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Oedipus Rex in the
Genomic Era
Human Behaviour, Law and Society
Yulia Kovas · Fatos Selita

Oedipus Rex in the Genomic Era
Yulia Kovas · Fatos Selita
Oedipus Rex
in the Genomic Era
Human Behaviour, Law and Society
Yulia Kovas Fatos Selita
School of Professional Studies, Barrister (England and Wales); Attorney
Science & Technology and Counselor at Law (NY State)
Goldsmiths University of London Goldsmiths University of London
London, UK London, UK
ISBN 978-1-349-96047-7 ISBN 978-1-349-96048-4 (eBook)

https://doi.org/10.1057/978-1-349-96048-4
© The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature
Limited 2021
The author(s) has/have asserted their right(s) to be identified as the author(s) of this work in
accordance with the Copyright, Designs and Patents Act 1988.
This work is subject to copyright. All rights are solely and exclusively licensed by the Publisher,
whether the whole or part of the material is concerned, specifically the rights of reprinting, reuse
of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way,
and transmission or information storage and retrieval, electronic adaptation, computer software, or
by similar or dissimilar methodology now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this
publication does not imply, even in the absence of a specific statement, that such names are exempt
from the relevant protective laws and regulations and therefore free for general use.
The publisher, the authors and the editors are safe to assume that the advice and information in this
book are believed to be true and accurate at the date of publication. Neither the publisher nor the
authors or the editors give a warranty, expressed or implied, with respect to the material contained
herein or for any errors or omissions that may have been made. The publisher remains neutral with
regard to jurisdictional claims in published maps and institutional affiliations.
Cover illustration: Daniel Hernanz Ramos/Getty Images
This Palgrave Macmillan imprint is published by the registered company Springer Nature Limited
The registered company address is: The Campus, 4 Crinan Street, London, N1 9XW, United
Kingdom
Acknowledgement
We dedicate this book to our parents.

This book is a product of years of wonderment, bits of sabbatical,
two lockdowns at the time of the pandemic and of infinite pleasure of
working together. An inexhaustible source of inspiration for this work
have been our students, who propel knowledge by their enthusiasm, their
questions and passion for knowledge.
We are grateful to Robert Chapman, Sasha Murray and Amanda Selita
for being the first readers and evaluators of our book. Your reassurance
and insightful comments have given us an impetus to finally complete
the work! A special thanks goes to Della Couling. It is an honour for us
to have received your edits on our manuscript. We are in awe of your eye
for detail, your wisdom, wit and energy.
v
Contents
1 A Journey into the Genomic Era: Sophocles Will Be

Our Guide… 1
Oedipus Rex and Behavioural Genetics 2
The Journey 4
Plot of the Three Theban Plays 7
Characters of the Theban Plays (and Their Equivalents
in the Genomic Era) 11
A Note on Citations and Translations 12
List of Translations Used in This Book 13
References 15
2 Genes, Environments and Life Trajectories 17
Familiality 20
Lineage 29
Chance, Genes and Crossroads 35
References 38
vii
viii Contents
3 Free Will in the Genomic Era 45

The Free Will Confusion, Illusion and Delusion 46
Free Will and Determinism 48
Free Will and Errors of Attribution 53
Free Will and Punishment 56
The Will of the Self 64
References 67
4 Prophesied Future and Redefined Past in the Genomic
Era 73
Destiny, Fatalism and Predetermination 74
Bringing Fate About 76
The New Pantheon of Gods in the Genomic Era 78
Prediction in the Digital World 79
Oracles and Prophecy in the Genomic Era 83
The Power of Expectation 88
The Dangers of Misinterpretation 90
Redefining the Past 95
Support in Dealing with ‘Prophecies’ 96
‘Quantum’ Prophecy 98
References 101
5 DNA: The Greatest Text of All 111
Length of Text and Depth of Meaning 112
Genes vs. Genome 118
Lost and Found in Translation 120
Alternative Splicing 122
Beyond Genes 126
Variation 128
Expression: Reading the Text 136
The Age of the Text 141
Particles, Atoms and Elements 141
Life Emerging from Non-Organic Universal Debris
and Star Dust 143
DNA—The Connector of All Species 145
Editing the Text 147
Storing New Meanings 149
Contents ix
Randomness, Feedback and the Absence of a Plan 154

Oedipus Rex in the Genomic Era 157
Epilogue 159
Chorus (Citizens of the Genomic Era) 160
Afterword 161
References 162
Glossary 173
Index 231
List of Figures
Fig. 1.1 Oedipus’ life journey. Oedipus’s life journey starts

in Thebes, from which he is taken as a baby
to Corinth where he grows up. In Delphi, the prophecy
from the Oracle ‘derails’ his life. After the eventful
journey he arrives in his birthplace—Thebes. In
the sequel to Oedipus Rex, Oedipus ends his days
in Colonus—the suburb of Athens (Source British
Library Flickr) 14
Fig. 5.1 DNA structure 114
xi
List of Boxes
Box 5.1 DNA Structure 114

Box 5.2 Genome size 116
Box 5.3 Transcription 121
Box 5.4 Mitosis 132
Box 5.5 Methylation 138
Box 5.6 Data storage 150
xiii
1
A Journey into the Genomic Era: Sophocles
Will Be Our Guide…
Why are we here? What are our worst fears? Is our will free? Why are
we our own worst enemies? Why are our judgements so flawed? Why
do we seek to know the future? What is fate and can we run away from
it? What is chance? This book explores the answers to these questions
with the help of two ancient texts that provide tremendous insights into
human behaviour.
The first text is Oedipus Rex (Oedipus Tyrannus) by Sophocles, written
in the fifth century BCE (Before Common Era). As many plays in
Sophocles’ time, Oedipus Rex was written for a competition and was
selected to be staged during a festival in front of possibly as many as
15,000 spectators. We can only imagine the impact Oedipus Rex had
on the spectators who gathered to watch its initial performance. The
play is so powerful that it was immediately recognised as a masterpiece
and its value did not depreciate with time. This international bestseller,
with millions of copies sold over the centuries, is perhaps one of the best
texts examining the complexities and the origins of human behaviour
and psyche. It masterfully explores themes of free will , fate and chance;
prediction, misinterpretation and the burden that comes with knowl-
edge of the future; self-fulfilling and self-defeating prophecies; the forces
© The Author(s), under exclusive license to Springer Nature 1
Limited 2021
Y. Kovas and F. Selita, Oedipus Rex in the Genomic Era,
https://doi.org/10.1057/978-1-349-96048-4_1
2 Y. Kovas and F. Selita
that contribute to similarities and differences among people; roots and

lineage; judgement of self and others; and much more.
The second text is Human DNA, with its origins around 4 billion years
ago and continuously revised by chance and evolution. This text, with
almost 8 billion slightly different copies in circulation (the number of
people living today), governs every cell of our body, contributes to every
action we take, every thought and every experience we have. DNA holds
stories of our past, present and future. Although DNA (deoxyribonucleic
acid ) is not literally a text, the analogy of text is most suitable. This is
because its function (‘meaning’) emerges from the specific sequences of
basic elements (DNA building blocks are like letters of the alphabet); and
because it is written (built) and is read (decoded) in a particular direction.
As with other texts, despite the fixed order of ‘letters’, different mean-
ings may emerge from the same ‘words’ (genes). The different meanings
depend on processes ‘above’ the text (epigenetics), such as ‘punctua-
tion’, ‘intonation’ and other elements (methylation and other epigenetic
processes in our cells)1 . Exploring DNA provides new insights into the
processes underlying human behaviour and mind that we discuss with
the help of Sophocles in this book. Knowledge of DNA is also becoming
more and more relevant to all people living in the Genomic Era.
Oedipus Rex and Behavioural Genetics

Today, Oedipus Rex continues to be widely read and included in litera-
ture curricula. Unfortunately, many people who study the play remember
not much more than ‘Oedipus killed his father and married his mother’.
This may be because it is indeed a striking element of the plot, but also
because Sigmund Freud used it as a metaphor for the concept of the
‘Oedipus complex’. Indeed, psychological processes that underscore the
Oedipus complex are evoked in the play. Jocasta (Oedipus’ wife/mother)
tells Oedipus that many men experience complex feelings towards the
mother.
…For that maternal wedding,

Have you no fear; for many men ere now
1 A Journey into the Genomic Era … 3
Have dreamed as much; but he who by such dreams

Sets nothing, has the easiest life of it2 .
Although this is an interesting and memorable theme, the play is about

much more. Through his characters, Sophocles has expressed many issues
and worries that we continue to have today about our nature, nurture
and will. The characters of the play are timeless, they are real. In the
words of George Young , the famous translator of Sophocles, ‘the good
are faulty, the bad are not without excuse’, ‘their actions are accounted
by adequate motives’2 (p. xi). The rulers, although smart and noble, are
consistently blind, ignoring the advice of those who ‘see’. The oracles
are confused, the gods are not all-powerful. The characters’ actions are
driven by a multitude of forces. They are empathised with, but they are
also judged and punished—by others and by themselves.
Oedipus Rex was not Sophocles’ only masterpiece. In his life (around
496–405 BCE), he is thought to have written over 100 plays. Most
did not survive to today and not much has been documented about his
life. However, we do know that Sophocles was revered for his plays. For
example, it is believed that Athenians appointed Sophocles a general of
their army, attesting to their admiration for his ‘Antigone’ (a play about
the fate of Oedipus’ daughter)2 (p. xi).
Among the most well-known plays of all times is Sophocles’ trilogy,
collectively known as the Theban plays: Oedipus Rex , Oedipus at Colonus
and Antigone—in which we find out about the fate of Oedipus and his
family. Throughout this book, we will provide quotes from Oedipus Rex ,
as well as a few quotes from Oedipus at Colonus and Antigone.
In modern times, many of the themes covered in Sophocles’ plays
have been formally explored by the branch of science called behavioural
genetics 3 . A major aim of this science is to determine genetic and environ-
mental conditions that make every person unique. It uncovers the origins
of differences among people through studying relatives, including twins,
adoptive and non-adoptive parents and siblings. To date, behavioural
genetics has generated many insights into complex processes that govern
behaviour and thought4,5 . In addition, in the last 60 years, genetics has
achieved unprecedented advances in decoding and understanding the
text of life in our cells—DNA. This text is of such immense complexity
that it will take a long time to fully comprehend all its intricacies.
However, the progress we have made already is staggering—attesting to
the magnificent power of the human brain.
Oedipus Rex in the Genomic Era examines the actions and thoughts
of Oedipus Rex’s characters in the context of the knowledge available
in today’s Genomic Era. It describes new twists to our quest to know
ourselves: genes and environments replacing ancient gods; genetic coun-
sellors replacing oracles. It considers how we defeat and fulfil prophecies
based on genetic (mis)information; and the ways gene editing technolo-
gies may enable us to ‘play’ evolution. The book also raises concerns and
dilemmas brought by the Genomic Era: How will the ability to read and
understand the text of life co-exist with our burning desires to know
and alter the future? How will we react to predictions of the future and
what support will we have to deal with our ever-accumulating knowl-
edge? How can we harness the new power brought by this knowledge so
that we can use it for individual and societal benefit?
The Journey
In Oedipus Rex in the Genomic Era we set out on a journey into the
Genomic Era. This journey is into the unknown where our wildest
wishes and worst fears may come true. We explore the horrors, the traps
and joys that may lie ahead. In this sense, our journey is like that of
Dante in the DivineComedy 6 , who visits hell, purgatory and paradise.
This journey helps him ‘to find his way’ after being lost. In the Divine
Comedy a Roman poet, Virgil, guides Dante on his journey. In this book
we take Sophocles as our guide, to whom we dedicate the words that
Dante addressed to Virgil: ‘O, glory and light to other poets, may that
long study and the great love, that made me scan your work, be worth
something now6 ’.
In this chapter we introduce the characters and the plot of the Theban
plays, that we will draw upon throughout the book.
Chapter 2 explores insights into human behaviour, generated by
behavioural genetics, illustrating genetic concepts with Sophocles’ text.
We look at adoption; blood relations and lineage; and similarities and
differences of family members. We explore gene-environment interplay

and how chance interacts with genes to shape our life trajectories. We
show how people have always known, worried and were in awe of genetic
effects; and explore the confusions that we have about the origins of
differences among us.
Chapter 3 delves into control, free will, attributional errors, judgement
and punishment—major themes of Oedipus Rex . Through the actions of
Oedipus and other characters we demonstrate confusions and delusions
that we hold about our will and show that free will is a powerful illu-
sion. We discuss threats that this illusion poses to sound judgement in
everyday life and in criminal justice.
Our courts today often resemble ‘spectacles’—as in Sophocles’ time,
when Athenians attended court hearings in much the same way that
they attended plays like Oedipus Rex . These real-life dramas unravelled
in front of spectators’ eyes. Plaintiffs and defendants were judged by
the ‘audience’, almost as if it were theatre. What was said and how it
was said and what emotional effect it had on the audience were major
elements affecting the courts’ decisions. The subjectivity, the persuasion,
the playing with the jurors’ emotions were all vital elements of the ‘show’.
These procedures have been criticised for as long as they have
existed, for example in the timeless comedy, The Wasps (422 BCE), by
Aristophanes7 , a contemporary of Sophocles. In the play, a trial of a
dog is staged. During the trial, all methods are used to soften the jury.
For example, at first the accused is described as a worthy member of
society: he is the ‘virtuoso’, the genius. However, when this fails, the
dog is described as disadvantaged by circumstances: he is uneducated
and had not had the opportunity to learn to play a musical instru-
ment. When that also fails, the dog’s puppies are brought in and asked
to cry and howl. Witnesses before the court are pots and pans, but,
because they cannot speak, they are represented by another character ‘on
behalf of the accused’. Finally, the characters simply cheat to obtain the
desired outcome. One character in The Wasps is a trialophile—addicted
to serving as a voluntary juror in a law court. As benefits of juror’s service
he lists attentions of powerful men who appeal to him for a favourable
verdict; and the freedom to interpret the law as he pleases. Our justice
system still operates with many of the same flaws; and continues with
numerous biases; erroneous assumptions about human behaviour; and

corruption.
In Oedipus Rex Sophocles uses an ingenious trick when the same crime
is unwittingly judged differently by the same judge (Oedipus), as his
perspective changes. We examine these judgements to demonstrate the
inherent flaws in the justice process. We suggest that in the Genomic
Era, we can improve our judgements to avoid tragedies experienced by
characters of Sophocles’ immortal plays.
Chapter 4 explores our desire to know the future while paradoxically
also fearing this knowledge. We follow Sophocles in his exploration of
fate and how it comes or does not come about through the processes of
self-fulfilling and self-defeating prophecies. The Genomic Era has brought
a new ‘pantheon of gods’, where polytheistic systems of ancient beliefs
are replaced with polygenic 8 and polyenvironmental systems that come
together to drive our behaviour. We now also have new ‘oracles’, such as
commercial genotyping companies that ‘prophesy the future’ based on
genetic risks and propensities. Genetic prediction brings new dilemmas,
creates new expectation effects and may even redefine our past. As
genomic information becomes ubiquitous, we need proper support and
advice on how to interpret this information and how to act on it.
Chapter 5 explores the knowledge that brought about the Genomic
Era—the knowledge about DNA—the unique, unparalleled master-
piece that we carry in our cells. The chapter describes the structure
and function of DNA; explores its age; and how it is linked to the
forming of the universe, exploding stars, meteorites and everything and
everyone on Earth. We explore how the meaning of DNA unravels over
time, interacting with environments—in similar ways as the meaning
of any text unravels through the mind of its reader. The mind of
today’s reader contains new knowledge about humans—gathered by a
host of sciences: biology, genetics, neuroscience, sociology, anthropology,
psychology, physics and mathematics. This knowledge leads to new
discoveries, allowing us to extract new meanings from the text of life and
to forecast a future that we cannot yet comprehend. The knowledge also
allows us to extract new meanings from masterpieces, such as Oedipus
Rex , created by giants on whose shoulders we stand.
In the Epilogue to this book, the Chorus of Citizens of the Genomic

Era pledges to abandon confusions and to use the marvellous tools
provided by knowledge for the benefit of all. Finally, the Afterword
introduces a new metaphor to the DNA—that of a galactic system.
We provide in-text references for some sources and mark in Italics
concepts, theories and scientific findings that are covered in more detail
in the Glossary. The Glossary also contains references to further reading.
The Glossary terms are also listed alphabetically in the Index.
Before we set out on our journey, we provide a brief summary of the
Theban plays; followed by the list of characters from the Theban plays
that will ‘come onto the stage’ in Oedipus Rex in the Genomic Era.
Plot of the Three Theban Plays

Laius and Jocasta are king and queen of Thebes. Laius receives a prophecy
from an oracle that, should Laius have a son with Jocasta, the son would
kill him. Allegedly when drunk (according to one version of the myth not
covered in the plays), he gets Jocasta pregnant. When the baby is born,
Laius casts him out, with ankles pinned, according to Jocasta ‘by hands of
others’. In reality (as described by a shepherd at the end of Oedipus Rex ),
Jocasta herself passes the baby to him to be killed. The shepherd spares
the baby’s life and passes him on to another shepherd for adoption. This
second shepherd presents the baby to the king and queen of Corinth—
Polybus and Merope, who are childless. They adopt the baby, now called
Oedipus (swollen foot) and raise him and love him as their own son. He
has a great life, held in high regard by all. This continues until one day
Oedipus hears from a drunken man that he is not a natural child of his
parents but is an adoptee. Oedipus is distraught by this and confronts
his parents, who are angry at the drunk and assure Oedipus that he is
their own son. Oedipus is comforted by this but cannot get the thought
out of his mind. In fact, the rumour that he was adopted spreads and he
hears this from other people too. Without the knowledge of his parents,
he goes to see the Oracle of Delphi to seek the truth about his birth. The
oracle does not clarify the matter of his birth, but instead prophesies that
Oedipus would kill his father, marry his mother and have children from
this incestuous relationship. Horrified, Oedipus flees Corinth to escape

this terrible fate. He is set to go as far as possible from what he thinks
is his homeland. The journey, however, ultimately takes him to his real
homeland—Thebes. On the way, he encounters a party of travellers, who
demand that he gives way to their carriage, pushing him out of the way.
Enraged, he strikes the man who pushed him. In turn, a man within
the carriage strikes him on the head. Enraged even further, Oedipus kills
4 of the 5 men, with the fifth man escaping. As it turns out later, the
man in the carriage who hit him was indeed his father, Laius, travelling
on state business; and the escaped man was the shepherd, who had once
saved Oedipus’ life by giving him for adoption rather than killing him as
ordered by the king and queen.
Not knowing any of this and unremorseful, Oedipus continues his
journey and eventually arrives in Thebes, after some time—possibly
months. When he arrives in Thebes, the city is in great misery: their
king (Laius) is believed to have been killed by highway robbers some
time ago; and the city is terrorised by a Sphinx, who guards the entrance
to the city and demands that someone solves her riddle, eating anyone
who fails at the task. Oedipus, being wise, solves the riddle and the evil
Sphinx kills herself. The city rejoices and offers Oedipus the hand of the
queen and to become Thebes’ ruler. He accepts and rules Thebes, jointly
with his wife Jocasta and her brother—Creon.
Oedipus and Jocasta have 4 children together, two boys and two girls.
The sons, Polynices and Eteocles, never appear in Oedipus Rex , but
feature in other plays—Oedipus at Colonus and Antigone. The daughters,
Antigone and Ismene, are still little children, when the city of Thebes is
stricken by a new trouble—a terrible plague, which causes animals and
women to be barren, kills people and crops. Oedipus sends his brother-
in-law, Creon, to see an oracle and find out how to rid the city of the
plague. This is where the narrative of Oedipus Rex actually begins. The
message Creon brings from the Oracle is that the city will be saved once
the murderer of Laius, the previous king, is identified and either killed
or exiled. The murder of the king had not been properly investigated at
the time, as it had happened in another country and Thebes had been
preoccupied with troubles caused by the evil Sphinx.
Unwittingly, Oedipus swears to do anything to find the murderer. Not

only for the sake of the city, but for his own sake as he believes that Laius’
murderer could kill him too. Oedipus, on Creon’s advice, calls for Tire-
sias—the blind wise man, a prophet—and demands that he tells him
how to find the murderer. Tiresias refuses to give any information and
advises Oedipus to drop the investigation. Enraged, Oedipus threatens
Tiresias, who gives in and reveals that the murderer he is looking for
is Oedipus himself. But Oedipus does not understand this revelation;
enraged even further he accuses Tiresias and Creon of plotting together
to overthrow him. Jocasta, who does not trust oracles as she believes that
their prophecy regarding Laius’ death was wrong and led to the murder
of her baby, tells Oedipus not to trust oracles. She tells Oedipus that
Laius was meant to die by his son’s hands, but instead he was killed by
robbers at a cross-road. She also describes Laius’ appearance, which not
only matches that of the man Oedipus murdered, but also in Jocasta’s
words is similar to Oedipus’ appearance. Jocasta’s story, instead of the
planned comforting effect, throws Oedipus into further anxiety and
pushes him to continue the investigation. He sends for the only witness
of the King’s murder—the now old shepherd. While they are waiting
for the shepherd to arrive, a messenger from Corinth appears, looking
for Oedipus, to announce to him that he is invited to rule Corinth, as
his father, king Polybus, has died. Oedipus is somewhat relieved that his
father died of natural causes rather than by his own hands, unless (as
Oedipus jokes) he died from missing his son. However, Oedipus is still
apprehensive, as his mother (Merope) is still alive and he continues to
worry about the prophesied incest. The messenger, who turns out to be
the shepherd who had passed Oedipus to Polybus and Merope, tells him
that he has information that will relieve Oedipus from his apprehension.
He reveals that Oedipus was adopted by his parents, but he does not
know where he actually came from. Only the other shepherd knows this.
Jocasta, now fully aware of who Oedipus is, begs Oedipus to give up
the investigation. Oedipus continues to be utterly confused and accuses
Jocasta of worrying about his lineage, suggesting that she, being a proud
woman, is afraid to find out that she may be married to a low class, or
worse—a slave. The shepherd arrives and it is finally revealed beyond any
doubt that Oedipus is indeed the murderer of his father, a husband to
his mother and a father to his sisters and brothers. Jocasta kills herself,
Oedipus blinds himself and asks Creon to exile him and to take care of
his daughters.
The story continues in Oedipus at Colonus—Sophocles’ ‘sequel’ to
Oedipus Rex . We learn that following the tragic ending of Oedipus Rex ,
Oedipus was not allowed to leave Thebes and later was banished from
the land by Creon and his own sons. We see Oedipus, accompanied by
his daughters, Antigone and Ismene, arriving at Colonus (the suburb of
Athens). He recognises the place as the place of his approaching death,
as prophesied by Apollo’s oracle when his unfortunate fate was foretold.
He asks the king of Athens, Theseus, to allow him to stay and to protect
him, in return for a future benefit. The oracle had foretold that the land
where Oedipus would be buried would be protected by the gods. Theseus
grants Oedipus and his daughters the right to stay and protection. Unex-
pectedly, Creon arrives at Colonus and asks Oedipus to return with
him to Thebes. He is determined that Oedipus should be buried near
Thebes, so that foreign lands do not benefit from his burial. Oedipus’s
refusal propels Creon to seize Antigone and Ismene as hostages. Theseus
sends troops and retrieves the girls, keeping his promise to protect them.
Oedipus is asked to receive another visitor, who turns out to be his son,
Polynices. Oedipus reluctantly sees him. His sons, who originally agreed
to rule Thebes in turn together with Creon, have now quarrelled. Polyn-
ices, the older son, asks Oedipus to help him to retake power in Thebes
on the basis that the younger son, Eteocles, thirsty for power himself,
had overthrown Polynices and exiled him from the land. Polynices, while
abroad, amassed an army and is planning to return to Thebes with the
army and take power again. However, he needs Oedipus’ backing, as
the result of the battle depends on which of the sons Oedipus supports.
Oedipus, unforgiving of both sons for not taking care of him, curses both
Polynices and Eteocles, foretelling that they would die by each-others’
hands. Shortly after, Oedipus dies, allowing only Theseus to know the
place of his burial. Antigone and Ismene are stricken by grief. Antigone
asks Theseus to let them go back to Thebes, as she hopes that they can
avert the terrible fate of her brothers.
The story continues further in Antigone. Creon is now the ruler of
Thebes, following the death of Oedipus’ sons, Eteocles and Polynices,
who killed each other in battle (as foretold by Oedipus). Creon orders
that Eteocles be buried as a hero and for Polynices to be left unburied
as a traitor. Antigone, now engaged to Creon’s son, Haemon, disregards
Creon’s order and buries Polynices. Ismene refuses to disobey Creon’s
order and does not participate in the forbidden burial. Tragedy ensues:
Antigone is punished (buried alive) by Creon, despite pleas from his son.
Tiresias reappears and with the help of the Senators manages to convince
Creon to free Antigone before it is too late; but by the time they make
it to the tomb, she is dead and so is Haemon who has killed himself.
Characters of the Theban Plays (and Their

Equivalents in the Genomic Era)
Chorus—senators of Thebes; citizens of Colonus (and citizens of the
Genomic Era).
Gods (polytheism)—the dwellers of Olympus (and multiple inter-
acting genetic and environmental forces).
Oracles—prophets of Delphi (and commercial genotyping compa-
nies).
Tiresias—the wise man; possesses information and has the power to
reveal or not to reveal it to people. Objective, mostly free of external
influences, ‘…who has in him the tongue that cannot lie’2 (p. 137).
Blind, but the only one who truly sees. Believes that knowledge about
the future in the absence of means to prevent the future is harmful.
Oedipus—biological son of king Laius and queen Jocasta; adopted
very soon after birth by king Polybus and queen Merope; son AND
husband of Jocasta.
Jocasta—wife of the late king Laius; mother AND wife of Oedipus.
Polybus—king of Corinth; adoptive father of Oedipus.
Merope—queen of Corinth; adoptive mother of Oedipus.
Creon—brother of queen Jocasta; uncle AND brother-in-law to
Oedipus; uncle to Antigone, Ismene, as well as Polynices and Eteocles,
with whom he briefly shares the throne after Jocasta’s death.
Eteocles—son of Jocasta and Oedipus; brother (possibly twin) of
Polynices; sides with his uncle Creon against his brother.
Polynices—son of Jocasta and Oedipus; brother (possibly twin) of

Eteocles; attempts to take power from his uncle Creon; posthumously,
proclaimed traitor and ordered by Creon not to be buried.
Antigone—the rebellious daughter of Jocasta and Oedipus; buries her
brother, Polynices, against her uncle’s orders; buried alive in a cave as
punishment by Creon.
Ismene—the timid daughter of Jocasta and Oedipus; tries to talk
Antigone out of her rebellious actions against Creon.
Shepherd—the man who received baby Oedipus from Jocasta to ‘make
away with’. The same man also accompanied Laius on the ominous
day of his death.
Messenger (the second shepherd)—the man who brings to Oedipus
the news that his father Polybus died. The same man earlier received
baby Oedipus from the Shepherd and presented Oedipus to Polybus
and Merope.
A Note on Citations and Translations

Throughout the book, we cite extracts from the Theban plays. We mostly
use the 1888 translation by Sir George Young, Edited by Ernest Rhys
in 1906 Everyman’s Library. In several places, we opt for other transla-
tions from different periods. Different translations can lead to ‘different
expression’—a genetic concept that we discuss in Chapter 5. Unfortu-
nately, neither of us reads Ancient Greek—and therefore we are entirely
in the hands of the translators. We have read the plays in seven different
translations and three different languages and have not found the ‘best’
translation. Some passages are particularly powerful in one translation;
other passages—in other translations. In places, interpretations of char-
acters’ actions and motives can be affected by translations. For example,
were Oedipus’ feet pierced by a pin or bound together? Was an episode
on the crossroads murder or manslaughter? We can only recommend that
the readers try different translations to appreciate how their effects differ.
We demonstrate the power of translation in this extract, where citizens
of Thebes are eager to hear the prophecy:
Sweet-voiced daughter of Zeus from O Prophecy of Jove, whose words

thy gold-paved Pythian shrine are sweet,
Wafted to Thebes divine, With what doom art thou sent
What dost thou bring me? To glorious Thebes, from Pytho’s
My soul is racked and shivers with gilded seat?
fear I am distraught with fearful
(Healer of Delos, hear!) wonderment,
Hast thou some pain unknown I thrill with terror and wait
before, reverently…
Or with the circling years renewest a Offspring of golden Hope,
penance of yore? immortal Oracle,
Offspring of golden Hope, thou Tell me, O tell!
voice immortal, O tell me
Sophocles. Oedipus the King; The Dramas of Sophocles in
Oedipus at Colonus; Antigone. English Verse. By Sir George
With an English translation by Young 1888. Everyman’s Library
Storr, Francis, BA. Edited by T. E. 1906. Edited by Ernest Rhys.
Page MA; and W. H. D. Rouse London: J.M. Dent & Sons Ltd.
Litt.D. 1912. London, William New York: E. P. Dutton & Co
Heinemann. New York: The
Macmillan Co.
List of Translations Used in This Book

1. Sophocles. Oedipus the King; Oedipus at Colonus; Antigone. With
an English translation by Francis Storr, BA. Edited by T. E. Page MA;
and W. H. D. Rouse Litt.D. 1912. London, William Heinemann.
New York: The Macmillan Co.
2. Sophocles. The Three Theban Plays: Antigone; Oedipus the King;
Oedipus at Colonus. Translated by Robert Fagles 1983. Penguin
Books.
3. Sophocles. The Theban Plays. Translated by E. F. Watling. Penguin
Books, 1947.
4. The Dramas of Sophocles in English Verse. By Sir George Young
1888. Everyman’s Library 1906. Edited by Ernest Rhys. London: J.
M. Dent & Sons Ltd. New York: E. P. Dutton & Co.
14
Y. Kovas and F. Selita
Fig. 1.1 Oedipus’ life journey. Oedipus’s life journey starts in Thebes, from which he is taken as a baby to Corinth
where he grows up. In Delphi, the prophecy from the Oracle ‘derails’ his life. After the eventful journey he arrives in
his birthplace—Thebes. In the sequel to Oedipus Rex, Oedipus ends his days in Colonus—the suburb of Athens (Source
British Library Flickr)
References
1. Gibney, E. R. & Nolan, C. M. Epigenetics and gene expression. Heredity
105, 4–13 (2010).
2. Sophocles. The dramas of Sophocles rendered in English verse, dramatic & lyric
(Translation by Sir George Young). (London, J.M. Dent, 1906), p. 157.
3. Knopik, V. S., Neiderhiser, J. M., DeFries, J. C. & Plomin, R. Behavioral
genetics (Worth Publishers, Macmillan Learning, 2017).
4. Plomin, R., DeFries, J. C., Knopik, V. S. & Neiderhiser, J. M. Top 10 repli-
cated findings from behavioral genetics. Perspectives on Psychological Science
11, 3–23 (2016).
5. Behavioural Genetics for Education. (Palgrave Macmillan UK, 2016). https://
doi.org/10.1057/9781137437327.
6. Dante’s Divine Comedy, Complete. https://www.gutenberg.org/files/8800/
8800-h/8800-h.htm.
7. Aristophanes (c.446–c.386 BC)—Wasps: Translated by George Theodoridis.
https://www.poetryintranslation.com/PITBR/Greek/Wasps.php.
8. Lvovs, D., Favorova, O. O. & Favorov, A. V. A polygenic approach to the
study of polygenic diseases. Acta Naturae 4, 59–71 (2012).
2
Genes, Environments and Life Trajectories
Ah mayst thou ne’er discover who thou art!1
I must pursue this trail to the end,

Till I have unravelled the mystery of my birth2 .
Sophocles, his characters and audiences would have called to divine

sources to understand the human condition. In actuality, all of them were
subject to gene-environment processes that we are only now starting to
understand3,4 . These processes have been intuitively described by people
for centuries and have been scientifically explored by behavioural genetics
since the nineteenth century. Behavioural genetics flourished in the twen-
tieth century, when solid scientific methods were developed to study
resemblance of family members with different degrees of genetic relat-
edness. Large-scale twin and adoption studies have shed much light on
the sources of individual differences5,6 . Unprecedented technological and
computational advances of the twenty-first century have led to the devel-
opment of new branches of behavioural genetics—molecular genetics 7 and
genomics. A huge body of scientific investigations have concluded that
© The Author(s), under exclusive license to Springer Nature 17
Limited 2021
Y. Kovas and F. Selita, Oedipus Rex in the Genomic Era,
https://doi.org/10.1057/978-1-349-96048-4_2
all human traits are on average 50% heritable and 50% environmental,
with some traits more heritable than others8 . This means that individual
differences in all aspects of human functioning are to some extent related
to differences in our DNA code and life circumstances.
Through Oedipus and other characters of the immortal Theban plays
we can observe gene-environment processes that contribute to differences
among people in their perceptions, thoughts, feelings and actions. In this
chapter we will follow Oedipus’ life, as it illustrates multiple behavioural
genetics concepts, including familiality, gene-environment interplay/co-
action 9,10 (including gene-environment correlations and gene-environment
interactions). Together with Sophocles, we will seek an answer to why
Oedipus behaves in such a destructive way, in the end literally turning
into his own worst enemy—initiating and conducting a criminal inves-
tigation against himself.
As an only child of a king, Oedipus’ life was meant to be one of priv-
ilege. For example, his prenatal development would have been protected
from many negative effects. Indeed, he did not experience maternal
malnutrition, lack of sanitation and other poverty-related circumstances.
However, Oedipus’ life begins with a series of traumatic events. The
prophecy, received by his father Laius—that he would die by his son’s
hands—triggers a host of cascading effects on Oedipus’ life, starting from
pre-birth. Jocasta’s pregnancy was not a happy one—knowing that if the
baby in her womb is a boy—he is destined either to be killed at birth
or to kill his own father. The stress experienced by her throughout preg-
nancy could have set in motion multiple processes, including an increase
in stress hormones, such as cortisol and norepinephrine. This, in turn,
could have led to inflammation which is now known to negatively affect
the foetal environment. Maternal psychological stress and anxiety can be
considered a teratogen—an agent that may have negative effects on the
baby’s brain development and the immune system, as well as increasing
the risk of developing somatic (physical) or psychiatric disorders later in
life.
Stress during pregnancy may also alter genetic expression in the devel-
oping baby, which may have long-lasting effects. Today we know that
many processes are involved in the action of our genes. For example,
a chemical called methyl can make our genes ‘silent’, preventing the
genes from making proteins (the process that happens ‘epi = above’ the
genes). Stress in pregnancy may lead to abnormal regulation of methy-
lation. These epigenome alterations due to in-utero stress exposure can
be long-term and have long-lasting effects—linking maternal stress with
problems in child functioning and into adult life11–13 .
The next traumatic event in Oedipus’ life happens very soon after
his birth: his ankles are pinned (that is how he later gets the name
Oedipus—swollen foot) and he is cast out by his father, Laius—destined
to certain death. At one point in the play, Jocasta describes Laius’ actions:
While from our son’s birth not three days went by

Before, with ankles pinned, he casts him out,
By hands of others, on a pathless moor14 .
We later find out from the shepherd that it was Jocasta herself who
handed baby Oedipus to him ‘to make away with’15 .
Oedipus then catches a lucky break. The man charged with executing
his terrible fate pities the baby and passes him to another shepherd from
Corinth, who, in turn, presents the baby to the childless king Polybus
and queen Merope. In behavioural genetics, this is an excellent example
not only of adoption, but also of so-called selective placement , when chil-
dren are adopted by families who in some ways resemble their biological
families16,17 . It is not clear in Oedipus’ case whether the second shep-
herd knew of the baby’s royal origin. In any case, being adopted by the
royal couple was a stroke of luck for Oedipus.
Adoption was not an uncommon event in ancient times, usually prac-
tised to ensure succession. Surrounded by love, growing up in the highest
socio-economic conditions, Oedipus’ childhood should have provided
him with much buffering against the effects of his unfortunate early life
experience. Indeed, he has done well and is held in great respect—until
an incident changes the course of his life yet again.
My sire was Polybus of Corinth and

My mother Merope, a Dorian;
And I was held the foremost citizen,
Till a strange thing befell me, strange indeed,
Yet scarce deserving all the heat it stirred18 .
Familiality
Oedipus Rex masterfully explores the theme of family resemblance and
the concept of familiality—when a particular trait/behaviour occurs with
greater frequency in a given family than in the general population. Does
Oedipus grow up resembling his adoptive parents who raised him from
the start? Or, does he resemble his biological parents—Laius and Jocasta,
with whom he has never lived? Are there some traits, some life patterns,
that members of Oedipus’ family have in common?
From large-scale adoption studies we know that children usually
resemble their genetic parents and siblings (with whom they do not live)
more than their adoptive parents and siblings, with whom they share
home/family environments. This suggests that genetic and early (possibly
prenatal) effects play a very important role in familyresemblance 19 .
Dissimilarities between family members are also explained by both
genetic and environmental factors. First degree relatives (parents and
children; full siblings) share only about 50% of the variable DNA
(sections of the DNA that differ across people), leaving much scope
for the DNA to create observable differences among them. In addition,
life events and circumstances that are unique to each individual (rather
than family wide) contribute to differences among family members. The
differences are further increased by the interplay between unique genetic
and environmental effects.
Throughout the Theban plays, we learn that Oedipus resembles his
biological father—Laius. Jocasta unknowingly tells Oedipus that he
looks very much like his father. The similarities extend to other traits
and behaviours, including both men committing serious crimes. We
don’t know much about Laius from the play but, as we will describe
later, Sophocles’ contemporaries would have known more about his past
crimes from mythology. Laius’ behaviour at the crossroads also suggests
that he suffered from temper and hubris—why else would he get so angry
with a stranger on the road?
Like Laius, Oedipus is uncompromising, irritable, impulsive and hot-
tempered. In Oedipus at Colonus, Creon describes Oedipus’ temper as
‘frenzy’, Oedipus’ perpetual bane.
Oedipus is also extremely arrogant and prone to hubris—often

boasting. For example, he tells the citizens, who advise him to consult
with Creon, that he had already sent for Creon (meaning others cannot
give him any useful advice).
Here too my zeal has nothing lagged, for twice

At Creon’s instance have I sent to fetch him,
And long I marvel why he is not here20 .
In another example of hubris, Oedipus implies that Tiresias is a useless

prophet because he could not stop the Sphinx, whereas Oedipus could
solve the riddle without any help from gods.
Till I came,
The unlettered Oedipus and ended her,
By slay of wit, untaught of augury ...21 .
The epitome of Oedipus’ hubris, his hamartia, is that throughout the

play he acts as if he has full knowledge when he has every reason to
suspect that he doesn’t. Killing a man old enough to be his father and
marrying a woman old enough to be his mother is a striking example of
his arrogance, his belief that he is in control of his fate.
Oedipus’ other flaw is that he often suspects others of ill intentions.
For example, he accuses Tiresias and Creon of plotting against him—
which was just a product of his suspicious mind:
Oedipus. Are these inventions Creon’s work, or yours?

Tiresias. Your bane is no-ways Creon, but your own self
...
Oedipus. I whom you now seek to cast out, in hope
To stand upon the steps of Creon’s throne22 .
Oedipus is also obsessive—for example, even having lived in a foreign

land for years, he continues to fear the prophecy, with this anxiety
affecting his whole life. He tells the messenger from Corinth, who came
to tell him that his father Polybus is dead, that he is still afraid to go
home because he fears he may still have incest with his mother.
Oedipus. Of course I fear lest Phoebus turn out true.

Messenger. Lest through your parents you incur foul stain?
Oedipus. Yes, father, yes; that is what always scares me23 .
In fact, both Oedipus and Laius are obsessive and stubborn, for example,
in seeking to know the future and trying to outsmart fate. Oedipus,
based on some rumours from a drunken man (that Oedipus was not
the son of his parents), decides to seek answers from an Oracle as to who
his biological parents are. In fact, he ignores assurances from his loving
parents that he is their own son and that they love him very much. He
decides to alter his prophesied future, abandoning his parents and fleeing
from his land and fate. This fleeing from fate repeats his father’s decision
to kill his own son to escape fate.
These examples of familiality—when physical and psychological traits
run in families—could in principle be explained by genetic factors as
well as how one is raised (shared environment ). However, in Oedipus’
case, his resemblance to his biological father is most likely explained by
genetic relatedness as he was not raised by Laius.
It is also possible that his similarity to his biological father was
increased by selective placement . In adoption studies, selective placement
complicates the interpretation of outcomes. For example, some of the
similarities in psychological characteristics that Oedipus shares with his
biological father could be due to similar environments in which they
developed as children: both sons of ruling kings, they would likely
have been encouraged to act in a dominant and arrogant manner. So,
being raised as a son of a king is a societal condition that may make
pride and hubris more likely. This demonstrates one way in which our
genes can correlate with our environments, a concept called passivegene-
environment correlation 24–26 . Laius might have passed on to Oedipus
some genes that contributed to both men’s impulsiveness and pride.
These characteristics might have been further cultivated by the regal
upbringing.
Another mechanism explaining why the adult Oedipus resembles his
biological father is known as an active gene-environment correlation. As we
go through life, we seek out and select environments that are consistent
with our genetic propensities. We can see how internal, largely uncon-
scious processes drive Oedipus to seek oracular advice; to get into a
fight at the crossroads, instead of opening the way; to threaten Tire-
sias when he refuses to prophesy; to ignore his wife when she begs
him not to pursue the search for the killer. These examples of Oedipus’
behaviour demonstrate how he is actively choosing, shaping and creating
his environments based on his genetic dispositions.
Not only does Oedipus consciously and unconsciously select certain
environments, he also perceives situations in a specific way. This
is another demonstration of active gene-environment correlation. For
example, he suspects Jocasta of being ashamed of his possible humble
origin, when she is not; he suspects Creon and Tiresias of plotting against
him, when they are not; he does not listen to advice, instead assigning
to others unreasonable intentions that are projections of his own mind.
Creon tells Oedipus that it would not make sense for him to plot against
him, as he is Oedipus’ brother-in-law and friend: why would he risk his
life?
But do not, on a dubious argument,

Charge me the beside the facts.
For just it is not,
To hold bad men for good, good men for bad,
To no good end: nay, ‘twere all one to me
To throw away a friend, a worthy one,
And one’s own life, which most of all one values27 .
The Chorus tries to convince Oedipus to listen to Creon, not to take

impulsive decisions:
His words sound fair – to one who fears to fall:

For swift in counsel is unsafe, by liege28 .
But Oedipus does not listen. His mind creates a menacing world that
in turn exacerbates his natural anxieties and obsessions. He cannot
stop himself from making harsh judgements of others. The Chorus
explains this behaviour by some internal stress (responding to Creon who
complains about Oedipus spreading nasty things about him).
Nay, the reproach, it may be, rather came

Through stress of anger than advisedly29 .
Another gene-environment process, called evocative gene-environment

correlation, also enhances the effects of our genes on our thoughts and
actions30 . We evoke particular reactions from people around us. Oedipus
may have evoked the incident at the crossroads, because he behaves as a
bully and is seemingly oblivious to his own flaws. He also evokes angry
responses from Tiresias and undermines his friendship with Creon. For
example, Tiresias tells him:
My offence
You censure: but your own, at home, you see not,
And yet blame me31 !
Sophocles continues to examine the themes of familiality and aetiology of

behaviour by following Oedipus to exile and telling us about the fate of
his children—in Oedipus at Colonus and in Antigone. Already in Oedipus
Rex we learn that Oedipus has had 4 children with Jocasta—2 sons (older
Polynices and younger Eteocles); and 2 younger daughters—Antigone
and Ismene. The children share much of their DNA with the parents and
with each other—due to incestuous conception. Does this make them
more similar in behaviour to each other than siblings on average? From
behavioural genetics we know that it is likely. Aside from incest, through
the process called assortative mating people tend to choose spouses who
are similar to them in physical attractiveness, verbal ability and some
other obvious traits32,33 . As these traits are partly influenced by genes,
spouses are usually more similar genetically than random people. In turn,
assortative mating leads to greater genetic and behavioural similarity of
offspring—so that some siblings are more similar to each other geneti-
cally than the average 50% of variable DNA (see Chapter 5 for more on
this).
From twin and adoption research, we also know that relatives who
share more DNA are usually more similar than relatives who share
less DNA. For example, monozygotic (MZ; identical) twins on average
are more similar than dizygotic (DZ; non-identical) twins in most
traits, including physical similarity, intelligence, motivation, academic
achievement and life satisfaction34–36 . However, we also know that

individual-specific events and experiences can contribute significantly to
differences—making even genetically identical MZ twins very different
from each other (although they are usually more similar than random
individuals).
We might expect that Ismene and Antigone will be very similar: they
share much of their genetic material; and both experience extremely
traumatic events, including their mother’s suicide, father’s exile and
extreme change in circumstances. Both daughters were very young when
Oedipus’ investigation into his origins ruins the lives of the entire family.
Oedipus describes his daughters in Oedipus at Colonus:
Young
Daughters of mine, twin curses37 !
But, in Antigone we learn that the girls are very different in thought and
behaviour—attesting to non-deterministic effects of genes and environ-
ments. Ismene is quiet, timid, accepting of the rules, accepting of inferior
status (in the men’s world), striving to avoid conflicts. Antigone is strong,
fiery and rebellious—resembling her father.
We don’t know much about Eteocles and Polynices except that they
fought against each other for power. In Oedipus at Colonus, Eteocles
sides with his uncle Creon against his brother. Polynices attempts to take
power from his uncle Creon. The brothers kill each other in battle.
Therefore, the whole family—Oedipus, his biological father, his
biological mother (also his wife), his sons and his daughters (who are also
his half-siblings)—all experience tragic life circumstances. In Oedipus at
Colonus, Oedipus suggests that the misfortunes, brought on his family,
may be the result of some ancient offence that his ancestors inflicted on
some deities. He says to Creon:
Such was the will of Heaven, that had some cause

For wrath it may be, with our house, of old38 .
And in Oedipus Rex he says to Creon:

Where to find it now,

the trail of the ancient guilt so hard to trace39 ?
Oedipus may be referring to events that unfolded before his conception,

when Laius had allegedly (according to the myth not mentioned in the
play, but mentioned in some other sources) committed a heinous crime
before he became king of Thebes, for which his whole lineage was cursed
by deities. Sophocles may also simply have referred to the fact that many
circumstances transmit from generation to generation. However, from
behavioural genetics we have learned that concrete biological processes,
rather than ancient curses, underlie this familial continuity.
From behavioural genetics we have learned that nothing is fully prede-
termined, as everyone’s genetic make-up and experiences are unique.
Through the processes described below, two siblings can turn out more
different from each other than two random individuals. Indeed, how
many of us have had the experience of meeting a stranger who is
a spitting image (in looks, mannerisms or features of behaviour) to
someone else we know? Such virtual twins result from them sharing
much of their variable DNA—simply by chance. As randomness tran-
scends geographical locations, we may all have virtual twins somewhere
in the world.
For one individual—we can never disentangle the effects of genes
and environments, as an individual is a complex product of the contin-
uous interplay between genetic and environmental factors. For example,
gene-environment interactions may partly explain differences between
Ismene and Antigone. Differences in their genetic codes provided them
with somewhat different sets of risk and resilience—leading to different
responses to the same traumatic events. Similarly, gene-environment inter-
actions may have contributed to shaping Oedipus’ character. We cannot
know how differently Oedipus (with his DNA code) would have devel-
oped, for example, if he had been raised by shepherds; or had he not
developed in the womb of a stressed mother.
Many studies have documented such interactive effects on both physi-
ology and psychology40,41 . For example, our bodies can withstand severe
attacks from pollutants, such as pesticides, flame retardants, lead and
mercury, that can enter our bodies by breathing polluted air, touching
contaminated surfaces or consuming contaminated foods. Although in
some people, these pollutants can lead to health problems and deficits in
cognitive function, other people are remarkably resilient to the effects
of toxins. Similarly, there are people who are remarkably resilient to
psychological ‘toxins’. Much research today is dedicated to understanding
genetic and environmental origins of such resilience42,43 .
Despite much research, our understanding of the processes involved in
shaping our behaviour and thought is far from complete. For example,
for centuries people have wondered what underscores differences among
people in personality characteristics. We often view personality as shaped
by our parents through discipline, personal example and training. It is
intuitively appealing to say that one’s mother’s example and instruc-
tion taught one to listen to people, to feel compassionate, to avoid
conflict and to find compromise. On the other hand, differences among
people in personality may reflect some initial biological differences. For
example, Sophocles’ contemporary and the father of modern medicine,
Hippocrates, theorised that personality traits and behaviour result from
an individual’s balance of the four bodily fluids (humours): blood, yellow
bile, black bile and phlegm44 . Although the specifics of this theory
have been refuted, the main premise (that personality is rooted in
biology) has been supported by modern science. Personality character-
istics, like any other trait, have been found to result from both genetic
and environmental influences45,46 .
Recent research also suggests that basic and unconscious processes,
such as our eye-movements, contribute to or reflect our personality.
For example, one study47 demonstrated that participants’ eye move-
ments, while running an errand on a university campus wearing an eye
tracking device, could be used as a window to their personality. Specif-
ically, eye-movement data were used to identify levels of some of the
so-called Big 5 personality dimensions: extraversion, neuroticism, consci-
entiousness, agreeableness and openness to experience. The authors of
the study concluded that personality greatly influences everyday eye
movement control. However, an alternative explanation is that some very
basic perceptual processes may underlie the development of different
personality characteristics. In turn, these basic perceptual processes differ
among people because of the differences in genetic and environmental
circumstances.
In fact, genes and environments always work together. Even traits

that are usually considered purely environmental are influenced by genes
to some extent. For example, social mobility—moving up from one’s
parent’s socio-economic status—is commonly viewed as an index of
environmental opportunity. Relatedly, lack of mobility (also known as
social persistence) is often thought to be fully explained by societal
limitations and unequal opportunities. However, today we know that
social mobility occurs (or does not occur) partly for genetic reasons48,49 .
Several studies, examining the DNA of family members, demonstrated
that inheriting a favourable combination of genes is associated with
upward social mobility; and inheriting a less favourable combination of
genes is associated with downward social mobility. For example, of two
brothers, the one with a greater number of education-increasing genetic
variants (higher polygenic score 50 ) is more likely to move up the social
ladder. Similarly, the daughter with a lower polygenic score than the father
and/or mother is at risk of moving down on the social ladder. Studies
have shown that children who achieved a greater educational or occupa-
tional level than their parents tended to have higher polygenic score than
their parents. Children whose social achievements were lower than their
parents’ tended to have lower polygenic scores.
Some research suggests that some of the persistence in social achieve-
ment within a family may be attributed to the intergenerational trans-
mission of genetic effects on cognitive, personality and motivational
traits51,52 . The research also suggests that some people who are born into
socially disadvantaged families, but manage to surpass the constraints
created by low socio-economic status (SES), do this in part because of
their genetic propensities. Recent research also indicates that in societies
with good environmental opportunities for all, educational attainment
depends less on social conditions and more on genetic make-up53,54 .
We generally tend to ignore gene-environment complexities and
selectively use genetic or environmental explanations (attributions) in
different contexts. An excellent example of this is when Oedipus takes
full credit for his ‘rising’ to prominence in Thebes: he boasts that
his strength and wisdom made him king in a foreign land. In reality
many factors contributed to his ‘rise’, including genetic predispositions,
affluent SES, ‘noble’ upbringing and a multitude of stochastic events 55–57 .
Yet, we lean towards simplistic, ‘parsimonious’ explanations of

behaviour. At worst, we attribute behaviour to a single causal source
(e.g. upbringing); at best—to a combination of traits, e.g. intelligence +
motivation + grit + parental support = success (each itself attributed to
some simplistic cause). This tendency can be likened to parsing out the
visible night sky into imaginary patterns of stars, which we call constel-
lations. In fact, human characteristics have often been described in terms
of underlying constellations of traits. Constellations are convenient for
describing what we see, but they do not represent any real systems, they
exclude many visible elements and they ignore the invisible patterns and
processes.
A better analogy for human behaviour and thought, unique to each
person, would be a galaxy, a gravitationally bound system of stars,
stellar remnants, interstellar gas, dust and dark matter. Each person is
a galaxy—a complex system of co-influencing processes unravelling over
time—a myriad of occurrences interacting with prior states, unravelling
on the genetic, epigenetic, neuronal and hormonal levels, to name just a
few. The processes of the mind are today discussed simplistically, refer-
ring to latent constructs (not directly measurable) such as motivation,
self-regulation, executive functions—into which we tap with our impre-
cise measures58,59 . These constructs resemble the ‘made up’ patterns of
constellations—limited by what pattern we choose to perceive in the
stars—person, object or creature. Similarly, our psychological concepts
are limited by our own definitions and are far from the real systems,
whose structure and laws we do not yet comprehend.
Lineage
Our confused views on influences of genes and environments on
behaviour manifest in our paradoxical obsession with ‘blood’, ‘lineage’
and ‘class/caste’. For example, Oedipus is devastated to discover that he
might not be of ‘noble’ blood. At first, he refuses to believe the messenger,
who, wanting to ‘free’ Oedipus from his fears of patricide and incest, tells
him that he wasn’t the biological son of Polybus and Merope:
Messenger. Since Polybus was naught to thee in blood.

Oedipus. What say’st thou? was not Polybus my sire?
Messenger. As much thy sire as I am, and no more.
Oedipus. My sire no more to me than one who is naught?
Messenger. Since I begat thee not, no more did he.
Know that he took thee from my hands, a gift.
Oedipus. Yet, if no child of his, he loved me well.
Messenger. A childless man till then, he warmed to thee.
Oedipus. A foundling or a purchased slave, this child?
Messenger. I found thee in Cithaeron’s wooded glens.
Oedipus. A vagrant shepherd journeying for hire?
Messenger. True, but thy savior in that hour, my son.
Oedipus. What, did another find me, not thyself?
Messenger. Not I; another shepherd gave thee me.
…
The same: he was a herdsman of the king60 .
Oedipus cannot believe that adoptive parents could have loved him as
much as biological parents. This is wonderfully expressed in another
translation as:
And loved me then, so much, at second hand61 ?
Why is this so difficult to believe? Why is ‘blood’ so important to

humans, despite our strong belief in the power of environmental influ-
ence on child development? In fact, the very act of adoption to ensure
succession, as in the case of Merope and Polybus, demonstrates that
blood is not important. However, the actions of Oedipus and other char-
acters throughout the Theban plays demonstrate the exact opposite. This
importance of ‘blood’/lineage has something to do with the unconscious
belief that something transmits within families, which goes beyond the
family environment (today we know that this something is the DNA).
This belief is beautifully expressed by Oedipus’ daughter in Antigone:
One husband dead, another might be mine;

Sons by another, did I lose the first;
But, sire and mother buried in the grave,
A brother is a branch that grows no more62 .
In Oedipus Rex , Oedipus is determined to discover his origins, he tells

Jocasta (who is trying to convince him to drop the investigation):
Nay, it cannot be
That having such a clue I should refuse
To solve the mystery of my parentage63 !
The gravity of his discoveries do not become apparent to Oedipus imme-

diately. In fact, for some time Oedipus believes that the worst that can
happen to him is that he would be found ‘slave born’ or of some other
humble origin. When Jocasta tells him: ‘Tis for thy sake I advise thee for
the best’, Oedipus says ‘I grow impatient of this best advice’, to which
Jocasta replies ‘Ah mayst thou ne’er discover who thou art!’64 .
It is an incredible delusion that a grown man can ‘discover who
he is’ through tracing his lineage. Oedipus is a king; he has his own
past, present and future—and yet the quest for his lineage becomes
all-consuming.
Oedipus is conflicted on the issue of lineage. He claims that it does
not matter to him what his roots are—that he will not be ashamed of
whatever his lineage is. He sees himself as ‘the child of Fortune’, some
deity that led to his growth from ‘small’ to greatness.
But I
Who rank myself as Fortune’s favourite child.
The giver of good gifts, shall not be shamed.
She is my mother and the changing moons
My brethren and with them I wax and wane.
Thus sprung why should I fear to trace my birth?
Nothing can make me other than I am65 .
This is supported by the Chorus, who suggests that Oedipus might have
been born of some nymph or deity.
Phoebus, may my words find grace!

Child, who bare thee, nymph or goddess? sure thy

sire was more than man66 .
At the same time, Oedipus obsesses over his origin and what others
may think about it. He engages in psychological projection—as he has
done with his accusations of Creon, Tiresias and the shepherd. It is his
mind that generates the concept of ‘unfit lineage’—but he unconsciously
defends his psyche against the harmful effects of this concept by denying
its existence in his mind and attributing it to Jocasta’s pride.
But she, perhaps, being, as a woman, proud,

Of my unfit extraction is ashamed67 .
Oedipus thinks that Jocasta tries to prevent him from investigating

his lineage because she is afraid to find out that she is married to a
non-noble—someone from a non ‘gilded lineage’, demonstrated in this
conversation between them:
Jocasta. For Heaven’s sake, if you care for your own life,
Don’t seek it! I am sick and that’s enough!
Oedipus. Courage! At least, if I be thrice a slave,
Born so three-deep, it cannot injure you!
Jocasta. Wretch, what thou art O might’st thou never know!
Oedipus. Will some one go and fetch the herdsman hither?
She is welcome to her gilded lineage!
Jocasta. Woe, woe, unhappy! This is all I have
To say to thee, and no word more, for ever68 !
Oedipus makes no attempt to work it out with his wife, to understand

her reasons. He is too self-obsessed—interpreting her words as if she does
not want to talk to him because he might not be a king by origin. He is
determined to trace ‘the source of him’ no matter how ‘low it may be’.
Chorus warns of the trouble that may come from Jocasta leaving with
grief. By now, Jocasta’s suspicions had grown into terrible certainty—
Oedipus was her son. Unable to stop him from his destructive enquiry,
she retreats to the palace to kill herself.
Why has the woman vanished, Oedipus,

Driven so wild with grief? I am afraid
Out of her silence will break forth some trouble.
But Oedipus is determined to continue with his quest, no matter what.
Break out what will, I shall not hesitate,

Low though it be, to trace the source of me69 .
When the herdsman (who had received baby Oedipus from Jocasta)
arrives, the messenger from Corinth asks him:
Well, thou mast then remember giving me

A child to rear as my own foster-son70 ?
The herdsman does not want to speak, knowing that this would only
lead to more trouble. This triggers Oedipus’ temper:
Oedipus. If thou lack’st grace to speak, I’ll loose thy tongue.

Herdsman. For mercy’s sake abuse not an old man.
Oedipus. Arrest the villain, seize and pinion him!
The herdsman gives in.
Herdsman. I had it from another, ’twas not mine.

Oedipus. From whom of these our townsmen, and what house?
Herdsman. Forbear for God’s sake, master, ask no more.
Oedipus. If I must question thee again, thou’rt lost.
Herdsman. Well then—it was a child of Laius’ house.
Oedipus. Slave-born or one of Laius’ own race?
Herdsman. Ah me!
I stand upon the perilous edge of speech.
Oedipus. And I of hearing, but I still must hear.
Herdsman. Know then the child was by repute his own,
But she within, thy consort best could tell71 .

And so, from the ‘edge of hearing’ Oedipus falls into the abyss of
knowing. He finally realises that it was his wife, Jocasta, also his
mother—who gave him as a baby to the herdsman ‘to make away with’.
Oedipus. What! she, she gave it thee?

Herdsman. ’Tis so, my king.
Oedipus. With what intent?
Herdsman. To make away with it.
Oedipus. What, she its mother.
Herdsman. Fearing a dread weird.
Oedipus. What weird?
Herdsman. ’Twas told that he should slay his sire.
Oedipus. What didst thou give it then to this old man?
Herdsman. Through pity, master, for the babe. I thought
He’d take it to the country whence he came;
But he preserved it for the worst of woes.
For if thou art in sooth what this man saith,
God pity thee! thou wast to misery born72 .
And so, his tragic quest comes to an end. It is meaningful that Oedipus
blinds himself, losing his sight, mirroring the metaphorical blindness of
his thoughts and actions throughout the play. It also is meaningful that
in his final speech he contrasts the happy past in his adoptive home with
his newly redefined past that makes him ‘ill’.
O Polybus,
And Corinth, and the old paternal roof
I once called mine, with what film of honour,
Corruption over-skinned, you fostered me,
Found ill myself, and from ill parents, now73 !
Oedipus falls from the greatest of great to the lowest of low—beautifully

summarised by the Chorus.
Who, shooting far beyond the rest,

Won wealth all-blest,
Slaying, Zeus, thy monster-maid,
Crook-taloned, boding; and
Who did arise and stand

Betwixt death and our land,
A tower of aid;
Yea, for this cause thou hast been named our king,
And honored in the highest, governing
The city of Thebes great
In royal state.
And now, who lives more utterly undone?

Who with sad woes, who with mischances rude
Stands closer yoked by life’s vicissitude74 ?
Chance, Genes and Crossroads

Chance is a major force that contributes to every aspect of our life.
In fact, life is possible because of stochastic events (στóχoς—stokhos
meaning to ‘guess’)—events that have no certainty, are random, non-
systematic, unpredictable and outside our control. Our uniqueness is a
product of stochastic (random) mutations that occur in DNA. These
random events are then followed by systematic evolutionary events, such
as spreading of the new mutations in the population via natural selec-
tion. Once we possess a certain genetic variant, it affects us via systematic
genetic processes, such as gene expression.
Environmental effects also come in two types: systematic environ-
mental processes, crafted by climatic and other natural phenomena and
culture; and stochastic events that can alter one’s life trajectory at any
moment75,76 . This is beautifully illustrated throughout Oedipus Rex .
Oedipus’ life is filled with vicissitude—fluctuations of favourable and
unfavourable events and situations that occur by chance. These changes
of circumstances and fortune could have happened to anyone, but
Oedipus’ reaction to them is unique to him. This is a clear demonstration
of how chance interacts with genetic propensities and other influences. By
the time we respond to chance we are already a product of endless gene-
environment processes. For example, a drunk man talking about Oedipus
and Oedipus overhearing this is a stochastic event. The consequent
obsession with tracing his lineage is not. It is a product of his character.
His adoptive parents’ decision not to reveal his origin to him might be a
product of societal norms or their own peculiarities.
In behavioural genetics, chance is included in the estimate of non-shared
environment —as it usually contributes to dissimilarities between family
members, including between identical (MZ) twins 77–79 . For example,
one sibling may overhear something that leads to the redefining of
her past (see Chapter 4), whereas another sibling may carry on ‘in the
dark’—diverging their psychological trajectories.
A wonderful description of a chance event that ‘derails’ all can be
found in Sophocles’ Electra—although ‘chance’ is attributed to some
deity’s harmful whim. In the play, one character (Guardian) is describing
to Electra how her brother Orestes died in a chariot event (bizarrely—
as Orestes was actually alive and planned this piece of temporary
misinformation with the Guardian):
‘But if some Deity is bent on harm,

It is not even a strong man can escape.
For he, another day at sunrise, when
Owners of horses met to try their speed,
With many other charioteers, went in.
… Then, in an instant, from one accident,
Car upon car began to crash and fall,
And the whole plain of Crisa became filled
With wreck of steeds and tackling80 .
Similarly, ‘in an instant, from one accident…’ Oedipus’s life turns into
a plain of ‘crashing chariots’, or as Oedipus describes to the citizens of
Colonus: ‘Wave upon wave of evils, numberless81 !’
Which ‘chance’ (or semi-chance) instance can be considered as life-
changing for Oedipus? Perhaps, the most obvious candidate for this is
the incident at the crossroads. Oedipus tells us:
As I journeyed on,
I found myself upon the self-same spot
Where, you say, this king perished. In your ears,
Wife, I will tell the whole. When in my travels
I was come near this place where three roads meet82 .
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Vitriniconus 16
Sitala 2
Kaliella 8
Trochomorpha 21
Endodonta 1
Plectopylis 3
Plectotropis 1
Aulacospira 3
Pupisoma 1
Satsuma 2
Dorcasia 2
Chloritis 7
Obbina 19
Papuina 1
Phoenicobius 7
Cochlostyla 247
Amphidromus 2
Hapalus (?) 4
Hypselostoma 1
Pupa 4
Clausilia 1
Subulina 3
Prosopeas 2
Opeas 4
Geostilbia 1
Tornalellina 1
Succinea 3
Vaginula 2
Ancylus 1
Limnaea 3
Planorbis 3
Physa 2
Melania 50
Pirena 2
Bithynia 1
Vivipara 7
Ampullaria 5
Acmella 2
Diplommatina 41
Arinia 6
Pupina 5
Registoma 7
Hargreavesia 1
Callia 2
Pupinella 3
Helicomorpha 4
Coptochilus 1
Alycaeus 1
Leptopoma 42
Lagochilus 11
Cyclophorus 31
Ditropis 7
Cyathopoma 5
Cyclotus 19
Omphalotropis 3
Helicina 18
Georissa 3
Anodonta 1
Cyrena 3
Corbicula 7
Islands adjacent to the Philippines.—The Philippines are
connected with Borneo by two distinct ridges or banks of elevation,
which enclose between them the Soo-loo or Mindoro Sea. There can
be little doubt that these ridges represent the ancient highway of
transit, by which Indo-Malay species passed into the Philippines. The
depth of the sea on either side is profound, ranging from an average
of about 1000 fathoms west of Palawan to 2550 off the south-west
coast of Mindanao.
It appears that the fauna of the Soo-loo ridge is definitely
Philippine up to and including Bongao, Sibutu, and Bilatan, the last
islands at the Bornean end of the ridge. On these are found two
species of Cochlostyla and an Obbina.
The Palawan ridge may also be described as more or less
Philippine throughout. One species of Cochlostyla occurs on
Balabac, just north of Borneo, and two on Palawan, but these are
perhaps counterbalanced by the definitely Indo-Malay Amphidromus
and Opisthoporus (1 sp. each). At the northern end of the ridge, on
Busuanga and Calamian, the Philippine element predominates.
Representatives of two remarkable groups of Helix (Camaena and
Phoenicobius) occur along the Palawan ridge and in Mindoro. The
Phoenicobius find their nearest allies in the curious small group
known as Obba, from N. Celebes, the Camaena possibly in a type of
Helix (Hadra) occurring in New Guinea and N.E. Australia. The only
other Helix from the whole of the E. Indies which bears any
resemblance to the Phoenicobius group is H. codonodes Pfr., which
is peculiar to the Nicobars. A few forms assigned to Camaena also
occur in Further India and Siam. It would appear possible, therefore,
that these two isolated groups are a sort of survival of a fauna which
perhaps had once a much more extended range.
(2) The Chinese Sub-region.—The Chinese Sub-region includes
the whole of China from its southern frontier up to and including the
basin of the Blue or Yang-tse River, together with the coast district,
including Corea, perhaps as far north as Vladivostok, and the
outlying islands of Hainan, Formosa, the Loo-Choo and Bonin
groups, and Japan to the north of Niphon. It may be divided into two
provinces, the Chinese and the Japanese.
(a) The fauna of the Chinese province proper bears, in many
respects, strong marks of relationship to that of India and Siam. Thus
Streptaxis, Helicarion, Macrochlamys, Kaliella, Sitala, Ariophanta,
Rhysota, Hemiplecta, Diplommatina, Opisthoporus, Pterocyclus,
Lagochilus, and Alycaeus all occur, especially in Southern China.
The two points in which the sub-region bears special marks of
individuality are Helix and Clausilia. The sub-genera of Helix which
have their metropolis in China are Satsuma, Cathaica, Aegista,
Acusta, Euhadra, Plectotropis, and Plectopylis. Sinistral forms
(compare Fig. 213) are rather prevalent. In several cases—e.g.
Trichia, Gonostoma, Fruticicola—there is a reappearance of forms
which appear to belong to well-known European sub-genera.
Clausilia here attains a kind of second centre of distribution, and is
represented by its finest forms, which belong to several peculiar sub-
genera. The carnivorous Mollusca are not abundant, and are
represented by Rathouisia (a peculiar genus of naked slug), Ennea,
and Streptaxis. In the western provinces Buliminus is abundant in
several sub-genera, one of which appears to be the European
Napaeus.
Fig. 213.—Helix
(Camaena) cicatricosa
Müll., China.
There is little which is striking in the operculates, which are most
abundant in the south, and appear to be mainly derived from Indian
and Siamese sources. The occurrence of Helicina (3 sp.),
Omphalotropis (1), Leptopoma (2), and Realia (2), is evidence of
some influence from the far East. Heudeia is a very remarkable and
quite peculiar form of Helicina with internal plicae, perhaps akin to
the Central American Ceres.
Fresh-water genera are exceedingly abundant, especially
Melania, Unio, and Anodonta. The occurrence of Mycetopus (a
South-American genus) is remarkable. There are several peculiar
forms of fresh-water operculates, whose exact position is hardly yet
assured.
Land and Fresh-water Mollusca of the Chinese Province
Rathouisia 1
Streptaxis 7
Ennea 12
Parmarion 2
Helicarion 15
Euplecta 3
Macrochlamys 19
Microcystina 2
Microcystis 7
Kaliella 16
Sitala 8
Ariophanta 1
Rhysota 5
Hemiplecta 1
Trochomorpha 2
Limax 1
Philomycus 1
Patula 2
Gonostoma 4
Metodontia 2
Vallonia 1
Plectotropis 9
Fruticicola 11
Satsuma 14
Trichia 10
Cathaica 22
Aegista 10
Armandia 3
Acusta 15
Obbina 1
Camaena 5
Euhadra 14
Plectopylis 19
Stegodera 6
Chloritis 1
Hel. Inc. sed. 39
Buliminus 21
Buliminopsis 3
Buliminidius 3
Napaeus 14
Rachis (?) 4
Pupa 10
Clausilia 102
Opeas 12
Euspiraxis 1
Subulina 5
Stenogyra (?) 12
Succinea 8
Vaginula 7
Limnaea 2
Planorbis 6
Melania 44
Paludomus 3
Bithynia 12
Lithoglyphus 3
Melantho (?) 1
Pachydrobia 1
Prososthenia 2
Stenothyra 2
Hydrobia 2
Mecongia 1
Oncomelania 9
Margaracya 1
Rivularia 4
Delavaya 1
Fenouillia 1
Vivipara 34
Diplommatina 20
Pupina 6
Alycaeus 23
Leptopoma 2
Lagochilus 10
Cyclophorus 18
Coelopoma 1
Pterocyclus 3
Opisthoporus 4
Cyclotus 10
Scabrina 4
Ptychopoma 12
Omphalotropis 1
Realia 2
Pseudopomatias 1
Helicina 3
Georissa 4
Heudeia 1
Cyclas 1
Corbicula 50
Unio 53
Monocondylaea 1
Anodonta 55
Mycetopus 12
Pseudodon 1
Dipsas 4
The island of Hainan, in the extreme south of the sub-region, has

40 species of Mollusca, 22 of which are peculiar, but there is no
peculiar genus.
The Mollusca of Formosa, although in many cases specifically
distinct, show close generic relationship with those of China. The
characteristic Chinese groups of Helix and Clausilia occur, and there
is still a considerable Indian element in several species of Streptaxis,
Macrochlamys, Kaliella, and Alycaeus. The occurrence of two
Amphidromus, a genus which, though Siamese, is not found in
China or Hainan, is remarkable.
The peninsula of Corea must undoubtedly be included in the
Chinese sub-region. It is true that the land operculates scarcely
occur, but there are still a number of Clausilia, and several of the
characteristic Chinese groups of Helix are reproduced. In some
points Corea appears to show more affinity to Japan than to China,
four of the Helices being specifically identical with those of Japan,
but the peninsula is at present too little explored for any
generalisations to be made as to its fauna in this respect.
(b) Japanese Province.—Kobelt distinguishes four groups of
Mollusca inhabiting Japan (a) circumpolar species, actually occurring
in Europe, Siberia, or N. America, or represented by nearly allied
species (these of course do not belong to the Japanese province as
such); (b) Indo-tropical species; (c) species which are Chinese or
akin to Chinese; (d) peculiar species, a mixture of two forms,
southern and northern, the latter being chiefly Hyalinia, Patula, and
Fruticicola. Out of a total of 193 Japanese species, at least 164 are
peculiar.
The Japanese Helices belong to sub-genera common to China
(Plectotropis 8, Euhadra 21, Acusta 23?); but the Naninidae scarcely
occur at all. The principal feature of the fauna is the development of
Clausilia, which presents some extraordinarily fine forms. One slug
(Philomycus) is identical with an Indian species. The operculates,
which consist mainly of a few species each of Diplommatina,
Cyclophorus, Pupinella, Pupina, Helicina, and Georissa, belong
almost exclusively to the southern islands Kiu-siu, Sikoku, and
southern Niphon. The three species usually reckoned as Japonia are
probably forms of Lagochilus.
C. The Australasian Region

This region includes all the islands of the Pacific east of the
Moluccas, and falls into three sub-regions—the Papuan, the
Australian, and the Polynesian.
1. The Papuan Sub-region may be divided into—(a) the Papuan
Province proper, which includes New Guinea, with the Aru Is. and
Waigiou, the Admiralty Is., New Ireland, New Britain, and the
d’Entrecasteaux and Louisiade Groups; (b) the Queensland
Province, or the strip of N.E. Australia from C. York to the Clarence
R. (about 29° S. lat.); (c) the Melanesian Province, which includes
the New Hebrides, New Caledonia, with the Loyalty Is. and the Viti
Is. The Solomons form a transition district between the Papuan and
Melanesian provinces, abounding on the one hand in characteristic
Papuan Helices, while on the other they form the north-western limit
of Placostylus, the group especially characteristic of the Melanesian
province.
(a) The Papuan Province.—The molluscan fauna of New Guinea
is the richest and by far the most original of all the Australasian
region. We find ourselves, almost in a moment, in a district full of
new and peculiar forms. New Guinea may be regarded as the
metropolis of the rich Helicidan fauna, which is also characteristic of
the Moluccas to the west, of N. and N.E. Australia to the south and
south-east, and of the Solomons and other groups to the north-east.
Here abound species of Papuina and Insularia (the latter being quite
peculiar), among which are found, if not the largest, certainly the
most finished forms of all existing Helices. Chloritis (13 sp.),
Planispira (5), and Cristigibba (9) are common with the Moluccas,
while a tropical Australian element is shown in Pedinogyra (1) and
Hadra (4). Very remarkable, too, is the occurrence of one species of
Obbina and Rhysota, genera which culminate in the Philippines and
here find their most eastward extension; while a single Corasia
serves to form a link between the Corasia of the Philippines and
those of the Solomon Is., if the latter are true Corasia.
We naturally find considerable traces of a Polynesian element,
which appears to be principally characteristic of the eastern part of
the island. Most noteworthy in this respect is the occurrence of
Partula (3), Tornatellina (1), Charopa (1), Thalassia (3). As compared
with the true Pulmonata, the operculates are feebly represented, and
the great majority are of a markedly Polynesian type. Not a single
Cyclophorus occurs; Lagochilus, Alycaeus, and all the tubed
operculates, so marked a feature of the Indo-Malay fauna, are
conspicuous by their absence, and the prevailing genera are
Cyclotus, Helicina, and a number of sections of Pupina. Leptopoma,
as in the Philippines, is strongly represented. Not that an Indo-Malay
element is altogether absent. We still have Xesta (5), Hemiplecta (8),
and even Sitala (2), but the great predominance of Helix seems to
have barred the progress, for the greater part, of the Indian
Naninidae.
The slugs appear to be represented by a solitary Vaginula. A
single Perrieria is a very marked feature of union with Queensland,
where the only other existing species (P. australis) occurs. The
solitary Rhytida, so far the only representative of the carnivorous
group of snails, emphasises this union still further. Little is known of
the fresh-water fauna. Melania (28 sp.) is predominant, but on the
whole the relations are Australian rather than Indo-Malay. Ampullaria
is wanting, while a decisive point of similarity is the occurrence of
Isidora (3 sp.), a genus entirely strange to the Oriental region, but
markedly characteristic of the Australasian.
Land and Fresh-water Mollusca of New Guinea
Rhytida 1
Helicarion 2
Rhysota 1
Hemiplecta 11
Xesta 2
Microcystis 3
Microcystina 2
Sitala 2
Oxytes (?) 2
Conulus 1
Trochomorpha 8
Nanina (?) 3
Charopa 1
Thalassia 3
Ochthephila(?) 1
Chloritis 13
Planispira 5
Cristigibba 9
Insularia 17
Obbina 1
Albersia 3
Hadra 4
Pedinogyra 1
Papuina 35
Corasia (?) 1
Bulimus (?) 1
Calycia 4
Partula 3
Pupa 1
Stenogyra 1
Tornatellina 1
Perrieria 1
Succinea 1
Vaginula 1
Limnaea 2
Isidora 3
Melania 28
Faunus 1
Vivipara 4
Diplommatina 1
Pupina 4
Pupinella 3
Omphalotropis 2
Bellardiella 2
Leptopoma 16
Cyclotus 5
Cyclotropis 5
Helicina 15
Unio 4
Cyrena 3
Corbicula 1
Batissa 8
Waigiou is practically a part of New Guinea. Twelve genera and
twenty species of Mollusca are known, eight of the latter being
peculiar. The occurrence of Papuina, Insularia, and Calycia
sufficiently attest its Papuan relationship. Two species each of
Albersia, Chloritis, and Planispira occur.[372]
The Aru Is. are, as we should expect from their position, and
particularly from the configuration of the adjacent sea bottom (see
map), markedly Papuan. At the same time they show unmistakable
signs of long-continued separation from the parent island, for of their
36 land Mollusca 15, and of their 20 fresh-water Mollusca 9 are
peculiar. The Papuan element consists in the presence of Papuina,
Albersia, and Cristigibba. Moluccan influence is not absent, for the
three Helicina, the Albersia, and one Cyclotus are all Moluccan
species. The fresh-water fauna appears to be a mixture of varied
elements. The single Segmentina is common to India, the
Glaucomya to Malacca and the Philippines, while the single Batissa
is also found in New Zealand.
Land and Fresh-water Mollusca of the Aru Islands
Xesta 4
Microcystis 1
Hyalinia(?) 1
Trochomorpha 1
Patula 1
Eulota 1
Chloritis 5
Cristigibba 2
Albersia 1
Papuina 4
Pupa 2
Stenogyra 2
Planorbis 1
Segmentina 1
Melania 14
Leptopoma 3
Moussonia 1
Realia 1
Cyclotus 3
Helicina 3
Cyrena 2
Glaucomya 1
Batissa 1
The Louisiades, the d’Entrecasteaux, and Trobriand Is., and
Woodlark I., are closely related to New Guinea, containing no
peculiar genera. Each group, however, contains a considerable
proportion of peculiar species, an indication that their separation
from New Guinea dates from a very distant period. From the
Louisiades are known 34 species in all, 22 of which are peculiar.
The fauna of the Admiralty Is., of New Hanover, and New Ireland
is markedly Papuan, without any especial feature of distinction. The
Admiralty Is. contain 15 sp. Papuina, 7 Chloritis, 1 Planispira, and 1
Corasia. A single Janella shows relationship with the New Hebrides
and with New Zealand. In New Ireland Planispira (which is specially
characteristic of W. New Guinea and the Moluccas) has
disappeared, but there are 7 Papuina and 6 Chloritis. The essentially
Polynesian Partula is present in both groups.
The prominent feature of the Mollusca of the Solomon Is. is the
extraordinary development of Papuina, which here culminates in a
profusion of species and singularity of form. The genus is arboreal,
crawling on the branches and attaching itself to the leaves of trees
and underwood. Of the 140 land Pulmonata known from the group,
no less than 50, or 36 per cent, are Papuina. Ten species of Corasia
occur, but whether the shells so identified are generically identical
with those of the Philippines, is not satisfactorily determined.
Trochomorpha, with 22 species, here attains its maximum. Chloritis
begins to fail, but still has 3 species. Indo-Malay influence still
appears, though feebly, in Hemiplecta (3), Xesta (1), and possibly
even Macrochlamys (1). The Rhytida, the 3 Hadra, and possibly the
Paryphanta represent the Australian element. The growing numbers
of Partula (13), the small and inconspicuous land operculates (only
22 in all, with Helicina very prominent), and the almost complete
absence of fresh-water bivalves, show signs of strong Polynesian
affinities. An especial link with the New Hebrides, New Caledonia,
and the Viti Is. is the occurrence of Placostylus (16 sp.). It is very
remarkable that this genus should occur in the Solomon Is. and not
in New Ireland. The occurrence of Streptaxis, if authentic, is very
noteworthy, the nearest species being from the Philippines.
Land and Fresh-water Mollusca of the Solomon Islands
Streptaxis (?) 1
Rhytida 1
Paryphanta (?) 1
Helicarion 2
Xesta 1
Macrochlamys 1
Hemiplecta 3
Microcystis 2
Trochomorpha 22
Nanina (?) 2
Patula 1
Thalassia 2
Chloritis 3
Philina 2
Hadra 3
Papuina 50
Merope 1
Corasia (?) 10
Placostylus 16
Partula 13
Succinea 1
Melania 18
Diplommatina 2
Pupina 4
Leptopoma 4
Omphalotropis 2
Cyclotus 1
Cyclotropis 2
Helicina 7
Unio 1
(b) The Queensland Province.—The strip of coast-line from Cape

York to the Clarence R. stands apart from the rest of Australia, and is
closely connected with New Guinea. There can be little doubt that it
has been colonised from the latter country, since an elevation of
even 10 fathoms would create (see map) a wide bridge between the
two. Many of the genera are quite strange to the rest of Australia.
Land operculates are abundant, and of a Papuan type. Several of
the characteristic Papuan genera of Helix (Papuina, Chloritis,
Planispira) occur, while Hadra attains its maximum. Panda,
Pedinogyra, and Thersites are three remarkable groups in a rich
Helix fauna. Parmacochlea is a peculiar form akin to Helicarion. The
carnivorous Mollusca are represented by Rhytida, Diplomphalus
(New Caledonia), and Elaea. One species of Janella, a slug peculiar
to this region, occurs. The predominant fresh-water genus is Bulinus
(Isidora). Ampullaria and Anodonta are entirely absent from Australia
and New Zealand.
Fig. 214.—Characteristic
Australian Helices: A, H.
(Hadra) pomum Pfr.; B, H.
(Thersites) richmondiana Pfr.
× ⅔.
Map D. To face page 322.
MAP
to illustrate the relations
OF THE LAND MOLLUSCA OF
NEW GUINEA WITH THOSE
OF NORTH AUSTRALIA.
The red line marks the 100 fathom line
London: Macmillan & Co.
Land Mollusca of the Queensland Province

Diplomphalus 1
Rhytida 10
Elaea 1
Parmacochlea 1
Helicarion 7
Nanina 3
Hyalinia 10
Thalassia 4
Charopa 5
Patula (?) 4
Macrocyclis (?) 1
Helicella 10
Planispira 8
Hadra 51
Chloritis 5
Pedinogyra 1
Thersites 1
Papuina 6
Panda 2
Helix (inc. sed.) 6
Bulimus (?) 1
Stenogyra 1
Tornatellina 4
Pupa 3
Vertigo 4
Perrieria 1
Succinea 3
Vaginula 1
Janella 1
Georissa 1
Pupina 16
Hedleya 1
Callia 1
Diplommatina 3
Ditropis 2
Dermatocera 1
Helicina 8
(c) The Melanesian Province includes those islands on which the

remarkable group Placostylus occurs, the metropolis of whose
distribution is New Caledonia. These islands are very possibly the
remains of what was once a much wider extent of land. A single
species of Placostylus occurs both on Lord Howe’s I. and in the
North I. of New Zealand, but this fact, while highly interesting as
indicating a possible former extension of land in a south-easterly
direction, is hardly sufficient to bring these islands within the
province as now limited. The Solomon Is., although containing
Placostylus as far to the west as Faro I., form, as has been already
stated, a transitional district to the Papuan province.
New Caledonia.—The chief features of the Mollusca are the
remarkable development of the helicoid carnivorous genera Rhytida
(30 sp.) and Diplomphalus (13 sp.), and of Placostylus (45 sp.).
There is a stray Papuina, and a peculiar form Pseudopartula, but
Helix has almost entirely disappeared. Polynesian influence is
represented by Microcystis (3 sp.), the so-called Patula (13 sp.,
many of which are probably Charopa), Tornatellina (2 sp.), and
Helicina (20 sp.). Partula does not reach so far south, but there are
two species of Janella. The recurrence of Melanopsis (19 sp.),
absent from the whole Oriental region, is curious, and forms another
link with New Zealand. The curious sinistral Limnaea (Isidora),
common with Australia and New Zealand, is abundant.
Fig. 215.—Placostylus
caledonicus Pet., New
Caledonia, × ⅔.
The New Hebrides link New Caledonia and the Solomons by their
possession of the typical heavy Placostylus (5 sp.) of the former, and
the lighter and more elegant Charis (2 sp.) of the latter. There are 4
Papuina, and Partula is abundant (18 sp.), but there is no evidence
at present that the carnivorous genera or the Melanopsis and Isidora
of New Caledonia occur.
The Fiji Is., by the possession of 14 Placostylus of the Charis
section, which is entirely absent from the adjacent Tonga group, form
the eastern limit of the province. There appears to be only a single
Partula, but the Polynesian element, especially as seen in Navicella
(8 sp.), Neritina (20 sp.), Helicina (11 sp.), and Omphalotropis (11
sp.), is very strong. The Microcystis (9 sp.) and Trochomorpha (14
sp.) are also of a Polynesian type.
(2) The Australian Sub-region includes the whole of Australia
(with the exception of the Queensland province) and Tasmania, with
New Zealand and the off-lying islands. The fauna, from the
prevalence of desert, is scanty, especially in genera. Land
operculates are almost entirely wanting. Limax is not indigenous,
though several species have become naturalised. The bulk of the
fresh-water species belong to Isidora, and it is doubtful whether
Physa occurs at all. Unio has a few species, and also Vivipara, but
neither Anodonta nor Ampullaria occur. There are a few Melania and
Neritina.
Tropical South Australia.—The Mollusca are scanty, and occur
chiefly in the neighbourhood of the rivers, the soil being arid, with no
shelter either of trees or rocks. Fresh-water species predominate,
and the rich land fauna of Queensland is totally wanting. There are
no land operculates, 6 Hadra, 1 Bulimus (?), 1 Stenogyra.
West Australia.—Owing to the deserts which bound it, the
Mollusca are very isolated, only one species being common with N.,
S., and E. Australia. The chief characteristics are Liparus, a form
intermediate between Helix and Bulimus, and, among the Helices,
the group Rhagada. There are no slugs, no carnivorous snails, and
only three land operculates.
Land Mollusca of West Australia
Lamprocystis 1
Hyalinia 1
Patula 7
Chloritis 2
Gonostoma 2
Trachia 3
Xerophila 1
Rhagada 8
Hadra 5
Liparus 10
Pupa 4
Succinea 3
Cyclophorus 2
Helicina 1
In Eastern and Southern Australia (New South Wales, Victoria,
and South Australia) the tropical element, so abundant in
Queensland, almost entirely disappears, the last operculate (a
Helicina) only reaching Port Macquarie, though several species of
Helicarion occur in the extreme south. Hadra is still abundant in New
South Wales (18 sp.) and S. Australia (10 sp.), but becomes scarce
in Victoria (2 sp.); New South Wales has also one Panda and two
Thersites. Cystopelta is common with Tasmania, and one of the
Janellidae (Aneitea) with Queensland. The carnivorous snails are
represented by Rhytida. Caryodes, a bulimoid group perhaps akin to
Liparus, is common with Tasmania only.
Tasmania.—About 80 species of land Mollusca are known, not
more than 10 being common with Australia. No land operculates
occur; Endodonta and Charopa are rare, and Hadra has entirely
disappeared, but Pupa and Succinea occur. Carnivorous genera are
represented by Paryphanta, Rhytida, and Rhenea. Anoglypta is a
peculiar section of Helix, while Caryodes, Cystopelta, and Helicarion
are common with Australia. Among the fresh-water Mollusca are a
Gundlachia (see p. 345), and some forms of Amnicola or Hydrobia,
one of which (Potamopyrgus) is common only with New Zealand.
[373]
The Neozealanian Province.—The Mollusca of New Zealand, with

the Kermadec, Chatham, and Auckland Is., are remarkably isolated.
Such genera as Nanina, Partula, Pupa, Stenogyra, Succinea,
Vaginula, Truncatella, Helicina, and Navicella, which might have
been expected to occur, are entirely absent. The bulk of the land
Mollusca are small and obscure forms, perhaps remains of a very
early type, and appear to belong to the Zonitidae, neither Patula nor
Helix occurring at all. The carnivorous forms are represented by
Schizoglossa, a peculiar genus akin to Daudebardia, by Paryphanta,
an extraordinary group of large shells with a thick leathery epidermis,
and by Rhytida and Rhenea. In spite of its extreme isolation, the
general relations of the fauna are partly with New Caledonia, partly
with E. Australia. The occurrence of Placostylus has already been
mentioned (p. 323), and three species of Janella, a genus which also
occurs in Queensland and New Caledonia, indicate the same affinity.

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Oedipus Rex in the Genomic Era Yulia

Genomic Politics: How the Revolution in Genomic Science

The Oedipus Plays of Sophocles: Philosophical

Applications of genetic and genomic research in cereals

Genomic Data Sharing Jennifer B. Mccormick

The Collaborative Era in Science: Governing the Network

Water Conservation in the Era of Global Climate Change

Risk and Resilience in the Era of Climate Change Vinod

Salman’s Legacy: The Dilemmas of a New Era in Saudi

Yulia Kovas · Fatos Selita

ISBN 978-1-349-96047-7 ISBN 978-1-349-96048-4 (eBook)

Cover illustration: Daniel Hernanz Ramos/Getty Images

We dedicate this book to our parents.

1 A Journey into the Genomic Era: Sophocles Will Be

3 Free Will in the Genomic Era 45

Randomness, Feedback and the Absence of a Plan 154

Fig. 1.1 Oedipus’ life journey. Oedipus’s life journey starts

Box 5.1 DNA Structure 114

that contribute to similarities and differences among people; roots and

Oedipus Rex and Behavioural Genetics

…For that maternal wedding,

Have dreamed as much; but he who by such dreams

Although this is an interesting and memorable theme, the play is about

differences of family members. We explore gene-environment interplay

numerous biases; erroneous assumptions about human behaviour; and

In the Epilogue to this book, the Chorus of Citizens of the Genomic

Plot of the Three Theban Plays

this incestuous relationship. Horrified, Oedipus flees Corinth to escape

Unwittingly, Oedipus swears to do anything to find the murderer. Not

Characters of the Theban Plays (and Their

Polynices—son of Jocasta and Oedipus; brother (possibly twin) of

A Note on Citations and Translations

Sweet-voiced daughter of Zeus from O Prophecy of Jove, whose words

List of Translations Used in This Book

Ah mayst thou ne’er discover who thou art!1

I must pursue this trail to the end,

Sophocles, his characters and audiences would have called to divine

While from our son’s birth not three days went by

My sire was Polybus of Corinth and

Oedipus is also extremely arrogant and prone to hubris—often

Here too my zeal has nothing lagged, for twice

In another example of hubris, Oedipus implies that Tiresias is a useless

The epitome of Oedipus’ hubris, his hamartia, is that throughout the

Oedipus. Are these inventions Creon’s work, or yours?

Oedipus is also obsessive—for example, even having lived in a foreign

Oedipus. Of course I fear lest Phoebus turn out true.

But do not, on a dubious argument,

The Chorus tries to convince Oedipus to listen to Creon, not to take

His words sound fair – to one who fears to fall:

Nay, the reproach, it may be, rather came

Another gene-environment process, called evocative gene-environment

Sophocles continues to examine the themes of familiality and aetiology of

achievement and life satisfaction34–36 . However, we also know that

Such was the will of Heaven, that had some cause

And in Oedipus Rex he says to Creon:

Where to find it now,

Oedipus may be referring to events that unfolded before his conception,

In fact, genes and environments always work together. Even traits