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Practical Guide to
Large Database
Migration
Preston Zhang

p,
p,
A SCIENCE PUBLISHERS BOOK
A SCIENCE PUBLISHERS BOOK
CRC Press
Taylor & Francis Group
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Boca Raton, FL 33487-2742

© 2019 by Taylor & Francis Group, LLC

CRC Press is an imprint of Taylor & Francis Group, an Informa business

No claim to original U.S. Government works

Printed on acid-free paper

Version Date: 20181226

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Title: Practical guide to large database migration / Preston Zhang,
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Description: Boca Raton, FL : CRC Press, Taylor & Francis Group, [2019] |
Includes bibliographical references and index.
Identifiers: LCCN 2018060369 | ISBN 9781138391628 (hardback : acid-free paper)
Subjects: LCSH: Systems migration. | Database management.
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 Preface

The success of any modern business relies on its ability to adapt to fast changing business
environments: for example, IT infrastructure, database version and servers need to be
upgraded every few years.
When upgrading existing hardware or servers or to a new system, a common task for an
IT team is how to migrate database from old system to new system. We need to make sure
that there is low business downtime during the migration. We also need to test new system
to avoid application failure.
As a database administrator I have done database migrations many times. For example,
migrating a MySQL database from a developer server to a production server or migrating
a SQL Server database from an older server to a new server. Usually, migrating data
within the same database system is not a difficult task. However, it is a challenging when
migrating database from one system to a different system. For example, migrating a SQL
Server database to Oracle database system.
I have tried to find a book about database migration in Oracle, SQL Server and MySQL
database systems, but I only found books about Oracle or AWS database migration. I
would like to share database migration experience with database developers and DBAs.
All the migration examples in this book use large databases in Oracle, SQL Server and
MySQL. This book is written in easy to read style with step-by-step examples.

Who This Book Is For

This book is for intermediate database developers, database administrators. If you are not
familiar with SQL syntax and installation of Oracle/SQL Server/MySQL please read my book
“Practical Guide to Oracle SQL, T-SQL and MySQL”. The SQL code in this book is fully tested
in Oracle 11g, Oracle 12c, SQL Server 2012, SQL Server 2016, MySQL 5.5 and MySQL 5.7.

How to Use this Book

To run the examples from this book you need to install the following database systems and
development tools:
Oracle 11g, Oracle 12c
Oracle SQL Developer
SQL Server 2012, SQL Server 2016
SQL Server Management Studio 2012 or above
MySQL Server 5.5, MySQL Server 5.7
MySQL Workbench 6.3
All the above software can be download from Oracle.com, Microsoft.com and MySQL.com
 Contents

Preface iii

Chapter 1 Introduction to Database Migration 1

1.1 What is Database Migration 1
1.2 Database Migration Stages 1
1.2.1 Migration Preparing Stage 1
1.2.2 Data Migration Stage 2
1.2.3 ETL (Extract, Transform and Local) Stage 3
1.2.4 Database and Application Testing Stage 3
1.3 Database Migration Paths 3
1.4 Database Migration Tools 4
1.4.1 ESF Database Migration Toolkit 5
1.4.2 Data Loader 5
1.4.3 MySQL Workbench Database Migration Wizard 6
1.4.4 SQL Server Migration Assistant Tools 7
1.5 Sample Large Databases 8
1.5.1 SQL Server Chinook Sample Database 8
1.5.2 SQL Server AdventureWorks2012 Sample Database 8
1.5.3 MySQL Employees Sample Database 9
1.5.4 MySQL Sakila Sample Database 9
1.5.5 Oracle Human Resources (HR) Sample Database 10
1.5.6 Oracle Sales History (SH) Sample Database 10
1.5.7 Microsoft Access MonthlySalesReports Sample Database 11
Summary 11
Chapter 2 SQL Server Database Migration 12
2.1 SQL Server to MySQL Migration Example 12
2.1.1 Using ESF Database Migration Toolkit 12
2.1.2 Using MySQL Workbench Migration Wizard 21
2.2 SQL Server to SQL Server Migration Example 29
2.3 SQL Server to Oracle Migration Example 36
Summary 42
Chapter 3 More About SQL Server Management Studio 43
3.1 Visual Database Design 43
3.2 SQL Server Query Designer 47
3.3 Generating Scripts 49
3.4 Activity Monitor 54
3.5 Query Options 55
vi Contents

3.6Template Browser 56
3.7Database Engine Tuning Advisor 57
3.8Disk Usage Report 61
3.9Database Properties 62
3.10
SQL Server Backup 63
3.10.1 Using T-SQL 63
3.10.2 Using Agent Job 63
3.10.3 Using Maintenance Plan 65
Summary 74
Chapter 4 MySQL Database Migration 75
4.1 MySQL to MySQL Migration Example 75
4.1.1 Using MySQL Export Commands and Import Wizard 75
4.1.2 Using MySQL Workbench Schema Transfer Wizard 80
4.2 MySQL to SQL Server Migration Example 86
4.2.1 Using Microsoft SQL Server Migration Assistant for 86
MySQL Tool
4.2.2 Using ESF Database Migration Tool 92
4.3 MySQL to Oracle Migration Example 99
Summary 108
Chapter 5 More About MySQL Workbench 109
5.1 MySQL Database Modeling 109
5.1.1 Visual Database Design 109
5.1.2 Reverse Engineering 114
5.1.3 A Visual Query Builder for MySQL & Other Databases 118
5.2 MySQL Server Management 121
5.2.1 Server Status 121
5.2.2 Server Logs 122
5.2.3 Server Performance Dashboard 123
5.3 Online Backup 123
5.4 MySQL Synchronization 125
5.5 Unique MySQL INSERT IGNORE Statement 130
Summary 130
Chapter 6 Oracle Database Migration 131
6.1 Oracle to MySQL Migration Example 131
6.2 Oracle to SQL Server Migration Example 137
6.3 Oracle to Oracle Migration Example 146
6.3.1 Using Oracle Data Dump Export and Import in 146
SQL Developer
6.3.2 Using ESF Database Migration Toolkit 159
Summary 168
Chapter 7 More About Oracle SQL Developer and Oracle EM 169
7.1 Oracle Query Builder 169
7.2 Oracle SQL Developer DBA Tool 171
7.3 Oracle 12c Enterprise Manager Database Express 173
Summary 179
 Contents vii

Chapter 8 Microsoft Access Database Migration 180

8.1 Microsoft Access to MySQL Migration Example 180
8.2 Microsoft Access to SQL Server Example 184
8.3 Microsoft Access to Oracle Migration Example 188
Summary 194
Index 195

About the Author 197

 Chapter 1

Introduction to Database Migration

1.1 What is Database Migration

Database migration is the process of moving a database from a vendor to another or
upgrading current version of database software. There are many reasons why organizations
would want to migrate their databases:
• High cost of traditional database platform ownership and maintenance cost
• Replacing legacy servers
• Updating storage equipment
• Company mergers
• Moving data to a cloud provider
• Application migration
According to DB-Engines, in June 2018, the most widely used systems are Oracle, MySQL,
Microsoft SQL Server, PostgreSQL, IBM DB2, Microsoft Access, and SQLite. All the
migration examples in this book use SQL Server, MySQL, Oracle and Access because those
are the top databases in the world.
Database migration is not an easy task. For example, it’s quite a challenge to migrate
Oracle database. This book provides step by step guides for migrating databases between
SQL Server, MySQL, Oracle and Microsoft Access. I hope that this book can help database
developers and DBAs to make database migration easier.

1.2 Database Migration Stages

Database migration usually involves several stages:
• Migration Preparing Stage
• Data Migration Stage
• ETL Stage
• Database and Application Testing Stage

1.2.1 Migration Preparing Stage

• Identify the source database tables, views, stored procedures and data
• Identify application and database users on the source system
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2 Chapter 1 Introduction to Database Migration

• Identify privileges and permissions on the source system

• Find application software that uses the source database
• Estimate migration risks and cost
• Check hardware capacity
• Evaluate migration tools

1.2.2 Data Migration Stage

Schema mapping between source database and target database. Data type mapping is easy
if there is no change in database format. For example, database migration from MySQL to
MySQL. However, data type mapping is a difficult task for large tables when migrating
from one database format to another database format. For example, database migration
from SQL Server to Oracle.
Database migration tools can help originations to accomplish migration projects. The tools
usually migrate views to tables. It’s hard to find a tool that can migrate triggers and stored
procedures. DBAs or database developers can create views in target database by using the
migrated tables. They also can generate SQL code for views from source database then
apply the code to target database. For triggers and stored procedures DBAs and database
developers need to get SQL code from the source database then convert the SQL code for
the target database.
Below is a data types mapping table. You can use this table to convert data types manually
or verify data types after database migration. In later chapter you will see that database
migration tools can map date types between a source and a target database automatically.

Table 1-1 Data types mapping table

SQL Server Type MySQL Type Oracle Type Access Type

INT INT NUMBER(10) LONG INTEGER
TINYINT TINYINT NUMBER(3) BYTE
SMALLINT SMALLINT NUMBER(5) INTEGER
BIGINT BIGINT NUMBER(20)
BIT TINYINT(1) NUMBER(3) YES/NO
FLOAT FLOAT FLOAT(53) DECIMAL
REAL FLOAT FLOAT(24) SINGLE
NUMERIC DECIMAL NUMBER(p, s) CURRENCY
DECIMAL DECIMAL NUMBER(p, s) CURRENCY
MONEY DECIMAL NUMBER(19, 4) CURRENCY
SMALLMONEY DECIMAL NUMBER(10, 4)
CHAR CHAR CHAR
NCHAR CHAR/LONGTEXT NCHAR
VARCHAR VARCHAR VARCHAR2 MEMO
NVARCHAR VARCHAR NVARCHAR TEXT

Table 1-1 contd. ...

 Chapter 1 Introduction to Database Migration 3

... Table 1-1 contd.

SQL Server Type MySQL Type Oracle Type Access Type

DATE DATE
DATETIME DATETIME DATE
SMALLDATETIME DATETIME DATE DATE/TIME
TIME TIME
TIMESTAMP TIMESTAMP TIMESTAMP
BINARY BINARY RAW BINARY(SIZE)
VARBINARY VARBINARY RAW
TEXT VARCHAR LONG MEMO
NTEXT VARCHAR LONG
IMAGE LONGBLOB LONG RAW
XML TEXT LONG

1.2.3 ETL (Extract, Transform, and Load) Stage (Optional)

• Sometimes organizations want to extract data from several sources and transform the
data based on business rules then load the data into target databases.
• Transforming process may include sorting, joining data and cleaning data, etc.
• Loading process includes loading the transformed data into target databases or data
warehouses.

1.2.4 Database and Application Testing Stage

• Verify the migrated data on target database
• Create the same application and database users on the target system
• Create the same privileges and permissions on the target system
• Test views, triggers, stored procedures and functions on the target database
• Test the database and application by the end users to make sure there are no errors

1.3 Database Migration Paths

You will see database migration examples from SQL Server to MySQL, SQL Server to SQL
Server and SQL Server to Oracle in Chapter 2. We will use SQL Server Chinook database
in this book.

Figure 1-1 SQL Server migration paths

4 Chapter 1 Introduction to Database Migration

You will see database migration examples from MySQL to MySQL, MySQL to SQL Server
and MySQL to Oracle in Chapter 4. We will use MySQL Employees database in this book.

Figure 1-2 MySQL database migration paths

You will see database migration examples from Oracle to MySQL, Oracle to SQL Server
and Oracle to Oracle in Chapter 6. We will use Oracle HR or SH database in this book.

Figure 1-3 Oracle database migration paths

You will see database migration examples from Microsoft Access to MySQL, Microsoft
Access to SQL Server and Microsoft Access to Oracle in Chapter 5. We will use Microsoft
Access MonthlySalesReports database in this book.

Figure 1-4 Access database migration paths

1.4 Database Migration Tools

There are a lot of database migration tools available. I have tested the following tools and I
will use those tools to demonstrate large database migration in this book.
• ESF Database Migration Toolkit
• Data Loader
• MySQL Workbench Migration Wizard
 Chapter 1 Introduction to Database Migration 5

• SQL Server Migration Assistant for MySQL

• SQL Server Migration Assistant for Oracle
• SQL Server Migration Assistant for Access

1.4.1 ESF Database Migration Toolkit

Website: https://www.easyfrom.net
ESF Database Migration Toolkit can help companies migrate data between Oracle, MySQL,
MariaDB, SQL Server, PostgreSQL, IBM DB2, SQLite, Microsoft Access, etc. It can migrate
most database objects except procedure, function and trigger. Database view will be
migrated to table. The trial version has a migration limit for 50,000 rows per table and it
will add an extra field in tables. That is good enough for testing database migration.
ESF Database Migration Toolkit is very easy to use. It saves database administrators a lot
of time for database migration projects.

1.4.2 Data Loader

Website: https://dbload.com
• Data Loader works well for database migration, but the trial version has 50 rows limit
per table, that’s why I will not use it as a migration tool in this book.
• Data Loader Standard Edition has 100,000 rows limit per table.
• Data Loader Professional Edition has 1,000,000 rows limit per table.
• Data Loader Enterprise Edition has unlimited rows per.
Data Loader supports MySQL, Oracle, MS SQL Server, MS Access, Excel, FoxPro/DBF,
CSV/text files:

Figure 1-5 Data Loader different versions

Figure 1-6 Data Loader database migration paths

6 Chapter 1 Introduction to Database Migration

1.4.3 MySQL Workbench Database Migration Wizard

The MySQL Workbench Migration Wizard allow users to convert an existing database to
MySQL in few steps. The source database might be MySQL, SQL Server, PostgreSQL and
Microsoft Access, etc.

Figure 1-7 MySQL Workbench Database Migration Wizard

1.4.4 SQL Server Migration Assistant Tools

Microsoft SQL Server Migration Assistant (SSMA) is a tool to help database migration to SQL
Server from MySQL, Oracle, Microsoft Access, DB2 and Sybase DB. We will use SQL Server
Migration Assistant for MySQL and SQL Server Migration Assistant for Oracle in this book.

Figure 1-8 SQL Server Migration Assistant for MySQL

 Chapter 1 Introduction to Database Migration 7

Figure 1-9 SQL Server Migration Assistant for Oracle

Figure 1-10 SQL Server Migration Assistant for Access

1.5 Sample Large Databases

1.5.1 SQL Server Chinook Sample Database
The Chinook Database is a sample database for SQL Server, Oracle, MySQL etc. It is being
used for application prototypes and website backend database. It includes the following
tables: Artists, Albums, Employee, Track, Playlist, Playlisttrack, Invoice, Invoiceline, Genre,
Customer and Mediatype. The maximum rows is 8,715 rows (Playlisttrack table).
Download the Chinook Database below:
https://github.com/jimfrenette/chinook-database
8 Chapter 1 Introduction to Database Migration

Figure 1-11 Tables in Chinook database

1.5.2 SQL Server AdventureWorks2012 Sample Database

The AdventureWorks sample database is very useful for testing and learning SQL Server
database. Below is AdventureWorks2012 table names and records (Figure 1-12):

Figure 1-12 Tables in AdventureWorks2012 database

 Chapter 1 Introduction to Database Migration 9

1.5.3 MySQL Employees Sample Database

The Employees sample database provides a large database with size about 160MB. It has
4 million records in total.

Figure 1-13 Sample records from MySQL Employees database salaries table

1.5.4 MySQL Sakila Sample Database

The Sakila sample database includes MySQL new features. It is good for tutorials and
examples. Figure 1-17 shows views, stored procedures and functions in Sakila database.

Figure 1-14 Sakila database structures

10 Chapter 1 Introduction to Database Migration

1.5.5 Oracle Human Resources (HR) Sample Database

Human Resources (HR) database is useful for introducing fundamental database topics.
Figure 1-15 shows that dept_emp table has 331,613 rows.

Figure 1-15 Oracle HR schema dept_emp table records

1.5.6 Oracle Sales History (SH) Sample Database

The sample database has many reports that evaluate past data trends. There are annual,
quarterly, monthly, and weekly sales reports. The database also has data for sales by
geographical area.

Figure 1-16 Oracle Sh schema tables

 Chapter 1 Introduction to Database Migration 11

1.5.7 Microsoft Access MonthlySalesReports Sample Database

Figure 1-17 shows that tblOrderDetails has 121,317 records.

Figure 1-17 Tables in Access MonthlySalesReports database

Summary
Chapter 1 covers the following:
Database migration definition
Database migration stages including Migration Preparing Stage, Database Migration
Stage, ETL Stage and Database and Application Testing Stage
Database migration sample paths
Database migration tools for Oracle SQL Server, MySQL and Microsoft Access databases
Database migration sample databases in Oracle, SQL Server, MySQL and Microsoft Access
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 Chapter 2

SQL Server Database Migration

The main topics in this chapter are illustrated in Figure 2-1: SQL Server database migration
to MySQL, SQL Server database migration to SQL Server and SQL Server database
migration to Oracle.
It’s important to understand the source database before migration. You can open SQL
Server database and view all the tables, views, stored procedures and functions. A decision
needs to be made weather your organization want to keep current database objects or
change them.

2.1 SQL Server to MySQL Migration Example

We will use ESF Database Migration Toolkit and MySQL Workbench Migration Wizard in
this section.

Figure 2-1 SQL Server database migration paths

2.1.1 Using ESF Database Migration Toolkit

Prerequisites: Windows XP/Vista/7/8/8.1/10
SQL Server 6.5 and above
MySQL 3.23 and above
Source Server and Database: SQL Server 2016 Chinook database
Target Server and Database: MySQL 5.7 Chinook database
Migration Tool: ESF Database Migration Toolkit
Database Transform: Yes
 Chapter 2 SQL Server Database Migration 13

Steps to migrate SQL Server to MySQL Using ESF Database Migration Toolkit:
Step 1:
Let us set up Chinook database on the source SQL Server. If you have not downloaded the
Chinook.sql file, please see Chapter 1 for the download link. Open the Chinook.sql file in
Query window and execute the commands. The Chinook database appears in the left pane
after refreshing the Object Explorer:

Figure 2-2 Opening Chinook.sql in a query window

Step 2:
Open table columns you will notice that all the fields have double quotes and brackets.
Remove the double quotes and brackets from the filed names.
14 Chapter 2 SQL Server Database Migration

Figure 2-3 Removing the double quotes and brackets from the filed names

Step 3:
Right Click Chinook database and select Reports -> Standard Reports -> Disk Usage by
Table:

Figure 2-4 Disk Usage by Table Report

 Chapter 2 SQL Server Database Migration 15

Step 4:
Figure 2-5 shows all the table records from Disk Usage by Table Report:

Figure 2-5 Disk Usage by Table Report

Step 5:
Open ESF Database Migration Toolkit. Choose Microsoft SQL Server (Windows
Authentication) as source server. Enter the SQL Server instance name. Username sa and
default port 1433 will be used. Enter SQL Server database name Chinook.
16 Chapter 2 SQL Server Database Migration

Figure 2-6 Choosing Microsoft SQL Server as source server

Step 6:
Create database Chinook on the target MySQL database by running a command:
create database Chinook;

Figure 2-7 Chinook database is created

Random documents with unrelated
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 Fig. 154.—Dissection of a male Teleost (Salmo fario) from the left side.
a.bl, Air-bladder opened; an, anus; au, auricle; b.a, bulbus
aortae; B.HY, basi-hyal; B.OC, basioccipital; cd.a, caudal artery;
cd.v, caudal vein; CN, centrum; crb, cerebellum; d.f.1, first dorsal
fin; D.F.R, dermal fin-rays; du, duodenum or anterior segment of
the intestine; FR, frontal; g.bl, gall-bladder; gul, oesophagus or
gullet; H.SP, haemal spine; int, intestine; kd, kidney; kd′, "head-
kidney"; lg, tongue; lr, liver; N.SP, neural spine; opt.l, optic lobes;
PA.SPH, parasphenoid; ph, pharynx; pn.b, pineal body; pn.d,
bristle passed into ductus pneumaticus; prsen, prosencephalon;
pty.b, pituitary body; PTG, pterygiophores, or radial elements of
dorsal and ventral fins; pv.f, pelvic fin; py.c, pyloric caeca; S.ETH,
supra-ethmoid; S.OC, supra-occipital; spl, spleen; st, stomach; ts,
testis; u.bl, urinary bladder; u.g.s, urino-genital sinus and its
external aperture; ur, ureter or kidney-duct; v, ventricle; v.ao,
ventral aorta; v.df, vas deferens; v.f, ventral fin; VO, vomer. (From
Parker and Haswell.)

In the remaining Fishes the degree of convolution varies within

rather wide limits. The oesophagus is usually straight and wide, but
in Lutodeira, among Teleosts, it is long and even convoluted, and in
the Plectognath Teleosts it gives off a large sac-like outgrowth ("air-
sac"), which extends anteriorly as far as the head, and posteriorly to
the beginning of the tail, and communicates with the oesophagus by
two apertures. The stomach may be U-shaped with the concavity
directed forwards, and consisting of a right limb passing backwards
from the oesophagus, and a left limb curving forwards to its junction
with the intestine (Fig. 153). In such instances as these the stomach
and the adjacent section of the intestine describe a characteristic
siphonal curve. In certain other Fishes (Fig. 160), the oesophageal
portion of the stomach terminates behind in a tubular or sac-like
dilatation at some distance posterior to the laterally situated pylorus,
which indicates the origin of the intestine. The intestine is straight,
or nearly so, in Elasmobranchs, Crossopterygii, and Dipnoi, and also
in a few Teleosts; but sometimes, and very generally in Teleosts, it is
more or less convoluted, notably in some of the Mugilidae, and in
the Loricariidae, where, as in Plecostomus, it is disposed in
numerous spiral coils like a watch-spring. The terminal portion of the
intestine or rectum either opens into a cloaca, which also receives
the urinary and genital ducts, as in Elasmobranchs (Fig. 153), and
Dipnoi (Fig. 155, A), or opens externally by an anus, situated in front
of the separate or united urinogenital ducts, as is the case with all
the remaining groups of Fishes (Fig. 154). The cloacal aperture is
invariably situated near the junction of the caudal and trunk regions,
and as a rule is median in position, rarely, as in the Dipnoi, displaced
to the right or left of the middle line; but the anus differs greatly in
position, sometimes retaining its primitive position at the hinder end
of the trunk, as in the Holocephali, Chondrostei, Crossopterygii,
Holostei, and many Teleosts, or occupying almost any position
between that point and, as in the "Electric Eels" (Gymnotidae), the
ventral surface of the throat (Fig. 351.)
Fig. 155.—A, alimentary canal and liver of a female Protopterus, from
the left side. Part of the left wall of the stomach and intestine,
and the peritoneal investment of the spleen have been removed.
a.p, Abdominal pore; b.d, bile-duct; b.ent, Bursa Entiana; cl,
cloaca; cl.ap, cloacal aperture; cl.c, caecum cloacae; c.m.a,
coeliaco-mesenteric artery; cy.d, bile duct; k.d, kidney duct; m.a,
mesenteric arteries; od, oviduct; pt.c, post-caval vein or inferior
vena cava; p.v, portal vein; the other reference letters as in B.
(From Newton Parker.) B, viscera of an adult female Lepidosteus,
ventral view. The oesophagus, the commencement of the
intestine and the rectum have been laid open. ab, air-bladder; an,
anus; b.d, intestinal aperture of the bile-duct; g.b, gall-bladder;
gl, oesophageal aperture of the air-bladder; h.d, hepatic duct; l,
liver; oes, oesophagus; py, pylorus; py.c; pyloric caeca; py.c′, the
four intestinal orifices of the pyloric caeca; r, rectum; s, spleen;
sp.v, spiral valve; st, stomach. (From Balfour and Newton Parker.)
 Fig. 156.—Transverse section of a Fish, diagrammatic. cn, Centrum;
coel, coelome; d.a, dorsal aorta; d.f, dorsal fin; d.m, dorsal
muscles; d.ms, dorsal mesentery; f.r, fin ray; gon, gonad; int,
intestine; l.v, lateral vein; msn, mesonephros; msn.d,
mesonephric duct; n.a, neural arch; p, parietal layer of the
peritoneum; p′, visceral layer; p.c.v, posterior cardinal vein; pn.d,
Müllerian duct; r, ventral rib; r′, dorsal rib; sp.c, spinal cord; t.p,
transverse process; v.m, ventral muscles; v.ms, ventral mesentery.
(Modified, after Parker and Haswell.)

The whole length of the alimentary canal from the oesophagus to

the rectum is invested externally by the visceral layer of the
peritoneum (Fig. 156), which histologically consists of a stratum of
connective tissue, supporting on its free surface an epithelial stratum
(coelomic epithelium). Primarily, the investing peritoneum is
continued both dorsally and ventrally into bilaminar suspensory
folds, the dorsal and ventral mesenteries (d.ms, v.ms), which extend
to the mid-dorsal or mid-ventral line of the abdominal cavity. The
two layers then separate and become continuous with the parietal
layer of the peritoneum lining the whole of the inner surface of the
body-wall. Embryologically, the two mesenteries owe their formation
to the fusion above and below the mesenteron of the contiguous
walls of two laterally situated and primitively distinct coelomic
cavities. The dorsal mesentery in the adult is occasionally complete,
as in the Myxinoid Cyclostomata and in the Elasmobranch Hypnos
subnigrum,[230] and also in some Dipnoi and in a few Teleosts, but
much more frequently it is reduced by absorption to anterior and
posterior remnants, or to a series of isolated bands, or even, as in
the Lamprey (Petromyzon), to a few filaments accompanying the
intestinal blood-vessels. The ventral mesentery, on the contrary, is
rarely present, and if present is never complete. In Lepidosteus[231]
a ventral mesentery is said to be present in connexion with that part
of the intestine which contains the spiral valve. In Protopterus,[232]
and also in Neoceratodus,[233] there is a well-developed ventral
mesentery in relation with the greater part of the length of the
intestine, although in the former Dipnoid its continuity is interrupted
by one or two vacuities, and in the latter the mesentery is
incomplete posteriorly. A ventral mesentery is also present in the
intestinal region of some of the Muraenidae among Teleosts.[234]

Fig. 157.—Transverse section through a portion of the wall of the

intestine, combined from the condition seen in both the higher
and the lower Vertebrata. Semi-diagrammatic. a.c, Epithelial cells
in the amoeboid state; b.v, blood-vessels; c.m, circular muscular
layer; g, one of Lieberkühn's glands in the higher Vertebrates;
i.ep, intestinal epithelium; l, leucocytes; l′, leucocytes in the
intestinal epithelium; l.f, lymph follicles; l.m, longitudinal muscular
layer; lym, lymphatic vessels; p, visceral layer of the peritoneum;
sm, the submucosa; v, villi of the higher Vertebrates. (From
Wiedersheim.)

Internal to its peritoneal investment the wall of the alimentary canal

consists in succession from without inwards of (1), a muscular coat,
(2) the submucosa, and (3) an epithelial stratum or mucous
membrane, the first two of these layers, with the addition of the
peritoneum, being derivatives of the inner or splanchnic portion of
the embryonic mesoblast.[235]
Excluding the oesophagus, where the muscular coat is mainly
composed of striated fibres, the musculature of the alimentary canal
usually consists solely of non-striated, spindle-shaped fibres
disposed in two layers, an external stratum of longitudinally
arranged fibres, and an inner stratum of circularly disposed fibres
(Fig. 157), with the addition, in the stomach, of an oblique layer
between the two. In the oesophagus the reverse arrangement may
exist, the circular layer being external and the longitudinal internal.
The muscular coat varies considerably in thickness in different
regions and in different Fishes, and in the Cyclostomata, the
Holocephali, some Teleosts, and the Dipnoi may be very feebly
developed, or even entirely absent, as in the intestine of the Hag-
Fish (Myxine). In the Gillaroo Trout (Salmo stomachicus),[236] on the
contrary, the distal section of the siphonal stomach has its
musculature unusually thickened, so as to form an incipient gizzard
for the crushing of the shells of the freshwater Molluscs on which the
Fish feeds. In some of the Mullets (Mugilidae),[237] a true gizzard is
developed by the enormous thickening of the muscular coat of the
caecal stomach, the cavity of which, in consequence, is reduced to a
mere vertical fissure, and is lined by an exceptionally thick, horny
epithelium.

There are a few exceptions to the rule that the muscular fibres are
of the non-striated variety. Thus in some Teleosts, as in the Tench
(Tinca vulgaris), striated fibres are continued from the oesophagus
into the walls of the stomach and intestine, and there form an outer
longitudinal and an inner circular layer, situated externally to the
corresponding layers of the non-striated stratum. In the Siluroid,
Amiurus, the striated fibres of the outer circular layer of the
oesophagus are continued, although but sparsely, into the inner
circular layer of the stomach.

The submucosa (Fig. 157) lies between the muscular layer externally
and the epithelial lining internally, and is characteristically developed
in the stomach, and even more so in the intestine. Histologically, it
consists of a framework of connective tissue, enclosing in its meshes
masses of leucocytes (lymphoid tissue), some of which are
amoeboid and migratory, and may even be found between the cells
of the intestinal epithelium (including in some instances the cloacal
epithelium), probably actively participating in the transmission of
food material from the alimentary canal to the lymphatics and blood-
vessels; while other and somewhat similar, but larger, leucocytes
(phagocytes), are concerned with the elimination of waste
substances or noxious micro-organisms. In addition to the diffused
lymphoid tissue of the submucosa, special rounded or oval, and
sometimes encapsuled, masses of this tissue (lymph follicles) are
common in the intestinal wall (Fig. 157) of Acipenser, the Dipnoi and
some Elasmobranchs, and are perhaps the only representatives in
Fishes of the solitary follicles or "Peyer's patches" of the higher
Vertebrates. A mass of lymphoid tissue exists in the axis of the spiral
valve of Acipenser, which has been compared with a similarly
situated structure in Lepidosiren.[238] In some Elasmobranchs a
large lymphoid organ is imbedded in the submucosa of the
oesophageal wall, while a local thickening of the tissue is met with in
the pyloric sphincter. Protopterus is remarkable among Vertebrates
for the extraordinary development of lymphoid tissue,[239] which,
apart from its distribution in the submucosa, is abundantly present
between the longitudinal and circular muscle layers, and the
peritoneal and muscular coats of the intestine.

In addition to the lymphoid tissue the submucosa contains non-

striated muscle cells and plexuses of capillary blood-vessels, which in
certain Loaches (e.g. Misgurnus), where intestinal respiration occurs,
extend between the cells of the intestinal epithelium. A network of
lymphatic spaces or vessels surrounds the blood-vessels. In some
Elasmobranchs the small arteries of the submucosa of the stomach
are provided with singular sphincter muscles, which occasionally
encircle both the artery and the corresponding vein.[240]
The lining epithelium differs considerably in character in different
portions of the alimentary canal. The epithelium of the mouth,
pharynx, and anterior section of the oesophagus is often squamous
and is succeeded in the hinder part of the oesophagus, and in the
stomach and intestine, by a columnar epithelium. As a rule the
epithelium of the rectum is also columnar, but in Elasmobranchs it
may become squamous. Goblet cells are of very frequent occurrence
throughout the whole length of the alimentary canal, from the
mouth to the rectum inclusive, interspersed between the superficial
epithelial cells; in the same position in the intestine migratory
leucocytes have been found. The primitive ciliation of the Vertebrate
alimentary canal is retained to a greater or less extent in many
Fishes, and is sometimes, but not always, associated with a feeble
development of the musculature. In the larval form of Petromyzon
(Ammocoetes), the whole canal is ciliated except the pharynx and
rectum; but in the adult ciliation is retained only in places which
gradually become fewer as the rectum is approached. In the
Myxinoids, however, cilia are said to be absent.

In the Dipnoi (e.g. Protopterus) the epithelium of the stomach and

intestine is largely ciliated, but in Elasmobranchs, ciliation is usually
restricted to the posterior portion of the oesophagus and the edge of
the spiral valve. Among the more generalised Teleostomi (e.g.
Acipenser, Lepidosteus, Amia), the oesophagus, stomach, and
intestine may be ciliated, but to an extent which varies in different
genera. The pyloric appendages, when present, are also more or
less extensively ciliated. In Teleosts, however, the recorded instances
of ciliation are relatively rare. Nevertheless, ciliated epithelium has
been found in the intestine of a few species (e.g. Rhombus
aculeatus and Syngnathus acus), and also in the pyloric
appendages; in the stomach (e.g. Perca and Esox), and even in the
oesophagus (e.g. Perca).

The mucous membrane, including the submucosa, is frequently

developed into variously arranged ingrowths projecting into the
lumen of the alimentary canal; these are generally of the nature of
longitudinal or transverse ridges, or a combination of the two, giving
rise to retiform structures. The simple longitudinal folds, which are
sometimes found in the oesophagus, stomach, and rectum, often
disappear on distension, and probably merely provide for the
enlargement of these cavities during the deglutition of relatively
large prey, or for the accumulation of faeces. On the other hand, the
permanent and often complicated folds of the intestinal mucous
membrane are probably related to an increase in the secretive or
absorptive area of this portion of the alimentary canal. In the
stomach the mucous membrane is usually smooth, rarely, as in the
"Electric Eel" (Gymnotus), reticulate. In the intestine the folds
assume a highly characteristic and often complicated disposition.
[241] In the Cyclostomata the folds are simple and longitudinally
arranged. In Elasmobranchs (Fig. 158, A), obliquely transverse folds
are present in addition, and, uniting with the longitudinal ridges,
bound linear depressions.

Fig. 158.—The intestinal mucous membrane of different Fishes, to show

the transition from simple longitudinal and transverse folds to
crypts. A, Of an Elasmobranch; B, C, and D, of various Teleosts.
(After Wiedersheim.)

In various Teleostomi (Fig. 158, B, C, D), the union of the two series
of folds becomes more or less retiform, and the network of
intersecting ridges bounds a series of deep tubular crypts which
appear to penetrate to a considerable distance into the intestinal
wall, and possibly foreshadow the characteristic Lieberkühn's glands
of Mammalia. Crypts may also be found in the stomach, where they
receive the apertures of the gastric glands, as in Amiurus, but more
usually they are restricted to the intestine. In the Dipnoi (e.g.
Protopterus) the mucous membrane of the stomach, and—excluding
the Bursa Entiana where a number of oblique folds are present—of
the intestine also, is, on the contrary, perfectly smooth.

In addition to transverse and longitudinal folds the mucous

membrane of the various sections of the alimentary canal is often
developed into outgrowths which are more or less linear.[242] In the
oesophagus these may be papilliform, as in Box and Caesio; obtuse
in Acipenser, hard and almost spine-like in species of Rhombus; or in
the form of pyramidal retroverted processes with jagged or fringed
edges, as in the Spiny Dog-Fish (Acanthias vulgaris). In the Basking
Shark (Selache) similar processes are present, which, near the
stomach, become unusually long and branched, so that the entrance
to that cavity is surrounded by a series of backwardly-directed
arborescent tufts. Peculiar papillose or tag-like processes of the
mucous membrane are frequently present on the spiral valve of
Elasmobranchs, in the intestine of such Teleosts as Balistes, Mugil
and some Pleuronectidae, and also in the rectum of Rhombus
maximus.

Of all the outgrowths from the mucous membrane of the alimentary

canal the so-called "spiral valve" of the Cyclostomata,
Elasmobranchs, Holocephali, Chondrostei, Crossopterygii, Amiidae,
Lepidosteidae and Dipnoi is the most characteristic. The first
appearance of this structure was probably in the form of a straight
longitudinal fold or ridge projecting into the cavity of the intestine,
similar, perhaps, to the typhlosole of many Invertebrata. This
primitive condition is not retained in any existing Fishes, although it
may be closely approached in the larval Cyclostome (Ammocoetes),
and is perhaps also indicated in the straight anterior portion of the
spiral valve of Polypterus. Absent altogether in the Myxinoids, the
valve is represented in its simplest condition, as in certain other
Cyclostomata (e.g. Petromyzon), by a ridge of mucous membrane
which commences anteriorly on the dorsal side, and, after describing
a partial spiral as it passes backwards, terminates posteriorly on the
ventral side, the width of the valve not exceeding half the diameter
of the intestine. This simple type of valve is repeated in embryo
Elasmobranchs, but in the adults of these Fishes the valve becomes
much more complicated, and exhibits a wide range of structural
variation. The increased complexity of the valve seems to depend on
several factors, the effect of which, in different Elasmobranchs, is
best studied in a series of valves of progressively higher
differentiation.[243]

In a hypothetical simple type of valve, easily derivable from the more

primitive type of Petromyzon, it may be conceived that, while not
exceeding in width the semi-diameter of the intestine, the valve
becomes disposed in several complete and more or less closely
approximated spiral turns, the free edge of the valve being on the
same level as its attached margin, and leaving an open axial canal
along the centre of the gut. The nearest approach to this
hypothetical type, which has been compared, not inaptly, to un
escalier tournant sans noyau, is perhaps to be found in the
Thresher-Shark (Alopecias vulpes).

The structure of the more complicated spiral valves of other

Elasmobranchs are well illustrated within the limits of the single
genus Raia.

In one specimen of Raia sp. (Fig. 159, A) the last four coils of the
valve are similar to those of the hypothetical type, but the more
anterior ones, owing to the greater width of the valve, which here
exceeds the semi-diameter of the intestine, have their free margins
deflected downwards, while that portion of the valve which forms
the first half turn is coiled inwards upon itself, so as to form a hollow
cone, open dorsally, and having its apex directed forwards. In other
examples a further modification is introduced by the increasing width
of the valve, which now, throughout its whole length, equals the
semi-diameter of the intestine; and by the formation of an axial
columella by the thickened free edge of the valve, which is traversed
by a central band of unstriped muscle, as well as by the intra-
intestinal artery and vein, and takes the place of the central canal of
the preceding types. The valve is, however, still regular, and its free
margin remains on the same level as the corresponding portion of
the attached edge. In other specimens, again, additional
complications are introduced by a still further increase in the width
of the valve, which now exceeds, often considerably, the semi-
diameter of the intestine, and the consequent deflection of the free
edge of the valve either forwards or backwards (C and D). As shown
in C the valve, in consequence of the backward deflection of its free
margin, presents the appearance of a nest of imperfect truncated
cones with their apices directed backwards, the successive cones
adhering so closely to one another that they combine to form a
central conical chamber with a spirally disposed cavity winding round
it. In D, on the contrary, the free edge of the valve is deflected
forwards, so that, as in C, a nest of cones is formed, but the apices
of the successive cones are directed forwards instead of backwards.
Notwithstanding these variations in the structure of the valve as a
whole, the first coil or half coil nearly always resembles that
described in A.

Fig. 159.—Examples of various types of the spiral valve in

Elasmobranchs. A, B, C, and D in specimens of Raia spp.; E, in
Sphyrna malleus. A, B, and D represents longitudinal sections of
 the intestine, the ventral portion of the valve being removed. In C
successive portions of the ventral wall of the intestine have been
cut out. In E the intestine has been opened along the mid-ventral
line and its wall reflected to the right and left; the ventral portion
of each coil of the "scroll" valve has been removed. In most of the
figures the pylorus is shown in the upper part, and the "rectal"
gland in the lower. (From T. Jeffery Parker.)

It is obvious that the structure of the valve varies considerably within

the limits of the genus, and it may be added that various
intermediate types of structure occur between A and B, A and C, and
A and D. The individual variations are perhaps even more
remarkable, and appear to be quite independent of age and sex. By
way of example it may be mentioned that valves approximating to
one or other of those represented by C and D occur in different
individuals of Raia maculata of the same sex and similar in size, even
in young specimens not more than three inches in length.

As regards other Elasmobranchs, the common Dog-Fish (Scyllium

canicula)[244] has a well-developed spiral valve disposed in twelve
coils, which structurally represents a more highly developed example
of the type D. The existence of considerable individual variation is
nevertheless indicated by the fact that in one specimen examined
the valve was intermediate between C and D, five of the eight cones
projecting forwards and three backwards. In a specimen of
Notidanus sp.[245] there were as many as twenty coils, which in
disposition were intermediate between B and C, approximating,
however, more nearly to B. In a specimen of the Port Jackson Shark
(Heterodontus)[246] the valve had eight coils, and in structure was
also intermediate between B and C, but approached more nearly to
C. Some of the Hammer-headed Sharks (e.g. Sphyrna malleus)[247]
possess a type of spiral valve which differs considerably from any of
those hitherto described, and is termed a "scroll" valve (Fig. 159, E).
The attached edge of the valve pursues a straight longitudinal
course, or at any rate only describes a half turn and back again in
passing from the pyloric to the cloacal extremity of the gut. In the
middle of its course the width of the valve is about equal to two-
thirds of its length, but towards either extremity it gradually
diminishes until the free and attached margins meet. The valve thus
constituted is rolled upon itself from left to right, the successive coils
being comparable to a series of cylinders placed one inside the other,
and becoming gradually larger both in length and diameter from
within outwards. A similar valve is present in some of the
Carchariidae.

In the Holocephali (e.g. Chimaera monstrosa)[248] the valve

describes only three and a half coils, and is further remarkable in
that the attached margin, for a considerable portion of its extent,
does not form a regular spiral but describes only a slightly sinuous
course. Posteriorly, the valve is more normal, and consists of about
two cones with their apices directed forwards.

In the Dipnoi the spiral valve is well developed, and in

Neoceratodus[249] describes nine coils, and in Protopterus[250] six or
seven. The structure of the valve in the latter Dipnoid resembles that
of Scyllium canicula, except for the smaller number of cones.

In the more generalised Teleostomi the valve is best developed in

the Sturgeon (Acipenser) and in Polypterus. In the former[251] the
valve is restricted to the posterior half of the total length of the
intestine, often extending to within an inch of the anal aperture, and
describing in its backward course about seven or eight coils. The
width of the valve is about equal to the semi-diameter of the
intestine, and the thickened free margin forms a well-marked axial
columella, round which the cavity of the gut winds, as in the type B,
except that the spiral is a more open one. In Polypterus the valve
begins close to the solitary pyloric caecum, and for some distance
pursues a straight longitudinal course, but eventually forms a few
spiral coils, ceasing, however, at a considerable distance from the
anus. The evidence afforded by petrified faeces or "coprolites"
proves that certain extinct Crossopterygii (e.g. Macropoma,
Megalichthys), like their living representative, Polypterus, possessed
a spiral valve.[252] In Amia and Lepidosteus[253] the valve is almost
vestigial, being restricted to the terminal portion of the intestine, and
is somewhat variable as to the precise number of its coils. In Amia
there are nearly four coils, extending over 3 cm., that is less than a
tenth of the total length of the intestine, but in some specimens the
coils do not exceed two and a half or three in number.
Lepidosteus[254] has a still shorter valve which, in specimens of 7-10
cm. in length, may not consist of more than three and a half coils,
and in much larger specimens may be reduced to less than two coils,
a variation which suggests that a reduction takes place in the
number of coils as the fish increases in age and size. The structure
of the valve in the three last-mentioned genera resembles that
described in Acipenser, and in none of them does the width of the
valve so far exceed the semi-diameter of the intestine as, by forward
or backward deflection, to give rise to the highly characteristic cones
of Elasmobranchs and Dipnoi.

In the more specialised Teleostomi (Teleostei) the spiral valve is

wholly wanting, except perhaps as a vestigial structure in certain
Clupeoids, as, for example, Chirocentrus,[255] and possibly also in
some Salmonidae.[256]

From what has been said as to the structure of the spiral valve in the
different groups of Fishes, it may be concluded that the valve most
nearly retains its primitive condition in the Cyclostomata; attains its
maximum development in the Elasmobranchs, especially in the
Notidanidae, and shows no indication of degeneration in the Dipnoi.
In the Holocephali and the lower Teleostomi, on the other hand, the
valve exhibits various stages of retrogressive modification, and in the
Teleosts is either absent altogether or persists only as a vestigial
structure in a very few species.
From a physiological point of view the object of the spiral valve is to
increase the absorptive inner surface of the intestine,[257] but, from
what has been said as to the structural variability of the valve, it is
obvious that its efficacy from a functional standpoint must be equally
variable. The value of the valve as an absorptive mechanism
necessarily depends on the area of absorption-surface which it
provides, as well as on the degree of resistance which it offers to the
passage of food material along the cavity of the intestine. These
factors will in turn depend on the number of coils, on the width of
the valve, and on the extent to which its free margin is deflected in
forming the series of cones, but these again are precisely the
structural features which are most liable to variation. The total
absorption area in the four types of valve characteristic of the genus
Raia has been calculated, and may be expressed in square
centimetres as follows:—A, 136.64; B, 143.82; C, 254.3; and D,
276.7.[258] Hence as regards mere absorption area a spiral valve of
the type D has twice the extent of a valve of the type A, and if, in
addition, account be taken of the retardation of the food due to the
increased obstruction offered by the columella and cones in D, it is
clear that the difference in physiological value between the two
types must be far more considerable than is indicated by a
comparison of their relative superficial areas alone.

The evolution of the spiral valve was probably due to the necessity
of increasing the absorptive area of an almost straight unconvoluted
intestine, a result which in other animals is often obtained by an
increase in the length and concurrent convolution of the intestine
itself. Any attempt to correlate the variations in the degree of
perfection or imperfection of the valve considered as an absorptive
mechanism with any special variations in the nature or quality of the
food is, however, a very difficult problem, and a satisfactory
explanation has yet to be found. The difficulty, moreover, is
increased by the fact that the majority of Fishes with a spiral valve
are mainly carnivorous; the Elasmobranchs, in which this structure is
at the same time most highly developed and most variable,
exclusively so. On the other hand, the term "carnivorous" covers a
multiplicity of minor differences in the nature and relative
digestibility of different forms of animal food, and it is quite possible
that it is with differences of this kind that the specific or individual
variations in the development of the spiral valve are associated. The
absence of the valve in the variously nourished Teleosts, save
perhaps as a vestige in one or two, is also difficult to account for,
although it is not improbable that compensating structural
modifications exist in this group. As a rule, the intestine is much
more convoluted in these Fishes, but to an extent which varies
greatly in different species, while the characteristic pyloric caeca and
the spiral valve appear to a certain extent to be developed in inverse
proportion to one another.

The Glands.

The glands associated with the alimentary canal in different Fishes

are (1) the gastric glands, (2) the liver, (3) the pancreas, (4) the
pyloric appendages, and (5) the "rectal" gland.

Oral salivary glands are wanting in all Fishes, the only secretory
structures in the mouth being numerous mucus-secreting goblet
cells, which here, as elsewhere throughout the alimentary canal, are
intermixed with the ordinary epithelial cells.

The Gastric Glands.—The Cyclostomata and Dipnoi do not possess

any specially differentiated gastric glands, and it is probable that in
these Fishes the secretion of the digestive fluids is effected by the
ordinary lining epithelium of the stomach or intestine, or both. In the
remaining groups gastric glands are generally present in the form of
simple caecal structures embedded in the submucosa and opening
on the surface of the mucous membrane into the cavity of the
stomach. The glands differ in different Fishes in the character of
their lining epithelium and in the extent to which their component
cells are differentiated from the epithelium of the stomach. There
does not appear, however, to be any distinction into "central"
(pepsin-forming) and "parietal" (acid-secreting) cells, as is the case
in the higher Vertebrata. Towards the pyloric end of the stomach the
true gastric glands are often replaced by mucous glands. There are,
nevertheless, not a few Teleosts in which special gastric glands are
absent, as, for example, Syngnathus acus, and several species of
Cyprinidae, Labridae, and Blenniidae, etc. In at least two genera
(Gastrosteus and Cobitis), belonging to widely different families,
gastric glands are present in certain species but absent in others. As
suggested by Edinger,[259] the absence of these glands may possibly
be due to degeneration.

It may be remarked that the formation of such digestive ferments as

pepsin and trypsin, which are associated with the stomach and
pancreas respectively, in the higher Vertebrates, is not nearly so
strictly localised in Cyclostomes and Fishes. So far from peptic
digestion being limited to the stomach, it may take place in the
pharynx, stomach, and intestine of Ammocoetes, and in some
Elasmobranchs (e.g. Scyllium), and in such Teleosts as the Pike, Eel,
and Carp, the peptic region extends from the stomach for some
distance along the intestine, while trypsin has been obtained from
the mucous membrane of the stomach, intestine and pyloric caeca,
as well as from the pancreas.[260]

Intestinal glands analogous to the glands of Lieberkühn in the higher

Vertebrates seem to be entirely wanting in Fishes, unless
represented by the sac-like or tubular crypts which are so generally
present in the Teleostomi.

The Liver.—Phylogenetically the oldest gland in connexion with the

Vertebrate alimentary canal, and in size by far the largest, the liver
arises as a caecal outgrowth from the embryonic mesenteron, and in
this primitive stage recapitulates a condition which is retained
throughout life in Amphioxus. By the subsequent division and
branching of this outgrowth the massive compound tubular gland of
the adult Fish is eventually formed.

The liver of Fishes (Figs. 153, 154) is very variable in size, shape,
colour, and degree of lobulation. Anteriorly, it is usually moulded to
the posterior face of the transverse septum between the pericardial
and abdominal portions of the coelom, and from thence extends
backwards in the abdominal cavity to a varying distance, in some
Sharks as far as the cloaca. Externally, the gland is invested by the
peritoneum, which extends on to it from the pericardial septum and
forms a suspensory fold, and also from the oesophagus and
stomach. The shape of the liver usually bears some relation to that
of the body, being, for example, longest in the Eels and broadest in
the Rays. In the great majority of Fishes the liver is bilobed,
consisting of two sub-equal lateral lobes, disposed longitudinally and
confluent anteriorly for a portion of their extent. From this normal
type there are a few minor variations.[261] In Petromyzon,
Lepidosteus (Fig. 155, B), and a few Teleosts (e.g. the
Gymnodontes, Lophobranchii, and some Salmonidae) the liver is
unilobed. In the Myxinoids and in the Dipnoi (e.g. Protopterus), the
organ is bilobed, but the small anterior lobe lies immediately in front
of the much larger posterior lobe, with the gall-bladder between the
two (Fig. 155, A). In some Teleosts (e.g. Scomber), the liver is
trilobed. A gall-bladder is invariably present in either the larval or
adult Cyclostomata, in the Chrondrostei, Holostei, Crossopterygii and
Dipnoi, and generally also in Elasmobranchs and Teleosts. In the
Elasmobranchs it is rarely entirely wanting, as in Sphyrna and Pristis,
and in the Teleosts in some of the Gurnards (Trigla). The gall-
bladder and bile-duct of Petromyzon fluviatilis atrophy after the
metamorphosis which follows the larval Ammocoetes stage, but in
Petromyzon marinus the duct, although usually absent, is sometimes
retained. In the Ammocoetes the epithelium lining the gall-bladder is
ciliated. In some Fishes, as, for example, in many Elasmobranchs,
the gall-bladder is more or less completely embedded in the
substance of the liver; in others, as in most Teleostomi, the organ is
quite distinct from the gland (Fig. 154).

A simple arrangement of the ducts from the liver and gall-bladder is

that found in the common Dog-Fish (Scyllium canicula). In this
Elasmobranch a cystic duct leaves the gall-bladder, and, after
receiving several hepatic ducts from the lobes of the liver, becomes
the bile-duct and opens into the commencement of the intestine. In
the Myxinoids and in the Dipnoi (e.g. Protopterus), there are but two
hepatic ducts, one from each lobe of the liver; these unite and then
meet the cystic duct to form the bile-duct (Fig. 155, A). The number
of hepatic ducts may, however, be considerably increased, as, for
example, in the Siluroid Amiurus,[262] where 8-10 separate ducts
join the cystic duct. In a few instances one of the hepatic ducts
opens directly into the intestine, independently of that which unites
with the cystic duct in forming the bile-duct. In the Dipnoi (e.g.
Protopterus),[263] and in some Teleostomi (e.g. Lepidosteus),[264]
the bile-duct receives the duct from the pancreas before opening
into the intestine.

The Pancreas.—In the Cyclostomes (e.g. Petromyzon, Bdellostoma,

Myxine) a rudimentary pancreas is apparently present, but the
evidence as to its identity is not wholly conclusive. A well-developed
pancreas occurs in Elasmobranchs, in at least one of the Dipnoi, and
probably in most Teleostomi.[265]

In Elasmobranchs the pancreas is a compact structure, uni- or bi-

lobed, and entirely distinct from the liver. In Scyllium canicula (Fig.
153), the bilobed gland lies in the angle between the distal limb of
the stomach and the adjacent portion of the intestine, and from the
smaller of its two lobes the duct issues to pass to its intestinal
aperture near the commencement of the spiral valve. In most of the
Teleostomi in which its existence has hitherto been recorded, the
pancreas is a singularly diffuse gland; and usually a considerable
portion, or even the whole of it, is embedded in the substance of the
liver, its lobules accompanying the ramifications of the hepatic artery
and duct, and the portal vein. The pancreatic duct usually opens into
the intestine near the aperture of the bile duct (e.g. Amiurus);
sometimes the two ducts open on the apex of a common papilla
(e.g. Acipenser and Amia), or by their union form a common duct
(e.g. Lepidosteus). Among the Dipnoi a well-developed pancreas is
present in Protopterus,[266] embedded in the wall of the stomach
and intestine, internal to the peritoneal investment of these organs,
and extending even into the first fold of the spiral valve. The gland is
traversed by fine ductules which unite together and open into the
bile-duct just before the latter enters the intestine. In the remaining
Dipnoi the existence of a pancreas has yet to be ascertained.
Developmentally, the pancreas resembles the liver, and,
histologically, is very similar to that of the higher Vertebrates,
consisting of terminal glandular alveoli continuous with intermediary
tubular portions, and eventually with the finer ductules, which, by
their union, form the main efferent duct.

The Pyloric Caeca.—These structures are caecal outgrowths from

the intestine, and are situated close to the pyloric extremity of the
stomach and the intestinal apertures of the bile and pancreatic
ducts. Wholly wanting in the Cyclostomata and Dipnoi, and, unless
represented by a pair of caeca opening into the long, tubular, non-
valvate anterior portion of the intestine in the Greenland Shark
(Laemargus borealis),[267] in the Elasmobranchs also, they are very
generally present in the Teleostomi, although extremely variable
both in number and arrangement in different families. In Amia there
is no trace of pyloric caeca. Polypterus has a single short caecum
with a thick muscular wall. In Acipenser, Polyodon, and Lepidosteus,
on the contrary, pyloric caeca are unusually well developed. In
Acipenser the caeca are not only numerous, but are so connected
together by connective tissue and blood-vessels, and so invested
externally by the peritoneum, as to form a large, compact, gland-like
mass, communicating with the intestine by a single wide duct. In
Polyodon the organ is essentially similar, but is lobed externally. In
Lepidosteus (Fig. 155, B, py.c), the caeca are also very numerous,
but relatively short, and, although united into a compact mass, open
by four pit-like orifices into the intestinal cavity. In Teleosts the caeca
are subject to extraordinary variations in number, size, and
arrangement.[268] In some families, and even in groups of higher
taxonomic value, they are entirely absent, as is the case with the
Siluridae, Esocidae, Cyprinodontidae, Labridae, Plectognathi, and
Lophobranchii. The "Sand-eel" (Ammodytes) has but a single
caecum; the Turbot (Rhombus maximus) two, and other
Pleuronectidae three to five; and the Perch (Perca), three (Fig. 160,
py.c).

In other Teleosts, on the contrary, these structures are much more

numerous. In Labrus labrax there are about 60, in the Whiting
(Gadus merlangus) 120, while in the Mackerel (Scomber scombrus)
there are no fewer than 191. If few in number the caeca open
separately into the intestine, but when numerous, more or fewer of
them may unite to form a smaller number of efferent ducts, as in the
Whiting, where four such ducts are formed. In some instances, as in
the Tunny (Thunnus), the union of the caeca by connective tissue
leads to the formation of a compact mass. As regards their
arrangement, the caeca may either be disposed in a whorl round the
intestine, as in the Whiting, or in a linear series, as in the Salmon
(Salmo) and in some of the Clupeidae.

The mucous membrane lining the anterior pyloric caeca is often

developed into a network of ridges, limiting crypt-like or tubular
depressions; and not infrequently the epithelium is ciliated.
 Fig. 160.—The alimentary canal of a Perch (Perca). an, Anus; in,
intestine; oes, oesophagus; py, pylorus; py.c, pyloric caeca; st,
stomach. (After Wiedersheim.)

The precise function of these organs, whether digestive or

absorptive, is still uncertain.[269] That they may be digestive is
suggested by the presence of certain amylolytic and proteolytic
enzymes, but this obvious conclusion is to some extent vitiated by
the close proximity of these organs to the stomach, and more
especially to the intestinal orifice of the pancreatic duct. It is by no
means improbable, however, that the caeca are both digestive and
absorptive organs. An attempt has been made to show that the
pyloric caeca and the spiral valve vary inversely as regards the
extent of their development in different groups of Fishes.[270] To
some extent the reciprocal variation of these structures supports this
view, but it is also evident that there are obvious objections to its
unqualified acceptance. Thus, in some Teleostomi (e.g. Acipenser,
Polyodon), exceptionally well-developed and numerous caeca and a
spiral valve are both present. Amia with an almost vestigial spiral
valve has no trace of pyloric caeca, and in Teleosts the absence of a
spiral valve is associated with the complete suppression of the caeca
in many large and important groups.

The Rectal Gland.—The "rectal" gland, or appendix digitiformis, is

a small organ of unknown function with complex glandular walls,
and a central duct opening dorsally into the terminal portion of the
intestine.[271] The organ is generally present in Elasmobranchs (Fig.
153, rct.gl), in which group the intestinal orifice of its duct may
either be close to the termination of the spiral valve, or, as in
Chlamydoselachus,[272] near the cloacal outlet of the gut. An
apparent representative of the gland, the "caecum cloacae," is also
present in the Dipnoi,[273] but communicates directly with the cloaca
(Fig. 155, A, cl.c). The "rectal" gland is perhaps homologous with
the intestinal caecum which is to be found in some Teleosts (e.g.
Box vulgaris), and possibly also with the "caecum" (caecum coli),
and its vermiform appendix in the higher Vertebrata.[274] The
caecum cloacae, on the contrary, is morphologically a urogenital
sinus, formed as a dilatation of the fused hinder portions of the
mesonephric ducts, and probably comparable with the sperm sacs of
male Elasmobranchs, and also with the urinary bladder of
Teleostomes.[275]

CHAPTER X

THE RESPIRATORY ORGANS

The principal respiratory organs consist of a series of pairs of

branchial clefts in the form of perforations in the side walls of the
throat, which place the pharynx in free communication with the
exterior. The first and most anterior of these clefts, the mandibulo-
hyoid cleft or "spiracle," is situated between the mandibular and
hyoid arches; the second, the hyo-branchial or hyoidean cleft,
between the hyoid arch and the first branchial arch; and the
remaining clefts between the succeeding branchial arches. On the
anterior and posterior walls of more or fewer of the clefts highly
vascular plate-like, or variously shaped filamentous outgrowths of
their lining membrane are developed, which subserve the purpose of
exposing the blood to the influence of the oxygen-containing water,
and are termed branchial lamellae or "gills." In addition to their
usual respiratory organs, the gills, a few Fishes utilise the air-bladder
either as a functional lung or as an oxygen reservoir, and in others
accessory breathing organs of various kinds are developed.

The arrangement of the branchial clefts and the gills may be

conveniently studied first in the Elasmobranchs. Excluding the
spiracles, there are usually in this group (Fig. 161, A), five pairs of
branchial clefts, but in certain primitive members of the group the
number may be larger. Thus, in Notidanus griseus (Hexanchus) and
in Chlamydoselachus there are six, and in Notidanus cinereus
(Heptanchus), seven clefts. The pharyngeal apertures of the clefts
are relatively wide, but their external openings, which are freely
exposed on the lateral surface of the head between the eye and the
pectoral fin, are usually narrow and slit-like.

Fig. 161.—A, Horizontal section through the head of an Elasmobranch;

B, similar section of a Teleost (diagrammatic). b.c, Branchial
cavity; b.l, branchial lamellae; c, coelom; e.b.a, external branchial
aperture; hy.a, hyoid arch; hy.c, hyo-branchial cleft; l.s,
interbranchial septum; n, nasal organ; oes, oesophagus; op,
operculum; p.q, palato-quadrate cartilage; Ph, pharynx; sp,
spiracle; s.ps, spiracular pseudobranch; 1-5, 1st to 5th branchial
arches. (From Boas, slightly altered.)

The successive clefts are separated from one another by a series of

inter-branchial septa, each of which consists of the lining membrane
of two contiguous clefts and a median fibrous sheet; it is further
strengthened on its pharyngeal margin by a branchial arch, and
more externally by the fringe of cartilaginous rods (branchial rays)
with which the outer convex edge of each arch is provided. The
anterior and posterior walls of each septum are produced into a
number of outwardly-radiating vascular plates or folds (branchial
lamellae or "gills"), which by their free edges project into the cavity
of the cleft (Fig. 161, A). Although slightly free at their outer
extremities, the lamellae do not extend so far as the external margin
of the septum to which they are attached (Fig. 164, B). Each series
of lamellae is termed a "hemibranch," and, from what has been said,
it is obvious that each inter-branchial septum and its supporting
branchial arch carry two hemibranchs, an anterior and a posterior,
the two forming a complete biserial gill or "holobranch." The hyoid
arch, however, has only a single hemibranch, viz. that pertaining to
the anterior wall of the hyo-branchial cleft, and as the fifth or last
cleft has a hemibranch only on its anterior wall, the fifth arch is gill-
less.[276] The spiracle is a vestigial cleft. At an early stage of
embryonic growth it differs but little from its fellows, but
subsequently degenerating it is represented in the adult by a tubular
passage between the oral cavity and the exterior, which, however, is
often complicated by the development of caecal outgrowths.[277]
The anterior wall of the spiracle often retains a rudiment of a
hemibranch in the shape of more or fewer vascular lamellae, which,
as they are supplied with arterial blood, and not with venous blood
like the ordinary gills, are said to form a mandibular or spiracular
"pseudobranch." The spiracle varies greatly in size in different
families, being largest in the Trygons and Torpedos, and very small,
or even absent in the Lamnidae. Its pseudobranch is best developed
in the Notidanidae, where it has the essential structure of a true
hemibranch, and, as in other Elasmobranchs, but to a greater
extent, probably aids in the additional aeration of the blood which is
distributed to the eye and brain. The characteristic opercular
covering of the external apertures of the gill-clefts in the Teleostomi
and Dipnoi is wanting in Elasmobranchs. It is interesting to note,
however, that in Chlamydoselachus[278] curious frilled cutaneous
folds are developed as extensions of the outer edges of the inter-
branchial septa, as well as of the hyoid region, and, like a series of
incipient opercula, project backwards over the successive branchial
clefts (Fig. 252).

While in many respects more primitive than in Elasmobranchs the

branchial system of the Cyclostomata presents certain special and
peculiar features. The branchial clefts assume the form of oval,
antero-posteriorly flattened pouches or sacs, varying, however, in
number, and in their mode of communicating with the exterior, in
different genera. In the Lamprey (Petromyzon) there are seven pairs
of obliquely-disposed gill-sacs opening externally by small rounded
orifices, and by similar apertures, not directly into the pharynx, but
into a branchial canal (Fig. 162, r.t), which underlies the
oesophagus, and, while ending blindly behind the last pair of sacs,
communicating in front with the oral cavity.[279] The first of the
series of gill-sacs corresponds to the hyo-branchial or hyoidean cleft
of Elasmobranchs and other Fishes. Spiracles are absent in the adult,
but in the embryo are represented by pouch-like outgrowths of the
hypoblast of the oral cavity, which subsequently undergo singular
changes.[280] Thus, the outgrowths become converted into the
lateral halves of a complete ciliated circum-oral groove, which is
retained even in the Ammocoetes stage, and recalls the ciliated
peripharyngeal ring of Ascidians. Another archaic feature is also to
be noted in the continuity of the groove with a ciliated mid-dorsal
pharyngeal ridge, which has been compared to the "dorsal lamina"
of Ascidians, and to the equally characteristic hyperbranchial groove
of Amphioxus.[281] Ventrally also, the lateral halves of the groove
unite to form a single groove, which, after receiving the median
aperture of the thyroid rudiment,[282] is continued backwards in the
mid-ventral line of the pharyngeal wall as far as the last branchial
arch. No trace of these ciliated structures is, however, to be met with
in the adult.
 Fig. 162.—Petromyzon marinus. Transverse section through the
branchial region (semi-diagrammatic). br.m, Branchial membrane;
d.ao, dorsal aorta; d.c, dorsal cartilage of the branchial basket;
d.m, dorsal muscles; e.a, external aperture of a gill-sac; f.t,
fibrous tissue enclosing neural canal; h, i, lateral longitudinal
cartilages of the branchial basket; i.a, internal aperture of a gill-
sac; i.ju, inferior jugular vein; ju, jugular vein (anterior cardinal);
my, spinal cord; nc, notochord; n.ca, neural canal; n.p, neural
process; oes, oesophagus; p.br, peri-branchial lymph sinus; r.m.t,
retractor muscle of the tongue; r.t, respiratory tube or branchial
canal; s, circum-oesophageal lymph sinus; v.ao, ventral aorta; v.c,
ventral cartilage of branchial basket; v.m, ventral muscles. (From
T. J. Parker.)

The branchial lamellae are represented by a series of vascular

horizontal and parallel ridges radiating outwards along the roof, floor,
and lateral walls of each gill-sac, and invested by an epithelium
which is partially ciliated. The inter-branchial septa are much thicker
than in Elasmobranchs, and include not only the walls of adjacent
sacs and the branchial muscles, but also contain cavernous
peribranchial lymph-sinuses. The cartilaginous branchial skeleton is
situated wholly external to the gill-sacs, the so-called branchial
arches lying between the external apertures of the sacs, and directly
beneath the superficial skin, or, in other words, on the outer margins
of the inter-branchial septa, and not on the inner, as is invariably the
case with the branchial arches of Fishes.
 Fig. 163—Dissection of Myxine glutinosa from the left side. au.c,
Auditory capsule; br.ap, left branchial aperture; br.b, rudiment of
branchial basket; br.s.1, first gill-sac; c.br.t, common branchial
tube; cn.c, cornual cartilage; gul, gullet; ht, heart; lg.m, lingual
muscles; m.v.c, median ventral cartilage; na.t, nasal tube; nch,
notochord; n.t, neural tube; oes.ct.d, oesophageo-cutaneous
duct; p.l.c, posterior lateral cartilage; sb.oc.a, subocular arch;
sp.c, spinal cord; st.p, styloid process. (After W. K. Parker, from
Parker and Haswell's Zoology.)

In the Hag-Fish (Myxine) (Fig. 163), there are usually six, very rarely
seven, pairs of gill-sacs, all of which open directly into the pharynx,
and not into a branchial canal as in the Lampreys. On the other
hand, Myxine is unique in having the outer extremities of its gill-sacs
produced into a corresponding number of tubular canals which, after
a longer or shorter course obliquely backwards and outwards, unite
to form on each side a ventrally-situated external aperture (Fig.
163). In the same genus a short canal, or oesophageo-cutaneous
duct, passes from the pharynx behind the last gill-sac of the left
side, and opens externally with the common external branchial
aperture of that side.

In Bdellostoma there are usually six or seven pairs of gill-sacs, but

some species have ten or even fourteen pairs.[283] They agree with
those of the Lamprey in having independent external apertures, but
resemble the corresponding organs in Myxine in opening directly into
the pharynx. An oesophageo-cutaneous duct is also present.[284]

In the Holocephali there are but four branchial clefts, the fifth cleft
being closed. Spiracles are absent in the adult, although present in
the young of Chimaera. The branchial lamellae resemble those of
Elasmobranchs, but the inter-branchial septa are somewhat shorter,
so that the lamellae project slightly beyond their outer margins (Fig.
164, B). A hyoidean hemibranch is present. A noteworthy feature is
the development of a cutaneous fold from the outer surface of the
hyoid arch, which grows backwards over the gill-clefts, and, uniting
above and below with the body-wall, terminates in a free posterior
margin, just behind the last gill-cleft. By the growth of this opercular
fold the gills become enclosed in a spacious branchial cavity, and the
clefts communicate with the exterior through a slit-like opening
between the free margin of the fold and the body-wall.

The reduction in the extent of the inter-branchial septa which is

initiated in the Holocephali is carried to a still further extent in the
Teleostomi. Commencing with the Chondrostei, and passing thence
to the more specialised Teleostei, the septa become gradually
reduced in length, and the branchial lamellae project freely beyond
their outer margins to an increasing extent.

This modification, least marked in Acipenser (Fig. 164, C) and

Polyodon, attains its maximum in the Teleosts (Fig. 164, D and E),
where the branchial lamellae take the form of a double series of free
filaments disposed along the convex outer margin of each branchial
arch, and attached by their bases only to the reduced and
inconspicuous septa. As a general rule each of the first four arches
supports two hemibranchs,[285] forming a biserial gill or holobranch.
In shape the branchial filaments are usually somewhat triangular,
and consist of an axial supporting cartilage or bone, invested
superficially by a highly vascular mucous membrane. As in most of
the preceding groups the fifth branchial arch is gill-less. All
Teleostomi possess a well-developed movable operculum, supported
by a more or less complete series of opercular bones, with or
without the addition of branchiostegal rays (Fig. 161, B). The size of
the external branchial aperture varies considerably. Usually the
hinder and lower margins of the operculum are free, and then the
aperture is spacious. Not infrequently, however, the more or less
extensive fusion of the ventral and hinder edges of the operculum
with the body-wall reduces the aperture to a narrow slit, as in the
Eels and some Siluridae, or to a small upwardly directed pore, as in
the "Sea-Horse" (Hippocampus). In the Symbranchidae the branchial