Beruflich Dokumente
Kultur Dokumente
Acta Astronautica
journal homepage: www.elsevier.com/locate/actaastro
a r t i c l e in f o abstract
Article history: Designing interstellar starships for human migration to exoplanets requires establishing
Received 27 September 2013 the starship population, which factors into many variables including closed-ecosystem
Received in revised form design, architecture, mass and propulsion. I review the central issues of population
10 December 2013
genetics (effects of mutation, migration, selection and drift) for human populations
Accepted 20 December 2013
Available online 30 December 2013
on such voyages, specifically referencing a roughly 5-generation (c. 150-year) voyage
currently in the realm of thought among Icarus Interstellar0 s Project Hyperion research
Keywords: group. I present several formulae as well as concrete numbers that can be used to help
Multigenerational space travel determine populations that could survive such journeys in good health. I find that
Space genetics
previously proposed such populations, on the order of a few hundred individuals, are
Space colonization
significantly too low to consider based on current understanding of vertebrate (including
Space settlement
human) genetics and population dynamics. Population genetics theory, calculations and
computer modeling determine that a properly screened and age- and sex-structured total
founding population (Nc) of anywhere from roughly 14,000 to 44,000 people would be
sufficient to survive such journeys in good health. A safe and well-considered Nc figure is
40,000, an Interstellar Migrant Population (IMP) composed of an Effective Population [Ne]
of 23,400 reproductive males and females, the rest being pre- or post-reproductive
individuals. This number would maintain good health over five generations despite (a)
increased inbreeding resulting from a relatively small human population, (b) depressed
genetic diversity due to the founder effect, (c) demographic change through time and (d)
expectation of at least one severe population catastrophe over the 5-generation voyage.
& 2013 IAA. Published by Elsevier Ltd. All rights reserved.
0094-5765/$ - see front matter & 2013 IAA. Published by Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.actaastro.2013.12.013
C.M. Smith / Acta Astronautica 97 (2014) 16–29 17
Fig. 1 indicates the timescale pertinent to this paper characteristics to preserve civilization and our species in
which models arrival at an exoplanet system roughly four the long term [3].
light years away in about 150 years (five human genera-
tions) if we achieve, in the next century, the same 1000- 2. Biological Issues in interstellar voyaging
fold increase in mechanized speed as achieved in the last
century from the locomotive to the Voyager probe (velo- Evolution is manifest any difference in the genetic proper-
cities from about 30 miles per hour to about 30,000 miles ties of parent and offspring generations. Particularly in
per hour, respectively). I emphasize that the point is not to sexually reproducing species which combine parental DNA
journey endlessly, but to colonize a distant planet, where in sperm and egg production, producing novelties, cloning is
the human population could grow again – this requires rare, and evolution occurs at each generation. Thus, inter-
arriving at such a destination with a genetically viable and stellar voyaging must grapple with genetic evolution. Four
healthy population. As I show below, one source of essential factors condition the genetic composition of any
humanity0 s longevity so far has been its large population, population. They are discussed below, with notes on their
and while we should not plan to grow beyond our implications for interstellar voyage populations.
resources, being widely spread (e.g. among more than
one solar system), genetically and biologically diverse 2.1. Mutation
(as in current large populations of humans) and populous
(as has been the case for our species for at least 5000 Mutation is the production of new genetic character-
years) are characteristics of species and genera that have istics, also known as diversity. Mutagenesis occurs in many
survived for long periods in evolutionary terms [2]. Inter- ways, including (a) mechanical deformation of DNA during
stellar migration is taken as a method to emulate these replication, (b) alteration of DNA by mutagens such as
Fig. 1. Space and time scales addressed in this paper. On the left axis is time in years and human generations, while on the right are ballpark figures for
significant changes in human biology and culture. Only biology is addressed in this paper.
18 C.M. Smith / Acta Astronautica 97 (2014) 16–29
radiation and certain chemicals, (c) the recombination of or out of that population [11]. Migration is driven by many
parental DNA in the production of gametes (sperm and egg motivators; two obvious kinds are “push” factors that
cells), (d) the failure of mutation-repair processes, and compel migration due to local competition, and “pull”
(more so in prokaryotes than eukaryotes), and (e) hori- factors including the perception of new, nonlocal oppor-
zontal gene transfer, the acquisition of heritable DNA tunities; note that human migration may be driven by
alteration during the course of life [4]. Whatever the cause, cultural imperatives not found in other life forms [12] and
mutations generate differences between parent and off- intent is not necessarily implied here. A human population
spring, and differences among members of any given aboard an interstellar vessel would not have in- or
generation. Currently, humans are estimated to have about out-migration until a new planet is reached, where the
38 DNA base pair mutations per human gamete (sex cell, population could grow again and might split up, resulting
either egg or sperm) as a result of recombination during in out-migration; therefore migration is not considered in
gamete formation, with each newborn carrying roughly this study.
50–100 total mutations (genetic differences among the 3
billion base pairs of the human genome) from its parents 2.3. Selection
and/or siblings [5, p. 966]. Mutations may be advantageous,
conferring an advantage to the carrier and making it more Selection is the favoring of certain individuals and
likely that the mutation will be passed on to the next characteristics in a population over others, often manifest
generation, disadvantageous, conferring a disadvantage in differential reproductive success of individuals such that
(e.g. poorer health) to the carrier, making it less likely that over time, a population of a species or an entire species
the mutation will be passed on to the next generation, or becomes tailored to its environment as poorer-fitted
neutral, having no measureable effect on the likelihood individuals become less numerous and more fit individuals
that it will be passed on to the next generation; this does become more numerous [13,14]. Selection can be natural
not mean, however, that neutral mutations never become (e.g. the effect of natural environment) or cultural (e.g.
more or less common in populations, as we will see below humans make careful mate choices, affecting genetics, and
in Section 2.4. cultural information changes by an evolutionary process as
The genetic diversity of a population (D), an important well, a process beyond the scope of this paper; see [15,16]).
quantity in long-term population planning, is a product of In nature the term “selection” is unfortunate in that it
a mutation rate (μ), the population size (Nc or Ne, intro- implies an intent or directionality for which we have no
duced below), and time (t), because mutations accumulate evidence. Certainly human beings select for and against
in a genome over time [6, p. 151]. Note that t indicates certain properties in our domesticated species, propagat-
generations (or intergenerational interval), not years. ing those we prefer and eventually terminating those that
Human generation time (the number of years between we do not. But most selection is not the result of such
birth and when an adult actually has offspring) has been choice; rather, factors such as temperature, precipitation,
estimated many times and within limits (ranging from c. radiation, the presence of predators and so on determine
25 to 30 among many sampled human populations) is which individuals (and thus which genes) will survive to
“rather consistent regardless of the cultural and environ- form the next generation, and which will not. Selection,
mental differences among human societies” according to then, is non-random differential survival of randomly vary-
Matsumura and Forster [7, p. 1504] and here I use 30 years, ing individuals in a population. Selection is not random
averaging their estimated 25–35 year generation times. because the better-suited tend to propagate while the less-
Mutation rate is quite variable among species, subpopula- fit propagate less, leading to order over time rather than
tions of species, families, individuals, sexes and even bodily disorder.
tissues [8] so no single mutation rate can be used in all It is commonly thought that modern medicine and
calculations. Having said this, an average nuclear mutation technology have all but ceased natural selection in modern
rate per generation for modern humans has been estimated at life, but this is not the case; selection continues, especially
1.3 10 8 per generation [9, p. 4]; as mentioned, such a figure where access to hygene and nutrients are not guaranteed.
is used to help calculate figures that will be introduced below. In the developed world there is also selection as in the
Compared to on Earth mutation by increased exposure cases of increased rickets among recent migrants from the
to cosmic radiation will likely increase during interstellar Indian subcontinent to the cloudy British Isles [17] and
voyaging, but will presumably be eased again when a synthetic compounds found in the uterus and fetus,
planet with a magnetic field serves as a shield in as on undoubtedly affecting the genome even early in life,
Earth. How much mutation will occur as a result of cosmic selection that we are only beginning to understand [18].
radiation is a topic for a separate paper, but current While interstellar migration populations will inhabit
estimates do not suggest that it clearly precludes multi- carefully constructed environments and will have superior
generational human interstellar travel [10]. As with pro- health care, it remains possible that new and unexpected
pulsion issues, for the purposes of this paper I am conditions related to interstellar travel (e.g. non-Earth-
assuming radiation hazards will be suitably mitigated. normal gas atmospheres and pressures, as well as gravita-
tion, in interstellar craft) could cause selection on the
2.2. Migration human population, especially in gastrulation, early in life
when things are quite fragile. Also, when an exoplanet is
Migration is a change in the genetic properties of a reached and settled, no matter what technology is used to
population as a result of individuals physically moving into adapt, some new selection will surely shape the human
C.M. Smith / Acta Astronautica 97 (2014) 16–29 19
150 1 30 49.83
Drift drives genetic change by a sampling phenomenon 150 2 60 49.67
unrelated to selection; it reflects a degree of randomness 150 3 90 49.50
affecting the likelihood that genes are propagated and is best 150 4 120 49.34
150 5 150 49.17
understood in the following example: “Imagine an organism
whose genotype is such that it expects, on average, to have 1000 1 30 49.98
1000 2 60 49.95
2.4 offspring. In fact, it might have no offspring, or one or two
1000 3 90 49.93
or three or more offspring. The outcome is simply the luck 1000 4 120 49.90
of the draw. Similarly, imagine an allele that never affects 1000 5 150 49.88
fitness, and that is present in 100 copies. In the next
2000 1 30 49.99
generation, maybe there will be only 90 copies, or 120 copies. 2000 2 60 49.98
The actual number is drawn from some probability distribu- 2000 3 90 49.96
tion with a mean value of 100” [20, p. 704]. Over time, then, 2000 4 120 49.95
2000 5 150 49.94
simply due to random sampling error, gene frequencies
change even absent strong selection. Drift is most active in
small populations that do not grow, and clearly a concern
in modeling multigenerational voyaging with a rather fixed below) and t¼elapsed generations. Table 1 shows that on
population over some generations. the five-generation timescale considered in project Hyper-
The principal effect of drift in closed populations can be ion, heterozygosity (beginning at 50%) would drop to
estimated by modeling the loss of heterozygosity, which is 44.05% in five generations in a population of 20 indivi-
a measure of the fact that in many genetic maladies, two duals; this six percent increase in homozygosity is roughly
copies of a deleterious gene (one from each parent and three times the amount allowed by animal breeders, who
each variant of the single gene called an allele) must be are familiar with the issues of inbreeding (an increase in
present for the deleterious effect to be expressed in homozygosity due to shared ancestry), and would be
an individual. For the individual organism, heterozygosity deleterious to the population. In populations even slightly
is a statistical concern regarding the probability that an larger, however, only one or less than one percent of
individual carries two different forms of a gene. For heterozygosity is lost in five generations; these results
populations, heterozygosity can also refer to the amount confirm O0 Rourke0 s estimation that from the perspective of
of allele diversity in the gene pool; for example, if hetero- heterozygotic decay and voyages on the order of 5–8
zygosity is 0.5, half of the alleles of a certain gene are generations in duration, populations over 150 or so indi-
deleterious and half not; this is an average for many viduals are entirely sufficient [24]. This is also confirmed in
deleterious genes in a human population. As a closed Strickberger0 s estimation that (under realistic conditions) a
population that does not substantially increase in size population of 1000 sexually reproducing life forms would
continues to inbreed (as on a starship), the statistics of take several hundred generations to lose just 10% of their
mate-selection dictate that the number of heterozygotes original heterozygosity [25, pp. 711–713].
decreases, increasing homozygosity, or the chance that For these reasons we can say that any population over
individuals will carry two copies of the deleterious allele 100 or so will be sufficient to avoid drift-related inbreed-
(one from each parent), leading to expression of such ing effects, including heterozygotic decay, on multigenera-
genetic maladies. Often seen in cases of incest, this results tional voyages in which population does not significantly
in the concentration of deleterious genes and reduced increase. Fig. 2 reiterates this point graphically, showing
overall health of the population; close incest, for example, that the drift-related loss of heterogeneity should be lower
causes mental disorders [21,22] and one study has shown than 5% – even in roughly double the time of any voyage
that first-cousin matings among Amish, Indian, French, currently conceived – for Ne populations of 150 or more,
Japanese, Pakistani, Swedish and Utah populations pro- though populations below this (e.g. 20) lose heterogeneity
duced up to 22% infant mortality, roughly double the much more rapidly.
percentage seen in offspring of unrelated individuals [23].
The reduction of heterozygosity in a population is 2.5. The founder effect
steady and can be determined from the equation:
The “Founder Effect” is the phenomenon in which
Ht ¼ H0ð1 1=2N e Þt
a subpopulation migrates away from the local origin,
where Ht¼heterozygosity after t generations, H0 is the establishing a new population that no longer interbreeds
initial heterozygosity, Ne ¼ Effective Population (discussed with members of the original population such that all
20 C.M. Smith / Acta Astronautica 97 (2014) 16–29
descendant characteristics will be strongly conditioned by Mutation and migration essentially introduce variation
the founding population only, which might not be a into gene pools, while selection and drift erode variation
statistically representative sample of the original popula- (there are exceptions to these generalization). The trick to
tion and will have its own unique characteristics, that will maintaining good genetic health is to at least balance these
further diverge from the donor population over time [26]. processes. In nonhuman populations this is largely done by
Clearly, the Founder Effect is of critical importance to instinct, whereas in humans, a small portion of instinct is
considering interstellar colonial populations because they elaborated by overriding cultural norms regulating repro-
will be a subset of the entire human gene pool and will be ductive activity. A review of the essential processes of
reproductively isolated from the human gene pool the population genetics suggests that for the timeframe dis-
moment they move away from Earth (using radio trans- cussed in this paper, selection, drift, mutation and migra-
missions to update genetic material aboard a starship, and/ tion are not as significant as population size, as reflected in
or using sperm and egg banks are both hypotheticals I do founder population demography. Long-term survival num-
not discuss here). The initial, founding genetic character- bers large enough that catastrophes can be sustained
istics of colony population, then, will have to be carefully without entering an extinction vortex. This is most clearly
considered to sample as much of Earth0 s human diversity addressed by ensuring that that the founding population is
as possible; the number of people required to reflect this large enough to persist over mutigenerational time.
diversity is a figure important to this study, and is returned
to below. 3. Estimating human population sizes for interstellar
craft
2.6. Inbreeding, Allee Effect and the extinction vortex
3.1. Quantifying population issues for interstellar voyaging:
Several other population genetic issues must be Effective Population (Ne), Minimum Viable Population (MVP)
addressed when considering an initial population of and census population (Nc)
humans for multigenerational health (two such population
figures, Ne or “Effective Population” and MCP or ‘Minimum Since Sewall Wright first explored specific population
Viable Population’, are discussed at length below). Breed- issues in his 1931 publication of “Evolution in Mendelian
ing among small populations has the effect of increasing Populations”, three figures (discussed below) have been
the frequency of deleterious genes in the population, devised to better undersand these issues. Importantly,
popularly known as inbreeding [27,28]. Thus, breeding these figures were largely theoretically considered until
population size and inbreeding concerns are directly the 1970s, when awareness of modern extinction rates in
C.M. Smith / Acta Astronautica 97 (2014) 16–29 21
wild populations led to the rise of the field of conservation are promoted), issues returned to below [6, p. 147]. Note
biology. Among this field0 s many concerns has been to that as humans are particularly active in making long-
identify the number of individuals, of a given species, distance mating networks, and are highly mobile, the
required to maintain that population in good genetic human Ne is well below the actual population of roughly
health for some time (such figures help to determine the six billion. Recent profiling of human genetic diversity has
size of wildlife refuges, for instance). greatly improved our understanding of the human Ne
One of these figures is Effective Population, or Ne. This [40,41].
figure, defined somewhat variably in the literature, The second figure, Minimum Viable Population (MVP)
broadly estimates the minimum number of individuals of includes all members of the population, whether or not they
a species that can interbreed while producing viable off- are members of the reproductive age cohort [42,43]. While
spring for some generations without suffering the effects Ne is often calculated based on surveys of the genetic
of inbreeding [25, p. 711]; another way to think of it is as diversity of species, MVP is normally a result of surveys of
an ideal figure representing the number of individuals of a the actual demographic structure (in terms of age and sex
species that demonstrate the same amount of genetic subdivisions or cohorts) of real-world populations of a
variation as the entire population, or, alternately “…the species. The Minimum Viable Population is always larger
number of individuals in an ideally behaving random- than Ne and depends on many variables including the
mating population that has the same magnitude of genetic species0 reproductive rate, gestation time, longevity per
drift as the actual population of interest” [32,8] (note that individual and so on [44]. Because MVP literature empha-
in all cases the sampled population is assumed to be sizes the minimum number of individuals to support endan-
genetically healthy on a multigenerational scale). Ne is gered nonhuman population for some generations (often a
normally lower than the actual number of organisms in a century, a figure of importance to humans attempting to
population because Ne only includes reproductively viable scale conservation efforts with human lifetimes and scales
individuals (that is, sexually mature but not senescent). of political attention), I prefer to use the third term, the
Ne (an harmonic rather than arithmetic mean) varies by theoretically rather synonymous Nc, or census population;
the reproductive biology of the species in consideration, this is similar to MVP but in its use does not emphasize the
the species0 individual average longevity and other factors minimum possible number, but rather a reasonable number,
(see [33] for cultural reasons for variation in the human Ne). to sustain the population over a given interval. Census
For many species, though, Effective Population ranges population, then, or Nc (which is often synonymous with
in the low thousands, and “…from bacteria to humans MVP in conservation biology literature), is strongly condi-
[Ne figures] are within an order of magnitude of each other.” tioned by Ne and is often expressed as the n of reproductive
[34, p. 2161, 35, p. 314]. In modern humans, who number individuals plus all nonreproductive members of the breed-
into the billions, Ne is often calculated to be around 10,000, a ing population [45]. In human populations, Nc is often twice
figure we will return to below. Ne is calculated in many the size of Ne in small, foraging-hunting-gathering popula-
subtly varying ways, but can be largely thought of as a tions (where populations are deliberately kept low to avoid
function dividing the amount of genetic variation in a sample exhausting limited resources), and three times the size of Ne
of a species by an estimate of the mutation rate and in agricultural populations (such as those of modern
population of that species. Also note that Ne is always an civilization supplied by agriculture, as will be the case on
estimate, as it differs over time as the demographic history interstellar craft).
changes of a species changes, and, further, that different Ne In summary, Effective Population (Ne) is the minimum
figures can be calculated for different genes or other traits of number of reproductive individuals required to ensure
a species. genetic health over multiple generations (in our case, five),
For all of these reasons Ne is always an estimate but so it must be large enough in the population of the first
remains a genetically and realistically well-informed mea- interstellar colonist generation to sample a wide variety of
sure of the of the genetic diversity of a species, which itself the diversity of the human genome, because genetic diver-
broadly indicates how many reproductive individuals are sity is key to the genetic health of a population over time.
needed, at minimum, to maintain species genetic health From Ne we can derive Census Population (Nc), the actual
(largely by avoiding the effects of inbreeding and loss of number of a real-world human population – distributed
heterozygosity) over multigenerational time [36,37, p. 2199, among a modern human sex- and age-cohort demographic
38, p. 325], clearly a figure of interest in the topics of structure – of the first generation of a Interstellar Migrant
this paper. Population (IMP), including reproductive and nonreproduc-
In nonhuman life Ne is often only 10–30% of the tive individuals. This figure should be selected to be sufficient
population [39], with nonreproductive young normally to avoid inbreeding and genetic drift effects according to the
composing the bulk of the remainder, and only a small duration of the projected voyage and its unique properties
percentage of nonreproductive elderly composing the final (compared to Earth conditions) of (a) low-to-no population
fraction (this fraction is normally higher in highly social growth and (b) relatively small breeding pool with no
animals, such as primates, where elders may provide opportunity for in-migration of new genes during its voyage.
protection or alloparenting for the young). For a number In this study I use five generations, each of 30 years
of reasons, Ne is normally 50% of the local breeding [46, pp. 420–422], for a total of 150 years. I adjust population
population in traditional human foraging cultures (where figures to account for various likelihoods of large-scale
low populations are promoted) and only about 30% of that disaster, which we can reasonably expect – or should at
population in farming cultures (where high populations least prudently model for – on such a voyage.
22 C.M. Smith / Acta Astronautica 97 (2014) 16–29
3.2. Survey of Effective Population (Ne), Minimum Viable studied (“Comp Sim” indicates a computer study only)
Population (MVP) and census population (Nc) Estimates and references. Many of these studies derived such figures
for various life Forms from DNA diversity analyses, while PVA (Population Viability
Analysis; see [44]) was often used in conjunction with
Before proceeding we must remember that Ne, Nc and demographic and population structure data to arrive at Nc,
MVP figures are calculated differently for different life forms, synonymous here with MVP as described above.
and there is active debate among scholars regarding methods Eight summary results are immediately apparent, some
to derive these figures. While none are calling to abandon tabulated for clarity in Table 3.
calculation of these figures, it is clear that there are no “magic
numbers” that apply to all species [47, p. 879], see also 1. There is no single universally applicable Ne or Nc; this is
[35,33]; at the same time, useful estimates can be made for expected because these figures are derived with
conservation efforts [48–50] analogous to the interests of slightly differing methods per study, for life forms with
project Hyperion0 s goal of sending viable human populations different life histories, reproductive biologies and life-
on intergenerational voyages. One reason for the variation in spans and because these figures themselves change
calculating these figures is that subpopulations of a single through time [53,27].
species vary genetically, such that they may have a different 2. Almost no study sets Ne below 1500, often setting
Ne than another sub-population; also, one variable used to minimal Nc/MVP0 s of at least 5000 in nonhumans (Reed
calculate Ne is the rate of mutation, that is the rate of new et al. [44, 30] suggest populations “for long-term persis-
variation in the life form per generation; but this also varies tence may range from 2000… to 413,000”) and 43,000
both among and within populations [9] and over time [51]; for modern humans; both, again, in the multiple thou-
finally, the Ne might itself have changed through time, as is sands rather than hundreds, for example, and in only one
suspected of the genus Homo [32], though it is still possible to of 61 cases below 100,000, setting the range between 103
make useful ballpark estimates. and 104 in many cases. The exception [54] is a highly
To estimate a genetically viable Ne and Nc for a five- abstracted model of Pleistocene hunter-gatherer popula-
generation (c. 150-year) timespan, I surveyed the research tions, which I discount from this analysis.
literature pertaining to these values published in the last 40 3. Ne figures average just under 20,000 for all life forms
years, highlighting research (including meta-analyses) pub- and computer simulations combined, but significantly
lished in the last decade to account for new genomic knowl- lower, just over 1500, for a few studies generalizing
edge that is reformulating evolutionary biology see [52]. upon “all animal life”. For hominoidea generally and
I focused on animal life (rather than including plants or hominins specifically, data derived largely from DNA
microbes) and, in particular, vertebrate life, and of the diversity studies indicates an Ne on the order of 17,000–
vertebrates, mammals, and of the mammals, primates, includ- 20,000, while for modern humans an Ne of just over
ing earlier and modern Homo. Such estimates include recent 10,000 is indicated by many studies, with attendant
studies using genomic data that have allowed high-resolution Nc0 s of roughly 20,000 if modeling after historical
understanding of modern humanity0 s Ne. Such estimates also foraging populations (who prefer smaller populations)
vary somewhat in their calculation of Ne for both methodo- or closer to 30,000 if modeling after historical pre-
logical reasons and because humanity deviates in several industrial agricultural populations (who prefer larger
ways from the Hardy–Weinberg equilibrium conditions that populations).
are the theoretical foundation of Ne estimates; for example, 4. Only 13 of 61 studies yielded Nc/MVP figures for the
humans exhibit (a) overlapping generations, (b) population sampled life forms under 5000.
subdivisions, (c) local population extinctions, recolonizations 5. Ne for large mammalian species only twice exceeds
and migrations and (d) different reproductive patterns 100,000 (this is also noted in a smaller sampling in
between the sexes and cultures. The variation in the figures Refs. [32,15]).
examined here, and the ways in which they are calculated by 6. Ne and Nc average (respectively) around 18,000 and
different researchers, are the reason for my not performing a 32,000 for the entire sample (n ¼61 studies), 14,500–
standard ANOVA on the data set presented in Table 2. How- 22,000 for all hominin estimates (n ¼34 studies) and
ever, there are significant patterns in the data set that are of 11,000–22,000 for all modern human estimates (n ¼18
use to the estimates sought in this paper, which are required studies, of which nearly half suggest an Ne approximat-
as estimates of magnitude regarding the scale of operation ing 10,000).
for interstellar voyaging. We see that demographically, in 7. The metastudies [44,55,47] all estimate Nc on the order
humans, scales over 150 are advisable from one perspective, of several thousand, ranging from around 4000 for
and we shall see that from other independent perspectives vertebrates and mammals specifically to 5000–7000
(e.g. Ne, Nc and associated figures) similar quanta are indicated, for samples of hundreds animal species from multiple
giving us a magnitude-level understanding of realistic figures genera ([44] studied 102 vertebrates while [55] studied
for multigenerational, closed-population voyaging. 212 varied species).
Note that human Ne figures vary in a rather tight range, 8. An important result not obvious on the table is that
as will be seen below, when we consider the range of in these meta-analyses, an Nc on the order of several
variation among all organisms sampled in this study. thousand ( 41500, o8000) was not found to vary
Results of the survey are found in Table 2. They include significantly by taxa (life form), trophic level
low and high range estimates for Ne and Nc (representing (e.g. carnivore or herbivore), or latitude (geographical
MVP), averages of these figures, an indication of the life form distribution) [44].
C.M. Smith / Acta Astronautica 97 (2014) 16–29 23
Table 2
Survey of effective and census population data.
While there is some significant variation in Ne and Nc the average Ne of roughly 10,000 individuals closely approx-
estimates for modern humans at large (including purely imates Relethford0 s similar conclusion, based on his own
theoretical modeling of ancient populations as well as survey of others0 estimates and his own mathematical esti-
empirically based estimates using genomic diversity data), mates [6]. This figure is high compared to other interstellar
24 C.M. Smith / Acta Astronautica 97 (2014) 16–29
voyaging population estimates, such as Moore0 s widely an ancient state of small, dispersed human populations, into
discussed population of 80–150 individuals (therefore an Ne the hundreds of thousands of years ago, and thus often
of roughly 40–75) [56]. Such small figures are based on considered a stable reproductive units over long time for
anthropological observation that in many preindustrial members of the genus Homo (e.g. [54]).
human cultures, tribal or band populations of individuals There is a significant problem with these figures,
are often of this scale – the “Dunbar Number” of c. 150 has however, for multigenerational interstellar voyaging. Even
also widely been publicized as a viable, self-contained though human groups might have fallen into such low-
human population. Such low-hundreds to low-thousands hundreds figures for social (e.g. ethnic) and logistical
figures of “natural human populations”, when observed in reasons, they are (and in the past, were) never completely
modern non-industrialized cultures are often taken to reflect isolated populations, as would be the case on a starship.
Human cultures have long-range mating networks and
maintain large breeding populations even if they reside
Table 3
Summary of population data surveyed.
in smaller subunits observed by anthropologists [57];
Eckhard notes that even among historically known fora-
Average Ne MVP ging peoples, though they lived and traveled in such small
groups, a larger collective gene pool (Ne) was maintained
All studies 18,187 31,810
by “matings between members of immediately adjacent
Animal 1638
Comp Sim 7813 populations…[and modern foragers] have systematic (and
Hominoidea 19,600 often complex) arrangements for mate exchange among
Insect 6521 adjacent bands, either to make up for a shortage of mates
Large vertebrate 5137
of appropriate demographic characteristics or to ensure
Later hominin 17,819 51,075
Mammal 3688
amicable relationships, or for other reasons” [58, p. 243].
Modern human 11,168 21,964 Gamble emphasizes such wide, integrating social and
Plant 4196 breeding networks as particularly distinctive of the mod-
Small vertebrate 3956 ern Homo sapiens sapiens as opposed to the Neanderthals
Various primates 49,000 83,846
(Homo sapiens neanderthalensis), who lived in smaller
Vertebrate 5090
All hominins 14,494 21,888 territories and might well have had smaller breeding
populations [59]. While some human populations have
Table 4
Effective and census populations and adjustments for safety considering various catastrophe probabilities and 30% mortality on interstellar vessels.
75 150 1.0 0.3 45 390 585 105 53 273 410 137 205
75 150 0.5 0.3 23 345 518 128 64 242 362 121 181
75 150 0.3 0.3 11 323 484 139 69 226 339 113 169
75 150 0.1 0.3 6 311 467 144 72 218 327 109 163
75 150 0.1 0.3 3 306 458 147 74 214 321 107 160
75 150 0.0 0.3 1 303 454 149 74 212 318 106 159
150 300 1.0 0.3 90 780 1170 210 105 546 819 273 410
150 300 0.5 0.3 45 690 1035 255 128 483 725 242 362
150 300 0.3 0.3 23 645 968 278 139 452 677 226 339
150 300 0.1 0.3 11 623 934 289 144 436 654 218 327
150 300 0.1 0.3 6 611 917 294 147 428 642 214 321
150 300 0.0 0.3 3 606 908 297 149 424 636 212 318
2000 4000 1.0 0.3 1200 10,400 15,600 2800 1400 7280 10,920 3640 5460
2000 4000 0.5 0.3 600 9200 13,800 3400 1700 6440 9660 3220 4830
2000 4000 0.3 0.3 300 8600 12,900 3700 1850 6020 9030 3010 4515
2000 4000 0.1 0.3 150 8300 12,450 3850 1925 5810 8715 2905 4358
2000 4000 0.1 0.3 75 8150 12,225 3925 1963 5705 8558 2853 4279
2000 4000 0.0 0.3 38 8075 12,113 3963 1981 5653 8479 2826 4239
3500 7000 1.0 0.3 2100 18,200 27,300 4900 2450 12,740 19,110 6370 9555
3500 7000 0.5 0.3 1050 16,100 24,150 5950 2975 11,270 16,905 5635 8453
3500 7000 0.3 0.3 525 15,050 22,575 6475 3238 10,535 15,803 5268 7901
3500 7000 0.1 0.3 263 14,525 21,788 6738 3369 10,168 15,251 5084 7626
3500 7000 0.1 0.3 131 14,263 21,394 6869 3434 9984 14,976 4992 7488
3500 7000 0.0 0.3 66 14,131 21,197 6934 3467 9892 14,838 4946 7419
11,000 22,000 1.0 0.3 6600 57,200 85,800 15,400 7700 40,040 60,060 20,020 30,030
11,000 22,000 0.5 0.3 3300 50,600 75,900 18,700 9350 35,420 53,130 17,710 26,565
11,000 22,000 0.3 0.3 1650 47,300 70,950 20,350 10,175 33,110 49,665 16,555 24,833
11,000 22,000 0.1 0.3 825 45,650 68,475 21,175 10,588 31,955 47,933 15,978 23,966
11,000 22,000 0.1 0.3 413 44,825 67,238 21,588 10,794 31,378 47,066 15,689 23,533
11,000 22,000 0.0 0.3 206 44,413 66,619 21,794 10,897 31,089 46,633 15,544 23,317
C.M. Smith / Acta Astronautica 97 (2014) 16–29 25
been very low for some centuries (e.g. castaway island row to model different contingencies.
populations; see [60,61]), these are anomalous situations L¼lethality of catastrophe¼percentage of Nc killed in a
in which the population size and social structure were not catastrophic disease sweep on the order of the Eur-
the product of long cultural and natural selection, but opean plague of the 14th Century AD; here I use
simply an historical contingency. I suggest rather that we 0.30 (30%).
opt for population sizes that have been the product of both CF ¼Catastrophe Factor¼ (Nc L)P ¼n of individuals to
cultural and biological evolution in our genus; such supplement the starship population, to offset cata-
populations, far exceeding the low hundreds, have so far strophe, based on the expected lethality of the disaster
provided longevity for our genome over many generations and one0 s comfort regarding modeling the likelihood of
– as many as 3000 if we estimate the origins of modern a disaster
humanity around 100,000 years ago. IMP, SF ¼ Interstellar Migrant Population with Safety
For all of these reasons, low-hundreds figures often Factor¼ (Nc þCF) [2 or 3]¼ the original population
proposed as populations for multigenerational interstellar (Nc) plus the number of replacements carried in event
voyages might work in theory, but likely underestimate of a catastrophe (CF), multiplied by two or three,
the actual size of a human gene pool required to maintain according to one0 s threshold for over-engineering.
good multigenerational health in absence of reproductive These two columns represent the total number of
links to other populations. Rather, I feel the Ne of roughly people (Nc) as the starship population, adjusted for
10,000 or 11,000 – reflecting and concurring with high- various probabilities of disaster; note that they always
resolution modern data on living populations as well as exceed Ne.
theoretical calculations – is a much more reasonable Nc-Cat ¼for reference, the Nc minus 30% as in the case
estimate. For these reasons, I proceed below using these of catastrophe.
more realistic population estimates based on the results of Ne-Cat ¼for reference, the Ne minus 30% as in the case
the survey reported in Table 2. The following investigation of catastrophe.
is heuristic, but not completely abstract: it is useful for The remaining four columns (on the right side of the
identifying, at least to an order of magnitude, reasonable table) are most useful for evaluating whether, after
estimates for the population issues of this paper. catastrophe, certain critical population size limits are
violated (discussed below).
NcSF2-CAT¼ the Nc multiplied by safety factor of 2,
3.3. Estimating and modeling a viable population for minus 30%, the population lost in a catastrophe. This
a five-generation interstellar voyage is the n of people remaining after a 30% depopulation if
the original population were doubled as a safety factor.
Table 4 can be used to evaluate the suitability of certain NcSF2-CAT¼ the Nc multiplied by safety factor of 3,
Ne and Nc figures for human populations on intergenera- minus 30%, the population lost in a catastrophe. This
tional, closed-system voyages. The columns indicate various is the n of people remaining after a 30% depopulation if
quantities described below. One factor built into the model- the original population were tripled as a safety factor.
ing is that of catastrophe. An undertaking as complex and NeSF2-CAT ¼the Nc multiplied by safety factor of 2,
long in duration as a 5-generation voyage to an exoplanet minus 30%, the population lost in a catastrophe. This
will undoubtedly be over-engineered in its lifeless compo- is the n of reproductively active people remaining after
nents, as are aircraft and cars today. The human population – a 30% depopulation if the original population were
the precious cargo – of such voyages should also be designed doubled as a safety factor.
to be resilient to shocks. Aside from mechanical failure in the NeSF2-CAT ¼the Ne multiplied by safety factor of 3,
habitats themselves, the most likely large-scale catastrophe minus 30%, the population lost in a catastrophe. This
that could occur in the envisioned voyage would be the is the n of reproductively active people remaining after
sweep of a plague through the population. Even the Earth0 s a 30% depopulation if the original population were
worst recorded human plagues have never had a mortality tripled as a safety factor.
greater than 2% of the entire human population, but in
smaller subpopulations – such as Europe during the Black
Plague of the 1300s – average mortality was as high as 30%
[62]. I use this mortality figure in planning for disaster in the 4. Results and discussion
calculations below.
The columns in Table 4 are arranged as follows, from 4.1. Overview
left to right;
Three lessons learned from the analysis of Table 2
Ne ¼ Effective Population ¼(n of reproductive indivi- are particularly important to understanding Table 3:
duals necessary to sustain genetic health while breed-
ing among the closed population). 1. Viability over several generations is rarely considered
Nc ¼census population (2Ne)¼total number of people possible in numbers (Ne or Nc) less than a few thou-
in the population, including reproductive and nonre- sand. While an Ne of 500 has been used for many
productive members. species in conservation efforts, it is not a “magic
P ¼probability of catastrophe on 5-generation voyage, number” Still, attendant Nc0 s normally measure in the
from 1.0 (100%) to.03 (3%), dropping somewhat in each low thousands at least, a figure acceptable to even
26 C.M. Smith / Acta Astronautica 97 (2014) 16–29
scholars most critical of population viability estimates about 10,000, while they might preserve Nc, are risky in
[35, p. 314]. the event of disaster regarding maintenance of Ne. For
2. A cautious approach does not allow Nc to dip below these reasons I discount an initial Ne of 2000 as too risky
about 7000 individuals; this is particularly important in for multigenerational success.
vertebrates [44]. With the initial Ne set at 3500 the recommendation
3. A cautious approach does not allow Ne to dip below that vertebrates in general have an Nc of about 7000 is met
about 10,000; this is particularly important in the [44]. In the column IMP, SF¼2 we see that any figure, from
genus Homo [6,63]. 18,200 to 14,131 (Nc0 s doubled) allows the roughly 7000
individuals thought to be a viable multigenerational verte-
brate population to flourish, even in the case of cata-
With these lessons in mind in Table 5 I review Ne and strophes, where we see 9800 to 19,110 individuals
associated figures of 75, 150, 2000, 3500 and 11,000. surviving. To preserve Ne around 10,000 after a cata-
With Ne (reproductive population) at 75 persons for an strophe (cell far right indicating 9555 survivors) can be
Nc (total population) of 150 persons – the oft-cited done by tripling the population (column IMP, SF¼3) for a
“Dunbar Number” – even doubling or tripling the Nc with founding population of Nc 27,300. This is a reasonable
safety factors never brings populations into even the low starting figure for an IMP.
thousands for either Ne or Nc. These populations would With the initial Ne set at 11,000 the average living
always be on the verge of disaster in the case of a Black- human Ne (as reported in Table 2) is approximated. With
plague-like catastrophe. The same applies to the estimates Nc0 s adjusted to accommodate catastrophe, ranging from a
with Ne at 150, giving an initial Nc of 300 persons, which is low of 44,413 to a high of 57,200 when adjusted for
little better; and in both cases the Ne, even after doubling catastrophe, recommendations for Nc0 s over c.5000 and
or tripling, is just at or even below 10% of the Ne of 10,000 Ne0 s around 11,000 after catastrophe are easily met by
that is apparently a good guide for human health. Though doubling the population (columns NcSF2-CAT and NeSF2-
these populations would not likely suffer from significant CAT). Tripling the population (column IMP, SF¼ 3) invokes
inbreeding effects (as discussed in Section 2.4) they are populations from nearly 70,000 to over 85,000, which
nevertheless demographically dangerous. For these rea- appears to be overkill as just mentioned; doubling is
sons I discount both of these founding populations (75 and sufficient to protect both Nc and Ne in a severe catastrophe.
150) as too risky to ensure multigenerational success. In sum, I would consider roughly 10,000 an absolute
With the initial Ne set at 2000 the recommendation minimum Ne population, with an Nc at or exceeding 30,000
that vertebrates over 1 kg body mass have an Nc of about for a safe IMP; this aligns well with the currently known
4000 [55] is met. In the first row we see that by adding modern human Ne, which ranges from 9000–17,000,
1200 individuals against the effects of a catastrophe, and averaging somewhere in-between [40, p. 1963], and would
then doubling the population, we achieve a population suffice to ensure enduring the modeled catastrophe.
(Nc) of 10,400; if this population were to be struck with
catastrophe of 30% lethality, 7820 individuals would sur-
vive, fulfilling the general guidance to keep the Nc at or 4.2. A note on demographic structure
above 10,400. Looking at the columns for Ne after cata-
strophe we see figures from a high of 5460 to a low of The demographic (age and sex) structure of such
2826; only one approaches even 50% of the recommended populations as explored above will be important to
human Ne, and most are far below it. For this reason,
founding populations on the order of anything under
Table 5 2000
Sample demographic structures for an interstellar vessel at various
founding populations.
1600
Population
[3] C.M. Smith, E.T. Davies, Emigrating Beyond Earth: Human Adapta- [35] F.H. Flather, G.D. Hayward, S.R. Beissinger, P.A. Stephens, Minimum
tion and Space Colonization, Springer, New York, 2012. viable populations: is there a ‘magic number0 for conservation
[4] E.C. Friedberg, Mutation as a phenotype, in: L.H. Caproale (Ed.), practicioners? Trends Ecol. Evol. 26 (6) (2011) 307–316.
The Implicit Genome, Oxford University Press, Oxford, 2006, [36] A. Caballero, Development of the prediction of effective population
pp. 39–56. size, Heredity 73 (1994) 657–679.
[5] M. Lynch, M. Rate, Molecular spectrum, and consequences of human [37] L.B. Jorde, W.S. Watkins, M.J. Bamshad, Population genomics: a
mutation, Proc. Natl. Acad. Sci. U.S.A. 107 (3) (2010) 961–968. bridge from evolutionary history to genetic medicine, Hum. Mol.
[6] J.H. Relethford, Genetics and the Search for Modern Human Origins, Genet. 10 (20) (2001) 2199–2207.
Wiley-Liss, New York, 2001. [38] N. Ryman, L. Lairke, Effects of supportive breeding on the gene-
[7] S. Matsamura, P. Forster, Generation time and effective population tically effective population size, Conserv. Biol. 5 (4) (1991) 325–329.
size in polar eskimos, Proc. R. Soc. B 275 (2008) 1501–1508. (1991).
[8] L. Duret, Mutation patterns in the human genome: more variable [39] R. Frankham, Effective population size/adult population size ratios
than expected, PLOS Biol. 7 (2) (2009) 217–219. in wildlife: a review, Genet. Res. 66 (1995) 95–107.
[9] E.M. Leffler, K. Bullaughey, D. Matute, W.K. Meyer, L. Segruel, [40] S. Harpending, M.A. Batzer, M. Gurven, L.B. Jorde, A.R. Rogers,
A. Venkat, P. Andolfatto, M. Przeworski, Revisting and old riddle: S.T. Sherry, Genetic traces of ancient demography, Proc. Natl. Acad.
what determines genetic diversity levels within species? PLOS Biol. Sci. U.S.A. 1998 (1995) 1961–1967.
10 (9) (2012) 1–9. [41] B.F. Voight, A.M. Adams, L.A. Frisse, Y. Qian, R.R. Hudson, A. Di
[10] G. Horneck, C. Mileikowski, H.J. Melosh, J.W. Wilson, F.A. Cucinotta, Rienzo, Interrogating multiple aspects of variation in a full rese-
B. Gladman, Viable transfer of microorganisms in the solar system quencing data set to infer human population size changes, Proc.
and beyond, in: G. Hornbeck, C. Baumstarck-Khan (Eds.), Astrobiol- Natl. Acad. Sci. U.S.A. 102 (51) (2005) 18508–18513.
ogy: The Quest for the Conditions of Life, Springer, New York, 2011, [42] D.H. Reed, E.H. Bryant, Experimental tests of minimum viable
pp. 57–76. population size, Anim. Conserv. 3 (2000) 7–14.
[11] E.M. Wijsman, L L Cavalti-Sforza, Migration and genetic population [43] M.L. Shaffer, Minimum population sizes for species conservation,
structure with special reference to human, Ann. Rev. Ecol. Syst. 15 Bioscience 31 (1991) 131–134.
(1984) 279–301. [44] D.H. Reed, J.J. O0 Grady, B.W. Brook, J.D. Ballou, R. Frankham, Estimates
[12] C. Gamble, Timewalkers, Harvard University Press, Cambridge, 2003. of minimum viable population sizes for vertebrates and factors
[13] J. Endler, Natural Selection in the Wild, Princeton University Press, influencing those estimates, Biol. Conserv. 113 (1) (2003) 23–34.
Princeton, NJ, 1986. [45] M.E. Soule, Viable Populations for Conservation, Cambridge Univer-
[14] D.L. Hull, R. Langman, S. Glenn, A general account of selection: sity Press, Cambridge, 1987.
biology, immunology and behavior, Behav. Brain Sci. 24 (2001) [46] J.N. Fenner, Cross-cultural estimation of the human generation
(2001) 511–528. interval for use in genetics-based population divergence studies,
[15] W. Durham, Coevolution: Genes, Culture and Human Diversity, 1st Am. J. Phys. Anthropol. 128 (2) (2005) 415–423.
ed. Stanford University Press, Stanford, 1993. [47] R. Lande, Mutation and conservation, Conserv. Biol. 9 (4) (1995)
[16] C.M. Smith, Comment on ‘an evolutionary framework for cultural 782–791.
change: selectionism versus communal exchange0 , Phys. Life Rev. 10 [48] U.K. Rai, Minimum sizes for viable population and conservation
(2013) 156–167. biology, Our Nat. 1 (2003) 3–9.
[17] J.B. Henderson, The importance of limited exposure to ultraviolet [49] C.D. Thomas, What do real population dynamics tell us about mini-
radiation and dietary factors in the etiology of Asian rickets: a risk- mum viable population sizes? Conserv. Biol. 4 (3) (1990) 324–327.
factor model, Q. J. Med. 63 (1987) 413–442. [50] M. Lynch, R. Lande., The critical effective size for a genetically secure
[18] J.G. Dorea, C.M. Donangelo, Early (in uterus and infant) exposure to population, Anim. Conserv. 1 (1) (1998) 70–72.
mercury and lead, Clin. Nutr. 25 (2006) 369–376. [51] J. Hey, On the number of new world founders: a population genetic
[19] J. Hawks, E.T. Wang, G.M. Cochran, H.C. Harpending, R.K. Moyzis, portrait of the peopling of the Americas, PLOS Biol. 3 (6) (2005)
Recent acceleration of human adaptive evolution. Proc. Natl. Acad. 0965–0975(New York, Springer (2002).
Sci. U.S.A. 104(52):20753–20758. [52] C.M. Smith, J. Ruppell, What anthropologists should know about the
[20] J. Masel, Rethinking Hardy–Weinberg and genetic drift in under- new evolutionary synthesis, Struct. Dyn. 5 (2) (2011) 1–13.
graduate biology, BioEssays 34 (2012) 701–710. [53] B. Charlesworth, Effective population size and patterns of molecular
[21] M.S. Adams, J.V. Neel, Children of incest, Pediatrics 40 (1) (1967) 55–62. evolution and variation, Nat. Rev. Gen. 10 (2009) 195–205.
[22] J. Jancar, S.J. Johnston, Incest and mental handicap, J. Intellectual Dis. [54] H.M. Wobst, Boundary conditions for palaeolithic social systems: a
Res. 34 (6) (2008) 483–490. simulation approach, Am. Antiq. 39 (1973) 303–309.
[23] L.B. Jorde, J.C. Carey, M.J. Bamshad, R.L. White., Medical Genetics, [55] L.W. Traill, C.J.A. Bradshaw, B.W. Brook, Minimum viable population
MosbySt. Louis, 1999. size: a meta-analysis of 30 years of published estimates, Biol.
[24] D. O0 Rourke, Genetic considerations in multi-generational space Conserv. 139 (2007) 159–166.
travel, in: Y. Kondo, F. Bruhweiler, J. Moore, C. Sheffield (Eds.), [56] J.H. Moore, Kin-based crews for interstellar multi-generational space
Interstellar Travel and Multi-Generational Space Ships, Apogee travel, in: Y. Kondo, F. Bruhweiler, J. Moore, C. Sheffield (Eds.),
Books, Wheaton, IL89–99. Interstellar Travel and Multi-Generational Space Ships, Apogee
[25] M.W. Strickberger, Genetics, Prentice Hall, New York, 1985. Books, Wheaton, IL81–88.
[26] N.H. Barton, B. Charlesworth, Genetic revolutions, founder effects [57] R.J. Kelly, The Foraging Spectrum: Diversity in Hunter-Gatherer
and speciation, Ann. Rev. Ecol. Syst. 15 (1994) 133–164. Lifeways, Eliot Warner Reprints, New York, 2007.
[27] L.J. Harmon, S. Braude., Conservation of small populations: effective [58] R.B. Eckhardt, Human Palaeobiology, Cambridge University Press,
population sizes, inbreeding, and the 50/500 rule, in: S. Braude, B. Cambridge, 2000.
S. Low (Eds.), An Introduction to Methods and Models in Ecology, [59] C. Gamble, The Palaeolithic Societies of Europe, Cambridge Univer-
Evolution, and Conservation Biology, Princeton University Press, sity Press, Cambridge, 1999.
Princeton, 125–138. [60] B. Finney and E.M. Jones. Interstellar Migration and the Human
[28] L. Scott Mills, P.E. Smouse, Demographic consequences of inbreeding Experience, Berkeley University of California Press (1985).
in remnant populations, Am. Nat. 144 (3) (1994) 412–431. [61] Y. Kondo, F. Bruhweiler, J. Moore, C. Sheffield (Eds.), Interstellar
[29] M. Kimura, T. Ohta, The average number of generations until fixation Travel and Multi-Generational Space Ships, Apogee Books, Wheaton,
of a mutant gene in a finite population, Genetics 61 (1969) 763–771. IL, 1991.
[30] R. Lande, Genetics and demography in biological conservation, [62] A.P. Galvani, M. Slatkin, Evaluating plague and smallpox as historical
Science 241 (1988) 1455–1460. selective pressures for the CCR5-Delta32 HIV-resistance allele, Proc.
[31] W.F. Fagan, E.E. Holmes, Quantifying the extinction vortex, Ecol. Lett. Natl. Acad. Sci. U.S.A. 100 (25) (2003) 15276–15279.
9 (2006) 51–60. [63] N. Takahata, Allelic geneology and human evolution, Mol. Biol. Evol.
[32] J. Hawks, K. HunleyS-H. leeM. Wolpoff, Population bottlenecks and 10 (1993) 2–22.
pleistocene human evolution, Mol. Biol. Evol. 17 (2000) 2–22. [64] S. Summerford, Colonized Interstellar Vessel: Conceptual Master
[33] J.W. Wood, The genetic demography of the gainj of papua new Planning. Internally-distributed paper available at 〈http://www.
guinea: determinants of effective population size, Am. Nat. 129 (2) icarusinterstellar.org/colonized-interstellar-vessel-conceptual-mas
(1987) 165–187. ter-planning〉, 2012.
[34] J.H. Gillespie, Is the population size of a species relevant to its [65] A. Hein, M. Pak, D. Putz, C. Buhler, P. Reiss, World ships—architectures
evolution? Evolution 55 (2001) 2161–2169. and feasibility revisited, J. Brit. Interplanet. Soc. 65 (2012) 119–133.
C.M. Smith / Acta Astronautica 97 (2014) 16–29 29
[66] J.R. Franklin, Evolutionary change in small populations, in: M.E. Soule, T. Topaloglou, E. Hubbell, E. Robinson, M. Mittmann, M.S. Morris,
B.A. Wilcox (Eds.), Conservation Biology: An Evolutionary-Ecological N. Shen, D. Kilburn, J. Rioux, C. Nusbaum, S. Rozen, T.J. Hudson,
Perspective: 135–150, Sinauer, Massachussetts, 1980, pp. 135–150. R. Lipshutz, M. Chee, E.S. Lander., Large-scale identification, map-
[67] S.T. Schultz, M. Lynch, Mutation and extinction: the role of variable ping, and genotyping of single-nucleotide polymorphisms in the
mutational effects, synergistic epistasis beneficial mutations, and human genome, Science 280 (5366) (1998) 1077–1082.
degree of outcrossing, Evolution 51 (1997) 1363–1371. [80] R.L. Dorit, H. Akashi, W. Gilbert, Absence of polymorphism at the
[68] M.C. Whitlock, Fixation of new alleles and the extinction of small AFY locus on the human Y chromosome, Science 268 (1995)
populations: drift load, beneficial alleles, and sexual selection, 1183–1185.
Evolution 54 (2000) 1855–1861. [81] M.F. Hammer, A recent common ancestry for human Y chromosome,
[69] J.R. Franklin, R. Frankham, How large must populations be to retain Nature 378 (1995) 376–378.
evolutionary potential? Anim. Conserv. 1 (1998) 69–70. [82] H. Huang, Y.X. Fu, B.H. Chang, X. Gu, L.B. Jorde, W.H. Li, Sequence
[70] N. Yu, M.I. Jensen-Seaman, L. Chemnick, O. Ryder, W-H. Li, Nucleo- variation in ZFX introns in human populations, Mol. Biol. Evol. 15 (2)
tide diversity in gorillas, Genetics 166 (2004) 1375–1383. (1998) 138–142.
[71] S.T. Sherry, A.R. Rogers, H. Harpending, H. Soodyall, T. Jenkins, [83] H. Kaesmann, V. Weibe, S. Paabo, Extensive nuclear DNA sequence
M. Stoneking., Alu evolution in human populations: using the
diversity among chimpanzees, Science 286 (1999) 1159–1162.
coalescent to estimate effective population size, Genetics 147 (1994)
[84] A. Tenesa, P. Navaro, B.J. Hayes, D.L. Duffy, G.M. Clarke, M.E. Goddard,
1977–1982.
P.M. Cisscher., Recent human effective population size estimated
[72] R.M. Harding, S.M. Fullerton, R.C. Griffiths, J. Bond, M.J. Cox,
from linkage disequilibrium, Genome Res. 17 (2007) 520–526.
J.A. Schneider, D.S. Moulin, J.B. Clegg, Archaic African and Asian
[85] F-G. Chen, W-H. Li., Genomic divergences between humans and
lineages in the genetic ancestry of modern humans, Am. J. Hum.
other hominoids and the effective population size of the common
Genet. (1997)772–789.
ancestor of humans and chimpanzees, Am. J. Hum. Genet. 68 (2)
[73] N. Takahata, Y. Satta, Footprints of intragenic recombination at HAL
locus, Immunogenetics 47 (1998) 430–441. (2001) 444–456.
[74] J. Zhao, W.J. Li, M.M. Xiong., Population-based linkage disequilibrium [86] G.A. Watterson, On the number of segregating sites in genetical
mapping of QTL: an application to simulated data in an isolated models without recombination, Theor. Popul. Biol. 7 (1975) 256–276.
population, Genet. Epidemiol. 21 (Suppl. 1) (2001) S655–S659. [87] F. Ayala, The myth of eve: molecular biology and human origins,
[75] J.D. Wall, M. Przeworski, When did the human population size start Science 270 (1995) 1930–1936.
increasing? Genetics 155 (2000) 1865–1874. [88] W.J.K. Bailey, K. Hayasaka, C.G. Skinner, S. Kehoe, L.C. Cieu,
[76] W.D. Newmark, A Land-Bridge Island, Perspective on mammalian J. Slightom, M. Goodman, Reexamination of the African hominoid
extinctions in Western North American parks, Nature 325 (1987) trichotomy with additional sequences from the primate beta-
430–432. globulin gene cluster, Mol. Phylogenet. Evol. 1 (1992) 97–135.
[77] M.E. Soule, Viable Populations for Conservation, Cambridge Univer- [89] M.F. Kinnaird, T.G. O0 Brien, Viable populations for an endangered
sity Press, Cambridge, 1987. forest primate, the Tana River crested mangabey (Cercocebus galer-
[78] M. Nei, D. Grauer, Extent of protein polymorphism and the neutral itus galeritus), Conserv. Biol. 5 (2) (1991) 203–213.
mutation theory, Evol. Biol. 27 (1984) 73–118. [90] A.H. Harcourt, Emprical estimates of minimum viable population
[79] D.G. Wang, J.-B. Fan, C.-J. Siao, P. Young, R. Sapolsky, G. Ghandour, sizes for primates: tens to tens of thousands? Anim. Conserv. 5
N. Perkins, E. Winchester, J. Spencer, L. Kruglyak, L. Stein, L. Hsie, (2002) 237–244.