International Journal of Darwin’s aliens

Astrobiology
Samuel R. Levin1, Thomas W. Scott1, Helen S. Cooper2 and Stuart A. West1
cambridge.org/ija
1
Department of Zoology, University of Oxford, South Parks Road, Oxford OX1 3PS, UK and 237 Beech Croft Road,
Oxford OX2 7AY, UK

Abstract
Review
Making predictions about aliens is not an easy task. Most previous work has focused on
Cite this article: Levin SR, Scott TW, Cooper extrapolating from empirical observations and mechanistic understanding of physics, chem-
HS, West SA. Darwin’s aliens. International istry and biology. Another approach is to utilize theory to make predictions that are not tied
Journal of Astrobiology https://doi.org/
10.1017/S1473550417000362
to details of Earth. Here we show how evolutionary theory can be used to make predictions
about aliens. We argue that aliens will undergo natural selection – something that should
Received: 30 May 2017 not be taken for granted but that rests on firm theoretical grounds. Given aliens undergo
Accepted: 8 September 2017 natural selection we can say something about their evolution. In particular, we can say some-
Key words:
thing about how complexity will arise in space. Complexity has increased on the Earth as a
aliens; astrobiology; evolution; extraterrestrial result of a handful of events, known as the major transitions in individuality. Major transi-
life; individuality; major transitions tions occur when groups of individuals come together to form a new higher level of the indi-
vidual, such as when single-celled organisms evolved into multicellular organisms. Both
Author for correspondence:
Samuel R. Levin, E-mail: [email protected].
theory and empirical data suggest that extreme conditions are required for major transitions
ac.uk to occur. We suggest that major transitions are likely to be the route to complexity on other
planets, and that we should expect them to have been favoured by similarly restrictive con-
ditions. Thus, we can make specific predictions about the biological makeup of complex
aliens.

Introduction
There are at least 100 billion planets in our Galaxy alone (Cassan et al. 2012), and at least 20%
of them are likely to fall in the habitable zone (Petigura et al. 2013), the region of space capable
of producing a biosphere. Even if 0.001% of those planets evolved life, that would mean
200 000 life-harbouring planets in our Galaxy; and it would only take one alien life form
for our conception of the Universe to change dramatically. It is no wonder, then, that hundreds
of millions of dollars have recently been invested in astrobiology research (Schneider 2016),
the USA and Europe have rapidly growing astrobiology initiatives (Des Marais et al. 2008;
Horneck et al. 2016), and myriad new work has been done to try and predict what aliens will
be like (Benner 2003; Davies et al. 2009; Rothschild 2009; Rothschild 2010; Shostak 2015). The
challenge, however, is that when trying to predict the nature of aliens, we have only one sample
– Earth – from which to extrapolate. As a result, making these predictions is hard.
So far, the main approach to making predictions about extra-terrestrial life has been rela-
tively mechanistic (Domagal-Goldman et al. 2016). We have used observations about how
things have happened on the Earth to make statistical statements about how likely they are
to have happened elsewhere. For example, certain traits have evolved many times on the
Earth, and so we posit that extraterrestrial life forms will converge on the same earthly
mechanisms. Because eye-like organs have evolved at least 40 times (von Salvini-Plawen &
Mayr 1977), and are relatively ubiquitous, we predict that they would evolve on other planets,
too (Conway Morris 2003; Flores Martinez 2014). Similarly, we have used a mechanistic
understanding of chemistry and physics to make predictions about what is most probable
on other planets. For example, carbon is abundant in the Universe, chemically versatile,
and found in the interstellar medium, so alien life forms are likely to be carbon-based
(Cohen & Stewart 2001). These kinds of predictions come from a mixture of mechanistic
understanding and extrapolating from what has happened on the Earth. There is no theoretical
reason why aliens could not be silicon-based and eyeless.
An alternative approach is to use theory. When making predictions about life on other pla-
nets, a natural theory to use would be evolutionary theory. Evolutionary theory has been used
to explain a wide range of features of life on the Earth, from behaviour to morphology. For
example, it has allowed us to predict when some organisms, especially insects, should manipu-
late the sex of their offspring, to produce an excess of sons or daughters, how some birds
© Cambridge University Press 2017 should forage for food, and why males tend to be larger than females (Darwin 1871;
Clutton-Brock & Harvey 1977; Davies & Houston 1981; West 2009; Davies et al. 2012). If
life arises on other planets, then the evolutionary theory should be able to make similar pre-
dictions about it. Neither approach – theoretical or mechanistic – is more or less valid than the
other. But each has different advantages and can be used to make different sorts of predictions.

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 2 Samuel R. Levin et al.

Here, we examine how theoretical and mechanistic approaches (1) heredity; (2) variation; and (3) differential success linked to
can be combined to better understand what to expect from alien variation, then natural selection will follow. The entities that are
life. We consider whether aliens will undergo natural selection, more successful will become more prevalent in the population,
and what implications would follow if they do. That aliens as a result of being ‘selected’. Natural selection does not depend
undergo natural selection is something often taken for granted, on a specific genetic system (Darwin knew nothing of modern
but which needs justification on firm theoretical grounds. We genetics) or a specific genetic material, elemental makeup or
then turn our attention to a specific subset of aliens: complex planet-type. Given that 1, 2 and 3 exist, natural selection occurs
ones. We examine how complexity has arisen on the Earth, and (Fig. 1).
make predictions about how complexity would arise elsewhere Natural selection not only explains evolutionary change, it also
in the Universe. Finally, we describe some biological features we explains adaptation. When we look around at the natural world,
would expect to find in complex extraterrestrial life. we cannot help but see what looks like design: a giraffe’s neck
is for reaching high up leaves, a stick insect’s body for camouflage,
a tree’s leaf for photosynthesizing. Organisms look designed or
Natural selection ‘adapted’ for the world in which they live. Through the gradual
selection of small improvements, traits associated with success
On Earth
in the environment accrue in the population. Consequently,
Darwin (1859) showed that just a few simple features of life on over time, natural selection will lead to organisms that appear
Earth lead to evolutionary change via natural selection. as if they were designed for success in the environment. The
Individual organisms differ in how they look and act – there is clause ‘as if’ is key here – natural selection leads to the appearance
natural variation. These differences are heritable – offspring of design (adaptation), without a designer (Grafen 2003; Gardner
tend to look and act like their parents. These heritable differences 2009).
are linked to differential success – some individuals, as a result of In fact, natural selection is the only explanation we have for
how they are made or behave, leave more offspring than others. the appearance of design without a designer (Gardner 2009).
These three features, with heritable variation leading to differen- Other processes can cause evolutionary change. For example,
tial success, result in natural selection (Darwin 1859; Fisher 1930). a mutation can cause a change from one generation to the
Any traits or behaviours linked to the greater production of off- next. But, without natural selection, random mutation is incred-
spring (higher fitness or success) will build up in the population ibly unlikely to produce the complex traits that we see around
over time. As the environment changes, different traits lead to us, like limbs or eyes. Things that appear purposeful, such as
higher success. This leads to changes in the population or evolu- limbs, organs and cells, require the gradual selection of
tionary change. improvements.
Thus, the ingredients required for natural selection are incred- Another way to say this is that natural selection is unique
ibly simple. Given a collection of entities (a population) that has: because it is a directional force. The entities that increase in

Fig. 1. Natural selection. Natural Selection operates if three conditions are satisfied: variation, differential success linked to variation and heredity. Here, we illus-
trate with an example: the evolution of long necks in giraffes. (i) Initially, there are natural variations in giraffes’ neck lengths. (ii) Longer-necked giraffes have access
to more food, high up in the trees and so live longer to have more offspring. (iii) Giraffes’ offspring resemble their parents. As a result of (i), (ii) and (iii), the popu-
lation gradually shifts to be dominated by long-necked giraffes.

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 International Journal of Astrobiology 3

representation in the population are a specific subset of the popu- its apparent purposiveness, leading to tasks such as replication
lation – those that are better at replicating. Natural selection and metabolism (Maynard Smith & Szathmáry 1995). Further,
increases fitness (Fisher 1930). As a result of these ‘successful’ without natural selection, entities cannot adapt to their environ-
entities accruing in the population, over time entities become ment, and are therefore transient and will not be discovered. If
adapted for the apparent purpose of success. They look like ‘well- we identified an extra-terrestrial entity that we deemed to be a for-
designed’ machines, with the ‘purpose’ of their ‘design’ being suc- eign life form, but that had no degree of adaptedness, this predic-
cessful replication. tion would not hold.
Picture an alien (Fig. 2). If what you are picturing is a simple
replicating molecule, then this ‘alien’ might not undergo natural
In space selection (Fig. 2a). For example, it could replicate itself perfectly
every time, and thus there would be no variation, and it would
Natural selection is the only way we know to get the kinds of life
never improve. Or it might have such a high error rate in replica-
forms we are familiar with, from viruses to trees. By familiar, we
tion that it quickly deteriorates. If we count things like that as life,
are not restricting ourselves to life forms that look earthly. Instead,
then there could be aliens that do not undergo natural selection.
they are familiarly life-like in the sense that they stand out from
But if you are picturing anything more complex or purposeful
the background of rocks and gases because they appear to be
than a simple molecule, then the alien you are picturing has
busy trying to replicate themselves. A simple replicator could
undergone natural selection (Fig. 2b). This is the kind of predic-
arise on another planet. But without natural selection, it won’t
tion that theory can make. Given heredity, variation and differen-
acquire apparently purposeful traits like metabolism, movement
tial success, aliens will undergo natural selection. Or, more
or senses. It won’t be able to adapt to its environment, and in
interestingly, without those three things, aliens could not be
the process, become a more complex, noticeable and interesting
more complicated than a replicating molecule. Given an adapted
thing.
alien, one with an appearance of design or purpose, it will have
We can ask, then, will aliens undergo natural selection?
undergone natural selection.
Evolutionary theory tells us that, for all but the most transient
and simple molecules, the answer is yes. Without a designer,
the only way to get something with the apparent purpose of rep- Complexity
licating itself (something like a cell or a virus), is through natural
What is complexity?
selection. Consequently, if we are able to notice it as life, then it
will have undergone natural selection (or have been designed by We have established that aliens will undergo natural selection. It
something that itself underwent natural selection). also seems reasonable that, given the sliding scale from replicating
It is easy to quibble about the definition of life, and as some molecules to large creatures with many ‘body parts’, and beyond,
authors have pointed out, trying to do so can reveal more about some alien discoveries would be more interesting than others. In
human language than about the external world (Cleland & particular, the more complex the aliens we find, the more interest-
Chyba 2002). Our goal here is not to thoroughly define life. We ing and exciting they will be, irrespective of whether they appear
adopt a functional stance – what separates life from non-life is anything like the life forms on the Earth. Something similar to a

Fig. 2. Picture an alien. These illustrations represent different levels of adaptive complexity we might imagine when thinking about aliens. (a) A simple replicating
molecule, with no apparent design. This may or may not undergo natural selection. (b) An incredibly simple, cell-like entity. Even something this simple has suf-
ficient contrivance of parts that it must undergo natural selection. (c) An alien with many intricate parts working together is likely to have undergone major
transitions.

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 4 Samuel R. Levin et al.

colony of Ewoks from Star Wars or the Octomite in Fig. 4 would complexity. For example, the evolution of multicellularity
likely be more interesting than a simple chemical replicator. involved a transition from an entity with one part (the single-
Complexity is difficult to define, and there is certainly no hard celled organism) working for the success of itself, to an entity
and fast rule about what is and is not complex. In biology, it is with many parts (the multicellular organism), working for the
common to define complexity in terms of functional parts. success of the whole group. The cells can now have very different
Things with more parts taking on more tasks and containing functions (a division of labour), as each is just a component of a
more functional interactions are more complex (Maynard Smith multicellular machine, sacrificing itself for the good of the group,
& Szathmary 1995; Corning & Szathmáry 2015). A tree is more to get a sperm or egg cell into the next generation. As a result,
complex than a virus, and a beehive is more complex than a pro- diverse specialized forms such as eyes, kidneys, and brains were
tein. Importantly, with organisms as with machines, the parts able to develop. The rise in complexity on Earth has been
need to be working towards a common purpose, such as assem- mediated by a handful of such jumps, when units with different
bling a car or surviving to reproduce. Again, our goal here is goals (genes, single cells, individual insects) became intricately
not to provide definitions. The challenge comes at the boundaries, linked collectives with a single common goal (genomes, multicel-
for example between a virus and a cell, where the definitions lular organisms, eusocial societies). Increases in complexity can
become murky. In the following sections, we are not focusing also occur through mutations, gene duplications, or even whole
on the boundaries, but things, like the vast majority of life on genome duplications, but these are not major transitions. These
the Earth, which clearly have a multitude of parts working in con- other changes tend to be reversible and gradual, while major tran-
cert. Astrobiology is a largely empirical field, and the kinds of sitions are irreversible and cause large leaps in complexity.
things programs like SETI are searching for are undeniably The identification of major evolutionary transitions was an
complex. empirical observation about how complexity has increased on
earth (Maynard Smith & Szathmáry 1995). The next step was to
use evolutionary theory to provide insight about when (or
Complexity on Earth under what conditions) we can expect major transitions to
What do we know about how complexity arises on the Earth? The occur (Maynard Smith & Szathmáry 1995; Queller 1997;
theory of natural selection itself is silent about whether complexity Gardner & Grafen 2009; Bourke 2011; West et al. 2015). Major
will arise. The theory is useful for making predictions about what transitions involve the original entities completely subjugating
kinds of conditions or environments will lead to what kinds of their own interests for the interests of the new collective. This
evolutionary adaptations – not for making long-term predictions represents an incredibly extreme form of cooperation. Think of
about the form of specific traits or creatures. However, recent the skin or liver cells in your body sacrificing for your sperm or
advances in the field of evolutionary biology have shed light on eggs, or the worker ants in a eusocial colony sacrificing for the
how complexity has arisen on the Earth, on what points on the queen. Evolutionary theory tells us what conditions lead to such
tree of life this has happened, and on what theoretical conditions extraordinary cooperation.
favour it (Maynard Smith & Szathmáry 1995; Queller 1997;
Bourke 2011; West et al. 2015). What conditions drive major transitions?
In particular, the evolution of complex life on the Earth
appears to have depended upon a small number of what have Consider a multicellular organism, such as yourself. Why don’t
been termed major evolutionary transitions in individuality. In your hand and heart cells try to reproduce themselves, as opposed
each transition, a group of individuals that could previously rep- to helping your sperm or egg cells? The answer involves genetic
licate independently cooperate to form a new, more complex life similarity or ‘relatedness’ (Hamilton 1964). Your hand cells con-
form or higher level organism. For example, genes cooperated to tain the same genes as your sperm cells because they are clonal
form genomes, different single-celled organisms formed the copies. A hand cell could in principle get the same fraction of
eukaryotic cell, cells cooperated to form multicellular organisms, its genes into the next generation (all of them) by either copying
and multicellular organisms formed eusocial societies (Maynard itself, or by helping copy the sperm cells. A similar phenomenon
Smith & Szathmáry 1995; Queller 1997; Bourke 2011; West occurs in eusocial insects, such as some ants, bees, wasps and ter-
et al. 2015). mites. A worker termite can pass on half her genes to her off-
spring. But a random sibling in the colony (her brother or
sister) also contains, on average, half her genes. Thus, a worker
Major transitions can get the same fraction of gene copies into the next generation
by reproducing or by helping her mother, the queen, to reproduce
Major transitions on the Earth
(Hamilton 1964; Boomsma 2009). Helping their mother is likely
Major evolutionary transitions are defined by two features. First, to be more efficient than reproducing on their own, and so our
entities that were capable of replication before the transition can termite can better get their genes into the next generation by help-
replicate only as part of a larger unit after it (interdependence). ing rather than reproducing (Hamilton 1964; Queller &
For example, the cells in our bodies cannot evolve back into Strassmann 1998; Bourke 2011).
single-celled organisms. Second, there is a relative lack of conflict These are two examples of alignment of interests. The ‘inter-
within the larger unit, such that it can be thought of as an organ- ests’ are evolutionary interests in getting genes into future genera-
ism (individual) in its own right (Queller & Strassmann 2009; tions. The hand and the sperm cells both act as if they ‘want’ to
West et al. 2015). For example, it is common to think of a single get copies of their genes into the next generation, because as we
bird as an individual, and not as a huge community of cells each discussed above, natural selection will have led to them being
doing their own thing. adapted in this way (Grafen 2003; Gardner 2009). The interests
Major transitions are important because the new higher-level between them are aligned because they share the same genes.
organisms that they produce can lead to a great jump in When individuals share genes, we say that they are genetically

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 International Journal of Astrobiology 5

related. Relatedness is a statistical measure of the extent to which mitochondria and the nucleus, the individual parts must be
individuals share genes (Grafen 1985). co-dependent for joint reproduction (Foster & Wenseleers 2006;
In the case of eusocial ant colonies and human bodies, the West et al. 2015) – which can be thought of as a different form
interests are aligned through genetic relatedness. But there are of bottleneck. The rarity of conditions like these – conditions
other ways for evolutionary interests to be aligned. Consider, for under which alignment is so complete – explains the rarity of
example, a mutualism between two species. Some aphids carry major transitions in individuality in the history of life.
bacteria in their gut (Moran 2007). The aphids provide the bac-
teria with sugars and other nutrients to survive and the bacteria
provide the aphids with vital amino acids missing from their Biology of organisms that have undergone major transitions
diet. The aphid and the bacteria do not share the same genes, Do the conditions required for major transitions tell us anything
but neither can reproduce without the other. To reproduce itself, about the biology of organisms that have undergone major transi-
the aphid has to help reproduce the bacteria and vice versa. Again, tions? Yes. Organisms are a nested hierarchy, where each nested
their evolutionary interests are aligned. level is the vestige of a former individual (Fig. 3). Eusocial ant col-
The very cells that make up our bodies – known as eukaryotic onies function as a single individual, but are made up of multicel-
cells – evolved through a similar kind of alignment of interests lular organisms. Those organisms themselves are made up of cells.
(Margulis 1970; Thiergart et al. 2012; Archibald 2015). Early in In turn, those cells resulted from the fusion of two simple species
the evolution of life, one bacterial species engulfed another. early in evolution. Each of those organisms had a genome that
Over time, the two species took on different roles, with one spe- evolved from the union of the individual, replicating molecules.
cializing in replication and the other in energy production. The Further, at each level of the hierarchy, there must be something
nucleus of our cells is the descendant of the former, and the mito- to align the interests of the parts. This usually happens through
chondria the latter. Neither can reproduce without the other. some form of population bottlenecking. When the parts are
Their interests are aligned through reproductive dependence on related, it is a relatedness bottleneck, such as the single-celled
each other. stage in multicellular organisms, or the singly mated female in
All cooperation in nature requires alignment of interests (West the social insects (Boomsma 2009, 2013; West et al. 2015).
et al. 2007). Consider, for example, flower pollination by bees. The When the parts are unrelated, it is usually another form of a
bee benefits by receiving food from the flower, and the flower bottleneck, such as enforced vertical transmission with joint
benefits by being pollinated. But major transitions are a particu- reproduction (Foster & Wenseleers 2006; West et al. 2015). We
larly extreme form of cooperation. Compare the pollination scen- use the term ‘bottleneck’ to refer to new generations being
ario to the cells within the flower or the bee. Major transitions founded by a strict unit (the zygote, the mutualist pair, etc.),
involve organisms cooperating so completely that they give up but another way to think of this is that the parts require each
their status as individuals, becoming parts of a whole (Queller other for reproduction (e.g. the soma and the germ line, or the
& Strassmann 2009). Unsurprisingly, then, major transitions mitochondria and the nucleus). Other, further aligners may be
require the extreme condition of effectively complete or perfect required (e.g. in multicellular organisms, there may need to be
alignment of interests (Gardner & Grafen 2009; West et al. 2015). a cap on somatic mutations), but these are more likely to be life-
It is also useful to consider the biology of organisms that do form specific.
not have interests sufficiently aligned, and thus where conflict To conclude so far, empirical observation tells us that com-
remains and major transitions have not occurred. For example, plexity has increased on earth through major transitions.
in single-celled organisms, we can compare non-clonal coopera- Evolutionary theory tells us that for major transitions to occur,
tive groups of things like slime moulds with clonal groups such the conflict must be eliminated. The theory also tells us what con-
as those that make up multicellular organisms such as humans ditions lead to the elimination of conflict. The empirical data
and trees. These non-clonal groups have evolved only relatively agree with the predictions of the theory, in that major transitions
limited division of labour, and never complex multicellular organ- have only occurred in the extreme conditions that effectively
isms (Fisher et al. 2013). Numerous experimental studies have remove conflict (Boomsma 2007; Hughes et al. 2008; Fisher
shown that this is because in non-clonal groups non-cooperative et al. 2013; West et al. 2015; Fisher et al. 2017).
‘cheats’ can spread, limiting the extent of cooperation (Griffin
et al. 2004; Diggle et al. 2007; Kuzdzal-Fick et al. 2011;
Rumbaugh et al. 2012; Pollitt et al. 2014; Popat et al. 2015; Complex aliens
Inglis et al. 2017).
Complexity and major transitions in space
Thus, there must be something in place to maintain the align-
ment of interests (Bourke 2011; West et al. 2015). Evolutionary We can now ask: what does the major evolutionary transition
theory can suggest what these somethings would have to be. In approach tell us about aliens? Will extraterrestrial life undergo
multicellular organisms, the something is the single-celled bottle- major transitions? Not necessarily. Natural selection cannot pre-
neck (Buss 1987; Queller 2000). Multicellular organisms start each dict a specific course of evolution. However, as we have said, we
new generation as a single-celled zygote, such that all the cells in might be particularly interested in complex aliens. Complexity
the resulting body are clonal (it could also be a spore giving rise to requires different parts or units working together towards a com-
a haploid cell). Eusocial insect colonies evolved from colonies mon goal or purpose. Under natural selection, units are selected
founded by a singly mated queen (Boomsma 2007, 2009, 2013; to be selfish, striving to replicate themselves at the expense of
Hughes et al. 2008). If the queen had multiple mating partners, others. Theory tells us that for units to unite under a common
a worker would have half-sisters, and be less related to her siblings purpose, the evolutionary conflict between them must effectively
than her offspring, breaking down the alignment. The monogam- eliminate (Gardner & Grafen 2009; West et al. 2015).
ous mating pair is the eusocial colony’s equivalent of a zygote or a Once again, picture an alien (Fig. 2). If you are picturing some-
bottlenecking event (Boomsma 2013). With unrelated units, like thing like unlinked replicating molecules or undifferentiated blobs

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 6 Samuel R. Levin et al.

of slime, then your aliens might not have undergone major tran- The biology of complex aliens
sitions. But if what you are picturing has different parts with spe-
Given that complex aliens will have undergone major transitions,
cialized functions, then your alien is likely to have undergone
we can make a number of predictions about their biology (Fig. 4).
major transitions (Fig. 2c). What matters is not that we call
them ‘major transitions’, but rather that complexity requires mul-
tiple parts of an organism striving to the same purpose, and that
theory predicts that this requires restrictive conditions (Gardner 1. They will be entities that are made up of smaller entities – a
& Grafen 2009; West et al. 2015). Consequently, if we find com- nested hierarchy of individuality with as many levels as com-
plex organisms, we can make predictions about what they will be pleted transitions. This could mean a collection of replicators,
like. like the first genomes on the Earth, or some hideously complex
Are there other ways to get complexity? To do so, natural nesting of groups on a planet where many more transitions
selection would have to sculpt separate parts with unique func- have occurred than on our own. For example, you might
tions out of a single replicator. Could, for example, the alien imagine a ‘society of societies’, where many different social col-
equivalent of a single copy of a gene, housed in one ‘cell’ generate onies collaborate, with each society specializing on different
the equivalent of limbs and organs? If so, it would disprove our tasks, such that they are completely dependent on each
prediction. However, both empirical (major transitions are how other. Versions of the simpler entities are likely to be found
complexity has increased on Earth) and theoretical (functional free-living on the planet as well.
parts requires the elimination of conflict) evidence support the 2. Whatever the number of transitions, there will be something
argument that complex aliens will have undergone major that aligns interests, or eliminates conflict within the entities,
transitions. at the level of each transition.

Fig. 3. Major Transitions. Life started with naked replicating molecules, and has since undergone a series of major transitions. Arrows show the occurrence of major
transitions in individuality. Dotted arrows represent transitions between dislike things and solid lines represent transitions between like things. Callouts show
examples of the present-day organisms that have undergone that transition but no further ones. (a) As we have not yet identified the earliest replicators,
Spiegelman’s monster, a simple replicating RNA molecule, is shown as an example candidate. (b) A single-celled bacteria, such as Escherichia coli. (c) A single-celled
eukaryote, like Blepharisma japonicum. (d) A multicellular organism, like frogs. (e) An obligate eusocial colony, such as honeybees. (f) Secondary endosymbiosis
events, such as the origin of the chloroplast. (g) Further endosymbiosis events, such as those leading to Dinoflagellates. (h) Obligate interspecific mutualisms, such
as between aphids and buchnera bacteria. (i) Obligate mutualisms between a multicellular organism and eusocial colony, such as between leaf-cutter ants and
fungi. All images courtesy of Wikipedia.

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 International Journal of Astrobiology 7

Fig. 4. Major transitions in space: ‘The Octomite’. A complex alien that comprises a hierarchy of entities, where each lower-level collection of entities has aligned
evolutionary interests such that conflict is effectively eliminated. These entities engage in a division of labour, with various parts specializing on various tasks, such
that the parts are mutually dependent.

3. Theory suggests that some sort of population bottlenecking argument is: (1) Extraterrestrial life will have undergone natural
will be key to aligning interests. Bottlenecking is not necessar- selection. (2) Knowing that aliens undergo natural selection, we
ily the only way to eliminate conflict, but it is probably the can make further predictions about their biology, based on the
easiest evolutionary route to take. In particular, it does not theory of natural selection. In particular, we can say something
require additional mechanisms of enforcement, such as kin about complex aliens – that they will likely have undergone
discrimination, policing or randomization. The specific kinds major transitions. (3) Theory tells us that restrictive conditions,
of bottlenecking will depend on whether like or dislike units which eliminate conflict, are required for major transitions. (4)
are united. Consequently, complex aliens will be composed of a nested hier-
a. When like entities come together, interests can be aligned archy of entities, with the conditions required to eliminate conflict
through a bottleneck similar to our single-celled bottle- at each of those levels.
neck in multicellular organisms or the single mating When making predictions about aliens, we must take advantage
pair in eusocial colonies, which maximizes relatedness of our entire scientific toolkit. Mechanistic understanding is a good
between entities. way to extrapolate from what we see on Earth. The theory is a good
b. If the organisms are made up different types of entities, way to make predictions that are independent of the details of the
we can expect something similar to the bottleneck that Earth. Combining both approaches is the best way to make predic-
forces mitochondria and nuclei to pass to the next gener- tions about the many hundreds, thousands or millions of hypo-
ation together, with joint reproduction. By trapping indi- thetical aliens. Now we just need to find them.
viduals together over evolutionary time, their interests
Acknowledgements. We thank The Clarendon Fund, Hertford College, and
become aligned.
the Natural Environment Research Council for funding; and Magdalen College
c. Some aliens, like us, may contain both types of conflict
for emergency housing.
reduction, for having both like and dislike types joined
within them. Author disclosure statement. No competing financial interests exist.

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