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S T EV E C L A R K E , HA Z E M Z O H N Y,
A N D J U L IA N S AV U L E S C U
1
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1
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Contents
Preface vii
Notes on Contributors xiii
1. Rethinking our Assumptions about Moral Status 1
Steve Clarke and Julian Savulescu
PA RT I : T H E I D E A O F M O R A L STAT U S
2. Suffering and Moral Status 23
Jeff McMahan
3. An Interest-Based Model of Moral Status 40
David DeGrazia
4. The Moral Status of Conscious Subjects 57
Joshua Shepherd
5. Moral Status, Person-Affectingness, and Parfit’s
No Difference View 74
F. M. Kamm
6. The Ever Conscious View and the Contingency of Moral Status 90
Elizabeth Harman
7. Moral Status and Moral Significance 108
Ingmar Persson
8. Moral Recognition and the Limits of Impartialist Ethics: On
Androids, Sentience, and Personhood 123
Udo Schuklenk
9. Is Moral Status Good for You? 139
Thomas Douglas
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vi Contents
PA RT I I : SP E C I F IC I S SU E S A B O U T M O R A L STAT U S
10. Toward a Theory of Moral Status Inclusive of Nonhuman
Animals: Pig Brains in a Vat, Cows versus Chickens, and
Human–Nonhuman Chimeras 159
Ruth R. Faden, Tom L. Beauchamp, Debra J. H. Mathews, and Alan
Regenberg
11. Revisiting Inexorable Moral Confusion About the Moral Status
of Human–Nonhuman Chimeras 179
Jason Scott Robert and Françoise Baylis
12. Chimeras, Superchimps, and Post-persons: Species Boundaries
and Moral Status Enhancements 197
Sarah Chan
13. Connecting Moral Status to Proper Legal Status 215
Benjamin Sachs
14. How the Moral Community Evolves 231
Rachell Powell, Irina Mikhalevich, and Allen Buchanan
15. Moral Status of Brain Organoids 250
Julian Koplin, Olivia Carter, and Julian Savulescu
16. How Much Moral Status Could Artificial Intelligence
Ever Achieve? 269
Walter Sinnott-Armstrong and Vincent Conitzer
17. Monkeys, Moral Machines, and Persons 290
David R. Lawrence and John Harris
18. Sharing the World with Digital Minds 306
Carl Shulman and Nick Bostrom
Index 327
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Preface
viii Preface
Preface ix
x Preface
challenges, since existing conceptual frameworks, and the criteria for moral
consideration they trigger, are still defensible and applicable.
Regardless of the specific characterization of moral status, an often-
neglected question is whether having it—and possibly having more of it—is
good for you. If it is, does losing it harm you? Rounding off the first part of
the volume, this question of the prudential value of moral status is precisely
what Thomas Douglas tackles in Chapter 9. Answering it is important in
helping us to decide whether or not we should enhance, or disenhance, the
cognitive and moral capacities of non-human animals. Doing either may
affect their moral status.
Part II, ‘Specific Issues about Moral Status’, begins with three chapters
focusing in particular on the prospect of interspecies chimeras. In Chapter 10,
Ruth Faden, Tom Beauchamp, Debra Mathews, and Alan Regenberg argue for
a theory of moral status that helps provide solutions to practical problems in
public policy, taking account of the interests of non-human animals. To illus-
trate this need, their chapter describes two contemporary problems, one in
science policy and one in food and climate policy. They sketch a way to think
about a tiered or hierarchical theory of moral status that could be fit for such
work, and then consider in some depth the problem of human non-human
chimeras.
In Chapter 11, Jason Roberts and Françoise Baylis revisit their earlier work
on the history, ethics, and future of stem cell research involving chimeras
made, in part, from human cells. In particular, they focus on the notion of
inexorable moral confusion: objections to the creation of chimeras are likely
motivated by a strong desire to avoid inexorable moral confusion about these
beings’ moral status. Here, they further specify and elaborate on the original
concept in light of recent scientific and technical developments as well as eth
ical insights. In Chapter 12, Sarah Chan explores the normative and concep-
tual challenges raised by the prospect of crossing both biological and moral
‘species boundaries’, including the implications of species transitions in rela-
tion to identity, obligations toward existing beings, and beings that might be
created via the species transition process. She reflects on how all of this might
advance our thinking about moral status.
In Chapter 13, Ben Sachs considers three proposals regarding the connec-
tion between an animal’s moral status and the legal status it ought to have.
The first proposal is this strong claim: if an act wrongs an animal then crimin
alizing it is justified. The second proposal is more moderate: if an act consti-
tutes an injustice to an animal then criminalizing it is justified. The third
proposal is the one Sachs defends: it is obligatory for legislators to eliminate
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Preface xi
any aspect of the law that facilitates the wronging of animals. The chapter
considers, in particular, the radical implications of this third proposal for ani-
mal ownership and state funding of medical research on animal subjects.
The chapter by Rachell Powell, Irina Mikhalevich, and Allen Buchanan
(Chapter 14) considers several evolved biases that distort our tendency to ascribe
moral status, focusing in particular on the example of invertebrates. These biases
include tendencies to deny moral standing, or to attribute lower moral status to
beings that elicit feelings of disgust or fear, as well as to those that are perceived
as less similar to us, less attractive, less individualized, and less disposed toward
reciprocal cooperation. These adaptive mechanisms may have served human
groups well in the evolutionary past, but in the modern world they pose an
obstacle to moral progress and play a key role in moral regression.
Chapter 15 examines brain organoids, which recapitulate the development
of the brain. Might these have moral status, or the potential for it? Julian
Koplin, Olivia Carter, and Julian Savulescu tackle this question head on. It is
plausible, they argue, that brain organoids could one day attain consciousness
and perhaps even higher cognitive abilities. Research on brain organoids
therefore raises difficult questions about their moral status—questions that
currently fall outside the scope of existing regulations and guidelines. The
chapter offers a novel moral framework for such research and outlines the
conditions under which brain organoids might attain moral status.
The final three chapters focus in particular on AI and the prospect of digi-
tal minds. Walter Sinnott-Armstrong and Vincent Conitzer ask, in Chapter 16,
just how much moral status artificial intelligence could ever achieve. They
suggest that different entities have different degrees of moral status with
respect to different moral reasons in different circumstances for different pur-
poses. Recognizing this variability will help resolve some debates about the
potential moral status of AI.
In Chapter 17 David Lawrence and John Harris argue that debates over
moral machines often make wide assumptions about the nature of future
autonomous entities, and frequently bypass the distinction between ‘agents’
and ‘actors’. The scope and limits of moral status, they suggest, are fundamen-
tally linked to this distinction, and position non-Homo sapiens great apes as
members of a particular moral status clade, which are treated in a similar
fashion to that proposed for so-called ‘moral machines’. They suggest that the
principles by which we ultimately decide how to treat great apes, and whether
or not we decide to act upon our responsibilities to them as moral agents, are
likely to be the same principles we use to determine our responsibilities to
moral AI in the future.
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xii Preface
Finally, Carl Shulman and Nick Bostrom (Chapter 18) conclude the volume
by investigating the moral status of digital minds. The minds of biological
creatures occupy a small corner of a much larger space of possible minds that
could be created. Their chapter focuses on one set of issues which are pro-
voked by the prospect of digital minds with superhumanly strong claims to
resources and influence. These could arise from the vast collective benefits
that mass-produced digital minds might derive from relatively small amounts
of resources. Alternatively, they could arise from individual digital minds
with superhuman moral status or ability to benefit from resources. Such
beings may contribute immense value to the world. Failing to respect their
interests could produce a moral catastrophe, but a naive way of respecting
them could be disastrous for humanity.
Work leading to this volume was supported by the Wellcome Trust under
Grant WT203132/Z/16/Z and the Uehiro Foundation on Ethics and
Education. Most of the chapters in the volume are revised versions of papers
that were originally presented at a conference on ‘Rethinking Moral Status’,
held at St Cross College, in Oxford, on 13 and 14 June 2019, organized by the
Wellcome Centre for Ethics and Humanities and the Oxford Uehiro Centre
for Practical Ethics, both at the University of Oxford. The chapter by Steve
Clarke and Julian Savulescu began life as a background paper, circulated to
participants before the event. We were fortunate enough to be able to add
chapters by Jeff McMahan, by Carl Schulman and Nick Bostrom, by Julian
Koplin, Olivia Carter, and Julian Savulescu, and by Rachell Powell, Irina
Mikhalevich, and Allen Buchanan to the collection.
As well as thanking the Wellcome Trust and Uehiro Foundation for their
generous support, the editorial team would like to thank Daniel Cohen, Alan
Crosier, Rachel Gaminiratne, Christa Henrichs, Guy Kahane, Neil Levy,
Morgan Luck, Mike Parker, Steven Tudor, Suzanne Uniacke, and Miriam
Wood, for helping us, in different ways, to put together the volume. We would
also like to thank Peter Momtchiloff for his expert editorial guidance. We
thank all of our contributors, as well as the various people who helped us with
the volume, for their forbearance during the COVID-19 global pandemic,
which led to the production of the volume taking somewhat longer than
originally anticipated.
Steve Clarke
Hazem Zohny
Julian Savulescu
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Notes on Contributors
Care Ethics and Law and a PhD in Bioethics from the University of Manchester, where
she was a Research Fellow in Bioethics from 2005 to 2015.
Steve Clarke is a Senior Research Fellow in Ethics and Humanities, in the Wellcome
Centre for Ethics and Humanities, the Uehiro Centre for Practical Ethics, and the
Faculty of Philosophy at the University of Oxford. He is also Associate Professor of
Philosophy in the School of Humanities and Social Sciences at Charles Sturt
University. He has broad research interests in philosophy and bioethics.
Vincent Conitzer is the Kimberly J. Jenkins University Professor of New Technologies
and Professor of Computer Science, Professor of Economics, and Professor of
Philosophy at Duke University. He is also the Head of Technical AI Engagement at the
Institute for Ethics in AI and Professor of Computer Science and Philosophy at the
University of Oxford. He received his PhD (2006) and MS (2003) degrees in Computer
Science from Carnegie Mellon University, and an AB (2001) degree in Applied
Mathematics from Harvard University. Conitzer works on artificial intelligence (AI).
More recently, he has started to work on AI and ethics.
David DeGrazia is Elton Professor of Philosophy at George Washington University.
DeGrazia’s nine books include Taking Animals Seriously: Mental Life and Moral
Status (Cambridge University Press, 1996) and, with Tom Beauchamp, Principles of
Animal Research Ethics (Oxford University Press, 2020)
Thomas Douglas is Professor of Applied Philosophy and Director of Research and
Development in the Oxford Uehiro Centre of Practical Ethics, University of Oxford.
He is also Senior Research Fellow at Jesus College, Oxford, and Editor of the Journal
of Practical Ethics, and Principal Investigator on the project ‘Protecting Minds’, funded
by the European Research Council. He trained in clinical medicine and philosophy
and works chiefly on the ethics of behaviour modification and neuroenhancement.
Ruth R. Faden is the founder of the Johns Hopkins Berman Institute of Bioethics. She
was the Berman Institute’s Director from 1995 until 2016, and the inaugural
Andreas C. Dracopoulos Director (2014–16). Dr Faden is the inaugural Philip Franklin
Wagley Professor of Biomedical Ethics. In the twenty years in which Dr Faden led the
Berman Institute, she transformed what was an informal interest group of faculty
across Johns Hopkins into one of the world’s premier bioethics programmes.
Elizabeth Harman is Laurance S. Rockefeller Professor of Philosophy and Human
Values at Princeton University. Her publications include ‘Creation Ethics’ (Philosophy
and Public Affairs), ‘“I’ll Be Glad I Did It” Reasoning and the Significance of Future
Desires’ (Philosophical Perspectives), ‘The Irrelevance of Moral Uncertainty’ (Oxford
Studies in Metaethics), ‘Morally Permissible Moral Mistakes’ (Ethics), and ‘Ethics is
Hard! What Follows?’ (forthcoming). She is co- editor of Norton Introduction to
Philosophy, Second Edition (2018), and Norton Introduction to Ethics (forthcoming).
John Harris is Professor Emeritus, University of Manchester, Visiting Professor
in Bioethics, Department of Global Health and Social Medicine, Kings
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Notes on Contributors xv
Ethics. His main publications are: The Retreat of Reason (OUP, 2005), From Morality
to the End of Reason (OUP, 2013), Inclusive Ethics (OUP, 2017), Reasons in Action
(OUP, 2019), Morality from Compassion (OUP, 2021), and with Julian Savulescu, Unfit
for the Future (OUP, 2012).
Rachell Powell is Associate Professor of Philosophy at Boston University. Her research
focuses on the philosophy of biological and biomedical science. Her books include
Contingency and Convergence: Toward a Cosmic Biology of Body and Mind (MIT
2019), and The Evolution of Moral Progress: A Biocultural Theory (OUP 2018, with
Allen Buchanan). She has published in such journals as Philosophy of Science, British
Journal for the Philosophy of Science, Journal of Philosophy, Ethics, and Journal of
Medicine and Philosophy.
Alan Regenberg is the Director of Outreach and Research Support and an associate
faculty member at the Johns Hopkins Berman Institute of Bioethics. He is engaged in
a broad range of research projects and programs, including the Berman Institute’s sci-
ence programs: The Stem Cell Policy and Ethics Program (SCOPE); the Program in
Ethics and Brain Sciences; and the Hinxton Group, an international consortium on
stem cells, ethics, and law.
Jason Scott Robert holds the Lincoln Chair in Ethics and a Dean’s Distinguished
(Associate) Professorship in the Life Sciences at Arizona State University. His work is
at the nexus of philosophy of biology and bioethics, focusing primarily on the justifi-
cation of good science in controversial areas of research in developmental biology and
the neurosciences.
Benjamin Sachs is a Senior Lecturer in Philosophy at the University of St Andrews.
His main interests are in applied ethics, coercion, political philosophy, and philoso-
phy of law. His first book, Explaining Right and Wrong, was published by Routledge in
2018. His second book, Contractarianism, Role Obligations, and Political Morality, is
forthcoming with Routledge. He is currently co-directing, with Alex Douglas, a
research network called The Future of Work and Income.
Julian Savulescu is Uehiro Chair in Practical Ethics, Director, Oxford Uehiro Centre
for Practical Ethics, and Co-Director, Wellcome Centre for Ethics and Humanities at
the University of Oxford. He is Visiting Professor in Biomedical Ethics, Murdoch
Children’s Research Institute, where he directs the Biomedical Ethics Research Group,
and Distinguished Visiting Professor in Law, Melbourne University.
Udo Schuklenk has taught at universities in Germany, Australia, the UK, and South
Africa before taking up the Ontario Research Chair in Bioethics at Queen’s University.
He has written or co-edited ten books and authored or co-authored more than 150
peer reviewed publications in journals and anthologies. His main research interests
are in the areas of end-of-life issues, public health, and medical professionalism.
Joshua Shepherd is Assistant Professor in Philosophy at Carleton University, Research
Professor at the Universitat de Barcelona (where he directs the ERC funded project
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Rethinking Conscious Agency), and Senior Fellow of the LOGOS Research Group.
He is the author of the book Consciousness and Moral Status.
Carl Shulman is a Research Associate at the Future of Humanity Institute, Oxford
Martin School, Oxford University, where his work focuses on the long-run impacts of
artificial intelligence and biotechnology. Previously, he was a Research Fellow at the
Machine Intelligence Research Institute. He attended New York University School of
Law and holds a degree in philosophy from Harvard University.
Walter Sinnott-Armstrong is Chauncey Stillman Professor of Practical Ethics at
Duke University in the Philosophy Department, the Kenan Institute for Ethics, the
Duke Institute for Brain Science, and the Law School. He publishes widely in ethics,
moral psychology and neuroscience, philosophy of law, epistemology, philosophy of
religion, and argument analysis.
Hazem Zohny is Research Fellow in Bioethics and Bioprediction at the Uehiro Centre
for Practical Ethics at Oxford University. He has a PhD in Bioethics from the
University of Otago and has published a number of academic papers related to
enhancement, disability, well-being. His current work focuses on the bioprediction of
behaviour and the ethics of using neurointerventions for crime prevention.
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1
Rethinking our Assumptions about
Moral Status
Steve Clarke and Julian Savulescu
When a being or entity has moral status its interests matter morally, for its
own sake (Jaworska and Tannenbaum 2018). If a being or entity has moral
status, then an act that is morally bad, in at least one respect, is committed
when an agent harms that being or entity. Any all-things-considered moral
justification for such an act must take into account the harm committed by
the agent against that being or entity. Ordinary adult humans are usually sup-
posed to have a specific and equal level of moral status—often referred to as
‘full moral status’ (FMS). Non-human animals are usually accorded some
moral status, but this is typically understood to be a lesser level or degree of
moral status than FMS.1
Statuses are often organized in hierarchies. In the peerage of Great Britain,
for example, an Earl has higher status than a Viscount, a Viscount ranks
higher than a Baron, and a Baron is the lowest-status British peer, ranking
only above commoners. Standard attributions of moral status form a partial
hierarchy. It is usually agreed that humans have a higher level of moral status
than non-human animals. However, there is no widely accepted ordering of
non-human animal moral status.2 Opinions vary about the relative levels of
moral status of different non-human animals, and about which species of ani-
mals have moral status. Most of us ascribe some moral status to non-human
primates. Many of us ascribe some moral status to other mammals. Some of
us ascribe some moral status to birds, reptiles, and fish and a few of us ascribe
some moral status to arachnids, insects, and crustaceans.3 Further disagree-
ment about the presence of moral status, or about the extent to which it is
possessed, becomes apparent when we consider humans other than ordinary
Steve Clarke and Julian Savulescu, Rethinking our Assumptions about Moral Status In: Rethinking Moral Status. Edited by:
Steve Clarke, Hazem Zohny, and Julian Savulescu, Oxford University Press. © Steve Clarke and Julian Savulescu 2021.
DOI: 10.1093/oso/9780192894076.003.0001
OUP CORRECTED AUTOPAGE PROOFS – FINAL, 19/06/21, SPi
adult humans. Do human foetuses and embryos have FMS, some lesser moral
status, or no moral status? What about infants? What about severely cogni-
tively impaired or unconscious adults?
Technological developments are throwing up new and controversial cases,
which will require our consideration. What are we to say about human non-
human chimeras,4 human brain organoids,5 or artificial intelligence?6 What
should we say about the moral status of a cyborg,7 a post-human,8 or a human
mind that has been uploaded into a computer, or onto the internet?9 To pro-
vide sensible answers to these questions we need to be able to think clearly
about what it is to have moral status, and about when and why we should
attribute moral status to beings and entities.
One way to help clarify our thinking is to try to define moral status.
However, when we attempt this, it can start to look like talk of moral status
doesn’t add anything to other, more familiar forms of moral discourse.
DeGrazia offers the following characterization of moral status:
To say that X has moral status is to say that (1) moral agents have obligations
regarding X, (2) X has interests, and (3) the obligations are based (at least
partly) on X’s interests. (DeGrazia 2008, p. 183)
We are already familiar with the language of interests and obligations, so why
not restrict ourselves to this terminology and forgo talk of moral status? An
answer to this question, defended by DeGrazia, is that reference to moral sta-
tus is a convenient form of shorthand, which is especially useful to us when
we want to generalize about moral obligations and interests (2008, p. 184).
Another answer is that moral status talk is well suited to play a specific
explanatory role that talk of moral interests and obligations is not well suited
to play. This is to relate the moral properties of beings to whom we have moral
obligations to the non-moral properties and capacities of those beings.10
If we are pushed to rethink our assumptions about moral status to accom-
modate artificial intelligence, cyborgs, human brain organoids, human non-
human chimeras, post- humans, and uploaded minds, then we should
consider the possibility that some of these beings and entities have a level of
moral status below FMS. We should also be open to the possibility that some
of these beings and entities might have a higher moral status than do ordinary
adult humans. The phrase ‘full moral status’ (FMS) suggests a threshold level
above which moral status cannot rise. However, as we will go on to discuss, it
seems possible that a being or entity could have a higher moral status than the
moral status of ordinary adult humans.
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In this chapter we consider some of the key philosophical issues that arise
when attempts are made to rethink our usual assumptions about moral status
to handle such new and controversial cases. In section 2 we critically examine
the widespread assumption that all ordinary adult humans have equal moral
status. In section 3 we subject to scrutiny the assumption that membership of
the species Homo sapiens somehow confers FMS. In section 4 we consider
some revisionary approaches to thinking about moral status that involve
rejecting the presupposition that there is a sharp distinction between the FMS
of ordinary adult humans and the partial moral status of non-human animals.
In section 5 we consider proposals to reject an almost universally accepted
assumption—that no beings or entities could have higher moral status than
the FMS usually attributed to ordinary adult humans. We also consider some
consequences that could follow from creating beings with higher moral status
than that of ordinary adult humans. In section 6 we turn our attention to a
practical concern: how to behave toward beings and entities when we find
ourselves uncertain about their moral status.
The assumption that all adult humans who are not severely cognitively
impaired have equal moral status is hardly ever challenged these days, at least
in Western liberal societies.11 It is a background assumption made by the
many of us who share liberal, democratic ideals. However, it would have been
rejected by most ordinary members of the various slave-owning societies that
flourished before the rise of modern liberal democracy.12 In slave-owning
societies, the enslaved were regarded as having fewer legal rights than the
free. The systematic difference between the expansive legal rights of the free
and the limited rights of the enslaved was provided with apparent justification
by the pervasive assumption made in many slave-owning societies that the
enslaved were of a lesser moral status than the free.13
Defenders of institutional slavery usually sought to justify the attribution
of different levels of moral status to different groups of people by appealing to
perceived natural differences between different types of humans. These differ-
ences were then invoked to try to justify the enslavement of humans of one
type by humans of another type. The best-known attempted philosophical
defence of slavery is from Aristotle, who argued that some humans lacked the
capacities for significant deliberation and foresight, and so were ‘natural
slaves’, in need of direction by natural masters who possessed the capacities
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that natural slaves lacked.14 Aristotle’s theory of natural slavery was based on
an assumption of systematic underlying differences between different types of
humans. However, unlike more recent, eighteenth- and nineteenth-century
defenders of slavery, Aristotle did not assume that these differences correlated
with racial differences. This should not be surprising. In Ancient Greece,
slaves were captured and traded from many different countries and had a
diverse range of ethnic origins. By the eighteenth and nineteenth centuries,
slavery in the West was, for the most part, restricted to specific ethnicities,
with blacks especially liable to be enslaved by whites. Eighteenth- and
nineteenth-century apologists for slavery often appealed to quasi-scientific
theories about racial differences, which lacked supporting evidence, in their
attempts to justify race-based slavery.15
The fact that institutional slavery was still practised in the US (and else-
where), as recently as 160 years ago is very disturbing to defenders of the view
that all adult humans who are not severely cognitively impaired have equal
moral status, including the authors of this chapter. It seems clear to us that
societies that fail to treat all ordinary adult humans as having equal moral
status ought to do so, because in fact all ordinary adult humans have equal
moral status. If all ordinary adult humans have equal moral status, as we are
convinced that they do, then this is presumably because of some or other
properties and capacities that they share, which may or may not be shared by
human infants, human foetuses and embryos, and severely cognitively
impaired humans. What properties or capacities might these be?
Almost all attempts to locate grounds for the moral status of ordinary adult
humans identify specific cognitive capacities as the basis for that moral status.
However, there is a lack of agreement in the literature regarding the cognitive
capacities necessary for FMS. Quinn suggests that the ability to will is neces-
sary for FMS (1984, p. 51), while Singer stresses the importance of future-
oriented planning (1993, pp. 116–17), McMahan suggests that self-awareness
is necessary for FMS (2002, p. 45). Baker (2000) argues that self-consciousness
is necessary,16 Metz (2012) suggests that the capacity to participate in com-
munal relationships is necessary,17 and Jaworska (2007) stresses the import
ance for FMS of having a capacity to care.
A different approach to grounding attributions of FMS is to argue that the
potential to go on to develop sophisticated cognitive capacities warrants the
attribution of FMS. Infants and severely cognitively impaired adults do not
possess sophisticated cognitive capacities, but many possess the potential to
develop—or recover—sophisticated cognitive capacities. Appeals to potential,
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as a basis for the attribution of FMS, are popular among opponents of abor-
tion and unpopular among proponents of abortion. Just as infants have the
potential to acquire sophisticated cognitive capacities, so do human foetuses
and embryos. If we are to grant FMS to human foetuses and embryos, then it
looks like we should ban most instances of abortion, as abortion will involve
killing beings who are acknowledged to have FMS.18
A major concern with appeals to potential as a basis for attributions of FMS
is that it is far from obvious what constraints there are on such appeals. An
unfertilized human ovum together with a human sperm have the potential to
become a human adult. However, anti-abortion activists do not usually want
to argue that unfertilized ovum-sperm pairs have FMS, in virtue of having the
potential to become adult humans. In response to objections along these
lines, opponents of abortion, such as Watt (1996) and Camosy (2008), draw
conceptual distinctions between the type of potential the unfertilized ovum-
sperm pair has and the potential that a fertilized ovum has to become a
human adult. It is not clear that such attempts to distinguish between differ-
ent types of potential are successful. Nor is it entirely apparent how any spe-
cific type of potential could confer moral status.19
A common way of supplementing accounts of the grounds necessary for
FMS is to assert that personhood is necessary and sufficient for FMS.20
Persons are said to have FMS, while non-persons are said to have either less-
than-full or no moral status.21 There is significant disagreement in the litera-
ture about which beings and entities are persons. Human foetuses are not
usually regarded as persons, at least by secular philosophers, but they are
considered to be legal persons in some jurisdictions.22 Human infants are
usually regarded as persons, but some scholars, such as Tooley (1972), argue
otherwise. Cognitively impaired human adults are usually regarded as per-
sons; however, it has been argued that once humans have become severely
cognitively impaired and have fallen into a persistent non-responsive state
they may no longer be persons (Callahan 1993). Non-human apes may be
persons, or at least ‘border-line persons’ (DeGrazia 2007, p. 323). Now-extinct
Neanderthals may have been persons (Buchanan 2009, p. 372), and some
intelligent machines that we might create in the future could be persons
(Bostrom and Yudkowsky 2014). It is not easy to see how the stipulation that
personhood is a criterion for FMS could assist us in identifying who and what
has FMS. If we treat personhood as a criterion for FMS then we transform the
problem of identifying who and what has FMS into the equally challenging
problem of figuring out who and what is a person.
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species membership. The same can be said of hybrid animals, such as mules
and ligers, as well as chimeras created by blending genetic material from more
than one species, such as sheep–goat chimeras. Artificial intelligence also
raises problems for species-membership accounts of moral status. In the
future, we may be able to create beings and entities with very sophisticated
cognitive capacities and intuitively it seems that there is a compelling case for
attributing FMS to at least some of them. But they will not be members of
species. Acceptance of species membership as a necessary condition for attri-
butions of moral status would preclude us from attributing moral status to
these beings and entities.
of interests’, and apply it to both humans and non-human animals. The most
important interests that beings have, according to Singer, are interests in
enjoyment and the avoidance of suffering. Some non-human animals will
have a greater capacity for enjoyment and a greater capacity for suffering
than do some cognitively impaired humans. So, application of the principle
of equal consideration of interests will lead us to prioritize the interests of
these animals over the interests of severely cognitively impaired humans
(Singer 2009, pp. 574–6).
Another revisionary view is due to Jeff McMahan. McMahan (2002) raises
the possibility of ‘intermediate moral status’. This a level of moral status some-
where between that of most non-human animals and FMS. McMahan (2002)
suggests that we should attribute intermediate moral status to human infants
and cognitively advanced non-human animals, such as ‘higher primates’
(2002, p. 265). While McMahan (2002) allowed for three distinct levels of
moral status, McMahan (2008, pp. 97–100) contemplates two levels of moral
status with a rising series of degrees of moral status between the two levels.
These rises take place in response to increases in ‘psychological capacity’ up
to a threshold of FMS (McMahan 2008, p. 99). On McMahan’s later view,
intermediate moral status still exists, but some beings with intermediate
moral status have higher moral status than others.30 Another possibility is
that moral status comes in degrees and rises consistently, before levelling off
at the threshold where personhood is attained (DeGrazia 2008).31 There are
also many other possible combinations of levels, or thresholds, of moral status
and continuous rises by degrees that we could invoke to depict the relation-
ship between increases in possession of the relevant properties and capacities
we take to underpin moral status and increases in moral status.32
The phrase ‘Full Moral Status’ suggests that there is a maximum level of moral
status that might be obtained. However, it seems possible that there could be
beings with higher moral status than the full moral status normally attributed
to ordinary adult humans. It may be difficult for us to conceive of such beings,
but it does not follow from the limitations of our imaginative capacities that
they cannot exist.33 If we think that humans have a higher level of moral sta-
tus than non-human animals, in virtue of possessing cognitive capacities that
are superior to those of non-human animals, then it looks like we should be
open to the possibility that beings with superior cognitive capacities to ours
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would have a higher level of moral status than us. Transhumanists, such as
Bostrom (2005), urge us to try to create ‘post-humans’ with superior cogni-
tive capacities to our own. If we manage to do so then we may also end up
creating beings with higher moral status than we possess (Agar 2013a;
Douglas 2013).
Suppose we could create beings with superior moral status to ours. Should
we do so? There are reasons that speak in favour of creating such beings. If
these beings have higher moral status than us, in virtue of having more devel-
oped cognitive capacities than we have, then, all things being equal, they will
be more capable of accurate and consistent moral reasoning than we are. If
they are more capable of accurate and consistent moral reasoning than us
then, all else being equal, they will be more likely to perform good acts and
less likely to perform bad acts than we are. From an impartial point of view, it
therefore seems that we should prefer the creation of post-humans with
superior moral status to the creation of mere humans. It is plausible to think
that the creation of post-humans will also be good for humans. All things
being equal, post-humans with more highly developed moral capacities than
humans possess will be more likely than humans to treat other beings, includ-
ing humans, in morally appropriate ways.
However, there are reasons to be concerned about the consequences for us
of creating post-humans with higher moral status than we possess. From the
point of view of post-humans with higher moral status than us, we humans
would be beings of lower moral status and morality could permit post-
humans to treat us in ways we would prefer not to be treated. To understand
this concern, it helps to start by thinking about the ways in which we regard it
as morally permissible to treat the non-human animals that we consider to
have lower moral status than us. Many of us regard it as morally permissible
to kill and eat non-human animals, sacrificing their lives for our nutrition
and our gustatory pleasure. Many also regard it as morally permissible to con-
duct harmful experiments on non-human animals if doing so leads to med
ical or cosmetic benefits to humans. Also, we generally regard it as morally
permissible, if not obligatory, to sacrifice non-human animals when human
lives are at stake.
Consider a ‘trolley case’ in which a runaway trolley is going to run over a
human unless we flick a switch, diverting the trolley down a side-track and
thereby killing five sheep that are stuck on the side-track. Most of us would
regard it as morally permissible, if not obligatory, to flick the switch and sacri-
fice the lives of several sheep to save one human life. If we are right about it
being morally acceptable for us to treat beings with lower moral status than
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ourselves, in the various ways that have been listed, then we should worry
about the ways in which beings with higher levels of moral status might
regard themselves as being entitled to treat us. By parity of reasoning, we can
infer that they may well regard themselves as being entitled to kill and eat us,
to conduct harmful experiments on humans, and to sacrifice the lives of many
of us in order to save just one of them.34
Agar (2013a) argues that we are not under a moral obligation to create
‘post-persons’—his preferred term for post-humans with higher moral status
than ourselves. He further argues that when we think about the potential
harms to us that may result from the creation of such beings, it becomes clear
that we have good reason not to do so. Persson (2013) offers a contrary view.
He argues that Agar is biased in favour of mere humans and against post-
persons. On Persson’s (2013) view, while we are not under a moral obligation
to create post-persons, it would be good, all things being equal, for us to cre-
ate post-persons. As post-persons will only be treating humans as morality
requires them to treat humans when they sacrifice human lives for the sake of
their own lives, there can be no moral objection to them sacrificing humans
for the sake of their own lives. Agar (2013b) disputes that he is merely biased
in favour of humans and against post-persons. His ‘Rawlsian’ conception of
justice leads him to put the interests of the worse off ahead of those of the
better off, and in a world in which mere humans and post-persons both exist
humans can reasonably be expected to be worse off and post-humans the better
off. To avoid this state of affairs it is better for humans not to create post-humans
in the first place, or so Agar (2013b) argues.
one of them but not both. If we knew that the post-human was of higher
moral status than the human then, all else being equal, we would be morally
obliged to save the post-human and allow the human to die.35 However, all we
know is that the post-human might have higher moral status than the human
or might have the same moral status as the human.
We know that, all things being equal, it is wrong to rescue a being with
lesser moral status if doing so involves allowing a being with higher moral
status to die. So, we know that we might be acting wrongfully by rescuing the
human rather than the post-human. However, all else being equal, there is no
chance that we will act wrongfully if we rescue the post-human and allow the
human to die. Even if they both have the same moral status, it is not wrong to
rescue the post-human and allow the human to die. We might be acting
wrongfully if we rescue the human, but all else being equal, we cannot be act-
ing wrongfully if we rescue the post-human. Therefore, it seems clear that we
ought to rescue the post-human and allow the human to die, even though we
are uncertain as to whether the post-human has superior moral status to
the human.
Issues of uncertainty about moral status don’t only arise when we think
about post-humans. They also arise when we think about human non-human
chimeras. Most of us regard it as clear that ordinary adult humans have higher
moral status than non-human animals. However, beings that are part human
and part non-human animal pose a threat to our intuitive sense of moral
clarity. Admittedly, current examples of human non-human chimeras do not
seem to present much of a challenge to our intuitive sense of moral clarity. An
ordinary adult human who has a pig valve transplant in her heart is technic
ally a chimera, but she would be regarded by the vast majority of us as having
the moral status of any other ordinary adult human. Similarly, most would
consider Oncomice (mice that are genetically engineered to contain a human
cancer-causing gene) as having the moral status of ordinary mice (Bok 2003).
In the future, however, we may be able to create chimeras that involve a more
extensive blending of human and non-human animal components and we
may find ourselves unable to determine the moral status of these more exten-
sively blended beings.
If we are uncertain about the moral status of a particular type of being and
are unable to figure out how to go about alleviating our uncertainty then we
are in a state of moral confusion. Robert and Baylis argue that the future cre
ation of human non-human chimeras threatens to place us in a state of moral
confusion and that this is an important reason for us to avoid creating such
beings (2003, p. 9). Whether or not one is put off creating human non-human
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Notes
1. A few scholars appear to proceed on the assumption that there is no level of moral sta-
tus other than FMS. For discussion of such views, see Hursthouse (2013, pp. 3425–7).
2. For a recent proposal to develop a hierarchy of non-human animal moral status, see
Kagan (2018).
3. A small number of us are inclined to attribute moral status to plants and perhaps eco-
systems. See, for example, Goodpaster (1978).
4. A human non-human chimera is a being created by combining cells that have human
origins with cells that have non-human origins. For discussion of the moral status of
human non-human chimeras, see Robert and Baylis (2003), Streiffer (2005), DeGrazia
(2007) and Hübner (2018).
5. Human brain organoids are artificially grown miniature organs resembling human
brains. They are created by culturing human pluripotent stem cells, and are used as rel-
atively simple in vitro models to assist in neurological experimentation. For discussion
of the moral status of human brain organoids, see Cheshire (2014) and Lavazza and
Massimini (2018).
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6. For discussion of the moral status of artificial intelligence, see Basl (2014) and
Søraker (2014).
7. For discussion of the moral status of cyborgs, see Gillett (2006) and Jotterand (2010).
8. Post-humans are humans who have radically enhanced capacities: for example, an IQ
above 200 or the power of telepathy or telekinesis. For discussion of the moral status
of post-humans, see Buchanan (2009), Agar (2013a; 2013b), and Douglas (2013).
9. For discussion of the moral status of uploaded minds, see Sandberg (2014).
10. Some authors worry that the apparent convenience and utility of talk of moral status
comes at a price and suggest that we should not pay that price. Sachs (2011) argues
that talk of moral status can be obfuscating. Horta argues that moral status talk can
distort our understanding of how we should behave toward some individuals in par-
ticular circumstances (2017, p. 909).
11. However, it is subject to the occasional sceptical challenge. For discussion of how
defenders of the assumption might try to respond to sceptical challenges, see
McMahan (2008).
12. There was a long delay between the initial rise of modern liberal democracy and the
end of institutional slavery. The US Declaration of Independence was signed in 1776,
and is surely one of the foundational documents of modern liberal society. It made the
unqualified assertion that it is self-evident that all men are created equal. However, it
would take 89 years and a hugely destructive civil war before institutional slavery was
abolished in all parts of the US.
13. For discussion of the challenge that slave-owning societies present, for those who
assume that humans have equal moral status, see Lindsay (2005).
14. Aristotle’s theory of natural slavery is much more complicated than this brief charac-
terization suggests. For further discussion, see Smith (1983).
15. An influential approach was to appeal to the now discredited theory of polygenesis,
which holds that different human races evolved separately, in different parts of the
world. The purportedly separate origins of different races was then appealed to, in an
attempt to explain why some races were better suited to freedom and others better
suited to slavery. For discussion of polygenetic defences of slavery amongst
eighteenth-century philosophers, see Watkins (2017). For a mid-nineteenth-century
polygenetic defence of slavery, see Nott and Gliddon (1854).
16. Shepherd (2017: 2018) has recently argued for the moral insignificance of
self-consciousness.
17. Metz’s (2012) appeal to the capacity to participate in communal relationships as
necessary for FMS is one component of a relational account of moral status he defends.
18. Not all arguments for abortion rely on appeals to the moral status of foetuses and
embryos. A famous argument for abortion that does not is due to Thomson (1971).
19. For arguments against the relevance of appeals to potential for moral status, see
McMahan (2002, pp. 308–29), Persson (2003), and Singer and Dawson (1988).
20. There are different senses of personhood discussed in the philosophical literature,
including metaphysical and legal personhood. The sense that is most relevant to dis-
cussions of moral status is ‘moral personhood’.
21. See, for example, Singer (1993), Baker (2000), Warren (1997, Chapter Four) and
McMahan (2002).
22. For further discussion, see Schroedel (2000).
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23. Those who take membership of our species as necessary for FMS are more likely to
employ theological rather than philosophical arguments to support their case. One
influential Christian theological argument for the conclusion that humanity is neces-
sary for FMS appeals to the biblical doctrine that humans are the only beings made in
the image of God. See Genesis 1:27.
24. For extended discussion of this line of reasoning, see Singer (2009, pp. 568–70).
25. For discussion of various ways to define species, see Ereshefsky (2017). See also
Robert and Baylis (2003, pp. 2–4).
26. The phenomenon of ring species makes further trouble for appeals to species member-
ship as a necessary condition for FMS. For discussion, see Persson and Savulescu (2010).
27. For discussion of the ‘species neutrality requirement’, see Liao (2010, pp. 159–63).
28. Metz reasons similarly (2012, pp. 399–400).
29. Regan doesn’t say exactly where the line is to be drawn between beings that are and
are not subjects-of-a-life. He does, however, mention that ‘mentally normal mammals
of a year or more’ are above that line (2004, p. xvi).
30. For discussion of the development of McMahan’s views between 2002 and 2008, see
Ebert (2018, pp. 84–8).
31. DeGrazia describes this position approvingly but does not explicitly endorse it. He
speculates that moral status may vary according to the extent of a being’s interests and
these may depend on ‘cognitive, affective and social complexity’ (2008, p. 192).
32. Douglas depicts six different possible relationships between moral status and the
underlying property of mental capacity (2003, p. 478).
33. Buchanan allows that it is possible for beings with higher moral status than humans
possess to exist. He thinks, however, that our inability to conceive of such beings
means that any argument that might be mounted to try to persuade us that particular
beings actually have higher moral status than humans would fail to be convincing to
us (2009, p. 363).
34. If they were to reason this way then they would be treating moral status as having
relative value. An alternative possibility is that they might regard human moral status
as an absolute value and then reason that their relatively superior moral status should
not be relevant to the moral permissibility of treating humans in particular ways.
35. Note, though, that if reasons can be agent-centred or if some forms of partiality are
justified, then it may be morally permissible to give priority to members of one’s own
group, even if they have lower moral status (Savulescu 1998).
36. Thanks to Alan Crosier, Katrien Devolder, Tom Douglas, John-Stewart Gordon and
Suzanne Uniacke for helpful comments on earlier versions of this chapter.
References
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Doing so is Wrong?’, Journal of Medical Ethics, 39, 2, 67–74.
Agar, Nicholas (2013b). ‘Still Afraid of Needy Post-Persons’, Journal of Medical
Ethics, 39, 2, 81–3.
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Quinn, Warren (1984). ‘Abortion: Identity and Loss’, Philosophy and Public Affairs,
13, 1, 24–54.
Regan, Tom (2004). The Case for Animal Rights. Berkeley: The University of
California Press.
Robert, Jason Scott and Baylis, Françoise (2003). ‘Crossing Species Boundaries’,
American Journal of Bioethics, 3, 3, 1–13.
Sachs, Benjamin (2011). ‘The Status of Moral Status’. Pacific Philosophical
Quarterly, 92, 87–104.
Sandberg, Anders (2014). ‘Ethics of Brain Emulations’, Journal of Experimental
and Theoretical Artificial Intelligence, 26, 3, 439–57.
Savulescu, Julian. (1998). ‘The Present-aim Theory: A Submaximizing Theory of
Rationality?’ Australasian Journal of Philosophy, 76, 229–43.
Savulescu, Julian (2009). ‘The Human Prejudice and the Moral Status of Enhanced
Beings: What Do We Owe the Gods?’ In Human Enhancement, edited by
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Schroedel, Jean Reith (2000). Is the Fetus a Person? A Comparison of Policies
Across the Fifty States. Ithaca, NY: Cornell University Press.
Shea, Matthew (2018). ‘Human Nature and Moral Status in Bioethics’, Journal of
Medicine and Philosophy, 43, 115–31.
Shepherd, Joshua (2017). ‘The Moral Insignificance of Self Consciousness’,
European Journal of Philosophy, 25, 2, 298-415.
Shepherd, Joshua (2018). Consciousness and Moral Status. London: Routledge.
Singer, Peter (1993). Practical Ethics. Cambridge: Cambridge University Press,
2nd Edition.
Singer, Peter (2009). ‘Speciesism and Moral Status’, Metaphilosophy 40,
3–4, 567–81.
Singer, Peter and Dawson, Karen (1988). ‘IVF Technology and the Argument
from Potential’, Philosophy and Public Affairs, 17, 2, 87–104.
Smith, Nicholas D. (1983). ‘Aristotle’s Theory of Natural Slavery’, Phoenix 37,
2, 109–22.
Søraker, Johnny Hart (2014). ‘Continuities and Discontinuities Between Humans,
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Streiffer, Robert (2005). ‘At the Edge of Humanity: Human Stem Cells, Chimeras
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Thomson, Judith Jarvis (1971). ‘A Defense of Abortion’. Philosophy and Public
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Warren, Mary Anne (1997). Moral Status: Obligations to Persons and Other Living
Things. New York: Oxford University Press.
Watkins, Margaret (2017). ‘ “Slaves among Us”: The Climate and Character of
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PART I
T HE IDE A OF MOR A L STAT U S
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2
Suffering and Moral Status
Jeff McMahan
1. Introduction
Jeff McMahan, Suffering and Moral Status In: Rethinking Moral Status. Edited by: Steve Clarke, Hazem Zohny, and
Julian Savulescu, Oxford University Press. © Jeff McMahan 2021. DOI: 10.1093/oso/9780192894076.003.0002
OUP CORRECTED AUTOPAGE PROOFS – FINAL, 19/06/21, SPi
24 Jeff M c Mahan
2. Unconnected Individuals
replaceable is just the claim that it makes no difference, or does not matter,
which of these two possibilities occurs.
An individual that is replaceable in this sense does not itself matter. It mat
ters, if at all, only insofar as it provides the physical basis for states of con
sciousness that may be intrinsically good or bad. Assuming that an
unconnected individual’s pleasurable and painful states of consciousness are
entirely constitutive of its well-being from moment to moment, it seems that
its well-being matters even though the individual does not in itself matter.
Some philosophers have argued, however, that an individual’s well-being mat
ters only because, and perhaps only to the extent that, the individual itself
matters. According to this view, if I am right that an unconnected individual
does not matter, it follows that its states of consciousness do not matter. It
does not matter, for example, whether its states of consciousness are pleasur
able or painful.
This cannot be right. Whether an unconnected individual’s experience is of
physical pleasure or physical pain—pain that is experienced as aversive—does
matter. If its experience is pleasurable, that is good in itself and arguably good
for the individual; and if its experience is painful, that is bad in itself and
arguably bad for the individual.
Does an individual that does not itself matter but whose states of con
sciousness do matter have moral status? That it does not matter in itself sug
gests that it does not have moral status. But that the character of its states of
consciousness does matter, and matter morally, suggests that it does have
moral status. There is, however, no substantive issue here, only a matter of
finding terms to draw these distinctions. It seems that while an unconnected
individual is experiencing suffering, it is bad for it to be in that state. There is
a moral reason to stop the suffering for the individual’s own sake. In this
respect, an unconnected individual is different from a plant, for there can be
no reason to do anything for the sake of a plant, even if there is a sense in
which plants can, as some philosophers claim, be benefited or harmed (for
example, by the provision or withholding of water or sunlight). Plants lack the
capacity for consciousness and therefore can have neither well-being nor ill-
being. But, like a plant, an unconnected individual does not matter in itself.
I suggest that we appropriate two terms from the philosophical literature
and assign them the following meanings. We can say that because there can
be reasons to act in certain ways for an unconnected individual’s own sake, it
has moral standing, which plants and all other non-conscious entities lack.
But because unconnected individuals are replaceable and thus do not matter
in themselves, but matter only as the experiencers of states of consciousness
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26 Jeff M c Mahan
that matter, they do not have moral status. I propose, in other words, to use
the term ‘moral status’ in such a way that all, but only, those individuals who
themselves matter for their own sake have it. Whereas plants lack both moral
standing and moral status, unconnected individuals have moral standing but
lack moral status. (If, as seems likely, unconnected individuals are the only
individuals that have moral standing but lack moral status, the notion of
moral standing as I understand it is of limited significance.)
Part of the explanation of why an unconnected individual is replaceable is
that it is not harmed by ceasing to exist, or benefited by being enabled to con
tinue to live. Because of its lack of any psychological connections to itself in
the future, it has no interest in continuing to live (in the sense of ‘having no
personal stake in’, rather than ‘not being interested in’). More precisely, it has
no ‘time-relative interest’ in continuing to live which is a function not only of
the magnitude of a benefit or harm an individual might receive but also of the
degree to which the individual that has the interest would be psychologically
related to itself at the time at which the benefit or harm would occur.1 An
unconnected individual’s continuing to live is no different from a different
unconnected individual’s coming into existence. Killing an unconnected indi
vidual is thus relevantly like preventing an unconnected individual from
coming into existence.
All this is consistent with the claim that unconnected individuals lack
moral status. But the fact that an unconnected individual would not be
harmed by being painlessly killed does not by itself entail that the individual
lacks moral status and thus would not be wronged by being killed. Imagine a
person whose subsequent life would unavoidably contain much more that
would be intrinsically bad for her than would be intrinsically good for her.
Such a person might not be harmed, and might indeed be benefited, by being
painlessly killed. But if she understood what her future life would be like and
still wanted to continue to live, and thus refused to consent to be killed, she
would be wronged by being killed. For she has a moral status that grounds
reasons to act in certain ways for her own sake that are nevertheless independ-
ent of what might be in or against her interests. They are reasons of respect for
her, as an individual who matters because of her intrinsic nature.
Reasons of this sort could in principle apply to one’s treatment of an uncon
nected individual. In addition to the reason deriving solely from the intrinsic
badness of suffering, there might be a further reason not to cause an uncon
nected individual to suffer that is grounded in a requirement of respect for its
nature. Or there might be a reason deriving from its nature not to kill an
unconnected individual, despite its having no interest in continuing to live.
Another random document with
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are characteristic for each species. The Nematus larvae that inhabit
galls possess all the characteristics of those that feed externally. As
a rule the skin of the larva is naked and free from hair, but it is often
minutely tuberculate, and in a few species it is armed with
remarkable forked spines. These spines may exist during part of the
larval life, and completely disappear at one of the moults. The
creatures are as a rule very sluggish, and move about much less
than Lepidopterous larvae; many of them, when alarmed, have the
power of exuding a disagreeable liquid, either from the mouth or
from pores in the skin; in the latter case it may be sent as a sort of
spray to some little distance from the body. This operation is said to
be very efficacious as a means of protecting the larvae from the
attacks of parasitic flies that are desirous of laying eggs in their
bodies. One peculiarity as to their colour has attracted the attention
of Réaumur and subsequent naturalists, namely, that in the case of
many species a great change takes place in the colour during the life
of the larva, and more especially at the period of the last moult. The
change to the pupal state usually takes place in a cocoon, and some
species have the peculiar habit of forming a double cocoon, the
outer one being hard and coarse, while the inner is beautifully
delicate. The cocoon is sometimes formed in the earth, and in that
case it may be to a large extent composed of earthy matter. The
Insect frequently remains a long time in its cocoon before emerging
as a perfect Insect; however long this time may be, it is nearly all of it
passed in the larval state; when the Insect does change to a pupa it
speedily thereafter emerges as a perfect Insect. In the pupa the
parts of the imago may be seen enveloped in a very delicate,
transparent skin.
1. The egg may be laid outside a larva, and the embryonic and larval
developments may both be passed on the exterior.
2. The egg may be laid and the embryonic development passed
through, outside the host, but the parasite on hatching may enter the
host, so that the post-embryonic development is passed in the lymph
of the host.
3. The egg may be laid inside the host, both embryonic and post-
embryonic developments being gone through in the fluids of the
host.
4. The egg may be laid inside another egg, the embryonic and post-
embryonic developments being passed therein.
We shall find that all these conditions exist in the Insects we are
about to consider.
Fam. I. Cynipidae—Gall-flies.
Wings with very few cells, with no dark patch (stigma) on the
anterior margin; pronotum fixed to the mesonotum, and at each
side extending back to the point of insertion of the front wing.
Antennae not elbowed but straight, composed of a moderate
number (12-15) of joints. Early stages passed either in galls or
as parasites in the bodies of other Insects.
The wings frequently bear fine hairs; the paucity of nervures and the
absence of the "stigma" are of importance in the definition of the
family. The most important of the cells is one called the radial cell,
situate just beyond the middle of the front part of the wing.
The exact mode in which the egg is brought to the requisite spot in
the plant is still uncertain. The path traversed by the ovipositor in the
plant is sometimes of considerable length, and far from straight; in
some cases before it actually pierces the tissues, the organ is thrust
between scales or through fissures, so that the terebra, or boring
part of the ovipositor, when it reaches the minute seam of cambium,
is variously curved and flexed. Now as the canal in its interior is of
extreme tenuity, and frequently of great length, it must be a very
difficult matter for the egg to reach the tissue where it should
develop. The eggs of Cynipidae are very remarkable bodies; they
are very ductile, and consist of a head, and of a stalk that in some
cases is five or six times as long as the head, and is itself somewhat
enlarged at the opposite end. Some other Hymenoptera have also
stalked eggs of a similar kind (Fig. 357, A, egg of Leucospis). It has
been thought that this remarkable shape permits of the contents of
the egg being transferred for a time to the narrower parts, and thus
allows the broader portion of the egg to be temporarily compressed,
and the whole structure to be passed through a very narrow canal or
orifice. It is, however, very doubtful whether the egg really passes
along the canal of the borer. Hartig thought that it did so, and Riley
supports this view to a limited extent. Adler, however, is of a different
opinion, and considers that the egg travels in larger part outside the
terebra. It should be remembered that the ovipositor is really
composed of several appendages that are developed from the
outside of the body; thus the external orifice of the body is
morphologically at the base of the borer, the several parts of which
are in longitudinal apposition. Hence there is nothing that would
render the view of the egg leaving the ovipositor at the base
improbable, and Adler supposes that it actually does so, the thin end
being retained between the divisions of the terebra. Riley is of
opinion that the act of oviposition in these Insects follows no uniform
system. He has observed that in the case of Callirhytis clavula,
ovipositing in the buds of Quercus alba, the eggs are inserted by the
egg-stalk into the substance of the leaf, and that the egg-fluids are at
first gathered in the posterior end, which is not inserted. "The fluids
are then gradually absorbed from this exposed portion into the
inserted portion of the egg, and by the time the young leaves have
formed the exposed [parts of the] shells are empty, the thread-like
stalk has disappeared, and the egg-contents are all contained within
the leaf tissue." He has also observed that in Biorhiza nigra the
pedicel, or stalk, only is inserted in the embryonic leaf-tissue, and
that the enlarged portion or egg-body is at first external. The same
naturalist also records that in the case of a small inquiline species,
Ceroptres politus, the pedicel of the egg is very short, and in this
case the egg is thrust down into the puncture made by the borer, so
that the egg is entirely covered.
Some Cynipidae bore a large number of the channels for their eggs
before depositing any of the latter, and it would appear that it is the
rule that the boring of the channel is an act separate from that of
actual oviposition. Adler distinguishes three stages: (1) boring of the
canal; (2) the passage of the egg from the base of the ovipositor,
where the egg-stalk is pinched between the two spiculae and the
egg is pushed along the ovipositor; (3) after the point of the
ovipositor is withdrawn, the egg-body enters the pierced canal, and
is pushed forward by the ovipositor until it reaches the bottom.[436]
About fifty years ago Hartig reared large numbers of certain species
of gall-flies from their galls, obtaining from 28,000 galls of Cynips
disticha about 10,000 flies, and from galls of C. folii 3000 or 4000
examples of this species; he found that all the individuals were
females. His observations were subsequently abundantly confirmed
by other naturalists, among whom we may mention Frederick Smith
in our own country, who made in vain repeated attempts to obtain
males of the species of the genus Cynips. On one occasion he
collected in the South of England 4410 galls of C. kollari (at that time
called C. lignicola), and from these he obtained 1562 flies, all of
which were females. A second effort was attended with similar
results. Hartig, writing in 1843, after many years' experience, stated
that though he was acquainted with twenty-eight species of the
genus Cynips, he had not seen a male of any one of them. During
the course of these futile attempts it was, however, seen that a
possible source of fallacy existed in the fact that the Insects were
reared from collected galls; and these being similar to one another, it
was possible that the males might inhabit some different gall. Adler
endeavoured to put the questions thus raised to the test by means of
rearing females from galls, and then getting these females to
produce, parthenogenetically, galls on small oaks planted in pots,
and thus completely under control. He was quite successful in
carrying out his project, and in doing so he made a most
extraordinary discovery, viz. that the galls produced by these
parthenogenetic females on his potted oaks, were quite different
from the galls from which the flies themselves were reared, and
were, in fact, galls that gave rise to a fly that had been previously
considered a distinct species; and of this form both sexes were
produced. Adler's observations have been confirmed by other
naturalists, and thus the occurrence of alternation of generations,
one of the two generations being parthenogenetic, has been
thoroughly established in Cynipidae. We may mention one case as
illustrative. A gall-fly called Chilaspis lowii is produced from galls on
oak-leaves at Vienna at the end of April, both sexes occurring. The
female thereafter lays eggs on the ribs of the leaves of the same
kind of oak, and thus produces a different gall from that which
nourished herself. These galls fall off with the leaves in the autumn,
and in July or August of the following year a gall-fly is produced from
them. It is a different creature from the mother, and was previously
known to entomologists under the name of Chilaspis nitida. Only
females of it occur, and these parthenogenetic individuals lay their
eggs in the young buds of the oak that are already present in the
autumn, and in the following spring, when the buds open and the
leaves develop, those that have had an egg laid in them produce a
gall from which Chilaspis lowii emerges in April or May. In this case
therefore the cycle of the two generations extends over two years,
the generation that takes the greater part of the time for its
production consisting only of females. Adler's observations showed
that, though in some species this alternation of generations was
accompanied by parthenogenesis in one part of the cycle, yet in
other species this was not the case. He found, for instance, that
some gall-flies of the genus Aphilothrix produced a series of
generations the individuals of which were similar to one another, and
were all females and parthenogenetic. In some species of the old
genus Cynips no males are even yet known to occur. A very curious
observation was made by the American, Walsh, viz. that of galls
gathered by him quite similar to one another, some produced
speedily a number of both sexes of Cynips spongifica, while much
later on in the season the remainder of the galls gave rise to females
only of an Insect called Cynips aciculata. It is believed that the galls
gathered by Walsh[437] were really all one species; so that parts of
the same generation emerge at different times and in two distinct
forms, one of them parthenogenetic, the other consisting of two
sexes. It has, however, been suggested that Cynips spongifica and
C. aciculata may be two distinct species, producing quite similar
galls.
Bassett recorded the first case of the kind in connexion with a North
American species, Cynips (Ceroptres) quercus-arbos Fitch. He says:
"On the first of June galls on Quercus ilicifolia had reached their full
size, but were still tender, quite like the young shoots of which they
formed part. Examining them on that day, I discovered on them two
gall-flies, which I succeeded in taking. They were females, and the
ovipositor of each was inserted into the gall so deeply that they could
not readily free themselves, and they were removed by force."
The great resemblance of the inquiline gall-fly to the fly that makes
the gall both dwell in, has been several times noticed by Osten
Sacken, who says "one of the most curious circumstances
connected with the history of two North American blackberry galls is,
that besides the Diastrophus, which apparently is the genuine
originator of the gall, they produce another gall-fly, no doubt an
inquiline, belonging to the genus Aulax, and showing the most
striking resemblance in size, colouring, and sculpture to the
Diastrophus, their companion. The one is the very counterpart of the
other, hardly showing any differences, except the strictly generic
characters! This seems to be one of those curious instances, so
frequent in entomology, of the resemblance between parasites and
their hosts! By rearing a considerable number of galls of D.
nebulosus I obtained this species as well as its parasite almost in
equal numbers. By cutting some of the galls open I ascertained that
a single specimen of the gall frequently contained both species, thus
setting aside a possible doubt whether these Insects are not
produced by two different, although closely similar galls."[438]
Not more than 500 species of Psenides and Inquiline Cynipidae are
known from all parts of the world; and of described Parasitic
Cynipidae there are only about 150 species. The British forms have
recently been treated by Cameron in the work we have already
several times referred to.[439]