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THE FROG:
AN INTRODUCTION TO

ANATOMY, HISTOLOGY, AND EMBRYOLOGY.

BY A.

MILNES MARSHALL,

t

M.D., D.Sc, M.A., F.R.S.,
COLLEGE, CAMBRIDGE;
;

LATE FELLOW OF

ST. JOHN's

PROFESSOR IN THE VICTORIA UNIVERSITY

BEYER PROFESSOR OF ZOOLOGY IN OWENS COLLEGE.

mttb

BDftion.

Revised and Illustrated.

MANCHESTER
CORNISH,
ITH ELDER, &
i6,

ST.

ANN'S SQUARE
PLACE.

LONDON CO., WATERLOO
1894.

.

.

have throughout been printed in is italics. from small size or for other reason. Where. The Owens sity. which forms part of the scheme of study prescribed by the Victoria Univerof a rather more extended and comprehensive nature than the courses held elsewhere under the same name.PREFACE TO THE FIRST EDITION. is College Course of Elementary Biology. and in order to definitely stamp the practical character of the work. fairly typical and a example of the great group its of Vertebrate animals.. etc. little is covered by no one of the It is to meet this want that the present work has been prepared. followed by the application of these methods to the examination. proved unsuitable. other animals have been substituted for For convenience of reference. the frog has it. the whole ground of which existing manuals. 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. both anatomical and of histological. containing practical instruction in the methods employed in biological investigation . It not expected that the student should do the whole of . first This instalment of the work consists of an Introduction. convenient to dissect. For this purpose the frog has been selected as being easy to obtain. an actual animal. directions for dissection.

1882. I am much indebted to Prof. Hartog. Waters for the important help they have given me in the histological portions. H. Demonstrator and from my friend and pupil Mr. In preparing this assistance College. also part I have received very valuable of Biology in the from Dr. Owens College. Hurst. C. August. Gamgee and to Mr.IV the work here given the section of the muscles first time he goes over it. The dis- and of the cranial nerves should only is be attempted if time remain after the other work first completed. .

revised. A. H. Giles. Since the appearance of the former editions. as the mode of treatment more elementary. I is concerned. my and former pupils. and in some ways different from that of the larger book. Hurst and myself has completed.PREFACE TO THE THIRD EDITION. Mr. 1888. have retained The Frog in is its original form. E. excellent I been added. Sidebotham. advantage of In preparing have had the of some series sections mounted by and Mr. . the pubhcation of the Junior Course of Practical Zoology by Dr. so far as Animal Morphology the Owens College Course of Elementary Biology. July. The present of the this edition has been carefully and an account examining friends development of the Frog has account. Owens College.

The principal changes in the present edition are in the chapter intro- on Embryology. In revising the book friends Dr.VI PREFACE TO THE FIFTH EDITION. in which some new figures have been duced and some mistakes corrected. 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. December^ 1893. of any Herbert Hurst. I have received valuable aid from my in Hurst and Mr. made no change C. . NOTE. In correcting the proofs I have importance. April. Gamble. Owens College. 1894. Assistant Lecturers Zoology in the College. Owens College.

Glands. Drawing. Peritoneum. I. Epithelium. Introduction. System. The Arteries.. General Anatomy of the Frog. Dissection. The Peripheral Nervous 78-96 Histology of Nerves . 38-49 Bone CHAP. Elementary Histology. Heart. Use of the Microscope. Cartilage. Digestive Organs Abdominal Viscera. The Vascular System of the Frog. 65-77 CHAP. Muscles of Head. Buccal Cavity. II. The Appendicular Skeleton 50-64 CHAP. The Structure of the Microscopical Examination of Blood 22-37 CHAP. IV. VI. The Heart. 13-21 CHAP. The Skeleton of the Frog. The Nervous System of the Frog. The Axial Skeleton. III. Laboratory Rules.. Apparatus required. Muscle. The Central Nervous System. V. Muscles of Trunk. The Veins. The Muscular System op the Frog.CONTENTS. Preparation of Microscopical Objects. External Characters. Muscles of Hind-limb. Connective Tissue. page Section cutting 1-12 CHAP.

VII.Vlll CHAP. The Gill Clefts and Arches. Development of the Sense Organs. Maturation of the of the Egg. The Eye of the Sheep The Ear of the Frog or Ox. The Vascular System. 106-108 CHAP. Formation of the Development of the Nervous Germinal Layers. Histology 97-105 CHAP. System. The Muscular System Development of the Skeleton. Development of the Frog. The Eye of the Frog. The Eye and Ear. The Male Frog. and the Ccelom. Egg. of the Eye. The Reproductive Organs and the Cloaca. IX. General Account. Segmentation. INDEX 157-163 . 109-155 Development of the Urinary System Formation Fertilisation. Development of the Alimentary Canal. The Female Frog VIII.

4. as explained in the next section. one large Two Two A B . Each student has the free use of the drawer and locker belonging to his seat the key may be obtained on j)ayment of a deposit of half a crown. : APPARATUS REaUIRED. The latter pair should have the blades bent at an angle (elbow scissors). which will be returned if the key is given up before the end of the term. firmly mounted in handles. Microscopes are provided by the College at a charge of five shillings per term. note book. fol- is required to provide himself with the or three scalpels or dissecting knives of different sizes. The Laboratory is open to members of the Biology Class from 10-30 to 5 on Tuesdays and Thursdays. and pencil. II. Each student has a definite seat assigned him in the Laboratory. 1. which he is not allowed to change without per- mission. 2. 4. All necessary reagents and specimens for dissection are provided by the Laboratory. pair of stout needles. I. Both pairs should be straight.-LABORATORY RULES. Tavo pairs of scissors . 3. Students are required to attend at least three hours on each of these days. but each student is required to furnish himself with dissecting instruments. pairs of forceps. one pair large and strong. 3. for cutting bone and other hard tissues . and one small. for fine dissections. but otherwise will be : forfeited. 2.INTRODUCTION. In selecting scissors be careful tosee that they cut quite up to the points of the blades. and should have the tips roughened in order to secure a firmer hold. the other pair small.-LIST OF Each student lowing 1.

. A pocket lens. The coverslips. Dissection consists mainly in removing the " connective tissue " which binds the several parts together. and a glass cannula with india-rubber 7. or when dissecting the muscles of the leg. and bent at an angle half an inch from the end. Never cut away anything until you are quite certain what it is you are removing.g. for drawing in an HB pencil. In cleaning bloodvessels or nerves always dissect along them and not across them . A seeker. but obliquely. 4. e. 9. 5. pin out the leg in such a position as to stretch the muscles you are cleaning. containing two or three lenses mounted in a handle. In pinning out a dissection stick the pins in. Slides and mens.. . ON DISSECTION. 1). Pin down the animal firmly to the dissecting board. 3. A metal blow ])ij)G of the finest : : A pair cap.-ON DISSECTION. 10. distend it by passing a small roll of paper or the handle of a seeker down the oesophagus . . A blank note book. for measuring the dissections. : 2. The object of dissection is to separate the several parts and organs from one another. and a piece of india-rubber.e. a blunt needle mounted in a handle. so as to define their boundaries and display clearly their mutual relations. 11. A III. Similarly when cleaning muscles. They are used for teasing histological preparations. so that their heads do not get in the way or obscure the dissection. Never attempt to dissect a specimen that is not so fixed. for mounting microscopical specicoverslips should be the thinnest sold (No. when dissecting the bloodvessels or nerves of the throat. The following rules should be carefully attended to 1. and avoid laying hold of them with the forceps. cheap pair of compasses. sewing needles. not vertically. mounted in handles only about a quarter of an inch of the needle should 2^1'oject. dissect along their fibres and not across them. Put the part you are dissecting slightly on the stretch . 6.2 5. and giving wdien combined a magnifying power of at least six diameters. 8. i.

4. and so on. IV. and draw every dissection you make... e. Draw on one side of the page only tion of your drawing on the opposite page. may be. 9. Keep certain colours for particular organs or tissues . when drawing the skeleton colour the cartilage . 6. if half the size. object mark it x 2 and write an explana5. which supports the parts and greatly facilitates the operation. If the object you are drawing is bilaterally symmetrical. stream' of water allowed to play gently on the dissection from time to time is often a valuable aid. A 8. stop and w^ash the dissection thoroughly under the tap before proceeding further. and this must on no account be omitted. i. 6 6. : . and still more of nerves. If you get in a muddle.g.ON DRAWING. and w^ater-colour paints still better. either the exact size of the natural object.e. The dissection of muscles. Fine dissections should be done under water. In commencing a drawing. and mark also the scale adopted. and sketch in lightly the whole outline before finishing any one part. Be careful not to blunt your fine scissors or scalpel by using them for cutting hard parts. Coloured pencils are very useful. 3. Make your drawing to scale. Keep a separate book for your Do not be drawings.-ON DRAWING. 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. Keep your instruments clean and sharp. discouraged if you find it difficult at first you will never regret : time spent on it. as the : case 2. or half or double or treble that size. is greatly aided by placing the specimens in spirit for a day before dissecting:. — . The following rules will be useful to those w^ho have not learnt drawing systematically 1. draw a faint line down the middle of your paper. 7. first determine by careful measurement the positions of the principal points. . Name on your drawing the several parts shown. If your drawing be of the if it be double the size of the natural size mark it thus x 1 x ^. It is absolutely essential to draw your dissections.

The higher the power employed the As the closer must the objective be brought to the object.4 THE MICROSCOPE. according to their magnifying power. directed on the object to be examined by means of a mirror Above t-he stage the attached to the stand below the stage. position of the object on the stage of the microscope is fixed. The stage is perforated in the middle by a hole. and can only be used Avhen low powers are being employed. known as the eyepiece. and to a slight extent with different observers. this distance varying with different objectives. consists essentially of a stand and a hody. the lower end of which is A little way attached to a heavy foot to ensure steadiness. in which the body of the microscope slides up and down. loiv 'power Similarly eyepieces are spoken of as high or low a liigh power. The body is a tube. stand supports a vertical tube. blue. : : . V. (1) By simply sliding the body up and down by hand is known as the coarse adjustment . while to the lower end is screwed another combination of lenses the objective. in the upper end of which is placed a combination of lenses. and the membrane bones either red or white .-THE USE OP THE MICROSCOPE. The microscope latter of — — . A microscope is usually provided with a couple of eyepieces and a cou]3le of objectives of different magnifying power. In order that an object may be seen clearl}^ the objective must be at a certain definite distance from the object. the size of which can be Through this hole light is varied by means of diaphragms. The stand is an upright pillar. this distance is regulated by moving the body of the microscope up and down in the tube in which it slides. when drawing the bloodvessels colour tlie arteries red and the veins blue. it should be performed with a slight screwing motion. above the foot the stand supports a horizontal plate the stage on which the object to be examined is placed. the cartilage bones yellow. This process of focussing is effected in two ways this. the which bears at its ends the lenses by which the magnifying power is obtained. An objective magnifying only a small number of times is called a one magnifying many times (200 diameters or more).

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. Should by any chance a drop of glycerine get on the face of the objective. Should Canada balsam be allowed to get on the objective. 3. but the clearness and definition of the object much increased. while focussing with the right hand. use a small diaphragm the amount of light will be somewhat diminished. 7. and then focus with the fine adjustment. . remove it. Focus accurately by means of the fine adjustment screw.THE USE OF THE MICROSCOPE. See that the body of the microscope slides smoothly in its If it does not. you move the object about slightly with the left hand. look down the microscope. When using a high power begin with the objective close to the cover-glass. and wipe it gently with a silk handkerchief or piece of chamois leather. and it will save : you much fatigue. Having adjusted the eyepiece and objective. and then place the object on the stage. : 6. It 1. direct the hght up the tube of the microscope by means of the mirror. and never to touch or allow any dirt to get on the face of the objective. and clean it by rubbing with a few drops of olive oil wipe off the oil before replacing the body in the tube. 2. By turning the head from right to left. and gradually twist the body up until the object becomes visible. do not attempt to clean it yourself. Twist down the body until the objective is about a quarter of an inch from the cover-glass. With a high power. will facilitate the process if. tube. (2) 5 close to the object that a is With high powers the objective has to be brought so more dehcate method of adjustment necessary. Take extreme care never to let the objective touch the cover-glass. a very Keep both eyes open when looking through the microscope little practice will enable you to do this. In using the microscope attend to the following rules : : Always examine an object first with the low power. but hand it at once to the assistant. Also get into the habit of using either eye. 5. wash it carefully with water from a washbottle. This ^flne adjust7nent is effected by a screw with a milled head placed at the top of the vertical pillar forming the stand. in the direction of the hands of a watch. The face of an objective cannot be cleaned without doing harm to it. 4.

A. 9. and drive out any airbubbles there may be in the fluid. 8. If the object appears dim or dirty. and above all things be careful not to let any of it get on the top of the cover-glass . Methods of Mounting. the top. it must be cleaned very carefully with a j)iece of silk or chamois : way While looking down the : leather. should this happen.. are the following. turn round the eyepiece with your right hand If the dirt turns round too. and arrange it with needles in Then place the cover glass carefully on the position desired. keej) one hand on the fine adjnstment. and keep screwing it up and down shghtly the whole time: in this way 2^'^^ts of the object at different depths are brought into focus successively. Slides should be labelled as soon as they are prepared. The method of procedure is much the same with all. remove the slide and clean it. VI.-THE PREPARATION OF MICROSCOPICAL OBJECTS. When examining an object. Put a. as they will jjrobably work out by themselves. If the fault is not in the eyepiece. The most important mounting media . There are various media in which objects may be mounted. if dirty. 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. if the dirt moves with the slide. small drop of the fluid in the middle of the slide. leave them alone.b THE PREPARATION OF MICROSCOPICAL OBJECTS. remove and clean the eyepiece. Be careful not to use too large a drop of yourmounting medium. place the object in the middle of the drojD. If the dirt does not move with either the eyepiece or the slide the fault is almost certainly in the objective. and a clearer idea of the object is obtained. the cover-glass must be removed at once and the specimen : mounted afresh with a clean one. If any air-bubbles still remain. and should be kept in a box or cabinet in w^hich they lie flat. find out where the fault lies in this microscope. which should be removed and examined. In mounting microscopical objects be careful that your slides and coverslips are thoroughly clean. move the slide about gently.

. and then torn up into shreds by means of a couple of needles held one ni each hand. before mounting. while preserving the individual cells.e. 2. The most important macerating fluids are as follows.THE PREPARATION OF MICROSCOPICAL OBJECTS.. if too thick. in 1. Canada Balsam is the most generally useful. a narrow ring of cement must be painted round the . i. C. requiring no cement. _ Maceration. which renders the particles easier to see.e. The process of teasing is in many cases facilitated by previously macerating the specimen. Take a very small fragment to commence with. may be diluted with chloroform.edge of the cover-glass to fix it to the slide. Normal Salt ^ mon salt in water. i. Canada balsam. it has practically no action on them. 2. be soaked for a few minutes in oil of cloves or turpentine in order to clear them. Specimens that are to be mounted in balsam must iirst be deprived of all water they may contain by placing for an hour or so in absolute alcohol. for making permanent preparations. soaking it in some fluid. and should then. Glycerine can be used either pure or diluted with its own bulk of water. 7 Solution: a 0-75 per cent. Tease it as finely as you can. It cannot be used. The process is often greatly facilitated by placing the slide on a piece of black paper. render them permeable by the balsam. unlike water. object of teasing is to separate the several parts of a tissue or organ from one another in order to show their minute structure. Teasing. as. however. If the preparations are intended to be permanent. 1. turpentine. a The two rules to be borne mind in teasing are the following. Your object is to separate the component parts from one another. _ The The fragment to be teased should be placed on a slide in a drop of the medium in which it is to be mounted. When torn cover-glass is placed on as before. up as finely as possible. 3. or benzole. B. which. solution of comThis is very useful in the examination of fresh specimens of animal tissues. tends to loosen them from one another..

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

dehydrated. alcohol. Such preparations are rarely permanent. A 1 per cent. or else washed. which binds together the several cells of a tissue. acid alcohol will vary. 7. according to the nature and size of the specimen. then washed thoroughly with distilled water. Picro-carmine answers best with specimens preserved in 70 per They should be left in the stain for a day. and to a certain extent a differential stain. Osmic Acid. and is therefore chiefly used when we wish to render prominent the The specimens should be placed outlines of the individual cells. Add some 5 grammes of picric acid.THE PREPARATION OF MICROSCOPICAL OBJECTS. and afterwards in 90 per cent. Magenta is stains very rapidly but diffusely : the colour also 5. only be left in it a few minutes. Some then placed in 70. solution of osmic acid in water forms an extremely useful staining reagent. when they may be mounted in glycerine. stirring it occasionally then evaporate it to dryness. and then to 90 per cent. Acetic Acid. much more strongly than the cells themselves. Silver Nitrate. should or black colour. : 4. h per cent. 3. acetic acid for an hour before transferring to alcohol. It is especially viseful for the detection of fat. and then decant from the excess of acid. and to every 2 grammes of the dried residue add 100 cc. of distilled water. specimens give better results when w^ashed freely with water on removal from the picro-carmine. as it colours the several tissues different Dissolve 1 gramme of carmine It may be prepared thus. of distilled water. continues until the specimens ultimately become too -dark to be of 6. A any use. which must be quite fresh. shake the mixture well for Leave the minutes. Although not strictly a staining agent . solution in water stains the intercellular substance. a very useful. alcohol. 9 transferred to 70. Picro-carmine is in 4 cc. and mounted in balsam. and may then be mounted in glycerine. and exposed to the light imtil stained sufficiently deeply. alcohol. not permanent. and cent. tints. fresh in the silver solution for from two minutes to a quarter of an hour. from a quarter of an hour up to a day or more. decanted liquid for some days. of liquid ammonia and 200 cc. in which The time of immersion in latter they may be left till required. and then placed in 1 per cent. to which the staining is due. which is stained by it a dark brown Specimens. as the reduction of the silver.

in which they may be left till required. and then transferred to 70 per cent. it has also the merit of staining It can. throughout the w^hole thickness of the object to be hardened.e. be employed when the specimens are very small.. alcohol and thence transferred after a couple of days to^ 90 per cent. The hardening reagents in most common use are as follows. . 2. as it hardens the surface layers so rapidly that it is unable to penetrate beyond a very slight depth. i. only the tissues as well as hardening them. Alcohol. Corrosive Sublimate. but the more usual The general action of these plan is by means of reagents. 3.. however. alcohol before staining. causes the protoplasm of cells to swell uj) and become transparent.e. of which a parts of the cells especially distinct.. . For this purpose a 1 per cent. There Hardening. After removal it is thoroughly washed with water or weak alcohol. 1 per cent. Hardening. cent. Before the object can be cut into sections it is necessary to harden it .-ON SECTION CUTTING. Many tissues and cutting organs can only be studied satisfactorily by into thin sections. hardening reagents is to coagulate the protoplasm of the tissues . special prominence. solution in water is used it acts almost instantaneously. Specimens may be placed at once in 70 per 1. A saturated solution in water is employed. VII. Osmic Acid. before the tissues can undergo any alteration thoroughly. and Cuttings are three chief stages in succession. A few minutes immersion is usually sufficient. in which the object is placed for half an hour or more. solution is employed. and this method of investigation is of such importance as to require special notice. : . inasmuch as it does not colour the specimens. To ensure the latter result it is always advisable to use small pieces of the substance to be cut. Imbedding. which will be noticed them : A. and so allows no change to occur in the tissues . acetic acid may conveniently be mentioned here as it is used for the same purpose as the true stains. i. and brings the nuclei into It is used with fresh specimens. for the sake of rendering certain Acetic acid.10 ON SECTION CUTTING. this may be effected by freezing. and the objects to be attained are to effect this coagulation and quickly.

and then transferred to 50 per cent. alcohol. if not too large. If the specimen is too large to stain whole. : : C. may now be stained with either hsematoxylin. It is then transferred to turpentine. it acts much more slowly 'than osmic acid.ON SECTION CUTTING. of water make a cold saturated solution of picric acid add 2 cc. except alcohol. Specimens that have been hardened in any of the preceding reagents. solution of chromic acid in water is a useful hardening reagent. the sections must be stained after they are cut. or picro-carmine they should then be replaced in 90 per cent. is A Mixture of often very useful. which is used in the following manner The stained S23ecimen is placed in absolute alcohol for an hour or two in order to completely dehydrate it. of which the best preparation is Kleinenberg's. 11 4. which should be changed daily until the specimens are free from the hardening reagent they may then be left in 90 per cent. A 0-25 to 0-5 per cent. of concentrated sulphuric acid filter. D. alcohol. on removal. 5. but penetrates to greater depths. In . as reagents. Dehydration. Staining. It is prepared thus with 100 cc. chromic acid with a few drops of osmic acid it combines the advantages of both 6. Imbedding^. but by far the most useful is paraffin. Specimens should be left in it from 12 to 24 hours. alcohol. alcohol on the following day they should be transferred to 70 per cent. borax-carmine. be placed for a few hours in 30 per cent. The preparation of sections is greatly facilitated by imbedding the specimen in some waxy substance. For this purpose various materials have been employed. Chromic Acid. alcohol until required. The hardened specimens. From the turpentine it is transferred to melted paraffin. Picric Acid is a very valuable hardening reagent. . which is kept by means of a water bath at a temperature just above its melting j^oint. and add to the filtrate three times its volume of water. Specimens should usually be left in the solution for one or more days. : : : B. in which it is left for half an hour or more until completely saturated. should.

The melted paraffin should then be washed off by turpentine. Instead of the mixture of collodion and oil of cloves. so as to form a ribbon. w^hen the sections will remain fixed to the slide by the collodion. When thoroughly set the block is removed from the box. By means of hot needles it can readily be arranged in any desired position . .12 ON SECTION CUTTING. but cut away from the sides at right angles to it. and to the edge of the razor. E. devised by Mr. just before they are used. and may be mounted in balsam in the usual manner. in order that it may be thoroughly permeated. These : — : spread over the slide in a thin layer by means of a glass rod. This depends on the fact that if the paraffin is of proper consistency the successive sections. will stick together at their edges. Section-Cutting. or other material. with a thin layer of a mixture of collodion and oil of cloves in equal parts. but a great saving of time is effected by the method of cutting continuous ribbons. as they are cut. or even for a whole day. a solution this should be of shellac in absolute alcohol may be used is The block sections. and the paraffin should then be cooled quickly. 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 paraffin cut If for parallel to one another. and allowed to dry. be coated with a layer of soft paraffin. so as to melt the paraffin and evaporate the oil of cloves. the block should before cutting. and the paraffin pared away with a knife until the specimen just comes into view. The slide is then heated by a water-bath to a temperature not exceeding 55° 60° C. Caldwell. Immediately before being used the slide should be brushed over with oil of cloves. left on the sides of the block parallel to the edge of the razor. any reason it is desirable to imbed the specimens in a paraffin too hard to form ribbons. and cut into thin may be transferred one by one to a slide. this it is left for several hours. placed on slides painted. It is then placed in a small box of paper. then placed in a microtome. by dipping it for a moment This outer coating should be in a dish of melted soft paraffin. filled with melted paraffin. when cut. The razor should be used dry and the sections.

The ventral surface. note. 1. tail. or belly. and hmbs . Fig. and the absence of neck and 2. or back. and make drawings^ The division into head.) : A. External Characters. trunk. The Common Frog {Rana temporaria) (from Ecker. is directed upwards when the frog is in the natural position.CHAPTER I. . GENERAL ANATOMY OP THE FROG. La^ the frog on a hoard before you showing the follovmig points 1. . a. The dorsal surface. The two great surfaces. h. is directed downwards towards the ground.

Each eye has two eyelids. There are tw^o pairs of limbs. The Mouth is a wide horizontal i. 5. 4. specially developed at the breeding season. The head is flat and triangular. fore and hind : each limb being composed of three segments. man. or openings on the surface of the body. a. and divided into the following parts : Thigh. just behind the bony projection formed by the posterior end of the urostyle. iii. GENERAL ANATOMY OP THE FROG. Hand. The Cloacal aperture is a small ii. The shortest Foot. The Hind-limb i. External apertures a. hole at the posterior : end of the body. The Fore-limb presents the following i. pigmented. Leg.14 3. and the longest to his fourth toe. of which the upper is thick. and claws. ii. . is much longer than the fore limb. The skin is moist and smooth and devoid of liairs. . In the male frog there is a thickening along the inner edge of the first digit. in the middle of which is a circular supported by a firm marginal area the tympanic membrane — — ring. ii. with a blunt apex directed forwards. while the lower is semi-transparent and freely moveable. toe corresponds to the big toe of man. The colour of the skin is variable in different specimens and at different times it is mottled on the dorsal surface. At the sides of the head are the eyes. slit. : 6. Arm. paler : on the ventral. with four fingers of corresponding to the four small and inconspicuous. Median apertures. Behind the eye on either side is an obliquely placed elongated patch of a dark colour. between the legs it lies slightly on the dorsal surface. The limbs. iii. digits. and almost immoveable. the thumb being very 1). which are large and prominent. divisions Forearm. with five toes webbed together. scales.

c. teeth in the fore part of the roof of the and near the middle line. of ivhich the hinder 'part or pharynx is continued hack into Note also the following structures the (esophagus. at the sides of the posterior part of the buccal cavity. Press down one of the eyes with your finger. The Teeth. 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. Two rounded jjrominences at the sides of the roof of the mouth are caused by the eyeballs. attached round the edge of the upper jaw. On a. i. 15 Paired apertures. the Eustachian tube into the mouth. b. Open the mouth to its : 1. the Roof of the Mouth.THE BUCCAL CAVITY. Perforate the tympanic membrane on one side ivith a needle. its The Buccal Cavity. The maxillary teeth are a row of fine teeth. and pass a bristle or seeker through the hole and — — down d. forms a bony margin to the floor of the mouth the rest of the floor is soft and fleshy. and note that it can be made to project very considerably into the buccal cavity. i. Pass bristles through the nostrils. The Eustachian tuhes or recesses are a pair of much larger holes. mouth h. but is slightly stiffened by a cartilaginous plate the body of the hyoid. which — . : The lower jaw. full extent : note the wide buccal or mouth cavity. The posterior nares are two small holes Ij^ing to the outer sides of and slightly in front of the two patches of vomerine teeth. The Nostrils or anterior nares are two small openings on the dorsal surface of the head. is devoid of teeth. and see that they come out through the posterior nares into the buccal cavity. close to anterior end. % On a. the Floor of the Mouth. B. The vomerine teeth are two small patches of sharp ii.

is attached to the front part of the floor of the mouth. Lay the frog on its hack and fasten it doimi to the dissecting hoard hy pins through the limhs. is a longitudinal sht in the floor of the posterior part of the mouth stiffened laterally by the arytenoid cartilages. GENERAL ANATOMY OF THE FROG. and the inner surface of the larger flap the anterior abdominal Pull the two flaps carefully dissect this from the flap. Separate the shin from the underlying parts. to the hinder end of the hody. inflate the bladder lungs with a blow-pipe through the glottis. The glottis. opj)Osite the fore-limbs. and through the cloacal aj^erture. The Abdominal Viscera. the ivhole length of the ventral surface. Pass bristles through the glottis into the lungs. The tongue. heing careful not to injure the anterior abdominal vein which runs along the inner surface of the hody wall in the middle line.16 b. which is thin and fleshy. Cut through the skin. and turn them hack so as to display the viscera. Note and draw the general arrangement of the viscera. cutting through them transversely at their posterior ends to facilitate the process. a?id taking care 7iot to injure the parts heneath. the tongue forwards with the forceps. under ivater. b. noticing its very loose attachment to these parts. and forwards to the jaw. and cut through hody cavity with scissors a little to one side of the median plane. or aperture of the larynx. cutting through the pectoral girdle ivith strong it Pinch up with forceps into the scissors. the muscular hody wall. or shoulder girdle : a bony arch running across the body. C. Continue the cut backwards. showing the following structures Inflate the . Notice — — — a. If any difficulty is experienced in finding the glottis snip through the angles of the mouth ivith scissors. so as to allow the mouth to he opened more widely. a^mrt. Note on vein. and the large space lymph cavity heneath it. The muscles The pectoral of the body w^all. Turn the flaps of skin outwards and pin them hack. along the middle line. and has its free bilobed end turned backwards towards the throats Turn C.

The ovaries two large bodies of irregular shape. and in the natural condition of the parts is covered by the pectoral girdle and the sternum. fold of skin at groin u. is situated in the middle line in front. lymph space between the skin and the muscular body wall 11.ABDOMINAL VISCERA. The Madder is a thin-walled bilobed sac at the posterior end of the body cavity. and usually come to the surface just behind the liver. dorsal a. The lungs are two thin-walled heart : elastic sacs at the sides of the they lie dorsal to the liver. oviduct p. behind and at the sides of the heart. The heart. kidney ov. 2. : A : : : : : : : : : : : 7. is a large reddish-brown bilobed organ. 6. they vary much in size. ureter v. lyi the female frog : note. or fat bodies. in the middle line behind is the much wider large intestine. 2. a. The corpora adiposa. spinal nerves o. 4. Fig. enclosed in the pericardium. are two bright yellow tufts of flattened processes attached to the dorsal wall of the body cavity . muscles of body wall aorta i. each consisting of a mass of spherical black and white eggs. urostyle d. posterior vena cava. skin : t. diagrammatic transverse section across the posterior part of the body of a female frog. C . and are often hidden by it Note the bristles already passed into the lungs through the glottis. large intestine h. The liver.a. in addition to the above jmrts. 3. 17 1. d. 5 The small intestine is a light coloured convoluted tube. peritoneum s. like small shot. ilium I.

E. bladder /i. a. spinal cord <. testis n. from the right side. ventricle urostyle cerelDellum y.s. 3. v. Fis. tongue pelvic symphysis . large intestine larynx : I. fat A-. small intestine d. Trace this to the mid-dorsal line where it is reflected downwards as a double layer the mesentery which embraces at its edge the alimentary canal. liver : : /. r. pancreas sp. attached to the dorsal wall of the body cavity. The Peritoneum. The Digestive Organs. Notice the thin pigmented membrane the peritoneum which lines the body cavity. lung body m. vesicula seminalis v. The testes: a pair bodies of a pale yellow colour. a. Eustachian recess z.s. In the male frog note. nasal sac. cloacal aperture : : : : : e. lying at the sides of the body 8.) binds its several coils together. and — — — (See Fig. bladder c. abdominal viscera are really outside Notice also that all the the peritoneum. long. bile duct : : g. cerebral hemisphere : General view of the viscera of the male frog.. stomach h. very much convoluted tubes with thick white walls. GENERAL ANATOMY OF THE FROG. gall : : : : : : : : u. kidney p. : 7/r. spleen ?. The oviducts: two cavity.18 "b. auricle lobe : a:. ureter o. of ovoid D. : : : ?/. optic : : . which forms a closed sac into which the viscera are as it were pushed from without. 2. d.

lobe. b. The stomach half in length a wide tubular sac about an inch and a it is narrower behind. and. and separated from the duodenum by a distinct pyloric constriction. remove the ovaries completely. e. is : C^it ojoen the stomach longitudinally along : its left side. wash out serted 771UC0US its its coiitents note the . f. divided into right and left lobes. Of the two lobes the left one is much the larger. The Liver. rather more than an inch in length beyond the pylorus it is bent back so as to lie parallel to the stomach. connected together by a narrow bridge of livertissue. The large intestine is a short straight tube about an inch and a quarter long it is very much wider than : the small intestine. damage the alimentary canal.DIGESTIVE ORGANS. Pass the and note the stomach lying beneath its handle of a seeker through the mouth and and oviducts down the oesophagus into the stomach. The small intestine is a slender convoluted tube about four and a half inches long. and is again subdivided into two. The cloaca in the frog is the last half inch of the large : which open the renal and genital ducts as well as the bladder it will be described more fully when considering the urinary and reproductive intestine. c. h mdle of the the seeker already in- through the mouth also longitudinal folds of the membrane lining the stomach. opening at its distal end by a small orifice into the large intestine. At its : further end d. taking care not to 1. into organs (see chapter 2. and opens behind to the exterior at the cloacal aperture. . The duodenum is the first part of the intestine. [If the specimen be a female. a.). The oesophagus is a short wide tube leading from the buccal cavity to the stomach. it is continuous with the small intestine. ivhich increase the extent of surface. viii. The liver is a large reddish-brown organ.] The Alimentary Canal. 19 Turn left the liver forwards.

p. "b. the strong wavy transverse folds of the onucous memhrane of the a hristle throicgh the duodenum. close to the middle line. Other Abdominal Viscera. and dissect with a scalpel cdong the line thus indicated. They lie in the large lymph space behind the peritoneum. and insert Notice also opening into the duct. 2. 1. The Pancreas.) . 17. The gall-Wadder is a small spherical sac lying between the riu'ht and left lobes of the liver. (See Fig. which passes through the pancreas to reach the duodenum. and spread it out on your dissecting hoard : measure the lengths of the several portions and draw them to scale. To trace the hile duct turn the liver forwards so that the point attachment of the gall-hladder is clearly see7i . The pancreas is a whitish 3. and best seen by The pancreatic ducts are turning the whole loop forwards.20 a. it has rather thick white the upper half is more more difficult to trace. and on the inner or concave side of the loop formed by the duodenum and stomach. and slightly stretch of Determine the the duodemcin hy a pin passed through the pylorus. The Kidneys are two flat elongated oval bodies of a red colour attached to the dorsal body wall. into which it opens about half an inch beyond the pylorus. numerous and open into the bile duct. slit up the first three quarters of an inch of the duodenum along its convex horder and wash out its contents : squeeze the gall-hladder so as to drive the hile along the duct into the duodenum : note the point at which it enters. F. leaving it attached at hoth ends. The distal half of the bile duct traverses the pancreas Avails and is easy to see slender and : . irregularly lobed mass lying in the loop between the stomach and duodenum. To see the opening of the hile duct. The bile duct is a slender tube leading from the liver and gall-bladder to the duodenum. position of the two ends of the hile duct from the description given above. GENERAL ANATOMY OF THE FROG. Cut through the mesentery along its attachment to the intestine : uncoil the intestine. one on each side of the back-bone or vertebral coliunn.

The Spleen is a small round dark-red body lying in the mesentery. opposite the commencement of the large intestine. and running back to open into the dorsal wall cloaca. or ducts of the kidneys. the vesicula seminalis. close to its opening into the cloaca. a.ABDOMINAL VISCERA. oi3posite the of the opening of the bladder. . 21 The ureters. are a pair of white tubes arising from the outer edges of the kidneys at about a quarter of their length from their hinder ends. In the male frog a pouch-like dilatation. 3. is present on the outer side of each ureter.

with the apex pointing backwards. i. hack wider water and open the body cavity as he/ore. and running obliquely forwards across the auricles. which by its contractions is continually driving the blood round and round the system of vessels (2) the arteries. iii. heai't in situ. ii. tahinr/ special care to preserve the anterior abdomiDissect this vein carefidly from the body wall. The auricles form the anterior and dorsal division of the heart they are thin. The truncns arteriosus is a cylindrical body arising from the right anterior border of the ventricle. the frog on its A. In freeing the pectoral girdle from the underlying muscles take care not to injure the neighbouring bloodvessels. 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. The Heart. dissect the pericardium the roots of the great vessels. from the heart and examine and draw the The divisions of the heart. On close examination the division into right and left auricles can be seen. Avhich are the vessels taking the blood from the heart to all parts of the body (3) the veins. Oi^en the pericardial cavity and.walled and appear dark in colour owing to the blood being seen : through their walls. a system of very small vessels connecting the arteries and veins together. owing to the greater thickness of its and is conical in shape. The ventricle is in colour posterior to the auricles : it is paler walls . . which carry the blood from those parts back to the heart and (4) the capillaries. 1.22 CHAPTER IL THE VASCULAR SYSTEM OF THE FROG. showing its several divisions. is : The vascular system : : : Pin down nal veiyi.

THE VEINS. or even after the heart is completely removed from the body. The sinus venosus to is a thin-walled sac. but not simultaneously. cardiac vein d. a. : b. f[S 4- Diagrammatic figure of the venous system of the frog.a. Lift up the ventricle sinus venosus. brachial vein :c. ngnt . 2. external jugular vein f.v.v.v. The pulsation of the heart.v. bladder: ?xv. innominate vein j. sinus venosus s.v. and finally the truncus arteriosus. C. ^ 23 so as to expose the and turn it forwards iv. large intestine e. vesical veins. from the a. Note the character of the heart's pulsations a regularly alternating series of contractions and dilatations. internal jugular vein Lp.r. : : : • • : : • : : : • : : : : : : : : : : : . right auricle v. Note that the contractions of the heart continue some time after the frog has been killed. gall bladder t. tongue t. stomach: a. ventricle v. femoral vein h.v. cloaca! aperture c. spleen g. it receives the three large venae cavae.p.v. truncus arteriosus u. The Veins.c. hepatic portal vein r. anterior alxlominal vein h. side. posterior vena cava i. then the two auricles. B.v.l. subclavian vein t. sciatic vein s.v. Note further that in each contraction or systole of the heart all four divisions of the heart contract.r. right pelvic vein r. musculo-cutaneous vein: o. lying dorsal the ventricle and behind the auricles . Lett pelvic vein m. liver e. kidney: ».c. then the ventricle. right renal portal vein -s. The sinus venosus contracts first.

I. ii. The mandibular vein. the thyroid gland. because. is formed by 3. returning blood from the interior of the skull. dissected before the arteries. from the fore-limb. at the posterior border of The subscapular vein. as nearer the surface and are therefore met with first. thinner walls. The lingual vein. a. it . In cleaning a vein take hold tcith the forceps^ not of the vein itself hut and of the tissue take especial care not to prick the vein^ as hy to surrounding doing so you also render allow the blood dissect escape and obscure the dissection^ to the loss and the vein itself difficult to see owing of colour. and returning to it the blood from the right side of the head and body. 1. The veins should be lie a rule. The external jugular vein is formed by i. The right anterior vena cava is a large vein opening into the right side of the sinus venosus. from the floor of the mouth and the tongue. Tlic i. subclavian vein. In close connexion with the ventral surface of the external jugular vein is a small round vascular body. they Dissect from the ventral surface. and pin out the as to stretch it slightly. Veins opening into the Sinus Venosus. Always 2^cirts so along and not across a bloodvessel. 2. a small vein from the back of the arm and shoulder. The innominate vein is formed by i. and darker colour. ii. The musculo-cutaneous vein: a very vein large returning blood from the skin and . the largest of the three. It is formed by the union of three veins. The brachial vein. due to the blood being seen through their walls. and from the right fore-limb. which it leaves by an aperture the orbit. The internal jugular vein.24 THE VASCULAR SYSTEM OF THE FROG. ii. from the margin of the lower jaw. The veins are further distinguished from the arteries by their larger size.

veins (in the male) II. commencing between the kidneys. from the liver these open into the jDOsterior vena cava just before it joins the sinus venosus. or spermatic returning blood from the ovaries or testes. (in the . It returns to the heart the blood from the liver and from the kidneys. III. It receives the following veins : i. Each pulmonary vein runs along the inner side of its lung. is one which. The Portal Systems. The left anterior vena cava corresponds and branches to the right one. "b. The posterior vena cava is a median vein which. or rather form by their union. The pulmonary vein is formed by the union of the right and left pulmonary veins. . and of the head as far forwards as the nose. portal vein laries of A some part. iii.THE VEINS. one supplying the kidneys. in its course c. and the other the liver. The right and left hepatic veins. 25 muscles of the side and back of the body. The ovarian veins female). The renal veins. and open into the posterior vena cava between the renal veins. In the frog there are two portal systems. They are usually four or five in number on each side. and indirectly from other viscera and from the hind limbs. the posterior vena The most anterior of these receive the cava. a. breaks . returning to the heart the blood from the right and left lungs respectively. veins from the fat bodies. from the kidneys of these there are four or five on each side which open into. runs forward to open into the posterior end of the sinus venosus. returning blood from the capilup 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. : ii. Vein opening into the Left Auricle.

The right ' - • sciatic vein. Vesical veins. .26 a. i. from the muscles and skin of the back of the thigh. This is formed partly by the anterior abdominal vein. THE VASCULAR SYSTEM OF THE FROG. receives the following veins. which enter the right and left lobes of During its course it the liver respectively. where it leaves the body-wall and divides into right and left branches. 1. in the they join the female. 1. before it reaches the kidney. It receives the following branches. of the ventral forwards along the middle line body-wall to the level of the liver. veins returning blood from the alimentary canal. where it ivill he seen to he formed. The renal portal system. divides. and. ' ii. from the oviduct renal portal vein opposite the kidney. . in its course The hepatic portal system. of the hind Hmh . Trace hack the anterior ahdominal vein to the hinder end of the body. The anterior abdominal vein is a median vein formed by the union of the two pelvic veins. The left renal portal vein corresponds and branches to the right vein. hy the union of the tivo Follow hack the pelvic vein of one side to the base pelvic veins. the large vein returning blood from the hind limb. ii. from the ventral body-wall. The other branch of the femoral vein is the renal portal vein. i. 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. the It runs ventral branches of the femoral veins. which is to be followed to the outer side of the kidney. The right dorso-lumbar veins : are small veins from the dorsal wall of the body. from the bladder Parietal veins. 2. here it ivill he seen to he one of two branches into which the femoral vein. which brings to the liver blood from the hind limbs and ]3artly by b. joins the renal portal vein close to its commencement.

' : : . It carries to the liver the blood from the walls of the alimentary canal. dorsal aorta: /. pelvic girdle 6. : : . pulmo-cutaneous 2. and is formed by the union of the following veins.^' small intestine c. its minal vein at and left branches a branch to the .n.'^ nnntJf* occijjito-vertehral "ostri^. before doing so.a. tongue: «f. from the stomach. p.s. sciatic artery: f. 11 Intestinal veins.r. o"' "rinogenital arteries v. f r. from the whole length of the intestine.a. from a network of vessels on the truncus arteriosus. The hepatic portal vein is a wide vein which runs in the mesentery and joins the anterior abdopoint of division into right giving off. kidney : o.THE ARTERIES. fromth^rSfSr''*'" nn»2>'f carotid T'"''!' gland -'' • : ^^"""^ ""^ ^^'^ '^'*''^'^^ '^'*'"' ^^ the male frog. ventricle: 1.i. testis: : : '' o. i. cutaneous artery: iptestine: d. femur: h.«. 27 A cardiac vein. iii.a. 111 The splenic vein. both small and large. artery . hepatic «vw? "?^^* !""§= ^'^ l"ig"al artery: m. C.jn coeliaco-mesenteric artery: c. sternum a.a. carotid arterv c. systemic arch arch. spleen: h. The gastric vein. pulmonary artery r. The Arteries. from the spleen : this usually joins one of the intestinal veins. subclavian artery: . carotid arch : o''''"^'''^^-'''°^"f 3.a. left lobe of the liver.

The carotid artery runs round the o'us to its side of the cesophaconnected with the systemic arch by a short branch. The external carotid artery. and the pulmo. The Carotid Arch is the most anterior of the three arches rims round tlie side of the a3sophagus. and then turns forwards beneath the base of the skull. are as follows a. The lingual artery is a small artery supplying the tongue. . The internal carotid artery. Branches given ofi' before the union of the two arches. the middle arch of the three. 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 The branches of the systemic arch into the two iliac arteries. 1.cutaneous arch. the ductus Botalli. and the orbit. 2. the carotid gland. The Systemic Arch. THE VASCULAR SYSTEM OF THE FROG. which in the adult frog is usually impervious . which enters the skull ii. the arches. supplying the roof and sides of the biiccal cavity. as so for as the to veins. each of which again divides into three aortic arches the carotid arch. Clean carefidly arteriosus. The laryngeal artery is a small branch arising from the inner side of the systemic arch near its origin from the truncus arteriosus. dorsal surface : it is : II. Pass a it roll stretch of paper down the the distend aortic and aortic the arches. and supplying the larynx. Immediately beyond the origin of the lingual artery the carotid arch presents a small spongy swelling. The oesophageal from the upi)er arteries are one or two branches arising ])art of the arch and qntering the dorsal wall of the oesophagus. commencing at truncus and follow the several arteries to their distributionj removing the veins arid other structures which overlie them. runs somewhat oblicpiely round the a3sophagus to the dorsal surface. : ' 2.28 Dissect (esophagus. and is connected dorsally with the second or systemic arch its chief branches are as : : follows. dividing in front into the two following vessels i. and supplies the brain. — it I. 1. Note the division of the truncus arteriosus in front into right and left branches. the systemic arch.

and supplying the stomach and intestines. The hepatic gall-bladder. 29 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. 1. or sometimes from the left arch just before the union. the distal part of the intestine. The vertebral artery a large artery w^hich runs back alongside of and above the vertebral column. and divides into two : : i. The occipital artery which runs forwards. /3. the reproductive organs and ducts. The cceliaco-mesenteric artery arising immediately is a large median artery beyond the point of union of the two arches. supplying 2. 3. supplying the stomach. i. and gives branches to the muscles of the bodywall and to the spinal cord. supplying the kidneys. The subclavian artery: ately behind the occipito-vertebral artery. supplying the spleen. splenic artery. Its branches are as follows : : The a. The lumbar arteries are small paired lateral branches supplying the body-walls. and immediately divide into right and left branches. supplying the proximal part of the intestine. artery. The 3.THE ARTERIES. The posterior mesenteric artery. : ii. The anterior mesenteric artery. suppl}-ing the side of the head and jaws. arises b. supplying the shoidder and fore-limb. Branches given off after the union of the two arches to form the dorsal aorta. y. supplying the liver and ii The mesenteric artery: which divides into a. /3. . from the arch immediand runs outwards. The nrinogenital arteries are four to six small arteries which arise from the ventral surface of the aorta between the kidneys. and the fat bodies. : 4. cceliac artery which divides into The gastric artery.

course along the outer side of the whole length of the lung. and left tyiterior vena? cavse open.30 4. giving epigastric branches to the ventrtil body-wall. Cut away with scissors the : to expose its cavity ivash out dorsal ivall of the sinus venosus so as any contained blood. The pulmonary artery runs with somewhat sinuous The Structure of the Heart. and supplying the hind-limbs. III. giving off branches into its substance. a small median artery and supplying the large intestine. THE VASCULAR SYSTEM OF THE FROG. The Pulmo-cutaneous Arch : aortic arches it 1. it is triangular in shape. c. p. which supplies the bladder. is the hindmost of the three divides about the level of the carotid gland into the following branches. supplying the skin of the back along the whole length and sending smaller branches to the sides of the head and to the skin of the ventral surface. the dorsal surface of the heart . D. The cutaneous artery is a large artery wdiich at first runs forwards and upwards and then turns backwards. vessels Place tJie heart at first with the dorsal surface upwards. A. remove the heart comIt is well to cut the pletely. and dividing at the knee into peroneal and tibial arteries supplying the leg and foot. cut them across. of nnequcd lengths on the two sides. . giving off branches to the muscles and skin of the thigh. and dissect it carefidly binder water. The Sinus Venosus (Fig. of the body. with Into its anterior angles the right the apex directed backwards. 23) is a thin-walled sac on 1. 2. as this loill facilitate the recognition of the sides of the heart during the dissection. Having completed the dissection of the blood-vessels. The haemorrhoidal artery arising is from the hinder end of the aorta. and then continues as the sciatic artery down the leg. 1. and into its posterior angle or apex the posterior vena cava. The iliac arteries are the two large arteries formed by the division of the aorta. Branches formed by the division of the aorta. about half an inch from the heart . Each gives off a hypogastric artery.

6. and very nearly in the median plane. The frog s heart seen from the ventral surface. The ventral walls of the truncus arteriosus and of the auricles and ventricle have heen removed. left carotid arch C. style passed down right carotid arch into the : : • : : : : : : right auricle : V. left auricle P. is a transversely oval opening. style. style passed down right systemic arch into the truncus arteriosus SV. aperture leading from ventricle to truncus arteriosus. guarded by imperfect anterior and posterior valves. 6. CSP Fig._ auriculo-ventricular aperture and one of its valves B. left systemic arch S'. rio-ht auricle S. Hurst. in the ventral wall of the sinus venosus. left pulmo-cutaneous arch P'P'.THE HEART. and dissected so as to show its structure.) * truncus arteriosus LA. opening from sinus venosus into : : A. . 31 The sinu-auricular aperture (Fig. ventricle. (From a drawin"- by Dr. close to its anterior end. passed down right pulmo-cutaneous arch into the truncus arteriosus PV.V. S. with one of its valves C. opening of pulmonary vein into left auricle RA.) leading from the sinous venosus to the right auricle.

than the right auricle. Wash out the blood from the auricles. very near surface. talcing care auricles with fine the truncus arteriosus ivhich lies across the right auricle. THE VASCULAR SYSTEM OF THE FROG. The spongy character is due to great development 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. The inter auricular septum is the thin partition between the right and left auricles. and are filled : with blood. The auriculo-ventricular aperture ventricle. sometimes much smaller. and the meshes of the spongework are really part of the cavity of the ventricle. 6. h.32 2. lies at the base of the and rather to the left side. The right auricle (Fig. 6 V) is conical in shape with the apex backwards. and has a small central cavity. is the opening of the pulmonary vein (Fig. which it resemIn its dorsal wall. The left auricle (Fig. 6 walls. with thick spongy walls. the left auricle lying rather more dorsally than the right. respectively. 6 LA) is smaller. It has thin the ventricle. admitting blood from the right and left auricles . C^it away carefully. opposite the auriculo-ventricular aperture. It is much thinner than the walls of the auricles. their edges by fine tendinous threads and it is divided by the free lower edge of the interauricular septum into right and left divisions. It is guarded by valves A) Avhich hang into the ventricle. Turn the heart over. and are tied down at (Fig. uptvards. taking care not to Cut aivay ivith scissors the ventral wall of damage the truncus arteriosiLs. 3. and is placed somewhat obliquely. to expose its cavity the ventral wall of the truncus arteriosus so as and the contained valves. bles in the structure of its walls. with Cut away the ventral ivall of both not to damar/e its ventral surface scissors. The Auricles. "b. The Ventricle (Fig. . tvith fine scissors. very close to the sinu-auricular aperture.. the median plane of the heart. 6 SY). c. 6 PV). is the aperture from the sinus venosus already described (Fig. RA) is the larger of the two. thickened by muscular strands which form interlacing reticular ridges on its inner In the dorsal wall of the auricle. The septum ends with a free posterior edge.

Pass bristles down these aortic arches. 33 4. very close together. a small left one. and note the points at which they severally open into the truncus arteriosus. where it fuses with the large right valve of the three between the pylangium and the synangium. Beyond this the synangium contains a wide cavity continued right and left into the two systemic arches S. across the aortic arches. The opening from the ventricle to the pylangiimi (Fig. which is a single vessel arising from the ventricle . is an aperture (Fig. and a distal part or synanginm. D . 6 P. 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. immediately beyond arteriosus. and note that though each Cut branch is apparently a single vessel its cavity is really divided into three vessels corresponding to the three aortic arches. C. a large right one.S'. The synangium is the distal part of the truncus : : In its dorsal wall. the valves separating it from the pylangium. which consists of the basal parts of the aortic arches closely united together. The opening from the pylangium to the synangium is also guarded by three semilunar valves which are of very unequal size. 6 P') leading to the right and left pulmo-cutaneous arches (Fig. and the valve is much wider at its anterior than at its posterior end. b. -. 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.C'. The ventral edge of the spiral valve is free and rounded. 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 Truncus Arteriosus consists of two parts a proximal part or pylangium.P'). 6 B) is guarded by three semilunar pocket valves. a. The pylangium (Fig. The spiral valve is a longitudinal ridge. and a still smaller dorsal valve. just heyond the division of the truncus into right and left branches.THE HEART. which lead into the right and left — : carotid arches.

The are the large along the dorsal surface of the bodyspaces cavity. "b. : The lymphatic system forms an accessory part of the vascular Its main divisions are as follows system. The lymphatic vessels are a series of thin. The body cavity itself also Fig. ventral to the kidneys. h. be seen in a living . have already been seen when removing which They are separated from one the skin. p. close to its hinder end. 1. another by narrow septa of connective tissue. frog. The suhcutaneous lymph sacs cavities are between the skin and the muscles. The posterior Ijrmph hearts lie at the sides of They the urostyle. E. a. and between {See the peritoneum and the body-walls. abdominal lymph sacs 3. 17. The lymph hearts tractile sacs are two pairs of small globular conplaced at points where the lymphatic vessels communicate with the veins. by short vessels wdth the communicate Their pulsations can easily femoral veins. The Lymphatic System.34 THE VASCULAR SYSTEM OF THE FROG. and can only be microscope.walled tubes.) commiuiicates with the lymphatic system through small openings or stomata in the peritoneum. The lymph sacs are large irregular spaces vessels. 2. recognised with the small size. very variable in diameter and irregular in shape. 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. cating with the lymphatic important are the following : communiThe rnost the large a. which bind the skin to the underlying bodywall. which traverse all the parts and organs of the body and are They are of in free communication with the veins. 2.

They seen when a corpuscle turns edgeways. ii. the liquor sanguinis or plasma. in which float the Wood corpuscles. These are much fewer in number and of smaller size they are colourless. red or yellowish red in colour. in . White corpuscles become clearer. and run a ring of oil round the edge to prevent evaporation : examine with the high power. and of a flattened oval shape. or about yyVo ^ TsVo" ^'^ ^^ inch. Action of acetic acid on blood. dilute Blood consists of a colourless fluid. Frog's Blood. Human Blood. and the red colour dis: appears. and exhibit " amoeboid width : movements. Red corpuscles the nuclei become much more apparent than before. pale corpuscles. Note the folloidng p)oints : Red corpuscles. tip of a. Normal. and examine as i. and examine with the high power : note the : changes produced. Prick the hlood on slide: your finger. ii. and show nuclei. 2. measure 0'0235 mm.BLOOD. granular. acetic acid i. F. the nucleus. with rounded edges and a central The flattened shape is best bulging. 1. sometimes more than one in a single corpuscle. Microscopic I. Normal. Place a fresh drop of blood on a clea7i slide : add a drop of cover. 1. These corpuscles are of two kinds. Sketch one half-a-dozen times at intervals of half a minute. 35 Examination of Blood. before. : II. Place on a slide a small drop of hlood from the heart of a frog it with a drop of normal salt solution (0"75 per cent) : put on a thin cover-glass. in length by 0*0 145 mm. cover. subspherical in shape. which are much smaller . These. White corpuscles. Red These are very numerous. i. and place a small drop of the add a drop of normal salt solution.

Action of acetic acid. ii. or their amoeboid ^^ ^^^ ^^^^ ^^ diameter "2 Wo" movements are not well seen unless the slide is : : warmed. of three kinds. it can readily web of Examine a frog prepared the foot. the direction of flow of the blood is from the larger trunk to its branches. 1. are in the form of circular with rounded edges. a. along which the blood flows from the arteries to the veins. is The web uniting the toes of the frog's foot so thin and transparent. unlike the frog's blood. : Treat tvith acetic acid as before note that. are distinguished from the arteries by the fact that the blood in them flows from smaller to larger vessels. These bloodvessels are The arteries. to show : the circulation in the Note the following points With a low power. iii. Circulation of the Blood in the Web of a Frog's Foot. .36 THE VASCULAR SYSTEM OF THE FROG. ii. or about -g-oVo ^^ ^^ inch. than in frog's blood. carrying the blood away from the web back towards the heart. but no nuclei. very thin-walled vessels.. "White corpuscles. The capillaries form a close network of very small. 2. The veins. carrying blood to the web. Their average diameter is 0*008 mm.. The irregularly branched pigment colour of the frog's skin b. no nuclei are visible in the red corpuscles. i. G. like piles of coins. biconcave discs They have a tendency to run together into rouleaux. averaging about 0"01 mm. is cells to which the due. are distinguished by the fact that when they divide. These are very similar to those they are slightly larger than the of the frog red corpuscles. that with the microscope the blood in be seen coursing along the capillaries. The fine meshwork of bloodvessels along wdiich the blood can be seen flowing.

This increased by the application of some irritant substance. : note the following points: The walls of the arteries and to see. 37 With a high power a. 2.CIRCULATION OF BLOOD. to the web. as a drop of weak acid. The seen best when turning the corners of the capillary network. may e. The tendency of the white corpuscles the walls of the to migrate through capillaries into the tissues outside. The white corpuscles have a marked tendency to creep along the sides of the vessels. : C. while the red corpuscles rush far more rapidly along the middle of the stream this is seen best in the small arteries. Owing to changes of size in adjacent vessels. The in calibre of the small arteries and whilst under observation an artery or capillary may be seen to change its size to a considerable extent. is much . which latter are often difficult b. variations : capillaries d. veins are much thicker than those of the capillaries. The indefinite character of the capillary circulation. they are elasticity of the red corpuscles : f. the direction of flow of the blood in a given capillary become reversed.

therefore. and structure in different tissues. A. and draw. or it may be so abundant as to separate them widely it is to be viewed as formed by the cells. the flattening is most marked in the superficial cells. and the At the external apertures various internal cavities of the body. The hlood corpuscles form most useful standards of measurement. vary much in shape. all the different tissues and organs of the body are found to consist of elementary bodies called cells and of an intercellular substance. which form the surface covering. 35). When examined under the microscope. Epithelium consists of cells placed side by side so as to form layers. Epithelium structure of I. The layers may be one or more cells in thickness . : . When drawing histological preparations. In this the component cells are flattened parallel to the surface they cover if the epithelium is stratified. of the body. . Squamous Epithelium. according to the shape and its component cells. : (p. or epidermis. ELEMENTARY HISTOLOGY. to the same scale as the rest of the drawing. it is well to look out for. and. of the body. in the latter stratified. as their dimensions are already known. a few red hlood corpuscles. it may be almost absent. substance varies very much in quantity. in the former case the epithelium is said to be simple.38 CHAPTER III. so that the several cells are practically in contact with one another . the epidermis is directly continuous with the epithelial lining of the internal cavities. is of different kinds. is a typical example. the blood vessels. The intercellular white blood corpuscle size. Epithelium. and line the alimentary canal. of which a of bricks cemented together with mortar. connecting the several cells together in much the same way as a wall is built These cells. but are to be considered as fundamentally equivalent to one another. as secondar}^ in importance to these.

narrow columwith nuclei packed together side by at their inner or deeper ends. and examine with a high power . showing the following points i. and then. and examine with the high power. Take a small piece of The cells are flattened. cover. a. the prepared sp>ecimen. cover. cells. Cells in situ cast skin of newt. ^ i. i. The superficial layer consists of long cells. 39 Isolated Cells. and examine with the high poiver. with nuclei near their inner or deeper ends. of dofs stomach which has been stained. Isolated cells isolated alcohol. section Take a prepared and then cleared in oil of cloves. . to form a continuous layer.EPITHELIUM. a. The cells. which has been stained in hematoxylin. Columnar Epithelium. are columnar in shape. : b. and examine with the high power. Cells in situ. ii. flattened and scale-like in shape. which often remain side by side in little groups. like a mosaic. "b. and fitted together at their edges. nar side. cleared ivith oil of cloves. i. from the small intestine of the frog in by maceration for 24 hours and stained with picro-carmine. often slightly curled up at their edges. The The cells are large. Scrape gently the inside of your cheek with the handle of a scalpel. Each cell has a large nucleus near its centre. If the epithelium is stratified the columnar character is most marked in the superficial cells. This consists of elongated rod-like placed vertically to the surface on which they rest. Mount in halsam. niicleus is oval and granular. . II. draiv. after treatment with alcohol. and put the scrapings on a slide. Kanvier's Mount a drop of prepared specimen in glycerine : paint a ring of cement round the cover-glass. and lies near the middle of the cell: it may be rendered more distinct by acetic acid or magenta. Mount the specimen in halsam .

and cleared in oil of cloves. ii. The movements of the individual cilia best seen when the specimen is beginning to die. bear at their free ends tufts of exceedingly fine hairlike processes cilia— which. often in a very complicated manner. a. when living. : Mount a small drop of the prepared specimen in glycerine: paint a ring of cement round the cover-glass. B. The shape of the cells : their nuclei cell. . Ciliated Epithelium. III. gland consists essentially of a layer of epithelial cells secreting some special fluid. This is characterised epithelium being several cells in thickness. which are usually columnar. ii. stained. and scraped into glycerine. From trachea of rabbit maceration for 24 hours in Ranvier's alcohol . The epithelial surface may be flat. of epithelium from the roof of the mouth of a freshly killed frog.40 ELEMENTARY HISTOLOGY. : IV. but is more usually folded or pitted. stained with picro-carmine. i. examine with the high power. Cells in situ: one end of each ciliary Sni}) off a small piece movement. or of conjunctiva of rabbit or pig. near the eyeball : mount in normal salt solution. A . examine with the high power. note — i. The transition from the deeper spherical or columnar cells to the superficial squamous cells. isolated by Isolated cells. and note : — The stratification of the epithelium. which has been hardened in chromic acid. exhibit active lashing niovements. so as to increase the extent of the secreting surface. and add a small drop of gamboge ivater to render the movements more clearly visible : examine with the high power . by the Tahe a prepared section of oesophagus of rabbit. and the tuft of cilia at b. and note — i. and the movements to slacken in speed. In this the cells. The currents due to the ciliary motion. Stratified Epithelium. Mount in balsam. Glands.

cavities are cut at various angles. 41 I. The gland cells. whose mouths serve to discharge the secretion on the free surface. •often in a very complicated manner. in which the several subdivisions are tubular. to which the special glandular epithelium is usually confined. in which the blind ends of the pits are dilated into globular chambers or alveoli. The glandular epithelium lining the pits is a single layer of short columnar granular cells. : : a. If cut transversely a tube appears as a circular ring : if cut obliquely. as a more or less elongated elliptical ring : if cut longitudinally. of frog : mount in balsarn. Compound tubular kidney/ glands. Take a prepared section of large intestine of rahhit which has been hardened in chromic acid. In simple glands the epithelial surface is increased by simple pit-like depressions. into which project little knots of capillary bloodvessels. The Malpighian bodies are spherical dilatations on the tubes. Their structure is most readily made in specimens in which the bloodvessels have out been injected with a coloured substance to make them more distinct. and cleared in oil of Mount in balsam. ii. The glands surface. stained. iii. The tubular gland. as two parallel rows of epithelial cells. Take a prepared section of and examine with both low and high i. cells form a single layer of cubical granular lining the tubes. Compound Glands.GLANDS. . which are swollen to form goblet cells. There are two chief varieties (I) tubular glands. and of tolerably uniform diameter throughout and (2) racemose glands. and examine first with the low power^ cloves. many of II. i. are simple tubular depressions of the ii. pioivers. Simple Glands. In compound glands each depression instead of being a simple pit is itself subdivided or branched. Note the following points then with the high.

Fig. longitudinal muscle fibres P. : membrane of the cardiac end of a dog's stomach. : : : : : : : . columnar cells CO. gland cavity cut cells GB. circular muscle fibres ML. connective tissue layer between the mucous membrane and the outer muscular walls of the stomach. Section through mucous x 140.42 ELEMENTARY HISTOLOGY. ovoid CM. mouth of gland MO. bloodvessel CC. cubic or peptic cells GC. fundus or bottom of gland cavity across: GM. B. 7.

and are said to secrete the acid of the gastric juice. There are three distinct kinds of epithelial cells found at different parts of the length of the gland. Gastric Glands. in the higher forms. They lie along the sides of each gland. and occur most abundantly a short way below the mouths of the glands. cells. i. loith a high power the section of the cardiac end ephitJielium. They are deep. Characters of the gland cells. The glands of the stomach are well adapted for a more minute examination of the histology of glands. owing to the glands being not quite straight. arranged in a somewhat radiate manner round the mouths of the glands. some of the glands may be seen cut along their entire length . imbedded vertically in the wall of the stomach. lining the deeper parts and the greater part of their length these are cubical granular cells with centrally placed nuclei. or the plane of section being oblique to the surface of the stomach. the dog's stomach already used for columnar Characters of the glands. very highly specialised.GLANDS OF STOMACH. : ii. the tubes will be cut more or less obliquely. of the glands cells large oval cells with large nuclei these are less numerous than the other two forms. Examine again of 1. or iii. Muscle. 2. cylindrical pits. but very narrow. In the microscopical sections. outside the cubical cells. Cubic peptic cells. but in most cases. or voluntary of which all : 3 . The gastric glands are good examples of simple or slightly branched tubular glands. The glands are lined by epithelial cells. or even transversely. In muscular tissue the component cells are much elongated and. and extending a short way down the tubes. with their open mouths discharging into its cavity. 43 III. Columnar cells. Ovoid : C. and are set very close together side by side. Muscular tissue is of two kinds (1) striated.

nucleated muscle-cells. and Mount in balsam. which chief exception to this rule. Each fibre is a single cell. I. or a. low and high powers : note — i. seen at places where the fibre has across. in normal salt solution high power : note Tease gently apiece of fresh frog^ s muscle cover. stai?ied. Tease in glycerine a small piece of craVs Crab's muscle. The transverse striations. been torn The alternate light and dark bands with which the muscle fibres are marked transversely.44 ELEMENTARY HISTOLOGY. The absence of transverse striation in the muscle. The sarcolemma: best seen by the specimen. fibres split iii. Frog's muscle. forms the of the heart. is derived. : slightly crushing iii. and is enclosed in a the sarcolemma which delicate elastic sheath wall be visible in but few cases it is most readily — — . 2~>encilled with a fine brush to remove the epithelium of the ijiner surface . : muscles that are under the control of the will consist and (2) non-striated or involuntary forming those muscles over whose The muscular tissue contractions the will has no direct control. ii. Voluntary Muscle. The elongated fibres of which the muscle consists. Take a prepared specimen of frog^s bladder II. ii. Non-striated. The readiness with which the tudinally into fibrils. up longi- b. elongated. ii. muscle that has been hardened in alcohol . and examine with cleared ivith oil of cloves. The formation of each band by a number of iii. striated muscle. Striated. and examine with the : — i. The bands of muscular fibre. fusiform. The nuclei in the fibres acid. or ivhich has been ^macerated in Ranvier^s alcohol for 2Jf hours . cover^ and examine with both low and high powers : note : — i. . though involuntary is striated. seen on addition of acetic Involuntary Muscle. and from which the name.

united together in very varying proportions in different situations (1) white fibrous tissue (2) : : yellow elastic tissue (3) connective tissue corpuscles. . 45 D. which are comparatively slightly altered cells. The fibres are arranged side by side in bundles. The fibrillse. White Fibrous Tissue. Tendon of rat's tail. and (4) ground substance. Histologically the connective tissues consist of elements of four kinds. usually branched . The fibres swell up and become transparent. Under the name " connective tissues tissue " are included various whose functions are mainly passive. and which serve to support. : a. note : — . with great power of resisting chemical reagents the fibres are formed from an intercellular matrix. and each fibre presents a number of longitudinal fibrillar striations. usually in small numbers. This consists of a number of fine transparent fibres of a more or less cylindrical shape. cover and examine ivith low and high powers . The fibres. and not from cells directly. The cellular origin of while fibrous tissue is difficult to determine. Yellow Elastic Tissue.. Pull from the tail of a rat : place out a small piece of tendon it on a slide in a drop of normal salt solution : spread it out with needles. by longitudinal wavy preparation : stria^ tions within the fibres. iv. Tease finely a small shred in water . Longitudinal become II. Ligamentum nuchaB of ox. indicated ii. This consists of fine branching homogeneous fibres. with wavy outlines. Connective Tissues. and with a very characteristic wavy outline between the fibres are connective tissue cells.MUSCLE : CONNECTIVE TISSUE. The fibres are believed to be formed by modification of the intercellular matrix rather than from the bodies of the cells themselves. note — i. : a. Add a drop of acetic acid to the note that iii. or intercellular substance. with nuclei. between the fibres. visible rows of tendon cells. examine with low and high powers . I. strengthen and bind together the various organs and parts of the body.

The fat cells large. composed of white fibrous tissue with wavy outlines. i. Connective tissue corpuscles. and with their nuclei near the surface. The branching fibres. with very sharp outlines. spherical. with nuclei. in which the fat cells. or from mutual pressure polyhedral. cells. The tendency of the branches to anastomose with one another and so form networks. : ii. The elastic tissue is unaltered. The white fibrous tissue swells up and becomes transparent. Add a iv. note: i. Mount a small piece of fresh omentum in 7iormal salt solution. of acetic add . rat. note that ii. The meshwork. III. The tendency droio of the fibres and branches to curl up at their broken ends. note that is V. examine with low and high powers . mingled with which are branched elastic fibres. iii. nuclei appear. This consists of a fine network of vascular connective tissue.46 i. Adipose tissue. Add acetic acid . The vascular connective lie tissue meshwork. This is a meshwork composed of both white fibrous and elastic tissues. and snip off a small piece of it Talce a fresldy hilled tissue the loose fibrous : which on a slide: add a drop of normal salt solution and examine ivith low and high powers . . connects the shin with the suhjace7it parts spread : — cover. distended with fatty matter. a.e. iv. ii. a. iii. Areolar tissue. Omentum of rabbit or kitten. in the meshes of which are fat cells.. IV. Subcutaneous tissue of mammal. note : — i. ELEMENTARY HISTOLOGY. No No alteration whatever produced in the fibres. connective tissue corpuscles in which large quantities of fatty or oily matter have accumulated. protect it from the pressure of the cover glass . become visible.

Cartilage of newt. Note the reduction of the osmic acid by the fat. The intercellular substance forms a dense translucent matrix resembling an extremely stiff jelly. Cartilage.CARTILAGE. 47 Osmic Acid specimen. Hyaline cartilage. mount in normal salt solution. and mount as a permanent preparation in glycerine . and occupies a cavity or lacuna in the matrix. Cartilage I. : — — . is either hyaline or faintly ii. and exami7ie with loiv and high the shoulder girdle i. and the matrix very slightly the layer of matrix immediately surrounding each cell the capsule stains more deeply than the other parts. in which are imbedded the cartilage cells. cartilage when free from other tissue is called hyaline from the clear or glassy appearance of the matrix. which becomes stained a dark brown or black colour. The cartilage cells are imbedded in the matrix. powers. either singly or in groups. and the cells consequently closer together than in most other older or more mature specimens. In places the cells are in groups of twos or fours owing to recent division. The cell nuclei are stained deeply. is greatly increased so as to far exceed in bulk the cells which it connects together. in contra-distinction to fibro-cartilage. The intercellular matrix granular. b. In young cartilage the intercellular substance is much less abundant. Take a small piece of cartilage from of a newt : scrape away gently any muscle or other tissue that may adhere to it . In cartilage or gristle the intercellular substance. E. in which the matrix is fibrous from admixture with white fibrous or elastic tissues. Wash the specimen thoroughly in water : stain with carmine. and note that iii. examine zvith the high p)ower. each cell is nucleated. a. which in tissues is only present in small quantity.

Towards the free surface the cells and cell groups become gradually flattened. iii.48 b. and the central canals. chiefly phosphate and carbonate of lime. lacuna) appear black. with its contained bone-cells. to the surface. and are readily recognised by the concentric arrangement of the lamellae. The cartilage cells. The matrix. . which penetrate the bone in bloodvessels. which form about two. the section being made perpendicular to the articular surface : examine ivith low and high powers. This forms caps covering the Articular cartilage. ELEMENTARY HISTOLOGY. and arranged paralled F. Bone. Examine with verse sections of i. 1. great numbers. up the spaces between They form parts of circles which are in many cases of much larger radius than the circles of the Haversian systems. in balsam the Mount head of i. ends of those bones which fit together to form moveable joints the caps act as elastic cushions to break the force of shocks. The lacunae are the spaces in the matrix in which In sections of dried bone the the bone-cells lie. A Haversian canal with its contained and its surrounding layers of matrix and cells. are together spoken of as a Haversian system. The matrix is richly impregnated with inorganic salts. is arranged in concentric layers or lamellse. through being filled either with air or with dirt. ii. around tubular passages. Bone consists of a dense fibrillar intercellular matrix. The Haversian systems form the greater part of the bone. The matrix is hyaline or faintly granular. the Haversian Canals. both low arid high powers preimred transa long bone.thirds by weight of the substance of the bone. and give it its great hardness and strength. in which lie the bloodvessels. in which are imbedded cells which lie in cavities connected with one another by fine branching canals. : the a prepared section of articular cartilage from femur. interstitial lamellae ii. The fill the Haversian systems.

which consists of adipose tissue. 49 The canaliculi are very fine branching canals connecting the lacunae together they are occupied while the bone is living by branching processes of the bone-cells. with very numerous bloodvessels and large nucleated reddish marrow large central occupied during cells. iv. layer. opening at both ends of the loop into the same lacuna. vi. are a series of concentric lamellae parallel to the surface and forming its most superficial vii. some of the canaliculi are looped. : V. At the outer part of each Haversian system. E .BONE. The perimeduUary lamellae tric lamellae lining are a series of concenthe central medullary cavity of the bone. The medullary cavity of the bone is life by the marrow. The peripheral or circumferential lamellae of the bone.

It forms a is composed partly of cartilage and partly of bone. including the limbs.e. The skeleton may conveniently be divided and make careful drauings to In your drawings colour the cartilage blue. have calcareous salts deposited in its matrix. framework giving definite shape to the body. Cartilage may also become calcified. notably the central In the early nervous system. including the skull and the vertebral column and (2) the appendicular portion. stages of its development the skeleton consists entirely of cartilage in the adult this primary cartilaginous skeleton is replaced to a greater or less extent by bone. Examine skeleton is clean. Membrane bones dermis or deeper layer of the skin in many fish they retain this primitive position. which forms the hard internal parts of the frog. without in any into (1) the axial portion. and is then skeleton. and serves also to protect from injury some of the more important and delicate organs. Bone may also be developed in places where there was no pre-existing cartilage. : arise in the first instance as ossifications in the way taking on the character of true bone.. in contradistinction to the former kind. and carefully brushing away the soft tissues until the the ^jrej^areci skeletons. and the limb-girdles which attach them to the body. Prepare skeletons for yourself by soaking the 2^<^^'ts in hot water. or cartilage-bone. : The called 7nemhrane-hone. the sense organs and the heart.50 CHAPTER IV. and precision to the movements. and the membrane hones white or red. the cartilage hones yellow. scale of the several parts. i. which replaces : cartilage. . but in the frog and most higher vertebrates they sink below the skin and graft themselves on to the more deeply placed cartilaginous skeleton. THE SKELETON OF THE FROG.

carpus m. humerus i. seen from the dorsal surface . urostyle c. the suprascapula and scapula have been removed. femurmetarcarpals h. : : . ilium :k. exoccipital /. maxilla 71. The skeleton of the frog. pterygoid maxilla: q. metatarsalsI. sacral vertebra t. 51 Fig. quadratojugal: r radio-ulna squamosal: sersphenethmoid: s.v. a. tibio-fibula u. suprascapula e.THE SKELETON OF THE FROG. nasal o. 8. pro-otic V7n mep. : : : : calcaneum : gr : : : : • : -s. frontoparietal : d. astragalus : left fp.

face downwards and slightly outwards. The transverse processes are a pair of large processes projecting horizontally outwards from the sides of the neural arch. The posterior articular processes." This is a bony tube which surrounds and protects the spinal cord . is is ii. and forms the sides and roof of the neural canal. The centrum or body tion of the ring : the thickened ventral porit it articulates with the centra of . and forms The neural arch consists of the lateral and dorsal portions of the ring .52 THE SKELETON OF THE FROG. it consists of an anterior part which is divided transversely into nine rings or vertebrae. or praezygin- upwards and slightly f3. and a posterior unsegmented portion of about equal length the urostyle. Examine one of the vertehrce. . The vertebra during life a bony ring . articulate with corresponding processes on the vertebrae in front and behind. I. showing the following points i. The anterior articular apophyses. — a. Structure of a vertebra. The Axial Skeleton. The Vertebral Column or "back bone. iv. more closely : draw it. A. or postzygapophyses. vi. The articular processes or zygapophyses. projecting upwards and backwards from the top of the neural arch. say the third. are the intervertebral foramina through which the nerves pass out from the spinal cord to the various parts of the body. and so serve to link the vertebrae together. a. the s]3inal cord lying in the central neural canal. face wards. on the anterior and posterior borders of the neural arch. v. processes. The spinous process or neural spine is a small blunt median process. At the sides of the tube. iii. the vertebrae in front of and behind the floor of the neural canal. between the successive vertebrse.

53 Special verteTjraB. The sacrum. The atlas or first vertel)ra articulates : in it front has no Note the large gap on the transverse processes. large tracts of unossified cartilage persisting in the adult. lo. the cranium. : : 1. and lodges the brain. body of the sacral vertebra by two surfaces. or ninth vertebra. enclosing the brain and forming an anterior continuation of the vertebral column: (2) the olfactory capsules and the auditory capsules. the central nervous system is divided and destroyed in the operation of pithing a frog. In the skull the original cartilage. Along its dorsal surface runs a prominent vertical ridge. The urostyle is the unsegmented posterior portion of It articulates in front with the the vertebral column.THE VERTEBRAL COLUMN. (1) an axial portion. is not so largely replaced by bone as in the vertebral column. are a pair of small holes through which nerves pass out. : II. dorsal surface between the skull and the neural arch of the atlas through this gap. i. 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. At the sides of the urostyle. whose cavity forms the anterior part of the neural canal. c. and about the length of a vertebra from its anterior end. which is closed by the strong occipito-atlantal membrane. : with the posterior end of the skull ii. The Cranium is an unsegmented cartilaginous tube. The skull consists of. The roof of the tube is imper- . has very stout backwardly directed transverse processes which support at their outer ends the pelvic arch. Besides the cartilage-bones the skull is further strengthened by the addition of numerous membrane-bones. and which therefore correspond to intervertebral foramina. The Skull. highest in front and gradually diminishing posteriorly the neural canal is continued down the anterior part of this ridge. or chondrocranium.

TH^ SKELETON OF THE FROG. pro-otic _?. quadratojugal se. They almost completely surround the foramen magnum or entrance to the cranial cavity . vomer pterygoid m. frontoparietal pa.. palatine ?i. i. pm.54 feet. The fronto-jparietals are two long flat bones on the top of the brain-case. : from the ventral : : surface. To study a. 9. in which lie the : olfactory sacs. and two smaller posterior fontanelles. The exoccipitals are two irregular bony masses at the sides of the posterior end of the skull. b. Cartilage-bones of cranium. and extends forwards into the olfactory region in front it is divided by a vertical partition into right and left cavities. sphenethmoid. a. . i. In the cartilage are developed cartiit and around membrane-bones. columella f. exoccipital c. parasphenoid e. which are closed by membrane lage-bones. the be stripped from one of the skulls membrane-hones should you have p)yepared for yourself. The : frog's skull.p. and bear on their posterior surfaces the occipital condyles. Fig. : : : : : ii. only. jjremaxilla q. covering the fontanelles. the cranium satisfactoyily. maxilla o. there l^eing one large anterior fontanelle. two oval convex processes which articulate with the first vertebra or atlas. Membrane-hones of cranium. The sphenethmoid or girdle-hone is a bony tube which encircles the anterior end of the cranial cavity.

and forms the inner boundary of the posterior narial opening of its side.THE SKULL. a diamond is shaped patch in which the sphenethmoid on the dorsal surface of the skull. i. with the anterior ends of the frontoparietals. its lateral processes underlying the auditory capsules. visible 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. ii. The auditory capsules are fused with the sides of the posterior end of the cranium. and forming also parts of their roof and floor. 2. The pro-otics are a pair of irregular shaped bones in the anterior walls of the capsules. h. Membrane-hones of olfactory capsules. a. and also with each other. extends forwards so as to invade the olfactory region. The Sense Capsules are cartilaginous and bony capsules which surround and protect the olfactory and auditory organs . Cartilage-hone of auditory capsules. The parasphenoid surface of the is a X shaped bone on the ventral cranium . Cartilage-bone of olfactory capsules. /3. They consist very largely of cartilage. while their enclose. they are fused with the cranium so as to form parts of the skull. . but does not properly belong to the olfactory capsules. to which they form wing-like projections: they consist largely of cartilage. 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 posterior and meet each other in ends diverge and front. a. i. ii. The sphenethmoid. The olfactory capsules of the are fused with the anterior end cranium. as already noticed. which is produced in front into the rhinal processes. 55 and overlapping the hinder end of the sphenethmoid.

In the case of the pterygoid and palatine bones ossification may extend into the underlying cartilage. consist of a. Each arch meets its fellow in the middle line in front. nostril E. A. MMi mentomeckelian M'. in connection with which cartilage -bones and membrane -bones are developed. angulosplenial parasphenoid FP. and the maxilliary arches. dentary C. anterior cornu of hyoid AS. bone. aperture for exit of ninth and tenth nerves ethmoid : T. The 'palatine is a slender transverse bone. exoccipital D. The Maxillary Arch. frontoparietal F. forming the upper jaw. squamosal SE. The iJterygoid is a large triradiate bone. maxilla L. maxillary and mandibular. are firmly connected with the cranium by anterior and posterior bony struts. nasal S. : B. ii. : : : : : : : : : : : : i. pterygoid O. quadratojugal P. . posterior cornu of hyoid. -PIVl MM Fig. The Jaws two cartilaginous arches on each side. 10. pro-otic N. : The : frog's skull from the right : : : side. connecting the upper jaw with the anterior end of the splienethmoid. body of hyoid aperture for exit of ojDtic nerve M. premaxilla Q. and the anterior limb forwards along the upper jaw to the palatine . the inner limb of which is connected with the auditory capsule while the posterior limb runs back to the angle of the mouth. aperture for exit of fifth and seventh nerves PM. THE SKELETON OP THE FROG. columella H. sphenR. All the bones of the upper jaw are membrane-bones except the quadratojugal.56 3.

the quadrate cartilage lies between the squamosal and pterygoid bones. The posterior limb of the is attached to the outer surface of the auditory T T capsule. The upper part of the arch remains unossified as the quadrate cartilage. In the adult frog. and so completes the margin of the upper jaw like the maxilla it bears teeth. where it is connected with the hinder end of the quadratojugal bone.THE SKULL. The angulosplenial ensheaths the inner and lower surfaces of Meckel's cartilage along the greater its length near its hinder end it produced upwards into the coronary process. It is connected behind with the quadra tojugal . and in front with the premaxilla. and with the body of the squamosal helps to support the annulus tympanicus. the stem of closely applied to the outer surface of the quadrate cartilage. and runs downwards and backwards to the angle of the mouth. which forms the suspensorium. : V. 1). 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. and is almost completely concealed by these. 57 The quadratojugal terior part of the is margin a short bone forming the posof the upper jaw. iii. The squamosal which is is a shaped bone. serves to connect the lower jaw with the skull this is a rod of cartilage which is fused above with the auditory capsule. The Mandibular Arch. It gives off on its dorsal surface a backwardly projecting process which forms part of the inner boundary of the nostril.e. : i. which forms the basis of the lower jaw.. : part of is . ii. and is ensheathed by cartilage-bones and membrane-bones. iv. about the middle of its length with the anterior limb of the pterygoid and with the palatine . The lower part of the arch persists in part unossified as Meckel's cartilage. i. The premaxilla is a small bone which meets : its fellow in the middle line in front.

vocal cord X. A. A transverse section across the posterior jiart of the frog's head. are represented black. quadratojugal S. : : : : : : : : : : I. and lies in the floor of the mouth. and the bones. and hyoid apparatus. The hyoid arch. columella D. : : : : : : : : : : : : 4. Eustachian tubes. except the columella. The mentomeckelian is a small ossification in Meckel's 'cartilage at the symphysis. The Hyoid Apparatus (Fig. auditory nerve cornu of hyoid L. pterygoid R. iv. quadrate cartilage T. pro-otic P. The cartilage is dotted. squamosal Q. tymi)anic membrane E. The columella (Figs. Eustachian tube F. together with a median ventral plate. which unites their lower ends together. annulus tympanicus V. The hyoid apparatus consists almost entirely of cartilage. . the union of the arches of the two sides at the chin. THE SKELETOX OF THE FROG. anterior vertical semicircvilar canal horizontal semicircular canal N.58 iii. anterior cornu of the hyoid FP. angulosplenial B. vestibule M. 10). parasi)henoid AS. On the right side the section passes through the tympanic cavity and the cohimella on the left side through the anterior cornu of the hyoid. to show the position and relations of the auditory organs. mid-brain. 10 and top of the hyoid arch : 11) it is is formed from the a small rod. i.e.. FP X K M Fig. frontoparietal G. glottis H. O. This consists of the hyoidean arch and the remains of the branchial arches of the two sides. a. as far forward as the mentonaeckelian bone. partly . i. the body of the hyoid. arytenoid cartilage posterior K. buccal cavity C. 11. The dentary is a flat bone covering the outer surface of the distal half of Meckel's cartilage.

formed by the fused ventral ends of the hyoid and branchial arches. which. bones developed independently of cartilage. 8." Each half ring bears in the middle of its hinder surface a cupshaped cavity. As in the case of the axial skeleton it consists at first entirely of cartilage. 51). while the ventral ends are united together in the median plane by the sternum or " breast bone. p. The posterior cornua of the hyoid are a pair of stout bony processes. while the outer end is attached to the tympanic mem- brane rather above ii. the clavicles being the only examples met with in the frog. 1. are very rare. The body of the hyoid is a flat squarish plate of cartilage. The anterior cornu of the hyoid (Fig. This comprises the limbs and the limb-girdles. which is divided into anterior and posterior divisions. the inner end of which inserted into the fenestra ovalis. bone and partly is 59 cartilage. (Fig. and lying in the floor of the mouth short processes are given off" from its angles. its middle. forms The part of the arch above the joint is the the shoulder joint. with the first bone of the fore-limb. 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. b.e. diverging from the hinder border of the body of the hyoid. which becomes afterwards replaced to a greater or less extent by cartilage-bone. The Pectoral Girdle. 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. the coracoid portion. an aperture in the outer wall of the auditory capsule . one on each side of the body. 10) is a long slender curved rod of cartilage.THE APPENDICULAR SKELETON. scapular portion: and the part below. attached above to the auditory capsule just below the fenestra ovalis.. c. : . The Appendicular Skeleton. This consists originally of two half rings of cartilage. i. Membrane-bones.

60 a. is an oblong bony plate. separated by the coracoid foramen. constricted in the middle. the upper : a thin expanded plate of cartilage overlapping the first four vertebrae it is partly calcified and partly ossified. but remain distinct It presents from before in the median portion. THE SKELETON OF THE FROG. The precoracoid a slender horizontal bar of caredge of the scapula with the sternum. is The suprascapula. closely applied to the anterior border of the precoracoid . backwards the following parts originally of : i. . iii. The clavicle is a slender bone. ii. portion. ii. iv. The omosternum is a slender bony rod projecting forwards in front of the clavicles. i. V. : C. The scapula 1). The scapular portion i. coracoids. or cavity of the shoulder joint. though very imperfectly. The epicoracoids are a pair of narrow strips of and lying between the ventral ends of the precoracoids and cartilage closely applied to each other. its outer or scapular end is bent forwards almost at a right angle. The episternum is a flat circular plate of cartilage. is tilage connecting the anterior ii. The xipMsternum is a broad cartilage at the hinder expanded plate of end of the sternum. The coracoid portion forms the lower : half of the glenoid cavity it is divided into anterior and posterior portions. it connects the posterior than its outer end edge of the scapula with the sternum. The coracoid wider at its inner is a stout bone. and forming the upper half of the glenoid cavity. is divided into two parts. The sternum lies in the mid-ventral line : it consists two lateral halves which fuse completely in front and behind. iii. ensheathed backwards behind the coracoids. The sternum proper is a rod in bone. projecting of cartilage.

The hand has four complete digits and a rudimentary poUex or thumb. in other animals it is single at its : proximal end. The pollex. is nearer to the body. i. hollowed out to articulate with the lower end of the humerus. is a spheroidal articular surface for the bone of and at either side of this a the fore-arm prominent condylar ridge. simply of a small metacarpal bone. 2. they are elongated^ The with enlarged ends capped with articular cartilage. the inner or postaxial one being the larger of the two. with which they do not unite until late in life. The fore-arm. i. d. c. each row having three bones. proximal and distal. frog. radius and ulna. and Its proximal end is posterior or ulnar portions. The arm. i. is called its proximal end. especially in the : male b. but in its distal half is imperfectly divided by a groove into anterior or radial. behind which it projects backwards as the olecranon process. The Fore-Limb. consists . enlarged ends or epiphyses ossify independently of the shaft of The the bone. In the arm there larged. and so form the elbow joint. The wrist consists of six small carpal bones arranged in two rows. end of a bone which. With the exception of the small bones of the wrist. the opposite extremity the distal end a. The radio-ulna corresponds to two bones. The proximal end or head is enand articulates with the glenoid cavity below the head is the of the pectoral girdle : strong deltoid ridge extending along the proximal At the distal end half of the anterior surface. when the limb is extended. beyond which are a variable number of phalanges.THE APPENDICULAR SKELETON. the anterior or preaxial digit. The bones of the fore-limb 61 are all cartilage-bones. Each digit consists of a proximal metacarpal bone. is only a single bone. The humerus.

The first complete digit. Girdle. about one-sixth. consists entirely of cartilage : ii. The Pelvic This consists primitively. fused together below' and attached above to the tips of the transverse processes of the sacrum. which is attached in front to the transverse process of the sacrum and bears along its dorsal surface a prominent vertical ridge of bone. Posteriorly the two ilia meet each other and are united together in the median plane to form the iliac symphysis. Each half presents on its outer aspect a cup-shaped cavity the acetabulum forming. the hip-joint we accordingly distinguish an iliac portion above the acetabulum and an ischio-pubic portion below it.62 ii. — : i. and form only a very small portion. and extends forwards as an elongated laterally-compressed bar. In the adult frog the girdle is pla. the iliac crest. The ilium forms the anterior and upper half of the acetabulum. vi. consists of a metacarpal and three phalanges. The third. The two pubes are completely fused together in the median plane. The second finger of man. corresponding respectively to the scapular and coracoid divisions of the pectoral girdle. consists corresponding to the middle of a metacarpal and two phalanges. like the pectoral girdle.ced very obliquely so as to be nearly parallel with the vertebral column instead of at right angles to it. iii. The postaxial digit. corresponding to the ring finger of consists of a metacarpal V. corresponding to the fore finger of man. The pubes it forms the anterior portion of the ventral division of the girdle. corresponding to the little finger of man. man. with the thigh bone. . and therefore corresponds to the precoracoid in the pectoral girdle. and three phalanges. of the acetabular cavities. ending behind in an abrupt vertical border. of a couple of half-rings of cartilage. phalanges. THE SKELETON OF THE FROG. consists of a metacarpal and two digit. 3.

metatarsal bone. i. C. 4. expanded at both ends.THE APBNDICULAR SKELETON. ii. The two ischia are completely fused together in the median plane. 63 of the ventral The ischium is the posterior portion division and corresponds therefore to the coracoid in the shoulder girdle. The foot has five complete digits. It presents along the greater part of its length a groove indicating its correspondence with two bones. which in man and many other animals remain distinct from each other. slightly curved. The ankle. The Hind-Limh. and a supernumerary Each digit consists of a proximal toe as well. and fits into the acetabulum to distal end is form the hip joint somewhat expanded laterally. The distal row of tarsal bones consists of two very small bones. iii. The bones have the same general characters as those of the fore-limb. and is is on the postaxial or fibular the larger of the two bones. and expanded laterally at both ends. corresponding consists of a. which tliey correspond very closely. leg. The i. The proximal row of tarsal bones consists of two elongated bones united together at both ends. tibia and fibula. : the h. two rows of tarsal bones. rather longer than the femur. i. beyond which are a variable number of phalanges. The proximal end or head is spheroidal. . The thigh. d. to the wrist in the fore-limb. The femur is a long slender bone. It forms the posterior third of the acetabulum. The astragalus The calcaneum side. The OS cruris or tibio-fibula is a single bone. to a. and curved slightly in a sigmoid manner. but widely separated in the middle. is on the preaxial or tibial side. /3.

consists of a metatarsal and three phalanges. ii. The in postaxial digit. corresponding to the little toe man. The second phalanges. The third consists of a metatarsal and three phalanges. iv.64 i. toe consists of a metatarsal and two iii. is the smallest of the series. The hallux or preaxial digit. corresponding to the great toe of man. V. THE SKELETON OF THE FROG. On the : inner side of the hallux is the calcar. The fourth. . supposed to be an additional or supernumerary toe it may have one or two joints in addition to a short metatarsal. consists of a metatarsal and four phalanges. the longest of the five. It consists of a metatarsal and two phalanges.

which is usually attached at each end by means of tendons to some'^hard Motion is effected by the muscle part. or both. and it : define test its action by pulling its fibres. are the direct means by which the movements of the body and of its several parts are brought about. but one or other end. the other to a more moveable and peripheral part the former attachment is called the origin The various : of the muscle. which are the only ones dealt with in this chapter. and separate them from one another.. are omitted. or of the alimentary canal. F . and consequently the parts to which the ends are attached. and not across its fibres and insertion of the muscle very clearly.e. i. A muscle consists of a fleshy belly.. as the muscles of the arm and (2) involuntary muscles. i. the latter its insertion. as the muscles of the heart and : : bloodvessels. Of the two attachments of a muscle one is usually to a more fixed and central part. those which are under the control of the will. Voluntary muscles. In the folloidng description some of the smaller muscles. those over which the will has no direct control. the stretch. may be attached to aponeuroses. very commonly to bone.e. muscles. take a frog that has been in spirit When cleaning a muscle be careful to put it on day or more. especially in the head. Always the gently with the forceps in the direction of have the skeleton in front of you so as to see accurately origins and insertions of the muscles. nearer together.e. and so bringing its two ends. For for a the dissection of the muscles. or flesh. the origin and to dissect along.. contracting. strong connective tissue membranes which ensheath the muscles and other parts. shortening. THE MUSCULAR SYSTEM OF THE FROG. are usually attached at both ends to bone .65 CHAPTER V. Muscles are of two kinds (1) voluntary muscles. i.

the fibres running obliquely downwards and backwards to end in an aponeurosis which passes dorsal to the rectus abdominis to be inserted into the linea alba. Muscles of the ventral body-wall. Pin out the frog on its hack. The obliquus externus which arises is a thin sheet of muscle from the aponeurosis of the back. the oesophagus and pericardium. consisting of a thoracic portion. The muscle arises from the pubes. I. the vertebrae from the fourth backwards. into which they are inserted. which must be removed in order to It arises from the transverse processes of it. verge to the deltoid ridge of the humerus. Muscles of the Trunk. the line of insertion extending down almost to the elbow. and clean i. and covers the whole of the side of the body.66 THE MUSCULAR SYSTEM OP THE FROG. sternum and an abdominal portion. The rectus abdominis runs longitudinally along the mid ventral wall. and are inserted into. runs forwards and is inserted into the dorsal surface of the sternum and coracoid. a short distance from the vertebral column. A. see : . and are inserted in front into the coracoid and sternum some of the fibres surround. the muscles. the muscles of the two sides being separated from each other in the median plane by the linea alba. immediately dorsal to which Each rectus lies the anterior abdominal vein. a longitudinal band of connective tissue. The obliquus internus lies beneath the obliquus externus. arising from the aponeurosis along the outer side of the rectus abdominis almost as far back as the From this extensive origin the fibres conpubes. muscle is divided into bellies by five transverse tendinous intersections. which arises from the whole length of the ventral surface of the . ii. iii. and from the ilium. iv. The pectoralis is a large fan-shaped muscle. The fibres run downwards and forwards. remove the skin.

viz. and converge to be inserted into the angle of the lower jaw. and is inserted into the dorsal border of the suprascapula. vi. vii. its origin being partly covered by the obliquus externus. The fibres run forwards and outwards. The cucuUaris is a small oblong muscle which. on one side. runs backwards and outwards. out the frog on i. in front. Remove the suprascapula and clean the median longitudinal muscles of the bach. like it. 2. : The depressor mandibulse ii. : Lift up the siqjrascajmla and note the muscles attaching it to the body. V. and clean the muscles in order. arising from the exoccipital near the middle line. iii. iv. mandibulae and latissimus dorsi and twn them down. The retrahens scapulae. into the linea alba. behind. Pin its belly : remove the skin. The levator anguli scapulae. 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. 67 The hinder two-thirds of the muscle pass dorsal to the tendon of the obliquus externus and are inserted. The muscle by its contraction opens the mouth. partly overlapped by the latissimus dorsi it runs outw^ards to be inserted into the deltoid ridge of the humerus its action is to elevate the arm. The extensor dorsi communis is a longitudinal mass of muscle arising from the urostyle. converging to be inserted by a long tendon into the deltoid ridge of the humerus. : Dissect away the depressor from their origins.MUSCLES OF THE TRUNK. Muscles of the Back. and running . The infraspinatus : arises from the dorsal surface of the suprascapula. The latissimus dorsi is a triangular muscle lying behind the depressor mandibulae it arises from the fascia dorsalis just behind the shoulder girdle.

it is inserted forwards and slightly outwards into the ilium. and lying beneath the extensor communis. and the sternal portion of the ii. Muscles of the Head. the tendon passing between the two divisions of the geniohyoid. It arises from the dorsal surface of the coracoid and clavicle. ix. and is inserted into the ventral surface of the body of the hyoid. sheet of muscle running across from one ramus of the mandible to the other. Muscles of the ventral surface of the head. and divides posteriorly into two portions which are inserted respectively into the bony and the cartilaginous posterior processes of the hyoid. The geniohyoid is a narrow longitudinal band a short distance from the middle line it arises from the lower jaw close to the chin. pectoralis. A narrow strip along the posterior border is commonly separated by a slight interval from the major or anterior part of the muscle. The mylohyoid or submandibular muscle is a flat i. The sternohyoid is practically the anterior continuation of the rectus abdominis. and runs backwards to be inserted into the trochanter of the femur. : iii.68 THE MUSCULAR SYSTEM OF THE FROG. The from the outer side of the posterior two-thirds of the ilium. g-lutseus arises B. runs back on the ventral surface of the body of the hyoid. 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 : . iv. and divided down the middle line by a tendinous intersection. into the transverse processes of the vertebrae. I. and in front into the posterior end The anterior part is divided by of the skull. transverse tendinous intersections. Remove the mylohyoid muscle. : viii. The intertransversales are small muscles. running between the transverse processes of the vertebrae.

(See p. ii. . and diverging in a fan -like manner. Depressors of the lower jaw i. and runs to the chin along it backwards to the tip. opening the mouth. m : V. These lie in the space between the auditory capsule and the eye. and behind into the side of the hyoid. and very close to the joint. and in front of the cartilaginous ring supporting the tympanic membrane. The pterygoideus it is a slender muscle placed just in front of the temporalis and partly covered by it arises from the side wall of the skull. iv.MUSCLES OF THE HEAD. Muscles of the side of the head.) b. forward the middle line as a stout band nearly it then enters the tongue. The depressor mandibulae has been already seen and dissected. Elevators of the lower jaw. shutting the mouth. noticing how much more closely it is attached to the underlying parts than ivas the case in the body. The first or most anterior band is a wide thin sheet of muscular tissue. the side of the head and jaws. pass round the floor of the pharynx and oesophagus to be inserted in front into the median ventral line of the pharynx. The temporalis arises from the upper surface of the auditory capsule. and passes outwards and downwards between the pterygoid and maxillary bones. : iii. and is inserted into the mandible further back than the temporalis. while the three posterior portions are very narrow slips. The petrohyoid muscles bands which are a set of four muscular from the outer surface of the auditory capsule. 67. from which some of its fibres arise it is inserted into the coronoid process of the lower jaw. Remove the skin carefully from. 69 In front of the larynx the muscle runs larynx. The masseter temporalis: is it arises a small muscle placed behind the from the quadratojugal and . a. arise close together 2.

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

it is inserted into the under surface seen best from below. The ohliquus inferior passes backwards beneath the rectus internus. inasmuch as the muscles which extend or straighten the leg lie along this edge the inner border is the flexor surface. as in the act : : : therefore the postaxial digit. we distinguish in it ventral and dorsal surfaces. it will be seen that the first digit or " big toe " is on the preaxial side. 71 The rectus inferior : is of the eyeball b. If the foot be examined carefully. iv. The obliqui muscles which arise close together are a group of two small muscles from the palatine bone at the anterior end of the orbit. and is If the frog's leg axis of the body. and run backwards to be inserted into the eyeball.MUSCLES OF THE EYEBALL. : c. and are better called preaxial and postaxial they correspond respectively to the inner and outer surfaces of the human leg. is a funnelshaped muscle which lies within the four recti and embraces the optic nerve it arises from the para: inserted into the eyeball. ii. i. which corresponds to the front of the leg in man. It is best exposed from below by carefully removing the recti muscles. and is inserted into the eyeball between it and the rectus inferior it is seen best from below. be stretched back parallel to the longitudinal of swimming. The obliquus superior : is inserted into the dorsal surface of the eyeball just in front of the rectus superior it is seen best from above. The outer border. The retractor bulbi. or choanoid muscle. and C. is sphenoid. Muscles of the Hind-limb. . an outer border in which is the projection of the knee. and hence may be called the preaxial digit: while the fifth or " little toe " is on the postaxial side. 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. is called the extensor surface. and an inner border along which is the bend of the knee.

just muscles of the extensor surface of the thigh. . and often completely hidden by the sartorius: it arises from the iliac symphysis beneath the sartorius. The rectus internus minor : is a narrow flat band of muscle running along the inner. and is inserted into the tibia just below its head. ii. iv. below its head. arises by three distinct origins. Superficial i.72 1. which will be described separately. It arises from the iliac symphysis below the acetabulum. and is inserted into the inner side of the tibia. The sartorius a long narrow muscular band which somewhat obliquely from the outer to the inner side. but passing beneath it at its distal end. the skin from one of the legs of the frog ^ and clean the muscles first of the j)reaxial and then of the i^ostaxial surfaces. Remove a. and unites a little way beyond the middle of the thigh with the adductor magnus. Superficial muscles of the preaxial (apparent ventral) is surface of the thigh. THE MUSCULAR SYSTEM OF THE FROG. Muscles of the thigh. V. i. b. It arises from the pubic and ischial symphyses. The adductor magnus iii. the great extensor muscle of the thigh. and passes under the sartorius to be inserted into the distal third of the femur. The adductor longus is a long narrow muscle lying along the outer side of the adductor magnus. or flexor margin of the thigh it arises from a tendinous expansion connected with the ischial symphysis. The triceps extensor femoris. crosses the thigh is a large muscle lying along the inner border of the sartorius. The rectus internus major a large muscle lying along the inner side of the adductor magnus and of the sartorius. and is inserted into the inner side of the head of the tibia. It arises from the ischial symphysis and is inserted into the head of the is tibia.

tibialis anticus TP. semimembranosus T. distal end of femur flexor tarsi GL. peroneus PY. tibialis Posticus VE. gastrocnemius P. 12.MUSCLES OF THE HIND-LIMB 1. biceps C. vastus externus VI. adductor brevis AM. sartorius SM. rectus inter]. triceps extensor femoris TF. AB. tendon of F. rectus anticus femoris Rl.- : : : : : . tibio-fibula TA. The superficial muscles of the frog's left hind-limb. cloaca! aperture EC. adductor magiius B.^'Xct^^ Fig. glutseus g. pyrilormis ra. from the preaxial surface. B from the postaxial surface. rectus internus major RN. extensor cruris FT. A : : : : : : : : : : : : : : : : : . vastus internus.ius minor S.

and is inserted by : a flattened tendinous expansion into the distal end of the femur and the head of the tibia. and lying in the thigh between the sartorius and the rectus anticus. is a large muscle arising from the ventral and anterior border of the acetabulum. The pyriformis . between it and the It arises from the dorsal rectus internus minor. Superficial muscles of the postaxial (apparent dorsal) surface of the thigh. : p. middle by an is a slender muscle which arises from the tip of the urostyle. and is inserted into the femur at the junction of its proximal and middle thirds. a. ii. i. angle of the ischial symphysis just beneath the cloacal opening. c. oblique tendinous intersection. The glutseus has been already noticed : it lies in the thigh between the rectus anticus and the vastus externus. and is inserted into the back of It is divided about its the head of the tibia. The semimembranosus is a stout muscle lying along the inner side of the biceps. iv. The vastus externus. in front of the acetabulum about half way down the thigh it joins the next division. and joins the other two divisions of the triceps about the junction of the middle and distal thirds of the thigh. the the triceps. The vastus internus. iii. the preaxial division of the triceps. passes backwards and outwards between the biceps and the semimembranosus. postaxial division of y. The rectus anticus femoris forms the middle division of the triceps: it arises from the ventral border of the posterior third of the ilium.74 THE MUSCULAR SYSTEM OF THE FROG. arises from the posterior edge of the dorsal crest of the ilium. 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.

The quadratus femoris of the is upper part of the thigh a small muscle on the back it arises from the : . lying beneath the upper end of the adductor magnus. an anterior one from the ischium close to the ventral angle of the ischial symphysis and the acetabulum . and is inserted into the back of the proximal half of the femur. : : V. : The semitendinosus arises is a long thin muscle which by two heads. The tendon of insertion is long and thin. The ilio-psoas arises by a wide origin from the inner surface of the acetabular portion of the it turns round the anterior border of the ilium ilium. d. so as to expose the ii. The pectineus is a rather smaller muscle. and crosses in front of the hip joint. iv. is and iii. Seijarate the adductor magnus and the rectus internus major with hlimt instruments so as to expose the following muscles i. Lay the frog on its hack and dissect the thigh from the preaxial surface. It arises from the pubic and ischial symphyses. The anterior head passes through a slit in the adductor magnus and unites with the posterior head in the distal third of the thigh. 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. inserted into the preaxial surface of the proximal half of the femur. lying along the outer (extensor) side of the adductor brevis. and is inserted like it into the proximal half of the femur. 75 Deep muscles of the thigh. It arises from the anterior half of the pubic symphysis in front of the adductor hrevis. and joins that of the rectus internus minor to be inserted into the tibia just below its head. and a posterior one from the ischial symphysis. following muscles: The adductor hrevis is a short wide muscle. . and turn Divide the adductor magnus and the sartorius in the middle the cut eyids backwards and forwards.MUSCLES OF THE HIND-LIMB.

while the smaller head. The gastrocnemius calf of the leg : the large muscle forming the has two heads of origin. The tibialis posticus arises from the whole length : it ends in a of the flexor surface of the tibia tendon which passes round the inner malleolus. of which the larger arises by a strong. corresponding to the inner and outer sides of the human leg. and is inserted into the head of the femur close to the glutaeus. being much thickened as it does so. flattened tendon from the flexor surface of the distal e. and divides about tibialis anticus lies : of the leg it arises . lying in a groove in the lower end of the tibia. i. and tapering posteriorly ends in the strong tendo AcMllis. arises from the edge of the tendon of the triceps extensor femoris where it covers The muscle is thickest in its upper the knee. we distinguish extensor and flexor surfaces. iii. which passes under the ankle joint.76 THE MUSCULAR SYSTEM OF THE FROG. 2. : of the iliac crest vi. and is inserted into the dorsal surface of the astragalus.ad of the femur . corresponding to the front and back of the leg in man . ilium above the acetabulum. it ii. As in the thigh. Lay the frog on its belly and commence is the dissection from the postaxial surface. and is inserted into the inner surface of the proximal third of the femur between the pyriformis and the ilio-psoas. The on the extensor surface by a long thin tendon from the lower end of the femur. The obturator arises is a deeply situated muscle which from the whole length of the ischial symphysis and the adjacent parts of the iliac and pubic symphyses. and also preaxial and postaxial surfaces. third. Muscles of the Leg. and from the base it lies beneath the pyriformis and behind the biceps. which joins the main muscle about one-fourth of its length below the knee. and ends in the strong plantar fascia of the foot.

77 the middle of the leg into two bellies which are inserted into the proximal ends of the astragalus and calcaneum iv. and is inserted into the outer malleolus of the tibia and the proximal end of the calcaneum. V. respectively. and is inserted into the extensor surface of the tibia along nearly its whole length. partly covered by this and partly by the strong fascia of the leg. between the tibialis anticus and the gastrocnemius. runs in a groove in the upper end of the tibia. The extensor cruris lies along the preaxial side of the tibialis anticus. .MUSCLES OF THE HIND-LIMB. It arises by a long tendon from the preaxial condyle of the femur. The peroneus is a stout muscle which lies along the postaxial surface of the leg. It arises from the distal end of the femur.

boundary between medulla oblongata and spinal cord 1-10. ramus anterior of glossopharyngeal 3-4. the nerves themselves. THE NERVOUS SYSTEM OF THE FROG. hyomandibular branch of facial: Vg. optic tract Lop. : : The nervous system of the (From Ecker. sciatic nerve No. trigeminal and facial nerves Va. ganglion of pneumogastric nerve He. and which is the centre where sensations are felt. conveying impulses to the brain or cord and (6) efferent or motor nerves. motor oculi nerve V. . I. maxillary branch of trigeminal Vd. sense organs.. mandibular branch of trigeminal Ve. A peripheral portion. VII. or motor impulses from the central organs to the muscles. abducens nerve. The nervous system 1. olfactory nerve IV.78 CHAPTER VI. X2. conveying impulses frojn the brain or cord. uiiper end of sympathetic trunk. optic lobe M. facial nerve G. in connection with Gasserian ganglion VI. muscles. These two functions are fulfilled by different nerves. viscera. the ventral surface. fourth II.) : edible frog {Rana esculenfa)^ : from F. the "brain in the cartilaginous causing the muscles to contract take their origin. trunk of sympathetic S 1-10. glossopharyngeal and pneumogastric nerves XI. continuation of : M : : : : : : : : symi)athetic into head. crural nerve o. etc. connection between fourth spinal nerve and sympathetic chain N. nasal sac Ni. cerebral hemisphere Lc. auditory nerve: A. optic nerve III. eyeball S. and serve to convey sensory impulses from these parts to the brain and cord. 2. lig. the spinal nerves: MS. Gasserian ganglion _Vs. 13. which lies and bony tube formed by the cranium and vertebral column. and whence motor impulses central portion. the sympathetic ganglia Sp. facial nerve VIII. ramus posterior of glossopharyngeal : : : : : : '• : : : : : : : : X branches of pneumogastric. consists of the following parts :-^^ A and spinal cord. ophthalmic branch of trigeminal Vc. which connect the central ]3ortion with the skin. which may accordingly be distinguished as (a) afferent or sensory nerves.

THE CENTRAL NERVOUS SYSTEM. 79 .

removing the pigmented membrane pia mater covering the several parts as you come to them. are united together in the median plane they give off the olfactory nerves from their anterior ends. A. a.80 THE NERVOUS SYSTEM OF THE FROG. on one side and then on the other. clear away the dorsal muscles from. portion of the brain. The cerebral hemispheres are a pair of smooth ovoid bodies which touch each other in the median plane but are not fused together. A special set of nerves in connection with the bloodvessels and viscera forms the sympathetic nervous system. and are separated behind by slight constrictions from the hemispheres. injure the brain beneath Introduce 07ie blade of the scissors into the cranial cavity. which are directly continuous with each other. through the occipito-atlantal slightly to make the both sides of the spine : cut membrane. ii. Turn the roof of the skull first forwards with forceys. This is divisible into in the cavity of the spinal cord which lies in the neural canal of the vertebral column. with the flat surface of the blade parallel to the back of the frog. The dorsal surface of the brain: note from before backwards the following parts.. There is no sharp line of demarcation between the two portions. which form the most anterior i. and in which the hrain has been exposed to the action of the spirit by removal of the roof of the skull. — an anterior portion the hrain lying cranium and a posterior portion the . Examine and draw the central nervous system in situ. — — : . The Central Nervous System. For the dissection of the nervous system specimens should he taken which have been in strong spirit for two or three days. its several parts. The Brain. The olfactory lobes. Similarly cut through and remove the neural arches of the vertebrce one by o7iefrom before backwards. showing as possible. and being careful not to it. and keeping as close to the roof of the skull Cut carefidly through the side walls of the skull. — — — If the brain and spinal cord have not already been exposed. — I. flexing the frog's head membrane tense. and remove it altogether.

Fig. fourth ventricle IN. or cavity of the thalamencephalon. body : : : : : : : : : roof of the thalamencephalon through which the vessels of the plexus pass into the third ventricle. tuber cinereum M.THE BRAIN. olfactory nerve ll. The optic lobes are a pair of prominent ovoid bodies G . cerebellum CH. and VIII. fourth nerve V. sixth nerve VII. dorsal surface. optic chiasma Fig. 15. cerebral hemisphere ventricle F. a small body adherent to and reOn removing roof of the skull. : The brain The brain : of the frog of the frog : : : : : : : : PB. CP. thalamencephalon. the choroid plexus a slit-like hole is left in the O. choroid plexus of third C. iv. olfactory lobe OC. P. third or motor oculi nerve I. medulla oblongata. The thickened sides of the thalemencephalon are the optic thalami. 14. i:)ituitary body T. stalk of pineal OL. ventral surface.L x 4. x 4. III. 81 The thalamencephalon is a lozenge-shaped portion lying immediately behind the hemispheres and it is between their diverging posterior ends covered by a thick vascular membrane the choroid plexus over which passes the stalk of the : — — moved with the pineal body. combined root of facial and auditory nerves IX and X. optic lobe O. fifth or trigeminal nerve VI. combined root of glossopharyngeal and pneumogastric nerves. iii. optic nerve IV.

82

THE NERVOUS SYSTEM OF THE FROG.
touching each other in the median plane, and forming the widest part of the brain the pia mater covering them is very strongly pigmented.
:

V.

The cerebellum

is a narrow transverse band immediately behind the optic lobes.

VI.

The medulla oblongata

is the part of the brain the cerebellum it is widest in front and behind gradually tapers towards its posterior end, wh^re It is it is continuous with the spinal cord. covered by a triangular and very vascular membrane, beneath which lies the fourth ventricle.
:

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

A

:

:

:

ventricle.

b.

The

cavities of the brain.
the upper surface

Slice

of

of the brain horizontally

so as to
it

expose the several cavities or ventricles, without removing

from

THE BRAIN.
the skull.

83

original central canal of

These cavities are merely 'parts of, or outgrowths of, the the neural tube of the embryo. {Gf.

Chap. IX.)
i.

The lateral ventricles extend through the whole length of the cerebral hemispheres and a short

way
ii.

into the olfactory lobes.
is situated in the thalamenopens in front through the foramina
:

The third ventricle
cephalon
:

^

it

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 aquaeductus Sylvii
ventriculum
is

or iter a tertio a quartum 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
vascular

is

in the medulla, already exposed

the large triangular cavity by removal of the
it.

membrane

coverino' O

c.

The ventral surface of the brain.
the medulla at the level of the hinder end of the carefully remove the brain from the cranial cavity, noting

Cut through
skidl
:

the several nerves arising

from
parts.

the brain as possible.

from it, and cutting through these as far Lay the brain on its dorsal surface,

and examine and draw
i.

the ventral surface, showing the following

The

optic

cMasma

is

formed by the decussation

; the point of crossing being opposite the hinder ends of the hemispheres, and immediately in front of the

of the roots of the optic nerves

infundibulum.
Trace back the optic nerves behind their point of crossing
their origins from the ojJtic lobes.
ii.

to

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
iii.

THE NERVOUS SYSTEM OF THE FROG.

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 infundibulum 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.
fissure of the brain is a median longitudinal 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.

V.

The ventral

II.

The Spinal Cord.
spinal cord

is a somewhat flattened band, presenting and lumbar enlargements opposite the points of origin brachial of the nerves for the fore and hind limbs respectively, and About the level slightly constricted between these two points.

The

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 making transverse sections of it. See p. 95.
B.
1.

is

best seen on

The Peripheral Nervous System.

from the each nerve arising by two roots, a sides of the spinal cord 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

The Spinal Nerves.

Ten

pairs of nerves arise

;

:

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 The roots opposite their points of origin from the spinal cord. 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
ivalls,

the frog on its hack : cut through and jom out the hodyand remove the abdominal viscera. Note the spinal nerves,

seen as white cords at the sides of the vertebral column. the nerves on one side and follow them to their distribution.

Clean

Each nerve

divides, directly after the

into a small dorsal branch,
1.

union of its two roots, and a much larger ventral branch.

The hypoglossal,

or first spinal nerve, leaves the vertebral canal between the first and second vertebrse, and then runs forwards on the under surface of the head beneath the mylohyoid and in the substance of the geniohyoid muscle to the
chin,

which it tongue and floor of the mouth, and also some of the muscles (Fig. 15.) of the back and shoulder.

where

it

enters the

tongue, in

ends.

It supplies the muscles of the

2 and 3. The second and third spinal nerves leave the canal between the second and third, and they third and fourth vertebrae respectively unite together to form the brachial nerve, which gives off a large cor aco clavicular 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, supplying the

forearm and hand.
4, 5,

and

6.

The

are small

fourth, fifth and sixth spinal nerves 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 vertebrae 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

Outside the vertebral canal the three nerves unite together opposite the middle of the urostyle to form the sciatic plexus. roots. and the mode of their union with one another are subject to considerable individual variation in different frogs. giving branches to it. 10. The ganglia on the dorsal roots as they pass through the intervertebral foramina these are covered on their ventral surfaces by whitish calcareous patches. Note the folloidng points i. which runs down the thigh. and other adjacent parts. bladder. scissors the centra of the vertehrre one hy one. b. from Avhich branches are given to the large intestine. The Cauda equina. The seventh nerve leaves the canal between the seventh and eighth vertebrae. iv. Just before joining the plexus the seventh nerves give off the ileohypogastric and crural nerves. to expose the spinal cord and the roots of the spinal nerves from the ventral surface. 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. canal form the main part of the Cauda equina. cloaca. the eighth nerve between the eighth and ninth vertebrte. The spinal nerves within the vertebral canal. formed by the roots of the hinder nerves together with the filum terminale. The coccygeal or tenth spinal nerve emerges through a small hole in the side of the urostyle near its anterior end. The obliquity of the middle and posterior iii. and the ninth nerve between the ninth or sacral vertebra and the urostyle. ii. It gives branches to the bladder. Cut away with The roots of the nerves : dorsal and ventral.86 THE NERVOUS SYSTEM OF THE FROG. supplying the muscles and skin of the abdomen and thigh. which form conspicuous objects : . etc. Beyond the plexus is the large sciatic nerve. oviducts.

close to the dorsal surface and alongside the vertebral column further back they are in close relation with the dorsal aorta. renal. The olfactory nerve. and fifth ganglia. : ii. Each sympathetic trunk receives a branch from each of the spinal nerves of its side. unlike the others. the chief ones being the following II. hsemorrhoidal. and is distributed to the membrane lining the nasal cavity. From the sympathetic ganglia nerves are given off to the bloodvessels and viscera. The cardiac plexus from the first is formed by nerves arising sympathetic ganglion the plexus is a meshwork of nerves on the auricles. and bladder respectively. : : i. large intestine. 87 Remove these patches carefulhj to see the ganglia. The Sympathetic Nervous System. The stomach solar plexus lies on the dorsal surface of the the nerves are derived mainly from the third. on either side of the vertebral column. (See Fgures 13 and 15). but is said to vary from two up to as many as twelve. fourth. arises from the anterior end and outer side of the olfactory lobe. and around the great vessels at their openings into the heart. The Cranial Nerves. connected by branches with the several spinal nerves. and vesical plexuses with the liver. This consists of a longitudinal nervous band on each side of the body. kidney. is connected with the sympathetic by more than one branch: the actual number of these branches is not constant. . reproductive organs. : Hepatic. in front.THE SYMPATHETIC NERVOUS SYSTEM. The coccygeal or tenth spinal nerve. the special nerve of smell. To dissect the cranial nerves expose the brain by removing the roof of the skull as already described. The two main sympathetic trunks lie. also exist in connection III. and then follow the special instructions given in the case of the more important nerves. which are numbered in order from before backwards. 1. alongside which they run. and at the junction of each of these branches with the main trunk there is a ganglionic enlargement. There are ten pairs of cranial nerves in the frog. genital.

2. Vila Fig. glossoi^haryngeal Vllb. right auricle Vb. arises from the side below the optic lobe. maxillary branch of trigemiVa. hypoglossal nerve a. second spinal nerve s. partially crosses over at the optic chiasma on the under surface of the brain.-^^^tX. The trigeminal. and then runs outwards to the eyeball. To see the course and distribution of the olfactory nerve. pidmonary branch of Xa. gastric pneumogastric Xc. The optic nerve. has been fully seen in previous 3. mandibular branch of trigeminal Vila. ]ialatine branch of nal IX. the nerve of sight. transverse i3rocess of second vertebra 3. Eustachian tube A. lung e. hyoidean branch of facial facial Xb. right anterior vena cava arteriosus u. squamosal t. ophthalmic branch of trigeminal V c. dissect from the dorsal surface. and pneumogastric nerves of the frog. dissected from the right side. of the brain just optic nerve The course of the dissections. 17. facial glossopharyngeal. 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. ventricle v. : : : : : : : : : : : : : : : : : : : : : : .88 THE NERVOUS SYSTEM OF THE FROG. It supplies four of the muscles XI 1 ^ e 1 3 m . laryngeal branch of pneumogastric Xd. transverse process of third vertebra. removing the roof of the anterior part of the skull.a. including the nasal bone. stomach m. cardiac branch of pneumogastric branch of i)neumogastric 2. i.c. truncus v. sinus venosus t.

The ramus ophthalmicus runs forwards through the orbit. At the end of the orbit it divides into two branches which pass through the walls of the nasal capsule. i. too small to he dissected satisfactorily in the The trigeminal frog. After a very short course it divides into the maxillary and mandibular nerves. ii. but above all the other muscles and the optic nerve. and obliquus inferior. It then passes through the skull wall immediately in front of the auditory capsule. between it and the eyeball. The fourth arising or pathetic nerve is a very slender nerve.THE CRANIAL NERVES. and divides at once into two main It arises : — branches. internus. Owing 4. and supply the skin of the fore of the eyeball anterior part of the head. The ramus maxillo . andjind the 7ierve running close alongside the skull wall. and find the nerve lying on the j^terygoid muscle and . in front of the auditory capsule and between the temporal and pterygoid muscles. viz.. and runs outwards and forwards to the skull wall just before reaching this it expands into a large swelling the Gasserian ganglion. and supplyiug the obliquus superior muscle of the eyeball. is the largest of the cranial nerves in the from the side of the anterior part of the medulla. rectus inferior. is The fourth nerve frog. to its small size the third nerve is 7iot easily made out in the frog. Trace it for- wards to the nose. lying close to its inner side. from the dorsal surface of the brain between the optic lobe and the cerebellum. To trace this branch.mandibularis runs directly outwards behind the eyeball. the rectus superior. To trace this nerve and its branches remove the squamosal bone carefully. dissect from the dorsal surface : cut away carefully with scissors the side ivall of the cranium : cut through and turn aside the rectus stq^erior. 5. between the skull wall and the eye. It passes beneath the rectus superior. 89 rectus moving the eyeball .

out- behind and wards eyeball. a. where it is very closely conIt passes through nected with the Gasserian ganglion. of the orbit a short distance from the side wall . the skull wall immediately behind and in close company with the trigeminal nerve. Follow the nerve behind the pterygoid immediately beJiind the eye. and then trace the branches outwards to their distribution. supplying the upper lip. : : 6. outwards. to the margin of the upper below the jaw. and a short way behind the pituitary body. The abducens is a very slender nerve which arises from the ventral surface of the medulla close to the median It line. and temporal muscles to the skull. the lower eyelid. which it reaches about midway along it then ends in branches which its length run along the jaw. and entering the orbit supplies the retractor bulbi and the rectus externus muscles. 7. and divides at once into its two main branches. The ramus palatinus runs forwards in the floor i. The ramus mandibularis runs parallel to and behind the ramus maxillaris as far as the outer border of the eyeball. The ramus maxillaris runs forwards and in the floor of the orbit. giving branches to the temporal and pterygoid muscles it then turns backwards. some forwards and some backwards. removing the muscles if necessary. passes either through. and downwards. the Gasserian ganglion. and passes forwards to the skull wall.90 THE NERVOUS SYSTEM OF THE FROG. and passing across the inner side of the upper jaw. is The abducens nerve the frog. supplying the lower lip and the muscles of the floor of the mouth. reaches the outer surface of the mandible just behind the it then insertion of the temporal muscle runs forwards along the outer side of the lower jaw to the chin. : /3. or in very close contact with. too small to be made out satisfactorily in The facial nerve arises from the side of the medulla immediately behind the trigeminal nerve. and other neighbouring parts.

from jaw: carefully remove the ii. The ramus mandibularis runs forwards floor of the in the mouth. The ramus hyomandibularis runs outwards and backwards round the front end of the auditory capsule. just above the level of the columella : fiiid the facial nerve rinuiing round the front end of the auditory capsule and in close contact ivith it. which is 7iot easy. and find the nerve lying on the ventral surface of the eyehall and its muscles. and immediately upon the mucous of the roof of the mouth.THE CRANIAL NERVES. Trace it backwards and forwards. and running parallel to and a short distance from the skull-wall. the outer end of the mouth. It supplies the mucous membrane of the roof of the mouth. one of which runs outwards and membrane anastomoses with the ramus maxillaris of the trigeminal nerve. may he depressor manclidulce the shoulder-girdle of one side . it then crosses over the inner end of the columella and turns downwards in the posterior wall of the Eustachian tube to the angle of the mouth. The exposure of performed thus cavity : —remove to the above nerve. a. and trace it hack over the columella and down to the angle of the columella. . 91 of the skull. Dissect this nerve the ventral surface. expose the braiyi : remove the tympanic membrane and clean Cut aivay carefully the 7'oofof the auditory capsule by a horizontal cut. It then divides into two branches. Near the anterior end of the orbit it divides into two branches. lying along the inner edge of the lower jaw and between the mylohyoid nmscle and the skin. Dissect from the ventral surface the floor under surface of of the : remove the skin from the mouth. also the and temporalis muscles : open the cranial as before. Cut aivay the lower mucous membrane of the roof of the mouth. and find the nerve running along the inner border of the mandible. while the other runs forwards to the anterior part of the roof of the mouth. giving branches to the tympanic membrane and to the articulation of the mandible. as far forward as the chin.

and is much facilitated by distending the oesophagus and pharynx with a cork or roll of pajjer. close to the middle and between the geniohyoid and hyoglossus muscles. which it enters and in which it ends. The ramus anterior runs downwards and forwards round the hinder border of the auditory capsule and beneath the depressor mandibulse muscle to join the facial nerve just after it has crossed over the columella. made for the ramus hymondihularis of show also the above branch of the glossothe facial nerve will The dissection already pharyngeal. The auditory nerve. from The exposure of the first part of the above nerve is best performed the side. arises from the the medulla immediately behind and in close side of contact with the root of the facial nerve it enters the auditory capsule and ends in the internal ear. by a root common it leaves the skull by an to it and the tenth nerve aperture immediately behind the auditory capsule. passing beneath the fourth division of the petrohyoid muscle but superficial to the others . The glossopharyngeal nerve arises from the side of the medulla behind the auditory nerve. is the two branches : 8. : i. it On reaching the floor of the mouth crosses the hypoglossal nerve obliquely. to the base of the tongue. and then runs forwards in a peculiarly sinuous course. the nerve of hearing. /3. ii. Its course along the floor of the mouth to the tongue should be dissected from the ventral surface. it runs just behind and parallel to the anterior cornu of the hyoid. It supplies the petrohyoid muscle. The ramus hyoideus the posterior and larger it runs forwards in the of floor of the mouth along the anterior cornu of the hyoid. .92 THE NERVOUS SYSTEM OF THE FROG. and divides behind the capsule into two branches. : 9. lying dorsal to it. The ramus posterior runs downwards and for- Avards to the ventral wall of the pharynx. and the mucous membrane of the pharynx line and tongue. supplying its muscles.

anterior fibres of the obliquus internus muscle. : ii. of each side extends forwards in The main sympathetic trunk front of the first ganglion. in which they end. 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. and immediately outside the skull presents a ganglionio enlargement it gives off dorsal branches to the muscles of the back. to see them 'properly. to the middle line. The rami pulmonales follow the course of the pulmonary artery to the lung. The dorsal portions of the several branches of the pneumogastric nerve are best exposed from the side. The ramus cardiacus passes dorsal to the pulmonary artery to the interauricular septum of the heart. which are as follows i. IV. and end in the walls of the stomach. The rami gastrici. usually two in number.THE CRANIAL NERVES. run through the partial diaphragm formed by the iii. 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. the shoulder girdle and fore limb must be removed and the cesophagus well distended : the terminal branches must be dissected from the ventral surface. iv. and to the sinus venosus. 93 The pneumogastric noticed. 10.. It leaves the skull by the same aperture as the ninth nerve. and then runs backwards and downwards round the side wall of the pharynx. The Cranial Portion of the Sympathetic Nervous System. in arises. dorsal to the artery. in which it ends. running along the hinder border of the fourth division of the petrohyoid or vagus nerve common with : muscle : behind this muscle it : divides into its main branches. where it ends in the larynx. as already the glossopharyngeal.

ii. iii. C. while the nerve fibres serve A local to convey these impulses from one part to another. The nerve cells are branching nucleated cells. called a ganglion. the essential part of the nerve fibre it is clearly : . and only in certain places. and the primitive sheath forms constrictions touching the axis cylinder. seen with difficulty. medullated and non- Medullated nerve fibres form almost the whole of the cranial and spinal nerves. Nervous matter consists histologically of elements of two nerve cells and nerve fibres. : ii. Take a small piece of the sciatic or some other nerve from a freshly killed frog : spread it out and tease it in a drop of normal salt solution: examine with low and high p)owers: note i. The nerve fibres : unbranched. a. fibre note or connective tissue binding the nerve fibres into bundles. very delicate or sheath of Schwann: a external investment. is . The medullary sheath quence of intervals its by is stained darkly in consefatty nature it is interrupted at the nodes of Ranvier. Histology of Nerves. and gives the nerve fibre its double contour.accumulation of nerve cells 1. kinds. The medullary sheath a thick fatty layer within the primitive sheath . or " nerves. Nerve Fibres are medullated. it is highly refractive. The nerve cells are the centres whence impulses originate. The axis cylinder is the central cylindrical rod. and a large part of the brain and spinal cord.94 THE NERVOUS SYSTEM OF THE FROG. Tease in glycerine a small jnece of nerve that has been treated with osmic acid : examine with the high power a single nerve fibre: note the following p)oints. of two kinds. The primitive sheath. i." In each nerve iii. : iv. connected by their processes with one another and with the nerve fibres. The perineurium. The nodes of Ranvier are spots where the medullary sheath is absent.

Nerve cells : Tease in glycerine a small fragmerit of the : ventral cornu of the spinal cord of the ox (lumbar region) and examine with low and high jjowers : note i. The primitive sheath. Non-medullated nerve fibres occur chiefly in the sympathey branch and anastomose . iii. and examine with low and high powers : 3. iv. in which are imbedded numerous nerve cells. thetic nerves 2. : or sheath of Schwann. tissue binding the several parts Structure of the Spinal Cord. or " anterior" fissure is a broad and shallow median cleft there is in the frog no : ii. . Take one of the prepared transverse sections of spinal cord of frog . and is chiefly composed of medullated nerve fibres. The section is bilaterally symmetrical. iv. The nerve fibres. cover The nerve cells : large nucleated cells with many branching processes. and oval in shape. 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. and is composed of a dense network of nonmedullated nerve fibres. HISTOLOGY OF NERVES. The fine connective together. The grey matter forms the central part of the cord. With the low power. visible at the 95 nodes. iii. and they have no medullary sheath. i. b. and is much less deeply stained than the medullary sheath. mount in balsam. The ventral distinct dorsal fissure. the transverse diameter considerably exceeding the vertical. note the following p)oints. ii. ii. The white matter forms the outer part of the cord. a. The axis cylinder.: . Nuclei are seen projecting into the medullary sheath about midway between the nodes.

it consists of a number of very slender bands of nerve fibres. ii. THE NERVOUS SYSTEM OF THE FROG. or "posterior" root is /3. to pass through their points of origin. lined by a ciliated epithelium." into which the grey matter is produced on either side.96 V. they enter from the pia mater. penetrating and supporting all parts of the cord. and continuous at the surface with the pia mater. . The cornua are the processes. The central canal The nerve a. The neuroglia a delicate network of connective tissue fibres and cells. The bloodvessels of the cord are small and numerous The central canal is V. it is a single thick band of nerve fibres. With the high power. The dorsal connected with the dorsal cornu of the grey matter. i."' and dorsal or "posterior. is iii. of the cord lies in the median ventral than the dorsal surface. The ganglion cells are large nucleated branched : they cells lying in groups in the grey matter and most numerous in the ventral are largest cornua. The pia mater is the delicate connective membrane closely ensheathing the cord. vi. tissue iv. nearer the vii. plane. The ventral or "anterior" root is connected with the ventral cornu of the grey matter . ventral or "anterior. roots are only seen if the section happens. b.

ii. is sclerotic formed of hyaline connective tissue. side. a firm solid transparent body. from a freslihj killed frog : snip off of the eyeball : note the following points. uitli The The shape. Place the eye under water and divide it ivith scissors into two cut passing through the middle of the cornea and through the sclerotic close to the optic nerve. is iv. The Eye of the Frog. 2. just iris and attached to its outer margin more convex on its inner than on its outer is : H . halves by a i.: 97 CHAPTER VII. the firm outer wall of the eyeball. Remove the eye scissors the muscles i. A. The pupil the aperture surrounded by the iris. THE EYE AND EAR. The optic nerve is seen piercing the sclerotic to enter the eyeball on its inner side. The ' iris is a pigmented ring placed behind the cornea and seen through it: it acts as a diaphragm. vi. and dense white The cornea it is side of the eye the transparent patch on the outer through which the light enters continuous at its margin with the sclerotic. 1. The lens it is behind the surface. cartilage. limiting the amount of light that enters the eye. is V. iii. The eyeball is flattened on its outer more convex on the inner or deeper side. so as to lay open completely the interior of the eye: note the following points. w^hich serves to admit the light to the interior of the eye.

a watery fluid.98 ii. white. The pupil The is the central. The conjunctiva ing the sclerotic. muscles of the eyeball. : iii. the frog. 1. except at the point where the optic nerve enters. . The choroid the iris. The eyeball is more spherical than in ii. transparent. and the fat which sur: note the following points. the choroid. its margin with the than the sclerotic. 0^ the rounds the optic nerve i. iv. THE EYE AND EAR. is sclerotic. tough. sixth of the The cornea. The sclerotic covers about five-sixths of the eyeball it is iii. the black pigmented layer lining and continuous in front with the V. vi. oval or dumb-bell shaped aperture surrounded by the iris. The iris is the oval pigmented ring seen through the cornea. Dissect The Eye of the Sheep or Ox. vii. is sclerotic : and continuous at it is more convex iv. The retina transparent membrane readily detached from It lining the eyeball. The posterior chamher of the eye is the large space behind the lens: it is filled by the vitreous humour. The anterior chamljer of the eye is the space between the cornea and the lens it is small and contains the aqueous humour. which covers the outer eyeball. a gelatinous body. and opaque. a delicate epithelial layer. is a delicate is B. coverfront of the cornea and part of the V. The shape. optic nerve is fibres piercing the sclerotic at the a thick white bundle of nerve back of the eye. is circular.

. Make four radial cuts. 99 Cut its from not to all round the cornea. 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. optic nerve P. : : : : : : : 3. choroid: CP. yellow spot. with stout scissors. cornea: Ch. equidistant from one another. posterior chamber R. 2.THE EYE. central artery of retina : B. : • : Fig. 11 The aqueous humour the transparent watery fluid fining the anterior chamber of the eye. or the lens will he driven out instantly: note: i. anterior chamber Ar. A. lens O. hyaloid membrane' enclosing the vitreous humour iris L. Pass the handle of a scalpel under the cut edge of the cornea. blind spot C. suspensory. ciliary processes : H. and escaping when the cornea is removed The lens: of entrance of the optic nerve. through the margin of the cornea and betiveen it a7id the iris. A diagrammatic section through the human eye passinothrough the centres of the cornea and lens. sclerotic SL. and through the yellow spot and point I. retina S. about J inch junction with the sclerotic : remove the cornea : take care squeeze the eye.ligament Y. 18.

ciliary vessels pierce the sclerotic to convey blood to and from the choroid. it from behind : note ii.100 the sclerotic^ THE EYE AND EAR. and pin them to the dissectijig board so as to fix the eye with the iris upwards : note the following points. iv. so couple of radial incisions^ about ^ inch apart. removal of the iii. the anterior half of the eye over. and Carefully peel off back towards the ojJtic nerve. The choroid The the dense black coat exposed by the sclerotic. the iris ciliary muscle. horizontal incision at half. . The The uvea ciliary processes. iii. the eye into tivo halves by the following parts. extend the cuts which the sclerotic is now divided from the the four flaps into underlying black choroid coat : turn them doivii. is ii. Make a and i. The ora serrata of is the indented anterior boundary the part of the retina sensitive to light: in front of this the retina becomes extremely thin. i. With a large pair its of scissors cut equator. taking care not to injure the deeper 2^ arts . \ . through and turn back the portion of the iris as to expose its between the hinder surface. Turn the following points. 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. Turn over the anterior or outer and examine i. so that its outer or corneal surface is uppermost : cut away the iris completely : note 6. but really extends forwards as far as the free edge of the iris. The ciliary nerves are seen passing through the sclerotic to the choroid while the flaps are being turned down. The ciliary processes are a series of densely pigmented and close set radial folds on the hinder surface of the outer margin of the iris . tivo cuts. lens. which is extremely vascular. is the layer of dense black pigment at the back of the iris and ciliary processes. they fit into corresponding folds in the ligament which a surrounds and supports the 5.

THE EYE. of the lens as a whole. Remove i. the lens of the retina and choroid should be shape. note its behind than in front : harden few minutes in water. The serrated edges of many of the cells. piece of fresh choriod on a examine with low and high powers. the vitreous humour from is the posterior half of the eye: note the following points. The Lens. a delicate pulpy membrane between humour and the choroid. cells are irregularly ii. i. The Choroid. The laminated character The elongated composed. The cut edges recognised. ii. 1. The pigment clear nuclei. iii. hardened by boiling . 8. examine with low and high powers : note : — i. epithelial cells of which the lens is iii. or zonule of the outer margin of the capsule of the it is marked with radiating folds into lens which the ciliary processes fit. branched. The suspensory ligament of the is lens. iii. 101 is The capsule of the lens membrane holding the Zinn. The retinal vessels enter with the optic nerve. Remove from its capsule it . Histology of the Eye. with 2. The retina the vitreous ii. . ii. Spread a small normal i. radiate from the blind spot. m. : an elastic transparent lens in its place. slide in salt solution : The Choroid is a network of bloodvessels thickly invested by pigment cells. . Tease in glycerine a small piece of lens. The blind spot nerve : is the point of entrance of the optic the retina adheres firmly to this spot. and C.ore convex or by boiling for a 7. though it can be readily separated from the choroid at all other parts. with spirit.

pigment cell PC. pigment of choroid R. nerve cell IL. 19.102 3. rod : RF. the cartilaginous sclerotic. red blood corpuscle C. THE EYE AND EAR. ill balsam one of the prepared sections of the j^osterior part of the frog's eye : examine with low aiid high powers. BC. thickness of the retina. outer molecular layer ON. radial or Miiller's fibre S. the choroid. inner limiting membrane IM. outer limiting membrane OM. inner molecular layer IN. Mount Fig. Vertical section through the posterior wall of the eye of a the section passes through the sclerotic. cone G. and the entire x 3(X). layer of nerve fibres OL. frog : : : : : : : : : : : : : . The Retina. inner nuclear layer: NF. outer nuclear layer P.

are a layer of columnar bodies placed vertically to the surface the rods are : ii.THE EYE. iv. or Miiller's fibres. sending processes between the rods and cones. It vii. The The layer of nucleated nerve cells is a single layer of large ix. branched cells. xi. with much pigment. The rods and cones far more numerous. The choroid is a vascular plexus. i. A layer of pigment cells. is a very thin layer. iii. formed by the branches X. finely granular or reticular in V. is layer of nerve fibres of the optic nerve. . layer. finely granular viii. The inner^ nuclear layer is thick and well stained. surrounded by thin layers of protoplasm. The outer molecular layer not stained structure. a. The radial fibres. well stained. commence with expanded ends in the layer of nerve fibres and outwards they can easily be traced through the inner molecular layer. and very closely connected with the rods and cones. The inner limiting membrane. not stained. 103 The sclerotic consists chiefly of hyaline cartilage. The outer nuclear layer is a moderately thick layer. C. consists of several layers of large nuclei. They consist of connective tissue and serve to strengthen and stretch : support the retina. b. The inner molecular layer is a thick. and much larger than the cones each consists of an inner and an outer : segment. The retina is composed of the following layers from without inwards. : it is vi. The outer limiting membrane as is seen in sections an exceedingly thin line separating the rods and cones from the outer nuclear layer. produced into radial processes.

ii. removed from the periotic cartilage and drawn from the outer surface. divisions. a. D. is The utriculus The sacculus the upper and larger division. and firmly fused with the hinder part of the cranium. in the following manner. The right internal ear of the frog. is the inferior and smaller division from it arise three small saccular dilatations. cut off the head with stout scissors. Then stain ivith horax carmine and imbed in paraffin. the utriculus. 2. solution of nitric acid. them. its ampulla s. which will take from a few hours to 3 or j^ days or more according to the strength of and acid employed. The frog's auditory organ is too small to dissect and is best studied by making transverse sections head. the sacculus u. its ampulla : h. The vestibule : is a membranous sac lying in the cavity of the periotic capsule. and cut into transverse sections with a microtome. the posterior vertical canal r. is The periotic capsule consists mainly of cartilage. remove the head transfer to from the decalcifying solution and weak alcohol and thence to strong alcohol. decalcify it hy placing in a 5 to 10 per cent. When the hones are thoroughly soft.104 THE EYE AND EAR. its ampulla p. Kill a frog ivitli of the entire chloroform . 20. or in a mixture of chromic acid with a few drops of nitric acid. showing the following points. the endolymph it is partly divided by a constriction into two main i. satisfactorily. and filled with a watery fluid. the horizontal canal i. : : : : : : . Fig. Mount the sections in series . The Ear of the Frog. the anterior vertical semicircular canal h. supposed to represent the cochlea of higher animals. examine and draw 1.

iii. The columella is a rod of bone and cartilage. See Chapter IX. a dilatation or ampulla. The horizontal or external canal has its ampulla at the anterior end. 11 p. i. The anterior vertical canal has anterior end. The Eustachian passage and tympanic cavity formed gill are connection with the hyomandibular cleft of the tadpole. into are three tubular offsets of the which they open at both ends. 5. The tympanic membrane closes the tympanic cavity on its outer side. 58). The essential organs the vestibule and its offsets. and are placed in planes at right angles to one another each has at one end. 3.e. which is deeply placed in the side of the head the communication with the surface is brought about by the accessory apparatus.— THE EAR. The posterior vertical canal has its ampulla at its outer end. and to the ampullae of the semicircular canals. in ii. of hearing i. It serves to communicate the auditory vibrations of the tympanic membrane to the vestibule. the outer end of which is attached to the tympanic membrane. while its inner end joins the posterior end of the anterior vertical canal to open into the vestibule by a common orifice. iii. 4. The auditory nerve leaves the cranial cavity through a hole in the inner w^all of the periotic capsule.. They lie in canals in the periotic cartilage. . The accessory auditory apparatus. its ampulla at its ii. close to its opening into the vestibule. and the auditory nerve are enclosed in the periotic cartilage. which consists of the following parts — : (Fig. 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. and divides into branches distributed to the sacculus and its diverticula. 105 The semicircular canals vestibule. : i.

The Cloaca. whence they escape by the ureter. Fig. 2. hack under water : open the body cavity from turn aside the alimentary canal and liver. 1. Within the testes are developed the male elements or spermatozoa. p. The testes are a pair of yellow ovoid bodies about half an inch long. and pin them out right and left. (Cf. . The Male Frog. lying on the ventral surface of the kidneys. the (Fig. the ventral surface. — — : iii. The vasa efPerentia are a number usually 10 to 12 of slender ducts. The vesicula seminalis is a large pouch-like dilatation on the outer side of the vas deferens. 18. p. Lay the frofi on its back: tvith a stout scalpel split the pelvic symphysis in the median plane : gently separate the tivo halves. 3.106 CHAPTER VIIT. 3. so as to expose the cloaca from. and then iv. The vas deferens backwards to the or ureter runs along the outer side of the posterior part of the kidney.) Pin frog on its : the ventral surface i. cloaca. The Reproductive Organs. 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. A. just behind the kidney and before reaching the cloaca. 18). which acts as vas deferens. essential ii. THE REPRODUCTIVE ORGANS AND THE CLOACA OF THE FROG.

along one side: wash out its contents and examine the opening into the bladder. Their shape and size vary much at different seasons of the year. The Eeproductive organs. The Female Frog. overhung by a slight valvular pro jection of the mucous membrane of the cloaca. THE REPRODUCTIVE ORGANS. like small shot these are ova in various stages of development r the smaller and younger ones are white . is intestine. B. its two lobes communicating freely with each other. The ovaries of the are a pair of black masses lying in folds peritoneum ventral to the kidneys. Innate the bladder with a hlow-pipe inserted through the cloacal aperture: pass a seeker up the cloaca to determine the exact position Cut up the cloaca of the opening from the bladder to the cloaca. i. into the terminal portion of the large which the ureters and the bladder ii. iv. and white or yellowish in the other. The bladder is a thin-walled bilobed muscular sac. Dissect as in the male. It is invested by peritoneum and attached to the sides of the body by special peritoneal folds. 107 The cloaca open. The openings of the two ureters are close together on the apices of two small papillae. Remove 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. 1. iii. The ureter or vas deferens is continued behind the vesicula seminalis as a very short tube. in very - the same position as the testes in the male. i. Each ovary much . opening into the dorsal wall of the cloaca almost exactly opposite the opening of the bladder on the ventral surface. lying on the ventral surface of the large intestine and cloaca. the larger and more mature ones black in one half. On their surfaces are numerous rounded projections..

apertures. twisted tubes. increa sing Their in size. They commence with thick gelatinous walls. The Cloaca. ii. consists of a couple of folds united along their ventral edges . i. iii. partition. The oviducts open separately into the dorsal wall of the cloaca. the female has genital walls. placed close together. The bladder like that of the male. hinder ends are greatly dilated. close to the outer side of the roots of the lungs . iv. by two wide apertures separated by a narrow median The ureters open by two small just behind the oviducts.108 REPBODUCTIVE ORGANS AND CLOACA. and becoming much convoluted. and run back. Dissect as in the male. ducts distinct from the ureters. is ii. just opposite the bladder. The cloaca is very similar to that of the male. 2. except that the urinary and genital products are discharged into it by separate ducts. but have thinner Unlike the male. The oviducts are a pair of white. into the dorsal wall of the cloaca . with open mouths at the extreme front end of the body cavity. the space between the folds is divided by partitions into about fifteen pouches.

Such spawn consists of a transparent gelatinous mass. and stick together to form the bulky masses we call frog's spawn. which may last several days. they are invested by thin coatings of an albuminous substance. I. in which are embedded the eggs these latter appear as small round bodies. THE DEVELOPMENT OF THE FROG. and as the eggs pass out from the cloaca of the female they are fertilised by spermatozoa discharged over them by the male. Each egg is at first spherical. the male frog clasps the female firmly. The frog's eggs. laid in this way and fertilised by the spermatozoa shed over them by the male. and the whites or albuminous investments of the yolks to the gelatinous matrix of the spawn. the externaL : : : . General Account. each presenting a black half and a white half. a mass would be produced similar to frog's spawn the yellow yolks corresponding to the frog's eggs. The whole animal becomes fish-like in appearance. and not from the white of a hen's egg. begin to develop at once. formed by the swollen albuminous matter. usually during March or the early part of April. in diameter. By the tenth day it is divided by slight constrictions into head. And just as the chicken is formed from the yolk. The frog's eggs are laid in water. During the act of oviposition.109 CHAPTER IX. but in about a week becomes ovoid in shape. and tail. The eggs. which swell up very greatly in the water. If a number of hen's eggs were broken into a basin. which are very numerous. the tail growing rapidly two pairs of branching tufts. and then rapidly increases in length. so also is the frog developed from the egg and not from the gelatinous investment. body. embracing her with his arms . care being taken not to rupture the yolks. are small spherical bodies about 1'75 mm.

.110 DEVELOPMENT OF THE FROG.

and forms a powerful swimming organ. the external gills at the same time decreasing in size and assuming a shrivelled appearance. 21. Fig. the gill clefts. 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. one on A anus . and the tadpole now feeds eagerly on confervse and other vegetable matter. a fold of skin. During this time the tadpole has been feeding freely.DEVELOPMENT OF THE FROG gills. and through which the water taken in at the mouth for respiration. The margins of these slits become folded. the is developed even before the mouth. so as to enclose them in a gill chamber. is small and but little used. the operculum. Very shortly afterwards rudiments of the hind-limbs can be seen as a pair of small papillae at the root of the tail. is broad and round. While the internal gills are developing.e. shortly after hatching. At the time of hatching the tadpole has no mouth. On the left side a spout-like opening remains which communicates with the gill chambers of both sides. and passed through the gill slits. the tail is much larger than before. few days after hatching the mouth appears. s. About the time of appearance of the mouth i. the liver and pancreas are formed. objects in the water. bordered by a pair of horny jaws.. and fringed with fleshy lips provided with horny papillae. while the sucker on the under surface of the head. and form the internal gills . and swim freely in the water. grow out from the sides 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. leading from the pharynx to the exterior. 111 by a third pair. becoming tubular and convoluted. and has increased greatly in size. The alimentary canal which has hitherto been wide and short. now rapidly increases in length. A horse-shoe shaped sucker is present on the under surface of the head. appear on each side of the neck. The body. by which the tadpole is enabled to attach itself to weeds or other followed shortly of the neck. and is dependent for food on granules of food yolk which are contained in large numbers in the egg. appears on each side of the head in front of the gills. and grows backwards over these. though still present. makes its escape to the exterior. and at the expense of which all the earlier processes of development are effected. a series of four slit-like openings.

112

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-hmbs 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 fishlike condition in which it has hitherto remained to the purely The tadpole air-breathing stage characteristic of the adult. a casting, or ecdysis, of the outer layer of the ceases to feed 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 the opercular fold, in which it leaves a ragged hole. abdomen shrinks ; the stomach and liver enlarge, but the intestine becomes considerably shorter than before, and of smaller the animal, previously a vegetable feeder, now diameter becomes carnivorovis. 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, ii, 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.
tadpole is really a fish ; not merely in its habits, but in its mode of breathing, in the arrangement of its heart and bloodvessels, 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 during its own development to repeat, or

A

;

FOKMATION OF THE EGG.
recapitulate, in a

113

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

own development.
II.

this history in the course of their

Detailed Account.

A. Formation of the Egg.
early stages in thp formation of the Qgg cannot be seen 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 ajDpear 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. Each genital ridge is at first due merely to slight modification

The

m the adult frog,

m the shape
flattened,

of the peritoneal epithelial cells which, elsewhere

become here cubical or slightly columnar. The ridges soon become more prominent, especially at their anterior ends j then^ growth being due partly to the epithelial cells increasingby 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 the male the testis, is developed ; while the anterior third undergoes degenerative changes, and becomes converted into the fat body.

m

early stage certain of the epiconspicuous by their around these larger cells, or primitive ova, as they are called, the smaller epithelial cells become arranged so as to form capsules or follicles, the follicles, with their contained primitive ova, forming small knoblike projections on the surface of the genital ridge. New primitive ova arise either directly from the surface epitheliiun, or by division of the already existing ones.
thelial cells of the genital ridge become larger size and more spherical shape ; and
I

The Primitive Ova.

At an

;

114

DEVELOPMENT OF THE FROG.

to this time there has been no and female, the processes described distinction between male

The Permanent Ova.

Up

Sexual differentiation appears occurring in all tadpoles alike. 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 in some cases each primitive ovum is the primitive ova 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.

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 suba number of small sharply-defined yellowish granules of stance 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

A permanent ovum

consists of
fluid

an elastic capsule or nuclear membrane, filled with 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 mode of origin of the vitelline membrane is the follicle. 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 The pigment is contained, and to one half or hemisphere. 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 This process of maturation concerns are ready to be fertilised. 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 Part of the nuclear fluid has a blackberry-like appearance. exudes through the nuclear membrane into the substance of the •egg ; 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 contraction 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 In the first, or thick-walled, part of the the roots of the lungs. oviduct the eggs acquire gelatinous investments, secreted by The terminal, or hinder, part of the oviduct glands in its walls. 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 tinally 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 hemisphere. 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.

About the time the egg is laid, but 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.
before
it is

The First Polar Body.
fertilised,

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 completion of the act of fertilisation.
C. Fertilisation

of the Egg.
strictly speaking, fusion of

Fertilisation, or impregnation, consists in fusion of the sper-

matozoon with the egg or, more 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
is

single spermatozoon is sufficient to fertilise an egg, and it 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.
:

about two and a half hours after fertilisation has commenced. to form the segmentation nucleus. which move away from each other.. that is known to result in both animal and vegetable kingdoms from cross fertilisation. from combining the energies of two distinct individuals in the act of reproduction. i. The -of earliest stages of development consist in repeated division the egg. The male and female prowhich are at first some little distance apart. whereby it becomes converted from the unicellular condition. : tage. fuse together. The female pronucleus may be regarded as an imperfect nucleus. To these early processes of development the name Segmentation is given. and after having increased considerably in size. the yoke granules at the same time showing a tendency to arrange themselves in lines radiating outwards from the ends of the spindle. divides the egg •spherical . come into close contact. which is permanent only in the lowest animals. and the upshot of the process of fertilisation is the completion of this nucleus . rapidly approach each other. Segmentation of the Egg. bisecting the egg. at first as a small pit and then as a groove. 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 nucleus now divides into two halves.e. the yolk granules tend to aggregate themselves around the two nuclei. replacing the part of the egg-nucleus which has been lost as the polar bodies. rapidly deepening. the nucleus of the spermatozoon. Very shortly after the completion of the act of fertilisation and formation of the segmentation nucleus this latter loses its form and becomes spindle-shaped. The segmentation nucleus is a large spherical vesicle imbedded in finely granular protoplasm. and. as regards vigour of offspring. The further explanation of the sexual process is probably to be found in the great advannuclei. D. At the upper or black pole of the egg a depression now appears. and the female pronucleus is the only part of the original egg nucleus still remaining. and a thin vertical plate of finely granular substance is left. which soon extends all round. to the multicellular state characteristic of all higher animals. 117 By this time the second polar body has been formed and extruded. or male pronucleus.FERTILISATION OF THE EGG.

and on its completion the egg consists of four precisely similar segments or cells. : : i 2 4 s Segmentation of the frog's egg. 22.) The numbers above the figures indicate the number of segments at The dotted lines mark the positions of the clefts the several stages. round which the cells are grouped during the later phases of segmentation this segmentation cavity. as it it is from the first is called. eight smaller upper ones. lying nearer the upper than the lower pole it divides each of the four cells into two. Two more vertical clefts next appear simultaneously at the upper pole. increases considerably in size : : . preceded by division of their nuclei. on the' completion of the third cleft. The third cleft is horizontal. i. giving sixteen cells in all. giving thirty-two in all.. At the stage when only eight cells are present. but in a plane at right angles to the first one . as from the first. Fig. Each of the two nuclei soon divides again into two. and eight larger lower ones. (From Haddon. Two more horizontal clefts then appear. that will next appear. midway between the two primary clefts. a small cavity appears in the centre of the egg. and a second cleft is formed in the same manner as before it also is vertical. an upper smaller and a lower larger one. From this stage segmentation proceeds in a less regular manner. but not equatorial. the upper and smaller cells dividing more rapidly than By means of radial and concentric the lower and larger ones.e. after Ecker.118 into DEVELOPMENT OF THE FBOG. the number of cells is rapidly increased. which again double the number of segments. each with a nucleus. two completely separate halves along a plane corresponding with the vertical plate mentioned above. division of the cells being in all cases. and extending downwards divide first the smaller and then the larger cells. clefts.

SEGMENTATION OF THE EGG. being themselves inert. while the cells of the lower half are almost colourless. occurring round the equator of the egg at the junction of its upper and lower halves. At the close of segmentation the egg has the structure shown in section in Fig. Vertical section through a frog's egg at the close of segmenX 28. 23. E. 23. a ring of cells more or less intermediate in size. The distinction between upper and lower cells is however not an absolute one. and depth of pigmentation. SC. These granules of food-yolk are more The process of . and It is a hollow ball with its walls regularly arranged. fairly uniform in size. 119 is situated nearer the upper than the lower pole of the egg. and of very unequal thickness. E Fig. The cells of the upper half are small. shape. lower layer or yolk cells. tation. epiblast : : segmentation is as mentioned above. must hinder the activity of the protoplasm in which they are imbedded. which. composed of three or four layers of cells. while those of the lower half are larger. and filled with fluid. 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. owing to the segmentation cavity lying in the upper half of the egg. simply one of cell-division . The superficial cells of the upper half are deeply pigmented. segmentation cavity Y. and more irregular both in shape and size.

inner or blastopore nervous layer of epiblast: SO. If it be true that an animal.e.: 120 DEVELOPMENT OF THE FROG. in E. and this unequal distribution of food-yolk is the direct cause of the unequal segmentation of the egg. as indicating that the higher animals are descended from forms which. outer or epidermic layer of epiblast E N. B. E w Longitudinal vertical section of a frog embryo. yolk cells.e. ^ E E. 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. segmentation cavity Y.. the most remote ancestors. mencing invagination. : . those of the upper half of the : Fig. during its own development repeats its ancestral history. On this view the imicellular condition of the e^g is of great interest as indicating a similar unicellular condition '. like the Protozoa nowadays.. abundant in the lower than the upper half of the egg. The Germinal Layers. climbs up its own genealogical tree. some very remote ancestor i. i. then the earliest phases of this development must represent the earliest. At the close of segmentation we have seen that the eg^ consists of cells of two kinds firstly. 24. remained throughout their lives single cells. 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. showing com: : x 28. The purpose of food-yolk is to afford a supply of nutriment at the expense of which the earlier developmental processes may be accomplished. such as a frog.

which is present in such quantity as to render them comparatively inert. which are smaller. yolk plug. less strongly pigmented. tilling up the blastopore. : : : : : tinuous and deeply pigmented layer . mesenteron H. but much distended by food-yolk. less regular. invaginate hypoblast chord SC. and in a few hours the pigmented epiblast cells have covered the whole of the egg with the exception of a . 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: : : Fig. 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. more regularly arranged. and at the close of segmentIn the ation these two areas are approximately equal in extent. while the deeper epiblast cells are more spherical. succeeding stages the black area increases rapidly at the expense of the white area. N. segmentation cavity YP. 25. and loosely arranged in a layer two or more cells deep. stage in the formation of the mesenteron. and almost free from pigment. 121 egg.FORMATION OF THE GERMINAL LAYERS. and comparatively free from food-yolk secondly those of the lower half of the egg which are larger. pigmented. mesoblast M N. Longitudinal vertical section through a frog embryo at a later x 30. notoM.

: . YP. It is at first short and ver}' shallow. yolk plug. 26. is formed by splitting apart of the yolk cells. nervous layer of epiblast M. The circular plug of yolk cells filling up the blastopore is spoken of as the : .122 DEVELOPMENT OF THE FROG. Figs. Fig. it is effected by the addition of cells cut out from the superficial layer of yolk-cells. 25. all round its margin . 24 and 25 . mesenteron : N. Fig. Longitudinal vertical section through a frog embryo showing the completion of the mesenteron. and not by the epiblast cells overlapping or spreading over the yolk-cells. small circular patch where alone the white yolk-cells come to This extension of the epiblast occurs the surface. 25. mesoblast H. but it rapidly extends further and further into the egg. the mesenteron. cavity. round which the epiblast turns inwards into the interior of the egg. invaginate hypoblast M N. : : : : : The circular aperture in the epiblast where the yolk-cells still come to the surface is called the blastopore Fig. MN. The Mesenteron. blastopore E E. epidermic layer of epiblast E N. and from its first appearance communicates with the exterior at the blastopore. notochord. It is situated at what will become the posterior end of the embryo and it is bordered by a distinct rim or lip. During this time a narrow slit-like cavity This has appeared in the interior of the egg Fig. B. and by depression of its lower w^all or floor becomes widened out into a space of considerable size. 26. B.

{Cf. mesoblast. definite layer of cells first at The Notochord. which gives rise to the greater part of the length aUmentary canal of the embryo. The Mesoblast. except at its hinder end. 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. is soon established. 25 and 26 N). so as to leave between them a narrow space. where the space between the epiblast and hypoblast is occupied by the notochord. From one or other of the three germinal layers. and is for some time the only skeleton which it possesses. and hypoblast. It is Between the epiblast and hypoblast a third cells.long the mid-dorsal line. formed by differentiation of the surface hypoblast and yolk cells as a separate layer. Fig. The mesenteron still communicates with the exterior through the blastopore. of the 123 MN. two layers. It extends all round the embryo except a. where it remains for some time in continuity with both hypoblast and epiblast at the lip of the blastopore.) In many specimens the mesobsast cells are from the first arranged in cells of The parallel layers or sheets. lying immediately beneath the epiblast. and is formed of a spoken of as the hypoblast: the floor is very thick. 26. but the permanent mouth and anus are not yet formed. incomplete in front. This is the notochord (Figs. or intermediate layer of the mesoblast. epiblast. The roof of the mesenteron is thin. which serves to slightly stiffen the back of the embryo. The epiblast. Fig. It very early splits off from the roof of the mesenteron.— FORMATION OF THE GERMINAL LAYERS. every part of the embryo is formed. or outer layer. Fate of the germinal layers. During the process of formation of the mesenteron by ingrowth of the hypoblast cells the segmentation cavity gets pushed out of place and ultimately obliterated. opposite the segmentation cavity. It is. for a time. which later on becomes the body cavity or coelom. to the nervous — . 27. but soon grows in from the sides so as to fill ujd the deficiency. but quite distinct from it. the mesoblast become early arranged in two which separate slightly from each other. and is formed by the mass of the yolk-cells. and to the various glandular and other structures derived from the epidermis. gives rise to the epidermis covering the body generally.

mation of the neural canal. somatopleuric layer of mesoblast MH. Transverse section through a frog embryo during the for. to the retina the other sensory organs . ccelom : : epiblast: H. splanchnopleuric layer of mesoblast: MN. The nervous system is a suitable one to commence with. It is convenient from the point we have now reached to deal with the several systems one by one. i. the bile ducts. gives rise to the epithelium lining the alimentary canal and its various diverticula. yolk cells. . 27. pancreatic ducts. the lungs. the bladder. as it appears at a very early N. The hypoblast. mesenteron NF. . blood-vessels and the urinary and reproductive organs. neural fold NG. From the mesoblast. mesoblast: ME. to the olfactory and system. tissue. EN. both central and peripheral and lens of the eye. stomach and intestine. and to auditory epithelium.e. the connective . IM'r notochord : : : . to the epithelial lining of the mouth and anus and to the pineal and pituitary bodies. or middle layer. including the glands of the oesophagus. (except the notochord). nervous layer ot EE. gall bladder.G Fig. are derived all structures between the epiblast and hypoblast .: 124 DEVELOPMENT OF THE FROG. epidermic layer of epiblast C. or inner layer. . muscles. neural groove : Y. and also the peritoneum. hypoblast: M. Development of the Nervous System. and the hepatic cells of the liver and the secreting cells of the pancreas the notochord is also formed from hypoblast.. skeleton ^nd lymphatics F.

and plays an important part. Fig. : : : BF. H s N Fig. proctodajum R. and along the flattened area the nervous layer of the epiblast thickens to form the neural plate. Longitudinal vertical section through a frog embryo shortly before the closure of the blastopore. mesoblast MN. {cf. Length of the embryo 2'5 mm. while the lower or nervous layer consists of ovoid or spherical cells. and two or three deep.: THE NERVOUS SYSTEM. central canal of spinal cord Y. fore-brain blastopore BH. 28. more loosely compacted. It is from the latter that the nervous system is developed. Of these the upper or epidermic (Fig. B. mid-brain hypoblast L. liver M. : : : : : : : : : : dorsal surface of the embryo now flattens slightly. neurenteric canal P. X H.. layer is a single stratum of closely fitting cubical cells . The epiblast consists almost from the first of two layers. ingrowth of epiblast to form pituitary body PD. yolk cells. notochord NC. in determining the shape and proportions of the embryo. 30. 2Q). 125 stage of development. The first trace of the nervous system is seen about a weekafter fertilisation. 23). hind-brain BM. rectal diverticulum of mesenteron S. especially in the younger stages. which is wide in front but The . mesenteron N. the distinction between which is already established at the close of segmentation. when the embryo is still spherical and the blastopore has become much reduced in size and difficult to see.

oesophagus P. The neural folds now grow rapidly the groove between them deepens. the neural folds. Length of the tadpole 8imm. 27) and finally meet and fuse. BH. pineal body SD. hind-brain: BM. vesicle of cerebral hemispheres C. bordering the sides of the neural plate and a longitudinal neural groove is formed along its dorsal surface in the median line. liver N. folds together. pituitary body PN. ventricle. truncus arteriosus V. soon appear. depression of floor of forebrain from which the optic vesicles arise OE. Longitudinal vertical section through the anterior end of a tadpole shortly after the time of hatching. narrows posteriorly towards the blastopore.: 126 DEVELOPMENT OF THE FROG. central canal of spinal cord T. : : : I. stomodaium S. slightly raising : A . Fig. thereby converting the neural groove into a tube. pericardial cavity bulum L. the blastopore. mid-brain: A. notochord O. extending forwards from . auricle of heart: BF. fore-hrain infundiCV. Slightly raised ridges. 29. and the folds becoming more and more prominent bend in towards each other (Fig. : : : : : : : : transverse fold connects the anterior ends of the neural up the anterior end of the neural plate.

30. liver N. x 14. stomach H. auricle of heart AD.^ between the nervous system and the alimentary . appear later. depression of floor of fore-brain from which the optic nerves arise OE. The last point at which fusion occurs is a little distance behind the anterior end of the tube.) The short channel of communication between the neural tube and the mesenteron. ventricle. infundibulum J. 12 legs. i. {Cf. Longitudinal vertical section through the head and anterior part of the body of a tadpole about the time of appearance of the hind BB. rudimentary cereC'. 28. A. choroid plexus of fourth ventricle CP'. 127 meet about the junction of the head and from which point the fusion extends rapidly in both directions. : Length of tadpole. fore-brain: BH. and is in free com: Fig. at the place where the pineal body will folds first The neural trunk of the future tadpole. lip L. mid-brain C. In front. .central canal of spinal cord T. basi-hranchial cartilage : BF. choroid plexus of third ventricle F. dorsal : : mm. pituitary body PN. pineal body S. Fig. notochord O. hind-brain BM. truncus arteriosus V.DEVELOPMENT OF THE FROG. coelom or body cavity CH. horny jaws K. lung I. aorta : : : : : : : : : : : : : : : : : : : : : munication with the mesenteron. forwards and backwards.e. pericardial cavity bellum CP. pharynx G. the neural tube ends blindly at its posterior end it opens to the exterior at the blastopore. oesophagus P. cerebral hemisj)here CB.

formed in this way. The Brain.) brain. BM. The neural tube. the mid-brain. is wide from side to side. : spoken of as the neurenteric canal it is only present and closes up before the tadpole hatches. immediately behind which the roof of the hind-brain is thickened transversely The cavity of the hind-brain to form the cerebellum. BH. and a lower and larger portion. 30 it is separated from the mid-brain on the dorsal surface by a well-marked groove. It will be seen that the rectangular bending of the brain. best marked at the sides. and still more largely by the formation of the mouth and the growth forwards of the face and lips. The further development of the brain is illustrated by Figs. which is produced laterally into a pair of hollow outgrowths. The hind-brain. 29 and 30. 128 canal. and the posterior part the The lumen or cavity of the tube persists as the spinal cord. has undergone but little change in Fig. which is known as cranial flexure. the optic vesicles. which forms the angle of the bend and lies opposite the anterior end of the notochord . or hindportion horizontal. is THE NERVOUS SYSTEM. BH. is apparent rather than real. however. which cause the brain to take a much less prominent share in determining the shape of the head. and that of the posterior The posterior portion. central canal of the spinal cord and the ventricles of the brain. except an increase in thickness of its floor and sides. : the spinal cord. the fore-brain. BF. and has moderately thick it is continuous behind with sides and floor. and which was so prominent a feature in the earlier stage. or rectification of the cranial flexure. . but a thin roof for a short time. soon separates from the surface epiblast. (Fig. that this straightening of the brain. remains as the fourth ventricle. into an upper or posterior part. and is brought about partly by the development of the cerebral hemispheres. It is divided by a slight constriction. 29. is no longer obvious a closer comparison of the figures will show.. which grow upwards and forwards from the fore-brain. 28. The anterior or vertical portion has walls of nearly uniform thickness in all parts. the roof of which is very thin : . At thestage represented in Fig. CB. and by thickening of its walls and other changes becomes converted into the central nervous system the anterior part forming 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.

THE BRAIN. and between the layers of which lie the blood vessels of the choroid plexus of the ventricle. In front of the pineal body. In front of the infundibulum is a transverse ridge projecting The : into the ventricle. 29 . this is formed at the spot where the final closure of the neural tube took place. BF. PN. becomes the thalamencephalon of the adult its cavity becomes the third ventricle. They arise as lateral outgrowths from the edges of the neural plate. the roof is thrown into folds which hang down into the ventricle forming a choroid plexus. and may be recognised at a . in connection with which is the pituitary body. arises as a median hollow diverticulum. very narrow from side to side. thickens on its floor to form the crura Its roof grows out laterally into a pair of hollow ovoid cerebri. which become fused together in the median plane. BM. 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. and the third ventricle. the former remaining as a small rounded body outside the skull. which by thickening of its walls to form the optic thalami is reduced to a vertical Its floor is produced downcleft. the wards and backwards into a infundibulum. CP'. from which at a slightly later stage the cerebral hemispheres. The anterior end of the fore-brain grows forwards as a median thin-walled cerebral vesicle. while the stalk persists as a slender pigmented : : : tract within the cranial cavity. similar to that in the medulla. the optic lobes: and its cavity persists as the aqueductus Sylvii. fore-brain. and formed by the roots of the optic nerves. processes. The peripheral nervous system. and at the anterior end of the fore-brain. hollow sac-like diverticulum. The cranial nerves and the dorsal roots of the spinal nerves are formed from the deeper or nervous layer of the epidermis. The roof of the fore-brain remains thin a little behind the middle of its length the pineal body. CP. which hang down into the ventricle. hemispheres grow forwards as the olfactory lobes. I. The mid-brain. arise as a pair of hollow outgrowths . Fig. CH. the foramina of Monro being the apertures of communication being the lateral ventricles or cavities of the The anterior ends of the hemispheres. 129 and thrown into numerous transverse folds.

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 . : G. these soon become constricted at their The Eye. therefore. and then thickens so greatly as almost to obliterate the cavity of the . and are They therefore necessarily formed on the surface of the body. necks so as to be connected with the brain by narrow stalks. from their first appearance conopen tinuous with the brain or spinal cord. From the inner or deeper end of each olfactory j^it a diverticulum. The outer surface of each optic vesicle. inasas the lens alone is formed directly from the surface epiblast. the mouths of the pits forming the nostrils or anterior nares. soon becomes flattened. at first solid. The organs of special sensation are developed from the deeper or nervous layer of the epiblast. 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. and the pits so formed become the nasal sacs . which ultimately become the optic nerves. but soon become connected with these. and become connected with their respective nerves at a very early stage of their formation.130 DEVELOPMENT OF THE FROG. Development of the Sense Organs. The derivation of the sense organs from the epiblast is explained by the fact that they are concerned with the appreciation of the presence and nature of external objects. in the angles between the fore-brain and the optic vesicles. A pitting-in of the surface. involving both layers of the epiblast. while the neural groove is still shallow and they are. much The eye differs from the other sense organs. may be regarded as specially modified portions of the epidermis. and the epiblast lining the pits giving rise to the olfactory epithelium. as the posterior nares. The Nose. soon appears in each of these thickenings. They are at first independent of the dorsal roots. very early stage. as outgrowths from the cord near its ventral surface. or retina. but soon becoming hollow. while the sensitive part of the eye. which is at first in close contact with the surface epiblast. grows downwards to the roof of the pharynx. very shortly after the formation of the mouth opening. into which it opens. The ventral roots of the spinal nerves arise later than the dorsal ones.

and may be spoken of : • as the lens vesicle. The inner wall of the optic cup gives rise to the retina. but soon becomes hollow and breaks away completely from the surface epiblast it becomes later on the lens of the eye. so that the auditory pits do not open to the exterior. and partly through growth of the optic vesicle itself. The optic cup enlarges considerably . : THE SENSE ORGANS. 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 eye developes very slowly. The ears are developed as a pair of pit-like invaginations of the nervous layer of the epiblast at the sides of The invaginations do not involve the epidermic the hind-brain.. and so converted into a cup the optic cup with double walls . The Ear. but elsewhere is separated from it by a space which becomes the posterior chamber of the eye. The choroid and sclerotic coats are formed from the mesoblast surrounding the optic cup. the inner wall being the thickened and originally outer wall of the optic vesicle. it remains in contact with the lens at its edge or lip. the semicircular canals arising — — : . and throughout the tadpole stage of existence is in a very rudimentary and imperfect condition. which proceeds so far as to finally obliterate the cavity. and the auditory vesicles so formed separate from the epiblast and lie in the mesoblast at the sides of the head. Throughout the tadpole stage of existence there is no further . or surface layer of the epiblast. The mouths of the pits very early narrow and close . projecting inwards from the surface this is at first solid. 131 of the deeper or ner- At the same time a thickening : vous layer of the surface epiblast takes place opposite the optic vesicle this grows rapidly and forms a spherical body. owing to thickening of its inner wall. From the optic cup and lens vesicle the adult eye is derived in the following way The lens becomes solid. this latter Ibecomes pitted on its outer surface. The vesicle becomes the vestibule of the adult ear . vesicle. and the outer wall of the cup being the original inner or deeper part of the wall of the vesicle.as outgrowths from it. Partly in consequence of the ingrowth of the lens vesicle. and in which the vitreous humour is iformed.

are found arranged in rows along the body. Fig. H. Development of the Alimentary Canal. The alimentary canal is developed in three lengths: (1) the mesenteron. while the Special Sense Organs. and dseum by epiblast. the mass of the food-yolk becomes confined to the ventral portion of the body region. which is a pitting-in at the anterior end of the body. it : : and from form the anal or cloacal opening. The mouth of the tadpole is also provided with small rounded (See Fig. as well as the notochord (2) the stomodseum. though much smaller. which are probably organs of taste. During the tadpole stage. expansion takes place. supplied by branches of the trigeminal and pneumo-gastric nerves. From teron the mode of their formation is lined by hypoblast. There is some reason.) papillse. this gives rise to nearly the whole length are developed the gill-slits. however. the thyroid. the lungs. is stated to widen somewhat and form the Eustachian passage. 58). The hypoblast. is considerably dilated from the first and at the hinder end of the embryo a similar. for thinking that the Eustachian passage developes independently in the frog. while the layer of integument closing its outer end becomes the tympanic membrane. animal is leading an aquatic life. which is a definite layer of cells. modification .132 DEVELOPMENT OF THE FROG. p. which has at no period opened to the exterior. which is formed by splitting apart of the yolk cells as described above . The mesenteron. 28). 11. The columella. at first confined : . and in other parts of These are lost at the time of the metamorphosis. special sense organs in the form of small epidermal papillae. and not from the hyomandibular cleft. The anterior end of the mesenteron. is formed still later. In this way {cf. follows that the mesenthe stomodseum and proctoit of the mesen- teron has already been described. the pancreas. which has been described with the skull. Fig. but shortly after the metamorphosis the hyomandibular cleft. the liver. which is a similar pitting-in at the hinder end of the body to of the alimentary canal. (fif. 30. the head. in the head region. The early development 1. and round the eyes. from which the mouth and pituitary body are formed and (3) the proctodseum. not extending into either the head or the tail. and the bladder.

. and later on is developed disappears altogether. but shortly before the opening of the mouth.. and the two folds meeting each other divide the canal into a dorsal tube or oesophagus. behind the liver in the later stages the ducts shift so as to open into the bile duct instead of directly : into the intestine. and the alimentary canal is completed as a tube. which from the first is slightly When the tadpole begins to feed. 28 as a is recognisable at a very early stage ventral and backwardly directed diverticulum of the anterior part of the mesenteron. 133 to the roof of the mesenteron. The bladder arises shortly before the metamorphosis as a ventral outgrowth from the hinder end of the mesenteron. which up to this time : The lungs. is definitely established. in tadpoles of about 8 mm. and the gall bladder arises as an out- The liver — — : growth from this. the alimentary canal narrows very considerably . About the time that the lungs first appear. and becomes coiled in a spiral manner. i. length. which soon becomes bind at its distal blind end. approximately uniform metamorphosis. and become thrown into folds between which the vascular mesoblast makes its way the diverticulum itself persists as the bile duct.e. the oesophagus. some time after hatching. small intestine. canal lengthens rapidly. length. forming the anterior boundary of the mass of food-yolk. which appears as this latter it becomes solid after a short time. The pancreas is developed as a pair of hollow outgrowths from the mesenteron. and a from this ventral one which forms the laryngeal chamber They latter the lungs arise as thin-walled lateral outgrowths. In the later stages the walls of the diverticulum thicken. and the distinction between stomach. the alimentary canal shortens considerably. Immediately behind the gill-bearing region or pharynx. it is of During the diameter throughout. The post-anal gut is an extension of the hinder end of the mesenteron into the base of the tail. its sides become folded inwards. gradually spreads round its sides until it encloses the whole of the food-yolk. the alimentary convoluted. in the gill-bearing region. appear first in young tadpoles of about 8 mm. It is perhaps to be regarded as formed mechanical drawing out of the intestine by the outgrowing by a : tail.THE ALIMENTARY CANAL. Fig. and large intestine. Except at the anterior end.

the two structures meet and the definitive anal or perforation occurs cloactl opening is formed. : : : : '. shortly after closure of the neural canal. Figs. 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 the tadpole begins to feed vigorously. length. After the perforation is effected. 28. and a slight depression of the surface epiblast opposite its base. P) is formed from the this rapidly ingrowing stalk of epiblast described above elongates. and slightly larger than the surrounding epiblast cells. The proctodseum or anal invagination appears before the Shortly before the neural folds have met to stomodfeum. 3. At the time of hatching. 28. the lips with the whole anterior part of the face grow forwards rapidly .) The pituitary body (Figs. in tadpoles of about 9 mm. and becomes applied to the ventral surface of the hinder end of the infundibulum to form the pituitary body. .134 DEVELOPMENT OF THE FROG. becomes solid. a little way below the blastopore. From the hinder end of the mesenteron a rectal diverticulum a (Fig. and some time before the tadpole hatches. 29 and 30. this septum becomes perforated. The stomodaeum. growing backwards between the brain and the roof of the mesenteron until it reaches the infundibulum its hinder end now becomes tubular. form the neural tube. Soon after hatching. 28. has been tubular. gives off a few lateral diverticula. The meaning of this curious point has not been ascertained. marks the position of the stomodaeum. this depression is a small shallow pit. 2. and the alimentary canal communicates with the exterior through this stomodaeal pit. separated from the anterior end of the mesenteron by a thin septum. . R) extends downwards towards the proctodteum little later. separates from the stalk. The cells lining it are rather strongly pigmented. For a short time the blastopore and the proctodseum are both open but very shortly after completion of the proctodceum the blastopore closes finally. 29. which soon disappears. the horny jaws are formed. . and remains so until a short time after the formation of the mouth. P. At the stage represented in Fig. the proctodseum is visible as a small median depression of the epiblast at the hinder end of the embryo. 30. {Cf. a conical ingrowth.

which extend outwards towards the surface of the neck as paired outgrowths. Of these outgrowths or pouches. The four hinder visceral clefts perforate the skin about the time of formation of the mouth opening.THE GILL CLEFTS AND ARCHES. and gives rise later on to the lower jaw: the second is the hyoid arch: and the succeeding four are the first. From mid-ventral line. 31). and are known as the : second. third and fourth branchial clefts. 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. though it does not actually open to the exterior. As the opercular folds develope. the hypoblastic epithelium at each side of the buccal cavity becomes thrown into folds. I. and a little later from the third branchial arches as well (Fig. 135 The Gill Clefts little and Arches. and lies between the mandibular and hyoid arches its outer end lies very close to the surface of the neck. third and fourth is hatched a series of on the sides of the head These are the visceral arches. there are five on each side. the hyoid arches a pair of opercular folds arise. on each side. Some vertical ridge-like thickenings appear and neck. The most anterior is the mandibular arch. The most anterior one is the hyomandibnlar cleft. and the branchial arches and The two opercular folds meet below the neck in the clefts. length. The hinder borders of the opercular folds fuse with the body wall except at one place on the left side. arranged in a double row along the ventral half of each of the first three . which grow back over the external gills.e. second. and are replaced functionally by the internal gills. where a spout-like opening remains through which the branchial chamber communicates with the exterior. and enclose the gills in a branchial chamber. in tadpoles of about 9 mm.. first and second branchial. At the same time. and third and fourth branchial arches respectively . and are six in number time before the tadpole branchial arches respectively. second and third branchial. lying between the visceral arches. These latter are delicate thin-walled vascular tufts. and open to the exterior as the gill clefts. the external gills gradually shrivel up. first. which are known as visceral clefts. i. These are slit-like openings lying between the hyoid and first branchial.

30. and produced into processes which unite to form a kind of filtering apparatus. branchial arches. The venous blood. The inner borders of the branchial arches are thickened. on reaching the anterior end of the pericardial cavity. 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. The Circulation during the time the tadpole is breathing by external gills. and 32). or sinus venosus from this it passes into the second or auricular chamber. or sieve. EF^ and EF^. and the course of the circulation in a 6j mm. tadpole. auricle is at first single. now aerated. through which the water. From this latter arise on each side the aortic arches. but later becomes divided by the downgrowth of a septum from its dorsal wall. The arrangement of the blood-vessels. The (Cf. but the internal gills have not yet is shown in Figs. The dorsal aortse gills formed. which carry it.136 DEVELOPMENT OF THE FROG. which unite above the alimentary canal to form the dorsal aorta. 29. is strained before being passed over the gills into the branchial cavity and so out. and in a single row on the fourth branchial arch. shape. : . becomes twisted into an S : : 1. at a time when the external are in full activity. blast of the ventral wall of the pharynx. Figs. which by its branches distributes the arterialised blood to all parts of the body. K. divides at once into right and left branches. thence to the ventricle. returned from the body generally. and divided by transverse constrictions into chambers. Each of these again divides into two. and from that to the truncus arteriosus. the afferent vessels for the first and second branchial arches. enters the posterior end of the heart. to the dorsal aorta in the roof of the pharynx. The Vascular System. The truncus arteriosus. which carry the venous blood to the gills to be aerated from the gills the blood is collected by efferent vessels. 31 and 32. The heart is at first a straight tube developed in the meso- This soon lengthens. While the tadpole is breathing by means of gills. its circulation is in all essential respects that of a fish. taken in through the mouth or nose.

first and third nephrostomes of pronephros RT. dorsal aorta AFi. external gill GM. efferent mandibular vessel GE. segmental or archinephric duct KP. 137 of the two sides run forwards as the carotid arteries. EF3. : : : : : : : : : : : : : : : about the junction of head and body to form the single systemic aorta which supplies arterial blood to all parts of the body. AF3. glomerulus KA. efferent branchial vessels of the first. and third brancliial arches AP. KS3. AF2. posterior comcerebral artery missural artery the arterial circle formed by these commissural vessels with the carotid arteries surrounds the infundibulum of the brain EFi. pulmonary artery AR. tr uncus arteriosus. The heart has been removed. efferent hyoidean vessel EM. second. AC. mm. head kidney or pronephros KSl. second. to supply the head and brain. the aortse of the two sides meeting and uniting a 6| Diagrammatic figure of the head and fore part of the body of tadpole. EF2. and fourth branchial arches EH. Besides the complete sets of afferent and efferent branchial .THE VASCULAR SYSTEM. and also run backwards in the roof of the pharynx. 31. EF4. seen from the ventral surface. A. Fig. afferent branchial vessels of the first. third. showing the arrangement of the branchial vessels as x 33. anterior commissural artery CP. anterior CA.

are present in the hinder arches as well. EF^ which opens into the dorsal aorta. The external gills have been removed. mandibular vein. which AP EF4 ef:3 ef. AF2. AB. 6^ mm. ventricle : auricle fourth branchial arch RA. hyoidean vein YM. efferent ])ranchial vessels of first. second. but as yet ends blindly.: 138 vessels in the first DEVELOPMENT OF THE FROG. carotid artery AFi. there is a short afferent branch AF^ from the afferent vessel of the second branchial arch. lacunar afferent RV. anterior AR. EF2. EF3. . vessel of VD^ Cuvierian vein VH. A. In the third branchial arch. Fig. In the fourth branchial arch there is no afferent vessel. hepatic veins VK. basilar artery AC. EF4. efferent hyoidean vessel EM. dorsal aorta afferent branchial vessels of first. glomerulus LV4.^ ef. efferent mandibular vessel GM. and fourth branchial arches EH. aorta. : : : : : : : : : : : : : : : There is also a well-developed efferent vessel. x 40. 32. and third branchial arches : AT. as yet incompletely developed. eh ab e^ AC Diagrammatic figure of the head and forepart of the body of a tadpole. and second branchial arches. l^alatine artery second. similar vessels. showing the heart. vein of sucker VY. AF3. pulmonary artery EFi. third. anterior cerebral artery AP. and vessels of the branchial arches from the right side.

There are thus six sets of branchial vessels on each side of the pharynx of these. EH. but which never attain full development. are present in the hyoid and mandibular arches. in the hyoid and mandibular arches. but incomplete. in the fourth branchial arch. and third branchial arches. which will become later the pulmonary artery. are rudimentary. Efferent branches. one. From this efferent vessel. and third branchial arches. as there are no afferent vessels corresponding to them. : aorta. are complete . The truncus arteriosus divides at once into right and left . in the first. in which the internal gills are fully established. and showing signs of degenerative changes. THE VASCULAR SYSTEM an efierent vessel. which clearly belong to the same category as the branchial vessels. opening into the dorsal it reaches the aorta. A similar set of vessels. EM. the condition of the blood-vessels is shown in Figs. connecting the heart with the aorta through the gill capillaries. The vessels in the hyoid and mandibular arches undergo further retrograde changes. and at a stage slightly later than that shown in Fig. probably owing to the fact that no gills are formed on these arches. while the tadpole is breathing by external gills. In front of the first branchial arch. a backwardly directed branch arises. A P. vessels are present in the hyoid and mandibular arches. just before — : : 2. The condition of the blood-vessels. On the formation of the internal gills. length. 31 in the third branchial arch as well. opening into the dorsal aorta. and the external gills are shrivelling up. are present in the first and second branchial arches. second. 33 and 34. may be summarised thus Complete systems of afferent and efferent vessels. is present in the fourth branchial arch and vessels formed on the same plan. In tadpoles of 12 mm. 139 EF^ is present. but still less complete. and two. and also a series of similar loops between the afferent and efferent vessels of the fourth branchial arch. second. is incomplete . additional loops of communication are formed in the gill tufts between the afferent and efferent vessels of the first. and need not be described in detail. are present in both hyoid and mandibular arches but these have no connection with the heart. three. The Circulation during the time the tadpole is breathing by internal gills..

hind nephrostomes of head kidney LI. AF. carotid gland branchial arches AL.3. lingual artery between afi"erent and efferent branchial vessels of first branchial arch EF. KS. EF.1. which is about double the length of the head and body. and the head kidneys and their ducts. afferent branchial vessels of first and third A. ventricle TO. has been removed.l. first and third LP. 33.3.140 DEVELOPMENT OF THE FROG. tadpole dissected from the ventral surface to show the heart. glomerulus tubules: KP.i. eJBferent branchial vessels of first and third branchial arches KM. aperture of opercular cavity OP. RT. the branchial vessels. archinephric or segmental duct GM. junction CG. x 22. upper lip LJ. oesophagus. lower lip limb OA. Fig. truncus arteriosus venosus TO. A 12 mm. ov)ercular cavity RS. Wolffian KA. the internal gills. sinus cloaca RV. dorsal aorta EA.3. The tail. rectal spout. : : : : : : : : ^ : : : : : : : : : : : . cut short TR. pronephros or head kidney: KS. AF.

AF^. and fourth branchial arches : GM. hepatic vein VP. branches. left auricle: RT. AF2. second. with which they are . of the first. this Each of these branches divides. anterior cerebral artery AP. In the four afferent branchial vessels AF^. pole from the right side. third and fourth branchial arches respectively are formed. cutaneous artery AT. second. second. anterior commissural vessel gland CP. AF^.. and fourth branchial arches AS. EF4. afi'erent branchial A. pulmonary vein. efferent branchial vessels of first. to show the heart and branchial vessels. tadFig. into three vessels. 34. posterior AR. 141 floor which run straight outwards in the of the pharynx.3 Efvi CP VD AF. pulmonary artery AU. after a short course. EF3. lingual artery vessels of first. gills and the gill capillaries are not represented. right auricle: RB. dorsal aorta AL. posterior vena cava : : : : A : : : : : : : : : : : : : : Each arch. AF^. way GM AB EF. glomerulus: RA. Cuvierian vein arteriosus RV. AF4. RB RA RV Af^a RT diagrammatic figure of the head and neck of a 12 mm. pharyngeal artery CA. VI. posterior commissural vessel EFi. ventricle VH. basilar artery AFi. AB. of the afferent vessel runs outwards and upwards in its own The efferent branchial vessels lie immediately in front corresponding afferent vessels.THE VASCULAR SYSTEM. anterior palatine artery palatine artery CG. carotid AY. truncus VD. third. EF2. The x 35. and the hindmost vessel again into two.

One other point of great importance remains to be noticed in the arrangement of the branchial vessels of the tadpole. length each efferent vessel becomes directly connected at its ventral end with the corresponding afferent vessel. . are branches from the efferent vessels of the fourth branchial arches. The venous blood in the heart is driven by the contraction of the ventricle into the truncus arteriosus. Fig. that the blood in any one of the afferent branchial to the gills. and therefore contain blood which has already passed which it : through the The blood from the lungs is gill capillaries. as already noticed. 3. 34. and all the blood is compelled to pass through the gills to reach the aorta. and thence to the dorsal aorta.along the afferent vessel. The lungs are by this time of considerable size they receive blood by the pulmonary arteries. as before. and the tail begins to shorten. through the capillary loops of the gills. these direct communications between afferent and efferent vessels. and so all over the body. The Changes in the Circulation at the time of the Meta- morphosis. so it may either (1) continue vessels has two paths open to it then reach the efferent vessel by . which. when the anterior limbs are protruded. These direct connections are situated ventrally Figs. in gets aerated. returned direct to the heart by two pulmonary veins which unite and open into the left auricle. At the time of the metamorphosis. to the efferent branchial vessels . connected by very numerous capillary loops in the substance of At their upper the internal gills. the single auricular cavity of the earlier stage being by this time divided by a vertical septum into right and left auricles. though cally present in all four branchial arches. and then along the afferent branchial vessels. and not shown in the figures. ends the efferent vessels open. however. are so small that practino blood passes through them. 33 and 34. and so reach the dorsal aorta without having passed through the gill at all.142 DEVELOPMENT OF THE FROG. The afferent and efferent vessels of each arch at first communicate only through the gill capillaries but in tadpoles of about 12 mm. So long as the tadpole is breathing by gills. into the dorsal aorta. AP. and 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.

so that an increasing of blood takes the direct short passage. arise. p. Fig. and reaches the aorta without having passed through the gills. until the end of the first year . The third aortic arch. the first. 124). though remnants of them usually persist. in the fourth branchial arch. The fourth aortic arch. and (2) a lateral plate. The further changes necessary to convert the circulation into that of the adult are slight. p. which is more dorsally situated. The splitting of the mesoblast into outer or somatopleuric. and finally disappears altogether. Development of the Muscular System and the Coelom. becomes the systemic arch of the frog. 27). Additional work is thus thrown on the lungs and skin. and is present from an early stage of development (Fig. and lies alongside the spinal cord and notochord. 27. also loses its connection with the aorta. lying along each side of the . from which both pulmonary and amount : : : cutaneous arteries L. and inner or splanchnopleuric layers has already been described. loses its connection with the aorta. 143 these direct communications enlarge. CG. in the first branchial arch. The second aortic arch. which form a row of hollow •and somewhat cubical bodies. but is formed by further elaboration of the direct communication between the afferent and efferent branchial vessels of 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 eflerent vessel of the arch. and the change from the gill-breathing to the air-breathing condition is completed. in the second branchial arch. as sometimes stated. Fig. In the body the mesoblast becomes very early divided on each side into (1) a vertebral plate. 5. Fig. is not. though the connection may in the adult become closed and ligamentous {Cf. Of the four aortic arches present at the metamorphosis. The vertebral plate very early becomes divided transversely into muscle-segments or myotomes. are already present. which consequently receive a larger supply of blood the gills rapidly atrophy. 34.THE MUSCULAR SYSTEM. in the third branchial arch. Its dorsal end remains connected with the carotid arch. 34) and the external and internal carotid arteries. in a functionless condition. {Cf. a persistent portion of a gill. The carotid gland. but persists as the pulmooutaneous arch of the adult. which surrounds the side of the body.

The Vertebral Column. the development of which from the hypoblast of the mid-dorsal wall of the mesenteron has already been It forms a cellular rod extending from the blastopore described.e. The myotomes may be well seen in the tail of the tadpole. to the pituitary body and as the tail is formed. but. remain comparatively thin. with tomes from each other are not transverse. The earliest skeletal structure. where they form the great lateral sheets of muscle on each side of the tail. This tube grows upwards at the sides of the spinal cord. 1. and become converted very largely into muscles . especially the inner walls. forms the wall of the The cells covering the alimentary canal and its diverticula. it extends back The notochord consist of vacuolated cells. the cavities of the two sides soon become continuous.) The outer or somatopleuric layer of mesoblast. but the space between them widens out This at considerably. first consists : M. with the hypoblast. and becomes the body cavity or coelom. while the cavities become obliterated. fluid. 29 and 30. anterior portion of the coslom is very early shut off from the hinder part as the pericardial cavity. Owing to the transparency of the tail. the ovaries and testes are formed. Figs. is formed round the notochord from the mesoblast. become the peritoneum. forms the body wall of the adult pleuric layer. their arrangement can be very readily made out the septa dividing the successive myoshaped. Aboiit the time of appearance of the hind legs. : > of two separate halves. owing to the splitting of the mesoblast extending down to the midventral The line. and separated from each other by connective tissue Later on. and time the only one. the cells lining the body cavity. somatopleuric and splanchnopleuric. free surfaces of both layers.. by which the swimming movements are effected. but forward towards the head. a delicate skeletal tube. septa. right and left. {Cf. filled with into it. the angles directed The lateral plates are also in part converted into muscle r the two layers. as we have already seen.144 spinal cord. DEVELOPMENT OF THE FROG. with the epithe inner or splanchnoblast. as a pair of longifor a . i. . from which. is the notochord. at first soft but soon becoming cartilaginous. Development of the Skeleton. the walls of the myotomes thicken considerably. and is invested by a delicate structureless sheath.

and ultimately completely absorbed. They extend into the septa between the myotomes. followed by a posterior unsegmented This transverse portion. The urostyle is the part of the axial skeleton behind the vertebrae . between the successive bony rings.THE SKELETON. the cartilaginous sheath of the notochord becomes cut up into a series of nine vertebrae. which later becomes the urostyle. which remains as a continuous structure until the complete absorption of the tail at the end of the metamorphosis. or even throughout life. and form the first rudiments of the vertebral centra. the notochord. which appeared at the sides of the spinal cord at a slightly earliei stage. and probably correspond to the ribs of other vertebrates. after the metamorphosis. the circumference. but very early fuse with them. are developed in the membrane investing the cartilaginous sheath of the notochord these correspond with the nine vertebrae already present. The transverse processes are at first independent of the corresponding vertebrae. and from the articular ends of each vertebra. very soon become continuous. it is not divided into vertebrae at any stage in development. ossification gradually spreads inwards . By the appearance of transverse lines of demarcation. imbedded in the base of the skull. but a small portion of notochord persists in the middle of each centrum for a long time. one pulling it forwards. divided into an anterior and a posterior portion. which fuse with the bony centra of adjacent vertebrae. but alternate with these . : and ossify to form their articular ends. 145 tudinal ridges with which a series of cartilaginous arches. The vertebrae are not placed opposite the myotomes. is gradually encroached on by the cartilage and bone around it. In the intervertebral regions. annular thickenings of the cartilaginous sheath occur. so as to be hour-glass-shaped in section. division does not affect the notochord. so that each vertebra is acted on by two myotomes on each side. slightly constricted in their centres. Shortly after the metamorphosis thin rings of bone. The anterior end of the notochord. From K . which grow inwards so as to constrict and ultimately obliterate Each of these intervertebral rings becomes. and the other backwards.

The first bone to be developed is the parasphenoid. while other bones primitively the membrane-bones distinct. the frontals and parietals. length these grow back alongside the notochord as a pair of horizontal parachordal rods. At their hinder ends the parachordals grow upwards to form the side walls of the cranium. The hinder ends of the trabeculae are some little distance apart. . and other bones soon follow . this cartilaginous skull is replaced to a considerable extent by cartilage-bone . The parachordals grow rapidly they extend inwards so as to meet each other both above and below the notochord. and a little later bend inwards so as to meet each other above the brain. 146 DEVELOPMENT OF THE FROG. the roof remaining im53) : : may perfect in this region. In the front part of the head a pair of longitudinal cartilaginous bars. which. form a plate of cartilage. all the bones of the adult cranium are present. except the sphenethmoid. becomes closed by a thin plate of cartilage. which underlies the anterior end of the brain. and the lateral margins of the parachordals and trabeculae grow upwards so as to form the side walls of the skull. and complete the occipital Further forwards the pituitary foramen part of the cranium. the trabeculse cranii appear in tadpoles of about 10 mm. enclosing the brain: it is developed as follows. dition is an unsegmented cartilaginous tube.— . which does not appear till some months later. The two parachordals soon fuse together to form the basilar plate. — graft themselves on to consists it. forms a firm cartilaginous floor to the brain case. 2. the trabeculse unite to lies. with the trabeculse. The exoccipitals. The Cranium or brain case. appearing until nearly the time of the metamorphosis. In the adult frog. The Skull. which are at first separate. and by the time the metamorphosis is complete and the tail absorbed. and is produced into blunt processes at its outer angles. distinct elements of which the skull with advantage be described separately. and between them is a space in which the pituitary body In front of this pituitary fossa. p. This in its fully-formed cona. and probably of dermal origin none of the bones of the skull. The skull of the tadpole consists almost entirely of cartilage with the exception of the parasphenoid. which they now completely surround. The three morphologically {cf.

The olfactory capsules are from their first appearance very closely connected with the anterior ends of the trabeculse. forming part They arise about the same time as the auditory capsules . parallel to and below the trabeculse. perpendicular to the long axis It very early. about the time of completion of the metamorphosis. it has altered its direction. both behind and in front of the eyeball. the mandibular arch. however. but before the completion of quite independent of the cranium. they remain distinct life. the most anterior part of the mandibular bar becomes segmented off as a short rod of cartilage. which is the largest of the series. In front of the palatopterygoid. which is directed upwards and forwards in the lower lip it is known as Meckel's cartilage. i. They develope later than the auditory and optic This consists of a series of cartic. and they develope in order from before backwards. b. and before the appearance of the opercular folds. The appear in tadpoles of about 12 cartilaginous auditory capsules mm.THE SKULL.e. the opercular folds they fuse with the upgrowing parachordals The pro-otic appears to form part of the side walls of the skull. and now runs almost horizontally forwards. lies i.. 147 The Sense Capsules. the latter union being effected by a short transverse bar of cartilage the palato-pterygoid. from the cranium throughout order to secure mobility of the eye-balls. Each hoop consists of right and left halves. at first parallel to the others. changes. length as thin shells of They are at first cartilage investing the auditory vesicles. hyoidean arch. undergoes important of the body. which are independent at their dorsal ends. and forms the basis of the lower jaw or mandible. and unlike the other sense The of the sclerotic coats of the eyes. and by the time the external gills are developed. and the four branchial arches respectively . laginous hoops developed within the visceral arches. The Visceral Skeleton. soon imites with the trabeculee. which grow up between them to form the median vertical internasal septum. capsules. in capsules. but fused or closely conThere are in all six of these hoops or bars in nected ventrally. It — : . The mandibular bar. and forming a framework which surrounds and stiffens the walls of the pharynx. optic capsules are thin shells of cartilage.

with corresponding lengthening of the upper and lower jaws.148 DEVELOPMENT OF THE FROG. which does not appear till some months after the completion of the metamorphosis. partly bone. and later on downwards and backwards as in the adult. which may be described as a rotation backwards of the bar. which appears in the lower and outer wall of the auditory capsule about the time the opercular folds are growing back over the gills. it now runs downwards and forwards. which are completed later on by the development of the pterygoid. and which fuses with the stapes at its inner end. ii. This rotation backwards of the distal end of the quadrate. so that in place of running horizontally forwards. which acquires a loose connection at its upper end with the cranium and with the quadrate cartilage. and connected 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 hyoid bar also undergoes important changes. proceeds rapidly during and after the metamorphosis. serving to support the horny jaws of the tadpole. It is a small the stapes consists of two elements. maxilla and other bones. 56. immediately behind the mandibular bar. and changes its direction. 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. Fig. while its outer end becomes : : . articulating at its upper end with the subocular part of the mandibular arch. When the mandibular arch becomes horizontal the hyoid forms a broad stout bar of cartilage. ((7/. At the commencement of the metamorphosis the hyoid bar becomes narrower. The development of the columella is imperfectly known. soon runs vertically downwards. causes lengthening of the these latter palato-pterygoid bar and of Meckel's cartilage become respectively the basis of the upper and lower jaws of the tadpole. 10. the fenestra ovalis. The other portion of the columella is a small rod. 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. instead of being directed downwards and forwards. so that the quadrate. — — . This change. squamosal. one of which plate of cartilage partially filling a hole. In connection with the lips two pairs of small labial cartilages appear. first it is a wide band of cartilage placed nearly vertically in the side wall of the pharynx. partly cartilage. p.

In tadpoles of about 12 mm. The branchial bars are at first simple flattened rods of cartilage. 58): this outer element of the columella is commonly regarded as formed from the uppermost part of the hyoid arch. and become bound together by connective tissue to form the compact Wolffian bodies or kidneys of : . iii. General Account. opening into the body cavity by three ciliated mouths or nephrostomes. As the hind-legs appear. p. and continued back along the dorsal wall as the archinephric or segmental duct. 1. and opens into the cloaca. but becoming early connected with a median basi-branchial cartilage. : N. 33. begin to form in the hinder part of the body as a series of paired tubules. where it joins with the corresponding duct of the opposite side. during the early stages are the head kidneys or pronephra. immediately behind the constricted neck region (Figs.THE URINARY SYSTEM. and together form a complex basket-work supporting the gills. Fig. Later on. and are the sole excretory organs of the tadpole during its early stages. These are a pair of globular organs imbedded in the dorsal wall of the body at its anterior end. Each head kidney is a The excretory organs of its existence. which appears in the floor of the mouth between the ventral ends of the first two pairs of bars. to the hinder end of the body. These Wolffian tubules rapidly increase in number. of the tadpole. The head kidneys and their ducts are well developed in the tadpole at the time of hatching they subsequently increase considerably in size. as well as in size and complexity. convoluted tube with glandular walls. the branchial bars become more slender: their dorsal ends disappear. The Development of the Urinary System. length the adult kidneys or Wolffian bodies (Fig. the branchial bars of each side coalesce with one another both at their dorsal and their ventral ends they also become strongly curved. while their ventral ends fuse with the basi-hyal and basi-branchial cartilages. independent of one another. and together give rise to the body of the hyoid and its posterior cornua. 149 connected with the tympanic membrane icf. as the gills begin to shrink. but appears to be really quite independent of it. 11. 33 and 35 KP). which grow towards and open into the segmental duct. KM). KA.

truncus arteriosus RV. oesophagus. the heart. ventricle TC. 40 mm. The tail has been cut oft. afferent branchial vessels of first and third AFl. lower lip L!. cut short TR. 35. now degenerating LA. fat body efterent branchial vessels of first and third branchial arches KM. A : : : : : : : : : : : : : : : : : : . carotid gland AL. glomerulus body KP. head kidneys diminish in size. tadpole dissected from the ventral surface to show Fig. LP. the branchial vessels. archinephric or segmental duct GM. bodies.: 150 DEVELOPMENT OF THE FROG At the same time the the fully-formed tadpole (Fig. 35 KM). upper lip still within opercular cavity limb OR. CG. and undergo degenerative changes. pronephros or head kidney. genital ridge RT. lingual artery branchial arches F. cloaca TO. Wolffian KA. and the head kidneys and Wolffian x 5. AF3. rectal spout. EF3. fore limb. hind LJ. dorsal aorta EFi. A.

and lying a little distance to the right and left of the notochord. the excretory organs thus consist on each side of (1) a head kidney. and opening into the anterior end of the body cavity by three ciliated openings. The head kidney continues to increase in size. The hinder part of the duct is the archinephric or segmental duct . and by a series of further changes the ureters and generative ducts of the adult become established. opening into the cloaca. in tadpoles of about 20 mm. 151 and by the time of the metamorphosis (Fig.^ some time before hatching. the anterior end of the duct becomes convoluted and twisted on itself to form a ball. The Wolffian bodies persist as the kidneys of the frog . The lips of each groove soon meet and fuse so as to convert the groove into a tube or duct. which is a convoluted tube lined by a glandular epithelium. 36) have almost completely disappeared. the three nephrostomes becoming at the same time lengthened out into short tubes. it remains straight. and it is from the blood of this vein that the epithelial cells of the head kidney tubules separate the excretory matters. and lateral diverticula arising from their sides. a pair of longitudinal grooves appear along the inner surface of the somatopleure. 2. begins to degenerate : . the tubules become obstructed length. indeed almost imbedded in.e. into which it opens. extending from the neck to the hinder end of the body. one behind another.: THE URINARY SYSTEM. or nearly so. The closure of the tube takes place from behind forwards. which is the posterior part of the tube. and the hind limbs are just commencing to appear. the posterior cardinal vein. three holes or nephrostomes. length. and shortly before the tadpole hatches acquires an In tadpoles of about 3 J mm. until the tadpole is about 12 mm. the nephrostomes . the tubules becoming still more convoluted. This convoluted mass is the head kidney or pronephros. i. which are then passed down the duct to the exterior. The head kidney is closely surrounded by. It remains stationary for a time and then. and runs back along the dorsal body wall nearly straight to the cloaca. As the embryo grows. in length. and (2) the archinephric or segmental duct. At the time of hatching. being left through which the tube opens into the body cavity. and at the anterior end is effected imperfectly. The Head Kidney and its duct.

dorsal aorta GM. glomerulus KA. It now shrinks rapidly. and in tadpoles of 40 mm. genital ridge O. others for a time irregularly the whole organ steadily diminishes in size. 35 KP) is not more than half its former size. head kidney. 36. x4. and at the time of the metamorphosis (Fig. ureter OR. near the close of the metamorphosis. : : : . A Tailed frog. tip of ventricle TO. 36 KP) has almost disappeared. disappearing: KU. (Fig. archinephric or F. all three some them become dilated Fig. fat body segmental duct: KM. A. oesophagus. of : collapsed.: 152 DEVELOPMENT OF THE FROG. mouth RV. Wolffian body: KP. cut short. : : : dissected from the ventral surface to shew the kidneys and rej)roductive organs.

: 3. point to a close physiological connection between the two organs. keep pace with those of the head kidney. as they are termed. arises from the aorta on each side (Figs. Opposite the head kidney an irregular sacculated outgrowth. and so give rise to a complicated system of glandular tubules. they also branch freely. though it is not easy to imagine what precise function the glomerulus subserves. though not touching them. It arises on each side as a series of small solid masses of mesoblast cells lying along the inner side of the segmental duct. and the most anterior ones about three segments behind the head kidney. dilate into bulb-like expansions. At their opposite ends these Wolffian tubules. and very close to these. It lies immediately opposite the nephrostomes. and the fact that its growth and subsequent degeneration. in length. and which soon disappear completely. These solid masses soon become elongated into twisted rods. and the organ being reduced to a few small pigmented and irregulary twisted tubules. From the necks of the Malpighiau bodies. 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. or kidney. between this and the aorta (Figs.THE URINARY SYSTEM. the hindmost pair being a short distance in front of the cloaca. 153 nephrostomes having closed up. and growing towards the segmental duct meet and open into it. and so form Malpighian bodies. form the . The Wolffian body. 36 GM) it is very small. when bound together by bloodvessels and connective tissue. 31 and 33 GM) this appears first about the time of hatching. Its close relation to the head kidney. which. and its development keeps pace with that of the head kidney. first appears in tadpoles of from 10 to 12 mm. These rods soon become hollow. and after the first year it can no longer be recognised. The Wolffian Body. which then become tubular. 33 and 35). which have separated from the duct. short solid rods of cells grow towards the peritoneal epithelium and fuse with it. At the time of the metamorphosis (Fig. derived from the dorsal aorta. The Wolffian tubules rapidly increase in number. the glomerulus. It begins to diminish in size about the same time as the head kidney. They develope from behind forwards. which become doubled up by ingrowth of little knots of bloodvessels.

which acts as the excretory duct first of the head kidney. The nephrostomes persist and in the adult frog as many as 200 or more are present on the ventral surface of the kidney. In the female frog it undergoes no further change of importance. In the opens. and will be called the Miillerian duct. male frog the Miillerian duct persists in this condition throughout life. while the posterior part becomes greatly convoluted and acquires thick glandular walls: the hindmost part of the oviduct remains thinner walled. in front of the Wolffian body. and then of the Wolffian body as well. which now ends blindly a short distance in front of the Wolffian body. and extending some distance in front In the female frog the Miillerian duct becomes the of it. becomes divided somewhat obliquely into two. About the time of the metamorphosis the head kidney. and receives the Wolffian tubules of the kidney. w^hich has become rudimentary. to the position characteristic of the peritoneal opening of the adult oviduct. The Wolffian duct becomes in both sexes the ureter.: 154 DEVELOPMENT OF THE FROG. and then by closure of the lips as a tube. The Wolffian and Miillerian ducts. the anterior opening being carried forward first as a groove. now to consider in what way the ureters and generative ducts of the adult frog are formed. which is simply the posterior part of the original segmental duct. but of much greater capacity. the segmental duct. which is now isolated from the Wolffian body. We have 4. this anterior end of the segmental duct. The Miillerian duct becomes connected in 'front with the peritoneal epithelium. In the male frog the hinder end of the Wolffian duct becomes . into which it So far the changes are the same in both sexes. So far we have only described one duct on each side. leading by short tubes into the renal veins. oviduct. 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 a posterior part the Wolffian duct. as minute funnel-like ciliated openings. separates completely from the duct. Wolffian body or kidney of the frog. At its hinder end it grows back along the outer side of the Wolffian duct to the cloaca. after completion of the metamorphosis and entire disappearance of the tail. and acquires an opening into the anterior end of the body cavity. A little later. an anterior part.

In both sexes at an early stage. The Vasa Efferentia. as at their other ends they open into the Wolffian tubules.: THE URINARY SYSTEM. they form passages along which the spermatozoa can get from the testis to the Wolffian duct or ureter. 5. and so out. 155 the glandular enlargement. those nearest to the genital ridges give off tubular branches from their capsules into the ridges. as the Malpighian bodies are forming in the Wolffian body. and. dilated into a much-branched vesicula seminalis. In the male frog these tubules become the vasa efferentia they become connected with the spermatic tubules. greatly. In the female frog these tubules are said to expand very and to give rise to the chambers or cavities present in the adult ovary but the point is not established with certainty. : .

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101 Blood. Blood plasma. 94 — Back-bone. 29 cceliaco-mesenteric. 35—37 Blood corpuscles. 46 Afferent nerves. 59 64 Aqueductus Sylvii. 83. 122. 107. 28 .2 157 INDEX. 29 cesophageal. l4. Abdominal viscera. 136 Aortic arch. 54 occipito-vertebral. 29 cutaneovis. 57 Aorta. 29 Blind spot. 48 —49 Bone angulo-splenial. 98. 123. 28. 99 Archinephric duct. 37 Body-cavity. 22. 29 pulmonary. 30. 28 lingual. 30 iliac. 57 astragalus. 133 Blastopore. 133 Bladder. 105. 20. 29 tibial. 48 Articular process. 29 Articular cartilage. 28 lumbar. external. 29 mesenteric. — peroneal. 14 median. 61 clavicle. 63 fronto-parietal. 19 development of. 148 Basilar plate. 105 Ankle. 78 Alimentary canal. 30 external carotid. 29 hsemorrhoidal. 1C8. 63 calcaneum. 52 59 Axis cylinder. 147 Auditory organ. 32 Axial skeleton. 151 Areolar tissue. 30 iirino-genital.. . 132—134 Ampulla. 146 Bile-duct. 28 cceliac. 136—143 Aperture. urinary. 148 coracoid. 58. 58 ex-occipital. 55. 30 splenic. 53 Auditory capsule. cloacal. 133 Bladder.. 46 Arteries. 28 gastric. 52 Atlas. 62 Adipose tissue. 60 dentary. 14 paired. 28 laryngeal.. 134 . 30 hepatic. 65 Apparatus. 139 sciatic. 28 epigastric. 30 dorsal aorta. 134 27—30 Artery anterior mesenteric. 20. 129 Aqueous humour. 17. 63 Annulus tympanicus. 29 hypogastric. 29 vertebral. 15 — 15 Aponeurosis. 54 femur. gall. 29 occipital. 36. 60 columella. 29 subclavian. 1 — Appendicular skeleton. 16 —18 Acetabulum. 30 internal carotid. . 29 carotid. structure of. 149 Basi-hyal. 52 Basi-branchial. 53. 104—105 Auricle. 30 posterior mesenteric. 63 carpal. 28. 54 girdle. 28—31. 143 Bone. 35 35.

113 114 segmentation of. 65 Coracoid foramen. 128 Cranial nerves. 62 INDEX. 131 Efferent nerves. 63 maxilla. 100 nerves. 28 Cartilage. 62 quadrato-jugal. 63 nasal. 54 Conjunctiva. 55 . 84. 54. 58 metacarpal. 123 epidermic layer of. 144—149 of urinary system. Distal. 104 — parasphenoid. 96 Centrum. 129 — Digestive organs. 97. 55 olfactory. 123. 117 120 Elastic tissue. 49 Capillaries. optic._61. 125 Epicoracoid. 53. 105 Canaliculi. 19 Ear. 136—143 of coelom. 22. 38 Central canal of cord. 55 pterygoid. 100 Circulation of blood. 101 Choroid plexus. 63 palatine. 57 mento-Meckelian. 55. 121. 50 Cauda equina. bile. 128—130 „ Branchial arch. 104-105. 40 ischium. 146 Crura cerebri. 78 Egg. 60 supra-scrapula. 135 Branchial cleft. 60 Epiphysis. 135 139 Branchial chamber^. 98.158 humerus. 11 Development. 148 Condyle. 113 155 of nervous system. 129 Chiasma. 36. 18—20 Dissection. 64 Canal. 98 Connective tissue. 100. 47—48 Cartilage-bone. 61 ilium. 61 scapula. 61 Drawing. 82.. 40 muscle. 63 vomer. 53—55. 36—37 Circulation in tadpole. auditory. 63. 130—132 of alimentary canal. semicircular. 55 omosternum. 100 processes. 52 Cerebellum. 129 — development of. 45^47 Contraction of muscle. 109—155 General account. 19. 83 Choroid. 143—144 Columella. 87 Carotid arch. 57 sternum. 128 Cerebral hemisphere. 136 143 Cloaca. 132— 134 of gill arches and clefts. Cilia. 55 Brain. 85 Cavities of brain. 98 Corpora. 82—83 Cell. 56^ Ciliary Ciliary Ciliary Ciliary Ciliary movement. 146 phalauges. 60 premaxilla. 60 Cornea. 124 130 — — of sense organs. 81. 61 — —117 — . 105. 134 Cochlea. Cardiac plexus. 60 tarsal. 135 Buccal cavity. 129 Cranium. 100 vessels. 143—144 of skeleton. 45 Epiblast. 146 squamosal. 116 formation of. 149 155 Calcar. 133 Duodenum. 14. 62. 80. 135— 136 of circulatory system. adiposa. 15—16 — Dehydration. occipital. 106—108 Cloaca! aperture. 109—113 Detailed Account. 53. 37 Capsule. 20. 109 fertilisation of. 80-84 Coelom. 64 pre-coracoid. 17 Cranial flexure. 125 nervous layer of. 56 pubes. 57 pro-otic. 60 sphen-ethmoid. 58. 3 2—3 Duct. 61 metatarsal. 63 tibio-fibula. 60 OS cruris. 129 Cerebral vesicle. 87 93. 57 radio-ulna.

131 Ligamentum nucha. of 54 83. 116—117 Fibrous tissue. 13—15 Eye. 17. 133 Lateral plate. 113 lower. 112 Lens. 135 internal. 45 Gland. 98 Iter. 38—43 ciliated. 41 simple. 111. 143. 48 Laryngeal chamber. 148 Gill arches. 60 Epithelium. 84 Fissure of cord. 10-=-ll Haversian system. Germinal Germinal Germinal layers. 41 Iris. 112. 60 Glottis. 111. 98-101 Fat-body. 136. 97-103. 102-103 Eye of ox. 57—58 56—57 149-154 Kidney. 116 117 Infundibulum. 34 Lymph heart. . 40 columnar. 40—43 159 squamous. 132 External characters. 58-59 Hyoid arch. 63—64 Glenoid cavity. 39 glandular. 129 Law of Recapitulation. 53. 35 Liver. 46 Female organs. 23 Hepatic plexus. racemose. 56-58. 111. 59 Fertilisation. 136 Eustachian passage. 87 Hepatic portal system. 40 Hallux. 130-131 Eye of frog. 99. 83. 34 Lymph sacs. 124 — Filum terminale. 94-96. external. upper. 87 Genital ridge. 128 Jaw. 64 Hardening. 133 Gasserian ganglion. 120—124 spot. 102—103 Hyaline cartilage. 38—39 stratified. 17. 135 Hypoblast. 17. 95 Hsemorrhoidal plexus. 147-148 Gall bladder. 111—112 of. 66 Lips. 114 Labial cartilage. 133 Lymphatic system. 123. 135—136 Gill clefts. 61—62 Insertion of muscle. 128 26—27 Hind-limb. 133 Lower layer cells. 30-33 development of. 19. 129 — Foramen. 129 Monro. intervertebral. 128. 65 Intercellular substance. 119 Imbedding. 40—43 carotid. 121 Lung. 135 Gills. 15. 149. 15. 83. 17. 101. 54 Food-yolk. Episterniim. 24 tubular. 116 Fenestra ovalis. 41 skeleton Linea alba. 47 Eat-cells. 53. 14. 28 compound. 105. 17. 20. 151—153 Heart. 97. 113 pulsation of. 22-23. 45 46 Hyoid. 144 Lateral ventricle. 14. 63-64 Histology. 109—111. 14. 149-150 Hyomandibular cleft. 114. 16 Grey matter. 43 Limbs. 107—108 Female pro-nucleus.INDEX. 83 Fourth ventricle. 97-98. 13—21 Genital plexus. 19. 114 vesicle. 87 Liquor sanguinis. 15. Hind-brain. Fore-brain. 41 gastric. 38-49. 135 Laboratory rules. 95 Fontanelle. 34 . 41 thyroid. 129 Fore-limb. 97. 89 General Anatomy. 48 Head. 52 magnum. 38 Intestine. 14 Head kidney. 20. 112 61—62. 83. 1 Lacuna. 132. 111. 11^-12 Impregnation.

74 rectus externus. 94 bone. 12 staining. 82 Medullary cavity. 67 depressor palpebrse inferioris. adductor brevis. 4 Mid-brain. 56—57 Maxillary teeth. 44 Muscular system. 14. 74 semitendinosus. 68 striated. 68. of head. 37 Mounting media. 123. 72 rectus superior. Male organs. — 135. 49 Medullary sheath. 75 infraspinatus. 103 Muscles. anterior. 71 rectus internus. 68 temporalis. 68 glutseus. 116 Malpighian body. 66 obliquus superior. 85' cranial. 86 coraco-clavicular. 44 structure of. 72 adductor magnus. 71 obturator. 130 Nephrostome. 76 tibialis posticus. 74 ciliary. 151. 7 mounting. 44 obliquus externus. 50 Mesenteron. methods Mouth. 128. 71 obliquus internus. 11 12 macerating. 100 coccygeal. 70 depressor mandibulse. 122—123. 71 Mounting. 92 brachial. 18 Mesoblast. 6—7 — obliquus inferior. 57—58. 66 rectus anticus femoris. 69 extensor cruris. 100 cucullaris. 68 71 of hind-limb. 74 vastus internus. 69 pyriformis. 77 petrohyoid. 67 levator anguli scapulae. 142—143 Methods. 67 gastrocnemius. 75 rectus abdominis. 66 68 Muscles. hardening. 15—16 of. 6 section-cutting. 8 10 teasing. 148 147— Marrow. 129 Migration of blood corpuscles. 41. 44 submandibular. 70 rectus inferior. 76 pectineus. 69 tibialis anticus. 67 sartorious. 72 rectus internus minor. 68 ilio-psoas. 76 triceps extensor femoris 72 vastvis externvas. 76 geniohyoid. 70 rectus internus major. 57.6 8 7 8 160 Maceration. 10 11 imbedding. 65—77 Myotome. 44 latissimus dorsi. 70 masseter. 67 levator bulbi. 71—77 of trunk. 77 extensor dorsi communis. 75 pectoralis. 66 Medulla oblongata. 72 biceps. 67 intertransversales. 66 peroneus. 143—144 Nares. 70 _ retractor bulbi. 69 mylohyoid. 94 Medullated nerves. 112. 74 voluntary. 106—107 Male. 147—148 involuntary. 72 semimembranosus. 132 Mesentery. 124 Metamorphosis. 87—93. 130 posterior. 129 i . 85 ciliary. 49 Maxillary arch. 154 Nerve. 90 auditory. 7 Membrane — — — — — Microscope. 68 — — retrahens scapulae. 153 Mandibular arch. 69 pterygoideus. 15. 75 sternohyoid. 15. 74 quadratus femoris. 154 Midler's fibres. 68 non-striated. abducens. 15 Meckel's cartilage. 7 INDEX. 6—7 Miillerian duct. 74 hyoglossus. 75 adductor longus. pro-nucleus.

61 Olfactory capsule. 94 Quadrate. 20. 81. 98. 19 Olecranon process. 86 spinal. 97. 81. 94 Proctodaeum. 52 Neural tube. 129 Peritoneum. 130 Operculum. 3 for dissection. 49 Perineurium. crural. 98 Pylorus. 129 Optic vesicle. 22. 129 Optic thalami. 65 Ovary. 87 Renal portal system. 131 Nostril. 26 Reproductive organs. 87. 96 Nerve fibres. 85 ileo-hypogastric. 85 sciatic. 124—130 histology of. 15 Notochord. 7 mounting. 133 Post-axial surface. 104 Peripheral lamellae. 85 vagus. 129 Olfactory organ. 30 Pupil. 144 PerimeduUary lamellae. 84-87. 116 Pollex. 94 Periotic capsule. 95. 84. 94—96 Neural arch. 128. 94. Origin of muscle. 6 — — staining. 129 Pituitary body. 93 radial. 123 Occipital condyle. 81. 61 development of. 2—3 for use of microscope. hardening. 49 Peripheral nervous system. 89 ulnar. 84—93. 92 hypoglossal. 53 _ Sarcolemma.7 8 INDEX. 130 Ora serrata. 53 Oesophagus. 151—153 Pro-nucleus. 147 Olfactory lobe. 88 pathetic. 103 1 Rules of Laboratory. 88 olfactory. 86 motor oculi. 18. 86 161 90 fourth. 86 pneumo-gastric. 135 Optic capsule. 133 Parachordal. 59—60 Pelvic girdle. 126 Neurenteric canal. 147 Optic chiasma. 71 Primitive sheath. 95 Nerve roots. 95 Nose. 93 Nerve cells. 52 Neural fold. 101. Rules for draM'ing. 61 Pulmo-cutaneous arch. 101 Sacrum. 8 — 10 Recapitulation. 52 Neural canal. 100 Reagents. 108 Palato-pterygoid. 127 Neural groove. 83 Optic cup. 93 tibial. 55 Rods and cones. 5—6 Sacculus. 149. 16. 102—103 Rhinal process. 71 Preaxial surface. J 8. 116 Proximal. 25 27 Post-anal gut. 107. 131 Optic lobe. 112 Renal plexus. 113 Oviduct. 148 Non-medullated nerves. 147 Pancreas. 86 trigeminal. 96 Nervous system. 57. 54 Occipito-atlantal membrane. 125 Neural spine. 134 Polar bodies. 55. 128 Neuroglia. 44 . 146 Pectoral girdle. 96 Portal system. 53. 10—11 macerating. 62 Pericardial cavity. 17. 89 peroneal. 19 — Nodes of Ranvier. 80 Pineal body. 89 glosso-pharyngeal. 78—96 facial. 130 sympathetic. 126 Neural plate. 132. 111. 87 optic. 106—108 Retina. 80. 144 Pia mater. 134 Pronephros.

12 Segmental duct. 26 sciatic. 25 parietal. 106. 28—29 Systole. 24 internal jugular. 59—64 axial. 26 Ventral fissure. 30-31 Skeleton. of brain. 123 Segmentation nucleus. 52 53 — Thalamencephalon. 83. 24 development of. 83 22. 8—10 Stapes. 108. 145 Utriculus. Symphysis. 109—155 Taste papillae. 24 subscapular. 24 Tongue. 27 lingual. 52—53 Vertebral column. 154 Urostyle. 143 Spleen. 52 Sclerotic. 21. 155 Vestibule. 57 Suspensory ligament. 84 Ventricle. 55. 86 Spinal nerves. 116 Splanchnopleure. 87 Vesicula sen)inalis. 19 Stomodc^um. 53-59 development of. 33 — Tympanic cavity. 16 Trabeculse cranii. 147 Truncus arteriosus. 107 Vasa efferentia. 113 spermatic. 105 Sense capsules. 146 development of. 97. 22—37 development Vein. 143 Vesical plexus. 106. 23. 130—132 Sheath of Schwann. 143 Solar plexus. Tuber cinereum. 52. 25—27 posterior vena cava. 27 innominate. 151 Segmentation cavity. 129 Third ventricle. 27 dorso-lumbar. 25 subclavian. 155 Vascular system. 144 Spinal cord. 105. 144—145 Vertebral plate. 16 Sinus venosus. 24 femoral. 123. 129 Ventricle. 101 Sympathetic nervous system. 25 hepatic portal. 146 —149 Somatopleure. 21. 7 Teeth. 84. 25 renal portal. 136 — 143 anterior vena cava. 25 pulmonary. 106. Transverse process. 76 Tendon. 134 Subcutaneous tissue. 32 Vertebro3. 111 Susi^ensorium. 95—96 layer cells. 148 Stomach. 84—86 Spinous process. 132. 109. 26 portal. 98. 21 Splitting of mesoblast. 83. 129 Thyroid gland. 18. 15. 106. 81. 14. 26 splenic. 125—126 Spinal ganglia. 27 Tendo Achillis. 87 Spawn. of heart. 118. 24 musculo-cutaneous. 53. 15. 52 59 Skin. 52 Staining reagents. 104 Uvea. 57 mandibular. 9c renal. 24 brachial. 24 vesical. 128. 62 Systemic arch. 120 Ureter. 102 Section cutting. 24-25. 117 120 Semicircular canal. 55. 15. 132 Upper Sense organs. 94 Shoulder girdle. 132 Teasing.162 INDEX. 117 Segmentation of the egg. 109 Spermatozoa. 22. 26 pelvic. 14 Skull. 26 external jugular. 36 of. 27 hepatic. 107. 45 Testis. 100 — Vas deferens. 25 87. 50-64 appendicular. 46 Subocular bar. 26 gastric. 106. 24 intestinal. 148 Sucker. 23 Tadpole. 105 Tympanic membrane. 24 cardiac. 106. 23—27. 24 ovarian. 104 .

114 Vitreous humour. Viscera. 15 163 Wrist. 52 Wolffian body. 124 Yolk plug. 16—17 Visceral arches. White matter. 135 Visceral clefts. 119-120 White fibrous tissue. 154—155 . 135 Visceral skeleton. abdominal. 95 45. 147—149 Vitelline membrane. 114. 98 Vomerine teeth. 121 Yolk hypoblast. 153—154 Wolffian duct. elastic tissue. 45 111. 61 Xiphisternum. 46 Yolk cells.INDEX. 122 Zonule of Zinn. 60 Yellow Yolk. 101 Zygapophysis.

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M. clear. A.. index greatly enhance the value of the mendingit. WORKS BY THE SAME AUTHOR. "This work in plan and execution is admirable." — Medical "Will be extremely useful to all teachers and students of biology . DR. M.. By A. . Cambridge. Milnes F. Cambridge. 15. Ph.. easy. Herbert Hurst. of its kind. 21s. . Beyer Professor of Zoology in Owens College . . he puts his facts in the clearest and shortest way possible. : 8vo. ELDER. With additional Illus- Crown "This book cannot fail to be of great value to those who are studying zoology in their laboratory work and to such we have great pleasure in strongly recom. . London : SMITH.D. written will render it most useful to the "We recommend the work with constudent and to the teacher. 8vo. Lecturer in the Victoria University. and the diagrams (largely original) are always perfect models of lucidity. D. Professor in the Victoria University. making reference Chronicle. Waterloo Place. Marshall. the descriptions The drawings and are well arranged. MiLNES MARSHALL.R. important illustrations are excellent. " The book is welcome. Vertebrate Embryology A Text-Book for students and Practitioners. University . A "In the art of writing text-books Professor Marshall has... M. "A book. .. trations. MILNES MARSHALL and . Manchester.D. M. Revised. and presented in The book. and we do not most valuable. . Sc. Such The an account is here provided final chapter deals with the human embryo. &c. and have been executed in a uniform style. "—Saturday Ee view. D.. and is one of the best books. in our opinion. " Nature. if not the best. The illustrations are medicine "Though embryology has been much throughout good they are mostly origi. John's College. in nal. S. Third Edition."— Edinburgh Medical Journal. .A. no complete account has been given of the development of the common frog or rabbit.— — — — ." Nature. the air of late years. no superior . a form demanding our sincere thanks. It is provided with an exceedingly good index. The directions for dissection are clear... & CO. A Junior Course of Practical Zoolog-y. By A. The mental history of leading types of care and accuracy with which it has beeu vertebrata. Demonstrator and Assistant Lecturer in Zoology."— London Medical Record. C. reflecting the greatest credit upon all concerned . 6d. John's College. Professor in the Victoria . HERBERT HURST. late Fellow of St. Owens College." The Spectator. Beyer Professor of Zoology in Owens College Assisted by late Fellow of St. With numerous Illustrations. most successful and . and sums up the present state of knowledge in a manner which will be equally valuable to the student of medicine and the practitioner. and masterly exposition of the known facts of the developdoubt but that it will obtain the popularity wliich it so well deserves. D. 10s. .S. B. C. with a uniform system of lettering.Sc." The fidence to students and practitioners of British Medical Journal. F.

B. M. each. ST.Sc.A. F.C. MARSHALL. F.D. . Studies from the Biological Laboratories of the VOLUMES I Owens and 2.S. F.D.. M.. CORNISH.. 16. Edited by C.R. MILNES MARSHALL. AND EDITED BY THE LATE Professor A. ANN'S SQUARE.R. LONDON DAVID NUTT & 1894. M.S. College. Price los. CO. published by the COUNCIL OF THE COLLEGE.Sc. J. D.: BIOLOGICAL LECTURES AND ADDRESSES DELIVERED BY THE LATE ARTHUR MILNES MARSHALL. MANCHESTER: E.

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