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C++ Programming: From Problem Analysis to Program Design, Eighth Edition 9-1

Chapter 9
Records (structs)
A Guide to this Instructor’s Manual:

We have designed this Instructor’s Manual to supplement and enhance your teaching
experience through classroom activities and a cohesive chapter summary.

This document is organized chronologically, using the same headings that you see in the
textbook. Under the headings, you will find lecture notes that summarize the section, Teacher
Tips, Classroom Activities, and Lab Activities. Pay special attention to teaching tips and
activities geared towards quizzing your students and enhancing their critical thinking skills.

In addition to this Instructor’s Manual, our Instructor’s Resources also contain PowerPoint
Presentations, Test Banks, and other supplements to aid in your teaching experience.

At a Glance

Instructor’s Manual Table of Contents

• Overview

• Objectives

• Teaching Tips

• Quick Quizzes

• Class Discussion Topics

• Additional Projects

• Additional Resources

• Key Terms

© 2018 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part, except for use as permitted in a
license distributed with a certain product or service or otherwise on a password-protected website for classroom use.
C++ Programming: From Problem Analysis to Program Design, Eighth Edition 9-2

Lecture Notes

Overview
In Chapter 9, students will be introduced to a data type that can be heterogeneous. They
will learn how to group together related values that are of differing types using records,
which are also known as structs in C++. First, they will explore how to create
structs, perform operations on structs, and manipulate data using a struct.
Next, they will examine the relationship between structs and functions and learn
how to use structs as arguments to functions. Finally, students will explore ways to
create and use an array of structs in an application.

Objectives
In this chapter, the student will:
• Learn about records (structs)
• Examine various operations on a struct
• Explore ways to manipulate data using a struct
• Learn about the relationship between a struct and functions
• Examine the difference between arrays and structs
• Discover how arrays are used in a struct
• Learn how to create an array of struct items
• Learn how to create structs within a struct

Teaching Tips
Records (structs)

1. Define the C++ struct data type and describe why it is useful in programming.

Discuss how previous programming examples and projects that used parallel
Teaching
arrays or vectors might be simplified by using a struct to hold related
Tip
information.

2. Examine the syntax of a C++ struct.

3. Using the examples in this section, explain how to define a struct type and then
declare variables of that type.

© 2018 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part, except for use as permitted in a
license distributed with a certain product or service or otherwise on a password-protected website for classroom use.
C++ Programming: From Problem Analysis to Program Design, Eighth Edition 9-3

Accessing struct Members

1. Explain how to access the members of a struct using the C++ member access
operator.
2. Use the code snippets in this section to illustrate how to assign values to struct
members.

Mention that the struct and class data types both use the member access
operator. Spend a few minutes discussing the history of the struct data type
and how it relates to C++ classes and object-oriented programming. Note that the
struct is a precursor to the class data type. Explain that the struct was
introduced in C to provide the ability to group heterogeneous data members
together and, for the purposes of this chapter, is used in that manner as well.
Teaching However, in C++, a struct has the same ability as a class to group data and
Tip
operations into one data type. In fact, a struct in C++ is interchangeable with
a class, with a couple of exceptions. By default, access to a struct from
outside the struct is public, whereas access to a class from outside the
class is private by default. The importance of this will be discussed later in the
text. Memory management is also handled differently for structs and
classes.

Quick Quiz 1
1. True or False: A struct is typically a homogenous data structure.
Answer: False

2. The components of a struct are called the ____________________ of the struct.

Answer: members

3. A struct statement ends with a(n) ____________________.

Answer: semicolon

4. True or False: A struct is typically defined before the definitions of all the functions
in a program.
Answer: True

Assignment

1. Explain that the values of one struct variable are copied into another struct
variable of the same type using one assignment statement. Note that this is equivalent to
assigning each member variable individually.

© 2018 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part, except for use as permitted in a
license distributed with a certain product or service or otherwise on a password-protected website for classroom use.
C++ Programming: From Problem Analysis to Program Design, Eighth Edition 9-4

Note how memory is handled in assignment operations involving struct

Teaching
variables of the same type; namely, that the values of the members of one
Tip
struct are copied into the member variables of the other struct.

Comparison (Relational Operators)

1. Emphasize that no relational aggregate operations are allowed on structs. Instead,

comparisons must be made member-wise, similar to an array.

Ask your students why they think assignment operations are permitted on
Teaching
struct types, but not relational operations. Discuss the issue of determining
Tip
how to compare a data type that consists of other varying data types.

Input/Output

1. Note that unlike an array, aggregate input and output operations are not allowed on
structs.

Mention that the stream and the relational operators can be overloaded to provide
Teaching
the proper functionality for a struct type and, in fact, that this is a standard
Tip
technique used by C++ programmers.

struct Variables and Functions

1. Emphasize that a C++ struct may be passed as a parameter by value or by reference,

and it can also be returned from a function.

2. Illustrate parameter passing with structs using the code snippets in this section.

Arrays versus structs

1. Using Table 9-1, discuss the similarities and differences between structs and arrays.

Spend a few minutes comparing the aggregate operations that are allowed on
Teaching structs and arrays. What might account for the differences? Use your previous
Tip exposition on the history of structs and memory management to facilitate this
discussion.

© 2018 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part, except for use as permitted in a
license distributed with a certain product or service or otherwise on a password-protected website for classroom use.
C++ Programming: From Problem Analysis to Program Design, Eighth Edition 9-5

Arrays in structs

1. Explain how to include an array as a member of a struct.

2. Using Figure 9-5, discuss situations in which creating a struct type with an array as a
member might be useful. In particular, discuss its usefulness in applications such as the
sequential search algorithm.

Ask your students to think of other applications in which using an array as a

member of a struct might be useful. For example, are there applications in
Teaching
which parameter passing might be reduced by using struct members in
Tip
conjunction with arrays? Also, are there other data members that would be useful
to include in the listType struct presented in this section?

3. Discuss situations in which a struct should be passed by reference rather than by

value. Use the sequential search function presented in this section as an example.

structs in Arrays

1. Discuss how structs can be used as array elements to organize and process data
efficiently.

2. Examine the employee record in this section as an example of using an array of

structs. Discuss the code for the struct as well as the array processing code. Use
Figure 9-7 to clarify the code.

Emphasize that using a structured data type, such as a struct or class, as the
Teaching element type of an array is a common technique. Using the vector class as an
Tip example, reiterate that object-oriented languages typically have containers such
as list or array types that in turn store objects of any type.

structs within a struct

1. Discuss how structs can be nested within other structs as a means of organizing
related data.

2. Using the employee record in Figure 9-8, illustrate how to reorganize a large amount of
related information with nested structs.

3. Encourage your students to step through the “Sales Data Analysis” Programming
Example at the end of the chapter to consolidate the concepts discussed in this chapter.

© 2018 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part, except for use as permitted in a
license distributed with a certain product or service or otherwise on a password-protected website for classroom use.
C++ Programming: From Problem Analysis to Program Design, Eighth Edition 9-6

Quick Quiz 2
1. What types of aggregate operations are allowed on structs?
Answer: assignment

2. Can struct variables be passed as parameters to functions? If so, how?

Answer: struct variables can be passed as parameters either by value or by reference.

3. True or False: A variable of type struct may not contain another struct.
Answer: False

4. True or False: A variable of type struct may contain an array.

Answer: True

Class Discussion Topics

1. With the advent of object-oriented programming, is it ever necessary to use C-type
structs rather than classes? If so, when? What are the advantages or disadvantages of
each approach?

2. Discuss how the object-oriented concept of reusability relates to structs, structs

within arrays, arrays within structs, and structs within structs. Ask students to
think of some applications in which defining these data types for later use would be
beneficial.

Additional Projects
1. Write a program that reads students’ names followed by their test scores. The program
should output each student’s name followed by the test scores and the relevant grade. It
should also find and print the lowest, highest, and average test score. Output the name
of the students having the highest test score.

Student data should be stored in a struct variable of type studentType, which has
four components: studentFName and studentLName of type string, testScore
of type int (testScore is between 0 and 100), and grade of type char. Suppose
that the class has 20 students. Use an array of 20 components of type studentType.

2. Write a program that lists all the capitals for countries in a specific region of the world.
Use an array of structs to store this information. The struct should include the
capital, the country, the continent, and the population. You might include additional
information as well, such as the languages spoken in each capital. Output the countries
with the smallest and largest populations.

© 2018 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part, except for use as permitted in a
license distributed with a certain product or service or otherwise on a password-protected website for classroom use.
C++ Programming: From Problem Analysis to Program Design, Eighth Edition 9-7

Additional Resources
1. Data Structures:
http://www.cplusplus.com/doc/tutorial/structures/

2. struct (C++):
https://msdn.microsoft.com/en-us/library/64973255.aspx

3. Classes, Structures, and Unions:

https://msdn.microsoft.com/en-us/library/4a1hcx0y.aspx

Key Terms
 Member access operator: the dot (.) placed between the struct and the name of one
of its members; used to access members of a struct
 struct: a collection of heterogeneous components in which the components are
accessed by the variable name of the struct, the member access operator, and the
variable name of the component

© 2018 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part, except for use as permitted in a
license distributed with a certain product or service or otherwise on a password-protected website for classroom use.
Another Random Scribd Document
with Unrelated Content
The development of the head-kidney in Amphibia has been fully
elucidated by the researches of W. Müller[434], Götte[435], and
Fürbringer[436], while to the latter we are indebted for a knowledge of
the development of the Müllerian duct in Amphibians. The first part
of the urinogenital system to develop is the segmental duct
(Vornieregang of Fürbringer), which is formed by a groove-like
invagination of the peritoneal epithelium. It becomes constricted into
a duct first of all in the middle, but soon in the posterior part also. It
then forms a duct, ending in front by a groove in free
communication with the body-cavity, and terminating blindly behind.
The open groove in front at first deepens, and then becomes
partially constricted into a duct, which elongates and becomes
convoluted, but remains in communication with the body-cavity by
from two to four (according to the species) separate openings. The
manner in which the primitive single opening is related to the
secondary openings is not fully understood. By these changes there
is formed out of the primitive groove an anterior glandular body,
communicating with the body-cavity by several apertures, and a
posterior duct, which carries off the secretion of the gland, and
which, though blind at first, eventually opens into the cloaca. In
addition to these parts there is also formed on each side of the
mesentery, opposite the peritoneal openings, a very vascular
projection into this part of the body-cavity, which is known as the
glomerulus of the head-kidney, and which very closely resembles in
structure and position the body to which we have assigned the same
name in the chick.
The primitive segmental duct is at first only the duct for the head-
kidney, but on the formation of the posterior parts of the kidney
(Wolffian body, &c.) it becomes the duct for these also.
After the Wolffian bodies have attained to a considerable
development, the head-kidney undergoes atrophy, and its peritoneal
openings become successively closed from before backwards. At this
period the formation of the Müllerian duct takes place. It is a solid
constriction of the ventral or lateral wall of the segmental duct,
which subsequently becomes hollow, and acquires an opening into
the body-cavity quite independent of the openings of the head-
kidney.
The similarity in development and structure between the head-
kidney in Amphibia and the body we have identified as such in Aves,
is to our minds too striking to be denied. Both consist of two parts—
(1) a somewhat convoluted longitudinal canal, with a certain number
of peritoneal openings; (2) a vascular prominence at the root of the
mesentery, which forms a glomerulus. As to the identity in position
of the two organs we hope to deal with that more fully in speaking
of the general structure of the excretory system, but may say that
one of us[437] has already, on other grounds, attempted to shew that
the abdominal opening of the Müllerian duct in the bird is the
homologue of the abdominal opening of the segmental duct in
Amphibia, Elasmobranchii, &c., and that we believe that this
homology will be admitted by most anatomists. If this homology is
admitted, the identity in position of this organ in Aves and Amphibia
necessarily follows. The most striking difference between Aves and
Amphibia in relation to these structures is the fact that in Aves the
anterior pore of the head-kidney remains as the permanent opening
of the Müllerian duct, while in Amphibia, the pores of the head-
kidney atrophy, and an entirely fresh abdominal opening is formed
for the Müllerian duct.

II.
The Growth of the Müllerian Duct.
Although a great variety of views have been expressed by different
observers on the growth of the Müllerian duct, it is now fairly
generally admitted that it grows in the space between a portion of
the thickened germinal epithelium and the Wolffian duct, but quite
independently of both of them. Both Braun and Egli, who have
specially directed their attention to this point, have for Reptilia and
Mammalia fully confirmed the views of previous observers. We were,
nevertheless, induced, partly on account of the à priori difficulties of
this view, and partly by certain peculiar appearances which we
observed, to undertake the re-examination of this point, and have
found ourselves unable altogether to accept the general account. We
propose first describing, in as matter-of-fact a way as possible, the
actual observations we have made, and then stating what
conclusions we think may be drawn from these observations.
We have found it necessary to distinguish three stages in the growth
of the Müllerian duct. Our first stage embraces the period prior to
the disappearance of the head-kidney. At this stage the structure we
have already spoken of as the rudiment of the Müllerian duct
consists of a solid rod of cells, continuous with the third groove of
the head-kidney. It extends through a very few sections, and
terminates by a fine point of about two cells, wedged in between the
Wolffian duct and germinal epithelium (described above, Nos. 7-10,
series A, Plate 27).
In an embryo slightly older than the above, such as that from which
series B was taken, but still belonging to our first stage, a definite
lumen appears in the anterior part of the Müllerian duct, which
vanishes after a few sections. The duct terminates in a point which
lies in a concavity of the wall of the Wolffian duct (Plate 27, Nos. 1
and 2, series G). The limits of the Wolffian wall and the pointed
termination of the Müllerian duct are in many instances quite
distinct; but the outline of the Wolffian duct appears to be carried
round the Müllerian duct, and in some instances the terminal point
of the Müllerian duct seems almost to form an integral part of the
wall of the Wolffian duct.
The second of our stages corresponds with that in which the atrophy
of the head-kidney is nearly complete (series D and H, Plate 28).
The Müllerian duct has by this stage made a very marked progress
in its growth towards the cloaca, and, in contradistinction to the
earlier stage, a lumen is now continued close up to the terminal
point of the duct. In the two or three sections before it ends it
appears as a distinct oval mass of cells (No. 10, series D, and No. 1,
series H), without a lumen, lying between and touching the external
wall of the Wolffian duct on the one hand, and the germinal
epithelium on the other. It may either lie on the ventral side of the
Wolffian duct (series D), or on the outer side (series H), but in either
case is opposite the maximum thickening of that part of the germinal
epithelium which always accompanies the Müllerian duct in its
backward growth.
In the last section in which any trace of the Müllerian duct can be
made out (series D, No. 11, and series H, No. 2), it has no longer an
oval, well-defined contour, but appears to have completely fused
with the wall of the Wolffian duct, which is accordingly very thick,
and occupies the space which in the previous section was filled by its
own wall and the Müllerian duct. In the following section the
thickening in the wall of the Wolffian duct has disappeared (Plate 28,
series H, No. 3), and every trace of the Müllerian duct has vanished
from view. The Wolffian duct is on one side in contact with the
germinal epithelium.
The stage during which the condition above described lasts is not of
long duration, but is soon succeeded by our third stage, in which a
fresh mode of termination of the Müllerian duct is found. (Plate 28,
series I.) This last stage remains up to about the close of the sixth
day, beyond which our investigations do not extend.
A typical series of sections through the terminal part of the Müllerian
duct at this stage presents the following features:
A few sections before its termination the Müllerian duct appears as a
well-defined oval duct lying in contact with the wall of the Wolffian
duct on the one hand and the germinal epithelium on the other
(series I, No. 1). Gradually, however, as we pass backwards, the
Müllerian duct dilates; the external wall of the Wolffian duct
adjoining it becomes greatly thickened and pushed in in its middle
part, so as almost to touch the opposite wall of the duct, and so
form a bay in which the Müllerian duct lies (Plate 28, series I, Nos. 2
and 3). As soon as the Müllerian duct has come to lie in this bay its
walls lose their previous distinctness of outline, and the cells
composing them assume a curious vacuolated appearance. No well-
defined line of separation can any longer be traced between the
walls of the Wolffian duct and those of the Müllerian, but between
the two is a narrow clear space traversed by an irregular network of
fibres, in some of the meshes of which nuclei are present.
The Müllerian duct may be traced in this condition for a considerable
number of sections, the peculiar features above described becoming
more and more marked as its termination is approached. It
continues to dilate and attains a maximum size in the section or so
before it disappears. A lumen may be observed in it up to its very
end, but is usually irregular in outline and frequently traversed by
strands of protoplasm. The Müllerian duct finally terminates quite
suddenly (Plate 28, series I, No. 4), and in the section immediately
behind its termination the Wolffian duct assumes its normal
appearance, and the part of its outer wall on the level of the
Müllerian duct comes into contact with the germinal epithelium
(Plate 28, series I, No. 5).
We have traced the growing point of the Müllerian duct with the
above features till not far from the cloaca, but we have not followed
the last phases of its growth and its final opening into the cloaca.
In some of our embryos we have noticed certain rather peculiar
structures, an example of which is represented at y in fig. K, taken
from an embryo of 123 hours, in which all traces of the head-kidney
had disappeared. It consists of a cord of cells, connecting the
Wolffian duct and the hind end of the abdominal opening of the
Müllerian duct. At the least one similar cord was met with in the
same embryo, situated just behind the abdominal opening of the
Müllerian duct. We have found similar structures in other embryos of
about the same age, though never so well marked as in the embryo
from which fig. K is taken. We have quite failed to make out the
meaning, if any, of them.
Our interpretation of the appearances we have described in
connection with the growth of the Müllerian duct can be stated in a
very few words. Our second stage, where the solid point of the
Müllerian duct terminates by fusing with the walls of the Wolffian
duct, we interpret as meaning that the Müllerian is growing
backwards as a solid rod of cells, split off from the outer wall of the
Wolffian duct; in the same manner, in fact, as in Amphibia and
Elasmobranchii. The condition of the terminal part of the Müllerian
duct during our third stage cannot, we think, be interpreted in the
same way, but the peculiarities of the cells of both Müllerian and
Wolffian ducts, and the indistinctness of the outlines between them,
appear to indicate that the Müllerian duct grows by cells passing
from the Wolffian duct to it. In fact, although in a certain sense the
growth of the two ducts is independent, yet the actual cells which
assist in the growth of the Müllerian duct are, we believe, derived
from the walls of the Wolffian duct.

III.
General considerations.
The excretory system of a typical Vertebrate consists of the following
parts:—
1. A head-kidney with the characters already described.
2. A duct for the head-kidney—the segmental duct.
3. A posterior kidney—(Wolffian body, permanent kidney, &c. The
nature and relation of these parts we leave out of consideration, as
they have no bearing upon our present investigations). The primitive
duct for the Wolffian body is the segmental duct.
4. The segmental duct may become split into (a) a dorsal or inner
duct, which serves as ureter (in the widest sense of the word); and
(b) a ventral or outer duct, which has an opening into the body-
cavity, and serves as the generative duct for the female, or for both
sexes.
These parts exhibit considerable variations both in their structure
and development, into some of which it is necessary for us to enter.
The head-kidney[438] attains to its highest development in the
Marsipobranchii (Myxine, Bdellostoma). It consists of a longitudinal
canal, from the ventral side of which numerous tubules pass. These
tubules, after considerable subdivision, open by a large number of
apertures into the pericardial cavity. From the longitudinal canal a
few dorsal diverticula, provided with glomeruli, are given off. In the
young the longitudinal canal is continued into the segmental duct;
but this connection becomes lost in the adult. The head-kidney
remains, however, through life. In Teleostei and Ganoidei (?) the
head-kidney is generally believed to remain through life, as the
dilated cephalic portion of the kidneys when such is present. In
Petromyzon and Amphibia the head-kidney atrophies. In
Elasmobranchii the head-kidney, so far as is known, is absent.
The development of the segmental duct and head-kidney (when
present) is still more important for our purpose than their adult
structure.
In Myxine the development of these structures is not known. In
Amphibia and Teleostei it takes place upon the same type, viz. by
the conversion of a groove-like invagination of the peritoneal
epithelium into a canal open in front. The head-kidney is developed
from the anterior end of this canal, the opening of which remains in
Teleostei single and closes early in embryonic life, but becomes in
Amphibia divided into two, three, or four openings. In
Elasmobranchii the development is very different.
“The first trace of the urinary system makes its appearance as
a knob springing from the intermediate cell-mass opposite the
fifth protovertebra. This knob is the rudiment of the
abdominal opening of the segmental duct, and from it there
 grows backwards to the level of the anus a solid column of
cells, which constitutes the rudiment of the segmental duct
itself. The knob projects towards the epiblast, and the column
connected with it lies between the mesoblast and epiblast.
The knob and column do not long remain solid, but the
former acquires an opening into the body-cavity continuous
with a lumen, which makes its appearance in the latter.”
The difference in the development of the segmental duct in the two
types (Amphibia and Elasmobranchii) is very important. In the one
case a continuous groove of the peritoneal epithelium becomes
constricted into a canal, in the other a solid knob of cells is
continued into a rod, at first solid, which grows backwards without
any apparent relation to the peritoneal epithelium[439].
The abdominal aperture of the segmental duct in Elasmobranchii, in
that it becomes the permanent abdominal opening of the oviduct,
corresponds physiologically rather with the abdominal opening of the
Müllerian duct than with that of the segmental duct of Amphibia,
which, after becoming divided up to form the pores of the head-
kidney, undergoes atrophy. Morphologically, however, it appears to
correspond with the opening of the segmental duct in Amphibia. We
shall allude to this point more than once again, and give our grounds
for the above view on p. 640.
The development of the segmental duct in Elasmobranchii as a solid
rod is, we hope to shew, of special importance for the elucidation of
the excretory system of Aves.
The development of these parts of Petromyzon is not fully known,
but from W. Müller's account (Jenaische Zeitschrift, 1875) it would
seem that an anterior invagination of the peritoneal epithelium is
continued backwards as a duct (segmental duct), and that the
anterior opening subsequently becomes divided up into the various
apertures of the head-kidney. If this account is correct, Petromyzon
presents a type intermediate between Amphibia and Elasmobranchii.
In certain types, viz. Marsipobranchii and Teleostei, the segmental
duct becomes the duct for the posterior kidney (segmental tubes),
but otherwise undergoes no further differentiation. In the majority of
types, however, the case is different. In Amphibia[440], as has already
been mentioned, a solid rod of cells is split off from its ventral wall,
which afterwards becomes hollow, acquires an opening into the
body-cavity, and forms the Müllerian duct.
In Elasmobranchii the segmental duct undergoes a more or less
similar division. “It becomes longitudinally split into two complete
ducts in the female, and one complete duct and parts of a second in
the male. The resulting ducts are (1) the Wolffian duct dorsally,
which remains continuous with the excretory tubules of the kidney,
and ventrally (2) the oviduct or Müllerian duct in the female, and the
rudiments of this duct in the male. In the female the formation of
these ducts takes place by a nearly solid rod of cells, being gradually
split off from the ventral side of all but the foremost part of the
original segmental duct, with the short undivided anterior part of
which duct it is continuous in front. Into it a very small portion of the
lumen of the original segmental duct is perhaps continued. The
remainder of the segmental duct (after the loss of its anterior
section and the part split off from its ventral side) forms the Wolffian
duct. The process of formation of the ducts in the male chiefly
differs from that in the female, in the fact of the anterior undivided
part of the segmental duct, which forms the front end of the
Müllerian duct, being shorter, and in the column of cells with which it
is continuous being from the first incomplete.”
It will be seen from the above that the Müllerian duct consists of two
distinct parts—an anterior part with the abdominal opening, and a
posterior part split off from the segmental duct. This double
constitution of the Müllerian duct is of great importance for a proper
understanding of what takes place in the Bird.
The Müllerian duct appears therefore to develop in nearly the same
manner in the Amphibian and Elasmobranch type, as a solid or
nearly solid rod split off from the ventral wall of the segmental duct.
But there is one important difference concerning the abdominal
opening of the duct. In Amphibia this is a new formation, but in
Elasmobranchii it is the original opening of the segmental duct.
Although we admit that in a large number of points, including the
presence of a head-kidney, the urinogenital organs of Amphibia are
formed on a lower type than those of the Elasmobranchii, yet it
appears to us that this does not hold good for the development of
the Müllerian duct.
The above description will, we trust, be sufficient to render clear our
views upon the development of the excretory system in Aves.
In the bird the excretory system consists of the following parts
(using the ordinary nomenclature) which are developed in the order
below.
1. Wolffian duct. 2. Wolffian body. 3. Head-kidney. 4. Müllerian duct.
5. Permanent kidney and ureter.
About 2 and 5 we shall have nothing to say in the sequel.
We have already in the early part of the paper given an account of
the head-kidney and Müllerian duct, but it will be necessary for us to
say a few words about the development of the Wolffian duct (so
called). Without entering into the somewhat extended literature on
the subject, we may state that we consider that the recent paper of
Dr Gasser[441] supplies us with the best extant account of the
development of the Wolffian duct.
The first trace of it, which he finds, is visible in an embryo with eight
protovertebræ as a slight projection from the intermediate cell mass
towards the epiblast in the region of the three hindermost
protovertebræ. In the next stage, with eleven protovertebræ, the
solid rudiment of the duct extends from the fifth to the eleventh
protovertebra, from the eighth to the eleventh protovertebra it lies
between the epiblast and mesoblast, and is quite distinct from both,
and Dr Gasser distinctly states that in its growth backwards from the
eighth protovertebra the Wolffian duct never comes into continuity
with the adjacent layers.
In the region of the fifth protovertebra, where the duct was
originally continuous with the mesoblast, it has now become free,
but is still attached in the region of the sixth and to the eighth
protovertebra. In an embryo with fourteen protovertebræ the duct
extends from the fourth to the fourteenth protovertebra, and is now
free between epiblast and mesoblast for its whole extent. It is still
for the most part solid though perhaps a small lumen is present in its
middle part. In the succeeding stages the lumen of the duct
gradually extends backwards and forwards, the duct itself also
passes inwards till it acquires its final position close to the peritoneal
epithelium; at the same time its hind end elongates till it comes into
connection with the cloacal section of the hind-gut. It should be
noted that the duct in its backward growth does not appear to come
into continuity with the subjacent mesoblast, but behaves in this
respect exactly as does the segmental duct in Elasmobranchii (vide
note on p. 634).
The question which we propose to ourselves is the following:—What
are the homologies of the parts of the Avian urinogenital system
above enumerated? The Wolffian duct appears to us morphologically
to correspond in part to the segmental duct[442], or what Fürbringer
would call the duct of the head-kidney. This may seem a paradox,
since in birds it never comes into relation with the head-kidney.
Nevertheless we consider that this homology is morphologically
established, for the following reasons:—
(1) That the Wolffian duct gives rise (vide supra, p. 631) to the
Müllerian duct as well as to the duct of the Wolffian body. In this
respect it behaves precisely as does the segmental duct of
Elasmobranchii and Amphibia. That it serves as the duct for the
Wolffian body, before the Müllerian duct originates from it, is also in
accordance with what takes place in other types.
(2) That it develops in a strikingly similar manner to the segmental
duct of Elasmobranchii.
We stated expressly that the Wolffian duct corresponded only in part
to the segmental duct. It does not, in fact, in our opinion,
correspond to the whole segmental duct, but to the segmental duct
minus the anterior abdominal opening in Elasmobranchii, which
becomes the head-kidney in other types. In fact, we suppose that
the segmental duct and head-kidney, which in the Ichthyopsida
develop as a single formation, develop in the Bird as two distinct
structures—one of these known as the Wolffian duct, and the other
the head-kidney. If our view about the head-kidney is accepted the
above position will hardly require to be disputed, but we may point
out that the only feature in which the Wolffian duct of the Bird
differs in development from the segmental duct of Elasmobranchii is
in the absence of the knob, which forms the commencement of the
segmental duct, and in which the abdominal opening is formed; so
that the comparison of the development of the duct in the two types
confirms the view arrived at from other considerations.
The head-kidney and Müllerian duct in the Bird must be considered
together. The parts which they eventually give rise to after the
atrophy of the head-kidney have almost universally been regarded
as equivalent to the Müllerian duct of the Ichthyopsida. By Braun[443],
however, who from his researches on the Lizard satisfied himself of
the entire independence of the Müllerian and Wolffian ducts in the
Amniota, the Müllerian duct of these forms is regarded as a
completely new structure with no genetic relations to the Müllerian
duct of the Ichthyopsida. Semper[444], on the other hand, though he
accepts the homology of the Müllerian duct in the Ichthyopsida and
Amniota, is of opinion that the anterior part of the Müllerian duct in
the Amniota is really derived from the Wolffian duct, though he
apparently admits the independent growth of the posterior part of
the Müllerian duct. We have been led by our observations, as well as
by our theoretical deductions, to adopt a view exactly the reverse of
that of Professor Semper. We believe that the anterior part of the
Müllerian duct of Aves, which is at first the head-kidney, and
subsequently becomes the abdominal opening of the duct, is
developed from the peritoneal epithelium independently of all other
parts of the excretory system; but that the posterior part of the duct
is more or less completely derived from the walls of the Wolffian
duct. This view is clearly in accordance with our account of the facts
of development in Aves, and it fits in very well with the development
of the Müllerian duct in Elasmobranchii. We have already pointed out
that in Elasmobranchii the Müllerian duct is formed of two factors—
(1) of the whole anterior extremity of the segmental duct, including
its abdominal opening; (2) of a rod split off from the ventral side of
the segmental duct. In Birds the anterior part (corresponding to
factor No. 1) of the Müllerian duct has a different origin from the
remainder; so that if the development of the posterior part of the
duct (factor No. 2) were to proceed in the same manner in Birds and
Elasmobranchii, it ought to be formed at the expense of the Wolffian
(i.e. segmental) duct, though in connection anteriorly with the head-
kidney. And this is what actually appears to take place.
So far the homologies of the avian excretory system are fairly clear;
but there are still some points which have to be dealt with in
connection with the permanent opening of the Müllerian duct, and
the relatively posterior position of the head-kidney. With reference to
the first of these points the facts of the case are the following:—
In Amphibia the permanent opening of the Müllerian duct is formed
as an independent opening after the atrophy of the head-kidney.
In Elasmobranchii the original opening of the segmental duct forms
the permanent opening of the Müllerian duct and no head-kidney
appears to be formed.
In Birds the anterior of the three openings of the head-kidney
remains as the permanent opening of the Müllerian duct.
With reference to the difficulties involved in there being apparently
three different modes in which the permanent opening of the
Müllerian duct is formed, we would suggest the following
considerations:
The history of the development of the excretory system teaches us
that primitively the segmental duct must have served as efferent
duct both for the generative products and kidney secretion (just as
the Wolffian duct still does for the testicular products and secretion
of the Wolffian body in Elasmobranchii and Amphibia); and further,
that at first the generative products entered the segmental duct
from the abdominal cavity by one or more of the abdominal
openings of the kidney (almost certainly of the head-kidney). That
the generative products did not enter the segmental duct at first by
an opening specially developed for them appears to us to follow
from Dohrn's principle of the transmutation of function
(Functionswechsel). As a consequence (by a process of natural
selection) of the segmental duct having both a generative and a
urinary function, a further differentiation took place, by which that
duct became split into two—a ventral Müllerian duct and dorsal
Wolffian duct.
The Müllerian duct without doubt was continuous with the head-
kidney, and so with the abdominal opening or openings of the head-
kidney which served as generative pores. At first the segmental duct
was probably split longitudinally into two equal portions, but the
generative function of the Müllerian duct gradually impressed itself
more and more upon the embryonic development, so that, in the
course of time, the Müllerian duct developed less and less at the
expense of the Wolffian duct. This process appears partly to have
taken place in Elasmobranchii, and still more in Amphibia; the
Amphibia offering in this respect a less primitive condition than
Elasmobranchii; while in Aves it has been carried even further. The
abdominal opening no doubt also became specialised. At first it is
quite possible that more than one abdominal pore may have served
for the generative products; one of which, no doubt, eventually
came to function alone. In Amphibia the specialisation of the
opening appears to have gone so far that it no longer has any
relation to the head-kidney, and even develops after the atrophy of
the head-kidney. In Elasmobranchii, on the other hand, the
functional opening appears at a period when we should expect the
head-kidney to develop. This state is very possibly the result of a
differentiation (along a different line to that in Amphibia) by which
the head-kidney gradually ceased to become developed, but by
which the primitive opening (which in the development of the head-
kidney used to be divided into several pores leading into the body-
cavity) remained undivided and served as the abdominal aperture of
the Müllerian duct. Aves, finally, appear to have become
differentiated along a third line; since in their ancestors the anterior
pore of the head-kidney appears to have become specialised as the
permanent opening of the Müllerian duct.
With reference to the posterior position of the head-kidney in Aves
we have only to remark, that a change in position of the head-
kidney might easily take place after it acquired an independent
development. The fact that it is slightly behind the glomerulus would
seem to indicate, on the one hand, that it has already ceased to be
of any functional importance; and, on the other, that the shifting has
been due to its having a connection with the Müllerian duct.
We have made a few observations on the development of the
Müllerian duct in Lacerta muralis, which have unfortunately led us to
no decided conclusions. In a fairly young stage in the development
of the Müllerian duct (the youngest we have met with), no trace of a
head-kidney could be observed, but the character of the abdominal
opening of the Müllerian duct was very similar to that figured by
Braun[445]. As to the backward growth of the Müllerian duct, we can
only state that the solid point of the duct in the young stages is in
contact with the wall of the Wolffian duct, and the relation between
the two is rather like that figured by Fürbringer (Pl. 1, figs. 14-15) in
Amphibia.

DESCRIPTION OF PLATES 27 AND 28.

Complete List of Reference Letters.
ao. Aorta. cv. Cardinal vein. gl. Glomerulus. gr1. First groove of head-kidney. gr2.
Second groove of head-kidney. gr3. Third groove of head-kidney. ge. Germinal
epithelium. mrb. Malpighian body. me. Mesentery. md. Müllerian duct. r1. First
ridge of head-kidney. r2. Second ridge of head-kidney. r3. Third ridge of head-
kidney. Wd. Wolffian duct. x. Fold in germinal epithelium.

Plate 27.

Series A. Sections through the head-kidney at our second stage. Zeiss 2, ocul. 3
(reduced one-third). The second and third grooves are represented with the ridge
connecting them, and the rod of cells running backwards for a short distance.
No. 1. Section through the second groove.
No. 2. Section through the ridge connecting the second and third grooves.
No. 3. Section passing through the same ridge at a point nearer the third groove.
Nos. 4, 5, 6. Sections through the third groove.
No. 7. Section through the point where the third groove passes into the solid rod
of cells.
No. 8. Section through the rod when quite separated from the germinal
epithelium.
No. 9. Section very near the termination of the rod.

No. 10. Last section in which any trace of the rod is seen.

Series B. Sections passing through the head-kidney at our third stage. Zeiss C,
ocul. 2. Our figures are representations of the following sections of the series,
section 1 being the first which passes through the anterior groove of the head-
kidney.

No. 1 Section 3.
" 2 " 4.
" 3 " 5.
" 4 " 6.
" 5 " 8.
" 6 " 10.
" 7 " 11.
" 8 " 13.
 " 9 " 15.
" 10 " 16.
" 11 " 17.
" 12 " 18.
" 13 " 19.
" 14 " 20.

The Müllerian duct extends through eleven more sections.

The first groove (gr1.) extends to No. 3.

The second groove (gr2.) extends from No. 4 to No. 7.

The third groove (gr3.) extends from No. 11 to No. 13.

The first ridge (r1.) extends from No. 2 to No. 5.

The second ridge (r2.) extends from No. 8 to No. 11.

The third ridge (r3.) extends from No. 13 backwards through twelve sections,
when it terminates by a pointed extremity.

Fig. C. Section through the ridge connecting the second and third grooves of the
head-kidney of an embryo slightly younger than that from which Series B was
taken. Zeiss C, ocul. 3 (reduced one-third).
The fold of the germinal epithelium, which gives rise to a deep groove (x.)
external to the head-kidney is well marked.

Series G. Sections through the rod of cells constituting the termination of the
Müllerian duct at a stage in which the head-kidney is still present. Zeiss C, ocul. 2.

Plate 28.

Series D. Sections chosen at intervals from a complete series traversing the

peritoneal opening of the Müllerian duct, the remnant of the head-kidney, and the
termination of the Müllerian duct. Zeiss C, ocul. 3 (reduced one-third).
Nos. 1 and 2. Sections through the persistent anterior opening of the head-kidney
(abdominal opening of Müllerian duct). The approach of the Wolffian duct to the
groove may be seen by a comparison of these two figures. In the sections in front
of these (not figured) the two are much more widely separated than in No. 1.
No. 3. Section through the Müllerian duct, just posterior to the persistent opening.
Nos. 4 and 5. Remains of the ridges, which at an earlier stage connected the first
and second grooves, are seen passing from the Müllerian duct to the peritoneal
epithelium.
No. 6. Rudiment of the second groove (gr2.) of the head-kidney.

Between 6 and 7 is a considerable interval.

No. 7. All traces of this groove (gr2.) have vanished, and the Müllerian duct is quite
disconnected from the epithelium.
No. 8. Rudiment of the third groove (gr3.).

No. 9. Müllerian duct quite free in the space between the peritoneal epithelium
and the Wolffian duct, in which condition it extends until near its termination.
Between Nos. 9 and 10 is an interval of eight sections.
No. 10. The penultimate section, in which the Müllerian duct is seen. A lumen
cannot be clearly made out.
No. 11. The last section in which any trace of the Müllerian duct is visible. No line
of demarcation can be seen separating the solid end of the Müllerian duct from the
ventral wall of the Wolffian duct.

Figs. E. and F. Sections through the glomerulus of the head-kidney from an

embryo prior to the appearance of the head-kidney. Zeiss B, ocul. 2. A comparison
of the two figures shows the variation in the thickness of the stalk of the
glomerulus. E. Section anterior to the foremost Malpighian body. F. Section
through both the glomerulus of the head-kidney and that of a Malpighian body.
The two are seen to be connected.

Series H. Consecutive sections through the hind end of the Müllerian duct, from an
embryo in which the head-kidney was only represented by a rudiment. (The
embryo was, perhaps, very slightly older than that from which Series D was
taken.) Zeiss C, ocul. 3 (reduced one-third).
No. 1. Müllerian duct is without a lumen, and quite distinct from the Wolffian wall.
No. 2. The solid end of the Müllerian duct is no longer distinct from the internal
wall of the Wolffian duct.
No. 3. All trace of the Müllerian duct has vanished.

Series I. Sections through the hinder end of the Müllerian duct from an embryo of
about the middle of the sixth day. Zeiss C, ocul. 2 (reduced one-third).
No. 1. The Müllerian duct is distinct and small.
No. 2. Is posterior by twelve sections to No. 1. The Müllerian duct is dilated, and
its cells are vacuolated.
No. 3. Penultimate section, in which the Müllerian duct is visible; it is separated by
three sections from No. 2.
No. 4. Last section in which any trace of the Müllerian duct is visible; the lumen,
which was visible in the previous section, is now absent.
No. 5. No trace of Müllerian duct. Nos. 3, 4, and 5 are consecutive sections.

Fig. K. Section through the hind end of the abdominal opening of the Müllerian
duct of a chick of 123 hours. Zeiss C, ocul. 2 (reduced one-third). It illustrates the
peculiar cord connecting the Müllerian and Wolffian ducts.

[424] From the Quarterly Journal of Microscopical Science, Vol. XIX.

1879.
[425] “Das Urogenital-System der Plagiostomen,” Arbeiten a. d.
zool.-zoot. Institut. Würzburg.
[426] “Zur vergl. Anat. u. Entwick. d. Excretionsorgane d.
Vertebraten,” Morphologisches Jahrbuch, Vol. IV.
[427] “On the Origin and History of the Urinogenital Organs of
Vertebrates,” Journal of Anat. and Phys., Vol. X. [This Edition No. VII.]
[428] Arbeiten a. d. zool.-zoot. Institut. Würzburg, Vol. IV.

[429] Beitr. zur Anat. u. Entwick. d. Geschlechtsorgane, Inaug. Diss.,

Zürich, 1876.
[430] Proceedings of Royal Society, 1878.
[431] A deep focus of the rather thick section represented in No. 3
shewed the body much more nearly in the position it occupies in No.
4.
[432] Beiträge zur Entwicklungsgeschichte d. Allantois der
Müller'schen Gange u. des Afters. Frankfurt, 1874.
[433] Loc. cit.
[434] Jenaische Zeitschrift, Vol. IX. 1875.
[435] Entwicklungsgeschichte d. Unke.
[436] Loc. cit.
[437] Balfour, “Origin and History of Urinogenital Organs of
Vertebrates,” Journal of Anat. and Phys. Vol. X., and Monograph on
Elasmobranch Fishes. [This edition Nos. VII. and X.]
[438] I am inclined to give up the view I formerly expressed with
reference to the head-kidney and segmental duct, viz. “that they
were to be regarded as the most anterior segmental tube, the
peritoneal opening of which had become divided, and which had
become prolonged backwards so as to serve as the duct for the
posterior segmental tubes,” and provisionally to accept the
Gegenbaur-Fürbringer view which has been fully worked out and
ably argued for by Fürbringer (loc. cit. p. 96). According to this view
the head-kidney and its duct are to be looked on as the primitive and
unsegmented part of the excretory system, more or less similar to
the excretory system of many Trematodes and unsegmented
Vermes. The segmental tubes I regard as a truly segmental part of
the excretory system acquired subsequently.—F. M. B.
[439] In a note on p. 50 of his memoir Fürbringer criticises my
description of the mode of growth of the segmental duct. The
following is a free translation of what he says: “In Balfour's, as in
other descriptions, an account is given of a backward growth, which
easily leads to the supposition of a structure formed anteriorly
forcing its way through the tissues behind. This is, however, not the
case, since, to my knowledge, no author has ever detected a sharp
boundary between the growing point of the segmental duct (or
Müllerian duct) and the surrounding tissues.” He goes on to say that
“the growth in these cases really takes place by a differentiation of
tissue along a line in the region of the peritoneal cavity.” Although I
fully admit that it would be far easier to homologise the development
of the segmental duct in Amphibia and Elasmobranchii according to
this view, I must nevertheless vindicate the accuracy of my original
account. I have looked over my specimens again, since the
appearance of Dr Fürbringer's paper, and can find no evidence of the
end of the duct becoming continuous with the adjoining mesoblastic
tissues. In the section, before its disappearance, the segmental duct
may, so far as I can make out, be seen as a very small but distinct
rod, which is much more closely connected with the epiblast than
with any other layer. From Gasser's observations on the Wolffian duct
in the bird, I am led to conclude that it behaves in the same way as
the segmental duct in the Elasmobranchii. I will not deny that it is
possible that the growth of the duct takes place by wandering cells,
but on this point I have no evidence, and must therefore leave the
question an open one.—F. M. B.
[440] Fürbringer, loc. cit.
[441] Arch. für Mic. Anat. Vol. XIV.

[442] The views here expressed about the Wolffian duct are nearly
though not exactly those which one of us previously put forward
(“Urinogenital Organs of Vertebrates,” &c., pp. 45-46) [This edition,
pp. 164, 165], and with which Fürbringer appears exactly to agree.
Possibly Dr Fürbringer would alter his view on this point were he to
accept the facts we believe ourselves to have discovered. Semper's
view also differs from ours, in that he believes the Wolffian duct to
correspond in its entirety with the segmental duct.
[443] “Urogenital-System d. Reptilien,” Arb. aus d. zool.-zoot. Inst.
Würzburg, Vol. IV.
[444] Loc. cit.
[445] Loc. cit.
 XIV. On the Early Development of the Lacertilia,
together with some Observations on the Nature and
Relations of the Primitive Streak[446].
(With Plate 29.)

Till quite recently no observations were recorded on the early

developmental changes of the reptilian ovum. Not long ago
Professors Kupffer and Benecke published a preliminary note on the
early development of Lacerta agilis and Emys Europea[447]. I have
myself also been able to make some observations on the embryo of
Lacerta muralis. The number of my embryos has been somewhat
limited, and most of those which I have had have been preserved in
bichromate of potash, which has turned out a far from satisfactory
hardening reagent. In spite of these difficulties I have been led on
some points to very different results from those of the German
investigators, and to results which are more in accordance with what
we know of other Sauropsidan types. I commence with a short
account of the results of Kupffer and Benecke.
Segmentation takes place exactly as in birds, and the resulting
blastoderm, which is thickened at its edge, spreads rapidly over the
yolk. Shortly before the yolk is half enclosed a small embryonic
shield (area pellucida) makes its appearance in the centre of the
blastoderm, which has, in the meantime, become divided into two
layers. The upper of these is the epiblast, and the lower the
hypoblast. The embryonic shield is mainly distinguished from the
remainder of the blastoderm by the more columnar character of its
constituent epiblast cells. It is somewhat pyriform in shape, the
narrower end corresponding with the future posterior end of the
embryo. At the narrow end an invagination takes place, which gives
rise to an open sac, the blind end of which is directed forwards. The
opening of this sac is regarded by the authors as the blastopore. A
linear thickening of epiblast arises in front of the blastopore, along
the median line of which the medullary groove soon appears. In the
caudal region the medullary folds spread out and enclose between
them the blastopore, behind which they soon meet again. On the
conversion of the medullary groove into a closed canal the
blastopore becomes obliterated. The mesoblast grows out from the
lip of the blastopore as four masses. Two of these are lateral: a third
is anterior and median, and, although at first independent of the
epiblast, soon attaches itself to it, and forms with it a kind of axis-
cord. A fourth mass applied itself to the walls of the sac formed by
invagination.
With reference to the very first developmental phenomena my
observations are confined to two stages during the segmentation[448].
In the earliest of these the segmentation was about half completed,
in the later one it was nearly over. My observations on these stages
bear out generally the statements of Kupffer and Benecke. In the
second of them the blastoderm was already imperfectly divided into
two layers—a superficial epiblastic layer formed of a single row of
cells, and a layer below this several rows deep. Below this layer
fresh segments were obviously being added to the blastoderm from
the subjacent yolk.
Between the second of these blastoderms and my next stage there
is a considerable gap. The medullary plate is just established, and is
marked by a shallow groove which becomes deeper in front. A
section through the embryo is represented in Pl. 29, Series A, fig. 1.
In this figure there may be seen the thickened medullary plate with
a shallow medullary groove, below which are two independent plates
of mesoblast (me.p.), one on each side of the middle line, very
imperfectly divided into somatopleuric and splanchnopleuric layers.
Below the mesoblast is a continuous layer of hypoblast (hy.), which
develops a rod-like thickening along the axial line (ch.). This rod
becomes in the next stage the notochord. Although this embryo is
not well preserved I feel very confident in asserting the continuity of
the notochord with the hypoblast at this stage.
At the hind end of the embryo is placed a thickened ridge of tissue
which continues the embryonic axis. In this ridge all the layers
coalesce, and I therefore take it to be equivalent to the primitive
streak of the avian blastoderm. It is somewhat triangular in shape,
with the apex directed backward, the broad base placed in front.
At the junction between the primitive streak and the blastoderm is
situated a passage, open at both extremities, leading from the upper
surface of the blastoderm obliquely forwards to the lower.
The dorsal and anterior wall of this passage is formed of a distinct
epithelial layer, continuous at its upper extremity with the epiblast,
and at its lower with the notochordal plate, so that it forms a layer
of cells connecting together the epiblast and hypoblast. The hinder
and lower wall of the passage is formed by the cells of the primitive
streak, which only assume a columnar form near the dorsal opening
of the passage (vide fig. 4). This passage is clearly the blind sac of
Kupffer and Benecke, who, if I am not mistaken, have overlooked its
lower opening. As I hope to show in the sequel, it is also the
equivalent of the neurenteric passage, which connects the neural
and alimentary canals in the Ichthyopsida, and therefore represents
the blastopore of Amphioxus, Amphibians, &c.
Series A, figs. 2, 3, 4, 5, illustrate the features of the passage and its
relation to the embryo.
Fig. 2 passes through the ventral opening of the passage. The
notochordal plate (ch´.) is vaulted over the opening, and on the left
side is continuous with the mesoblast as well as the hypoblast. Figs.
3 and 4 are taken through the middle part of the passage (ne.),
which is bounded above by a continuation of the notochordal plate,
and below by the tissue of the primitive streak. The hypoblast (hy.),
in the middle line, is imperfectly fused with the mesoblast of the
primitive streak, which is now continuous across the middle line. The
medullary groove has disappeared, but the medullary plate (mp.) is
quite distinct.
In fig. 5 is seen the dorsal opening of the passage (ne.). If a section
behind this had been figured, as is done for the next series (B), it
would have passed through the primitive streak, and, as in the chick,
all the layers would have been fused together. The epiblast in the
primitive streak completely coalesces with the mesoblast; but the
hypoblast, though attached to the other layers in the middle line,
can always be traced as a distinct stratum.
Fig. B is a surface view of my next oldest embryo. The medullary
groove has become much deeper, especially in front. Behind it
widens out to form a space equivalent to the sinus rhomboidalis of
the embryo bird. The amnion forms a small fold covering over the
cephalic extremity of the embryo, which is deeply embedded in the
yolk. Some somites (protovertebræ) were probably present, but this
could not be made out in the opaque embryo.
 Fig. 1. Diagrammatic longitudinal section of an embryo of Lacerta.
pp. body-cavity. am. Amnion. ne. Neurenteric canal. ch. Notochord.
hy. Hypoblast. ep. Epiblast. pr. Primitive streak.

The woodcut (fig. 1) represents a diagrammatic longitudinal section

through this embryo, and the sections belonging to Series B illustrate
the features of the hind end of the embryo and of the primitive
streak.
As is shown in fig. 1, the notochord (ch.) has now throughout the
region of the embryo become separated from the subjacent
hypoblast, and the lateral plates of mesoblast are distinctly divided
into somatic and splanchnic layers. The medullary groove is
continued as a deepish groove up to the opening of the neurenteric
passage, which thus forms a perforation in the floor of the hinder
end of the medullary groove (vide Series B, figs. 2, 3, and 4).
The passage itself is somewhat shorter than in the previous stage,
and the whole of it is shown in a single section (fig. 4). This section
must either have been taken somewhat obliquely, or else the
passage have been exceptionally short in this embryo, since in an
older embryo it could not all be seen in one section.
The front wall of the passage is continuous with the notochord,
which for two sections or so in front remains attached to the
hypoblast (figs. 2 and 3). Behind the perforation in the floor of the
medullary groove is placed the primitive streak (fig. 5), where all the
layers become fused together, as in the earlier stage. Into this part a
narrow diverticulum from the end of the medullary groove is
continued for a very short distance (vide fig. 5, mc.).
The general features of the stage will best be understood by an
examination of the diagrammatic longitudinal section, represented in
woodcut, fig. 1. In front is shown the amnion (am.), growing over
the head of the embryo. The notochord (ch.) is seen as an
independent cord for the greater part of the length of the embryo,
but falls into the hypoblast shortly in front of the neurenteric
passage. The neurenteric passage is shown at ne., and behind it is
shown the primitive streak.
In a still older stage, represented in surface view on Pl. 29, fig. C,
the medullary folds have nearly met above, but have not yet united.
The features of the passage from the neural groove to the hypoblast
are precisely the same in the embryo just described, although the
lumen of the passage has become somewhat narrower. There is still
a short primitive streak behind the embryo.
The neurenteric passage persists but a very short time after the
complete closure of the medullary canal. It is in no way connected
with the allantois, as conjectured by Kupffer and Benecke, but the
allantois is formed, as I have satisfied myself by longitudinal sections
of a later stage, in the manner already described by Dobrynin,
Gasser, and Kölliker for the bird and mammal.
The general results of Kupffer's and Benecke's observations, with the
modifications introduced by my own observations, are as follows:—
After the segmentation and the formation of the embryonic shield
(area pellucida) the blastoderm becomes distinctly divided into
epiblast and hypoblast[449]. At the hind end of the shield a somewhat
triangular primitive streak is formed by the fusion of the epiblast and
hypoblast with a number of cells between them, which are probably
derived from the lower rows of the segmentation cells. At the front
end of the streak a passage arises, open at both extremities, leading
obliquely forwards through the epiblast to the space below the
hypoblast. The walls of the passage are formed of a layer of
columnar cells continuous both with epiblast and hypoblast. In front
of the primitive streak the body of the embryo becomes first
differentiated by the formation of a medullary plate, and at the same
time there grows out from the primitive streak a layer of mesoblast,
which spreads out in all directions between the epiblast and
hypoblast. In the axis of the embryo the mesoblast plate is stated by
Kupffer and Benecke to be continuous across the middle line, but
this appears very improbable. In a slightly later stage the medullary
plate becomes marked by a shallow groove, and the mesoblast of
the embryo is then undoubtedly constituted of two lateral plates,
one on each side of the median line. In the median line the
notochord arises as a ridge-like thickening of the hypoblast, which
becomes very soon quite separated from the hypoblast, except at
the hind end, where it is continued into the front wall of the
neurenteric passage. It is interesting to notice the remarkable
relation of the notochord to the walls of the neurenteric passage.
More or less similar relations are also well marked in the case of the
goose and the fowl (Gasser)[450], and support the conclusion
deducible from the lower forms of vertebrata, that the notochord is
essentially hypoblastic.
The passage at the front end of the primitive streak forms the
posterior boundary of the medullary plate, though the medullary
groove is not at first continued back to it. The anterior wall of this
passage connects together the medullary plate and the notochordal
ridge of the hypoblast. In the succeeding stages the medullary
groove becomes continued back to the opening of the passage,
which then becomes enclosed in the medullary folds, and forms a
true neurenteric passage. It becomes narrowed as the medullary
folds finally unite to form the medullary canal, and eventually
disappears.
I conclude this paper with a concise statement of what appears to
me the probable nature of the much-disputed organ, the primitive
streak, and of the arguments in support of my view.
In a paper on the primitive streak in the Quart. Journ. of Mic. Sci., in
1873 (p. 280) [This edition, p. 45], I made the following statement
with reference to this subject:—“It is clear, therefore, that the
primitive groove must be the rudiment of some ancestral feature....
It is just possible that it is the last trace of that involution of the
epiblast by which the hypoblast is formed in most of the lower
animals.”
At a later period, in July, 1876, after studying the development of
Elasmobranch fishes, I enlarged the hypothesis in a review of the
first part of Prof. Kölliker's Entwicklungsgeschichte. The following is
the passage in which I speak of it[451]:
"In treating of the exact relation of the primitive groove to the
formation of the embryo, Professor Kölliker gives it as his
view that though the head of the embryo is formed
independently of the primitive groove, and only secondarily
unites with this, yet that the remainder of the body is without
doubt derived from the primitive groove. With this conclusion
we cannot agree, and the very descriptions of Professor
Kölliker appear to us to demonstrate the untenable nature of
his results. We believe that the front end of the primitive
groove at first occupies the position eventually filled by about
the third pair of protovertebræ, but that as the protovertebræ
are successively formed, and the body of the embryo grows
in length, the primitive groove is carried further and further
back, so as always to be situated immediately behind the
embryo. As Professor Kölliker himself has shewn it may still
be seen in this position even later than the fortieth hour of
incubation.
"Throughout the whole period of its existence it retains a
character which at once distinguishes it in sections from the
medullary groove.
"Beneath it the epiblast and mesoblast are always fused,
though they are always separate elsewhere; this fact, which
was originally shewn by ourselves, has been very clearly
brought out by Professor Kölliker's observations.
"The features of the primitive groove which throw special
light on its meaning are the following:—
"(1) It does not enter directly into the formation of the
embryo.
"(2) The epiblast and mesoblast always become fused
beneath it.
"(3) It is situated immediately behind the embryo.
"Professor Kölliker does not enter into any speculations as to
the meaning of the primitive groove, but the above-
mentioned facts appear to us clearly to prove that the
primitive groove is a rudimentary structure, the origin of
which can only be completely elucidated by a knowledge of
the development of the Avian ancestors.
"In comparing the blastoderm of a bird with that of any
anamniotic vertebrate, we are met at the threshold of our
investigations by a remarkable difference between the two.
Whereas in all the lower vertebrates the embryo is situated at
the edge of the blastoderm, it is in birds and mammals
situated in the centre. This difference of position at once
suggests the view that the primitive groove may be in some
way connected with the change of position in the blastoderm
which the ancestors of birds must have undergone. If we
carry our investigations amongst the lower vertebrates a little
further, we find that the Elasmobranch embryo occupies at
first the normal position at the edge of the blastoderm, but
that in the course of development the blastoderm grows
round the yolk far more slowly in the region of the embryo
than elsewhere. Owing to this, the embryo becomes left in a
bay, the two sides of which eventually meet and coalesce in a
linear fashion immediately behind the embryo, thus removing
the embryo from the edge of the blastoderm and forming
behind it a linear streak not unlike the primitive streak. We
would suggest the hypothesis that the primitive groove is a
rudiment which gives the last indication of a change made by
the Avian ancestors in their position in the blastoderm, like
that made by Elasmobranch embryos when removed from the
edge of the blastoderm and placed in a central situation
similar to that of the embryo bird. On this hypothesis the
situation of the primitive groove immediately behind the
embryo, as well as the fact of its not becoming converted into
any embryonic organ would be explained. The central groove
might probably also be viewed as the groove naturally left
between the coalescing edges of the blastoderm.
“Would the fusion of epiblast and mesoblast also receive its
explanation on this hypothesis? We are of opinion that it
would. At the edge of the blastoderm which represents the
blastopore mouth of Amphioxus all the layers become fused
together in the anamniotic vertebrates. So that if the primitive
groove is in reality a rudiment of the coalesced edges of the
blastoderm, we might naturally expect the layers to be fused
there, and the difficulty presented by the present condition of
the primitive groove would rather be that the hypoblast is not
fused with the other layers than that the mesoblast is
indissolubly united with the epiblast. The fact that the
hypoblast is not fused with the other layers does not appear
to us to be fatal to our hypothesis, and in Mammalia, where
the primitive and medullary grooves present precisely the
same relations as in birds, all three layers are, according to
Hensen's account, fused together. This, however, is denied by
Kölliker, who states that in Mammals, as in Birds, only the
epiblast and mesoblast fuse together. Our hypothesis as to
the origin of the primitive groove appears to explain in a fairly
satisfactory manner all the peculiarities of this very
enigmatical organ; it also relieves us from the necessity of
accepting Professor Kölliker's explanation of the development
of the mesoblast, though it does not, of course, render that
explanation in any way untenable.”
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