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A Guide to Experiments in Quantum Optics
A Guide to Experiments in Quantum Optics

Hans-A. Bachor and Timothy C. Ralph

Third Edition
Authors All books published by Wiley-VCH
are carefully produced. Nevertheless,
Hans-A. Bachor authors, editors, and publisher do not
The Australian Nat. University warrant the information contained in
Research School of Physics these books, including this book, to
Science Road be free of errors. Readers are advised
Canberra ACT 0200 to keep in mind that statements, data,
Australia illustrations, procedural details or other
items may inadvertently be inaccurate.
Dr. Timothy C. Ralph
University of Queensland Library of Congress Card No.:
Department of Physics applied for
St. Lucia QLD 4072
Australia British Library Cataloguing-in-Publication
Data
Cover A catalogue record for this book is
The artwork was prepared by B. Hage available from the British Library.
and is based on an experimental
demonstration of photon number states Bibliographic information published by
published in: the Deutsche Nationalbibliothek
The Deutsche Nationalbibliothek lists
Chrzanowski H.M., Assad S.M., Bernu J., this publication in the Deutsche
and Hage B. et al. (2013). Reconstruction Nationalbibliografie; detailed
of Photon Number Conditioned States bibliographic data are available on the
Using Phase Randomized Homodyne Internet at <http://dnb.d-nb.de>.
Measurements, J. Phys. B At. Mol. Opt.
Phys. 46: 104009. © 2019 Wiley-VCH Verlag GmbH &
Co. KGaA, Boschstr. 12, 69469
Weinheim, Germany

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Print ISBN: 978-3-527-41193-1

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10 9 8 7 6 5 4 3 2 1
 v

Contents

Preface xv
Acknowledgments xix

1 Introduction 1
1.1 Optics in Modern Life 1
1.2 The Origin and Progress of Quantum Optics 3
1.3 Motivation Through Simple and Direct Teaching Experiments 7
1.4 Consequences of Photon Correlations 12
1.5 How to Use This Guide 14
References 16

2 Classical Models of Light 19

2.1 Classical Waves 20
2.1.1 Mathematical Description of Waves 20
2.1.2 The Gaussian Beam 21
2.1.3 Quadrature Amplitudes 24
2.1.4 Field Energy, Intensity, and Power 25
2.1.5 A Classical Mode of Light 26
2.1.6 Light Carries Information 28
2.1.7 Modulations 30
2.2 Optical Modes and Degrees of Freedom 32
2.2.1 Lasers with Single and Multiple Modes 32
2.2.2 Polarization 33
2.2.2.1 Poincaré Sphere and Stokes Vectors 35
2.2.3 Multimode Systems 36
2.3 Statistical Properties of Classical Light 37
2.3.1 The Origin of Fluctuations 37
2.3.1.1 Gaussian Noise Approximation 38
2.3.2 Noise Spectra 39
2.3.3 Coherence 40
2.3.3.1 Correlation Functions 44
2.4 An Example: Light from a Chaotic Source as the Idealized Classical
Case 46
2.5 Spatial Information and Imaging 50
vi Contents

2.5.1 State-of-the-Art Imaging 50

2.5.2 Classical Imaging 52
2.5.3 Image Detection 55
2.5.4 Scanning 56
2.5.5 Quantifying Noise and Contrast 58
2.5.6 Coincidence Imaging 59
2.5.7 Imaging with Coherent Light 60
2.5.8 Image Reconstruction with Structured Illumination 60
2.5.9 Image Analysis and Modes 61
2.5.10 Detection Modes and Displacement 61
2.6 Summary 62
References 63
Further Reading 64

3 65
Photons: The Motivation to Go Beyond Classical Optics
3.1 Detecting Light 65
3.2 The Concept of Photons 68
3.3 Light from a Thermal Source 70
3.4 Interference Experiments 73
3.5 Modelling Single-Photon Experiments 78
3.5.1 Polarization of a Single Photon 79
3.5.1.1 Some Mathematics 80
3.5.2 Polarization States 81
3.5.3 The Single-Photon Interferometer 83
3.6 Intensity Correlation, Bunching, and Anti-bunching 84
3.7 Observing Photons in Cavities 88
3.8 Summary 90
References 90
Further Reading 92

4 Quantum Models of Light 93

4.1 Quantization of Light 93
4.1.1 Some General Comments on Quantum Mechanics 93
4.1.2 Quantization of Cavity Modes 94
4.1.3 Quantized Energy 95
4.1.4 The Creation and Annihilation Operators 97
4.2 Quantum States of Light 97
4.2.1 Number or Fock States 97
4.2.2 Coherent States 99
4.2.3 Mixed States 101
4.3 Quantum Optical Representations 102
4.3.1 Quadrature Amplitude Operators 102
4.3.2 Probability and Quasi-probability Distributions 104
4.3.3 Photon Number Distributions 108
4.3.4 Covariance Matrix 111
4.3.4.1 Summary of Different Representations of Quantum States and
Quantum Noise 112
 Contents vii

4.4 Propagation and Detection of Quantum Optical Fields 113

4.4.1 Quantum Optical Modes in Free Space 114
4.4.2 Propagation in Quantum Optics 115
4.4.3 Detection in Quantum Optics 117
4.4.4 An Example: The Beamsplitter 118
4.5 Quantum Transfer Functions 120
4.5.1 A Linearized Quantum Noise Description 121
4.5.2 An Example: The Propagating Coherent State 123
4.5.3 Real Laser Beams 123
4.5.4 The Transfer of Operators, Signals, and Noise 124
4.5.5 Sideband Modes as Quantum States 126
4.5.6 Another Example: A Coherent State Pulse Through a Frequency
Filter 129
4.5.7 Transformation of the Covariance Matrix 130
4.6 Quantum Correlations 131
4.6.1 Photon Correlations 131
4.6.2 Quadrature Correlations 132
4.6.3 Two-Mode Covariance Matrix 133
4.7 Summary 134
4.7.1 The Photon Number Basis 134
4.7.2 Quadrature Representations 135
4.7.3 Quantum Operators 135
4.7.4 The Quantum Noise Limit 136
References 136
Further Reading 137

5 Basic Optical Components 139

5.1 Beamsplitters 140
5.1.1 Classical Description of a Beamsplitter 140
5.1.1.1 Polarization Properties of Beamsplitters 142
5.1.2 The Beamsplitter in the Quantum Operator Model 143
5.1.3 The Beamsplitter with Single Photons 144
5.1.4 The Beamsplitter and the Photon Statistics 146
5.1.5 The Beamsplitter with Coherent States 149
5.1.5.1 Transfer Function for a Beamsplitter 149
5.1.6 Comparison Between a Beamsplitter and a Classical Current
Junction 151
5.1.7 The Beamsplitter as a Model of Loss 152
5.2 Interferometers 153
5.2.1 Classical Description of an Interferometer 154
5.2.2 Quantum Model of the Interferometer 155
5.2.3 The Single-Photon Interferometer 156
5.2.4 Transfer of Intensity Noise Through the Interferometer 156
5.2.5 Sensitivity Limit of an Interferometer 157
5.2.6 Effect of Mode Mismatch on an Interferometer 160
5.3 Optical Cavities 162
5.3.1 Classical Description of a Linear Cavity 164
viii Contents

5.3.2 The Special Case of High Reflectivities 169

5.3.3 The Phase Response 170
5.3.4 Spatial Properties of Cavities 172
5.3.4.1 Mode Matching 172
5.3.4.2 Polarization 174
5.3.4.3 Tunable Mirrors 175
5.3.5 Equations of Motion for the Cavity Mode 175
5.3.6 The Quantum Equations of Motion for a Cavity 176
5.3.7 The Propagation of Fluctuations Through the Cavity 177
5.3.8 Single Photons Through a Cavity 180
5.3.9 Multimode Cavities 181
5.3.10 Engineering Beamsplitters, Interferometers, and Resonators 182
5.4 Other Optical Components 184
5.4.1 Lenses 184
5.4.2 Holograms and Metasurfaces 185
5.4.3 Crystals and Polarizers 187
5.4.4 Optical Fibres and Waveguides 188
5.4.5 Modulators 189
5.4.5.1 Phase and Amplitude Modulators 191
5.4.6 Spatial Light Modulators 193
5.4.7 Optical Noise Sources 195
5.4.8 Non-linear Processes 195
References 196

6 Lasers and Amplifiers 199

6.1 The Laser Concept 199
6.1.1 Technical Specifications of a Laser 201
6.1.2 Rate Equations 203
6.1.3 Quantum Model of a Laser 207
6.1.4 Examples of Lasers 209
6.1.4.1 Classes of Lasers 209
6.1.4.2 Dye Lasers and Argon Ion Lasers 209
6.1.4.3 The CW Nd:YAG Laser 210
6.1.4.4 Diode Lasers 213
6.1.4.5 Limits of the Single-Mode Approximation in Diode Lasers 213
6.1.5 Laser Phase Noise 214
6.1.6 Pulsed Lasers 215
6.2 Amplification of Optical Signals 215
6.3 Parametric Amplifiers and Oscillators 218
6.3.1 The Second-Order Non-linearity 219
6.3.2 Parametric Amplification 220
6.3.3 Optical Parametric Oscillator 221
6.3.3.1 Noise Spectrum of the Parametric Oscillator 222
6.3.4 Pair Production 223
6.4 Measurement-Based Amplifiers 224
6.4.1 Deterministic Measurement-Based Amplifiers 225
6.4.2 Heralded Measurement-Based Amplifiers 228
 Contents ix

6.5 Summary 230

References 231

7 Photon Generation and Detection 233

7.1 Photon Sources 236
7.1.1 Deterministic Photon Sources 239
7.2 Photon Detection 240
7.2.1 Detecting Individual Photons 240
7.2.1.1 Photochemical Detectors 241
7.2.1.2 Photoelectric Detectors 241
7.2.1.3 Photo-thermal Detectors 243
7.2.1.4 Multipixel and Imaging Devices 243
7.2.2 Recording Electrical Signals from Individual Photons 245
7.3 Generating, Detecting, and Analysing Photocurrents 247
7.3.1 Properties of Photocurrents 247
7.3.1.1 Beat Measurements 247
7.3.1.2 Intensity Noise and the Shot Noise Level 248
7.3.1.3 Quantum Efficiency 249
7.3.1.4 Photodetector Materials 250
7.3.2 Generating Photocurrents 251
7.3.2.1 Photodiodes and Detector Circuit 251
7.3.2.2 Amplifiers and Electronic Noise 252
7.3.2.3 Detector Saturation 254
7.3.3 Recording of Photocurrents 255
7.3.4 Spectral Analysis of Photocurrents 257
7.3.4.1 Digital Fourier Transform 257
7.3.4.2 Analogue Fourier Transform 258
7.3.4.3 From Optical Sidebands to the Current Spectrum 258
7.3.4.4 The Operation of an Electronic Spectrum Analyser 259
7.3.4.5 Detecting Signal and Noise Independently 260
7.3.4.6 The Decibel Scale 261
7.3.4.7 Adding Electronic AC Signals 262
7.4 Imaging with Photons 263
References 264
Further Reading 267

8 Quantum Noise: Basic Measurements and Techniques 269

8.1 Detection and Calibration of Quantum Noise 269
8.1.1 Direct Detection and Calibration 269
8.1.1.1 White Light Calibration 273
8.1.2 Balanced Detection 273
8.1.3 Detection of Intensity Modulation and SNR 275
8.1.4 Homodyne Detection 275
8.1.4.1 The Homodyne Detector for Classical Waves 275
8.1.5 Heterodyne Detection 279
8.1.5.1 Measuring Other Properties 280
8.2 Intensity Noise 281
x Contents

8.2.1 Laser Noise 281

8.3 The Intensity Noise Eater 282
8.3.1 Classical Intensity Control 282
8.3.2 Quantum Noise Control 285
8.3.2.1 Practical Consequences 289
8.4 Frequency Stabilization and Locking of Cavities 290
8.4.1 Pound–Drever–Hall Locking 292
8.4.2 Tilt Locking 293
8.4.3 The PID Controller 294
8.4.4 How to Mount a Mirror 295
8.4.5 The Extremes of Mirror Suspension: GW Detectors 296
8.5 Injection Locking 296
References 299

9 Squeezed Light 303

9.1 The Concept of Squeezing 303
9.1.1 Tools for Squeezing: Two Simple Examples 303
9.1.1.1 The Kerr Effect 304
9.1.1.2 Four-Wave Mixing 307
9.1.2 Properties of Squeezed States 310
9.1.2.1 What Are the Uses of These Various Types of Squeezed Light? 312
9.2 Quantum Model of Squeezed States 314
9.2.1 The Formal Definition of a Squeezed State 314
9.2.2 The Generation of Squeezed States 317
9.2.3 Squeezing as Correlations Between Noise Sidebands 319
9.3 Detecting Squeezed Light 322
9.3.1 Detecting Amplitude Squeezed Light 322
9.3.2 Detecting Quadrature Squeezed Light 322
9.3.3 Using a Cavity to Measure Quadrature Squeezing 324
9.3.4 Summary of Different Representations of Squeezed States 325
9.3.5 Propagation of Squeezed Light 325
9.4 Early Demonstrations of Squeezed Light 330
9.4.1 Four Wave Mixing 330
9.4.2 Optical Parametric Processes 333
9.4.3 Second Harmonic Generation 339
9.4.4 The Kerr Effect 343
9.4.4.1 The Response of the Kerr Medium 343
9.4.4.2 Optimizing the Kerr Effect 345
9.4.4.3 Fibre Kerr Squeezing 346
9.4.4.4 Atomic Kerr Squeezing 348
9.4.4.5 Atomic Polarization Self-Rotation 349
9.5 Pulsed Squeezing 349
9.5.1 Quantum Noise of Optical Pulses 349
9.5.2 Pulsed Squeezing Experiments with Kerr Media 352
9.5.3 Pulsed SHG and OPO Experiments 353
9.5.4 Soliton Squeezing 354
9.5.5 Spectral Filtering 355
 Contents xi

9.5.6 Non-linear Interferometers 356

9.6 Amplitude Squeezed Light from Diode Lasers 358
9.7 Quantum State Tomography 360
9.8 State of the Art of CW Squeezing 363
9.9 Squeezing of Multiple Modes 365
9.9.1 Twin-Photon Beams 365
9.9.2 Polarization Squeezing 367
9.9.3 Degenerate Multimode Squeezers 368
9.10 Summary: Quantum Limits and Enhancement 370
References 371
Further Reading 376

10 Applications of Quantum Light 377

10.1 Quantum Enhanced Sensors 377
10.1.1 Coherent Sensors and Sensitivity Scaling 377
10.1.2 Practical Examples of Sensors 380
10.1.3 Ultimate Sensing Limits 382
10.1.4 Adaptive Phase Estimation 384
10.2 Optical Communication 384
10.3 Gravitational Wave Detection 389
10.3.1 The Origin and Properties of GW 389
10.3.1.1 Concept and Design of an Optical GW Detector 392
10.3.2 Quantum Properties of the Ideal Interferometer 393
10.3.2.1 Configurations of Interferometers 396
10.3.2.2 Recycling 397
10.3.2.3 Modulation Techniques 398
10.3.3 The Sensitivity of GW Observatories 400
10.3.3.1 Enhancement Below the SQL 402
10.3.4 Interferometry with Squeezed Light 405
10.3.4.1 Quantum Enhancement Beyond the SQL 410
10.4 Quantum Enhanced Imaging 411
10.4.1 Imaging with Photons on Demand 411
10.4.2 Quantum Enhanced Coincidence Imaging 412
10.5 Multimode Squeezing Enhancing Sensors 414
10.5.1 Spatial Multimode Squeezing 414
10.6 Summary and Outlook 419
References 419

11 QND 425
11.1 QND Measurements of Quadrature Amplitudes 425
11.2 Classification of QND Measurements 427
11.3 Experimental Results 430
11.4 Single-Photon QND 432
11.4.1 Measurement-Based QND 434
References 437
xii Contents

12 Fundamental Tests of Quantum Mechanics 441

12.1 Wave–Particle Duality 441
12.2 Indistinguishability 446
12.3 Non-locality 453
12.3.1 Einstein–Podolsky–Rosen Paradox 453
12.3.2 Characterization of Entangled Beams via Homodyne Detection 458
12.3.2.1 Logarithmic Negativity and Two-Mode Squeezing 459
12.3.2.2 Entanglement of Formation 460
12.3.3 Bell Inequalities 461
12.3.3.1 Long-Distance Bell Inequality Violations 466
12.3.3.2 Loophole-Free Bell Inequality Violations 466
12.4 Summary 468
References 468

13 Quantum Information 473

13.1 Photons as Qubits 473
13.1.1 Other Quantum Encodings 475
13.2 Post-selection and Coincidence Counting 475
13.3 True Single-Photon Sources 477
13.3.1 Heralded Single Photons 477
13.3.2 Single Photons on Demand 480
13.4 Characterizing Photonic Qubits 482
13.5 Quantum Key Distribution 484
13.5.1 QKD Using Single Photons 485
13.5.2 QKD Using Continuous Variables 489
13.5.3 No Cloning 492
13.6 Teleportation 492
13.6.1 Teleportation of Photon Qubits 493
13.6.2 Continuous Variable Teleportation 495
13.6.3 Entanglement Swapping 502
13.6.4 Entanglement Distillation 502
13.7 Quantum Computation 505
13.7.1 Dual-Rail Quantum Computing 506
13.7.1.1 Quantum Circuits with Linear Optics 507
13.7.1.2 Cluster States 511
13.7.1.3 Quantum Gates with Non-linear Optics 513
13.7.2 Single-Rail Quantum Computation 514
13.7.2.1 Quantum Random Walks 515
13.7.2.2 Boson Sampling 516
13.7.3 Continuous Variable Quantum Computation 518
13.7.3.1 Cat State Quantum Computing 519
13.7.3.2 Continuous Variable Cluster States 521
13.7.4 Large-Scale Quantum Computation 522
13.8 Summary 525
References 526
Further Reading 531
 Contents xiii

14 The Future: From Q-demonstrations to Q-technologies 533

14.1 Demonstrating Quantum Effects 533
14.2 Matter Waves and Atoms 535
14.3 Q-Technology Based on Optics 537
14.3.1 Applications of Squeezed Light 537
14.3.2 Quantum Communication and Logic with Photons 539
14.3.3 Cavity QED 542
14.3.4 Extending to Other Wavelengths: Microwaves and Cryogenic
Circuits 542
14.3.5 Quantum Optomechanics 542
14.3.6 Transfer of Quantum Information Between Different Physical
Systems 543
14.3.7 Transferring and Storing Quantum States 544
14.4 Outlook 544
References 545
Further Reading 547

Appendices 549
Appendix A: List of Quantum Operators, States, and Functions 549
Appendix B: Calculation of the Quantum Properties of a Feedback
Loop 551
Appendix C: Detection of Signal and Noise with an ESA 552
Reference 554
Appendix D: An Example of Analogue Processing of
Photocurrents 554
Appendix E: Symbols and Abbreviations 556

Index 559
 xv

Preface

The Idea Behind This Guide

Some of the most interesting and sometimes puzzling phenomena in optics are
those where the quantum mechanical nature of light is apparent. Recent years
have seen a rapid expansion of quantum optics. Beautiful demonstrations and
applications of the quantum nature of light are now common, and optics has
been shown to be one of the key areas of physics to build quantum technolo-
gies based on the concepts behind quantum mechanics. The field of photonics
has already spawned major industries and changed the way we communicate,
gather data, and handle information. Future applications of quantum informa-
tion will enhance this trend and will introduce even more quantum aspects into
our everyday activities.
Why should we still write and read books at the age of search engines, blogs,
and Wikipedia? This question did not arise when the first edition of this book
was published in 1998, which is a short step back in history but a long time ago
in terms of the changes in information technology and the way we live, learn, and
do research. You are likely to read this on a screen and can follow up links within
seconds, tapping into an amazing wealth of information. This calls even more on
this Guide to Experiment in Quantum Optics to explain the concepts, introduce
reliable techniques, and help to select from the enormous amount of information
that is available.
We intend to guide you through the many experiments that have been pub-
lished and to present and interpret them in one common style. This book provides
a practical basis for exploring the ever-expanding technology in optics and elec-
tronics and provides a practical link to the rapidly expanding field of quantum
information, which is now being used in many different platforms. Finally, this
book is an attempt to communicate our personal insights and experiences in this
field of science. Science is driven by creative people, and some insight in the way
researchers advance the field should find interest with you the reader.
Many excellent textbooks are available on this topic. In the same way as this
field of research has been initiated by theoretical ideas, most of these books are
written from a theoretical point of view. They provide fascinating, solid, and
reliable descriptions of the field. However, we found the literature frequently
leaves out some of the important simpler pictures and intuitive interpretations
of the experiments. In this guide we focus on both the first and the most recent
xvi Preface

experiments. The guide explains the underlying physics in the most intuitive
way we can find. It addresses questions such as the following: What are the
limitations of the equipment? What can be measured and what remains a goal
for the future?
The answers will prepare the readers to form their own mind map of quantum
optics. It gives them the tools to make independent predictions of the outcome
of their own experiments. For those who want to use quantum optics in practical
ways, it tries to give an understanding of the design concepts of the machines
that exploit the quantum nature of light. We assume that most of our readers
will already have a fairly good understanding of optical phenomena and the way
light propagates and how it interacts with components such as lenses, mirrors,
detectors, etc. They can imagine the processes in action in an optical instrument.
It is useful in the everyday work of a scientist to have practical pictures of what is
going on inside the experiment or the machine. Such pictures have been shaped
over many years of systematic study and through one’s own experimentation. Our
guide provides such pictures, which can be filled in with many more details from
theory and technology.
In quantum optics three different pictures, or interpretations, are used simul-
taneously: light as waves, light as photons, and light as the solution to operator
equations. To these we add the picture of fluctuations propagating through an
optical system, a description using noise transfer functions. All of them are useful;
all of them are correct within their limits. They are based on specific interpreta-
tions of the one formalism, and they are mathematically equivalent. In this guide
all these four descriptions are introduced rigorously and are used to discuss a
series of experiments. We start with simple demonstrations, gradually getting
more complex and include most of the landmark experiments in single-photon
and quantum noise detection, photon pair generation, squeezed states, quantum
non-demolition, and entanglement measurements published to date. All four pic-
tures – waves, photons, operators, and noise spectra – are used in parallel. In each
case the most appropriate and intuitive interpretation is highlighted, and the lim-
itations of all four interpretations are stated. In this way we hope to maintain and
present a lot of the fascination of quantum optics that has motivated us and many
of our colleagues, young and old, over so many years.

The Expansion of Quantum Optics and the Third Edition

of This Guide
The development of the field has been rapid in recent years, and many new tech-
nical advances have been reported: vastly improved sources for photon pairs,
squeezed and entangled states. We have better detectors and nonlinear media.
With the ability to record and process much more information the era of analogue
technology is gone. Post-processing is now the norm, quantum gates and quan-
tum memories have been reported. We can generate and process multiple optical
modes in frequency, space and time, demonstrate multimode entanglement and
design and build reliable complex optical systems.
 Preface xvii

The field of quantum information is growing rapidly, and new concepts for
quantum communication, quantum logic, and quantum internet are emerging.
Optics will play a major role in this. The field has broadened to include both
more fundamental concepts about the quantum nature of information and
new ideas for the engineering of complex optical designs. Measurements using
non-classical light are now well established and have become a routine operation
in some select fields such as gravitational-wave detectors and sensors. Entangle-
ment has grown from an abstract concept to a technical tool. At the same time
commercial companies using quantum techniques for secure communication
have already carved out a large market for themselves.
The ideas in this guide, and the examples chosen, draw attention to both the
techniques of single-photon detection and single-event logic and the techniques
of continuous variable logic and the use of continuous coherent and squeezed
beams. There are now far more practitioners of the former, and the latter has
become a specialized niche, mainly used for metrology. However, both will play
a role in the development of future applications and in the understanding the
concepts of the quantum world. In addition, quantum optics based on pairs of
photons has been combined with the technology of strong laser beams and con-
tinuous variables in a series of exciting hybrid experiments.
This third edition focusses on the latest landmark experiments, and examples
that are now outdated have been deleted. The coverage of quantum information
concepts and protocols has been expanded to keep pace with the evolution of the
field. Reference is made to the state-of-the-art technologies, hinting at potential
future developments. This guide follows the well-tested idea of providing inde-
pendent building blocks in theory and experimental techniques that are used in
later chapters to discuss complete experiments and applications. We found that
most of them were still valid and thus remain unchanged.
Optics is no longer dominant as the medium of choice for quantum technolo-
gies. Technical advances have allowed the same concepts now to be used in areas
such as cryogenic electric circuits, superconducting systems, trapped ions and
neutral atoms, Bose–Einstein condensates (BECs) and atom laser beams, and
optomechanical systems, and other examples yet to be tested. This has posed a
challenge: how to define the limits of this guide? To keep the book compact, we
decided to focus entirely on optical systems and to point to links with other sys-
tems where required, but not to venture into the technical details of these fields. It
is clear that future quantum technology will be using many complementary plat-
forms. Optics will remain a major contributor and quite likely the most direct
approach to the fascinating and sometimes counterintuitive effects of the quan-
tum world.

Australia, 2018 Hans Bachor and Timothy Ralph

 xix

Acknowledgments

This book would not have been possible without the teaching we received from
many of the creative and motivating pioneers of the field of quantum optics. We
have named them in their context throughout the book. HAB is particularly in
debt to Marc Levenson, who challenged him first with ideas about quantum noise.
From him he learned many of the tricks of the trade and the art of checking one's
sanity when the experiment produces results that seem to be impossible. TCR
wishes particularly to thank Craig Savage for instilling the importance of doing
theory that is relevant to experimentalists. Both of us owe a great deal to Dan
Walls and Gerard Milburn who provided such beautiful vision of what might
be possible.
Over the years we learned from and received help for this book from a large
international team of colleagues and students. Too many to name individually.
All of you made important contributions to the our understanding and the way
we communicate our knowledge and sight. We enjoy the feedback and are keen
to discuss the mysteries and the past and future of quantum optics. We like to
thank Aska Dolinska for the creative graphics work.
Most importantly this Third Edition would not have been possible without the
support of our loved ones. TCR wishes to thank his wife Barbara for her support
and son Isaac for putting up with a distracted father. And HAB wishes to thank
his wife Cornelia for the patience she showed with an absent-minded and many
times frustrated writer.

Hans Bachor
Timothy Ralph 2018
 1

Introduction

1.1 Optics in Modern Life

The invention of the laser and the use of the special properties of laser light
created one of the great technical revolutions of the twentieth century and has
impact on many aspects of our modern lives. It allows us to manufacture the
gadgets we love to use, including the computer with which this has been written.
It allows us to communicate literally at the speed of light and with an information
bandwidth that is unprecedented. Optics has been for a while the technology of
choice for data storage, with DVD and Blu-ray still being used as a medium for
data and 3D optical storage that might emerge in future. The modern alternatives,
downloading and streaming, depend in many ways on photonics.
In addition, there is a plethora of other applications of light, from laser weld-
ing in manufacturing to laser fusion for power generation and from laser-based
microscopes to laser surgery. Imaging and display technologies are in many gad-
gets. The developments of optical technology have been relentless and will con-
tinue to be so for many more years. This has allowed us to access new levels
of power, controllability, accuracy, complexity, and low cost that were deemed
impossible only a few years ago. This is a convincing reason to study and teach
quantum optics, as a scientist or engineer.
Whilst all of the techniques in photonics use the quantum concept of stimu-
lated emission and the laser [1], most of these applications still use quantum ideas
in a quite simple and peripheral way. Most of the designs are based on conven-
tional optics and can be understood on the basis of wave optics and traditional
physical optics [2]. An interpretation of light as a classical wave is perfectly ade-
quate for the understanding of effects such as diffraction, interference, or image
formation. Non-linear optics, such as frequency doubling or wave mixing, is well
described by classical theory. Even most of the properties of coherent laser light
can be understood and modelled in this way. There is a wealth or new ideas and
applications emerging, exploiting the regimes of wave optics that were previously
not accessible, with new theories and devices. This includes the many fascinating
new devices made possible through new engineered materials, waveguide struc-
tures, photonic bandgap materials, linear and non-linear metamaterials. They all
hold the promise of even higher performance, better specifications, and even new
applications, such as cloaking or 3D storage, optical data processing at unheard

A Guide to Experiments in Quantum Optics, Third Edition. Hans-A. Bachor and Timothy C. Ralph.
© 2019 Wiley-VCH Verlag GmbH & Co. KGaA. Published 2019 by Wiley-VCH Verlag GmbH & Co. KGaA.
2 1 Introduction

speeds, or optical data processing. Conferences and research reviews all docu-
ment that in 2018 a bright future is ahead in these areas of optics.
However, which role will optics play in a future that will use quantum
technologies?
Firstly, the question of what ultimately limits the performance of optical devices
lies outside the classical theory of light. We have to focus on a different aspect of
optics. We follow the ideas that come directly from the quantum nature of light,
the full properties of photons, and the subtle effects of quantum optics [3]. Ini-
tially, optics was recognized as a pretty ideal medium to study the finer effects
of the quantum world. Photons in the visible spectrum were easily generated,
detected with high efficiency, and not masked by thermal noise at room temper-
ature, and any correlations could be recorded very well.
Non-linear effects allowed the generation of a variety of different forms of light.
This established optics as the medium of choice to explore the quantum world,
and the early improvements in the technology in the 1980s led to great demon-
strations, such as the Bell inequalities, signalling the beginning of experimental
quantum optics. This exploration of the ‘weirdness’ of the quantum world is still
today one of the frontiers in quantum optics and discussed in this guide.
It was recognized that quantum effects pose a limit to the sensitivity of many
measurements, with the now successful detectors for gravitational waves the
pre-eminent example. A closer analysis showed that there are ways of, at least
partially, circumventing such limitations. Optical quantum metrology was born
and is a major topic in this guide. It will continue to evolve – and we now see the
first routine applications of quantum-enhanced metrology, with few photons and
with laser beams. The ideas are spreading and can be realized in other systems
such as coherent groups of atoms or ions or in circuits. Quantum-enhanced
metrology in its various forms is becoming more widespread, from the definition
of the SI units to very practical situations.
Optics technology and our knowledge of protocols and devices have steadily
grown in time. Almost gone are the analogue devices of the 1990s, and we have
direct access to digital data. For example, we can now detect, record, process, and
store information in our experiments in much better ways, with A/D converters,
multichannel analysers, memories of Terabit size, etc. This means that we can
access much more information and process and analyse data more thoroughly.
The next step might well include the use of artificial intelligence to optimize data
analysis and also the performance of devices.
The detection of quantum correlations are a case in hand: we can now directly
explore the correlation properties, use different approaches and measures to
evaluate the quality of entanglement, and post-select data, or use heralded
information, to select the best quantum data. This provides a path to optical
quantum gates and more elaborate quantum logic. We can post-process the
information and make it conditional on other measurements. This allows us to
not only describe the state with tomography in great detail but also provide a
path for looking inside the statistical properties of a beam of light. An example
is the Wigner function shown on the front cover, which shows strong negative
values that are contained in one beam from an entangled pair of beam – after
 1.2 The Origin and Progress of Quantum Optics 3

conditional detection and letting information from one beam influence the
reading from the other entangled beam [4].
Furthermore, so-called hybrid detection uses entangled beams, where the
information from a single photon detector in one beam is used to post-select the
most relevant data from stream of photon recorded from the other beam. Using
this form of hybrid detection, it is possible to filter out states of light with specific
quantum properties, for example, Schrödinger cat states [5]. They represent a
state that is a quantum superposition of two orthogonal states and is one of the
icons and hallmarks of quantum physics, as recognized in the 2012 Nobel Prize
of Physics [6].
On the other hand, the last few years has seen an ever-increasing interest in the
information that is contained within a beam of light, whether it be images, ana-
logue pulses, or digital data encoded in the light. The emergence of the concept of
quantum information has created a rising interest in the ability to communicate,
store, and process more complex information than available in classical digital
electronics, all by using beams of light or individual photons. This has created
a great deal of interest, an expanding research field, and a demand for practical
quantum optics-based devices. Optics overlaps more and more with condensed
matter and soft matter physics, materials science, and mechanical engineering.
Ever-improving technology drives these fields, as well as now insights into quan-
tum logic and quantum control. Optics is in many cases a precursor, sometimes
a testing ground, and increasingly a component of such quantum devices.

1.2 The Origin and Progress of Quantum Optics

Ever since the quantum interpretation of the black body radiation by M. Planck
[7], the discovery of the photoelectric effect by W. Hallwachs [8] and P. Lennard
[9] and its interpretation by A. Einstein [10] has the idea of photons been used to
describe the origin of light. For the description of the generation of light, a quan-
tum model is essential. The emission of light by atoms, and equally the absorption
of light, requires the assumption that light of a certain wavelength 𝜆, or frequency
𝜈, is made up of discrete units of energy each with the same energy hc∕𝜆 = h𝜈.
This concept is closely linked to the quantum theory of the atoms themselves.
Atoms are best described as quantum systems; their properties can be derived
from the wave functions of the atoms. The energy eigenstates of atoms lead
directly to the spectra of light which they emit or absorb. The quantum theory of
atoms and their interaction with light has been developed extensively; it led to
many practical applications [11] and has played crucial role in the formulation
of quantum mechanics itself. Spectroscopy relies almost completely on the
quantum nature of atoms. However, in this guide we will not be concerned with
the quantum theory of atoms, but concentrate on the properties of the light, its
propagation, and its applications in optical measurements.
Historically, special experiments with extremely low intensities and those that
are based on detecting individual photons raised new questions, for instance, the
famous interference experiments by G.I. Taylor [12] where the energy flux corre-
sponds to the transit of individual photons. He repeated Th. Young’s double-slit
4 1 Introduction

experiment [13] with typically less than one photon in the experiment at any one
time. In this case, the classical explanation of interference based on electromag-
netic waves and the quantum explanation based on the interference of probability
amplitudes can be made to coincide by the simple expedient of making the prob-
ability of counting a photon proportional to the intensity of the classical field.
The experiment cannot distinguish between the two explanations. Similarly, the
modern versions of these experiments, using the technologies of low noise cur-
rent detection or, alternatively, photon counting, show the identity of the two
interpretations.
The search for uniquely quantum optical effects continued through experi-
ments concerned with intensity, rather than amplitude interferometry. The focus
shifted to the measurement of fluctuations and the statistical analysis of light.
Correlations between the arrival of photons were considered. It started with the
famous experiment by R. Hanbury-Brown and R. Twiss [14] who studied the cor-
relation between the fluctuations of the two photocurrents from two different
detectors illuminated by the same light source. They observed with a thermal
light source an enhancement in the two-time intensity correlation function for
short time delays. This was a consequence of the large intensity fluctuations of
the thermal light and was called photon bunching. This phenomenon can be
adequately explained using a classical theory that includes a fluctuating electro-
magnetic field. Once laser light was available, it was found that a strong laser
beam well above threshold shows no photon bunching; instead the light has a
Poissonian counting statistics. One consequence is that laser beams cannot be
perfectly quiet; they show noise in the intensity that is called shot noise, a name
that reminds us of the arrival of many particles of light. This result can be derived
from both classical and quantum models.
Next, it was shown by R.J. Glauber [15] that additional, unique predictions
could be made from his quantum formulation of optical coherence. One such
prediction is photon anti-bunching, where the initial slope of the two-time cor-
relation function is positive. This corresponds to greater than average separa-
tions between the arrival times of photons; the photon counting statistics may
be sub-Poissonian, and the fluctuations of the resulting photocurrent would be
smaller than the shot noise. It was shown that a classical theory based on fluc-
tuating field amplitudes would require negative probabilities in order to predict
anti-bunching, and this is clearly not a classical concept but a key feature of a
quantum model.
It was not until 1976, when H.J. Carmichael and D.F. Walls [16] predicted that
light generated by resonance fluorescence from a two-level atom would exhibit
anti-bunching, that a physically accessible system was identified that exhibits
non-classical behaviour. This phenomenon was observed in experiments by
H.J. Kimble et al. [17], where individual atoms were observed. This opened the
era of quantum optics. It is now possible to track and directly display the evolu-
tion of the photon number inside a cavity, thanks to the innovation introduced
by S. Haroche and coworkers. They have designed better and better supercon-
ducting cavities to store microwave photons and can carry out non-demolition
experiments in a cavity with very long decay time. They can now directly
observe the evolution of the photon number in an individual mode [18]. See
 1.2 The Origin and Progress of Quantum Optics 5

for details Figure 3.7. This is a most beautiful demonstration of the theoretical
concepts.
The concept of modes will be a central feature in this guide. This concept has
evolved in time; it now has several distinct meanings and is playing a pivotal
role in quantum optics. For an experimentalist, a single mode can mean a per-
fect beam shape without any distortion. A single-mode laser means a device that
is emitting a beam with exactly one frequency, in one polarization, and with a
perfect beam shape. In the theory, a mode is the fundamental entity that can be
quantized and be described by simple operators. We have a reliable and direct
model for the behaviour of such a single mode, and up to recently most quantum
optics were formulated in this single-mode model.
At the same time, in optical technology, such as communication or imaging,
the concept of a mode is associated with an individual channel of information.
This could be a specific modulation frequency, as we are familiar with from radio
transmission, or it could be a specific spatial image. In the world of optical sens-
ing, we wish to detect and transmit changes of various parameters, such as lengths
or position, magnetic or electric fields, pressure or temperature, to name a few.
The evolution of these parameters are then encoded into the light and transmit-
ted to the receiver. In this case, each parameter, and each degree of freedom for
the change of the parameter, corresponds to one specific mode. It has now been
recognized that a sensor can be optimized and will show the best possible perfor-
mance, when the optical mode is fully matched to the information. This can be a
simple mode, as mentioned in the case of the traditional single-mode laser, or it
can actually be a much more complicated spatial and temporal distribution.
The first extension in quantum optics has been to multiple beams of light, each
containing one mode each. We can have many separate beams, all originating
from one complex optical apparatus. However, a single optical beam can also con-
tain multiple modes, for example, spatial modes, temporal modes, or frequency
modes. They correspond to separate degrees of freedom or separate information
channels within the beam. Such a multimode beam might contain only a few, or
the occasional, photon, or it might have the full flux of a laser beam. In either case,
the underlying physics is the same. The information transmitted is best described
in terms of a set of independent orthogonal modes. Each can be quantized individ-
ually and has their own independent statistics. The signals, and the fluctuations,
transmitted by these modes do not interact with each other, and the full quantum
description of these devices is straightforward using this concept.
There has been great progress in the technology to generate individual photons
in a controlled way. The technology has evolved to a stage where many of the ini-
tial thought experiments dealing with individual atoms can now be performed
with real machines. In addition, a whole range of new types of single photon
emitters has been developed and studied by many groups around the world. They
include quantum dots that show the required quantum properties and are con-
tinuously improving in their robustness and reliability. They are now being used
regularly in a variety of applications where small size matters. This includes appli-
cations for quantum communication with quantum dots placed in specific places
or in the form of nanodiamonds imbedded as emitters in biological samples that
respond to the local conditions in living cells.
6 1 Introduction

Initially, these single photon effects were a pure curiosity. They showed the
difference between the quantum and the classical world and were used to test
quantum mechanics. In particular, the critique by A. Einstein et al. and their EPR
paradox [19] required further investigation. Through the work of J.S. Bell [20], it
became possible to carry out quantitative tests. They are all based on the quan-
tum concept of entanglement. In the simplest case, this means pairs of quantum
systems that have a common origin and have linked properties, such that the
detection of one allows the perfect prediction of the other. Such systems include
not only individual photons or atoms, or coherent states of light, such as laser
beams but also physical matter, such as was demonstrated with Bose–Einstein
condensates. See Chapter 14.
Starting with the experiments by A. Aspect et al. [21] with atomic sources, and
later with the development of sources of twin photon pairs and experiments by
A. Zeilinger [22] and many others, the quantum predictions have been extremely
well tested. The concept of entanglement has now been demonstrated and tested
with single photons and also with intense beams of light, over large distances of
many kilometres and through optical fibres. There has been a long series of exper-
iments with the aim to demonstrate the validity and necessity of quantum physics
beyond any doubt, closing step by step every loophole. It has even been possible
to extend the concept of entanglement from two entities, such as two particles, to
many particles, most prominently to many trapped ions. Such multipartite entan-
glement is at the core of plans to build quantum simulators and quantum logic
devices. Similarly there has been an extension from two modes of light to several
modes. This leads to the idea of multimode entanglement, which is a new topic of
research, with both interesting physics and the potential for new applications.
In summary, the progress in quantum optics, which initially was concerned
with the quality of the quantum demonstrations, has recently led to the con-
quering of complexity and the study of the robustness, or decoherence, of such
complex systems. The peculiarities of single photon quantum processes have led
to many new technical applications, mainly in the area of communication. For
example, it was found that information sent by single photons can be made secure
against listening in; the unique quantum properties do not allow the copying of
the information. Any eavesdropping will introduce noise and can therefore be
detected. Quantum cryptography and quantum key distribution (QKD) [23] are
now viable technologies, based on the inability to clone single photons or quan-
tum modes.
There is great anticipation of other future technological applications of pho-
tons. The superposition of different states of photons can be used to send more
complex information, allowing the transfer not only of classical bits of informa-
tion, such as 0 and 1, but also of qubits based on the superposition of states 0
and 1. The coherent evolution of several such quantum systems could lead to
the development of quantum logic connections, quantum gates, and eventually
whole quantum computers that promise to solve certain classes of mathemat-
ical problems much more efficiently and rapidly [24]. Just as astounding is the
concept of teleportation [25], the disembodied communication of the complete
quantum information from one place to another. This has already been tested in
experiments (see Chapter 13).
 1.3 Motivation Through Simple and Direct Teaching Experiments 7

Optics is now a testing ground for future quantum information technologies

and for data communication and processing. At the same time, the concepts and
ideas of quantum optics are emerging in other related quantum technologies,
for example, based on quantum circuits or mechanical systems, which are all
quantized and show in principle the same physics. We can see the equivalence
of the physics of photons, phonons, polaritons, and other particles that describe
the excitation of quantum systems. Technology is moving very fast, and we can
expect over the next decade to see machines that utilize any of these individual
quantum systems or even combine several of them. The rules of quantum optics,
as described in this guide, will become important in many different other types of
instruments. Optics could become one of the best ways to visualize the broader
quantum concepts that apply to many other types of instruments.

1.3 Motivation Through Simple and Direct Teaching

Experiments
The best starting point for the demonstration of photons in current technology
is to start with images that are generated by CCD (charge-coupled device) cam-
eras. It is worthwhile to note that basically all information we have about light
and optical effects is based on either some form of photodetector or the human
eye. The first generates an electric signal, either pulses or a continuous photon
current. Whilst the eye produces, in a clever and complex way, a signal in the
neurones in the brain, even they can be interpreted as a form of electric signal.
It is the characteristics of these signals and their statistics that provides access to
the quantum properties of light.
In the extreme case we would have a detector that is capable of detecting one
individual photon at a time. This is now possible with special CCD cameras,
where the sensitivity has been lifted to such a high level and thermal noise has
been reduced so much that it hardly interferes with the observation. The num-
ber of dark counts, that is, clicks that are generated at random without photons
present, can be neglected. In this way we can actually detect and record images
at the single photon level as shown in Figure 1.1.
The human eye is actually capable of seeing at this level, and many animals
exceed our capability. At this level of illumination, the world looks very grainy. We
can see individual events. The image is composed out of individual local detec-
tion events. An example, taken with a special CCD, is shown in Figure 1.1. At
some level this can be regarded as trivial – obviously it will be like this. We have
photons. On the other this image demonstrates several subtle points. We can see
that the concept of intensity, bright and dark, is directly related to the concept of
probability. Areas with a high local photon flux, or high probability, correspond
to high intensities, that is, a high amplitude of the electromagnetic field. Con-
versely, dark areas of the image correspond to a low intensity. The central concept
in quantum science of a probability amplitude become directly obvious.
Two, or many, waves with different amplitudes and relative phases can inter-
fere, and this is a central concept of the classical wave picture of light. The example
8 1 Introduction

(a)

300

250
Number of photons

200

150

100

50

0
0 50 100 150 200
(b) Pixel

Figure 1.1 Image of the fringes from a double-slit experiment recorded at the single photon
level. (a) Distribution of individual photons. (b) Histogram of photons in each vertical line. The
full movie can be seen at https://www.lcf.institutoptique.fr/Groupes-de-recherche/Gaz-
quantiques/Membres/Permanents/Alain-Aspect. Source: After https://www.lcf.institutoptique
.fr/Groupes-de-recherche/Gaz-quantiques/Membres/Permanents/Alain-Aspect.

shown in Figure 1.1 is a real interference fringe system, generated with coherent
light and a double-slit apparatus. This is one of the simplest arrangements that
shows the wave nature of light – and yet in this case it is detected by a num-
ber of individual photons. There is no contradiction here – the two explanations,
waves and particles, are fully complementary. The shape of the fringes, the over-
all intensity function, is best described with the wave model. At the same time,
the statistics of the light, the visibility that is observed, and thus the quality of the
fringes can best be described with the photon model. This very simple example
immediately shows the strengths and weaknesses of both models. It demonstrates
how we want to use these two models for different aspects of the experiment. We
should choose the pictures in our mind that is most suitable to understand the
specific aspect of the experiment we wish to investigate.
The next level would be to quantify the statistics. What would be an appro-
priate statistical model for the fluctuations, detected in one specific area of the
image and averaged over many time intervals? Equivalently we could ask: what
is the statistics of the number of photons detected in many separate areas of
equal size, all receiving the same intensity? In the first case one could imagine
listening to the number of clicks being generated. It would sound like a random
sequence of clicks, as we know them from a Geiger counter detecting 𝛼 or 𝛾 rays.
 1.3 Motivation Through Simple and Direct Teaching Experiments 9

A proper analysis would reveal that we have a Poissonian distribution for the
different time intervals. The same result applies to the analysis of the number of
photons in many image areas of equal size. The graininess we see in Figure 1.1
is a direct representation of the counting statistics, which can be derived from
simple statistical arguments, assuming random processes.
The randomness in the arrival times of the photons in a weak beam of light is
so perfect that it is now used as the gold standard for randomness, exceeding in
quality the widely used random number generators based on electronic processes
or imbedded in computers. This has led to commercial activities where optically
generated random numbers now can be accessed via the internet and the gener-
ators are for profit. The demand is substantial – and already a small niche market
for quantum random numbers has emerged.
The 1960s saw a rapid development of new laser light sources and improve-
ments in light detection techniques. This allowed the distinction between inco-
herent (thermal) and coherent (laser) light on the basis of photon statistics. The
groups of F.T. Arecchi [26], L. Mandel, and R.E. Pike all demonstrated in their
experiments that the photon counting statistic goes from super-Poissonian at the
lasing threshold to Poissonian far above threshold. The corresponding theoretical
work by R.J. Glauber [15] was based on the concept that both the atomic variables
and the light are quantized and showed that light can be described by a coherent
state, the closest quantum counterpart to a classical field. The results are essen-
tially equivalent to a quantum treatment of a classical oscillator. However, it is
an important consequence of the quantum model that any measurement of the
properties of this state, intensity, amplitude, or phase of the light, will be limited
by these fluctuations. This is the optical manifestation of Heisenberg’s uncertainty
principle.
Quantum noise can impose a limit to the performance of lasers, sensors, and
communication systems, and near the quantum limit the performance can be
quite different. To illustrate this, consider the result of a very simple and practical
experiment: use a laser, such as a laser pointer with a few milliwatt of power,
and detect all of the light with a photodiode. The average number of detection
events is now very large. Listening to the output, we can no longer distinguish
between the individual clicks; we will hear a continuous noise. The level of
noise will increase with the intensity, as the width of the Poissonian distribution
increases with the average number of photons detected. This quantum noise
is the direct experimental evidence of the quantum statistics of light. If we
record the current from the photodetector on an oscilloscope, we would see
a trace with random fluctuations, above and below a mean level, representing
the average intensity. The excursions have a Gaussian distribution around the
mean value, which is fully consistent with the transition from a Poissonian to a
Gaussian distribution. Relatively speaking this effect gets smaller if we average
the signal over more photons, that means using a brighter laser or measuring for
a longer time. As we will see, this effect cannot be avoided completely. Quantum
noise is not an effect that can be eliminated by good traditional engineering.
Decades ago quantum noise was called shot noise and was regarded by many as
a consequence of the photodetection process, as a randomness of the stream of
electrons produced in the photodetector. This view prevailed for a long time,
10 1 Introduction

Modulator

A
Laser

B X2

X1

Figure 1.2 Experiment to investigate the correlation between two parts of a laser beam, using
a laser pointer, modulator, two detectors, and an x–y oscilloscope.

particularly in engineering textbooks. However, this view is misleading as we

find in the squeezing experiments described in this book. It cannot account for
situations where non-linear optical processes modify the quantum noise, whilst
the detector remains unchanged. In the squeezing experiments the quantum
noise is changed optically, and consequently we have to assume it is a property
of the light, not of the detector.
Today it is straightforward to see these effects of quantum noise with simple
equipment, accessible to undergraduate students. The light source can be a laser
pointer, carefully selected to operate in a single mode. The laser beam is split
by a beamsplitter into two beams of equal power that are detected by two indi-
vidual detectors, A and B, at different locations. Each photocurrent is amplified,
and they are both displayed via the x and y axes of one oscilloscope. We want to
see both the average (DC) value of the photocurrent, which represents the time
averaged intensity of the beams A and B, and the fluctuations (AC) of the two
photocurrents, as they represent any modulation and the fluctuations, or noise,
of the intensity of the two beams. This can easily be done by looking at the signals
at a few megahertz, which is typically the bandwidth of the RF amplifiers used.
This is shown in Figure 1.2.
The laser can be modulated, through periodic variations of the laser current.
Classical optics predicts that the intensity of both beams A and B will be mod-
ulated, and this will show up clearly as a line at 45∘ on the oscilloscope. That
means the two photocurrents change synchronously in time, an increase in one
is accompanied by an increase in the other etc. In practice some care is required to
achieve a clear 45∘ line: there should be no delay in the signals from the detector
to the oscilloscope and the size of the signals has been matched. This result shows
a strong correlation between the two beams. A correlation of the photocurrents
means a correlation of the flux of photons.
We can also look at the fluctuations in more detail. If we look at beams A and
B individually, we see fluctuations of the current above and below the long-term
average value. Closer inspection would show a Gaussian distribution of the excur-
sions below and above, as we expect it for random noise in the beam. The mag-
nitude of the fluctuations will follow the gain curve of the RF amplifiers. With
careful calibration of the frequency response of our equipment, we find that the
noise in the light is not frequency dependent; it is white noise. The magnitude of
the fluctuations is the same for a laser beam with or without a small modulation.
 1.3 Motivation Through Simple and Direct Teaching Experiments 11

A A C
C
X2 X2

X1 X1

(a) (b)
A
C
X2

X1

(c)

Figure 1.3 The noise of beams A and B displayed on an oscilloscope. (a) No beam, technical
background noise. (b) The laser beam has no modulation, quantum noise. (c) The laser beam
with modulation. The signal from the two detectors is displayed in the coordinates X1 and X2 ,
correlations appear in the direction C, and uncorrelated signal appears in both directions A
and C.

The noise increases with intensity, and careful measurements show that the vari-
ance of the noise is proportional to the intensity.
Is the noise from A correlated with the noise from B? The oscilloscope image
immediately provides the answer. The noise from A and B results in a fuzzy blob
on the screen. See Figure 1.3b. The fluctuations in the two photocurrents are
independent. The point on the screen of the oscilloscope traces out a random
walk independently in both directions. This is in stark contrast to the modulation
where the photocurrent fluctuates synchronously, as shown in Figure 1.3c. The
lack of correlation is immediately evident. This confirms the special properties of
quantum noise; it does not obey the classical rules.
One simple explanation is that for small fluctuations we would notice the effect
that one photon cannot be split into halves at the beamsplitter. It will appear in
one or the other beam, with a random choice which beam it will join. We will see
in Chapter 3 how this simple statement leads to an explanation of the properties
of quantum noise in this situation. If we have no modulation, the blob is right in
the middle of the screen. The blob is circular, given there is no additional noise in
the system and the photon noise dominates over the electronic noise generated
in the apparatus. For a laser beam with modulation, we will see a fuzzy area tilted
at 45∘ , as shown in Figure 1.3c. This is a combination of the two effects, a line
12 1 Introduction

in direction C for the modulation and a fuzzy blob for the laser noise. Classical
modulations and quantum noise have different properties, even when they have
similar magnitudes, and this simple experiment can show the difference.

1.4 Consequences of Photon Correlations

It is worthwhile to explore the properties of quantum noise somewhat further.
It was found that, in clear distinction to classical noise, no technical trick can
eliminate quantum noise. This becomes evident from our experiment. A com-
mon conventional idea for noise suppression would be to make two copies of the
signal, both containing the noise, and to subtract them for each other. We have all
the necessary components in the apparatus used in Figure 1.2. We have made two
copies of the beam. We can use one beam, say, A, for our experiment, record it
with detector A, and then subtract the electronic fluctuations generated by detec-
tor B. Based on our observations we now see that we can subtract the classical
modulation, which is common to both beams. However, we cannot subtract the
quantum noise, which is independent and uncorrelated.
The subtraction works for any modulation, for harmonic modulation demon-
strated here, and for random intensity modulations, such as those generated by a
low-quality, noisy current source for the laser. That means we can distinguish
between classical laser noise and quantum noise that is a consequence of the
quantum properties of the photons. The result remains unchanged if the sub-
traction is replaced by a sum or if the two currents, or if the two currents are
added with an arbitrary short time delay, such as introduced be an extra length of
cable. The resulting noise is always the quadrature sum of the two-input signal. It
is independent of the sign, or phase, of the summation. This is equivalent to the
statement that the noise in the two photocurrents is not correlated.
At this stage it is not easy to identify the point in the experiment where this
uncorrelated noise is generated. One interpretation assumes that the noise in
the photocurrents is generated in the photodetectors. A different interpretation
assumes that the beamsplitter is a random selector for photons and consequently
the intensities of the two beams are random and thus uncorrelated. A distinc-
tion can only be made by further experiments with squeezed light, which are
discussed later in this book.
An alternative scheme for noise suppression is the use of feedback control, as
shown in Figure 1.4. It achieves equivalent results to difference detection. The

Modulator

Laser

Feedback control

Figure 1.4 The second attempt to eliminate laser noise: an improved apparatus using a
feedback controller, or ‘noise eater’.
Another random document with
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Mouth-Frame.—In its broad outlines there is practically no variation
in this organ throughout the group, but in respect of the spines,
which are borne on the flanks of the jaws (mouth-papillae) and on
their apices (teeth and tooth-papillae) there is very great variation.
Teeth are always present. Mouth-papillae are very frequently
present, tooth-papillae are rarer, and it is only in a restricted number
of genera (Ophiocoma and its allies) that both mouth-papillae and
tooth-papillae are present at the same time.

Fig. 216.—A portion of an arm of Ophiohelus umbella, near the distal extremity,
treated with potash to show the skeleton, × 55. The vertebrae are seen to
consist of two curved rods united at their ends. The triangular side-plates
bear a row of movable hooks which articulate with basal outgrowths of the
plate. (After Lyman.)

Skeleton of the Disc.—This is typically composed of a mosaic of

plates of different sizes, but in some cases (Ophiomyxa, most
Streptophiurae, and Cladophiurae) these, with the exception of the
radials and genitals, are entirely absent, and the disc is then quite
soft and covered with a columnar epithelium, the persistent
ectoderm. Even the scuta buccalia may disappear. Radial shields
are absent in Ophiohelus. In many cases (Ophiothrix and
Ophiocoma) all the dorsal plates except the radials are concealed
from view by a covering of small spines. In some genera
(Ophiopyrgus) there are five large plates in the centre of the upper
part of the disc, which have been termed "calycinals" from a
mistaken comparison with the plates forming the cup or calyx of the
Pelmatozoa, but there is no connexion between the two sets of
structures.
The madreporite is usually quite rudimentary, but in Cladophiurae
there may be five madreporites, each with about 200 pores, and, of
course, five stone-canals.

The number of genital organs varies very much. In the small

Amphiura squamata there are two gonads, an ovary and a testis,
attached to each bursa, but in the larger species there may be very
many more.

We follow Bell's classification,[463] according to which the

Ophiuroidea are divided, according to the manner in which the
vertebrae move on one another (cf. Fig. 210), into three main orders,
since these movements are of prime importance in their lives.

(1) Streptophiurae, in which the faces of the vertebrae have

rudimentary knobs and corresponding depressions, so that the arms
can be coiled in the vertical plane. These are regarded as the most
primitive of Ophiuroidea.

(2) Zygophiurae, in which the vertebral faces have knobs and pits
which prevent their coiling in a vertical plane.

(3) Cladophiurae, in which the arms can be coiled as in (1) and are
in most cases forked. No teeth; the arm-spines are papillae, the
covering plates of the arms are reduced to granules.

Order I. Streptophiurae.
This is not a very well defined order; it includes a few genera
intermediate in character between the Cladophiurae and the
Zygophiurae, and believed to be the most primitive Ophiuroids living.
It is not divided into families. The vertebrae have rudimentary
articulating surfaces, there being two low bosses and corresponding
hollows on each side, and so they are capable of being moved in a
vertical plane, as in the Cladophiurae; the arms never branch, and
further, they always bear arm-spines and lateral arm-plates at least.
No species of this order are found on the British coast, but
Ophiomyxa pentagona, in which the dorsal part of the disc is
represented only by soft skin, is common in the Mediterranean.

Ophioteresis is devoid of ventral plates on the arms, and appears to

possess an open ambulacral groove, though this point has not been
tested in sections. Ophiohelus and Ophiogeron have vertebrae in
which traces of the double origin persist (see p. 491).

Order II. Zygophiurae.

This group includes all the common and better-known British forms.
They are divided into five families, all of which are represented in
British waters.

Fig. 217.—Aboral view of Ophioglypha (Ophiura) bullata. × 3. (From Wyville

Thomson.)
Fam. 1. Ophiolepididae.[464]—Arm inserted in a definite cleft in the
disc, or (expressing the same fact in another way) the interradial
lobes out of which the disc is composed are not completely united.
Radial shields and dorsal plates naked. Arm-spines smooth and
inserted on the posterior border of the lateral arm-plates.

Fig. 218.—Oral view of Ophioglypha (Ophiura) bullata. × 5. (From Wyville

Thomson.)

This family includes all the Brittle Stars of smooth porcelanous

aspect and provided with only short spines. Forbes[465] called them
Sand-stars, since their short spines render these animals incapable
of burrowing or of climbing well, and hence they appear to move
comparatively rapidly over firm ground, sand, gravel, or muddy sand,
and they are active enough to be able to capture small worms and
Crustacea. The prey is seized by coiling one of the arms around it.

One genus, Ophiura, is fairly common round the British coast, being
represented by O. ciliaris and O. albida; the former is the commoner.
An allied species dredged by H.M.S. "Challenger" is represented in
Figs. 217 and 218.

Ophiomusium (Fig. 219) is a very peculiar genus. The mouth-

papillae on each side of each mouth-angle are confluent, forming a
razor-like projection on each side of each mouth-angle (Fig. 220).
The arms are short, and the podia are only developed at the bases
of the arms. Ophiopyrgus has the dorsal surface raised into a conical
elevation protected by a central plate surrounded by five large
plates.

Fig. 219.—Aboral view of Ophiomusium pulchellum. × 7. (From Wyville

Thomson.)

In the remaining four families the arms are inserted on the under
surface of the disc; in other words, the interradial lobes which make
up the disc have completely coalesced dorsally; and the spines
stand out at right angles to the arm.

Fam. 2. Amphiuridae.—Mouth-papillae present, but no tooth-

papillae; radial shields naked; small scuta buccalia.

The most interesting Brittle Star belonging to this family is Amphiura

squamata (elegans), a small form, with a disc about ¼ inch in
diameter covered with naked plates. It is hermaphrodite and
viviparous, the young completing their development inside the
bursae of the mother. Occasionally the whole disc, with the
exception of the mouth-frame, is thrown off and regenerated. This
appears to be a device to enable the young to escape. Three other
species of Amphiura are found in British waters.

Fig. 220.—Oral view of Ophiomusium pulchellum. × 7. (From Wyville Thomson.)

Ophiactis is another genus belonging to this family, distinguished

from Amphiura by its shorter arms and smoother arm-spines. It lives
in the interstices of hard gravel. The British species, O. balli,
presents no special features of interest, but the Neapolitan O. virens
is an extraordinary form. It has six arms, three of which are usually
larger than the other three, for it is always undergoing a process of
transverse division, each half regenerating the missing part. It has
from 1 to 5 stone-canals, the number increasing with age; numerous
long-stalked Polian-vesicles in each interradius, and in addition a
number of long tubular canals which spring from the ring-canal, and
entwine themselves amongst the viscera.[466] All the canals of the
water-vascular system, except the stone-canals, contain non-
nucleated corpuscles, carrying haemoglobin,[467] the respiratory
value of which compensates for the loss of the genital bursae, which
have entirely disappeared.

Ophiopholis is distinguished from the foregoing genera by the

granular covering of its dorsal plates; whilst in Ophiacantha these
granules develop into prominent spinelets, and the arm-spines are
also thorny. Ophiopholis aculeata occurs in swarms in the branches
of the Firth of Clyde, and presents a most remarkable series of
variations in colour. Ophiopsila is a closely allied form, distinguished
by its large peristomial plates.

Fig. 221.—Oral view of Ophiacantha chelys. × 4. (From Wyville Thomson.)

Fam. 3. Ophiocomidae.—Both mouth-papillae and tooth-papillae

are present;[468] the arm-spines are smooth, and the disc is covered
with granules.

Ophiocoma nigra is the only common British representative of this

family. In this species the plates of the dorsal surface are completely
hidden from view by a covering of granules. Ophiarachna.

Fam. 4. Ophiothricidae.—Tooth-papillae alone present, mouth-

papillae absent; arm-spines roughened or thorny.

This family is represented only by Ophiothrix fragilis, which is

perhaps the most abundant of all British Ophiuroids, and has been
selected as the type for special description.

The back is covered with spinules, having, however, the triangular

radial plates bare. This produces a contrast-effect, which suggested
the name pentaphyllum, formerly used by some naturalists for the
species. It occurs in swarms, and presents variations in colour nearly
as marked as those of Ophiopholis. Ophiopteron is probably a
swimming Ophiuroid, as the lateral spines of each segment of the
arm are connected by a web of skin.

Order III. Cladophiurae.

Fig. 222.—Aboral view of young Astrophyton linckii, slightly enlarged. (From

Wyville Thomson.)

These, like the Streptophiurae, have the power of rolling the arms in
a vertical plane, but the articulating surfaces of the vertebrae are
well-developed and saddle-shaped. The dorsal surface of the disc
and arms is covered with a thick skin with minute calcifications.
Upper-arm plates wanting. Radial plates always present, though
occasionally represented by lines of scales. The order is divided into
three families, two of which are represented in British waters.

Fam. 1. Astroschemidae.—Arms unbranched. Astronyx is

comparatively common in the sea-lochs of Scotland. There are a
series of pad-like ridges on the arms, representing the side-plates
and bearing the spines. Astroschema.
Fam. 2. Trichasteridae.—Arms forked only at the distal ends.
Trichaster, Astrocnida.

Fam. 3. Euryalidae.—Arms forked to their bases. Gorgonocephalus

is occasionally taken in deep water off the north coast of Scotland. In
it the arms repeatedly fork, so that a regular crown of interlacing
arms is formed. The animal obviously clings to external objects with
these, for it is often taken in fishermen's nets with its arms coiled
around the meshes. The genital bursae are said to be represented
by slits which open directly into the coelom. (Lyman describes the
coelom as divided into ten compartments by radiating septa; it is
possible—even probable—that these are really the bursae.) An allied
species is common in the Bay of Fundy, being found in
comparatively shallow water. Astrophyton (Fig. 222) is closely allied
to Gorgonocephalus, differing only in trifling points. It is doubtful
whether the separation of these two genera is justified.

Fossil Ophiuroidea.—The Ophiuroidea are rather sparsely

represented among fossils, but in the Silurian and Devonian a series
of very interesting forms occur which are intermediate in character
between Starfish and Brittle Stars, and which were therefore in all
probability closely allied to the common ancestors of modern
Ophiuroids and Asteroids. Jaekel[469] has recently added largely to
our knowledge of these primitive forms, and has described a number
of new genera. Thus Eophiura from the Lower Silurian has an open
ambulacral groove, and the vertebrae are represented by an
alternating series of quadrate ossicles, each deeply grooved on its
under surface for the reception of the tentacle, which was not yet (as
in modern forms) enclosed in the vertebra. The lateral or
adambulacral plates extended horizontally outwards, and each bore
a series of spines at its outer edge.

A remarkable fact is that where the halves of the vertebrae (i.e. the
ambulacral ossicles) diverge in order to form the mouth-angles, no
less than five or six vertebrae are thus affected, instead of only two
as in modern forms. The actual "jaw," however, seems, as in modern
forms, to consist only of the first adambulacral fused to the second
ambulacral, so that instead of concluding with Jaekel that the "jaws"
of modern forms result from the fusion of five or six vertebrae, a
conclusion which would require that a number of tentacles had
disappeared, we may suppose that the gaping "angles" of these old
forms have, so to speak, healed up, except at their innermost
portions.

In Bohemura, which belongs to a somewhat younger stratum, the

structure is much the same, but the groove in the ambulacral ossicle
for the tentacle has become converted into a canal, and the
ambulacral groove itself has begun to be closed at the tip of the arm
by the meeting of the adambulacrals.

In Sympterura, a Devonian form described by Bather,[470] the two

ambulacral plates of each pair have thoroughly coalesced to form a
vertebra, but there is still an open ventral groove, and no ventral
plates.

In the Trias occurs the remarkable form Aspidura, which had short
triangular arms, in which the tentacle pores were enormous and the
ventral plates very small. The radial plates formed a continuous ring
round the edge of the disc. Geocoma from the Jurassic is a still more
typical Ophiuroid; it has long whip-like arms, and the dorsal skeleton
of the disc is made of fifteen plates, ten radials, and five interradials.
In the Jurassic the living genus Ophioglypha, appears.

The Cladophiurae are represented already in the Upper Silurian by

Eucladia, in which, however, the arms branch not dichotomously, as
they do in modern forms, but monopodially. There is a large single
madreporite.

Onychaster, with unbranched arms, which occurs in the

Carboniferous, is a representative of the Streptophiurae.
It will therefore be seen that the evolution of Ophiuroidea must have
begun in the Lower Silurian epoch. The Streptophiurae are a few
slightly modified survivors of the first Ophiuroids. By the time the
Devonian period had commenced, the division of the group into
Zygophiurae and Cladophiurae had been accomplished.

CHAPTER XVIII

ECHINODERMATA (CONTINUED): ECHINOIDEA = SEA-URCHINS

CLASS III. ECHINOIDEA

The Sea-urchins or Echinoidea (Gr. ἐχῖνος, Hedgehog or Sea-
urchin), which constitute the third class of the Eleutherozoa, have
derived both their popular and scientific names from the covering of
long spines with which they are provided. At first sight but little
resemblance is to be discerned between them and the Starfish and
Brittle Stars. They are devoid of any outgrowths that could be called
arms; their outline is generally either circular or that of an equilateral
pentagon, but as their height is almost always smaller than their
diameter, they are never quite spherical; sometimes it is so small
that the animals have the form of flattened discs.

All doubt as to the relationship of the Echinoidea to the Starfish is at

once dispelled in the mind of any one who sees one of the common
species alive. The surface is beset with delicate translucent tube-
feet, terminated by suckers resembling those of Starfish, although
capable of much more extension. The animal throws out these
organs, which attach themselves by their suckers to the substratum
and so pull the body along, whilst the spines are used to steady it
and prevent it from overturning under the unbalanced pull of the
tube-feet. When moving quickly the animal walks on its spines, the
tube-feet being little used. The tube-feet are distributed over five
bands, which run like meridians from one pole of the animal to the
other. These bands are termed "radii," and they extend from the
mouth, which is situated in the centre of the lower surface, up to the
neighbourhood of the aboral pole. The radii must be compared to the
ambulacral grooves on the oral surface of the arms of Starfish, and
hence in Urchins the aboral surfaces of the arms have, so to speak,
been absorbed into the disc, so that the oral surfaces have become
bent in the form of a semicircle. The radii are separated from one
another by meridional bands called "interradii," which correspond to
the interradial angles of the disc of a Starfish and to the sides of its
arms. The small area enclosed between the upper terminations of
the radii is called the "periproct," and this corresponds to the entire
dorsal surface of the Starfish, including that of the arms.

One of the commonest species of British Sea-urchin is Echinus

esculentus. In sheltered inlets, such as the Clyde, it is often left
exposed by the receding tide, whilst everywhere on the coast in
suitable localities it may be obtained by dredging at moderate depths
on suitable ground. In the Clyde it is easy to observe the habits of
the animal through the clear still water. It is then seen to frequent
chiefly rocky ground, and to exhibit a liking for hiding itself in
crevices. Often specimens will be seen clinging to the rock by some
of their tube-feet, and, as it were, pawing the under surface of the
water with the others. In the Clyde it feeds chiefly on the brown
fronds of Laminaria, with which the rocks are covered. In more
exposed situations, such as Plymouth Sound, it does not occur in
shallower water than 18 to 20 fathoms. At this depth it occurs on a
rocky ridge; but in 1899, after a south-west gale, all the specimens
had disappeared from this ridge, showing at what a depth wave
disturbance is felt.

A full-grown specimen is as large as a very large orange; its under

surface is flattened, and it tapers somewhat towards the aboral pole.
The outline is that of a pentagon with rounded angles. The spines in
Echinus esculentus are short in comparison to the diameter of the
body, and this is one of the characteristics of the species.
The animal is provided with a well-developed skeleton, consisting of
a mail of plates fitting closely edge to edge, and carrying the spines.
This cuirass bears the name "corona" (Fig. 227). It has two
openings, an upper and a lower, which are both covered with flexible
skin. The upper area is known as the "periproct" (Fig. 227, 2); it has
in it small isolated plates, and the anus, situated at the end of a small
papilla, projects from it on one side of the centre. The lower area of
flexible skin surrounds the mouth, and is called the "peristome" (Fig.
229), though it corresponds to considerably more than the peristome
of Asteroidea. In the mouth the tips of the five white chisel-like teeth
can be seen.

The plates forming the corona are, like all the elements of the
skeleton of Echinodermata, products of the connective tissue which
underlies the ectoderm, which in Echinoidea remains in a fully
developed condition covering the plates, and does not, as in
Ophiuroidea, dry up so as to form a mere cuticle. The ectoderm
consists of the same elements as that of Asteroidea, viz. delicate
tapering sense-cells with short sense-hairs, somewhat stouter
supporting cells and glandular cells. It is everywhere underlaid by a
plexus of nerve fibrils, which, in part, are to be regarded as the basal
outgrowths of the sense-cells and partly as the outgrowths of a
number of small bipolar ganglion-cells, found intermixed with the
fibres.

Fig. 223.—Aboral view of Echinus esculentus. × ½. (After Mortensen.)

Just as the muscular arm has been the determining factor in the
structure of the Ophiuroidea, so the movable spine has been the
leading factor in the evolution of Echinoidea. The spines have cup-
shaped basal ends, which are inserted on special projections of the
plates of the skeleton called tubercles. The tubercle is much larger
than the cup, and hence the spine has a great range of possible
motion. The spines differ from those of Starfish and Brittle Stars in
being connected with their tubercles by means of cylindrical sheaths
of muscle fibres, by the contraction of which they can be moved in
any direction. The muscles composing the sheath consist of an outer
translucent and an inner white layer. The former are easily
stimulated and soon relax; they cause the movements of the spines.
The latter require stronger stimulation, but when aroused respond
with a prolonged tetanus-like contraction, which causes the spines to
stand up stiffly in one position; these muscles can be torn across
sooner than forced to relax. Uexküll[471] has appropriately named
them "block musculature." These sheaths, like everything else, are
covered with ectoderm, which is, however, specially nervous, so that
we may say that the muscular ring is covered by a nerve-ring from
which stimuli are given off to the muscles.

The spines are, speaking generally, of two sizes, the larger being
known as "primary spines" and the smaller as "secondary." In many
Echinoidea these two varieties are very sharply contrasted, but in
Echinus esculentus there is not such a great difference in length,
and intermediate kinds occur. The forest of spines has an
undergrowth of pedicellariae. All Echinoidea possess pedicellariae,
which are much more highly developed than those of any Asteroid.
With few exceptions all the pedicellariae of Echinoidea possess
three jaws and a basal piece. This latter is, however, drawn out so as
to form a slender rod, which articulates with a minute boss on a plate
of the skeleton.

Of these pedicellariae there are in E. esculentus four varieties, viz.

(1) "tridactyle" (Fig. 225, C; Fig. 226, B): large conspicuous
pedicellariae with three pointed jaws, each armed with two rows of
teeth on the edges. There is a flexible stalk, the basal rod reaching
only half way up. These are scattered over the whole surface of the
animal.

(2) "Gemmiform" (Fig. 225, A, B; Fig. 226, A), so called from the
translucent, almost globular head. The appearance of the head is
due to the fact that there is on the outer surface of each jaw a sac-
like gland developed as a pouch of the ectoderm. From it are given
off two ducts which cross to the inner side of the blades and, uniting
into one, run in a groove to near the tip. The gland secretes a
poisonous fluid. The basal rod reaches up to the jaws, so that this
form of pedicellaria has a stiff stalk. On the inner side of each blade,
near the base, there is a slight elevation (Fig. 225, B, s), consisting
of cells bearing long cilia; this is a sense-organ for perceiving
mechanical stimuli. The gemmiform pedicellariae are particularly
abundant on the upper surface of the animal.

Fig. 224.—View of the apical region of Echinus esculentus, showing spines and
pedicellariae; drawn from the living specimen, × 3. a, Anus; g.p, genital
pore; i, interradius; mp, madreporite; per, periproct; p.gemm, gemmiform
pedicellaria; pod, podia; p.trid, tridactyle pedicellaria; p.trif, trifoliate
pedicellaria; r, radius; t.t, pore for terminal tentacle of the radial water-
vascular canal.
(3) "Trifoliate" (Fig. 225, E; Fig. 226, D): these are very small
pedicellariae, in which the jaws are shaped like leaves with the broad
end projecting outwards. They are scattered over the whole surface
of the body.

(4) "Ophicephalous" (Fig. 225, D; Fig. 226, C): pedicellariae in which

the jaws have broad rounded distal ends fringed with teeth; these
ends bear a resemblance to a snake's head, whence the name. The
bases are also broad and thin, with a strong median rib and a
peculiar semicircular hoop beneath the spot where they articulate
with one another. The three hoops of the three jaws work inside each
other in such a way as to cause the jaws to have a strong grip and to
be very difficult to dislocate from their mutual articulation.

The ophicephalous pedicellariae are in Echinus the most abundant

of all; and they alone extend on to the peristome, where a special
small variety of them is found.

A thorough investigation of the functions and reactions of the

pedicellariae has quite recently been made by von Uexküll.[472] He
showed, first of all, that there is a nervous centre in the stalk of each
pedicellaria (see below), which causes the organ to incline towards a
weak stimulus, but to bend away from a stronger stimulus. In the
head there is an independent nervous centre, which regulates the
opening and closing of the valves, and causes these to open on
slight stimulus and close when a stronger one is applied. The
amount of stimulus necessary to cause the pedicellariae to retreat
varies with the kind of pedicellariae, being least with the tridactyle
and most with the gemmiform, so that when a chemical stimulus,
such as a drop of dilute ammonia, is applied to the skin, the
tridactyle pedicellariae may be seen to flee from and the gemmiform
to approach the point of stimulation. In a living Sea-urchin, if the
attempt is made to seize the tridactyle pedicellariae they will evade
the forceps, but the ophiocephalous are easy to catch.
The tridactyle pedicellariae open with the very slightest mechanical
stimulus and close with rather greater mechanical stimuli or with
exceedingly slight chemical ones. Uexküll calls them "Snap-
pedicellariae," and their function is to seize and destroy the minute
swimming larvae of various sessile parasitic animals, which would
otherwise settle on the delicate exposed ectoderm of the Sea-urchin.

The gemmiform pedicellariae are brought into action when a more

serious danger threatens the Sea-urchin, such as an attack of a
Starfish. The corrosive chemical influence, which it can be proved
exudes not only from the stomach but even from the tube-feet of the
Starfish, causes the gemmiform pedicellariae to approach and open
widely. When the foe approaches so closely as to touch the sense-
organs (Fig. 225, B, s) situated on the inner side of the valves of
these pedicellariae, the blades close violently, wounding the
aggressor and causing its juice to exude, thus producing a renewed
and severe chemical stimulation which irritates the poison glands
and causes the poison to exude. The virulence of the poison may be
gauged from the fact that the bite of a single gemmiform pedicellaria
caused a frog's heart to stop beating.

Fig. 225.—The pedicellariae of Echinus acutus, drawn from a living specimen. A,

gemmiform pedicellaria, closed. B, gemmiform pedicellaria, open; g, poison
gland; s, sense-organ, × 3. C, tridactyle pedicellaria, × 6. D, ophicephalous
pedicellaria, × 9. E, trifoliate pedicellaria, × 12; a (in all figures), axial rod of
the stalk. (After Uexküll.)

Prouho[473] has described a combat between a Sea-urchin and a

Starfish. When the latter approached, the spines of the Sea-urchin
diverged widely (strong form of reaction to chemical stimulus),
exposing the gemmiform pedicellariae. These at once seized the
tube-feet of the enemy and the Starfish retreated, wrenching off the
heads of these pedicellariae; then the Starfish returned to the attack
and the same result followed, and this was repeated till all the
pedicellariae were wrenched off, when the Starfish enwrapped its
helpless victim with its stomach.

The minute trifoliate pedicellariae are brought into play by any

prolonged general irritation of the skin, such as bright light or a rain
of particles of grit or mud. They have the peculiarity that not all the
blades close at once, so that an object may be held by two blades
and smashed by the third. They may be seen in action if a shower of
powdered chalk is poured on the animal, when they seize the
particles and by breaking up any incipient lumps reduce the whole to
an impalpable powder, which the cilia covering the skin speedily
remove. In thus assisting in the removal of mechanical "dirt" they
earn the name which Uexküll has bestowed on them, of "cleaning
pedicellariae."

Fig. 226.—Views of a single blade of each kind of pedicellaria. A, blade of

gemmiform pedicellaria of Echinus elegans; g, groove for duct of poison
gland; B, blade of tridactyle pedicellaria of the same species; C, blade of
ophicephalous pedicellaria of the same species; r, ring for clamping this
blade to the other blades; D, blade of trifoliate pedicellaria of E. alexandri.
(After Mortensen.)

The ophicephalous pedicellariae, with their powerful bull-dog grip,

assist in holding small animals, such as Crustacea, till the tube-feet
can reach them and convey them to the mouth.

The number and variety of the pedicellariae, then, is an eloquent

testimony to the dangers to which the soft sensitive skins of the Sea-
urchin and other Echinodermata are exposed, and afford
confirmatory evidence in support of the view expressed above, that
the method adopted to defend the skin was one of the great
determining features which led to the division of the Asteroidea into
different races.

Fig. 227.—Dried shell of Echinus esculentus, showing the arrangement of the

plates of the corona. × 1. 1, The anus; 2, periproct, with irregular plates; 3,
the madreporite; 4, one of the other genital plates; 5, an ocular plate; 6, an
interambulacral plate; 7, an ambulacral plate; 8, pores for protrusion of the
tube-feet; 9, tubercles of the primary spines, i.e. primary tubercles.

The corona consists of five radial or "ambulacral" bands of plates

and five interradial, or as they are usually termed, "interambulacral"
bands of plates—ten in all. Each of the ten consists of two vertical
rows of plates throughout most of its extent, and each plate is
studded with large bosses, or "primary tubercles" for the primary
spines, smaller bosses called "secondary tubercles" for the
secondary spines, and finally, minute elevations called "miliary
tubercles" for the pedicellariae.
 Fig. 228.—The so-called calyx and the periproct of Echinus esculentus. × 4. 1,
Genital plates with genital pores; 2, ocular plates with pores for terminal
tentacles of the radial water-vascular canals; 3, madreporite; 4, periproct
with irregular plates; 5, anus. (After Chadwick.)

Even in the dried skeleton, however, the ambulacral plates can be

discriminated from the interambulacral by the presence of pores to
permit the passage of the tube-feet. These pores are arranged in
pairs, and each pair corresponds to a single tube-foot, since the
canal connecting the ampulla with the external portion of the tube-
foot is double in the Echinoidea. In Echinus esculentus there are
three pairs of such pores in each plate, in Strongylocentrotus
droëbachiensis four pairs. The ambulacral plate is really made up of
a series of "pore-plates," each carrying a single pair of pores, and
these become united in threes in Echinus and fours in
Strongylocentrotus, while in primitive forms like the Cidaridae they
remain separate. Each ambulacral and interambulacral area ends at
the edge of the periproct with a single plate. The plate terminating
the ambulacral band is pierced by a single pore for the exit of the
median tentacle, which, as in Asteroids, terminates the radial water-
vascular canal. Thus the aboral end of the radius in an Echinoid
corresponds to the tip of the arm in an Asteroid. The plate is termed
"ocular," because the terminal tentacle has a mass of pigmented
cells at its base; but no eye-cups can be seen, and there is no
evidence that this spot is specially sensitive to light. Species which
show special sensitiveness to light have often a large number of
what we may perhaps term secondary eyes. The plate terminating
the interambulacral series is termed the "genital plate," because it is
pierced by the duct of one of the five genital organs. One of the