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Introduction to Quantum Field Theory
with Applications to Quantum Gravity
 Introduction to Quantum Field Theory
with Applications to Quantum Gravity

Iosif L. Buchbinder
Department of Theoretical Physics, Tomsk State Pedagogical University, Tomsk, 634061,
Russia

Ilya L. Shapiro
Departamento de Fı́sica – Instituto Ciências Exatas, Universidade Federal de Juiz de Fora,
Juiz de Fora, CEP 36036-330, MG, Brazil

1
 3
Great Clarendon Street, Oxford, OX2 6DP,
United Kingdom
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c Iosif. L. Buchbinder and Ilya Shapiro, 2021
The moral rights of the author have been asserted
First Edition published in 2021
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DOI: 10.1093/oso/9780198838319.001.0001
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for information only. Oxford disclaims any responsibility for the materials
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Preface

For many decades, quantum field theory has played an important role in the successful
description of the interactions of elementary particles. Besides, this area of theoretical
physics has been always important due to the exchange of new ideas and methods with
other branches of physics, such as statistical mechanics, condensed matter physics,
gravitational physics, and cosmology. The last applications are becoming more impor-
tant nowadays, especially because the amount of experimental and observational data
demonstrates a fast growth and requires more detailed and reliable theoretical back-
ground. One of the most evident examples is the study of dark energy. Every few years,
the estimates of its equation of state (EoS) become more precise and it can not be
ruled out that, at some point, the EoS of the cosmological constant may be excluded
from the list of phenomenologically acceptable possibilities. Does this necessarily mean
that there is some special fluid (quintessence or alike) in the Universe? Or that the
situation can be explained by the variable cosmological constant, e.g., some quantum
effects? This is a phenomenologically relevant question, which should be answered at
some point. On the other hand, this is a theoretical question, that can be answered
only within a correctly formulated framework of quantum or semiclassical gravity.
In gravitational theory, general relativity represents a successful theory of relativis-
tic gravitational phenomena, confirmed by various experiments in the laboratories and
astronomical observations. Starting from the seventies and eighties, there has been a
growing interest in the idea of the unification of all fundamental forces, including
electroweak and strong interactions. Also, there is a general understanding that the
final theory should also include gravitation. An important component of such unifi-
cation is the demand for a quantum description of the gravitational field itself or, at
least, a consistent formulation of the quantum theory of matter fields on the classical
gravitational background, called semiclassical gravity.
The application of quantum field theory methods to gravitational physics, in both
semiclassical and full quantum frameworks, requires a careful formulation of the fun-
damental base of quantum theory, with special attention to such important issues as
renormalization, the quantum theory of gauge theories and especially effective action
formalism. The existing literature on these subjects includes numerous review papers
and also many books, e.g., [172, 56, 80, 150, 199, 240]. At the same time, the experience
of the present authors, after giving many courses on the subject worldwide, shows that
there is a real need to have a textbook with a more elementary introduction to the
subject. This situation was one of the main motivations for writing this book which
ended up being much longer than originally planned.
The textbook consists of two parts. Part I is based on the one-semester course
given by I.B. in many places, including the Tomsk State Pedagogical University and
the Federal University of Juiz de Fora. It includes a detailed introduction to the general
vi Preface

methods of quantum field theory, which are relevant for quantum gravity, including its
semiclassical part. Part II is mainly based on the one-semester course given regularly
by I.Sh. in the Federal University of Juiz de Fora and on the numerous mini-courses
in many countries. We did not pretend to do the impossible, that is, produce a com-
prehensive course of quantum field theory or quantum gravity. Instead, our purpose
was to give a sufficiently detailed introduction to the fundamental, basic notions and
methods, which would enable the interested reader to understand at least part of the
current literature on the subject and, in some cases, start original research work.
It is a pleasure for us to acknowledge the collaborations on various subjects dis-
cussed in this book with M. Asorey, R. Balbinot, E.V. Gorbar, A. Fabbri, J.C. Fab-
ris, J.-A. Helaël-Neto, P.M. Lavrov, T.P. Netto, S.D. Odintsov, F.O. Salles and A.A.
Starobinsky. We would like also to thank many colleagues, especially A.O. Barvin-
sky, A.S. Belyaev, E.S. Fradkin, V.P. Frolov, S.J. Gates, E.A. Ivanov, D.I. Kazakov,
S.M. Kuzenko, O. Lechnetfeld, H. Osborn, B.A. Ovrut, N.G. Pletnev, K. Stelle, A.A.
Tseytlin, I.V. Tyutin, and G.A. Vilkovisky for fruitful discussions of the problems of
quantum field theory.
We are grateful to Guilherme H.S. Camargo, Eduardo A. dos Reis, and especially
to Wagno Cesar e Silva for communicating to us misprints and corrections; and also
to Andreza R. Rodrigues and Yackelin Z. R. López for typing certain parts of the
manuscript, and to Vadim Zyubanov for valuable technical assistance.
The main work on Part I of the book was done during the long-term visit of J.B.
to the Federal University of Juiz de Fora (UFJF). The authors are grateful to UFJF
and especially to the Physics Department for providing both kind hospitality and the
conditions for productive work during this visit. Throughout the preparations of the
manuscript, the work of the authors has been supported by a special project APQ-
01205-16 from the Fundação de Amparo á Pesquisa de Minas Gerais (FAPEMIG). On
the top of that, the scientific activity of I.Sh. was partially supported by the Conselho
Nacional de Desenvolvimento Cientı́fico e Tecnológico (CNPq/Brazil). The authors
are also grateful to the Russian Ministry of Science and High Education and Russian
Foundation for Basic Research for their long-term support of the Center of Theoretical
Physics at the Tomsk State Pedagogical University.
Contents

PART I INTRODUCTION TO QUANTUM FIELD THEORY

1 Introduction 3
1.1 What is quantum field theory, and some preliminary notes 3
1.2 The notion of a quantized field 3
1.3 Natural units, notations and conventions 5
2 Relativistic symmetry 8
2.1 Lorentz transformations 8
2.2 Basic notions of group theory 11
2.3 The Lorentz and Poincaré groups 15
2.4 Tensor representation 17
2.5 Spinor representation 20
2.6 Irreducible representations of the Poincaré group 26
3 Lagrange formalism in field theory 31
3.1 The principle of least action, and the equations of motion 31
3.2 Global symmetries 34
3.3 Noether’s theorem 36
3.4 The energy-momentum tensor 39
4 Field models 42
4.1 Basic assumptions about the structure of Lagrangians 42
4.2 Scalar field models 43
4.3 Spinor field models 46
4.4 Models of free vector fields 52
4.5 Scalar and spinor filelds interacting with an electromagnetic field 55
4.6 The Yang-Mills field 59
5 Canonical quantization of free fields 70
5.1 Principles of canonical quantization 70
5.2 Canonical quantization in field theory 74
5.3 Canonical quantization of a free real scalar field 77
5.4 Canonical quantization of a free complex scalar field 82
5.5 Quantization of a free spinor field 85
5.6 Quantization of a free electromagnetic field 90
6 The scattering matrix and the Green functions 99
6.1 Particle interactions and asymptotic states 99
6.2 Reduction of the S-matrix to Green functions 104
6.3 Generating functionals of Green functions and the S-matrix 109
viii Contents

6.4 The S-matrix and the Green functions for spinor fields 112
7 Functional integrals 117
7.1 Representation of the evolution operator by a functional integral 117
7.2 Functional representation of Green functions 123
7.3 Functional representation of generating functionals 127
7.4 Functional integrals for fermionic theories 128
7.5 Perturbative calculation of generating functionals 134
7.6 Properties of functional integrals 137
7.7 Techniques for calculating functional determinants 143
8 Perturbation theory 147
8.1 Perturbation theory in terms of Feynman diagrams 147
8.2 Feynman diagrams in momentum space 151
8.3 Feynman diagrams for the S-matrix 154
8.4 Connected Green functions 155
8.5 Effective action 158
8.6 Loop expansion 162
8.7 Feynman diagrams in theories with spinor fields 166
9 Renormalization 171
9.1 The general idea of renormalization 171
9.2 Regularization of Feynman diagrams 172
9.3 The subtraction procedure 178
9.4 The superficial degree of divergences 194
9.5 Renormalizable and non-renormalizable theories 197
9.6 The arbitrariness of the subtraction procedure 204
9.7 Renormalization conditions 205
9.8 Renormalization with the dimensional regularization 209
9.9 Renormalization group equations 212
10 Quantum gauge theories 223
10.1 Basic notions of Yang-Mills gauge theory 223
10.2 Gauge invariance and observables 226
10.3 Functional integral for gauge theories 228
10.4 BRST symmetry 235
10.5 Ward identities 236
10.6 The gauge dependence of effective action 247
10.7 Background field method 249
10.8 Feynman diagrams in Yang-Mills theory 252
10.9 The background field method for Yang-Mills theory 254
10.10 Renormalization of Yang-Mills theory 257

PART II SEMICLASSICAL AND QUANTUM GRAVITY MODELS

11 A brief review of general relativity 271
11.1 Basic principles of general relativity 271
11.2 Covariant derivative and affine connection 272
 Contents ix

11.3 The curvature tensor and its properties 274

11.4 The covariant equation for a free particle: the classical limit 277
11.5 Classical action for the gravity field 280
11.6 Einstein equations and the Newton limit 283
11.7 Some physically relevant solutions and singularities 284
11.8 The applicability of GR and Planck units 289
12 Classical fields in curved spacetime 294
12.1 General considerations 294
12.2 Scalar fields 295
12.3 Spontaneous symmetry breaking in curved space
and induced gravity 300
12.4 Spinor fields in curved space 304
12.5 Massless vector (gauge) fields 312
12.6 Interactions between scalar, fermion and gauge fields 313
13 Quantum fields in curved spacetime: renormalization 314
13.1 Effective action in curved spacetime 314
13.2 Divergences and renormalization in curved space 321
13.3 Covariant methods: local momentum representation 328
13.4 The heat-kernel technique, and one-loop divergences 342
14 One-loop divergences 363
14.1 One-loop divergences in the vacuum sector 363
14.2 Beta functions in the vacuum sector 372
14.3 One-loop divergences in interacting theories 374
15 The renormalization group in curved space 388
15.1 The renormalization group based on minimal subtractions 388
15.2 The effective potential from a renormalization group 391
15.3 The global conformal (scaling) anomaly 395
16 Non-local form factors in flat and curved spacetime 397
16.1 Non-local form factors: simple example 397
16.2 Non-local form factors in curved spacetime 407
16.3 The massless limit and leading logs vs. the infrared limit 412
17 The conformal anomaly and anomaly-induced action 414
17.1 Conformal transformations and invariants 414
17.2 Derivation of the conformal anomaly 416
17.3 Anomaly-induced action 420
18 General notions of perturbative quantum gravity 425
18.1 Symmetries of the classical gravitational models 425
18.2 Choice of the action and gauge fixing for quantum gravity 427
18.3 Bilinear forms and linear approximation 434
18.4 Propagators of quantum metric and Barnes-Rivers projectors 438
18.5 Gravitational waves, quantization and gravitons 443
x Contents

18.6 Gauge-invariant renormalization in quantum gravity 446

18.7 Power counting, and classification of quantum gravity models 447
19 Massive ghosts in higher-derivative models 455
19.1 How to meet a massive ghost 455
19.2 The dangers of having a ghost or a tachyon 457
19.3 Massive ghosts in polynomial models 460
19.4 Complex poles and the unitarity of the S-matrix 462
19.5 Ghosts in nonlocal models 463
19.6 Effective approach to the problem of ghosts 465
19.7 Stable solutions in the presence of massive ghosts 467
20 One-loop renormalization in quantum gravity 474
20.1 Preliminary considerations 474
20.2 Gauge-fixing dependence in quantum GR 475
20.3 Gauge-fixing dependence in higher-derivative models 476
20.4 One-loop divergences in quantum general relativity 478
20.5 One-loop divergences in a fourth-derivative model 481
20.6 One-loop divergences in superrenormalizable models 486
21 The renormalization group in perturbative quantum gravity 488
21.1 On-shell renormalization group in quantum GR 489
21.2 The renormalization group in fourth-derivative gravity 490
21.3 The renormalization group in superrenormalizable models 492
22 The induced gravity approach 494
22.1 Gravity induced from the cut-off 495
22.2 Gravity induced from phase transitions 498
22.3 Once again on the cosmological constant problem 500
23 Final remarks on Part II 503
References 504
Index 522
Part I
Introduction to Quantum Field
Theory
1
Introduction

1.1 What is quantum field theory, and some preliminary notes

Quantum field theory (QFT) is part of the broader field of theoretical physics and is the
study of quantum effects in continuous physical systems called fields. One can say that
quantum field theory represents the unification of quantum mechanics and classical
field theory. Since a natural and consistent description of fundamental interactions can
be achieved in the framework of special relativity, it is also true to say that relativistic
quantum field theory represents the unification of quantum mechanics and special
relativity.
The main application of quantum field theory is the description of elementary
particles and their interactions. However, QFT has also extensive applications in other
areas of physics, including cosmology. Furthermore, quantum field theory plays an
important role in the theoretical condensed matter physics, especially in the description
of ensembles of a large number of interacting particles. The progress made in the theory
of superconductivity, the theory of phase transitions and other areas of condensed
matter physics is characterized by the consistent use of quantum field theory methods,
and vice versa.
The first part of this book is devoted to the basic notions and fundamental elements
of modern QFT formalism. In the second part, we present an introduction to the QFT
in curved space and quantum gravity, which are less developed and essentially more
complicated subjects.
The reader will note that the style of the two parts is different. In almost all of Part
I and in most of Part II, we tried to give a detailed presentation, so that the reader
could easily reproduce all calculations. However, following this approach for the whole
topic of quantum gravity would enormously increase the size of the book and make it
less readable. For this reason, in some places we avoided giving full technical details
and, instead, just provided references of papers or preprints where the reader can find
intermediate formulas. The same approach concerns the selection of the material. Since
we intended to write an introductory textbook, in Part II we gave only the need-to-
know information about quantum gravity. For this reason, many advanced subjects
were not included. In addition, in some cases, only qualitative discussion and minimal
references have been provided.

1.2 The notion of a quantized field

The field φ(x) = φ(t, x) is defined as a function of time t and the space coordinates,
that form a three-dimensional vector, x. It is assumed that the values of the space

Introduction to Quantum Field Theory with Applications to Quantum Gravity. Iosif L. Buchbinder
and Ilya L. Shapiro, Oxford University Press (2021). © Iosif L. Buchbinder and Ilya Shapiro.
DOI: 10.1093/oso/9780198838319.003.0001
4 Introduction

coordinates correspond to a bounded or unbounded domain of the three-dimensional

space. From the physical point of view, the field φ(t, x) can be treated as a dynamical
object with an infinite number of degrees of freedom, marked by the three-dimensional
vector index x.
The notion of a field naturally arises in the framework of special relativity. Since
there exists a maximal speed of propagation for any type of interaction, the physical
bodies separated by space intervals can not affect each other instantly. Therefore, there
should be a physical object responsible for transmitting perturbation from one body
to another. Such an object is a field that fills the space between the bodies and carries
perturbation from one body to another. The simplest example is an electromagnetic
field that carries interaction between electrically charged bodies.
Taking into account quantum mechanical universality, it is natural to assume that
fields should be quantized, like any other physical system. This means that quantum
states are given by wave functions, while dynamical variables are given by operators
acting on wave functions. Thus, in quantum theory, a field becomes an operator φ̂(t, x),
which is called a field operator.
As we have mentioned (and will discuss in more detail later on), a field is a system
with an infinite number of degrees of freedom. However, it turns out that the state of
the quantum field can be described in terms of either particles or fields. It turns out
that the quantum field is a physical notion that is most suitable for the description of
systems with an arbitrary number of particles.
It is well known that, in relativistic theory, there is a relation between momentum
p and the energy ε = ε(p) of a free particle,

ε 2 = m 2 c 4 + c 2 p2 , (1.1)

where c is the speed of light, and m is the mass of the particle. If the field can describe
particles, it must take into account the relation (1.1) between energy and momentum.
Let us try to clarify how the relation (1.1) can be implemented for the field. Let φ̂(t, x)
be the field operator, associated with the free particle. We can write the expansion as
a Fourier integral,
Z
i
φ̂(t, x) = d3 p dε e ~ (p·x−εt) φ̂(ε, p) , (1.2)

where ~ is the Planck constant. According to the standard interpretation, the vector
p is treated as the momentum of a particle, and the quantity ε as the energy of the
particle. Then, since, for each Fourier mode, ε and p are related by Eq. (1.1), the
quantity ε under the integral (1.2) is not an independent variable but is a function of
p. In order to satisfy this condition, one can write

φ̂(ε, p) = δ(ε2 − ε2 (p))φ̂∗ (p), (1.3)

where φ̂∗ (p) depends only on p. As a result, we arrive at the representation

Z
i
φ̂(t, x) = d3 p dε e ~ (px−εt) δ(ε2 − ε2 (p))φ̂∗ (p). (1.4)
 Natural units, notations and conventions 5

Consider the following expression showing a d’Alembert operator acting on the

field (1.2):
 1 ∂2 
φ̂(t, x) = 2 2
− ∆ φ̂(t, x)
Z c ∂t
i 1 1 
= d3 p dε e ~ (p·x−εt) 2 p2 − 2 ε2 δ(ε2 − ε2 (p))φ̂∗ (p)
~ c
Z h i
i 1 1
= d3 p e ~ (p·x−ε(p)t) 2 p2 − 2 ε2 (p) δ(ε2 − ε2 (p))φ̂∗ (p)
~ c
Z
3 i
(p·x−ε(p)t) 1h 2 1 i
= d p e~ 2
p − 2 (c2 p + m2 c4 ) δ(ε2 − ε2 (p))φ̂∗ (p)
~ c
2 2
m c
= − 2 φ̂(t, x).
~

Thus, we find that the free field operator should satisfy

 1 ∂2 m2 c2   m2 c2 
− ∆ + φ̂(t, x) =  + φ̂(t, x) = 0, (1.5)
c2 ∂t2 ~2 ~2

the Klein–Gordon equation. Equation (1.5) is a direct consequence of the relativistic

dispersion relation between the energy and the momentum of the particle.
If the field corresponds to a massless particle, the parameter m in Eq. (1.5) is zero.
Therefore, the field operator of a free massless field satisfies the wave equation

φ̂(t, x) = 0. (1.6)

Thus, any kind of a free relativistic quantum field is a spacetime-dependent oper-

ator satisfying the Klein–Gordon equation. In the case of interacting quantum fields,
their dynamics is described by much more complicated equations which will be dis-
cussed in the following chapters.

1.3 Natural units, notations and conventions

It is evident that the units of measurements of physical quantities should correspond to
the scales of phenomena where these units are used. For example, it is not reasonable
to measure the masses of elementary particles in tons or grams, or the size of atomic
nuclei in kilometers or centimeters.
When we consider the relativistic high-energy quantum phenomena in the fun-
damental quantum physics of elementary particles, it is natural to employ the units
related to the fundamental constants of nature. This means that we have to choose
the system of units where the speed of light is c = 1, and the Planck constant (which
has the dimension of the action) is ~ = 1. As a result, we obtain the natural sys-
tem of units based only on the fundamental constants of nature. In these units, the
action is dimensionless, the speed is dimensionless and the dimensions of energy and
momentum coincide. As in quantum theory, there is a Planck formula, relating energy
6 Introduction

and frequency as ε ∼ ~ω, and ω ∼ 1t , where t is time, and the dimensions satisfy the
relation

[ε] = [p] = [m] = [l]−1 = [t]−1 . (1.7)

Thus, we have only one remaining dimensional quantity, the unit of energy. Usually,
the energy in high-energy physics is measured in electron-volts, such that the unit of
energy is 1 eV , or 1 GeV = 109 eV . The dimensions of length and time are identical. In
what follows, we shall use this approach and assume the natural units of measurements
described above, with ~ = c = 1.
Other notations and conventions are as follows:
1) Minkowski space coordinates xµ ≡ (x0 , x) ≡ (t, x) ≡ (x0 , xi ), where Greek letters
represent the spacetime indices α, . . . , µ = 0, 1, 2, 3, while Latin letters are reserved for
the space indices, i, j, k, · · · = 1, 2, 3.
2) Functions in Minkowski space are denoted as φ(x) ≡ φ(x0 , xi ) ≡ φ(x0 , x) ≡ φ(t, x).
3) The Minkowski metric is
 
1 0 0 0
 0 −1 0 0 
ηµν = 
 0 0 −1 0  ≡ diag(1, −1, −1, −1), (1.8)
0 0 0 −1

and the same is true for the inverse metric, η µν = diag(1, −1, −1, −1). One can easily
check the relations η µν ηνρ = δ µ ρ and ηµν η νρ = δµ ρ .
Furthermore, εµναβ is the four-dimensional, totally antisymmetric tensor. The sign
convention is that ε0123 = 1 and hence ε0123 = −1.
4) Partial derivatives are denoted as

∂ ∂2
≡ ∂µ , ≡ ∂µ ∂ν , etc. (1.9)
∂xµ ∂xµ ∂xν

5) Rising and lowering the indices looks like

Aµ = η µν Aν , Aµ = ηµν Aν , ∂ µ = η µν ∂ν , ∂µ = ηµν ∂ ν , etc.

Let us note that these and some other rules will be changed in Part II, when we start
to deal with curved spacetime.
6) The scalar product is as follows:

AB = Aµ Bµ = A0 B0 + Ai Bi = A0 B0 − Ai Bi .

In particular,
px = pµ xµ = p0 x0 + pi xi = p0 x0 − p · x,
where pµ ≡ (p0 , p).
 Comments 7

7) The integral over four-dimensional space is

Z Z Z
d4 x = d3 x dx0 ,
R
while the integral over three-dimensional space is d3 x.
8) Dirac’s delta function in Minkowski space is
′ ′ ′
δ 4 (x − x′ ) ≡ δ(t − t′ )δ(x − x′ ) ≡ δ(t − t′ )δ(x1 − x 1 )δ(x2 − x 2 )δ(x3 − x 3 ). (1.10)
R
In particular, this means d4 x δ 4 (x − x′ )φ(x′ ) = φ(x).
9) The d’Alambertian operator is

 = ∂02 − ∂12 − ∂22 − ∂32 = η µν ∂µ ∂ν = ∂0 2 − ∆, (1.11)

where the Laplace operator is

∆ = ∂02 + ∂22 + ∂32 . (1.12)

10) The convention is that repeated indices imply the summation in all cases, i.e.,
N
X
XI YI = XI YI . (1.13)
I=1

Comments
There are many books on quantum field theory that differ in their manner and level of
presentation, targeting different audiences that range from beginners to more advanced
readers. Let us present a short list of basic references, which is based on our preferences.
The standard textbooks covering the basic notions and methods are those by J.D.
Bjorken and S.D. Drell [57], C. Itzykson and J.-B. Zuber [187], M.E. Peskin and D.V.
Schroeder [250], M. Srednicki [304] and M.D. Schwartz [274].
A brief and self-contained introduction to modern quantum field theory can be
found in the books by P. Ramond [256], M. Maggiore [215] and L. Alvarez-Gaume and
M.A. Vazquez-Mozo [155].
Comprehensive monographs in modern quantum field theory, with extensive cover-
age but aimed for advanced readers are those by J. Zinn-Justin [356], S. Weinberg [345],
B.S. DeWitt [106, 109] and W. Siegel [293].
There are also very useful lecture notes available online, e.g., those by H. Osborn
[235]. For mathematical and axiomatical aspects and approaches to quantum field
theory see, e.g., the book by N.N. Bogolubov, A.A. Logunov, A.I. Oksak and I. Todorov
[60].
2
Relativistic symmetry

In this chapter, we briefly review special relativistic symmetry, which will be used
in the rest of the book. In particular, we introduce basic notions of the Lorentz and
Poincaré groups, which will be used in constructing classical and quantum fields.
In general, the principles of symmetry play a fundamental role in physics. One
of the most universal symmetries of nature is the one that we can observe in the
framework of special relativity.

2.1 Lorentz transformations

According to special relativity, a spacetime structure is determined by the following
general principles:
1) Space and time are homogeneous.
2) Space is isotropic.
3) There exists a maximal speed of propagation of a physical signal. This maximal
speed coincides with the speed of light. In all inertial reference frames the speed of
light has the same value, c.
Let P1 and P2 be two infinitesimally separated events that are points in spacetime.
In some inertial reference frame, the four-dimensional coordinates of these events are
xµ and xµ + dxµ . The interval between these two events is defined as

ds2 = ηµν dxµ dxν . (2.1)

In another inertial reference frame, the same two events have the coordinates x′µ and
x′µ + dx′µ . The corresponding interval is

ds′2 = ηµν dx′µ dx′ν . (2.2)

The two intervals (2.1) and (2.2) are equal, that is, ds′2 = ds2 , reflecting the indepen-
dence of the speed of light on the choice of the inertial reference frame. Thus,

ηµν dxµ dxν = ηαβ dx′α dx′β . (2.3)

Eq. (2.3) enables one to find the relation between the coordinates x′α and xµ .
Let x′α = f α (x), with some unknown function f α (x). Substituting this relation into
Eq. (2.3), one gets an equation for the function f α (x) that can be solved in a general
form. As a result,

x′α = Λα µ xν + aα , (2.4)

Introduction to Quantum Field Theory with Applications to Quantum Gravity. Iosif L. Buchbinder
and Ilya L. Shapiro, Oxford University Press (2021). © Iosif L. Buchbinder and Ilya Shapiro.
DOI: 10.1093/oso/9780198838319.003.0002
 Lorentz transformations 9

where Λ ≡ (Λα µ ) is a matrix with constant elements, and aα is a constant four-vector.

Substituting Eq. (2.4) into Eq. (2.3), we get
ηαβ Λα µ Λβ ν = ηµν . (2.5)
The coordinate transformation (2.4) with the matrix Λα µ , satisfying Eq. (2.5), is called
the non-homogeneous Lorentz transformation. One can say that the non-homogeneous
Lorentz transformation is the most general coordinate transformation preserving the
form of the interval (2.1). If in Eq. (2.4) the vector aα = 0, the corresponding coordi-
nate transformation is called the homogeneous Lorentz transformation, or simply the
Lorentz transformation. Such a transformation has the form
x′µ = Λµ ν xν (2.6)
with the matrix Λµ ν satisfying Eq. (2.5).
It is convenient to present the relation (2.5) in a matrix form. Let us introduce the
matrices η ≡ (ηαβ ) and Λ ≡ (Λα µ ). Then Eq. (2.5) can be written as
ΛT η Λ = η, (2.7)
T T α α
where Λ is the transposed matrix with the elements (Λ )µ = Λ µ . One can regard
Eq. (2.7) as a basic relation. Any homogeneous Lorentz transformation is characterized
by the matrix Λ satisfying the basic relation, and vice versa. Therefore, the set of
all homogeneous Lorentz transformations is equivalent to the set of all matrices Λ,
satisfying (2.7).
Let us consider some important particular examples of Lorentz transformations:
1. Matrix Λ has the form
 
1 0
Λ= (2.8)
0 Ri j
where the matrix R = (Ri j ) transforms only space coordinates, x′i = Ri j xj . Substi-
tuting eq.(2.8) into the basic relation (2.7), we obtain the orthogonality condition
R T R = 13 , or Ri k δij Rj l = δkl , (2.9)
where 13 is a three-dimensional unit matrix with elements δij . Relation (2.9) defines
the three-dimensional rotations
x′0 = x0 , x′i = Ri j xj . (2.10)
If matrix R satisfies Eq. (2.9), then the transformation (2.10) is the Lorentz transfor-
mation. Thus, the three-dimensional rotations represent a particular case of Lorentz
transformation.
2. Consider a matrix Λ with the form
 v 
q 1 00 q c
2 2
 1− vc2 1− vc2 
 
 0 10 0 
Λ= , (2.11)
 0 01 0 
 v 
q c
00 q 1
2 2
1− vc2 1− vc2
10 Relativistic symmetry

where (v/c)2 < 1. It is easy to show that this matrix satisfies the basic relation.
Therefore, this matrix describes a Lorentz transformation,

x0 + v x1 x3 + v x0
x′0 = q c , x′1 = x1 , x′2 = x2 , x′3 = q c . (2.12)
2 2
1 − vc2 1 − vc2

This is the standard form of the Lorentz transformation for the case when one inertial
frame moves with respect to another one in the x3 direction. Indeed, one can construct
a similar matrix describing relative motion in any other direction. Transformations of
the type (2.12) are called boosts.
3. The matrix Λ corresponding to the time inversion, or T -transformation, is
 
−1 0 0 0
 0 1 0 0
Λ = ΛT =   0 0 1 0.


0 001

This matrix corresponds to the Lorentz transformation

x′0 = −x0 , x′i = xi . (2.13)

4. Let the matrix Λ have the form

 
1 0 0 0
 0 −1 0 0 
Λ = ΛP =  
 0 0 −1 0  .
0 0 0 −1

It is easy to check that the basic relation (2.7) is fulfilled in this case. This matrix
corresponds to the following Lorentz transformation:

x′0 = x0 x′i = −xi , (2.14)

which is called the space reflection or parity (P) transformation.

5. The matrix Λ with the form
 
−1 0 0 0
 0 −1 0 0 
Λ = ΛP T = ΛP ΛT = ΛT ΛP =  
 0 0 −1 0 
0 0 0 −1

corresponds to the following Lorentz transformation:

x′µ = −xµ , (2.15)

which is called the full reflection.

Eqs. (2.13), (2.14), (2.15) are called discrete Lorentz transformations.
 Basic notions of group theory 11

We will mainly need only the subclass of all Lorentz transformations that can be
obtained by small deformations of the identical transformation. Let the transformation
matrix have the form Λ = I, where I is the unit 4 × 4 matrix with elements δ µ ν .
Matrix I satisfies the basic relation (2.7). This matrix realizes the identical Lorentz
transformation

x′µ = xµ .

Stipulating small deformations of identical transformations means that we consider

matrices Λ of the form

Λ = I + ω, (2.16)

where ω is a matrix with infinitesimal elements ω µ ν . Requiring that the matrix Λ from
(2.16) correspond to a Lorentz transformation, we arrive at the relation

(I + ω)T η(I + ω) = η.

Taking into account only the first-order terms in ω, one gets

ω T η + ηω = 0.

Recovering the indices, we obtain

α
(ω T )µ ηαν + ηµα ω α ν = 0 =⇒ ω α µ ηαν + ηµα ω α ν = ωµν + ωνµ = 0. (2.17)

One can see that the matrix ω is real and antisymmetric, and hence it has six inde-
pendent elements. The matrix Λ (2.16) corresponds to the coordinate transformation

x′µ = xµ + ω µ ν xν ,

which is called the infinitesimal Lorentz transformation.

2.2 Basic notions of group theory

Group theory is a branch of mathematics devoted to the study of the symmetries.
In this subsection, we consider the basic notions of group theory that will be used in
the rest of the book. It is worth noting that this section is not intended to replace a
textbook on group theory. In what follows, we consequently omit rigorous definitions
and proofs of the theorems and concentrate only on the main notions of our interest.
A set G of the elements g1 , g2 , g3 , . . . , equipped with a law of composition (or
product of elements, or multiplication rule, or composition law), e.g., g1 g2 , is called a
group if for each pair of elements g1 , g2 ∈ G, the composition law satisfies the following
set of conditions:
1) Closure, i.e., ∀g1 , g2 ∈ G: g1 g2 ∈ G.
2) Associativity, i.e., ∀g1 , g2 , g3 ∈ G, for the product g1 (g2 g3 ) = (g1 g2 )g3 .
3) Existence of unit element, i.e., ∃e ∈ G, such that ∀g ∈ G : ge = eg = g.
4) Existence of inverse element, i.e., ∀g ∈ G, ∃g −1 ∈ G such that gg −1 = g −1 g = e.
12 Relativistic symmetry

Using these conditions, one can prove the uniqueness of the unit and inverse ele-
ments.
A group is called Abelian or commutative if, ∀g1 , g2 ∈ G, the product satisfies
g1 g2 = g2 g1 . In the opposite case, the group is called non-Abelian or non-commutative,
i.e., ∃g1 , g2 ∈ G such that g1 g2 6= g2 g1 .
A subset H ⊂ G is said to be a subgroup of group G if H itself is the group under
the same multiplication rule as group G. In particular, this means if h1 , h2 ∈ H, then
h1 h2 ∈ H. Also, e ∈ H, and if h ∈ H, then h−1 ∈ H.
A group consisting of a finite number of elements is called finite. In this case, it is
possible to form a group table gi gj . A finite group is sometimes called a finite discrete
group.
Let us consider a few examples:
1. Let G be a set of n × n real matrices M such that det M 6= 0. It is evident that
if M1 , M2 ∈ G, then det M1 M2 = det M1 det M2 6= 0 and hence M1 , M2 ∈ G. Thus,
this set forms a group under the usual matrix multiplication. The unit element is the
unit matrix E, and the element inverse to the matrix M is the inverse matrix M −1 . We
know that the multiplication of matrices is associative. Thus, all group conditions are
fulfilled. This group is called a general linear n-dimensional real group and is denoted
as GL(n|R). Consider a subset H ⊂ GL(n|R) consisting of matrices N that satisfy the
condition det N = 1. It is evident that det (N1 N2 ) = det N1 det N2 = 1. Hence

N1 , N2 ∈ H =⇒ N1 N2 ∈ H.

Consider other properties of this group. It is evident that E ∈ H. On the top of this,


N ∈ G =⇒ det N −1 = ( det N )−1 = 1.

The last means N −1 ∈ H. Hence H is a subgroup of the group GL(n|R). Group H is

called a special linear n-dimensional real group and is denoted as SL(n|R). In a similar
way, one can introduce general and special complex groups GL(n|C) and SL(n|C),
respectively, where C is a set of complex numbers.
2. Let G be a set of complex n × n matrices U such that U + U = U U + = E, where
E is the unit n × n matrix. Here, as usual, (U + )ab = (U ∗ )ba or U † = (U ∗ )T , where ∗
means the operation of complex conjugation, and T means transposition. Evidently,
E ∈ G and, for any U1 , U2 ∈ G, the following relations take place:

(U1 U2 )+ (U1 U2 ) = U2+ (U1+ U1 )U2 = U2+ U2 = E,

(U1 U2 )(U1 U2 )+ = U1 (U2 U2+ )U1+ = U1 U1+ = E. (2.18)

In addition, if U ∈ G, then (U −1 )+ U −1 = (U U + )−1 = U −1 (U −1 )+ = (U + U )−1 =

E. Therefore, if U ∈ G, then U −1 ∈ G too. As a result, the set of matrices under
consideration form a group. This group is called the n-dimensional unitary group
U (n).
The condition U + U = E leads to | det U |2 = 1. Hence det U = eiα , where α ∈ R.
One can also consider a subset of matrices U ∈ U (n), that satisfy the relation det U =
1. This subset forms a special unitary group and is denoted SU (n).
 Basic notions of group theory 13

Since the multiplication of matrices is, in general, a non-commutative operation,

the matrix groups GL(n, R), SL(n, R), U (n) and SU (n) are, in general, non-Abelian.
A group G is called the Lie group if each of its element is a differentiable function
of the finite number of parameters, and the product of any two group elements is a
differentiable function of parameters of each of the factors. That is, consider, ∀g ∈ G,
(1) (1)
(2) (2)

and for g1 = g ξ1 , . . . , ξN and g2 = g ξ1 , . . . , ξN , g = g1 g2 = g(ξ1 , . . . , ξN ).
Then
(1) (1) (2) (2)

ξI = fI ξ1 , . . . , ξN , ξ1 , . . . , ξN , (2.19)

where I = 1, 2, . . . , N are the differentiable functions of the parameters

(1) (1) (2) (2)
ξ1 , . . . , ξN , ξ1 , . . . , ξN . The Lie group is called compact if the parameters ξ1 , . . . , ξN
vary within a compact domain. One can prove that the parameters ξ1 , . . . , ξN can be
chosen in such a way that g(0, . . . , 0) = e, where e is the unit element of the group.
All matrix groups described in the examples above are the Lie groups, where the
role of parameters is played by independent matrix elements.
The two groups G and G′ are called homomorphic if there exists a map f of the
group G into the group G′ such that, for any two elements g1 , g2 ∈ G, the following
conditions take place: f (g1 g2 ) = f (g1 )f (g2 ), and if f (g) = g ′ , then f (g −1 ) = g ′−1 ,
where g ′−1 is an inverse element in the group G′ . Such a map is called homomorphism.
One can prove that f (e) = e′ , where e′ is the unit element of the group G′ . One-to-
one homomorphism is called isomorphism, and the corresponding groups are called
isomorphic. We will write, in this case, G = G′ .
Let G be some group, and V be a real or complex linear space. Consider a map
R such that, ∀g ∈ G, there exists an invertible operator DR (g) acting in the space V .
Furthermore, let the operators DR (g) satisfy the following conditions:
1) DR (e) = I, where I is a unit operator in the space V ; and 2) ∀g1 , g2 ∈ G, we
have DR (g1 g2 ) = DR (g1 )DR (g2 ).
The map R is called a representation of the group G in the linear space V . Operators
DR (g) are called the operators of representation, and the space V is called the space
−1
of the representation. One can prove that, ∀g ∈ G, there is DR (g −1 ) = DR (g), where
−1
DR (g) is the inverse operator for DR (g). Thus, the set of operators DR (g) forms a
group where a multiplication rule is the usual operator product.
We will mainly concern ourselves with matrix representations, where the operators
DR (g) are the n×n matrices DR (g)i j , i, j = 1, 2, . . . , n. Let v be a vector in a space of
representation with the coordinates v 1 , v 2 , . . . , v n , in some basis. The matrices DR (g)i j
generate the coordinate transformation of the form

v ′i = DR (g)i j v j .

Let R be a representation of the group G in the linear space V , and Ṽ be a

subspace in V , i.e., Ṽ ⊂ V . We assume that, for any vector ṽ ∈ Ṽ and for any
operator DR (g), the condition DR (g)ṽ ∈ Ṽ takes place. Then, the subspace Ṽ is
called the invariant subspace of the representation R. Any representation always has
two invariant subspaces, which are called trivial. These are the subspace Ṽ = V , and
the subspace Ṽ = {0}, which consists of a single zero element. All other invariant
14 Relativistic symmetry

subspaces, if they exist, are called non-trivial. A representation R is called reducible

if it has non-trivial invariant subspaces, and irreducible if it does not. In other words,
the representation R is called irreducible if it has only trivial invariant subspaces.
A representation is called completely irreducible if all representation matrices DR (g)
have the block-diagonal form. This means that, in a certain basis,
 
D1 (g) 0 0 ... 0
 
 0 D2 (g) 0 ... 0 
 
DR (g) =  .. .
 . ... 
 0 0 0 
0 0 0 ... Dk (g)

This situation means that the representation space has k non-trivial invariant sub-
spaces. In each of such subspaces, one can define an irreducible representation, Dk (g).
A given Lie group can have different representations, where the matrices DR (g) may
have different forms. However, some properties are independent of the representation.
Some of these properties can be formulated, e.g., in terms of Lie algebra. Let DR (g)
be the operators of representation, and g = g(ξ). Then, the operators DR (g) will be
the functions of N parameters ξ 1 , ξ 2 , . . . , ξ N , i.e., DR (g) = DR (ξ) and DR (ξ)|ξI =0 =
DR (e) = 1, where 1 is a unit matrix in the given representation space. One can prove
that, in an infinitesimal vicinity of the unit element, operators DR (ξ) can be presented
in the form
∂DR (ξ)
DR (ξ) = 1 + iξ I TR I , where TR I = −i . (2.20)
∂ξ I ξ=0

The operators TR I are called the generators of the group G in the representation R.
One can show that any operator DR (ξ) which is obtained by the continuous deforma-
tion from the unit element can be written as
I
DR (ξ) = eiξ TR I
. (2.21)
+ +
If the operator DR (ξ) is unitary, i.e., DR (ξ)DR (ξ) = DR (ξ)DR (ξ) = 1, then the
+
generators TR I are Hermitian, i.e., TR I = TR I . The generators of our interest satisfy
the following relation in terms of commutators:

[TRI , TRJ ] = ifIJ K TRK , (2.22)

where fIJ K are the structure constants of the Lie group G. It is evident that
fIJ K = −fJI K . In general, the form of the matrices TR I depends on the represen-
tation. However, one can prove that the structure constants do not depend on the
representation. Thus, these constants characterize the group G itself.
The group generators are closely related to the notion of Lie algebra. Let A be a
real or complex linear space with the elements a1 , a2 , . . . . A linear space A is called
Lie algebra, if for each two elements a1 , a2 ∈ A, there exists a composition law (also
called multiplication or the Lie product) [a1 , a2 ], such that
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internal lymphatic portion, which forms the main mass of the ovarian
ridge (Pl. 24, figs. 2, 3, and 6).
(2) At first the thickened germinal epithelium is sharply separated by
a membrane from the subjacent stroma (Pl. 24, figs. 1, 2, and 3),
but at about the time when the follicular epithelium commences to
be formed round the ova, numerous strands of stroma grow into the
epithelium, and form a regular network of vascular channels
throughout it, and partially isolate individual ova (Pl. 24, figs. 7 and
8). At the same time the surface of the epithelium turned towards
the stroma becomes irregular (Pl. 24, fig. 9), owing to the
development of individual ova. In still later stages the stroma
ingrowths form a more or less definite tunic close to the surface of
the ovary. External to this tunic is the superficial layer of the
germinal epithelium, which forms what has been spoken of as the
pseudo-epithelium. In many instances the protoplasm of its cells is
produced into peculiar fibrous tails which pass into the tunic below.
(3) Primitive ova.—Certain cells in the epithelium lining the dorsal
angle of the body-cavity become distinguished as primitive ova by
their abundant protoplasm and granular nuclei, at a very early period
in development, even before the formation of the genital ridges.
Subsequently on the formation of the genital ridges these ova
become confined to the thickened germinal epithelium on the outer
aspect of the ridges (Pl. 24, fig. 1).
(4) Conversion of primitive ova into permanent ova.—Primitive ova
may in Scyllium become transformed into permanent ova in two
ways—the difference between the two ways being, however, of
secondary importance.
(a) A nest of primitive ova makes its appearance, either by continued
division of a single primitive ovum or otherwise. The bodies of all the
ova of the nest fuse together, and a polynuclear mass is formed,
which increases in size concomitantly with the division of its nuclei.
The nuclei, moreover, pass through a series of transformations. They
increase in size and form delicate vesicles filled with a clear fluid, but
contain close to one side a granular mass which stains very deeply
with colouring reagents. The granular mass becomes somewhat
stellate, and finally assumes a reticulate form with one more highly
refracting nucleoli at the nodal points of the reticulum. When a
nucleus has reached this condition the protoplasm around it has
become slightly granular, and with the enclosed nucleus is
segmented off from the nest as a special cell—a permanent ovum
(figs. 13, 14, 15, 16). Not all the nuclei in a nest undergo the whole
of the above changes; certain of them, on the contrary, stop short in
their development, atrophy, and become employed as a kind of
pabulum for the remainder. Thus it happens that out of a large nest
perhaps only two or three permanent ova become developed.
(b) In the second mode of development of ova the nuclei and
protoplasm undergo the same changes as in the first mode; but the
ova either remain isolated and never form part of a nest, or form
part of a nest in which no fusion of the protoplasm takes place, and
all the primitive ova develop into permanent ova. Both the above
modes of the formation continue through a great part of life.
(5) The follicle.—The cells of the germinal epithelium arrange
themselves as a layer around each ovum, almost immediately after
its separation from a nest, and so constitute a follicle. They are at
first flat, but soon become more columnar. In Scyllium they remain
for a long time uniform, but in large eggs they become arranged in
two or three layers, while at the same time some of them become
large and flask-shaped, and others small and oval (fig. 29). The
flask-shaped cells have probably an important function in the
nutrition of the egg, and are arranged in a fairly regular order
amongst the smaller cells. Before the egg is quite ripe both kinds of
follicle cells undergo retrogressive changes (Pl. 25, fig. 23).
In Raja a great irregularity in the follicle cells is observable at an
early stage, but as the ovum grows larger the cells gradually assume
a regular arrangement more or less similar to that in Scyllium (Pl.
25, figs. 30-33).
(6) The egg membranes.—Two membranes are probably always
present in Elasmobranchii during some period of their growth. The
first formed and outer of these arises in some instances before the
formation of the follicular epithelium, and would seem to be of the
nature of a vitelline membrane. The inner one is the zona radiata
with a typical radiately striated structure. It is formed from the
vitellus at a much later period than the proper vitelline membrane. It
is more developed in Scyllium than in Raja, but atrophies early in
both genera. By the time the ovum is nearly ripe both membranes
are very much reduced, and when the egg (in Scyllium and
Pristiurus) is laid, no trace of any membrane is visible.
(7) The vitellus.—The vitellus is at first faintly granular, but at a later
period exhibits a very distinct (protoplasmic) network of fibres,
which is still present after the ovum has been laid.
The yolk arises, in the manner described by Gegenbaur, in ova of
about 0.06 mm. as a layer of fine granules, which stain deeply with
osmic acid. They are at first confined to a stratum of protoplasm
slightly below the surface of the ovum, and are most numerous at
the pole furthest removed from the germinal vesicle. They are not
regularly distributed, but are aggregated in small masses. They
gradually grow into vesicles, in the interior of which oval solid bodies
are developed, which form the permanent yolk-spheres. These oval
bodies in the later stages exhibit a remarkable segmentation into
plates, which gives them a peculiar appearance of transverse
striation.
Certain bodies of unknown function are occasionally met with in the
vitellus, of which the most remarkable are those figured at x on Pl.
25, fig. 25A.
(8) The germinal vesicle.—A reticulum is very conspicuous in the
germinal vesicle in the freshly formed ova, but becomes much less
so in older ova, and assumes, moreover, a granular appearance. At
first one to three nucleoli are present, but they gradually increase in
number as the germinal vesicle grows older, and are frequently
situated in close proximity to the membrane.

The Mammalian Ovary (Pl. 26).

The literature of the mammalian ovary has been so often dealt with
that it may be passed over with only a few words. The papers which
especially call for notice are those of Pflüger[400], Ed. van
Beneden[401], and especially Waldeyer[402], as inaugurating the newer
view on the nature of the ovary, and development of the ova; and of
Foulis[403] and Kölliker[404], as representing the most recent
utterances on the subject. There are, of course, many points in
these papers which are touched on in the sequel, but I may more
especially here call attention to the fact that I have been able to
confirm van Beneden's statement as to the existence of polynuclear
protoplasmic masses. I have found them, however, by no means
universal or primitive; and I cannot agree in a general way with van
Beneden's account of their occurrence. I have found no trace of a
germogene (Keimfache) in the sense of Pflüger and Ed. van
Beneden. My own results are most in accordance with those of
Waldeyer, with whom I agree in the fundamental propositions that
both ovum and follicular epithelium are derived from the germinal
epithelium, but I cannot accept his views of the relation of the
stroma to the germinal epithelium.
In the very interesting paper of Foulis, the conclusion is arrived at,
that while the ova are derived from the germinal epithelium, the
cells of the follicle originate from the ordinary connective tissue cells
of the stroma. Foulis regards the zona pellucida as a product of the
ovum and not of the follicle. To both of these views I shall return,
and hope to be able to shew that Foulis has not traced back the
formation of the follicle through a sufficient number of the earlier
stages. It thus comes about that though I fully recognise the
accuracy of his figures, I am unable to admit his conclusions.
Kölliker's statements are again very different from those of Foulis.
He finds certain cords of cells in the hilus of the ovary, which he
believes to be derived from the Wolffian body, and has satisfied
himself that they are continuous with Pflüger's egg-tubes, and that
they supply the follicular epithelium. To the general accuracy of
Kölliker's statements with reference to the relations of these cords in
the hilus of the ovary I can fully testify, but am of opinion that he is
entirely mistaken as to their giving rise to the follicular epithelium, or
having anything to do with the ova. I hope to be able to give a fuller
account of their origin than he or other observers have done.
My investigations on the mammalian ovary have been made almost
entirely on the rabbit—the type of which it is most easy to procure a
continuous series of successive stages; but in a general way my
conclusions have been controlled and confirmed by observations on
the cat, the dog, and the sheep. My observations commence with an
embryo of eighteen days. A transverse section, slightly magnified,
through the ovary at this stage, is represented on Pl. 26, fig. 35, and
a more highly magnified portion of the same in fig. 35A. The ovary is
a cylindrical ridge on the inner side of the Wolffian body, composed
of a superficial epithelium, the germinal epithelium (g.e.), and of a
tissue internal to this, which forms the main mass of it. In the latter
two constituents have to be distinguished—(1) an epithelial-like
tissue (t), coloured brown, which forms the most important element,
and (2) vascular and stroma elements in this.
The germinal epithelium is a layer about 0.03 - 0.04 mm. in
thickness. It is (vide fig. 35A, g.e.) composed of two or three layers
of cells, with granular nuclei, of which the outermost layer is more
columnar than the remainder, and has elongated rather than
rounded nuclei. Its cells, though they vary slightly in size, are all
provided with a fair amount of protoplasm, and cannot be divided
(as in the case of the germinal epithelium of Birds, Elasmobranchii,
&c.), into primitive ova, and normal epithelial cells. Very occasionally,
however, a specially large cell, which, perhaps, deserves the
appellation primitive ovum, may be seen. From the subjacent tissue
the germinal epithelium is in most parts separated by a membrane-
like structure (fluid coagulum); but this is sometimes absent, and it
is then very difficult to determine with exactness the inner border of
the epithelium. The tissue (t), which forms the greater mass of the
ovary at this stage, is formed of solid columns or trabeculæ of
epithelial-like cells, which present a very striking resemblance in size
and character to the cells of the germinal epithelium. The
protoplasm of these cells stains slightly more deeply with osmic acid
than does that of the cells of the germinal epithelium, so that it is
rather easier to note a difference between the two tissues in osmic
acid than in picric acid specimens. This tissue approaches very
closely, and is in many parts in actual contact with the germinal
epithelium. Between the columns of it are numerous vascular
channels (shewn diagrammatically in my figures) and a few normal
stroma cells. This remarkable tissue continues visible through the
whole course of the development of the ovary, till comparatively late
in life, and during all the earlier stages might easily be supposed to
be about to play some part in the development of the ova, or even
to be part of the germinal epithelium. It really, however, has nothing
to do with the development of the ova, as is easily demonstrated
when the true ova begin to be formed. In the later stages, as will be
mentioned in the description of those stages, it is separated from
the germinal epithelium by a layer of stroma; though at the two
sides of the ovary it is, even in later stages, sometimes in contact
with the germinal epithelium.
In most parts this tissue is definitely confined within the limits of the
ovary, and does not extend into the mesentery by which the ovary is
attached. It may, however, be traced at the anterior end of the ovary
into connection with the walls of the Malpighian bodies, which lie on
the inner side of the Wolffian body (vide fig. 35B), and I have no
doubt that it grows out from the walls of these bodies into the ovary.
In the male it appears to me to assist in forming, together with cells
derived from the germinal epithelium, the seminiferous tubules, the
development of which is already fairly advanced by this stage. I shall
speak of it in the sequel as tubuliferous tissue. The points of interest
in connection with it concern the male sex, which I hope to deal with
in a future paper, but I have no hesitation in identifying it with the
segmental cords (segmentalstränge) discovered by Braun in Reptilia,
and described at length in his valuable memoir on their urogenital
system[405]. According to Braun the segmental cords in Reptilia are
buds from the outer walls of the Malpighian bodies. The bud from
each Malpighian body grows into the genital ridge before the period
of sexual differentiation, and sends out processes backwards and
forwards, which unite with the buds from the other Malpighian
bodies. There is thus formed a kind of trabecular work of tissue in
the stroma of the ovary, which in the Lacertilia comes into
connection with the germinal epithelium in both sexes, but in
Ophidia in the male only. In the female, in all cases, it gradually
atrophies and finally vanishes, but in the male there pass into it the
primitive ova, and it eventually forms, with the enclosed primitive
ova, the tubuli seminiferi. From my own observations in Reptilia I
can fully confirm Braun's statements as to the entrance of the
primitive ova into this tissue in the male, and the conversion of it
into the tubuli seminiferi. The chief difference between Reptilia and
Mammalia, in reference to this tissue, appears to be that in
Mammalia it arises only from a few of the Malpighian bodies at the
anterior extremity of the ovary, but in Reptilia from all the
Malpighian bodies adjoining the genital ridge. More extended
observations on Mammalia will perhaps shew that even this
difference does not hold good.
It is hardly to be supposed that this tissue, which is so conspicuous
in all young ovaries, has not been noticed before; but the notices of
it are not so numerous as I should have anticipated. His[406] states
that the parenchyma of the sexual glands undoubtedly arises from
the Wolffian canals, and adds that while the cortical layer (Hulle)
represents the earlier covering of a part of the Wolffian body, the
stroma of the hilus, with its vessels, arises from a Malpighian body.
In spite of these statements of His, I still doubt very much whether
he has really observed either the tissue I allude to or its mode of
development. In any case he gives no recognisable description or
figure of it.
Waldeyer[407] notices this tissue in the dog, cat, and calf. The
following is a free translation of what he says, (p. 141):—“In a full
grown but young dog, with numerous ripe follicles, there were
present in the vascular zone of the ovary numerous branched
elongated small columns (Schläuche) of epithelial cells, between
which ran blood-vessels. They were only separated from the egg
columns of the cortical layer by a row of large follicles. There can be
no doubt that we have here remains of the sexual part of the
Wolffian body—the canals of the parovarium—which in the female
sex have developed themselves to an extraordinary extent into the
stroma of the sexual gland, and perhaps are even to be regarded as
homologues of the seminiferous tubules (the italics are my own). I
have almost always found the above condition in the dog, only in old
animals these seminiferous canals seem gradually to atrophy. Similar
columns are present in the cat, only they do not appear to grow so
far into the stroma.” Identical structures are also described in the
calf.
Romiti gives a very similar description to Waldeyer of these bodies in
the dog[408]. Born also describes this tissue in young and embryonic
ovaries of the horse as the Keimlager[409]. The columns described by
Kölliker[410] and believed by him to furnish the follicular epithelium,
are undoubtedly my tubuliferous tissue, and, as Kölliker himself
points out, are formed of the same tissue as that described by
Waldeyer.
Egli gives a very clear and accurate description of this tissue, though
he apparently denies its relation with the Wolffian body.
My own interpretation of the tissue accords with that of Waldeyer. In
addition to the rabbit, I have observed it in the dog, cat, and sheep.
In all these forms I find that close to the attachment of the ovary,
and sometimes well within it, a fair number of distinct canals with a
large lumen are present, which are probably to be distinguished
from the solid epithelial columns. Such large canals are not as a rule
present in the rabbit. In the dog solid columns are present in the
embryo, but later they appear frequently to acquire a tubular form,
and a lumen. Probably there are great variations in the development
of the tissue, since in the cat (not as Waldeyer did in the dog) I have
found it most developed.
In the very young embryonic ovary of the cat the columns are very
small and much branched. In later embryonic stages they are
frequently elongated, sometimes convoluted, and are very similar to
the embryonic tubuli seminiferi. In the young stages these columns
are so similar to the egg tubes (which agree more closely with
Pflüger's type in the cat than in other forms I have worked at) that
to any one who had not studied the development of the tissue an
embryo cat's ovary at certain stages would be a very puzzling object.
I have, however, met with nothing in the cat or any other form
which supports Kölliker's views.
My next stage is that of a twenty-two days' embryo. Of this stage I
have given two figures corresponding to those of the earlier stage
(figs. 36 and 36A).
From these figures it is at once obvious that the germinal epithelium
has very much increased in bulk. It has a thickness 0.1 - 0.09 mm.
as compared to 0.03 mm. in the earlier stage. Its inner outline is
somewhat irregular, and it is imperfectly divided into lobes, which
form the commencement of structures nearly equivalent to the nests
of the Elasmobranch ovary. The lobes are not separated from each
other by connective tissue prolongations; the epithelium being at
this stage perfectly free from any ingrowths of stroma. The cells
constituting the germinal epithelium have much the same character
as in the previous stage. They form an outer row of columnar cells
internal to which the cells are more rounded. Amongst them a few
large cells with granular nuclei, which are clearly primitive ova, may
now be seen, but by far the majority of the cells are fairly uniform in
size, and measure from 0.01 - 0.02 mm. in diameter, and their nuclei
from 0.004 - 0.006 mm. The nuclei of the columnar outer cells
measure about 0.008 mm. They are what would ordinarily be called
granular, though high powers shew that they have the usual nuclear
network. There is no special nucleolus. The rapid growth of the
germinal epithelium is due to the division of its cells, and great
masses of these may frequently be seen to be undergoing division at
the same time. Of the tissue of the ovary internal to the germinal
epithelium, it may be noticed that the tubuliferous tissue derived
from the Malpighian bodies is no longer in contact with the germinal
epithelium, but that a layer of vascular stroma is to a great extent
interposed between the two. The vascular stroma of the hilus has,
moreover, greatly increased in quantity.
My next stage is that of a twenty-six days' embryo, but the
characters of the ovary at this stage so closely correspond with
those of the succeeding one at twenty-eight days that, for the sake
of brevity, I pass over this stage in silence.
Figs. 37 and 37A are representative sections of the ovary of the
twenty-eighth day corresponding with those of the earlier stages.
Great changes have become apparent in the constitution of the
germinal epithelium. The vascular stroma of the ovary has grown
into the germinal epithelium precisely as in Elasmobranchii. It
appears to me clear that the change in the relations between the
stroma and epithelium is not due to a mutual growth, but entirely to
the stroma, so that, as in the case of Elasmobranchii, the result of
the ingrowth is that the germinal epithelium is honeycombed by
vascular stroma. The vascular growths generally take the paths of
the lines which separated the nests in an earlier condition, and
cause these nests to become the egg tubes of Pflüger. It is obvious
in figure 37 that the vascular ingrowths are so arranged as
imperfectly to divide the germinal epithelium into two layers
separated by a space with connective tissue and blood-vessels. The
outer part is relatively thin, and formed of a superficial row of
columnar cells, and one or two rows of more rounded cells; the
inner layer is much thicker, and formed of large masses of rounded
cells. The two layers are connected together by numerous
trabeculæ, the stroma between which eventually gives rise to the
connective tissue capsule, or tunica albuginea, of the adult ovary.
The germinal epithelium is now about 0.19 to 0.22 mm. in thickness.
Its cells have undergone considerable changes. A fair number of
them (fig. 37A, p.o.), especially in the outer layer of the epithelium,
have become larger than the cells around them, from which they are
distinguished, not only by their size, but by their granular nucleus
and abundant protoplasm. They are in fact undoubted primitive ova
with all the characters which primitive ova present in Elasmobranchii,
Aves, &c. In a fairly typical primitive ovum of this stage the body
measures 0.02 mm. and the nucleus 0.014 mm. In the inner part of
the germinal epithelium there are very few or no cells which can be
distinguished by their size as primitive ova, and the cells themselves
are of a fairly uniform size, though in this respect there is perhaps a
greater variation than might be gathered from fig. 37A. The cells are
on the average about 0.016 mm. in diameter, and their nuclei about
0.008 to 0.001 mm., considerably larger, in fact, than in the earlier
stage. The nuclei are moreover more granular, and make in this
respect an approach to the character of the nuclei of primitive ova.
The germinal epithelium is still rapidly increasing by the division of
its cells, and in fig. 37A there are shewn two or three nuclei in the
act of dividing. I have represented fairly accurately the appearance
they present when examined with a moderately high magnifying
power. With reference to the stroma of the ovary, internal to the
germinal epithelium, it is only necessary to refer to fig. 37 to observe
that the tubuliferous tissue (t) forms a relatively smaller part of the
stroma than in the previous stage, and is also further removed from
the germinal epithelium.
My next stage is that of a young rabbit two days after birth, but to
economise space I pass on at once to the following stage five days
after birth. This stage is in many respects a critical one for the ovary,
and therefore of great interest. Figure 38 represents a transverse
section through the ovary (on rather a smaller scale than the
previous figures) and shews the general relations of the tissues.
The germinal epithelium is very much thicker than before—about
0.38 mm. as compared with 0.22 mm. It is divided into three
obvious layers: (1) an outer epithelial layer which corresponds with
the pseudo-epithelial layer of the Elasmobranch ovary, average
thickness 0.03 mm. (2) A middle layer of small nests, which
corresponds with the middle vascular layer of the previous stage;
average thickness 0.1 mm. (3) An inner layer of larger nests;
average thickness 0.23 mm.
The general appearance of the germinal epithelium at this stage
certainly appears to me to lend support to my view that the whole of
it simply constitutes a thickened epithelium interpenetrated with
ingrowths of stroma.
The cells of the germinal epithelium, which form the various layers,
have undergone important modifications. In the first place a large
number of the nuclei—at any rate of those cells which are about to
become ova—have undergone a change identical with that which
takes place in the conversion of the primitive into the permanent ova
in Elasmobranchii. The greater part of the contents of the nucleus
becomes clear. The remaining contents arrange themselves as a
deeply staining granular mass on one side of the membrane, and
later on as a somewhat stellate figure: the two stages forming what
were spoken of as the granular and stellate varieties of nucleus. To
avoid further circumlocution I shall speak of the nucleus undergoing
the granular and the stellate modifications. At a still later period the
granular contents form a beautiful network in the nucleus.
The pseudo-epithelium (fig. 38A) is formed of several tiers of cells,
the outermost of which are very columnar and have less protoplasm
than in an earlier stage. In the lower tiers of cells there are many
primitive ova with granular nuclei, and others in which the nuclei
have undergone the granular modification. The primitive ova are
almost all of the same size as in the earlier stage. The pseudo-
epithelium is separated from the middle layer by a more or less
complete stratum of connective tissue, which, however, is traversed
by trabeculæ connecting the two layers of the epithelium. In the
middle layer there are comparatively few modified nuclei, and the
cells still retain for the most part their earlier characters. The
diameter of the cells is about 0.012 mm., and that of the nucleus
about 0.008 mm. In the innermost layer (fig. 38B), which is not
sharply separated from the middle layer, the majority of the cells,
which in the previous stage were ordinary cells of the epithelium,
have commenced to acquire modified nuclei. This change, which first
became apparent to a small extent in the young two days after birth,
is very conspicuous at this stage. In some of the cells the nucleus is
modified in the granular manner, in others in the stellate, and in a
certain number the nucleus has assumed a reticular structure
characteristic of the young permanent ovum.
In addition, however, to the cells which are becoming converted into
ova, a not inconsiderable number may be observed, if carefully
looked for, which are for the most part smaller than the others,
generally somewhat oval, and in which the nucleus retains its
primitive characters. A fair number of such cells are represented in
fig. 38B. In the larger ones the nucleus will perhaps eventually
become modified; but the smaller cells clearly correspond with the
interstitial cells of the Elasmobranch germinal epithelium, and are
destined to become converted into the epithelium of the Graafian
follicle. In some few instances indeed (at this stage very few), in the
deeper part of the germinal epithelium, these cells commence to
arrange themselves round the just formed permanent ova as a
follicular epithelium. An instance of this kind is shewn in fig. 38B, o.
The cells with modified nuclei, which are becoming permanent ova,
usually present one point of contrast to the homologous cells in
Elasmobranchii, in that they are quite distinct from each other, and
not fused into a polynuclear mass. They have around them a dark
contour line, which I can only interpret as the commencement of the
membrane (zona radiata?), which afterwards becomes distinct, and
which would thus seem, as Foulis has already insisted, to be of the
nature of a vitelline membrane.
In a certain number of instances the protoplasm of the cells which
are becoming permanent ova appears, however, actually to fuse, and
polynuclear masses identical with those in Elasmobranchii are thus
formed (cf. E. van Beneden[411]). These masses become slightly more
numerous in the succeeding stages. Indications of a fusion of this
kind are shewn in fig. 38B. That the polynuclear masses really arise
from a fusion of primitively distinct cells is clear from the description
of the previous stages. The ova in the deeper layers, with modified
granular nuclei, measure about 0.016 - 0.02 mm., and their nuclei
from 0.01 - 0.012 mm.
With reference to the tissue of the hilus of the ovary, it may be
noticed that the tubuliferous tissue (t) is relatively reduced in
quantity. Its cells retain precisely their previous characters.
The chief difference between the stage of five days and that of two
days after birth consists in the fact that during the earlier stage
comparatively few modified nuclei were present, but the nuclei then
presented the character of the nuclei of primitive ova.
I have ovaries both of the dog and cat of an equivalent stage, and in
both of these the cells of the nests or egg tubes may be divided into
two categories, destined respectively to become ova and follicle
cells. Nothing which has come under my notice tends to shew that
the tubuliferous tissue is in any way concerned in supplying the
latter form of cell.
In a stage, seven days after birth, the same layers in the germinal
epithelium may be noticed as in the last described stage. The
outermost layer or pseudo-epithelium contains numerous developing
ova, for the most part with modified nuclei. It is separated by a well
marked layer of connective tissue from the middle layer of the
germinal epithelium. The outer part of the middle layer contains
more connective tissue and smaller nests than in the earlier stage,
and most of the cells of this layer contain modified nuclei. In a few
nests the protoplasm of the developing ova forms a continuous
mass, not divided into distinct cells, but in the majority of instances
the outline of each ovum can be distinctly traced. In addition to the
cells destined to become ova, there are present in these nests other
cells, which will clearly form the follicular epithelium. A typical nest
from the middle layer is represented on Pl. 26, fig. 39A.
The nests or masses of ova in the innermost layer are for the most
part still very large, but, in addition to the nests, a few isolated ova,
enclosed in follicles, are to be seen.
A fairly typical nest, selected to shew the formation of the follicle, is
represented on Pl. 26, fig. 39B.
The nest contains (1) fully formed permanent ova, completely or
wholly enclosed in a follicle. (2) Smaller ova, not enclosed in a
follicle. (3) Smallish cells with modified nuclei of doubtful
destination. (4) Small cells obviously about to form follicular
epithelium.
The inspection of a single such nest is to my mind a satisfactory
proof that the follicular epithelium takes its origin from the germinal
epithelium and not from the stroma or tubuliferous tissue. The
several categories of elements observable in such a nest deserve a
careful description.
(1) The large ova in their follicles.—These ova have precisely the
character of the young ova in Elasmobranchii. They are provided
with a granular body invested by a delicate, though distinct
membrane. Their nucleus is large and clear, but traversed by the
network so fully described for Elasmobranchii. The cells of their
follicular epithelium have obviously the same character as many
other small cells of the nest. Two points about them deserve notice
—(a) that many of them are fairly columnar. This is characteristic
only of the first formed follicles. In the later formed follicles the cells
are always flat and spindle-shaped in section. In this difference
between the early and late formed follicles Mammals agree with
Elasmobranchii. (b) The cells of the follicle are much more columnar
towards the inner side than towards the outer. This point also is
common to Mammals and Elasmobranchii.
Round the completed follicle a very delicate membrana propria
folliculi appears to be present[412].
The larger ova, with follicular epithelium, measure about 0.04 mm.,
and their nucleus about 0.02 mm., the smaller ones about 0.022
mm., and their nucleus about 0.014 mm.
(2) Medium sized ova.—They are still without a trace of a follicular
epithelium, and present no special peculiarities.
(3) The smaller cells with modified nuclei.—I have great doubt as to
what is the eventual fate of these cells. There appear to be three
possibilities.
(a) That they become cells of the follicular epithelium; (b) that they
develop into ova; (c) that they are absorbed as a kind of food by the
developing ova. I am inclined to think that some of these cells may
have each of the above-mentioned destinations.
(4) The cells which form the follicle.—The only point to be noticed
about these is that they are smaller than the indifferent cells of the
germinal epithelium, from which they no doubt originate by division.
This fact has already been noticed by Waldeyer.
The isolated follicles at this stage are formed by ingrowths of
connective tissue cutting off fully formed follicles from a nest. They
only occur at the very innermost border of the germinal epithelium.
This is in accordance with what has so often been noticed about the
mammalian ovary, viz. that the more advanced ova are to be met
with in passing from without inwards.
By the stage seven days after birth the ovary has reached a
sufficiently advanced stage to answer the more important question I
set myself to solve, nevertheless, partly to reconcile the apparent
discrepancy between my account and that of Dr Foulis, and partly to
bring my description up to a better known condition of the ovary, I
shall make a few remarks about some of the succeeding stages.
In a young rabbit about four weeks old the ovary is a very beautiful
object for the study of the nuclei, &c.
The pseudo-epithelium is now formed of a single layer of columnar
cells, with comparatively scanty protoplasm. In it there are present a
not inconsiderable number of developing ova.
A layer of connective tissue—the albuginea—is now present below
the pseudo-epithelium, which contains a few small nests with very
young permanent ova. The layer of medium sized nests internal to
the albuginea forms a very pretty object in well stained sections,
hardened in Kleinenberg's picric acid. The ova in it have all assumed
the permanent form, and are provided with beautiful reticulate
nuclei, with, as a rule, one more especially developed nucleolus, and
smaller granular bodies. Their diameter varies from about 0.028 to
0.04 mm. and that of their nucleus from 0.016 to 0.02 mm. The
majority of these ova are not provided with a follicular investment,
but amongst them are numerous small cells, clearly derived from the
germinal epithelium, which are destined to form the follicle (vide fig.
40Aand B). In a few cases the follicles are completed, and are then
formed of very flattened spindle-shaped (in section) cells. In the
majority of cases all the ova of each nest are quite distinct, and each
provided with a delicate vitelline membrane (fig. 40A) In other
instances, which, so far as I can judge, are more common than in
the previous stages, the protoplasm of two or more ova is fused
together.
Examples of this are represented in Pl. 26, fig. 40A. In some of these
the nuclei in the undivided protoplasm are all of about the same size
and distinctness, and probably the protoplasm eventually becomes
divided up into as many ova as nuclei; in other cases, however, one
or two nuclei clearly preponderate over the others, and the smaller
nuclei are indistinct and hazy in outline. In these latter cases I have
satisfied myself as completely as in the case of Elasmobranchii, that
only one or two ova (according to the number of distinct nuclei) will
develop out of the polynuclear mass, and that the other nuclei
atrophy, and the material of which they were composed serves as
the nutriment for the ova which complete their development. This
does not, of course, imply that the ova so formed have a value other
than that of a single cell, any more than the development of a single
embryo out of the many in one egg capsule implies that the embryo
so developing is a compound organism.
In the innermost layer of the germinal epithelium the outlines of the
original large nests are still visible, but many of the follicles have
been cut off by ingrowths of stroma. In the still intact nests the
formation of the follicles out of the cells of the germinal epithelium
may be followed with great advantage. The cells of the follicle,
though less columnar than was the case at an earlier period, are
more so than in the case of follicles formed in the succeeding
stages. The previous inequality in the cells of the follicles is no
longer present.
The tubuliferous tissue in the zona vasculosa appears to me to have
rather increased in quantity than the reverse; and is formed of
numerous solid columns or oval masses of cells, separated by
strands of connective tissue, with typical spindle nuclei.
It is partially intelligible to me how Dr Foulis might from an
examination of the stages similar to this, conclude that the follicle
cells were derived from the stroma; but even at this stage the
position of the cells which will form the follicular epithelium, their
passage by a series of gradations into obvious cells of the germinal
epithelium and the peculiarities of their nuclei, so different from
those of the stroma cells, supply a sufficient series of characters to
remove all doubt as to the derivation of the follicle cells. Apart from
these more obvious points, an examination of the follicle cells from
the surface, and not in section, demonstrates that the general
resemblance in shape of follicle cells to the stroma cells is quite
delusory. They are in fact flat, circular, or oval, plates not really
spindle-shaped, but only apparently so in section. While I thus
fundamentally differ from Foulis as to the nature of the follicle cells,
I am on this point in complete accordance with Waldeyer, and my
own results with reference to the follicle cannot be better stated
than in his own words (pp. 43, 44).
At six weeks after birth the ovary of the rabbit corresponds very
much more with the stages in the development of the ovary, which
Foulis has more especially studied, for the formation of the follicular
epithelium, than during the earlier stages. His figure (Quart. Journ.
Mic. Sci., Vol. XVI., Pl. 17, fig. 6) of the ovary of a seven and a half
months' human fœtus is about the corresponding age. Different
animals vary greatly in respect to the relative development of the
ovary. For example, the ovary of a lamb at birth about corresponds
with that of a rabbit six weeks after birth. The points which may be
noticed about the ovary at this age are first that the surface of the
ovary begins to be somewhat folded. The appearances of these folds
in section have given rise, as has already been pointed out by Foulis,
to the erroneous view that the germinal epithelium (pseudo-
epithelium) became involuted in the form of tubular open pits. The
folds appear to me to have no connection with the formation of ova,
but to be of the same nature as the somewhat similar folds in
Elasmobranchii. A follicular epithelium is present around the majority
of the ova of the middle layer, and around all those of the inner layer
of the germinal epithelium. The nests are, moreover, much more cut
up by connective tissue ingrowths than in the previous stages.
The follicle cells of the middle layers are very flat, and spindle-
shaped in section, and though they stain more deeply than the
stroma cells, and have other not easily characterised peculiarities,
they nevertheless do undoubtedly closely resemble the stroma cells
when viewed (as is ordinarily the case) in optical section.
In the innermost layer many of the follicles with the enclosed ova
have advanced considerably in development and are formed of
columnar cells. The somewhat heterodox view of these cells
propounded by Foulis I cannot quite agree to. He says (Quart. J.
Mic. Sci., Vol. XVI., p. 210): “The protoplasm which surrounds the
vesicular nuclei acts as a sort of cement substance, holding them
together in the form of a capsular membrane round the young
ovum. This capsular membrane is the first appearance of the
membrana granulosa.” I must admit that I find nothing similar to
this, nor have I met with any special peculiarities (as Foulis would
seem to indicate) in the cells of the germinal epithelium or other
cells of the ovary.
Figure 41 is a representation of an advanced follicle of a six weeks'
rabbit, containing two ova, which is obviously in the act of dividing
into two. Follicles of this kind with more than one ovum are not very
uncommon. It appears to me probable that follicles, such as that I
have figured, were originally formed of a single mass of protoplasm
with two nuclei; but that instead of one of the nuclei atrophying,
both of them eventually developed and the protoplasm subsequently
divided into two masses. In other cases it is quite possible that
follicles with two ova should rather be regarded as two follicles not
separated by a septum of stroma.
On the later stages of development of the ovary I have no complete
series of observations. The yolk spherules I find to be first developed
in a peripheral layer of the vitellus. I have not been able definitely to
decide the relation of the zona radiata to the first formed vitelline
membrane. Externally to the zona radiata there may generally be
observed a somewhat granular structure, against which the follicle
cells abut, and I cannot agree with Waldeyer (loc cit., p. 40) that this
structure is continuous with the cells of the discus, or with the zona
radiata. Is it the remains of the first formed vitelline membrane? I
have obtained some evidence in favour of this view, but have not
been successful in making observations to satisfy me on the point,
and must leave open the question whether my vitelline membrane
becomes the zona radiata or whether the zona is not a later and
independent formation, but am inclined myself to adopt the latter
view. The first formed membrane, whether or no it becomes the
zona radiata, is very similar to the vitelline membrane of
Elasmobranchii and arises at a corresponding stage.

Summary of observations on the mammalian ovary.—The general

results of my observations on the mammalian ovary are the
following:—
(1) The ovary in an eighteen days' embryo consists of a cylindrical
ridge attached along the inner side of the Wolffian body, which is
formed of two parts; (a) an external epithelium—two or three cells
deep (the germinal epithelium); (b) a hilus or part forming in the
adult the vascular zone, at this stage composed of branched masses
of epithelial tissue (tubuliferous tissue) derived from the walls of the
anterior Malpighian bodies, and numerous blood-vessels, and some
stroma cells.
(2) The germinal epithelium gradually becomes thicker, and after a
certain stage (twenty-three days) there grow into it numerous
stroma ingrowths, accompanied by blood-vessels. The germinal
epithelium thus becomes honeycombed by strands of stroma. Part of
the stroma eventually forms a layer close below the surface, which
becomes in the adult the tunica albuginea. The part of the germinal
epithelium external to this layer becomes reduced to a single row of
cells, and forms what has been spoken of in this paper as the
pseudo-epithelium of the ovary. The greater part of the germinal
epithelium is situated internal to the tunica albuginea, and this part
is at first divided up by strands of stroma into smaller divisions
externally, and larger ones internally. These masses of germinal
epithelium (probably sections of branched trabeculæ) may be
spoken of as nests. In the course of the development of the ova
they are broken up by stroma ingrowths, and each follicle with its
enclosed ovum is eventually isolated by a layer of stroma.
(3) The cells of the germinal epithelium give rise both to the
permanent ova and to the cells of the follicular epithelium. For a
long time, however, the cells remain indifferent, so that the stages,
like those in Elasmobranchii, Osseous Fish, Birds, Reptiles, &c., with
numerous primitive ova embedded amongst the small cells of the
germinal epithelium, are not found.
(4) The conversion of the cells of the germinal epithelium into
permanent ova commences in an embryo of about twenty-two days.
All the cells of the germinal epithelium appear to be capable of
becoming ova: the following are the stages in the process, which are
almost identical with those in Elasmobranchii:—
(a) The nucleus of the cells loses its more or less distinct network,
and becomes very granular, with a few specially large granules
(nucleoli). The protoplasm around it becomes clear and abundant—
primitive ovum stage. It may be noted that the largest primitive ova
are very often situated in the pseudo-epithelium. (b) A segregation
takes place in the contents of the nucleus within the membrane, and
the granular contents pass to one side, where they form an irregular
mass, while the remaining space within the membrane is perfectly
clear. The granular mass gradually develops itself into a beautiful
reticulum, with two or three highly refracting nucleoli, one of which
eventually becomes the largest and forms the germinal spot par
excellence. At the same time the body of the ovum becomes slightly
granular. While the above changes, more especially those in the
nucleus, have been taking place, the protoplasm of two or more ova
may fuse together, and polynuclear masses be so formed. In some
cases the whole of such a polynuclear mass gives rise to only a
single ovum, owing to the atrophy of all the nuclei but one, in others
it gives rise by subsequent division to two or more ova, each with a
single germinal vesicle.
(5) All the cells of a nest do not undergo the above changes, but
some of them become smaller (by division) than the indifferent cells
of the germinal epithelium, arrange themselves round the ova, and
form the follicular epithelium.
(6) The first membrane formed round the ovum arises in some cases
even before the appearance of the follicular epithelium, and is of the
nature of a vitelline membrane. It seems probable, although not
definitely established by observation, that the zona radiata is formed
internally to the vitelline membrane, and that the latter remains as a
membrane, somewhat irregular on its outer border, against which
the ends of the follicle cells abut.

General Observations on the Structure and Development of the Ovary.

In selecting Mammalia and Elasmobranchii as my two types for
investigation, I had in view the consideration that what held good for
such dissimilar forms might probably be accepted as true for all
Vertebrata with the exception of Amphioxus.
The structure of the ovary.—From my study of these two types, I
have been led to a view of the structure of the ovary, which differs
to a not inconsiderable extent from that usually entertained. For
both types the conclusion has been arrived at that the whole egg-
containing part of the ovary is really the thickened germinal
epithelium, and that it differs from the original thickened patch or
layer of germinal epithelium, mainly in the fact that it is broken up
into a kind of meshwork by growths of vascular stroma. If the above
view be accepted for Elasmobranchii and Mammalia, it will hardly be
disputed for the ovaries of Reptilia and Aves. In the case also of
Osseous Fish and Amphibia, this view of the ovary appears to be
very tenable, but the central core of stroma present in the other
types is nearly or quite absent, and the ovary is entirely formed of
the germinal epithelium with the usual strands of vascular
stroma[413]. It is obvious that according to the above view Pflüger's
egg-tubes are merely trabeculæ of germinal epithelium, and have no
such importance as has been attributed to them. They are present in
a more or less modified form in all types of ovaries. Even in the adult
Amphibian ovary, columns of cells of the germinal epithelium, some
indifferent, others already converted into ova, are present, and, as
has been pointed out by Hertwig[414], represent Pflüger's egg-tubes.
The formation of the permanent ova.—The passage of primitive ova
into permanent ova is the part of my investigation to which the
greatest attention was paid, and the results arrived at for Mammalia
and Elasmobranchii are almost identical. Although there are no
investigations as to the changes undergone by the nucleus in other
types, still it appears to me safe to conclude that the results arrived
at hold good for Vertebrates generally[415]. As has already been
pointed out the transformation which the so-called primitive ova
undergo is sufficient to shew that they are not to be regarded as ova
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