so bringing its two ends, and
consequently the parts to which the ends are attached, nearer
together. Of the two attachments of a muscle one is usually
to a more fixed and central part, the other to a more moveable
and peripheral part: the former attachment is called the origin
of the muscle, the latter its insertion.
Muscles are of two kinds: (1) voluntary muscles, 7.¢., those
which are under the control of the will, as the muscles of the
arm: and (2) involuntary muscles, i.c., those over which the
will has no direct control, as the muscles of the heart and
bloodvessels, or of the alimentary canal.
Voluntary muscles, which are the only ones dealt with in this
chapter, are usually attached at both ends to bone; but one or
other end, or both, may be attached to aponeuroses, strong
connective tissue membranes which ensheath the muscles and
other parts, and separate them from one another.
For the dissection of the muscles, take a frog that has been in sprit
for a day or more. When cleaning a muscle be careful to-put tt on
the stretch, and to dissect along, and not across tts fibres: define
the origin and insertion of the muscle very clearly, and test tts
action by pulling it gently with the forceps in the direction of
its fibres. Always have the skeleton in front of you so as to see
accurately the origins and insertions of the muscles. In the
following description some of the smaller muscles, especially in the
head, are omitted.
F
66 THE MUSCULAR SYSTEM OF THE FROG.

A. Muscles of the Trunk.

I. Muscles of the ventral body-wall.
Pin out the frog on tts back, remove the skin, and clean the muscles.
i. The rectus abdominis runs longitudinally along
the midventral wall, the muscles of the two sides
being separated from each other in the median
plane by the linea alba, a longitudinal band of
connective tissue, immediately dorsal to which ™
lies the anterior abdominal vein. Each rectus
muscle is divided into bellies by five transverse
tendinous intersections.
The muscle arises from the pubes, runs forwards
and is inserted into the dorsal surface of the
sternum and coracoid.
ii. The pectoralis is a large fan-shaped muscle, consist-
ing of a thoracic portion, which arises from
the whole length of the ventral surface of the
sternum ; and an abdominal portion, arising from
the aponeurosis along the outer side of the
rectus abdominis almost as far back as the
pubes. From this extensive origin the fibres con-
verge to the deltoid ridge of the humerus, into
which they are inserted, the line of insertion
extending down almost to the elbow.
iii. The obliquus externus is a thin sheet of muscle
which arises from the aponeurosis of the back, a
short distance from the vertebral column, and
covers the whole of the side of the body, the
fibres running obliquely downwards and_back-
wards to end in an aponeurosis which passes
dorsal to the rectus abdominis to be inserted
into the linea alba.
iv. The obliquus internus lies beneath the obliquus
externus, which must be removed in order to
see it. It arises from the transverse processes of
the vertebree from the fourth backwards, and
from the ilium. The fibres run downwards and
forwards, and are inserted in front into the cora-
coid and sternum: some of the fibres surround, and
are inserted into, the cesophagus and pericardium.
 MUSCLES OF THE TRUNK. 67

The hinder two-thirds of the muscle pass dorsal

to the tendon of the obliquus externus and are
inserted, like it, into the linea alba.
2. Muscles of the Back.
Pin out the frog on its belly: remove the skin, and clean the
muscles in order.
i. The depressor mandibule is a broad triangular
muscle which arises from the fascia covering the
dorsal surface of the suprascapula: the fibres run
downwards behind the tympanic membrane, and
converge to be inserted into the angle of the
lower jaw. The muscle by its contraction opens
the mouth.
iii The cucullaris is a small oblong muscle which,
arising from the exoccipital near the middle line,
runs backwards and outwards, and is inserted
into the dorsal border of the suprascapula.
iii, The latissimus dorsi is a triangular muscle lying
behind the depressor mandibule: it arises from
the fascia dorsalis just behind the shoulder girdle,
its origin being partly covered by the obliquus
externus. The fibres run forwards and outwards,
converging to be inserted by a long tendon into
the deltoid ridge of the humerus.
Dissect away the depressor mandibule and latissimus dorsi
from their origins, and turn them down.
iv. The infraspinatus arises from the dorsal surface of
the suprascapula, partly overlapped by the
latissimus dorsi: it runs outwards to be inserted
into the deltoid ridge of the humerus: its action
is to elevate the arm.
Lvft up the suprascapula and note the muscles attaching it to
the body, viz. :
v. The retrahens scapule, behind.
vi. The levator anguli scapule, in front.
Remove the suprascapula on one side, and clean the median
longitudinal muscles of the back.
vii. The extensor dorsi communis is a longitudinal mass
of muscle arising from the urostyle, and running
68 THE MUSCULAR SYSTEM OF THE FROG.

forwards and slightly outwards: it is inserted

into the ilium, into the transverse processes of
the vertebre, and in front into the posterior end
of the skull. The anterior part is divided by
transverse tendinous intersections.
viii. The intertransversales are small muscles, running
between the transverse processes of the vertebree,
and lying beneath the extensor communis.
ix. The gluteus arises from the outer side of the pos-
terior two-thirds of the ilium, and runs backwards
to be inserted into the trochanter of the femur.

B. Muscles of the Head.

I. Muscles of the ventral surface of the head,
i. The mylohyoid or submandibular muscle is a flat
sheet of muscle running across from one ramus
of the mandible to the other, and divided down
the middle line by a tendinous intersection. A
narrow strip along the posterior border is
commonly separated by a slight interval from the
major or anterior part of the muscle.
Remove the mylohyoid muscle, and the sternal portion of the
pectoralis.
ii. The geniohyoid is a narrow longitudinal band a
short distance from the middle line: it arises from
the lower jaw close to the chin, runs back on the
ventral surface of the body of the hyoid, and
divides posteriorly into two portions which are
inserted respectively into the bony and the car-
tilaginous posterior processes of the hyoid.
iii. The sternohyoid is practically the anterior con-
tinuation of the rectus abdominis. It arises from
the dorsal surface of the coracoid and clavicle,
and is inserted into the ventral surface of the
body of the hyoid, the tendon passing between
the two divisions of the geniohyoid.
iv. The hyoglossus arises on either side from the
posterior bony horn of the hyoid: the two muscles
converge and meet each other in front of the
 MUSCLES OF THE HEAD. 69

larynx. In front of the larynx the muscle runs

forward in the middle line as a stout band nearly
to the chin: it then enters the tongue, and runs
along it backwards to the tip.
y. The petrohyoid muscles are a set of four muscular
bands which arise close together from the outer
surface of the auditory capsule, and diverging in
a fan-like manner, pass round the floor of the
pharynx and cesophagus to be inserted in front
into the median ventral line of the pharynx, and
behind into the side of the hyoid. The first or
most anterior band is a wide thin sheet of mus-
cular tissue, while the three posterior portions
are very narrow slips.
2. Muscles of the side of the head.
Remove the skin carefully from the side of the head and jaws,
noticing how much more closely it is attached to the underlying
parts than was the case in the body.
a. Depressors of the lower jaw ; opening the mouth.
i, The depressor mandibulz has been already seen
and dissected. (See p. 67.)
b. Elevators of the lower jaw; shutting the mouth.
These lie in the space between the auditory capsule
and the eye.
ii. The temporalis arises from the upper surface of
the auditory capsule, and passes outwards and
downwards between the pterygoid and maxillary
bones, and in front of the cartilaginous ring sup-
porting the tympanic membrane, from which
some of its fibres arise: it is inserted into the
coronoid process of the lower jaw.
ili. The pterygoideus is a slender muscle placed just in
front of the temporalis and partly covered by it:
it arises from the side wall of the skull, and is
inserted into the mandible further back than the
temporalis, and very close to the joint.
iv, The masseter is a small muscle placed behind the
temporalis: it arises from the quadratojugal and
70 THE MUSCULAR SYSTEM OF THE FROG.

runs downwards and slightly backwards to be

inserted into the outer surface of the mandible,
just in front of the joint.
To see the insertions of these last three muscles the mouth should
be opened widely.
3. Muscles of the eyeball.
Remove the temporal aud pterygoid muscles carefully, dissecting
them away from their origins, and then turning the muscles down
and cutting them short close to their insertions. Remove also the
lower jaw ; pin the frog out on its back and dissect away carefully
the mucous membrane of the roof of the mouth.
i. The levator bulbi is a thin sheet of muscle lying
between the mucous membrane and the eye. Its
fibres arise from the side of the skull, run out-
wards underneath the eye, and are inserted into
the upper jaw. The muscle by its contraction
serves to lift up the eyeball and so make it more
prominent. Some of its fibres are inserted into
the lower eyelid, which they serve to depress,
acting as a depressor palpebre inferioris
Remove the levator bulbi and clean the remaining muscles,
dissecting them partly from the dorsal and partly from the
ventral surface.
a. The recti muscles are a group of four small muscles
which arise close together from the inner and posterior
angle of the orbit close to the optic foramen, and run
forwards and outwards, diverging from one another, to
be inserted into the bulb of the eye.
i, The rectus superior is inserted into the dorsal
surface of the eyeball: it is seen best from above.
ii. The rectus externus, the most posterior of the
four, is inserted into the posterior surface of the
eyeball: it is seen best from the side or from
below.
ili. The rectus internus, the longest of the four, runs
forwards between the skull wall and the eyeball,
and is inserted into the inner or median surface
of the eyeball: it is seen best from below.
 MUSCLES OF THE EYEBALL. 71

iv. The rectus inferior is inserted into the under surface

of the eyeball : it is seen best from below.

b. The obliqui muscles are a group of two small muscles

which arise close together from the palatine bone at the
anterior end of the orbit, and run backwards to be
inserted into the eyeball.
i, The obliquus superior is inserted into the dorsal
surfaceof the eyeball just in front of the rectus
superior: it is seen best from above.
ii, The obliquus inferior passes backwards beneath
the rectus internus, and is inserted into the eye-
ball between it and the rectus inferior: it is seen
best from below.
c. The retractor bulbi, or choanoid muscle, is a funnel-
shaped muscle which lies within the four recti and
embraces the optic nerve: it arises from the para- -
sphenoid, and is inserted into the eyeball. It is best
exposed from below by carefully removing the recti
muscles.

C. Muscles of the Hind-limb.

If the frog’s leg be stretched back parallel to the longitudinal
axis of the body, as in the act of swimming, we distinguish in
it ventral and dorsal surfaces, an outer border in which is the
projection of the knee, and an inner ‘border along which is the
bend of the knee. The outer border, which corresponds to the
front of the leg in man, is called the extensor surface, inas-
much as the muscles which extend or straighten the leg lie
along this edge : the inner border is the flexor surface. The
“ventral” and “dorsal” surfaces only appear to be such in
consequence of the extreme obliquity of the pelvic girdle: they
are really anterior and posterior, and are better called preaxial
and postaxial: they correspond respectively to the inner and
outer surfaces of the human leg. If the foot be examined care-
fully, it will be seen that the first digit or “big toe” is on the
preaxial side, and hence may be called the preaxial digit:
while the fifth or “little toe” is on the postaxial side, and is
- therefore the postaxial digit.
72 THE MUSCULAR SYSTEM OF THE FROG.

1. Muscles of the thigh.

Remove the skin from one of the legs of the frog, and clean the
muscles first of the preaxtal and then of the postawial surfaces,
a. Superficial muscles of the preaxial (apparent ventral)
surface of the thigh.
i. The sartorius is a long narrow muscular band which
crosses the thigh somewhat obliquely from the
outer to the inner side. It arises from the iliac
symphysis below the acetabulum, and is inserted
into the inner side of the head of the tibia.
ii. The adductor magnus is a large muscle lying along
the inner border of the sartorius, but passing
beneath it at its distal end. It arises from the
pubic and ischial symphyses, and passes under
the sartorius to be inserted into the distal third
of the femur.
iii. The adductor longus is a long narrow muscle lying
along the outer side of the adductor magnus,
and often completely hidden by the sartorius:
it arises from the iliac symphysis beneath the
sartorius, and unites a little way beyond the
middle of the thigh with the adductor magnus.
iv. The rectus internus major is a large muscle lying
along the inner side of the adductor magnus
and of the sartorius. It arises from the ischial
symphysis and is inserted into the head of the
tibia.
v. The rectus internus minor is a narrow flat band of
muscle running along the inner, or flexor margin
of the thigh: it arises from a tendinous expansion
connected with the ischial symphysis, and is
inserted into the inner side of the tibia, just
below its head.
b. Superficial muscles of the extensor surface of the thigh.
i. The triceps extensor femoris, the great extensor
muscle of the thigh, arises by three distinct
origins, which will be described separately, and
is inserted into the tibia just below its head.
 MUSCLES OF THE HIND-LIMB. 73

GL Nuiti

Fig. 12. The superficial muscles of the frog’s left hind-limb. A : from
the preaxial surface. B : from the postaxial surface.
AB, adductor brevis: AM, adductor magnus: B, biceps: C, cloacal
aperture : EC, extensor cruris: F, distal end of femur: FT, tendon of
flexor tarsi: G, gastrocnemius: GL, glutzus : P, peroneus: PY, pyri-
formis: RA, rectus anticus femoris: RI, rectus internus major: RN,
rectus internus minor : S, sartorius :SM, semimembranosus; T, triceps
extensor femoris: TA, tibialis anticus : TF, tibio-fibula: TP, tibialis
Posticus : VE, vastus externus : VI, vastus internus.
74 THE MUSCULAR SYSTEM OF THE FROG.

a. The rectus anticus femoris forms the middle

division of the triceps: it arises from the ven-
tral border of the posterior third of the ilium,
in front of the acetabulum: about half way
down the thigh it joins the next division.
3. The vastus internus, the preaxial division of
the triceps, is a large muscle arising from
the ventral and anterior border of the
acetabulum, and lying in the thigh between
the sartorius and the rectus anticus.
y. The vastus externus, the postaxial division of
the triceps, arises from the posterior edge
of the dorsal crest of the ilium, and joins
the other two divisions of the triceps about
the junction of the middle and distal thirds
of the thigh.
c. Superficial muscles of the postaxial (apparent dorsal)
surface of the thigh.
i. The gluteus has been already noticed: it lies in
the thigh between the rectus anticus and the
vastus externus.
ii. The biceps is a long slender muscle which arises
from the crest of the ilium just above the
acetabulum : it lies in the thigh along the inner
border of the vastus externus, and is inserted by
a flattened tendinous expansion into the distal
end of the femur and the head of the tibia.
iii. The semimembranosus is a stout muscle lying along
the inner side of the biceps, between it and the
rectus internus minor. It arises from the dorsal
angle of the ischial symphysis just beneath the
cloacal opening, and is inserted into the back of
the head of the tibia. It is divided about its
middle by an oblique tendinous intersection.
iv. The pyriformis is a slender muscle which arises from
the tip of the urostyle, passes backwards and
outwards between the biceps and the semi-
membranosus, and is inserted into the femur at
the junction of its proximal and middle thirds.
 MUSCLES OF THE HIND-LIMB, 75

d. Deep muscles of the thigh.

Lay the frog on its back and dissect the thigh from the
preaxial surface. Separate the adductor magnus and the rectus
internus major with blunt instruments so as to expose the follow-
ing muscles :
i. The semitendinosus is a long thin muscle which
arises by two heads; an anterior one from the
ischium close to the ventral angle of the ischial
symphysis and the acetabulum ; and a posterior
one from the ischial symphysis. The anterior
head passes through a slit in the adductor mag-
nus and unites with the posterior head in the
distal third of the thigh. The tendon of inser-
tion is long and thin, and joins that of the rectus
internus minor to be inserted into the tibia just
below its head,
Divide the adductor magnus and the sartorius in the middle
and turn the cut ends backwards and forwards, so as to expose the
following muscles:
ii. The adductor brevis is a short wide muscle, lying
beneath the upper end of the adductor magnus.
It arises from the pubic and ischial symphyses,
and is inserted into the preaxial surface of the
proximal half of the femur.
ii. The pectineus is a rather smaller muscle, lying along
the outer (extensor) side of the adductor brevis.
It arises from the anterior half of the pubic sym-
physis in front of the adductor brevis, and is
inserted like it into the proximal half of the femur.
iv. The ilio-psoas arises by a wide origin from the
inner surface of the acetabular portion of the
ilium : it turns round the anterior border of the
ilium, and crosses in front of the hip joint, where
for a short part of its course it is superficial
between the heads of the vastus internus and of
the rectus anticus femoris: it then passes down
the thigh beneath these muscles, and is inserted
into the back of the proximal half of the femur.
v. The quadratus femoris is a small muscle on the back
of the upper part of the thigh: it arises from the .
76 THE MUSCULAR SYSTEM OF THE FROG.

ilium above the acetabulum, and from the base

of the iliac crest: it lies beneath the pyriformis
and behind the biceps, and is inserted into the
inner surface of the proximal third of the femur
between the pyriformis and the ilio-psoas.
vi. The obturator is a deeply situated muscle which
arises from the whole length of the ischial sym-
physis and the adjacent parts of the iliac and
pubic symphyses, and is inserted into the head
of the femur close to the gluteus.
2. Muscles of the Leg.
As in the thigh, we distinguish extensor and flexor surfaces,
corresponding to the front and back of the leg in man ; and also
preaxial and postaxial surfaces, corresponding to the inner and
outer sides of the human leg.
Lay the frog on its belly and commence the dissection from the
postaxial surface.
i. The gastrocnemius is the large muscle forming the
calf of the leg: it has two heads of origin, of
which the larger arises by a strong, flattened
tendon from the flexor surface of the distal end
of the femur; while the smaller head, which joins
the main muscle about one-fourth of its length
below the knee, arises from the edge of the tendon
of the triceps extensor femoris where it covers
the knee. The muscle is thickest in its upper
third, and tapering posteriorly ends in the strong
tendo Achillis, which passes under the ankle joint,
being much thickened as it does so, and ends in
the strong plantar fascia of the foot.
ii. The tibialis posticus arises from the whole length
of the flexor surface of the tibia: it ends in a
tendon which passes round the inner malleolus,
lying in a groove in the lower end of the tibia,
and is inserted into the dorsal surface of the
astragalus.
iii. The tibialis anticus lies on the extensor surface
of the leg: it arises by a long thin tendon from
the lower end of the femur, and divides about
 Ld
MUSCLES OF THE HIND-LIMB. i7

the middle of the leg into two bellies which are

inserted into the proximal ends of the astragalus
and calcaneum respectively.
iv. The extensor cruris lies along the preaxial side of
the tibialis anticus, partly covered by this and
partly by the strong fascia of the leg. It arises
by a long tendon from the preaxial condyle of
the femur, runs in a groove in the upper end of
the tibia, and is inserted into the extensor surface
of the tibia along nearly its whole length.
v. The peroneus is a stout muscle which lies along
the postaxial surface of the leg, between the
tibialis anticus and the gastrocnemius. It
arises from the distal end of the femur, and is
inserted into the outer malleolus of the tibia
and the proximal end of the calcaneum.
78

CHAPTER VI.

THE NERVOUS SYSTEM OF THE FROG.

The nervous system consists of the following parts :—

1, A central portion, the brain and spinal cord, which lies
in the cartilaginous and bony tube formed by the
cranium and vertebral column, and which is the centre
where sensations are felt, and whence motor impulses
causing the muscles to contract take their origin,
2. A peripheral portion, the nerves themselves, which
connect the central portion with the skin, sense
organs, muscles, viscera, etc., and serve to convey
sensory impulses from these parts to the brain and
cord, or motor impulses from the central organs to
the muscles. These two functions are fulfilled by
different nerves, which may accordingly be distin-
guished as (a) afferent or sensory nerves, conveying
impulses to the brain or cord and (0) efferent or
motor nerves, conveying impulses from the brain
or cord.
Fig. 13. The nervous system of the edible frog (Rana esculenta), from
the ventral surface. (From Ecker.)
F, facial nerve: G, ganglion of pneumogastric nerve: He, cerebral
hemisphere: Le, optic tract: Lop, optic lobe: M, boundary between
medulla oblongata and spinal cord : M 1-10, the spinal nerves: MS, con-
nection between fourth spinal nerve and sympathetic chain: N, nasal
sac: Ni, sciatic nerve: No, crural nerve: o, eyeball: S, trunk of
sympathetic: § 1-10, the sympathetic ganglia: Sp, continuation of
sympathetic into head.
I, olfactory nerve: II, optic nerve: III, motor oculi: IV, fourth
nerve: V, trigeminal and facial nerves: Va, ophthalmic branch of
trigeminal: Vc, maxillary branch of trigeminal : Vd, mandibular branch
of trigeminal : Ve, hyomandibular branch of facial: Vg, Gasserian gang-
lion.: Vs, upper end of sympathetic trunk, in connection with Gasserian
genenon : VI, abducens nerve, : VII, facial nerve : VIII, auditory nerve:
, glossopharyngeal and pneumogastric nerves: XI, ramus anterior of
pees haryngeal: X2, ramus posterior of glossopharyngeal: X 3-4,
ranches of pneumogastric.
THE CENTRAL NERVOUS SYSTEM. 79

i /A
\ oe S
NU); \
p) \ !\

~ Ma
Mo!
80 THE NERVOUS SYSTEM OF THE FROG.

A special set of nerves in connection with the bloodvessels

and viscera forms the sympathetic nervous system.
For the dissection of the nervous system specimens should be
taken which have been in strong spirit for two or three days, and
in which the brain has been exposed to the action of the spirit by
removal of the roof of the skull.

A. The Central Nervous System.

This is divisible into an anterior portion—the brain—lying
in the cavity of the cranium; and a posterior portion—the
spinal cord—which lies in the neural canal of the vertebral
column. There is no sharp line of demarcation between the two
portions, which are directly continuous with each other.
If the brain and spinal cord have not already been exposed,
clear away the dorsal muscles from both sides of the spine: cut
through the occipito-atlantal membrane, flexing the frog’s head
slightly to make the membrane tense, and bemg careful not to
injure the brain beneath it. Introduce one blade of the scissors
into the cranial cavity, with the flat surface of the blade paraliel
to the back of the frog, and keeping as close to the roof of the skull
as possible, Cut carefully through the side walls of the skull,
first on one side and then on the other. Turn the roof of the skull
forwards with forceps, and remove tt altogether.
Similarly. cut through and remove the neural arches of the
vertebree one by one,f rom before backwards.
Examine and draw the central nervous system in situ, showing
its several parts.
I. The Brain.
a. The dorsal surface of the brain: note from before
backwards the following parts, removing the pigmented
membrane—pia mater—covering the several parts as
you come to them.
i. The olfactory lobes, which form the most anterior
portion of the brain, are united together in the
median plane: they give off the olfactory nerves
from their anterior ends, and are separated behind
by slight constrictions from the hemispheres.
ii. The cerebral hemispheres are a pair of smooth
ovoid bodies which touch each other in the median
plane but are not fused together.
 THE BRAIN. 7 81

iii. The thalamencephalon is a lozenge-shaped portion

lying immediately behind the hemispheres and
between their diverging posterior ends: it is
covered by a thick vascular membrane-—the
choroid plexus—over which passes the stalk of the
pineal body, a small body adherent to and re-
moved with the roof of the skull, On removing
the choroid plexus a slit-like hole is left in the

Fig. 14. The brain of the frog: dorsal surface. x 4.

Fig. 15. The brain of the frog : ventral surface. x 4.
C, cerebellum : CH. cerebral hemisphere : CP, choroid plexus of third
ventricle : F, fourth ventricle : IN, tuber cinereum : M, medulla oblongata,
O, olfactory lobe: OC, optic chiasma: OL, optic lobe: P, stalk of pineal
body : PB, pituitary body: T, thalamencephalon.
I, olfactory nerve: II, optic nerve: III, third or motor oculi nerve :
IV, fourth nerve: V, fifth or trigeminal nerve: VI, sixth nerve: VII,
and VIII, combined root of facial and auditory nerves: IX and X, °
combined root of glossopharyngeal and pneumogastric nerves.

roof of the thalamencephalon through which the

vessels of the plexus pass into the third ventricle,
or cavity of the thalamencephalon. The thickened
sides of the thalemencephalon are the optic
thalami.
iv. The optic lobes are a pair of prominent ovoid bodies
G ,
 82 THE NERVOUS SYSTEM OF THE FROG.

touching each other in the median plane, and form-

ing the widest part of the brain: the pia mater
covering them is very strongly pigmented.
v. The cerebellum is a narrow transverse band im-
mediately behind the optic lobes.
vi. The medulla oblongata is the part of the brain
_ behind the cerebellum: it is widest in front and
gradually tapers towards its posterior end, where
it is continuous with the spinal cord. It is
covered by a triangular and very vascular mem-
brane, beneath which lies the fourth ventricle.

Fig. 16. A horizontal section through the brain of the frog, to show
the internal cavities. (From Ecker.)
Aq, ventricles of the optic lobes, and Sylvian aqueduct: Dv, third
ventricle: MF, foramen of Monro: Sv, lateral ventricle : Vv, fourth
ventricle.

b. The cavities of the brain.

Slice off the upper surface of the brain horizontally so as to
expose the several cavities or ventricles, without removing it from
 THE BRAIN. 83

the skull. These cavities are merely paris of, or outgrowths of, the
original central canal of the newral tube of the embryo. (Cf.
Chap. IX.) ue
i. The lateral ventricles extend through the whole
length of the cerebral hemispheres and a short
way into the olfactory lobes.
ii. The third ventricle is situated in the thalamen-
cephalon: it opens in front through the foramina
of Monro into the lateral ventricles: the stalk
of the pineal body: opens into it above ; and in
the hinder part of its floor is a conical depression,
the infundibulum.
iii. The aqueductus Sylvii or iter a tertioa quartum
ventriculum is a narrow passage leading from the
third to the fourth ventricle: it communicates
above with the cavities or ventricles of the optic
lobes, which are hollow.
iv. The fourth ventricle is the large triangular cavity
in the medulla, already exposed by removal of the
vascular: membrane covering it.
c. The ventral surface of the brain.
Cut through the medulla at the level of the hinder end of the
skull: carefully remove the brain from the cranial cavity, noting
the several nerves arising from it, and cutting through these as far
From the brain as possible. Lay the brain on its dorsal surface,
and examine and draw the ventral surface, showing the following
parts.
i. The optic chiasma is formed by the decussation
of the roots of the optic nerves; the point of
crossing being opposite the hinder ends of the
hemispheres, and immediately in front of the
infundibulum, |
Trace back the optic nerves behind their point of crossing to
their origins from the optic lobes.
ii, The tuber cinereum is a small median swelling
immediately behind the optic chiasma, caused by
the depression of the floor of the third ventricle
to form the infundibulum. It is divided by a
median groove into right and left halves.
84 THE NERVOUS SYSTEM OF THE FROG,

iii. The pituitary body is a flattened ovoid sac, lying

behind, and continuous with, the tuber cinereum.
It is almost certain to be left behind in the skull
on removing the brain, in which case the infundi-
bulum will be seen torn across.
iv. The crura cerebi are two dense white columns of
nervous matter, lying at the base of the optic
lobes, and partly hidden by the pituitary body:
they serve to connect the hemispheres with the
medulla and spinal cord.
v. The ventral fissure of the brain is a median longi-
tudinal groove on the ventral surface of the hinder
part of the brain: it is continuous with a similar
groove on the ventral surface of the spinal cord.

II. The Spinal Cord.

The spinal cord is a somewhat flattened band, presenting
brachial and lumbar enlargements opposite the points of origin
of the nerves for the fore and hind limbs respectively, and
slightly constricted between these two points. About the level
of the sixth or seventh vertebra the cord narrows rapidly to
form a fine thread, the filum terminale, which extends back into
the canal of the urostyle. |
The tubular character of the spinal cord is best seen on
making transverse sections of it. See p. 95.
B. The Peripheral Nervous System.
1. The Spinal Nerves. Ten pairs of nerves arise from the
sides of the spinal cord; each nerve arising by two roots, a
ventral or ‘‘anterior,” and a dorsal or “posterior,” which unite
at their point of exit from the vertebral canal through the
intervertebral foramen: just before their union the dorsal root
bears a ganglionic swelling.
Within the vertebral canal the roots of the anterior spinal
nerves run nearly transversely outwards, so as to leave the canal
opposite their points of origin from the spinal cord. The roots
of the middle and posterior nerves, owing to the vertebral
column being of greater length than the part of the cord
belonging to it, pass obliquely backwards to their points of exit:
and in the case of the hindmost nerves, the roots run backwards
 THE SPINAL NERVES. 85

within the vertebral canal some distance before reaching their

foramina of exit: the bundle formed by these roots, together
with the filum terminale, is spoken of as the cauda equina.

a. The spinal nerves outside the vertebral canal.

Lay the frog on tts back: cut through and pin out the body-
walls, and remove the abdominal viscera. Note the spinal nerves,
seen as white cords at the sides of the vertebral column. Clean
the nerves on one side and follow them to their distribution.
Each nerve divides, directly after the union of its two roots,
into a small dorsal branch, and a much larger ventral branch.
1, The hypoglossal, or first spinal nerve, leaves the
vertebral canal between the first and second ver-
tebre, and then runs forwards on the under sur-
face of the head beneath the mylohyoid and in
the substance of the geniohyoid muscle to the
chin, where it enters the tongue, in which it
ends. It supplies the muscles of the tongue and
floor of the mouth, and also some of the muscles
of the back and shoulder. (Fig. 15.)
2and 3. The second and third spinal nerves leave
the canal between the second and third, and
third and fourth vertebrze respectively : they
unite together to form the brachial nerve, which
gives off a large coracoclavicular branch to the
shoulder muscles and then runs down the arm,
supplying it with muscular and cutaneous
branches, and divides just above the elbow into
the radial and ulnar nerves, supply the
forearm and hand.
4,5, and 6. The fourth, fifth and sixth spinal nerves
are small and supply the muscles and skin of
the body-wall. They leave the vertebral canal
between the fourth and fifth, fifth and sixth, and
sixth and seventh vertebrze respectively.
7,8, and 9. The seventh, eighth, and ninth spinal
nerves together form the sciatic plexus. The
roots of these three nerves within the vertebral
86 THE NERVOUS SYSTEM OF THE FROG.

canal form the main part of the cauda equina.

The seventh nerve leaves the canal between
the seventh and eighth vertebre, the eighth
nerve between the eighth and ninth vertebre,
and the ninth nerve between the ninth or
sacral vertebra and the urostyle. Outside the
vertebral canal the three nerves unite together
opposite the middle of the urostyle to form
the sciatic plexus, from which branches are
given to the large intestine, bladder, oviducts,
etc. Just before joining the plexus the seventh
nerves give off the ileohypogastric and crural
nerves, supplying the muscles and skin of the
abdomen and thigh. Beyond the plexus is the
large sciatic nerve, which runs down the thigh,
giving branches to it, and dividing a short distance
above the knee into the tibial and peroneal nerves
supplying the leg and foot
The relative sizes of the nerves forming the
sciatic plexus, and the mode of their union with
one another are subject to considerable individual
variation in different frogs.
10. The coccygeal or tenth spinal nerve emerges
through a small hole in the side of the urostyle
near its anterior end. It gives branches to the
bladder, cloaca, and other adjacent parts.
b. The spinal nerves within the vertebral canal.
Cut away with scissors the centra of the vertebree one by one, to
expose the spinal cord and the roots of the spinal nerves from the
ventral surface. Note the following points :
i, The roots of the nerves: dorsal and ventral.
ii. The obliquity of the middle and posterior roots.
ii. The cauda equima, formed by the roots of the
hinder nerves together with the filum terminale,
iv. The ganglia on the dorsal roots as they pass
through the intervertebral foramina: these are
covered on their ventral surfaces by whitish cal-
careous patches, which form conspicuous objects
 THE SYMPATHETIC NERVOUS SYSTEM. 87

on either side of the vertebral column. Remove

these patches carefully to see the ganglia.
II. The Sympathetic Nervous System. This consists of a
longitudinal nervous band on each side of the body, connected
by branches with the several spinal nerves. The two main sympa-
thetic trunks lie, in front, close to the dorsal surface and alongside
the vertebral column: further back they are in close relation
with the dorsal aorta, alongside which they run.
Each sympathetic trunk receives a branch from each of the
spinal nerves of its side, and at the junction of each of these
branches with the main trunk there is a ganglionic enlargement.
The coccygeal or tenth spinal nerve, unlike the others, is
connected with the sympathetic by more than one branch:
the actual number of these branches is not constant, but is
said to vary from two up to as many as twelve.
From the sympathetic ganglia nerves are given off to the
bloodvessels and viscera, the chief ones being the following:
i. The cardiac plexus is formed by nerves arising
from the first sympathetic ganglion: the plexus
is a meshwork of nerves on the auricles, and
around the great vessels at their openings into
the heart.
ii. The solar plexus lies on the dorsal surface of the
stomach : the nerves are derived mainly from the
third, fourth, and fifth ganglia.
Hepatic, renal, genital, hemorrhoidal, and vesical plexuses
also exist in connection with the liver, kidney, reproductive
organs, large intestine, and bladder respectively.
III. The Cranial Nerves. There are ten pairs of cranial
nerves in the frog, which are numbered in order from before
backwards. (See Fgures 13 and 15).
To dissect the cranial nerves expose the brain by removing the
roof of the skull as already described, and then follow the special
instructions given in the case of the more important nerves.
1. The olfactory nerve, the special nerve of smell, arises
from the anterior end and outer side of the olfactory lobe,
and is distributed to the membrane lining the nasal
cavity.
88 THE NERVOUS SYSTEM OF THE FROG,

To see the course and distribution of the olfactory nerve, dissect

from the dorsal surface, removing the roof of the anterior part of
the skull, including the nasal bone.
2. The optic nerve, the nerve of sight, arises from the side
of the brain just below the optic lobe, partially crosses
over at the optic chiasma on the under surface of the
brain, and then runs outwards to the eyeball.
The course of the optic nerve has been fully seen in previous
dissections,
3. The motor oculi is a small nerve arising from the ventral
surface of the brain close to the median line and between
the crura cerebri. It supplies four of the muscles

Fig. 17. The trigeminal, facial glossopharyngeal, and pneumogastric

nerves of the frog, dissected from the right side.
a, stomach: e, Eustachian tube: h, hypoglossal nerve: 7, lung:
m, second spinal nerve: s, sinus venosus: ¢, squamosal: ¢.a, truncus
arteriosus : wu, right auricle: v, ventricle: v.c, right anterior vena cava:
Va, ophthalmic branch of trigeminal: Vb, maxillary branch of trigemi-
nal: Ve, mandibular branch of trigeminal: VIIa, palatine branch of
facial: VIIb, hyoidean branch of facial: IX, glossopharyngeal :
Xa, laryngeal branch of pneumogastric : Xb, pulmonary branch of
neumogastric : Xc, cardiac branch of pneumogastric: Xd, gastric
Van of pneumogastric : 2, transverse process of second vertebra:
3, transverse process of third vertebra.
 THE CRANIAL NERVES. 89

moving the eyeball; wz., the rectus superior, rectus

internus, rectus inferior, and obliquus inferior.
Owing to its small size the third nerve is not easily made out
an the frog.
4, The fourth or pathetic nerve is a very slender nerve,
arising from the dorsal surface of the brain between the
optic lobe and the cerebellum, and supplying the obliquus
superior muscle of the eyeball.
The fourth nerve is too small to be dissected satisfactorily in the
frog.
5. The trigeminal is the largest of the cranial nerves in the
frog. It arises from the side of the anterior part of the
medulla, and runs outwards and forwards to the skull
wall: just before reaching this it expands into a large
swelling—the Gasserian ganglion. It then passes
through the skull wall immediately in front of the
auditory capsule, and divides at once into two main
branches.
i. The ramus ophthalmicus runs forwards through the
orbit, lying close to its inner side, between the
skull wall and the eye. It passes beneath the
rectus superior, but above all the other muscles
of the eyeball and the optic nerve. At the
anterior end of the orbit it divides into two
branches which pass through the walls of the
nasal capsule, and supply the skin of the fore
part of the head.
To trace this branch, dissect from the dorsal surface: cut away
carefully with scissors the side wall of the cranium: cut through
and turn aside the rectus superior, and find the nerve running close
alongside the skull wall, between it and the eyeball. Trace it for-
wards to the nose.
ii. The ramus maxillo-mandibularis runs directly
outwards behind the eyeball, in front of the
auditory capsule and between the temporal and
pterygoid muscles, After a very short course it
divides into the maxillary and mandibular nerves.
To trace this nerve and tts branches remove the squamosal bone
carefully, and find the nerve lying on the pterygoid muscle and
90 THE NERVOUS SYSTEM OF THE FROG.

immediately behind the eye. Follow the nerve behind the pterygoid
and temporal muscles to the skull, removing the muscles if necessary;
and then trace the branches outwards to their distribution.
a, The ramus maxillaris runs forwards and out-
wards in the floor of the orbit, behind and
below the eyeball, to the margin of the upper
jaw, which it reaches about midway along
its length: it then ends in branches which
run along the jaw, some forwards and some
backwards, supplying the upper lip, the lower
eyelid, and other neighbouring parts.
GB. The ramus mandibularis runs parallel to
and behind the ramus maxillaris as far as
the outer border of the eyeball, giving
branches to the temporal and pterygoid
muscles: it then turns backwards, outwards,
and downwards, and passing across the inner
side of the upper jaw, reaches the outer
surface of the mandible just behind the
insertion of the temporal muscle: it then
runs forwards along the outer side of the
lower jaw to the chin, supplying the lower
lip and the muscles of the floor of the mouth.
6. The abducens is a very slender nerve which arises from
the ventral surface of the medulla close to the median
line, and a short way behind the pituitary body. It
passes either through, or in very close contact with, the
Gasserian ganglion, and entering the orbit supplies the
retractor bulbi and the rectus externus muscles.
The abducens nerve is too small to be made out satisfactorily in
the frog.
7. The facial nerve arises from the side of the medulla
immediately behind the trigeminal nerve, and passes
forwards to the skull wall, where it is very closely con-
nected with the Gasserian ganglion. It passes through
the skull wall immediately behind aud in close company
with the trigeminal nerve, and divides at once into its
two main branches.
i. The ramus palatinus runs forwards in the floor
of the orbit a short distance from the side wall
 THE CRANIAL NERVES. 91

of the skull, and immediately upon the mucous

membrane of the roof of the mouth. Near the
anterior end of the orbit it divides into two
branches, one of which runs outwards and
anastomoses with the ramus maxillaris of the
trigeminal nerve, while the other runs forwards
to the anterior part of the roof of the mouth. It
supplies the mucous membrane of the roof of
the mouth.
Dissect this nerve from the ventral surface. Cut away the lower
jaw: carefully remove the mucous membrane .of the roof of the
mouth, and find the nerve lying on the ventral surface of the eye-
ball and its muscles, and running parallel to and a short distance
from the skull-wall. Trace it backwards and forwards.
ii. The ramus hyomandibularis runs outwards and
backwards round the front end of the auditory
capsule; it then crosses over the inner end of the
columella and turas downwards in the posterior
wall of the Eustachian tube to the angle of the
mouth, giving branches to the tympanic mem-
brane and to the articulation of the mandible.
It then divides into two branches.
The exposure of the above nerve, which is not easy, may be
performed thus :—remove the shoulder-girdle of one side ; also the
depressor mandidule and temporalis muscles: open the cranial
cavity as before, to expose the brain: remove the tympanic
membrane and clean the outer end of the columella, Cut away
carefully the roof of the auditory capsule by a horizontal cut, just
above the level of the columella: find the facial nerve running
round the front end of the auditory capsule and in close contact
with tt, and trace it back over the columella and down to the angle
of the mouth.
a. The ramus mandibularis runs forwards in the
floor of the mouth, lying along the inner edge
of the lower jaw and between the mylohyoid
muscle and the skin, as far forward as the
chin.
Dissect from the ventral surface: remove the skin from the
under surface of the floor of the mouth, and find the nerve running
along the inner border of the mandible.
93 THE NERVOUS SYSTEM OF THE FROG.

(3. The ramus hyoideus is the posterior and larger

of the two branches: it runs forwards in the
floor of the mouth along the anterior cornu
of the hyoid, supplying its muscles.
8. The auditory nerve, the nerve of hearing, arises from the
side of the medulla immediately behind and in close
contact with the root of the facial nerve: it enters the
auditory capsule and ends in the internal ear.
9. The glossopharyngeal nerve arises from the side of the
medulla behind the auditory nerve, by a root common
to it and the tenth nerve: it leaves the skull by an
aperture immediately behind the auditory capsule, and
divides behind the capsule into two branches.
i. The ramus anterior runs downwards and forwards
round the hinder border of the auditory capsule
and beneath the depressor mandibule muscle to
join the facial nerve just after it has crossed over
the columella.
The dissection already made for the ramus hymondibularis of
the facial nerve will show also the above branch of the glosso-
pharyngeal.
ii. The ramus posterior runs downwards and _for-
wards to the ventral wall of the pharynx, passing
beneath the fourth division of the petrohyoid
muscle but superficial to the others; it runs
just behind and parallel to the anterior cornu of
the hyoid. On reaching the floor of the mouth
it crosses the hypoglossal nerve obliquely, lying
dorsal to it, and then runs forwards in a
peculiarly sinuous course, close to the middle
line and between the geniohyoid and hyoglossus
muscles, to the base of the tongue, which it enters
and in which it ends. It supplies the petrohyoid
muscle, and the mucous membrane of the pharynx
and tongue.
The exposure of the first part of the above nerve is best performed
from the side, andis much facilitated by distending the esophagus
and pharynx with a cork or roll of paper. Its course along the
floor of the mouth to the tongue should be dissected from the
ventral surface.
 THE CRANIAL NERVES. . 93

10. The pneumogastric or vagus nerve arises, as already

noticed, in common with the glossopharyngeal. It
leaves the skull by the same aperture as the ninth nerve,
and immediately outside the skull presents a ganglionic
enlargement : it gives off dorsal branches to the muscles
of the back, and then runs backwards and downwards
round the side wall of the pharynx, running along the
hinder border of the fourth division of the petrohyoid
muscle: behind this muscle it divides into its main
branches, which are as follows:
i. The ramus laryngeus loops round the posterior
cornu of the hyoid and round the pulmocutaneous
artery close to its origin from the aortic trunk :
it then passes inwards, dorsal to the artery, to the
middle line, where it ends in the larynx.
ii, The ramus cardiacus passes dorsal to the pul-
monary artery to the interauricular septum of
the heart, and to the sinus venosus.
ii. The rami pulmonales follow the course of the.
pulmonary artery to the lung, in which they end.
iv, The rami gastrici, usually two in number, run
through the partial diaphragm formed by the
anterior fibres of the obliquus internus muscle,
and end in the walls of the stomach,
Lhe dorsal portions of the several branches of the pneumogastric
nerve are best exposed from the side; to see them properly, the
shoulder girdle and fore limb must be removed and. the esophagus
well distended : the terminal branches must be dissected From the
ventral surface,

IV. The Cranial Portion of the Sympathetic Nervous

System.
The main sympathetic trunk of each side extends forwards in
front of the first ganglion, and enters the skull at the foramen
in the exoccipital bone through which the glossopharyngeal
and pneumogastric nerves pass out: it is connected with the
pneumogastric nerve, and then runs forwards within the skull to
the Gasserian ganglion of the trigeminal nerve, in which it ends.
 94 THE NERVOUS SYSTEM OF THE FROG,

C. Histology of Nerves.
Nervous matter consists histologically of elements of two
kinds, nerve cells and nerve fibres. The nerve cells are
branching nucleated cells, connected by their processes with one
another and with the nerve fibres. ‘The nerve cells are the
centres whence impulses originate, while the nerve fibres serve
to convey these impulses from one part to another. A local
accumulation of nerve cells is called a ganglion.
1. Nerve Fibres are of two kinds, medullated and non-
medullated.
a, Medullated nerve fibres form almost the whole of the
cranial and spinal nerves, and a large part of the brain
and spinal cord.
Take a small piece of the sciatic or some other nerve from a
freshly killed frog: spread tt out and tease tt in a drop of normal
salt solution: examine with low and high powers - note
i. The nerve fibres: unbranched.
ii. The perineurium, or connective tissue binding the
nerve fibres into bundles, or ‘ nerves.”
In each nerve fibre note
iii. The primitive sheath, or sheath of Schwann: a
very delicate external investment, seen with
difficulty, and only in certain places.
iv. The medullary sheath: a thick fatty layer within
the primitive sheath ; it is highly refractive, and
gives the nerve fibre its double contour.
Tease in glycerine a small piece of nerve that has been treated
with osmic acid: examine with the high power a single nerve
fibre ; note the following points.
i. The medullary sheath is stained darkly in conse-
quence of its fatty nature: it is interrupted at
intervals by the nodes of Ranvier.
ii, The nodes of Ranvier are spots where the medul-
lary sheath is absent, and the primitive sheath
forms constrictions touching the axis cylinder.
iii, The axis cylinder is the central cylindrical rod,
the essential part of the nerve fibre: it is clearly
 HISTOLOGY OF NERVES. 95

visible at the nodes, and is much less deeply

stained than the medullary sheath.
iv. Nuclei are seen projecting into the medullary sheath
about midway between the nodes.
Tease a small piece of fresh nerve in chloroform : this will
partially dissolve the fatty medullary sheath and so render the
' primitive sheath and axis cylinder more clearly visible: note -—
i. The primitive sheath, or sheath of Schwann.
ii. The axis cylinder.
b. Non-medullated nerve fibres occur chiefly in the sympa-
thetic nerves: they branch and anastomose; and they have
no medullary sheath.
2. Nerve cells: Tease in glycerine a small fragment of the
ventral cornu of the spinal cord of the ox (lumbar region) : cover,
and examine with low and high powers: note
i. The nerve cells: large nucleated cells with many
branching processes.
ii. The nerve fibres.
ili, The fine connective tissue binding the several parts
together.
3. Structure of the Spinal Cord.
Take one of the prepared transverse sections of spinal cord of
Jrog ; mount in balsam, and exanune with low and high powers :
note the following points.
a. With the low power.
i. The section is bilaterally symmetrical, and oval in
shape; the transverse diameter considerably ex-
ceeding the vertical.
ii, The ventral or “anterior” fissure is a broad and
shallow median cleft: there is in the frog no
distinct dorsal fissure.
iii. The white matter forms the outer part of the cord,
and is chiefly composed of medullated nerve fibres.
iv. The grey matter forms the central part of the cord,
and is composed of a dense network of non-
medullated nerve fibres, in which are imbedded
numerous nerve cells.
96 THE NERVOUS SYSTEM OF THE FROG.

y. The cornua are the processes, ventral or ‘‘anterior,”

and dorsal or ‘posterior,’ into which the grey
matter is produced on either side.
vi. The central canal of the cord lies in the median
plane, nearer the ventral than the dorsal surface.
vii. The nerve roots are only seen if the section happens
to pass through their points of origin.
a. The dorsal or “ posterior” root is connected with
the dorsal cornu of the grey matter; it is a
single thick band of nerve fibres.
B. The ventral or “anterior” root is connected
with the ventral cornu of the grey matter ; it
consists of a number of very slender bands of
nerve fibres.
b. With the high power.
i. The ganglion cells are large nucleated branched
cells lying in groups in the grey matter: they
are largest and most numerous in the ventral
cornua.
ii. The neuroglia is a delicate network of connective
tissue fibres and cells, penetrating and supporting
all parts of the cord, and continuous at the sur-
face with the pia mater.
iii. The pia mater is the delicate connective tissue
membrane closely ensheathing the cord.
iv. The bloodvessels of the cord are small and numerous:
they enter from the pia mater.
v. The central canal is lined by a ciliated epithelium.
 97

CHAPTER VII.

THE EYE AND EAR.

A. The Eye of the Frog.

1. Remove the eye from a freshly killed frog : snip off with
scissors the muscles of the eyeball - note the following points,
i. The shape. The eyeball is flattened on its outer
side, more convex on the inner or deeper side.
ii. The sclerotic is the firm outer wall of the eyeball,
formed of hyaline cartilage, and dense white
connective tissue.
ili. The cornea is the transparent patch on the outer
side of the eye through which the light enters:
it is continuous at its margin with the sclerotic.
iv. The iris is a pigmented ring placed behind the
cornea and seen through it: it acts as a dia-
phragm, limiting the amount of light that enters
- the eye.
v. The pupil is the aperture surrounded by the iris,
which serves to admit the light to the interior
of the eye.
vi. The optic nerve is seen piercing the sclerotic to
enter the eyeball on its inner side.
2. Place the eye under water and divide it with scissors into two
halves by a cut passing through the middle of the cornea and
through the sclerotic close to the optic nerve, so as to lay open com-
pletely the interior of the eye: note the following points.
i, The lens is a firm solid transparent body, just
behind the iris and attached to its outer margin:
it is more convex on its inner than on its outer
surface.
98 THE EYE AND EAR.

ii. The anterior chamber of the eye is the space between

the cornea and the lens: it is small and contains
the aqueous humour, a watery fluid.
iii. The posterior chamber of the eye is the large space
behind the lens: it is filled by the vitreous
humour, a gelatinous body.
iv. The choroid is the black pigmented layer lining
the sclerotic, and continuous in front with the
iris.
vy. The retina is a delicate transparent membrane
lining the eyeball. It is readily detached from
the choroid, except at the point where the optic
nerve enters.

B. The Eye of the Sheep or Ox.

1. Dissect off the muscles of the eyeball, and the fat which sur-
rounds the optic nerve; noté the following points.
i. The shape. The eyeball is more spherical than in
the frog.
ii. The sclerotic covers about five-sixths of the eyeball:
it is tough, white, and opaque.
iii. The cornea, which covers the outer sixth of the
eyeball, is circular, transparent, and continuous at
its margin with the sclerotic: it is more convex
than the sclerotic.
iv. The conjunctiva is a delicate epithelial layer, cover-
ing the front of the cornea and part of the
sclerotic.
v. The iris is the oval pigmented ring seen through
the cornea.
vi. The pupil is the central, oval or dumb-bell shaped
aperture surrounded by the iris.
vii. The optic nerve is a thick white bundle of nerve
fibres piercing the sclerotic at the back of the
eye.
 THE EYE, 99

2. Cut all round the cornea, with stout scissors, about 4 inch
from its junction with the sclerotic: remove the cornea: take care
not to squeeze the eye, or the lens will be driven out instantly: note:
i, The aqueous humour: the transparent watery fluid
filling the anterior chamber of the eye, and
escaping when the cornea is removed
ii The lens.

Fig. 18. A diagrammatic section through the human eye passing

through the centres of the cornea and lens, and through the yellow spot
and point of entrance of the optic nerve.
A, anterior chamber: Ar, central artery of retina: B, blind spot:
=
C, cornea: Ch, choroid: CP, ciliary processes: H, hyaloid membrane,
enclosing the vitreous humour: |, iris: L, lens: O, optic nerve: P, posterior
chamber : R, retina : S, sclerotic : SL, suspensory,ligament: Y, yellow spot.
3. Pass the handle of a scalpel under the cut edge of the cornea,
between it and the iris, and carefully separate the sclerotic From
the choroid the whole way round for a distance of about half an
inch beyond the edge of the cornea. Make four radial cuts, equt-
distant from one another, through the margin of the cornea and
100 THE EYE AND EAR.

the sclerotic, taking care not to injure the deeper parts; and
extend the cuts back towards the optic nerve. Carefully peel off
the four flaps into which the sclerotic is now divided from the
underlying black choroid coat: turn them down, and pin them to
the dissecting board so as to fia the eye with the iris upwards » note
the following points.
i. The ciliary muscle is a whitish ring of unstriped
muscle connecting the outer margin of the iris
with the junction of the cornea and sclerotic.
ii. The choroid is the dense black coat exposed by the
removal of the sclerotic.
iii, The ciliary vessels pierce the sclerotic to convey
blood to and from the choroid, which is extremely
vascular.
iv. The ciliary nerves are seen passing through the
sclerotic to the choroid while the flaps are being
turned down.
4. Make a couple of radial incisions, about 4 inch apart, through
the iris and ciliary muscle, and turn back the portion of the tris
between the two cuts, so as to expose its hinder surface.
i. The ciliary processes are a series of densely pig-
mented and close set radial folds on the hinder
surface of the outer margin of the iris: they fit
into corresponding folds in the ligament which
surrounds and supports the lens.
5. With a large pair of scissors cut the eye into two halves by a
horizontal inciston at tts equator. Turn over the anterior or outer
half, and examine tt from behind : note the following parts.
i. The ciliary processes.
ii. The uvea is the layer of dense black pigment at
the back of the iris and ciliary processes.
iii. The ora serrata is the indented anterior boundary
of the part of the retina sensitive to light:
in front ofthis the retina becomes extremely
thin, but really extends forwards as far as the
free edge of the iris.
6. Turn the anterior half of the eye over, so that its outer or
corneal surface is uppermost: cut away the tris completely : note
the following points,
 THE EYE. 101

i. The capsule of the lens is an elastic transparent

membrane holding the lens in its place.
_ii. The suspensory ligament of the lens, or zonule of
Zinn, is the outer margin of the capsule of the
lens: it is marked with radiating folds into
which the ciliary processes fit.
iii, The cut edges of the retina and choroid should be
recognised.
7. Remove the lens from tts capsule ;note its shape, more convex
behind than in front: harden tt with spirit, or by boiling for a
Jew minutes in water.
8. Remove the vitreous humour from the posterior half of the
eye: note the following points.
i. The retina is a delicate pulpy membrane between
the vitreous humour and the choroid.
ii. The blind spot is the point of entrance of the optic
nerve: the retina adheres firmly to this spot,
though it can be readily separated from the
choroid at all other parts.
iii, The retinal vessels enter with the optic nerve, and
radiate from the blind spot.

C. Histology of the Eye.

1, The Choroid. Spread a small piece of fresh choriod on a
slide in normal salt solution + examine with low and high powers.
i, The Choroid is a network of bloodvessels thickly
invested by pigment cells.
ii. The pigment cells are irregularly branched, with
clear nuclei.
2. The Lens. Tease in glycerine a small piece of lens, hardened
by boiling ; examine with low and high powers : note :—
i. The laminated character of the lens as a whole.
ii. The elongated epithelial cells of which the lens is
composed,
iii, The serrated edges of many of the cells.
102 THE EYE AND EAR,

3. The Retina.
Mount in balsam one of the prepared sections of the posterior
part of the frog’s eye: examine with low and high powers,

PG

BC

NF

Be
Fig. 19. Vertical section through the posterior wall of the eye of a
frog: the section passes through the sclerotic, the choroid, and the entire
thickness of the retina. 300.
BC, red blood corpuscle: C, cone: G, nerve cell: IL, inner limiting mem-
brane : IM, inner molecular layer: IN, inner nuclear layer: NF, layer of
nerve fibres: OL, outer limiting membrane: OM, outer molecular layer:
ON, outer nuclear layer: P, pigment cell : PC, pigment of choroid : R, rod :
RF, radial or Miiller’s fibre: Ss, the cartilaginous sclerotic.
 THE EYE. 103

a. The sclerotic consists chiefly of hyaline cartilage.

b. The choroid is a vascular plexus, with much pigment.
c. The retina is composed of the following layers from
without inwards.
i, A layer of pigment cells, sending processes between
the rods and cones.
ii, The rods and cones are a layer of columnar bodies
placed vertically to the surface: the rods are
far more numerous, and much larger than the
cones: each consists of an inner and an outer
segment.
iii, The outer limiting membrane is seen in sections
as an exceedingly thin line separating the rods
and cones from the outer nuclear layer.
iv. The outer nuclear layer is a moderately thick layer,
well stained, and very closely connected with the
rods and cones.
a The outer molecular layer is a very thin layer,
not stained: it is finely granular or reticular in
structure.
vi. The inner nuclear layer is thick and well stained.
It consists of several layers of large nuclei,
surrounded by thin layers of protoplasm, produced
into radial processes.
vil. The inner molecular layer is a thick, finely granular
layer, not stained.
vill, The layer of nucleated nerve cells is a single layer
of large branched cells.
ix, The layer of nerve fibres is formed by the branches
of the optic nerve.
a The inner limiting membrane.
xi. The radial fibres, or Muller’s fibres, commence with
expanded ends in the layer of nerve fibres and
stretch outwards: they can easily be traced
through the inner molecular layer. They consist
of connective tissue and serve to strengthen and
Support the retina.
104 THE EYE AND EAR.

D. The Ear of the Frog.

The frog’s auditory organ is too small to dissect satisfactorily,
and is best studied by making transverse sections of the entire
head, in the following manner.
Kull a frog with chloroform; cut off the head with stout scissors,
and decaleify wt by placing im a 5 to 10 per cent. solution of
nitric acid, or in a meature of chromic acid with a few drops of
nitric acid. When the bones are thoroughly soft, which will take
From a few hours to 3 or 4 days or more according to the strength of
acid employed, remove the head from the decalcifying solution and
transfer to weak alcohol and thence to strong alcohol. Then stain
with borax carmine and imbed in paraffin, and cut into transverse
sections with a microtome. Mount the sections in series ; examine
and draw them, showing the following points.
1. The periotic capsule consists mainly of cartilage, and is
firmly fused with the hinder part of the cranium.
2. The vestibule is a membranous sac lying in the cavity of
the periotic capsule, and filled with a watery fluid, the
endolymph: it is partly divided by a constriction into
two main divisions.
i. The utriculus is the upper and larger division.
ii, The sacculus is the inferior and smaller division:
from it arise three small saccular dilatations,
supposed to represent the cochlea of higher
animals,

Fig. 20. The right internal ear of the frog, removed from the periotic
cartilage and drawn from the outer surface.
a, the anterior vertical semicircular canal: 6, its ampulla: Ah, the
horizontal canal: 7, its ampulla: p, the posterior vertical canal: 7, its
ampulla: s, the sacculus: w, the utriculus.
 THE EAR. 105

3. The semicircular canals are three tubular offsets of the

vestibule, into which they open at both ends. They lie
in canals in the periotic cartilage, and are placed in
planes at right angles to one another: each has at one
end, close to its opening into the vestibule, a dilatation
or ampulla,
i. The anterior vertical canal has its ampulla at its
anterior end.
ii. The posterior vertical canal has its ampulla at its
outer end, while its inner end joins the posterior
end of the anterior vertical canal to open into the
vestibule by a common orifice.
iii, The horizontal or external canal has its ampulla
at the anterior end.
4. The auditory nerve leaves the cranial cavity through a
hole in the inner wall of the periotic capsule, and divides
into branches distributed to the sacculus and its diverti-
cula, and to the ampullee of the semicircular canals.
5, The accessory auditory apparatus. The essential organs
of hearing—z.e., the vestibule and its offsets, and the
auditory nerve—are enclosed in the periotic cartilage,
which is deeply placed in the side of the head: the com-
munication with the surface is brought about by the
accessory apparatus, which consists of the following parts
(Fig. 11 p. 58).
i. The Eustachian passage and tympanic cavity are
formed in connection with the hyomandibular
gill cleft of the tadpole. See Chapter IX.
ii. The tympanic membrane closes the tympanic cavity
on its outer side.
iii, The columella is a rod of bone and cartilage, the
outer end of which is attached to the tympanic
membrane, while its inner end is inserted into a
hole in the outer wall of the periotic capsule so
as to lie in close contact with the vestibule. It
serves to communicate the auditory vibrations
of the tympanic membrane to the vestibule.
106

CHAPTER VIII.

THE REPRODUCTIVE ORGANS AND THE CLOACA OF -

THE FROG.

A. The Male Frog.

1. The Reproductive Organs. (Fig. 3, p. 18.)
Pin the frog on tts back under water : open the body cavity from
the ventral surface: turn aside the alimentary canal and liver.
i. The testes are a pair of yellow ovoid bodies about
half an inch long, lying on the ventral surface of
the kidneys. Within the testes are developed the
essential male elements or spermatozoa.
ii. The vasa efferentia are a number—usually 10 to
12—of slender ducts, connecting the testis of
each side with the inner or median border of the
corresponding kidney: they serve to convey the
spermatozoa from the testis into the tubules of
the kidney, whence they escape by the ureter,
which acts as vas deferens.
iii. The vas deferens or ureter runs along the outer
side of the posterior part of the kidney, and then
backwards to the cloaca.
iv. The vesicula seminalis is a large pouch-like dilata-
tion on the outer side of the vas deferens, just
behind the kidney and before reaching the cloaca.
2. The Cloaca. (Cf. Fig. 3, p. 18).
Lay the frog on its back: with a stout scalpel split the pelvic
symphysis in the median plane : gently separate the two halves, and
pin them out right and left, so as to expose the cloaca from the
ventral surface.
 THE REPRODUCTIVE ORGANS, 107

i, The cloaca is the terminal portion of the large

intestine, into which the ureters and the bladder
open. - |
ii. The bladder is a thin-walled bilobed muscular sac,
lying on the ventral surface of the large intestine
and cloaca, its two lobes communicating freely
with each other. It is invested by peritoneum
and attached to the sides of the body by special
peritoneal folds.
Intlate the bladder with a blow-pipe inserted through the cloacal
aperture: pass a seeker up the cloaca to determine the exact position
of the opening from the bladder to the cloaca. Cut up the cloaca
along one side: wash out its contents and examine the opening into
the bladder.
iii, The ureter or vas deferens is continued behind
the vesicula seminalis as a very short tube, open-
ing into the dorsal wall of the cloaca almost
exactly opposite the opening of the bladder on
the ventral surface. The openings of the two
ureters are close together on the apices of two
small papille, overhung by a slight valvular pro
jection of the mucous membrane of the cloaca.
iv. Lemove a small piece of the testes: place it on a slide
in a drop of salt solution: press it slightly: cover and
examine with a high power to see the spermatozoa.

B. The Female Frog.

1. The Reproductive organs.
Dissect as in the male.
i. The ovaries are a pair of black masses lying in folds
of the peritoneum ventral to the kidneys, in very
much the same position as the testes in the male.
Their shape and size vary much at different
seasons of the year. On their surfaces are
- numerous rounded projections, like small shot ;
these are ova in various stages of development:
the smaller and younger ones are white; the
larger and more mature ones black in one half,
and white or yellowish in the other. Each ovary
108 REPRODUCTIVE ORGANS AND CLOAGA.

consists of a couple of folds united. along their

ventral edges; the space between the folds is
divided by partitions into about fifteen pouches.
ii. The oviducts are a pair of white, twisted tubes,
with thick gelatinous walls. They commence
with open mouths at the extreme front end
of the body cavity, close to the outer side of
the roots of the lungs; and run back, increasing
in size, and becoming much convoluted. Their
hinder ends are greatly dilated, but have thinner
walls. Unlike the male, the female has genital
ducts distinct from the ureters.
2. The Cloaca.
Dissect as in the male.
i. The cloaca is very similar to that of the male,
except that the urinary and genital products are
discharged into it by separate ducts.
ii. The bladder is like that of the male.
iii. The oviducts open separately into the dorsal wall
of the cloaca, just opposite the bladder, by two
wide apertures separated by a narrow median
partition.
iv. The ureters open by two small apertures, placed
close together, into the dorsal wall of the cloaca
just behind the oviducts.
 109:

CHAPTER IX.

THE DEVELOPMENT OF THE FROG.

I. General Account.
The frog’s eggs are laid in water, usually during March or
the early part of April.
During the act of oviposition, which may last several days,
the male frog clasps the female firmly, embracing her with his
arms; and as the eggs pass out from the cloaca of the female
they are fertilised by spermatozoa discharged over them by the
male.
The eggs, which are very numerous, are small spherical bodies.
about 1°75 mm. in diameter; they are invested by thin coatings
of an albuminous substance, which swell up very greatly in the
water, and stick together to form the bulky masses we call
frog’s spawn. Such spawn consists of a transparent gelatinous.
mass, formed by the swollen albuminous matter, in which are:
embedded the eggs: these latter appear as small round bodies,.
-each presenting a black half and a white half.
If a number of hen’s eggs were broken into a basin, care:
being taken not to rupture the yolks, a mass would be produced
similar to frog’s spawn: the yellow yolks corresponding to the
frog’s eggs, and the whites or albuminous investments of the
yolks to the gelatinous matrix of the spawn. And just as the
chicken is formed from the yolk, and not from the white of a
hen’s egg, so also is the frog developed from the egg and not
from the gelatinous investment.
The frog’s eggs, laid in this way and fertilised by the sperma-
tozoa shed over them by the male, begin to develop at once.
Each egg is at first spherical, but in about a week becomes ovoid
in shape, and then rapidly increases in length. By the tenth
day it is divided by slight constrictions into head, body, and
tail. The whole animal becomes fish-like in appearance, the
tail growing rapidly: two pairs of branching tufts, the external.
 OF
THE110
DEVELOPMENT
FROG.

‘Sly"1G SNOTIVASoHe4S
UL oY} JUoUIdooA
Jo oy ep “sor WoI) SMYysIg (‘wagapiary,]
‘T ‘soonqysnl "prey ‘z, ‘soBqq A]qIOYsS
Jaqze ‘SUIAYT
‘e ‘opodpey,Apyaroys asogoq
qsnl “poyozey
“suryoye
“F y ‘sojodpey,
e puv‘9 sojodpey
YIM, [eUto}xXe
‘s]jIs
f pue‘8 sopodpey
YAM, AT[NF
¢ pure ‘OT poutsoy Ae]No1ed
‘SPl[Of
o
sojodpey
Y4IM
, [Tom pedojeasp ‘sso]-puryATZIOYS a1ofeq
ot4
ety ‘sIsoydi
‘TT oweqouropodpey, sulinp
‘sISoydi
‘zy oureqout
SunoXsory YFIMoy4 Trey Ayuo ATperyred ‘poqaosqe
 DEVELOPMENT OF THE FROG Tit

gills, followed shortly by a third pair, grow out from the sides
of the neck, and in about a fortnight from the time of laying of
the eggs the young tadpoles make their way out of the gelatinous
mass of the spawn, and swim freely in the water.
At the time of hatching the tadpole has no mouth, and is
dependent for food on granules of food yolk which are contained
in large numbers in the egg, and at the expense of which all the
earlier processes of development are effected. A horse-shoe
shaped sucker is present on the under surface of the head, by
which the tadpole is enabled to attach itself to weeds or other
objects in the water.
A few days after hatching the mouth appears, bordered by a
pair of horny jaws, and fringed with fleshy lips provided with
horny papille. The alimentary canal which has hitherto been
wide and short, now rapidly increases in length, becoming
tubular and convoluted; the liver and pancreas are formed; the
anus is developed even before the mouth, and the tadpole now
feeds eagerly on conferve and other vegetable matter.
About the time of appearance of the mouth i.e. shortly
after hatching, a series of four slit-like openings, the gill clefts,
appear on each side of the neck, leading from the pharynx to the
exterior. The margins of these slits become folded, and form
the internal gills; the external gills at the same time decreasing
in size and assuming a shrivelled appearance.
While the internal gills are developing, a fold of skin, the
operculum, appears on each side of the head in front of the gills,
and grows backwards over these, so as to enclose them in a gill
chamber. ‘Towards the end of the fourth week the hinder edges
of the opercular folds fuse with the body wall on the ventral
surface and along the right side. On the left side a spout-like
opening remains which communicates with the gill chambers of
both sides, and through which the water taken in at the mouth
for respiration, and passed through the gill slits, makes its
escape to the exterior.
During this time the tadpole has been feeding freely, and
has increased greatly in size. The body, Fig. 21, s, is broad and
round; the tail is much larger than before, and forms a powerful
Swimming organ; while the sucker on the under surface of the
head, though still present, is small and but little used.
Very shortly afterwards rudiments of the hind-limbs can be
seen as a pair of small papille at the root of the tail, one on
113 DEVELOPMENT OF THE FROG.

each side of the anus: these steadily increase in size: about

the seventh week they become divided into joints; and a week
later the toes appear.
The fore-limbs arise about the same time as the hind ones,
but are covered by the opercular folds, and hence do not become
visible till a later stage.
Towards the end of the second month the lungs come into
use, and the tadpoles which now have the form shown in Fig. 21,
9 and 10, frequently come to the surface of the water to breathe.
The gills now begin to degenerate, but for a time respiration is
effected both by the gills and the lungs.
A fortnight or three weeks later a distinct metamorphosis
occurs, whereby the tadpole becomes transformed from the fish-
like condition in which it has hitherto remained to the purely
air-breathing stage characteristic of the adult. The tadpole
ceases to feed: a casting, or ecdysis, of the outer layer of the.
skin takes place: the gills are gradually absorbed: the horny
jaws are thrown off; the large frilled lips shrink up; the mouth
loses its rounded suctorial form and becomes much wider; the
tongue, previously small, increases considerably in size; the
eyes become larger and more prominent ; the fore-limbs appear,
the left one being pushed through the spout-like opening of the
branchial chamber, and the right one forcing its way through
the opercular fold, in which it leaves a ragged hole. The
abdomen shrinks ; the stomach and liver enlarge, but the intes-
tine becomes considerably shorter than before, and of smaller
diameter: the animal, previously a vegetable feeder, now
becomes carnivorous. The gill clefts close up; and important
modifications accompanying the change in breathing, occur in
the bloodvessels.
The tail, which is still of great length, Fig. 21, 11, now begins
to shorten, and is soon completely absorbed: the hind-limbs
lengthen considerably, and the animal leaves the water as a Frog.
Explanation of the metamorphosis.
A tadpole is really a fish ; not merely in its habits, but in its
mode of breathing, in the arrangement of its heart and blood-
vessels, and, indeed, in almost every detail of its organisation.
The fact that a frog should commence its life as a fish is
explained by the Law of Recapitulation, according to which every
animal is constrained dnring its own development to repeat, or
 FORMATION OF THE EGG. Lig

recapitulate, in a more or less modified or abbreviated manner,

the past ancestral history of the species to which it belongs ;
is compelled, in fact, to climb up its own genealogical tree.
That frogs should in their early Stages be fish is, therefore,
to be understood as meaning that frogs are descended from
fish-like ancestors, and repeat this history in the course of their
own development.

II. Detailed Account.

A. Formation of the Egg.
The early stages in the formation of the egg cannot be seen
in the adult frog, but must be studied in tadpoles. In tadpoles
of about 10 mm. length, shortly after the opening of the mouth,
a pair of longitudinal ridge-like thickenings of peritoneum
appear along the dorsal surface of the body cavity, close to the
mesentery, and along the inner borders of the developing kidneys.
These genital ridges are found in all tadpoles of this age, no
distinction of sex appearing until a much later period.
Kach genital ridge is at first due merely to slight modification
in the shape of the peritoneal epithelial cells which, elsewhere
flattened, become here cubical or slightly columnar. The ridges
soon become more prominent, especially at their anterior ends ;
their growth being due partly to the epithelial cells increasing
by division so as to form a layer several cells thick, and partly
to the ingrowth of an axial core of connective tissue from the
basal membrane of the peritoneum, along which the bloodvessels
gain access to the ridge.
From the posterior two-thirds of the genital ridge the ovary,
or in the male the testis, is developed ; while the anterior third
undergoes degenerative changes, and becomes converted into the
fat body. 7
The Primitive Ova. At an early stage certain of the epi-
thelial cells of the genital ridge become conspicuous by their
larger size and more spherical shape; and around these larger
cells, or primitive ova, as they are called, the smaller epitheli
al
cells become arranged so as to form capsules or follicles, the
follicles, with their contained primitive ova, forming small knob-
like projections on the surface of the genital ridge. New
primitive ova arise either directly from the surface epithelium,
or by division of the already existing ones.
I
114 DEVELOPMENT OF THE FROG.

The Permanent Ova. Up to this time there has been no

distinction between male and female, the processes described
occurring in all tadpoles alike. Sexual differentiation appears
about the time of the metamorphosis. In the female the change
consists essentially in a great increase in the size of the genital
ridge, which now becomes the ovary, and in the formation of the
permanent ova, or eggs. The permanent ova are derived from
the primitive ova: in some cases each primitive ovum is
directly converted into a permanent ovum, but in others two
or more primitive ova appear to be concerned in the formation
of a single permanent ovum.
A permanent ovum is enclosed in a follicle or capsule like the
primitive ovum, and differs from this latter in the following
points :—(1) it is of larger size; (2) it contains within its sub-
stance a number of small sharply-defined yellowish granules of
food yolk, which are elaborated by the follicle cells and passed
on from them to the ovum; these yolk granules increase
rapidly in number, and to them the greater size and opacity of
the permanent ovum are chiefly due; (3) important changes
have occurred in the nucleus; in the primitive ovum the
nucleus is small, granular in appearance, and apparently solid ;
in the permanent ovum the nucleus, or germinal vesicle, is of
very large size, up to half the diameter of the entire ovum, and
consists of an elastic capsule or nuclear membrane, filled with
fluid and traversed by a protoplasmic recticulum enlarged at its
nodes to form the nucleoli, or germinal spots.
When the permanent ovum has reached a diameter of about
0-5 mm., an exceedingly thin structureless investment, the
vitelline membrane, is formed immediately around it, within
the follicle. The mode of origin of the vitelline membrane is
not clearly made out, but it seems to be formed from the ovum
itself rather than from the follicular epithelium.
A little later still a layer of black pigment appears on the
surface of the ovum; it is at first irregularly distributed over
the whole surface, but as the ovum ripens it becomes restricted
to one half or hemisphere. The pigment is contained, and
apparently formed within the ovum itself, but it is not clear
how it is formed or what purpose it fulfils.
B. Maturation of the Egg.
The eggs have now reached their full size, and project from
 MATURATION OF THE EGG. 115

the surface of the ovary like small shot; but they have still to
pass through the process of maturation, or ripening, before they
are ready to be fertilised. This process of maturation concerns
the nucleus almost exclusively.
The nucleus, which at its full size we have seen to be quite
half the diameter of the egg itself, begins to shrink; the
nuclear membrane becomes wrinkled, its surface presenting a
number of small wart-like projections, so that the whole nucleus
has a blackberry-like appearance. Part of the nuclear fluid
exudes through the nuclear membrane into the substance of the
ege; a great part of the nuclear reticulum disappears, or
becomes broken up into isolated globules or nucleoli, but a very
small part remains in the centre as a slender intricately-coiled
thread, the nuclear skein.
About this time the eggs are discharged from the ovary, the
follicles rupturing, and the eggs falling into the body cavity of
the frog; along this they pass forwards, directed partly by con-
traction of the muscular body-walls, partly by the action of the
cilia of the peritoneum, to the mouths of the oviducts, which
are situated at the anterior end of the body cavity opposite
the roots of the lungs. In the first, or thick-walled, part of the
oviduct the eggs acquire gelatinous investments, secreted by
glands in its walls. The terminal, or hinder, part of the oviduct
forms a thin-walled sac capable of great distension, within
which the eggs accumulate in large numbers. Finally the eggs
are passed out through the cloaca into water, in which the
albuminous investments of the eggs speedily swell up to form
the gelatinous mass of the frog’s spawn.
During the discharge of the egg from the ovary, and its
passage down the oviduct, further changes occur in its nucleus.
The nuclear membrane becomes still further collapsed, and
finally disappears completely ; the nuclear fluid and nucleoli
become distributed through the substance of the egg, and of the
original large nucleus the exceedingly minute nuclear skein
alone remains.
This nuclear skein moves from the centre of the egg to its
‘surface, which it reaches opposite the centre of the black hemi-
sphere. The skein, previously an irregularly tangled thread, now
assumes the definite form and arrangement of a nuclear spindle,
‘such as may be seen in the nucleus of an epithelial or other cell
immediately before division of the cell occurs.
116 DEVELOPMENT OF THE FROG.

The First Polar Body. About the time the egg is laid, but
before it is fertilised, the egg becomes slightly flattened at its
upper or black pole, a certain amount of fluid being exuded
between the egg and the vitelline membrane. The nuclear
spindle now divides into two equal parts, one of which remains
within the egg, while the other is extruded from it as the
first polar body, a minute ovoidal white globule, which lies on
the surface of the egg in the exuded peri-vitelline fluid.
The Second Polar Body. The half of the nuclear spindle
that remains within the egg retreats from the surface a little
distance, and then divides into two equal parts, one of which
remains within the egg as the female pronucleus, while the
other is extruded as the second polar body, a minute white
globule very similar to the first polar body, and like this lying
freely in the perivitelline fluid on the top of the egg.
In the case of most animals in which the formation of polar:
bodies has been observed, both the first and second polar bodies.
are extruded before fertilisation is effected. In the frog the
extrusion of the second polar body does not occur until after
the spermatozoon has entered the egg, though before the com-
pletion of the act of fertilisation.
C. Fertilisation of the Egg.
Fertilisation, or impregnation, consists in fusion of the sper-
matozoon with the egg; or, more strictly speaking, fusion of
the nuclei of these two bodies.
The spermatozoa, after being shed over the spawn by the
male, swim actively by means of their long tails, penetrate the:
gelatinous investment of the eggs, bore their way through the
vitelline membrane, and so penetrate into the eggs themselves,
which they enter at or close to the upper or black pole.
A single spermatozoon is sufficient to fertilise an egg, and it.
is doubtful whether more than one is ever concerned in the
process.
In about an hour after the spermatozoon has entered, a pig-
mented process may be seen projecting inwards from the surface
of the egg, with a clear spot in its centre. This spot is the:
nucleus of the spermatozoon, and is spoken of as the male pro-
nucleus: it penetrates further into the egg, carrying the pigment
with it, so that it appears surrounded by a pigmented capsule:
connected with the surface of the egg by a pigmented stalk.
 FERTILISATION OF THE EGG. 117

By this time the second polar body has been formed and
extruded, and the female pronucleus is the only part of the
original egg nucleus still remaining. The male and female pro-
nuclei, which are at first some little distance apart, rapidly
approach each other, come into close contact, and after having
increased considerably in size, fuse together, about two and a
half hours after fertilisation has commenced, to form the
segmentation nucleus.
The segmentation nucleus is a large spherical vesicle im-
bedded in finely granular protoplasm, and surrounded by an
ill-defined capsule of pigment: its formation by the fusion of the
male and female pronuclei completes the act of fertilisation.
The female pronucleus may be regarded as an imperfect
nucleus, and the upshot of the process of fertilisation is the
completion of this nucleus ; the nucleus of the spermatozoon, or
male pronucleus, replacing the part of the egg-nucleus which
has been lost as the polar bodies. The further explanation of
the sexual process is probably to be found in the great advan-
tage, as regards vigour of offspring, that is known to result
in both animal and vegetable kingdoms from cross fertilisation,
2.€., from combining the energies of two distinct individuals in
the act of reproduction.

D. Segmentation of the Egg.

The earliest stages of development consist in repeated division
of the egg, whereby it becomes converted from the unicellular
condition, which is permanent only in the lowest animals, to the
multicellular state characteristic of all higher animals. To these
early processes of development the name Segmentation is given.
Very shortly after the completion of the act of fertilisation
and formation of the segmentation nucleus this latter loses its
spherical form and becomes spindle-shaped, the yoke granules at
the same time showing a tendency to arrange themselves in lines
radiating outwards from the ends of the spindle. The nucleus
now divides into two halves, which move away from each other;
the yolk granules tend to aggregate themselves around the two
nuclei, and a thin vertical plate of finely granular substance is
left, bisecting the egg.
At the upper or black pole of the egg a depression now
appears, at first as a small pit and then as a groove, which
soon extends all round, and, rapidly deepening, divides the egg
118 DEVELOPMENT OF THE FROG.

into two completely separate halves along a plane corresponding

with the vertical plate mentioned above.
Each of the two nuclei soon divides again into two, and a
second cleft is formed in the same manner as before: it also is
vertical, but in a plane at right angles to the first one; and on
its completion the egg consists of four precisely similar segments
or cells, each with a nucleus.
The third cleft is horizontal, but not equatorial, lying nearer
the upper than the lower pole: it divides each of the four cells
into two, an upper smaller and a lower larger one.
Two more vertical clefts next appear simultaneously at the
upper pole, midway between the two primary clefts, and extend-
ing downwards divide first the smaller and then the larger cells,
giving sixteen cells in all, eight smaller upper ones, and eight
larger lower ones. Two more horizontal clefts then appear, which
again double the number of segments, giving thirty-two in all.

Fig. 22. Segmentation of the frog’s egg. (From Haddon, after Ecker.)
The numbers above the figures indicate the number of segments at
the several stages. The dotted lines mark the positions of the clefts
that will next appear.
From this stage segmentation proceeds in a less regular
manner, the upper and smaller cells dividing more rapidly than
the lower and larger ones. By means of radial and concentric
clefts, the number of cells is rapidly increased, division of the
cells being in all cases, as from the first, preceded by division of
their nuclei.
At the stage when only eight cells are present, 7.e., on the
completion of the third cleft, a small cavity appears in the
centre of the egg, round which the cells are grouped: during .
the later phases of segmentation this segmentation cavity, as it
is called, increases considerably in size: it is from the first
 SEGMENTATION OF THE EGG. 119

situated nearer the upper than the lower pole of the egg, and is
filled with fluid.
At the close of segmentation the egg has the structure shown
in section in Fig. 23.
It is a hollow ball with its walls composed of three or four
layers of cells, and of very unequal thickness, owing to the
segmentation cavity lying in the upper half of the egg. The
cells of the upper half are small, fairly uniform in size, and
regularly arranged, while those of the lower half are larger,
and more irregular both in shape and size. The superficial
cells of the upper half are deeply pigmented, while the cells of
the lower half are almost colourless.
The distinction between upper and lower cells is however not
an absolute one, a ring of cells more or less intermediate in size,
shape, and depth of pigmentation, occurring round the equator
of the egg at the junction of its upper and lower halves.
E

Fig. 23. Vertical section through a frog’s egg at the close of segmen-
tation. x 28
E, epiblast : SC, segmentation cavity: Y, lower layer or yolk cells.
The process of segmentation is as mentioned above, simply
one of cell-division; and the unequal rates at which the different
parts of the egg segment are to be regarded as due to the
retarding influence of the granules of food-yolk, which, being
themselves inert, must hinder the activity of the protoplasm in
which they are imbedded. These granules of food-yolk are more
120 DEVELOPMENT OF THE FROG.

abundant in the lower than the upper half of the egg, and this
unequal distribution of food-yolk is the direct cause of the
unequal segmentation of the egg. The purpose of food-yolk is to
afford a supply of nutriment at the expense of which the earlier
developmental processes may be accomplished, until the young
animal is sufficiently advanced to obtain food for itself: and the
direct influence of this food-yolk will be to hinder rather than
to help these processes.
We have seen above that the history of development of
an animal is to be regarded as a recapitulation of its pedigree :
and this explanation applies to the earliest stages equally with
the later ones. If it be true that an animal, such as a frog,
during its own development repeats its ancestral history, climbs
up its own genealogical tree, then the earliest phases of this
development must represent the earliest, z.e., the most remote
ancestors. On this view the unicellular condition of the egg
is of great interest as indicating a similar unicellular condition
in some very remote ancestor ; 2.¢., as indicating that the higher
animals are descended from forms which, like the Protozoa
nowadays, remained throughout their lives single cells.
E. The Germinal Layers.
At the close of segmentation we have seen that the egg con-
sists of cells of two kinds: firstly, those of the upper half of the
ry | pO Ye Sat
d LSaeUNeM, ~~
p "' > x 4 (i

MAG

Fig. 24. Longitudinal vertical section of a frog embryo, showing com-

mencing invagination. x 28.
B, blastopore: EE, outer or epidermic layer of epiblast: EN, inner or
nervous layer of epiblast: SC, segmentation cavity: Y, yolk cells.
 FORMATION OF THE GERMINAL LAYERS, [21

egg, which are smaller, pigmented, more regularly arranged,

and comparatively free from food-yolk: secondly those of the
lower half of the egg which are larger, less regular, and almost
free from pigment, but much distended by food-yolk, which is
present in such quantity as to render them comparatively inert.
The former are the epiblast cells: the latter may conveniently
be spoken of as the lower layer cells or yolk-cells,
The epiblast shows almost from the first a distinction into
two layers: the most superficial cells being somewhat cubical
in shape and closely applied side by side so as to form a con-

Bali es
NT i in
Eri ll
ml Se

cone

SS ~] ‘
OO pas i
SOS Mod
URC SS lsSeen

Fig. 25. Longitudinal vertical section through a frog embryo at a later

stage in the formation of the mesenteron. x 30.
H, invaginate hypoblast: M, mesoblast: MN, mesenteron: N, noto-
chord: SC, segmentation cavity : YP, yolk plug, filling up the blastopore.
tinuous and deeply pigmented layer ; while the deeper epiblast
cells are more spherical, less strongly pigmented, and loosely
arranged in a layer two or more cells deep.
The limit between the epiblast and the lower layer cells is
indicated on the surface of the egg by the boundary line between
the black and white areas of the egg, and at the close of segment-
ation these two areas are approximately equal in extent. In the
succeeding stages the black area increases rapidly at the expense
of the white area, and in a few hours the pigmented epiblast
cells have covered the whole of the egg with the exception of a
139 DEVELOPMENT OF THE FROG.

small circular patch where alone the white yolk-cells come to

the surface, Fig. 25, YP. This extension of the epiblast occurs
all round its margin; it is effected by the addition of cells cut
out from the superficial layer of yolk-cells, and not by the

iy| iit
oei
f

tl

Fig. 26. Longitudinal vertical section through a frog embryo showing

the completion of the mesenteron.
B, blastopore : EE, epidermic layer of epiblast: EN, nervous layer of
epiblast : H, invaginate hypoblast: M. mesoblast: MN, mesenteron :
N, notochord.
The circular aperture in the epiblast where the yolk-cells still
come to the surface is called the blastopore: Fig. 26, B. It is
situated at what will become the posterior end of the embryo ;
and it is bordered by a distinct rim or lip, round which the
epiblast turns inwards into the interior of the egg. The circular
plug of yolk cells filling up the blastopore is spoken of as the
yolk plug.
The Mesenteron. During this time a narrow slit-like cavity
has appeared in the interior of the egg: Fig. 25,MN. This
cavity, the mesenteron, is formed by splitting apart of the
yolk cells, and from its first appearance communicates with
the exterior at the blastopore. It is at first short and very
shallow, Figs. 24 and 25; but it rapidly extends further and
further into the egg, and by depression of its lower wall or
floor becomes widened out into a space of considerable size,
 FORMATION OF THE GERMINAL LAYERS. 123

Fig. 26, MN, which gives rise to the greater part of the length
of the alimentary canal of the embryo. The mesenteron still
communicates with the exterior through the blastopore, but
the permanent mouth and anus are not yet formed.
The roof of the mesenteron is thin, and is formed of a
definite Jayer of cells spoken of as the hypoblast: the floor is
at first very thick, and is formed by the mass of the yolk-cells.
During the process of formation of the mesenteron by in-
growth of the hypoblast cells the segmentation cavity gets
pushed out of place and ultimately obliterated.
The Notochord. Along the roof of the mesenteron in the
mid-dorsal line a rod-like thickening of the hypoblast is formed
at a very early stage. This is the notochord (Figs. 25 and 26 N),
which serves to slightly stiffen the back of the embryo, and is
for some time the only skeleton which it possesses.
It very early splits off from the roof of the mesenteron, except
at its hinder end, where it remains for some time in continuity
with both hypoblast and epiblast at the lip of the blastopore.
The Mesoblast. Between the epiblast and hypoblast a third
or intermediate layer of cells, the mesoblast, is soon established.
It is formed by differentiation of the surface hypoblast and
yolk cells as a separate layer, lying immediately beneath the
epiblast, but quite distinct from it. It extends all round the
embryo except along the mid-dorsal line, where the space
between the epiblast and hypoblast is occupied by the notochord.
It is, for a time, incomplete in front, opposite the segmentation
cavity, but soon grows in from the sides so as to fill up the
deficiency.
The cells of the mesoblast become early arranged in two
parallel layers or sheets, which separate slightly from each other,
so as to leave between them a narrow space, which later on
becomes the body cavity or celom. (Cf. Fig. 27.) In many
Specimens the mesobsast cells are from the first arranged in
two layers.
Fate of the germinal layers. From one or other of the three
germinal layers,—epiblast, mesoblast, and hypoblast,—every
part of the embryo is formed.
The epiblast, or outer layer, gives rise to the epidermis
covering the body generally, and to the various glandular and
other structures derived from the epidermis; to the nervous
124 DEVELOPMENT OF THE FROG.

system, both central and peripheral; to the olfactory and

auditory epithelium, to the retina and lens of the eye, and to
the other sensory organs; to the epithelial lining of the mouth
and anus; and to the pineal and pituitary bodies.
The hypoblast, or inner layer, gives rise to the epithelium
lining the alimentary canal and its various diverticula, including
the glands of the esophagus, stomach and intestine, the lungs,
the bladder, the bile ducts, gall bladder, pancreatic ducts, and
the hepatic cells of the liver and the secreting cells of the
pancreas ; the notochord is also formed from hypoblast.
From the mesoblast, or middle layer, are derived all struc-
tures between the epiblast and hypoblast ; z.e., the connective
tissue, muscles, skeleton (except the notochord), blood-vessels
and lymphatics ; and also the peritoneum, and the urinary and
reproductive organs.
F. Development of the Nervous System.
It is convenient from the point we have now reached to deal
with the several systems one by one. The nervous system is a
suitable one to commence with, as it appears at a very early
NG
NF

N NO
2 |Ean Por ae “G'
e ane TS
: SEES
r ea (y%
SOB yest ME
omLQ oy O>.%
ey
MN ms) MH
:
j jas
TORK
sts
Vy
Ge

So ‘) (D
on
e.

EN
Fig. 27. Transverse section through a frog embryo during the for-
mation of the neural canal.
C, celom: EE, epidermic layer of epiblast: EN, nervous layer of
epiblast :H, hypoblast :M, mesoblast: ME, somatopleuric layer of meso-
blast: MH, splanchnopleuric layer of mesoblast: MN, mesenteron:
N; notochord : NF, neural fold: NG, neural groove: Y, yolk cells.
 THE NERVOUS SYSTEM. 125

stage of development, and plays an important part, especially

in the younger stages, in determining the shape and proportions:
of the embryo. .
The epiblast consists almost from the first of two layers, the
distinction between which is already established at the close of
segmentation. (Fig. 23). Of these the upper or epidermic
layer is a single stratum of closely fitting cubical cells ; while
the lower or nervous layer consists of ovoid or spherical cells,
more loosely compacted, and two or three deep. It is from the
latter that the nervous system is developed.
The first trace of the nervous system is seen about a week.
after fertilisation, when the embryo is still spherical and the:
blastopore has become much reduced in size and difficult to see..
(of, Fig. 26).

ECE
Ti
seeaal'aas oeELI |ill
| ]
D> \ a
<7)

L.
Fig. 28. Longitudinal vertical section through a frog embryo shortly
bere the closure of the blastopore. Length of the embryo 2°5 mm.
x 30.
B, blastopore: BF, fore-brain: BH, hind-brain: BM, mid-brain :.
H, hypoblast: L, liver: M, mesoblast: MN, mesenteron: N, notochord:
NC, neurenteric canal: P, ingrowth of epiblast to form pituitary body:
PD, proctodzeum : R, rectal diverticulum of mesenteron : S, central canal
of spinal cord: Y, yolk cells.
The dorsal surface of the embryo now flattens slightly, and
along the flattened area the nervous layer of the epiblast.
thickens to form the neural plate, which is wide in front but.
126 DEVELOPMENT OF THE FROG.

narrows posteriorly towards the blastopore. Slightly raised

ridges, the neural folds, soon appear, bordering the sides of the
neural plate ; and a longitudinal neural groove is formed along
its dorsal surface in the median line, extending forwards from
the blastopore.

BH

v
Fig. 29. Longitudinal vertical section through the anterior end of
S tadpole shortly after the time of hatching. Length of the tadpole
Imm.
A, auricle of heart: BF, fore-brain : BH, hind-brain: BM, mid-brain:
C’, pericardial cavity: CV, vesicle of cerebral hemispheres: |, infundi-
bulum: L, liver: N, notochord: O, depression of floor of forebrain from
which the optic vesicles arise: OE, cesophagus: P, pituitary body:
PN, pineal body: Ss, central canal of spinal cord: SD, stomodzeum:
T, truncus arteriosus: V, ventricle.
A transverse fold connects the anterior ends of the neural
folds together, slightly raising up the anterior end of the neural
plate. The neural folds now grow rapidly : the groove between
them deepens, and the folds becoming more and more prominent
bend in towards each other (Fig. 27) and finally meet and fuse,
thereby converting the neural groove into a tube.
 DEVELOPMENT OF THE FROG. 127

The neural folds first meet about the junction of the head and
trunk of the future tadpole, from which point the fusion extends
rapidly in both directions, forwards and backwards. The last
point at which fusion occurs is a little distance behind the
anterior end of the tube, at the place where the pineal body will
appear later.
In front, the neural tube ends blindly: at its posterior end
it opens to the exterior at the blastopore, and is in free com-

Fig. 30. Longitudinal vertical section through the head and anterior
art of the body of a tadpole about the time of appearance of the hind
egs. Length of tadpole, 12mm. x 14.
A, auricle of heart: AD, dorsal aorta: BB, basi-branchial cartilage:
BF, fore-brain: BH, hind-brain : BM, mid-brain : C, ccelom or body cavity:
Cc’, pericardial cavity :CH, cerebral hemisphere: CB, rudimentary cere-
bellum: CP, choroid plexus of fourth ventricle: CP’, choroid plexus of
third ventricle: F, pharynx: G, stomach: H, lung: |, infundibulum:
J, horny jaws: K, lip: L, liver: N, notochord: 0, depression of floor of
fore-brain from which the optic nerves arise : OE, cesophagus : P, pituitary
dy: PN, pineal body: §, .central canal of spinal cord: T, truncus
arteriosus: V, ventricle.

munication with the mesenteron. (Cf. Fig. 28.) The short

channel of communication between the neural tube and the
mesenteron, 2.¢., between the nervous system and the alimentary
 128 THE NERVOUS SYSTEM.

canal, is spoken of as the neurenteric canal: it is only present:

for a short time, and closes up before the tadpole hatches.
The neural tube, formed in this way, soon separates from the
surface epiblast, and by thickening of its walls and other
changes becomes converted into the central nervous system ;
the anterior part forming the brain, and the posterior part the
spinal cord. The lumen or cavity of the tube persists as the
central canal of the spinal cord and the ventricles of the brain.
The Brain, At the time of its first appearance the brain is
bent at right angles about the middle of its length ; the axis of
the anterior portion being vertical, and that of the posterior
portion horizontal. (Fig. 28.) The posterior portion, or hind-
brain, BH, is wide from side to side, and has moderately thick
sides and floor, but a thin roof: it is continuous behind with
the spinal cord.
The anterior or vertical portion has walls of nearly uniform
thickness in all parts. It is divided by a slight constriction,
best marked at the sides, into an upper or posterior part, the
mid-brain, BM, which forms the angle of the bend and lies
opposite the anterior end of the notochord; and a lower and
larger portion, the fore-brain, BF, which is produced laterally
into a pair of hollow outgrowths, the optic vesicles.
The further development of the brain is illustrated by Figs.
29 and 30. It will be seen that the rectangular bending of the
brain, which is known as cranial flexure, and which was so
prominent a feature in the earlier stage, is no longer obvious:
a closer comparison of the figures will show, however, that this
straightening of the brain, or rectification of the cranial flexure,
is apparent rather than real, and is brought about partly by the
development of the cerebral hemispheres, which grow upwards
and forwards from the fore-brain, and still more largely by the
formation of the mouth and the growth forwards of the face and
lips, which cause the brain to take a much less prominent share
in determining the shape of the head.
The hind-brain, BH, has undergone but little change in Fig.
29, except an increase in thickness of its floor and sides. At the
stage represented in Fig. 30 it is separated from the mid-brain
on the dorsal surface by a well-marked groove, immediately
behind which the roof of the hind-brain is thickeued transversely
to form the cerebellum, CB. The cavity of the hind-brain
remains as the fourth ventricle, the roof of which is very thin
 THE BRAIN. 129

and thrown into numerous transverse folds, GP, which hang

down into the ventricle, and between the layers of which lie the
blood vessels of the choroid plexus of the ventricle.
The mid-brain, BM, thickens on its floor to form the crura
cerebri. Its roof grows out laterally into a pair of hollow ovoid
processes, the optic lobes: and its cavity persists as the
aqueductus Sylvii.
The fore-brain, BF, becomes the thalamencephalon of the
adult: its cavity becomes the third ventricle, which by thicken-
ing of its walls to form the optic thalami is reduced to a vertical
cleft, very narrow from side to side. Its floor is produced down-
wards and backwards into a hollow sac-like diverticulum, the
infundibulum, |, in connection with which is the pituitary body.
In front of the infundibulum is a transverse ridge projecting
into the ventricle, and formed by the roots of the optic nerves.
The roof of the fore-brain remains thin: a little behind the
middle of its length the pineal body, PN, arises as a median
hollow diverticulum, Fig. 29; this is formed at the spot where
the final closure of the neural tube took place, and is at first
directed backwards: in the later stages it grows forwards and
forms a rounded vesicle connected with the brain by a long
pigmented stalk: when the skull develops it cuts off the vesicle
from the stalk, the former remaining as a small rounded body
outside the skull, while the stalk persists as a slender pigmented
tract within the cranial cavity.
In front of the pineal body, and at the anterior end of the
fore-brain, the roof is thrown into folds which hang down into
the ventricle forming a choroid plexus, CP’, similar to that in
the medulla.
The anterior end of the fore-brain grows forwards as a median
thin-walled cerebral vesicle, from which at a slightly later stage
the cerebral hemispheres, CH, arise as a pair of hollow out-
growths; the foramina of Monro being the apertures of com-
munication being the lateral ventricles or cavities of the
hemispheres, and the third ventricle. The anterior ends of the
hemispheres grow forwards as the olfactory lobes, which become
fused together in the median plane.
The peripheral nervous system. The cranial nerves and the
dorsal roots of the spinal nerves are formed from the deeper or
nervous layer of the epidermis. They arise as lateral outgrowths
from the edges of the neural plate, and may be recognised at a
J
130 DEVELOPMENT OF THE FROG.

very early stage, while the neural groove is still shallow and
open: they are, therefore, from their first appearance con-
tinuous with the brain or spinal cord.
The ventral roots of the spinal nerves arise later than the
dorsal ones, as outgrowths from the cord near its ventral surface.
They are at first independent of the dorsal roots, but soon
become connected with these.
G. Development of the Sense Organs.
The organs of special sensation are developed from the deeper
or nervous layer of the epiblast, and become connected with
their respective nerves at a very early stage of their formation.
The derivation of the sense organs from the epiblast is
explained by the fact that they are ‘concerned with the appre-
ciation of the presence and nature of external objects, and are
therefore necessarily formed on the surface of the body. They
may be regarded as specially modified portions of the epidermis.
The Nose. The olfactory organs appear at a very early stage
as paired thickenings of the nervous layer of the epiblast at the
anterior end of the head, in the angles between the fore-brain
and the optic vesicles. A pitting-in of the surface, involving
both layers of the epiblast, soon appears in each of these
thickenings, and the pits so formed become the nasal sacs; the
mouths of the pits forming the nostrils or anterior nares, and
the epiblast lining the pits giving rise to the olfactory epithelium.
From the inner or deeper end of each olfactory pit a diverti-
culum, at first solid, but soon becoming hollow, grows down-
wards to the roof of the pharynx, into which it opens, as the
posterior nares, very shortly after the formation of the mouth
opening.
The Eye. The eye differs from the other sense organs, inas-
much as the lens alone is formed directly from the surface epi-
blast, while the sensitive part of the eye, or retina, arises as an
outgrowth from the brain. The optic vesicles have already been
described as arising at a very early period as lateral outgrowths
from the fore-brain; these soon become constricted at their
necks so as to be connected with the brain by narrow stalks,
which ultimately become the optic nerves.
The outer surface of each optic vesicle, which is at first in close
contact with the surface epiblast, soon becomes flattened, and
then thickens so greatly as almost to obliterate the cavity of the
 THE SENSE ORGANS, 131

vesicle. At the same time a thickening of the deeper or ner-

vous layer of the surface epiblast takes place opposite the optic
vesicle : this grows rapidly and forms a spherical body, projecting
inwards from the surface: this is at first solid, but soon becomes
hollow and breaks away completely from the surface epiblast :
it becomes later on the lens of the eye, and may be spoken of
as the lens vesicle.
Partly in consequence of the ingrowth of the lens vesicle,
and partly through growth of the optic vesicle itself, this latter
becomes pitted on its outer surface, and so converted into a cup
—the optic cup—with double walls ; the inner wall being the
thickened and originally outer wall of the optic vesicle, and the
outer wall of the cup being the original inner or deeper part of
the wall of the vesicle.
From the optic cup and lens vesicle the adult eye is derived
in the following way: The lens becomes solid, owing to thicken-
ing of its inner wall, which proceeds so far as to finally oblite-
rate the cavity. The optic cup enlarges considerably ; it remains
in contact with the lens at its edge or lip, but elsewhere is
separated from it by a space which becomes the posterior
chamber of the eye, and in which the vitreous humour is
formed. The inner wall of the optic cup gives rise to the
retina, the rods and cones growing out from its outer surface ;
while the outer and thinner wall of the optic cup forms the
layer of pigment cells in which the rods and cones are imbedded.
‘The choroid and sclerotic coats are formed from the mesoblast
surrounding the optic cup.
The eye developes very slowly, and throughout the tadpole
stage of existence is in a very rudimentary and imperfect
condition.
The Ear. The ears are developed as a pair of pit-like in-
vaginations of the nervous layer of the epiblast at the sides of
the hind-brain. The invaginations do not involve the epidermic
or surface layer of the epiblast, so that the auditory pits do not
-open to the exterior.
The mouths of the pits very early narrow and close; and the
auditory vesicles so formed separate from the epiblast and lie in
the mesoblast at the sides of the head. The vesicle becomes
the vestibule of the adult ear; the semicircular canals arising
-as outgrowths from it.
Throughout the tadpole stage of existence there is no further
132 DEVELOPMENT OF THE FROG.

modification ; but shortly after the metamorphosis the hyoman-

dibular cleft, which has at no period opened to the exterior,
is stated to widen somewhat and form the Eustachian passage,
while the layer of integument closing its outer end becomes the
tympanic membrane. There is some reason, however, for thinking
that the Eustachian passage developes independently in the frog,
and not from the hyomandibular cleft. The columella, which
has been described with the skull, is formed still later. (Cf
Fig. 11, p. 58).
Special Sense Organs. During the tadpole stage, while the
animal is leading an aquatic life, special sense organs in the
form of small epidermal papillae, supplied by branches of the
trigeminal and pneumo-gastric nerves, are found arranged in
rows along the body, and round the eyes, and in other parts of
the head. These are lost at the time of the metamorphosis.
The mouth of the tadpole is also provided with small rounded
papille, which are probably organs of taste. (See Fig. 30.)
H. Development of the Alimentary Canal.
The alimentary canal is developed in three lengths: (1) the
mesenteron, which is formed by splitting apart of the yolk cells
as described above ; this gives rise to nearly the whole length
of the alimentary canal; and from it are developed the gill-slits,
the lungs, the thyroid, the liver, the pancreas, and the bladder;
as well as the notochord: (2) the stomodzum, which is a
pitting-in at the anterior end of the body, from which the
mouth and pituitary body are formed : and (3) the proctodeum,
which is a similar pitting-in at the hinder end of the body to
form the anal or cloacal opening.
From the mode of their formation it follows that the mesen-
teron is lined by hypoblast, and the stomodzum and procto-
deeum by epiblast.
1. The mesenteron, The early development of the mesen-
teron has already been described.
The anterior end of the mesenteron, in the head region, is
considerably dilated from the first :and at the hinder end of the
embryo a similar, though much smaller, expansion takes place.
In this way (ef. Fig. 28), the mass of the food-yolk becomes
confined to the ventral portion of the body region, not extend-
ing into either the head or the tail.
The hypoblast, which is a definite layer of cells, at first confined.
 THE ALIMENTARY CANAL. 133

to the roof of the mesenteron, gradually spreads round its sides

until it encloses the whole of the food-yolk, and the alimentary
canal is completed as a tube, which from the first is slightly
convoluted. When the tadpole begins to feed, the alimentary
canal lengthens rapidly, and becomes coiled in a spiral manner.
Except at the anterior end, in the gill-bearing region, it is of
approximately uniform diameter throughout. During the
metamorphosis, the alimentary canal shortens considerably, and
the distinction between stomach, small intestine, and large
intestine, is definitely established.
The liver is recognisable at a very early stage—Fig. 28—as a
ventral and backwardly directed diverticulum of the anterior
part of the mesenteron, forming the anterior boundary of the
mass of food-yolk. In the later stages the walls of the diver-
ticulum thicken, and become thrown into folds between which
the vascular mesoblast makes its way: the diverticulum itself
persists as the bile duct, and the gall bladder arises as an out-
erowth from this.
The pancreas is developed as a pair of hollow outgrowths
from the mesenteron, behind the liver: in the later stages the
ducts shift so as to open into the bile duct instead of directly
into the intestine.
The bladder arises shortly before the metamorphosis as a
ventral outgrowth from the hinder end of the mesenteron, which
soon becomes bifid at its distal blind end.
The post-anal gut is an extension of the hinder end of the
mesenteron into the base of the tail, which appears as this latter
is developed: it becomes solid after a short time, and later on
disappears altogether. It is perhaps to be regarded as formed
by a mechanical drawing out of the intestine by the outgrowing
tail.
The lungs. Immediately behind the gill-bearing region or
pharynx, the alimentary canal narrows very considerably ; its
sides become folded inwards, and the two folds meeting each
other divide the canal into a dorsal tube or cesophagus, and a
ventral one which forms the laryngeal chamber: from this
latter the lungs arise as thin-walled lateral outgrowths. They
appear first in young tadpoles of about 8 mm. length, 7.e., some
time after hatching, but shortly before the opening of the
mouth. About the time that the lungs first appear, in tadpoles
of about 8 mm. length, the cesophagus, which up to this time
134 DEVELOPMENT OF THE FROG.

has been tubular, becomes solid, and remains so until a short

time after the formation of the mouth, The meaning of this
curious point has not been ascertained.
2. The stomodeum. At the stage represented in Fig. 28,
shortly after closure of the neural canal, a conical ingrowth, P,
of the nervous layer of the epidermis is formed at the anterior
end of the body immediately below the fore-brain: from this
ingrowth the pituitary body is developed, and a slight depres-
sion of the surface epiblast opposite its base, marks the position
of the stomodzeum.
At the time of hatching, this depression is a small shallow
pit, separated from the anterior end of the mesenteron by a thin
septum. Soon after hatching, in tadpoles of about 9 mm.
length, this septum becomes perforated, and the alimentary
canal communicates with the exterior through this stomodeeal
pit. After the perforation is effected, the lips with the whole
anterior part of the face grow forwards rapidly ; the horny jaws
are formed, and the tadpole begins to feed vigorously. (Cf Figs.
29 and 30.)
The pituitary body (Figs. 28, 29, 30, P) is formed from the
ingrowing stalk of epiblast described above: this rapidly
elongates, growing backwards between the brain and the roof
of the mesenteron until it reaches the infundibulum : its hinder
end now becomes tubular, gives off a few lateral diverticula,
separates from the stalk, which soon disappears, and becomes
applied to the ventral surface of the hinder end of the infundi-
bulum to form the pituitary body,
3. The proctodeum or anal invagination appears before the
stomodeum. Shortly before the neural folds have met to
form the neural tube, the proctodseum is visible as a small
median depression of the epiblast at the hinder end of the
embryo, a little way below the blastopore. The cells lining it
are rather strongly pigmented, and slightly larger than the
surrounding epiblast cells.
From the hinder end of the mesenteron a rectal diverticulum
(Fig. 28, R) extends downwards towards the proctodeum: a
little later, and some time before the tadpole hatches, the two
structures meet ; perforation occurs; and the definitive anal or
cloacal opening is formed. For a short time the blastopore
and the proctodeum are both open; but very shortly after
completion of the proctodzeum the blastopore closes finally.
 THE GILL CLEFTS AND ARCHES. 135

I. The Gill Clefts and Arches.

Some little time before the tadpole is hatched a series of
vertical ridge-like thickenings appear on the sides of the head
and neck. These are the visceral arches, and are six in number
on each side.
The most anterior is the mandibular arch, and gives rise
later on to the lower jaw: the second is the hyoid arch:
and the succeeding four are the first, second, third and fourth
branchial arches respectively.
About the time of hatching the external gills grow out as
branching and richly ciliated processes from the outer surfaces
of the first and second branchial arches, and a little later from
the third branchial arches as well (Fig. 31).
_ At the same time, the hypoblastic epithelium at each side of
the buccal cavity becomes thrown into folds, which extend out-
wards towards the surface of the neck as paired outgrowths, lying
between the visceral arches. Of these outgrowths or pouches,
which are known as visceral clefts, there are five on each side.
The most anterior one is the hyomandibular cleft, and lies
between the mandibular and hyoid arches: its outer end lies
very close to the surface of the neck, though it does not actually
open to the exterior.
The four hinder visceral clefts perforate the skin about the
time of formation of the mouth opening, 2.e., in tadpoles of about
9 mm. length, and open to the exterior as the gill clefts. These
are slit-like openings lying between the hyoid and first branchial,
first and second branchial, second and third branchial, and third
and fourth branchial arches respectively ; and are known as the
first, second, third and fourth branchial clefts.
From the hyoid arches a pair of opercular folds arise, which
grow back over the external gills, and the branchial arches and
clefts. The two opercular folds meet below the neck in the
mid-ventral line, and enclose the gills in a branchial chamber.
The hinder borders of the opercular folds fuse with the body
wall except at one place on the left side, where a spout-like
opening remains through which the branchial chamber com-
municates with the exterior.
As the opercular folds develope, the external gills gradually
shrivel up, and are replaced functionally by the internal gills.
These latter are delicate thin-walled vascular tufts, arranged in
a double row along the ventral half of each of the first three
136 _ DEVELOPMENT OF THE FROG.

branchial arches, and in a single row on the fourth branchial

arch.
The inner borders of the branchial arches are thickened, and
produced into processes which unite to form a kind of filtering
apparatus, or sieve, through which the water, taken in through
the mouth or nose, is strained before being passed over the gills
into the branchial cavity and so out.

K. The Vascular System.

The heart is at first a straight tube developed in the meso-
blast of the ventral wall of the pharynx. This soon lengthens,
becomes twisted into an § shape, and divided by transverse
constrictions into chambers, (Cf. Figs. 29, 30, and 32). The
auricle is at first single, but later becomes divided by the down-
growth of a septum from its dorsal wall.
While the tadpole is breathing by means of gills, its circula-
tion is in all essential respects that of a fish. The venous blood,
returned from the body generally, enters the posterior end of
the heart, or sinus venosus: from this it passes into the second
or auricular chamber, thence to the ventricle, and from that to
the truncus arteriosus. From this latter arise on each side the
aortic arches, which carry the venous blood to the gills to be
aerated : from the gills the blood is collected by efferent vessels,
which unite above the alimentary canal to form the dorsal aorta,
which by its branches distributes the arterialised blood to all
parts of the body.
1. The Circulation during the time the tadpole is breathing
by external gills.
The arrangement of the blood-vessels, and the course of the
circulation in a 64 mm. tadpole, at a time when the external
gills are in full activity, but the internal gills have not yet
formed, is shown in Figs. 31 and 32.
The truncus arteriosus, on reaching the anterior end of the
pericardial cavity, divides at once into right and left branches.
Each of these again divides into two, the afferent vessels for the
first and second branchial arches, AF* and AF*, which carry
blood into the external gills and their branches: from these the
blood passes through short wide capillary loops into the efferent
branchial vessels, EF! and EF*, which carry it, now aerated, to
the dorsal aorta in the roof of the pharynx. The dorsal aortz
 THE VASCULAR SYSTEM. 137

of the two sides run forwards as the carotid arteries, AC, to

supply the head and brain, and also run backwards in the roof
of the pharynx, the aortz of the two sides meeting and uniting

EM
CA kok Ve EH
CE9
2 ee
ee
i F =
‘
rae E Fit

uM AUT =

UZ

(un

™ (
(I
it
mitt
A
Tul
TTA
meta
\(

a A
Fig. 31. Diagrammatic figure of the head and fore part of the body of
a 64 mm. tadpole, showing the arrangement of the branchial vessels as
seen from the ventral surface. The heart has been removed. x 33.
A, dorsal aorta: AF1, AF2, AF3, afferent branchial vessels of the first,
second, and third branchial arches: AP, pulmonary artery: AR, anterior
cerebral artery: CA, anterior commissural artery: CP, posterior com-
missural artery : the arterial circle formed by these commissural vessels
with the carotid arteries surrounds the infundibulum of the brain:
EF1, EF2, EF3, EF4, efferent branchial vessels of the first, second, third,
and fourth branchial arches: EH, efferent hyoidean vessel: EM, efferent
mandibular vessel: GE, external gill: GM, glomerulus: KA, segmental or
archinephric duct: KP, head kidney or pronephros: KS1, KS3, first and
third nephrostomes of pronephros: RT, truncus arteriosus.

about the junction of head and body to form the single

systemic aorta which supplies arterial blood to all parts of the
body.
Besides the complete sets of afferent and efferent branchial
138 DEVELOPMENT OF THE FROG.

vessels in the first and second branchial arches, similar vessels,

as yet incompletely developed, are present in the hinder arches
as well.
In the third branchial arch, there is a short afferent branch
AF® from the afferent vessel of the second branchial arch, which
c, ERS EF2 EFT EH. Ap

“o>
cra eee
a Tea
ale nD
ANS va ay
Fi=
=
E= PATTON
w=
3|
Ej=SFj7;; F
z
lh ppt
3
= Va NO Xe
8

=Ss

AF3 AF2 pry VK

Fig. 32. Diagrammatic figure of the head and forepart of the body of a
64 mm. tadpole, showing the heart, aorta, and vessels of the branchial
arches from the right side. The external gills have been removed. x 40.
A, dorsal aorta: AB, basilar artery : AC, carotid artery: AF1, AF2, AF3,
afferent branchial vessels of first, second, and third branchial arches :
AP, pulmonary artery: AR, anterior cerebral artery: AT, anterior
palatine artery: EF1, EF2, EF3, EF4, efferent branchial vessels of first,
second, third, and fourth branchial arches: EH, efferent hyoidean vessel :
EM, efferent mandibular vessel: GM, glomerulus : LV4, lacunar afferent
vessel of fourth branchial arch: RA, auricle: RV, ventricle:
VD, Cuvierian vein : VH, hepatic veins : VK, vein of sucker : VY, hyoidean
vein: YM, mandibular vein.

as yet ends blindly. There is also a well-developed efferent

vessel, EF*, which opens into the dorsal aorta.
In the fourth branchial arch there is no afferent vessel, but
 THE VASCULAR SYSTEM 139

an efferent vessel, EF*, is present, opening into the dorsal

aorta. From this efferent vessel, just before it reaches the aorta,
a backwardly directed branch arises, which will become later
the pulmonary artery, AP.
In front of the first branchial arch, vessels are present in the
hyoid and mandibular arches, which clearly belong to the same
category as the branchial vessels, but which never attain full
development, probably owing to the fact that no gills are formed
on these arches. LEfferent branches, EH, EM, opening into the
dorsal aorta, are present in both hyoid and mandibular arches ;
but these have no connection with the heart, as there are no
afferent vessels corresponding to them.
The condition of the blood-vessels, while the tadpole is
breathing by external gills, may be summarised thus :—Com-
plete systems of afferent and efferent vessels, connecting the
heart with the aorta through the gill capillaries, are present in
the first and second branchial arches, and at a stage slightly
_later than that shown in Fig. 31 in the third branchial arch as
well. <A similar set of vessels, but incomplete, is present in the
fourth branchial arch: and vessels formed on the same plan, but
still less complete, and showing signs of degenerative changes,
are present in the hyoid and mandibular arches.
There are thus six sets of branchial vessels on each side of
the pharynx: of these, three, in the first, second, and third
branchial arches, are complete; one, in the fourth branchial
arch, is incomplete; and two, in the hyoid and mandibular
arches, are rudimentary.
2. The Circulation during the time the tadpole is breathing
by internal gills.
On the formation of the internal gills, additional loops of
communication are formed in the gill tufts between the afferent
and efferent vessels of the first, second, and third branchial
arches, and also a series of similar loops between the afferent
and efferent vessels of the fourth branchial arch. The vessels
in the hyoid and mandibular arches undergo further retrograde
changes, and need not be described in detail.
In tadpoles of 12 mm. length, in which the internal gills are
fully established, and the external gills are shrivelling up, the
condition of the blood-vessels is shown in Figs. 33 and 34.
The truncus arteriosus divides at once into right and left
140 DEVELOPMENT OF THE FROG.

Ev (on
! = \ KP
KS.s~ | 5 |
Ao 40] || ~~ KA
go} j

jf> kM

LP

TR aA.
Fig. 33. A 12mm. tadpole dissected from the ventral surface to show
the heart, the internal gills, the branchial vessels, and the head kidneys
and their ducts. The tail, which is about double the length of the head
and body, has been removed. x 22.
A, dorsal aorta: AF,1, AF,3, afferent branchial vessels of first and third
branchial arches: AL, lingual artery: CG, carotid gland: EA, junction
between afferent and efferent branchial vessels of first branchial arch:
EF,1, EF,3, efferent branchial vessels of first and third branchial arches :
GM, glomerulus: KA, archinephric or segmental duct: KM, Wolffian
tubules: KP, pronephros or head kidney: KS,1, KS,3, first and third
nephrostomes of head kidney: LI, upper lip: Lu, lower lip: LP, hind
limb : OA, aperture of opercular cavity : OP, opercular cavity : RS, sinus
venosus: RT, truncus arteriosus: RV, ventricle: TC, cloaca:
TO, cesophagus, cut short: TR, rectal spout.
 THE VASCULAR SYSTEM. 141

branches, which run straight outwards in the floor of the

pharynx. Each of these branches divides, after a short course,
into three vessels, and the hindmost vessel again into two. In
this way the four afferent branchial vessels AF*, AF*, AF®, AF‘, of
the first, second, third and fourth branchial arches respectively
are formed.
A GM AB AU EFs Efe CP EF, cp
ere ea aie enehyuA sf

(I
TUTeTTITMAATETTTTTA
irr
Ee ET,
/
QED

VO AFe RB RA RV Ake RT
Fig. 34. A diagrammatic figure of the head and neck of a 12 mm. tad-
pole from the right side, to show the heart and branchial vessels. The
gills and the gill capillaries are not represented. x 35.
A, dorsal aorta: AB, basilar artery: AF1, AF2, AF4, afferent branchial
vessels of first, second, and fourth branchial arches: AL, lingual artery:
AP, pulmonary artery: AR, anterior cerebral artery: AS, posterior
palatine artery: AT, anterior palatine artery: AU, cutaneous artery:
AY, pharyngeal artery: CA, anterior commissural vessel: CG, carotid
gland: CP, posterior commissural vessel: EF1, EF2, EF3, EF4, efferent
branchial vessels of first, second, third, and fourth branchial arches:
GM, glomerulus: RA, right auricle: RB, left auricle: RT, truncus
arteriosus: RV, ventricle: VD, Cuvierian vein: VH, hepatic vein:
VI, posterior vena cava: VP, pulmonary vein.

Each afferent vessel runs outwards and upwards in its own

arch. The efferent branchial vessels lie immediately in front
of the corresponding afferent vessels, with which they are
142 DEVELOPMENT OF THE FROG.

connected by very numerous capillary loops in the substance of

the internal gills, and not shown in the figures. At their upper
ends the efferent vessels open, as before, into the dorsal aorta,
Fig. 34.
The venous blood in the heart is driven by the contraction of the
ventricle into the truncus arteriosus, and then along the afferent
branchial vessels, through the capillary loops of the gills, in
which it gets aerated, to the efferent branchial vessels; and
thence to the dorsal aorta, and so all over the body.
The lungs are by this time of considerable size: they receive
blood by the pulmonary arteries, AP, which, as already noticed,
are branches from the efferent vessels of the fourth branchial
arches, and therefore contain blood which has already passed
through the gill capillaries. The blood from the lungs is
returned direct to the heart by two pulmonary veins which
unite and open into the left auricle, the single auricular cavity
of the earlier stage being by this time divided by a vertical
septum into right and left auricles.
One other point of great importance remains to be noticed
in the arrangement of the branchial vessels of the tadpole.
The afferent and efferent vessels of each arch at first com-
municate only through the gill capillaries: but in tadpoles of
about 12 mm. length each efferent vessel becomes directly con-
nected at its ventral end with the corresponding afferent vessel,
Figs. 33 and 34. These direct connections are situated ventrally
to the gills, so that the blood in any one of the afferent branchial
vessels has two paths open to it: it may either (1) continue
along the afferent vessel, and then reach the efferent vessel by
passing through the connecting loops afforded by the gill capil-
laries; or (2) it may pass at once into the efferent vessel
through the direct communication, and so reach the dorsal aorta
without having passed through the gill at all.
So long as the tadpole is breathing by gills, these direct com-
munications between afferent and efferent vessels, though
present in all four branchial arches, are so small that practi-
cally no blood passes through them, and all the blood is com-
pelled to pass through the gills to reach the aorta.
3. The Changes in the Circulation at the time of the Meta-
morphosis.
At the time of the metamorphosis, however, when the
anterior limbs are protruded, and the tail begins to shorten,
 THE MUSCULAR SYSTEM. 143

these direct communications enlarge, so that an increasing

amount of blood takes the direct short passage, and reaches the
aorta without having passed through the gills. Additional
work is thus thrown on the lungs and skin, which consequently
receive a larger supply of blood: the gills rapidly atrophy,
though remnants of them usually persist, in a functionless con-
dition, until the end of the first year ; and the change from the
gill-breathing to the air-breathing condition is completed.
The further changes necessary to convert the circulation into
that of the adult are slight. Of the four aortic arches present
at the metamorphosis, Fig. 34, the first, in the first branchial
arch, persists as the carotid arch of the adult frog: the lingual
artery is a branch from the ventral end of the efferent vessel of
the arch, and is present from an early stage of development
(Fig. 34): and the external and internal carotid arteries, are
already present. The carotid gland, CG, is not, as sometimes
stated, a persistent portion of a gill, but is formed by further
elaboration of the direct communication between the afferent
and efferent branchial vessels of the first branchial arch.
The second aortic arch, in the second branchial arch, becomes
the systemic arch of the frog. Its dorsal end remains connected
with the carotid arch, though the connection may in the adult
become closed and ligamentous (Cf. Fig. 5, p. 27).
The third aortic arch, in the third branchial arch, loses its
connection with the aorta, and finally disappears altogether.
The fourth aortic arch, in the fourth branchial arch, also
loses its connection with the aorta, but persists as the pulmo-
cutaneous arch of the adult, from which both pulmonary and
cutaneous arteries arise.
L. Development of the Muscular System and the Celom.
The splitting of the mesoblast into outer or somatopleuric,
and inner or splanchnopleuric layers has already been described.
(Cf. Fig. 27, p. 124).
In the body the mesoblast becomes very early divided on
each side into (1) a vertebral plate, which is more dorsally
situated, and lies alongside the spinal cord and notochord; and
(2) a lateral plate, which surrounds the side of the body.
The vertebral plate very early becomes divided transversely
into muscle-segments or myotomes, which form a row of hollow
and somewhat cubical bodies, lying along each side of the
Sf
144 DEVELOPMENT OF THE FROG,

spinal cord, and separated from each other by connective tissue

septa. Later on, the walls of the myotomes thicken consider-
ably, especially the inner walls, and become converted very
largely into muscles ; while the cavities become obliterated.
The myotomes may be well seen in the tail of the tadpole,
where they form the great lateral sheets of muscle on each side
of the tail, by which the swimming movements are effected.
Owing to the transparency of the tail, their arrangement can be
very readily made out: the septa dividing the successive myo-
tomes from each other are not transverse, but > shaped, with
the angles directed forward towards the head.
The lateral plates are also in part converted into muscle:
the two layers, somatopleuric and splanchnopleuric, remain
comparatively thin, but the space between them widens out
considerably, and becomes the body cavity or coelom. ‘This at
first consists of two separate halves, right and left; but, owing to
the splitting of the mesoblast extending down to the midventral
line, the cavities of the two sides soon become continuous. The
anterior portion of the ccelom is very early shut off from the
hinder part as the pericardial cavity. (Cf Figs. 29 and 30.)
The outer or somatopleuric layer of mesoblast, with the epi-
blast, forms the body wall of the adult: the inner or splanchno-
pleuric layer, with the hypoblast, forms the wall of the
alimentary canal and its diverticula. The cells covering the
free surfaces of both layers, 7.e., the cells lining the body cavity,
become the peritoneum, from which, as we have already seen, the
ovaries and testes are formed.
M. Development of the Skeleton.
1. The Vertebral Column.
The earliest skeletal structure, and for a time the only one,
is the notochord, the development of which from the hypoblast.
of the mid-dorsal wall of the mesenteron has already been
described. It forms a cellular rod extending from the blastopore:
to the pituitary body ; and as the tail is formed, it extends back
into it. The notochord consist of vacuolated cells, filled with
fluid, and is invested by a delicate structureless sheath.
About the time of appearance of the hind legs, a delicate
skeletal tube, at first soft but soon becoming cartilaginous, is.
formed round the notochord from the mesoblast. This tube
grows upwards at the sides of the spinal cord, as a pair of longi--
 THE SKELETON, 145

tudinal ridges with which a series of cartilaginous arches, which

appeared at the sides of the spinal cord at a slightly earlier
stage, very soon become continuous.
By the appearance of transverse lines of demarcation, the
cartilaginous sheath of the notochord becomes cut up into a
series of nine vertebra, followed by a posterior unsegmented
portion, which later becomes the urostyle. This transverse
division does not affect the notochord, which remains as a
continuous structure until the complete absorption of the tail
at the end of the metamorphosis.
Shortly after the metamorphosis thin rings of bone, slightly
constricted in their centres, so as to be hour-glass-shaped in
section, are developed in the membrane investing the cartila-
ginous sheath of the notochord : these correspond with the nine
vertebree already present, and form the first rudiments of the
vertebral centra.
In the intervertebral regions, between the successive bony
rings, annular thickenings of the cartilaginous sheath occur,
which grow inwards so as to constrict and ultimately obliterate
the notochord. Each of these intervertebral rings becomes,
after the metamorphosis, divided into an anterior and a posterior
portion, which fuse with the bony centra of adjacent vertebree,
and ossify to form their articular ends.
From the circumference, and from the articular ends of each
vertebra, ossification gradually spreads inwards; but a small
portion of notochord persists in the middle of each centrum
for a long time, or even throughout life.
The vertebree are not placed opposite the myotomes, but
alternate with these ; so that each vertebra is acted on by two
myotomes on each side, one pulling it forwards, and the other
backwards.
The transverse processes are at first independent of the
corresponding vertebre, but very early fuse with them. They
extend into the septa between the myotomes, and probably
correspond to the ribs of other vertebrates.
The urostyle is the part of the axial skeleton behind the
vertebree ; it is not divided into vertebre at any stage in
development,
The anterior end of the notochord, imbedded in the base of
the skull, is gradually encroached on by the cartilage and bone
around it, and ultimately completely absorbed.
K
146. DEVELOPMENT OF THE FROG.

2. The Skull.
The skull of the tadpole consists almost entirely of cartilage ;
none of the bones of the skull, with the exception of the para-
sphenoid, appearing until nearly the time of the metamorphosis.
In the adult frog, this cartilaginous skull is replaced to a con-
siderable extent by cartilage-bone; while other bones primitively
distinct, and probably of dermal origin—the membrane-bones—
graft themselves on to it.
The three morphologically distinct elements of which the skull
consists (cf. p. 53) may with advantage be described separately.
a. The Cranium or brain case. This in its fully-formed con-
dition is an unsegmented cartilaginous tube, enclosing the brain:
it is developed as follows.
In the front part of the head a pair of longitudinal cartilagi-
nous bars, the trabecule cranii appear in tadpoles of about 10
mm. length: these grow back alongside the notochord as a
pair of horizontal parachordal rods.
The hinder ends of the trabecule are some little distance
apart, and between them is a space in which the pituitary body
lies. In front of this pituitary fossa, the trabeculze unite to
form a plate of cartilage, which underlies the anterior end of the
brain, and is produced into blunt processes at its outer angles.
The parachordals grow rapidly: they extend inwards so as to
meet each other both above and below the notochord, which
they now completely surround. The two parachordals soon fuse
together to form the basilar plate, which, with the trabecule,
forms a firm cartilaginous floor to the brain case. At their
hinder ends the parachordals grow upwards to form the side
walls of the cranium, and a little later bend inwards so as to
meet each other above the brain, and complete the occipital
part of the cranium. Further forwards the pituitary foramen
becomes closed by a thin plate of cartilage, and the lateral
margins of the parachordals and trabecule grow upwards so as
to form the side walls of the skull, the roof remaining im-
perfect in this region.
The first bone to be developed is the parasphenoid. The
exoccipitals, the frontals and parietals, which are’ at first
separate, and other bones soon follow; and by the time the
metamorphosis is complete and the tail absorbed, all the bones
of the adult cranium are present, except the sphenethmoid,
which does not appear till some months later.
 THE SKULL. 147

b. The Sense Capsules. The cartilaginous auditory capsules

appear in tadpoles of about 12 mm. length as thin shells of
cartilage investing the auditory vesicles. They are at first
quite independent of the cranium, but before the completion of
the opercular folds they fuse with the upgrowing parachordals
to form part of the side walls of the skull. The pro-otic appears
about the time of completion of the metamorphosis.
The optic capsules are thin shells of cartilage, forming part
of the sclerotic coats of the eyes. They arise about the same
time as the auditory capsules; and unlike the other sense
- capsules, they remain distinct from the cranium throughout life,
in order to secure mobility of the eye-balls.
The olfactory capsules are from their first appearance very
closely connected with the anterior ends of the trabecule, which
grow up between them to form the median vertical internasal
septum. They develope later than the auditory and optic
capsules.
c. The Visceral Skeleton. This consists of a series of carti-
laginous hoops developed within the visceral arches, and forming
a framework which surrounds and stiffens the walls of the
pharynx. Each hoop consists of right and left halves, which
are independent at their dorsal ends, but fused or closely con-
nected ventrally. There are in all six of these hoops or bars in
the mandibular arch, hyoidean arch, and the four branchial
arches respectively ; and they develope in order from before
backwards.
i. The mandibular bar, which is the largest of the series, lies
at first parallel to the others, i.e., perpendicular to the long axis
of the body. It very early, however, undergoes important
changes, and by the time the external gills are developed, and
before the appearance of the opercular folds, it has altered its
direction, and now runs almost horizontally forwards, parallel
to and below the trabecule.
It soon unites with the trabecule, both behind and in front of
the eyeball, the latter union being effected by a short transverse
bar of cartilage—the palato-pterygoid. In front of the palato-
pterygoid, the most anterior part of the mandibular bar becomes
segmented off as a short rod of cartilage, which is directed
upwards and forwards in the lower lip: it is known as Meckel’s
cartilage, and forms the basis of the lower jaw or mandible.
 148 DEVELOPMENT OF THE FROG,

In connection with the lips two pairs of small labial cartilages

appear, serving to support the horny jaws of the tadpole.
In the later stages the subocular or quadrate portion of the
mandibular bar acquires a very close connection at its hinder
end with the auditory capsule, and changes its direction, so
that in place of running horizontally forwards, it now runs
downwards and forwards. This change, which may be described
as a rotation backwards of the bar, causes lengthening of the
palato-pterygoid bar and of Meckel’s cartilage: these latter
become respectively the basis of the upper and lower jaws of the
tadpole, which are completed later on by the development of
the pterygoid, squamosal, maxilla and other bones.
This rotation backwards of the distal end of the quadrate,
with corresponding lengthening of the upper and lower jaws,
proceeds rapidly during and after the metamorphosis, so that
the quadrate, instead of being directed downwards and forwards,
soon runs vertically downwards, and later on downwards and
backwards as in the adult. (Cf. Fig. 10, p. 56.)
ii. The hyoid bar also undergoes important changes. At
first it is a wide band of cartilage placed nearly vertically in
the side wall of the pharynx, immediately behind the mandibular
bar. When the mandibular arch becomes horizontal the hyoid
forms a broad stout bar of cartilage, articulating at its upper
end with the subocular part of the mandibular arch, and con-
nected at its ventral end with the hyoid bar of the other side
by a small median basi-hyal plate in the floor of the mouth,
At the commencement of the metamorphosis the hyoid bar
becomes narrower, and begins to extend upwards towards the .
auditory capsule: and by the end of the metamorphosis this
upper part of the hyoid has become the long slender anterior
cornu of the hyoid, which acquires a loose connection at its
upper end with the cranium and with the quadrate cartilage.
The development of the columella is imperfectly known. It
consists of two elements, one of which—the stapes—is a small
plate of cartilage partially filling a hole, the fenestra ovalis,
which appears in the lower and outer wall of the auditory cap-
sule about the time the opercular folds are growing back over
the gills, The other portion of the columella is a small rod,
partly cartilage, partly bone, which does not appear till some
months after the completion of the metamorphosis, and which
fuses with the stapes at its inner end, while its outer end becomes
 THE URINARY SYSTEM. 149

connected with the tympanic membrane (cf. Fig. 11, p. 58): this
outer element of the columella is commonly regarded as formed
from the uppermost part of the hyoid arch, but appears to be
really quite independent of it.
iii. The branchial bars are at first simple flattened rods of
cartilage, independent of one another, but becoming early con-
nected with a median basi-branchial cartilage, which appears in
the floor of the mouth between the ventral ends of the first two
pairs of bars.
As the hind-legs appear, the branchial bars of each side coalesce
with one another both at their dorsal and their ventral ends:
they also become strongly curved, and together form a complex
basket-work supporting the gills. Later on, as the gills begin to
shrink, the branchial bars become more slender: their dorsal ends
disappear, while their ventral ends fuse with the basi-hyal and
basi-branchial cartilages, and together give rise to the body of
the hyoid and its posterior cornua.
N. The Development of the Urinary System.
1. General Account.
The excretory organs of the tadpole, during the early stages
of its existence, are the head kidneys or pronephra. These
are a pair of globular organs imbedded in the dorsal wall of the
body at its anterior end, immediately behind the constricted
neck region (Figs. 33 and 35 KP). Each head kidney is a
convoluted tube with glandular walls, opening into the body
cavity by three ciliated mouths or nephrostomes, and continued
back along the dorsal wall as the archinephric or segmental
duct, KA, to the hinder end of the body, where it joins with
the corresponding duct of the opposite side, and opens into the
cloaca.
The head kidneys and their ducts are well developed in the
tadpole at the time of hatching: they subsequently increase
considerably in size, and are the sole excretory organs of the
tadpole during its early stages. In tadpoles of about 12 mm.
length the adult kidneys or Wolffian bodies (Fig. 33, KM), begin
to form in the hinder part of the body as a series of paired
tubules, which grow towards and open into the segmental duct.
These Wolffian tubules rapidly increase in number, as well as
in size and complexity, and become bound together by connec-
tive tissue to form the compact Wolffian bodies or kidneys of
150 DEVELOPMENT OF THE FROG

the fully-formed tadpole (Fig. 35 KM). At the same time the

head kidneys diminish in size, and undergo degenerative changes,
LJ
Ry an ainiah

Fig. 35. A 40mm. tadpole dissected from the ventral surface to show
the heart, the branchial vessels, and the head kidneys and Wolffian
bodies. The tail has been cut off. x 5
A, dorsal aorta: AF1, AF3, afterent branchial vessels of first and third
branchial arches: AL, lingual artery: CG, carotid gland: EF1, EF3,
efterent branchial vessels of first and third branchial arches : F, fat body:
GM, glomerulus: KA, archinephric or segmental duct : KM, Wolffian
body : KP, pronephros or head idaney. now degenerating : LA, fore limb,
still within opercular cavity: L!, upper lip: Lu, lower lip: LP, hind
limb : OR, genital ridge : RT, truncus arteriosus : RV, ventricle : TC, cloaca:
TO, cesophagus, cut short : TR, rectal spout.
 THE URINARY SYSTEM. 151

and by the time of the metamorphosis (Fig. 36) have almost

completely disappeared. ‘The Wolffian bodies persist as the
kidneys of the frog; and by a series of further changes the
ureters and generative ducts of the adult become established.
2. The Head Kidney and its duct.
In tadpoles of about 34 mm. length, 7.e., some time before
hatching, a pair of longitudinal grooves appear along the inner
surface of the somatopleure, extending from the neck to the
hinder end of the body, and lying a little distance to the right
and left of the notochord. The lips of each groove soon meet
and fuse so as to convert the groove into a tube or duct. The
closure of the tube takes place from behind forwards, and at the
anterior end is effected imperfectly, three holes or nephrostomes,
one behind another, being left through which the tube opens
into the body cavity. As the embryo grows, the anterior end
of the duct becomes convoluted and twisted on itself to form a
ball, the three nephrostomes becoming at the same time
lengthened out into short tubes. This convoluted mass is the
head kidney or pronephros. The hinder part of the duct is
the archinephric or segmental duct ; it remains straight, or
nearly so, and shortly before the tadpole hatches acquires an
opening into the cloaca.
At the time of hatching, the excretory organs thus consist on
each side of (1) a head kidney, which is a convoluted tube
lined by a glandular epithelium, and opening into the anterior
end of the body cavity by three ciliated openings, the nephro-
stomes; and (2) the archinephric or segmental duct, which is
the posterior part of the tube, and runs back along the dorsal
body wall nearly straight to the cloaca, into which it opens.
The head kidney is closely surrounded by, indeed almost
imbedded in, the posterior cardinal vein, and it is from the
blood of this vein that the epithelial cells of the head kidney
tubules separate the excretory matters, which are then passed
down the duct to the exterior.
The head kidney continues to increase in size, the tubules
becoming still more convoluted, and lateral diverticula arising
from their sides, until the tadpole is about 12 mm. in length,
and the hind limbs are just commencing to appear. It remains
stationary for a time and then, in tadpoles of about 20 mm.
length, begins to degenerate: the tubules become obstructed:
152 DEVELOPMENT OF THE FROG.

some of them become collapsed, others for a time irregularly

dilated: the whole organ steadily diminishes in size, and in
tadpoles of 40 mm. (Fig. 85 KP) is not more than half its
former size. It now shrinks rapidly, and at the time of the
metamorphosis (Fig. 36 KP) has almost disappeared, all three
0

Fig. 36. A Tailed frog, near the close of the metamorphosis, dissected
a the ventral surface to shew the kidneys and reproductive organs.
x 4,
A, dorsal aorta: F, fat body: GM, glomerulus: KA, archinephric or
segmental duct: KM, Wolffian body: KP, head kidney, disappearing:
KU, ureter: O, mouth: OR, genital ridge: RV, tip of ventricle:
TO, cesophagus, cut short.
 THE URINARY SYSTEM. 153

nephrostomes having closed up, and the organ being reduced to a

few small pigmented and irregulary twisted tubules, which have
separated from the duct, and which soon disappear completely.
Opposite the head kidney an irregular sacculated outgrowth,
the glomerulus, arises from the aorta on each side (Figs. 31
and 33 GM): this appears first about the time of hatching, and
its development keeps pace with that of the head kidney. It
lies immediately opposite the nephrostomes, and very close to
these, though not touching them. It begins to diminish in size
about the same time as the head kidney. At the time of the
metamorphosis (Fig. 36 GM) it is very small, and after the first
year it can no longer be recognised. Its close relation to the
head kidney, and the fact that its growth and subsequent
degeneration, keep pace with those of the head kidney, point
to a close physiological connection between the two organs,
though it is not easy to imagine what precise function the
glomerulus subserves.
3. The Wolffian Body.
The Wolffian body, or kidney, first appears in tadpoles of
from 10 to 12 mm. in length. It arises on each side as a series
of small solid masses of mesoblast cells lying along the inner
side of the segmental! duct, between this and the aorta (Figs.
33 and 35). They develope from behind forwards, the hindmost
pair being a short distance in front of the cloaca, and the most
anterior ones about three segments behind the head kidney.
These solid masses soon become elongated into twisted rods,
which then become tubular, and growing towards the segmental
duct meet and open into it. At their opposite ends these
Wolffian tubules, as they are termed, dilate into bulb-like
expansions, which become doubled up by ingrowth of little
knots of bloodvessels, derived from the dorsal aorta, and so
form Malpighian bodies. From the necks of the Malpighian
bodies, short solid rods of cells grow towards the peritoneal
epithelium and fuse with it. These rods soon become hollow,
and open into the body cavity by ciliated funnel-shaped mouths
or nephrostomes: their opposite ends break away from the
Wolffian tubules and open directly into the renal veins on the
ventral surface of the kidney. The Wolffian tubules rapidly
increase in number; they also branch freely, and so give rise to
a complicated system of glandular tubules, which, when bound
together by bloodvessels and connective tissue, form the
154 DEVELOPMENT OF THE FROG.

Wolffian body or kidney of the frog. The nephrostomes persist:

and in the adult frog as many as 200 or more are present on
the ventral surface of the kidney, as minute funnel-like ciliated
openings, leading by short tubes into the renal veins.
4, The Wolffian and Mullerian ducts.
So far we have only described one duct on each side, the
seomental duct, which acts as the excretory duct first of the
head kidney, and then of the Wolffian body as well. We have
now to consider in what way the ureters and generative ducts of
the adult frog are formed.
About the time of the metamorphosis the head kidney, which
has become rudimentary, separates completely from the duct,
which now ends blindly a short distance in front of the Wolffian
body.
A little later, after completion of the metamorphosis and
entire disappearance of the tail, this anterior end of the
segmental duct, in front of the Wolffian body, becomes divided
somewhat obliquely into two; an anterior part, which is now
isolated from the Wolffian body, and will be called the
Mullerian duct; and a posterior part the Wolffian duct, which
is simply the posterior part of the original segmental duct, and
receives the Wolffian tubules of the kidney.
The Mullerian duct becomes connected infront with the
peritoneal epithelium, and acquires an opening into the anterior
end of the body cavity. At its hinder end it grows back along
the outer side of the Wolffian duct to the cloaca, into which it
opens. So far the changes are the same in both sexes. In the
male frog the Miillerian duct persists in this condition through-
out life, and may be recognised as a slender longitudinal streak
lying in the thickness of the peritoneum a short distance to the
outer side of the kidney, and extending some distance in front
of it. In the female frog the Miillerian duct becomes the
oviduct, the anterior opening being carried forward first as a
groove, and then by closure of the lips as a tube, to the position
characteristic of the peritoneal opening of the adult oviduct;
while the posterior part becomes greatly convoluted and acquires
thick glandular walls: the hindmost part of the oviduct remains
thinner walled, but of much greater capacity.
The Wolffian duct becomes in both sexes the ureter. In
the female frog it undergoes no further change of importance.
In the male frog the hinder end of the Wolffian duct becomes
 THE URINARY SYSTEM. 155

dilated into a much-branched glandular enlargement, the

vesicula seminalis.
5. The Vasa Efferentia.
In both sexes at an early stage, as the Malpighian bodies
are forming in the Wolffian body, those nearest to the genital
ridges give off tubular branches from their capsules into the
ridges.
In the female frog these tubules are said to expand very
greatly, and to give rise to the chambers or cavities present in
the adult ovary: but the point is not established with certainty.
In the male frog these tubules become the vasa efferentia:
they become connected with the spermatic tubules, and, as at
their other ends they open into the Wolffian tubules, they form
passages along which the spermatozoa can get from the testis
to the Wolffian duct or ureter, and so out.
 ; ie
AV rae
~
“s
iV = oe
“ % .:

ap ane
4 j
a pe
CF a : ia |b

d
j
.
Tai
a: "4
s

W's
Jf »

i LY
}

, ‘

iv

>

' b
Ye .

’ ar

A »
. Sy
ayer \** oF
 157

INDEX.

Abdominal viscera, 16—18 peroneal, 30

Acetabulum, 62 posterior mesenteric, 29
Adipose tissue, 46 pulmonary, 30, 139
Afferent nerves, 78 sciatic, 30
Alimentary canal, 19 splenic, 29
development of, 132—134 subclavian, 29
Ampulla, 105 tibial, 30
Ankle, 63 urino-genital, 29
Annulus tympanicus, 57 vertebral, 29
Aorta, 28, 136 Articular cartilage, 48
Aortic arch, 28—31, 136—143 Articular process, 52
Aperture, cloacal, 14, 134 Atlas, 53
bP) external, 14—15 Auditory capsule, 53, 55, 147
a9 median, 14 Auditory organ, 1 105
3 paired, 15 Auricle, 22, 32
Aponeurosis, 65 Axial skeleton, 52—59
Apparatus, 1—2 Axis cylinder, 94.
Appendicular skeleton, 59—64
Aqueductus Sylvii, 83, 129 Back-bone, 52
Aqueous humour, 98, 99 Basi-branchial, 149
Archinephric duct, 151 Basi-hyal, 148
Areolar tissue, 46 Basilar plate, 146
Arteries, 27—30 Bile-duct, 20, 133
Artery Bladder, gall, 20, 133
anterior mesenteric, 29 Bladder, urinary, 17, 107, 108, 133
carotid, 28 Blastopore, 122, 134
coeliac, 29 Blind spot, 101
cceliaco-mesenteric, 29 Blood, 85—37
cutaneous, 30 Blood corpuscles, 35, 36, 37
dorsal aorta, 28 Blood plasma, 35
epigastric, 30 Body-cavity, 123, 143
external carotid, 28 Bone, structure of, 48—49
gastric, 29 Bone
hemorrhoidal, 30 angulo-splenial, 57
hepatic, 29 astragalus, 63
hypogastric, 30 calcaneum, 63
iliac, 28, 30 carpal, 61
internal carotid, 28 clavicle, 60
laryngeal, 28 columella, 58, 105, 148
lingual, 28 coracoid, 60
lumbar, 29 dentary, 58
mesenteric, 29 ex-occipital, 54
occipital, 29 femur, 63
occipito-vertebral, 29 fronto-parietal, 54
cesophageal, 28 girdle, 54
158 INDEX.

humerus, 61 Cilia, 40
ilium, 62 Ciliary movement, 40
ischium, 63 Ciliary muscle, 100
maxilla, 57 Ciliary nerves, 100
mento-Meckelian, 58 Ciliary processes, 100
metacarpal, 61 Ciliary vessels, 100
metatarsal, 63 Circulation of blood, 36—37
nasal, 55 Circulation in tadpole, 136—143
omosternum, 60 Cloaca, 19, 106—108
os cruris, 63 Cloacal aperture, 14, 134
palatine, 56 Cochlea, 104
parasphenoid, 55, 146 Ccelom, 123, 143—144
phalanges, 61, 62, 63, 64 Columella, 58, 105, 148
pre-coracoid, 60 Condyle, occipital, 54
premaxilla, 57 Conjunctiva, 98
pro-otic, 55 Connective tissue, 45—47
pterygoid, 56 Contraction of muscle, 65
pubes, 62 Coracoid foramen, 60
quadrato-jugal, 57 Cornea, 97, 98
radio-ulna, 61 Corpora, adiposa, 17
scapula, 60 Cranial flexure, 128
sphen-ethmoid, 54, 146 Cranial nerves, 87—93, 129
squamosal, 57 Cranium, 53—55, 146
sternum, 60 Crura cerebri, 84, 129
supra-scrapula, 60
tarsal, 63 Dehydration, 11
tibio-fibula, 63 Development, 109—155
vomer, 55 General account, 109—113
Brain, 80—84 Detailed Account, 113—155
5, development of, 128—130 of nervous system, 124—130
Branchial arch, 135—139 of sense organs, 130—132
Branchial chamber, 135 of alimentary canal, 132—134
Branchial cleft, 135 of gill arches and clefts,135—136
Buccal cavity, 15—16 of circulatory system, 186—143
of ccelom, 143—144
Calcar, 64 of skeleton, 144—149
Canal, semicircular, 105 of urinary system, 149—155
Canaliculi, 49 Digestive organs, 18—20
Capillaries, 22, 36, 37 Dissection, 2—3
Capsule, auditory, 53, 55 Distal, 61
, olfactory, 53, 55 Drawing, 3
Cardiac plexus, 87 Duct, bile, 20, 133
Carotid arch, 28 Duodenum, 19
Cartilage, 47—48
Cartilage-bone, 50 Ear, 104—105, 131
Cauda equina, 85 Efferent nerves, 78
Cavities of brain, 82—83 Egg, 109
Cell, 38 fertilisation of, 116—117
Central canal of cord, 96 formation of, 113—114
Centrum, 52 segmentation of, 117—120
Cerebellum, 82, 128 Elastic tissue, 45
Cerebral hemisphere, 80, 129 Epiblast, 121, 123
Cerebral vesicle, 129 epidermic layer of, 125
Chiasma, optic, 83 nervous layer of, 125
Choroid, 98, 100, 101 Epicoracoid, 60
Choroid plexus, 81, 129 Epiphysis, 61
 INDEX. 159

Episternum, 60 Hallux, 64
Epithelium, 38—43 Hardening, 10—11
ciliated, 40 Haversian system, 48
columnar, 39 Head, 14
glandular, 40—43 Head kidney, 149, 151—153
squamous, 38—39 Heart, 17, 22—23, 30—33
stratified, 40 development of, 136
Eustachian passage, 15, 105, 132 pulsation of, 23
External characters, 13—15 Hepatic plexus, 87
Kye, 97—103, 130—131 Hepatic portal system, 26—27
Eye of frog, 97—98, 102—103 Hind-brain, 128
Eye of ox, 98—101 Hind-limb, 14, 63—64
Histology, 38—49, 94—96, 102—103
Fat-body, 17, 113 Hyaline cartilage, 47
Fat-cells, 46 Hyoid, 15, 538, 58—59
Female organs, 107—108 Hyoid arch, 136, 149—150
Female pro-nucleus, 116 Hyomandibular cleft, 132, 135
Fenestra ovalis, 59 Hypoblast, 123, 124
Fertilisation, 116—117
Fibrous tissue, 45—46 Imbedding, 11-12
Filum terminale, 84
Impregnation, 116—117
Fissure of cord, 95 Infundibulum, 83, 129
Fontanelle, 54
Insertion of muscle, 65
Food-yolk, 111, 114, 119 Intercellular substance, 38
Foramen, intervertebral, 52 Intestine, 17, 19, 41
magnum, 54
of Monro, 83, 129
Tris, 97, 98
Fore-brain, 128, 129 Iter, 83
Fore-limb, 14, 61—62
Fourth ventricle, 83, 128 Jaw, 53, 56—58, 111, 147—148
lower, 15, 57—58
Gall bladder, 20, 133 upper, 56—57
Gasserian ganglion, 89
General Anatomy, 13—21
Genital plexus, 87 Kidney, 20, 149—154
Genital ridge, 113
Germinal layers, 120—124 Labial cartilage, 148
Germinal spot, 114 Laboratory rules, 1
Germinal vesicle, 114 Lacuna, 48
Gill arches, 135—136 Laryngeal chamber, 133
Gill clefts, 111, 135 Lateral plate, 143, 144
Gills, external, 109—111, 135 Lateral ventricle, 83, 129
internal, 111, 135 Law of Recapitulation, 112
Gland, 40—43 Lens, 97, 99, 101, 131
carotid, 28 Ligamentum nuche, 45
compound, 41 Limbs, 14, 111—112
gastric, 43 skeleton of, 61—62, 683—64
racemose, 41 Linea alba, 66
simple, 41 Lips, 112
thyroid, 24 Liquor sanguinis, 35
tubular, 41 Liver, 17, 19, 133
Glenoid cavity, 60 Lower layer cells, 121]
Glottis, 16 Lung, 17, 112, 133
Grey matter, 95 Lymphatic system, 34
Lymph heart, 34
Hemorrhoidal plexus, 87 _ Lymph sacs, 34
160 INDEX,

Maceration, 7—8 involuntary, 44

Male organs, 106—107 latissimus dorsi, 67
Male, pro-nucleus, 116 levator anguli scapule, 67
Malpighian body, 41, 153 levator bulbi, 70 »
Mandibular arch, 57—58, 135, 147— masseter, 69
148 mylohyoid, 68
Marrow, 49 non-striated, 44
Maxillary arch, 56—57 obliquus externus, 66
Maxillary teeth, 15 obliquus inferior, 71
Meckel’s cartilage, 57, 147—148 obliquus internus, 66
Medulla oblongata, 82 obliquus superior, 71
Medullary cavity, 49 obturator, 76
Medullary sheath, 94 pectineus, 75
Medullated nerves, 94 pectoralis, 66
Membrane bone, 50 peroneus, 77
Mesenteron, 122—123, 132 petrohyoid, 69
Mesentery, 18 pterygoideus, 69
Mesoblast, 123, 124 pyritormis, 74
Metamorphosis, 112, 142—143 quadratus femoris, 75
Methods, hardening, 10—11 rectus abdominis, 66
imbedding, 11—12 rectus anticus femoris, 74
macerating, 7—8 rectus externus, 70
mounting, 6—7 rectus inferior, 71
section-cutting, 12 rectus internus, 70
staining, 8—10 rectus internus major, 72
teasing, 7 rectus internus minor, 72
Microscope, 4—6 rectus superior, 70
Mid-brain, 128, 129 retractor bulbi, 71
Migration of blood corpuscles, 37 retrahens scapule, 67
Mounting media, 6—7 sartorious, 72
Mounting, methods of, 6—7 semimembranosus, 74
Mouth, 14, 15—16 semitendinosus, 75
Miillerian duct, 154 sternohyoid, 68
Miiller’s fibres, 103 striated, 44
Muscles, of head, 68—71 structure of, 44
of hind-limb, 71—77 submandibular, 68
of trunk, 66—68 temporalis, 69
Muscles, adductor brevis, 75 tibialis anticus, 76
adductor longus, 72 tibialis posticus, 76
adductor magnus, 72 tricepsextensor femoris 72
biceps, 74 vastus externus, 74
ciliary, 100 vastus internus, 74
cucullaris, 67 voluntary, 44
depressor palpebre infer- Muscular system, 65—77
ioris, 70 Myotome, 143—144
depressor mandibule, 69
extensor cruris, 77 Nares, anterior, 15, 130
sed dorsi communis, posterior, 15, 130
Nephrostome, 151, 154
gastrocnemius, 76 Nerve, abducens, 90
geniohyoid, auditory, 92
gluteus, 68, 74 brachial, 85
hyoglossus, 68 ciliary, 100
ilio-psoas, 75 coccygeal, 86
infraspinatus, 67 coraco-clavicular, 85:
intertransversales, 68 cranial, 87—93, 129;
 INDEX. 161

crural, 86 Origin of muscle, 65

facial, 90 Ovary, 17, 107, 113
fourth, 89 - | Oviduct, 18, 108
glosso-pharyngeal, 92
hypoglossal, 85 Palato-pterygoid, 147
ileo-hypogastric, 86 Pancreas, 20, 1é
motor oculi, 88 Parachordal, 146
olfactory, 87 Pectoral girdle, 16, 59—60
optic, 88 Pelvic girdle, 62
pathetic, 89 Pericardial cavity, 22, 144
peroneal, 86 Perimedullary lamelle, 49
pneumo-gastric, 93 Perineurium, 94.
radial, 85 Periotic capsule, 104
sciatic, 86 Peripheral lamelle, 49
spinal, 84—87, 130 Peripheral nervous system, 84—93,
sympathetic, 87, 93 129
tibial, 86 Peritoneum, 18, 144
trigeminal, 89 Pia mater, 80
ulnar, 85 Pineal body, 81, 129
vagus, 93 Pituitary body, 84, 134
Nerve cells, 95, 96 Polar bodies, 116
Nerve fibres, 94, 95 Pollex, 61
Nerve roots, 96 Portal system, 25—27
Nervous system, 78—96 Post-anal gut, 133
development of, 124—130 Post-axial surface, 71
histology of, 94—96 Preaxial surface, 71
Neural arch, 52 Primitive sheath, 94
Neural canal, 52 Proctodeum, 132, 134
Neural fold, 127 Pronephros, 149, 151—153
Neural groove, 126 Pro-nucleus, 116
Neural plate, 125 Proximal, 61
Neural spine, 52 Pulmo-cutaneous arch, 30
Neural tube, 126 Pupil, 97, 98
Neurenteric canal, 128 Belorus, 19
Neuroglia, 96
Nodes of Ranvier, 94 Quadrate, 57, 148
- Non-medullated nerves, 95
Nose, 131 Reagents, hardening, 10—11
Nostril, 15 macerating, 7—8
Notochord, 123 mounting, 6—7
staining, 8—10
Occipital condyle, 54 Recapitulation, 112
Occipito-atlantal membrane, 53 Renal plexus, 87
Oesophagus, 19 Renal portal system, 26
Olecranon process, 61 Reproductive organs, 106—108
Olfactory capsule, 53, 55, 147 Retina, 98, 101, 102—103
Olfactory lobe, 80, 129 Rhinal process, 55
Olfactory organ, 130 Rods and cones, 103
Operculum, 111, 135 Rules of Laboratory, 1
Optic capsule, 147 Rules for drawing, 3
Optic chiasma, 83 for dissection, 2—3
Optic cup, 131 for use of microscope, 5—6
Optic lobe, 81, 129
Optic thalami, 81, 129 Sacculus, 104
Optic vesicle, 128, 130 Sacrum, 53
Ora serrata, 100 Sarcolemma, 44
L
162 INDEX.

Sclerotic, 97, 98, 102 Transverse process, 52

Section cutting, 12 Truncus arteriosus, 22, 33
Segmental duct, 151 Tuber cinereum, 83
Segmentation cavity, 118, 123 Tympanic cavity, 15, 105
Segmentation nucleus, 117 Tympanic membrane, 14, 15, 105, 132
Segmentation of the egg, 117—120
Semicircular canal, 105 Upper layer cells, 120
Sense capsules, 55, 147 Ureter, 21, 106, 107, 108, 154
Sense organs, 130—132 Urostyle, 52, 53, 145
Sheath of Schwann, 94 Utriculus, 104
Shoulder girdle, 16 Uvea, 100
Sinus venosus, 23, 24—25, 30—31
Skeleton, 50—64 Vas deferens, 106, 107
appendicular, 59—64 Vasa efferentia, 106, 155
axial, 52—59 Vascular system, 22—37
Skin, 14 development of, 136—143
Skull, 53-59 Vein, 23—27, 36
development of, 146—149 anterior vena cava, 24
Somatopleure, 143 brachial, 24
Solar plexus, 87 cardiac, 27
Spawn, 109 dorso-lumbar, 26
Spermatozoa, 106, 109, 116 external jugular, 24
Splanchnopleure, 143 femoral,
Spleen, 21 gastric, 27
Splitting of mesoblast, 123, 144 hepatic, 25
Spinal cord, 84, 95—96 hepatic portal, 27
development of, 125—126 innominate,
Spinal ganglia, 86 internal jugular, 24
Spinal nerves, 84—86 intestinal, 27
Spinous process, 52 lingual, 24
Staining reagents, 8—10 mandibular, 24
Stapes, 148 musculo-cutaneous, 24
Stomach, 19 ovarian, 25
Stomodzeum, 132, 134 parietal, 26
Subcutaneous tissue, 46 pelvic, 26
Subocular bar, 148 portal, 25—27
Sucker, 111 posterior vena cava, 25
Suspensorium, 57 pulmonary, 25
Suspensory ligament, 101 renal, 25
Sympathetic nervous system, 87, 93 renal portal, 26
Symphysis, 62 sciatic, 26
Systemic arch, 28—29 splenic, 27
Systole, 23 spermatic, 25
subclavian, 24
subscapular, 24
vesical, 26
Ventral fissure, 84
Teeth, 15, 55, 57 Ventricle, of brain, 83, 128, 129
Tendo Achillis, 76 Ventricle, of heart, 22,
Tendon, Vertebree, 52—53
Testis, 18, 106, 113 Vertebral column, 52-—53
Thalamencephalon, 81, 129 development of, 144—145
Third ventricle, 83, 129 Vertebral plate, 143
Thyroid gland, 24 Vesical plexus, 87
Tongue, 16 Vesicula seminalis, 21, 106, 155
Trabeculz cranii, 146 Vestibule, 104
 INDEX. 163
Viscera, abdominal, 16—17 Wrist, 61
Visceral arches, 135
Visceral clefts, 135 Xiphisternum, 60
Visceral skeleton, 147—149
Vitelline membrane, 114 Yellow elastic tissue, 45
Vitreous humour, 98 Yolk, 111, 114, 119—120
Vomerine teeth, 15 Yolk cells, 121
Yolk hypoblast, 124
White fibrous tissue, 45, 46 Yolk plug, 122
White matter, 95
Wolffian body, 153—154 Zonule of Zinn, 101
Wolffian duct, 154—155 Zygapophysis, 52
cya,
WORKS BY THE SAME AUTHOR.
With numerous Illustrations. 8vo. 21s.

Vertebrate Embryology: A Text-Book for Students and

Practitioners. By A. MILNES MARSHALL, M.D., D.Sc., M.A.,
F.R.S., Professor in the Victoria University ; Beyer Professor
of Zoology
aS)
in Owens College; late Fellow of St. John’s College,
Cambridge, Xe.
‘* The book is welcome, and we do not most valuable, clear, and masterly expo-
doubt but that it will obtain the popu- sition of the known facts of the develop-
larity which it so well deserves. The mental history of leading types of
care and accuracy with which it has been vertebrata.”—NATURE.
written will render it most useful to the ‘“We recommend the work with con-
student and to the teacher.”—THE fidence to students and practitioners of
BRITISH MEDICAL JOURNAL. medicine . The illustrations are
“Though embryology has been much throughout good; they are mostly origi-
in the air of late years, no complete nal, and have been executed in a uniform
account has been given of the develop- style, with a uniform system of lettering,
ment of the common frog or rabbit. Such making reference easy.” — MEDICAL
an account is here provided..... The CHRONICLE.
final chapter deals with the human ‘Tn the art of writing text-books Pro-
embryo, and sums up the present state of fessor Marshall has, in our opinion, no
knowledge in a manner which will be superior; he puts his facts in the clearest
equally valuable to the student of medicine and shortest way possible, and the dia-
and the practitioner.” THE SPECTATOR. grams (largely original) are always perfect
“Will be extremely useful to all models of lucidity.” —SATURDAY REVIEW.
teachers and students of biology ...A

DR. MILNES MARSHALL and C. HERBERT HURST.

A Junior Course of Practical Zoology. By A. Mitnus
MARSHALL, M.D., D.Sce., M.A., F.R.S., Professor in the Victoria
University ; Beyer Professor of Zoology in Owens College;
late Fellow of St. John’s College, Cambridge. Assisted by
C. HERBERT Hurst, Ph.D., Lecturer in the Victoria University ;
Demonstrator and Assistant Lecturer in Zoology, Owens College,
Manchester. Third Edition, Revised. With additional [lus-
trations. Crown 8vo. 10s. 6d.
“This book cannot fail to be of great a form demanding our sincere thanks.”—
value to those who are studying zoology NATURE.
in their laboratory work; and to such we
have great pleasure in strongly recom-
‘‘This work in plan and executionis
mendingit.”—Lonpon MEpIcAL RECORD. admirable, and is one of the best books,
if not the best, of its kind. The directions
“A most successful and important for dissection are clear, the descriptions
book. . . . The illustrations are excellent, are well arranged. The drawings and
reflecting the greatest credit upon all index greatly enhance the value of the
concerned . .. It is provided with an book.” EDINBURGH MEDICALJOURNAL.
exceedingly good index, and presented in

Lonpon: SMITH, ELDER, & CO., 15, Watertoo Puace.

 BIOLOGICAL
LECTURES AND- ADDRESSES
DELIVERED BY THE LATE

AKTHURK MILNES MARSHALL,

M.A., M.D., D.Sc., F.R.S.

EDITED BY

CovFo MARSBALL, M.D. B.dcy PR eae

LONDON :
DAVID NUTT & CO.
1894.

Price Ios. each.

Studies from the Biological

Laboratories of the Owens College.
VOLUMES 1 and 2.

PUBLISHED BY THE
COUMUNGIL, OF; THE COLLEGE
AND EDITED BY THE LATE

Proressor A. MILNES MARSHALL.

MANCHESTER: J. E. CORNISH, 16, ST. ANN’S SQUARE.

 y
as |
Wikre Oe
y
oa
 ti a
anes a
Ma

oe ie
 v if ie Be 1 4Dix

man ii ti iA i
ai i

4
ae
)
pao
oe é)
 nee
nie
Pi
oe M
on
- ¥ ce .
Os Al y ‘
EN hs ‘¢ “” ’ ¥

ule © —s
_ \ a 4 i 4 Orsi
ar fon ee oe ; eu ake I ‘ aes ‘Sie bs ‘ a? f Py
ort. wa¥
omy, ie, Whig eyes iatye?
Me! 7 } hla
ve 5 eh, i, " ae
Pea- {3
Agee)Se ik t. Ama
ve iy
; . v at bh en ry ;
MONEx ¥ ‘ : eae a | :
Ary =
oe achat. ied peed
‘
ty 1s} ; Ay “ra i: tty aX *
ae ‘
‘, AL v7
* ma SNae
s ne ed aa“sa
. hy, y wai ‘