Review Article

Published: 2024-02-08
https://doi.org/10.20935/AcadBiol6158

Evolution by natural selection is a scientific law and not

just a theory
Daniel J.M. Crouch1,*, Walter F. Bodmer2

Academic Editor(s): Cesar Rogério Leal do Amaral, Andre J. van Wijnen

Abstract
The concept of evolution by natural selection was developed primarily by Darwin and Wallace in the 19th century as an explanation for
the diversity and origin of complex organisms. They, however, did not have access to a mechanism of inheritance, which was needed
for a proper understanding of how evolution by natural selection could work. Mendel’s discovery of the basic laws of inheritance in
diploid sexual organisms enabled Fisher, Haldane, and Wright to provide a theoretical framework for understanding the selective
process of evolution by natural selection. The developing knowledge of prokaryotic microorganisms and the discovery of DNA or RNA
as the fundamental basis of inheritance in all living organisms, together with Mendel’s laws, now provide the four basic conditions for
evolution by natural selection, namely reproduction that is stable, but allows for variation that can increase fitness in the broadest
sense. These definitive conditions establish an exponential growth law for evolution by natural selection that applied initially for the
longest period of time to the evolution of asexual clonal organisms, and through Mendel's laws, extends to bisexual multicellular
organisms. It is mistaking evolution by natural selection to be a scientific theory rather than a law that has led to unnecessary
disagreements over its fundamental validity and explanatory power.

Keywords: natural selection, evolution, mendelian inheritance

Citation: Crouch DJM, Bodmer WF. Evolution by natural selection is a scientific law and not just a theory. Academia Biology 2024;2.
https://doi.org/10.20935/AcadBiol6158

1. Introduction and historical background

The concept of evolution by natural selection developed in the
19th century as an explanation for the diversity and origin of
complex organisms. It is widely described as a scientific theory
[1, 2], a term with various definitions, but which typically refers
to a set of logical relations whereby some basic, assumed proper-
ties of a natural system lead to predicted outcomes, which have
been verified by experiment or observation. A scientific theory
can therefore only be consistent with an increasing number of
observations or experimental results, but is never guaranteed to
be true, as there might always be another, perhaps unknown,
theory which predicts the same outcomes and others. Describing
evolution by natural selection as a theory was appropriate during
the 19th century, but the discovery of Mendelian genetics
signaled the point at which all of the essential conditions logically
entailing natural selection were confirmed. These conditions,
when present in a given environment, are as follows:
1) The entity in question has the ability to reproduce, i.e., to
produce identical or similar descendants of itself.
3) The inheritance system must not be 100% stable, allowing for
new variations of entities.
4) It is possible for new variations to exhibit increased rates of
reproduction or increased lifespan.
Mendelian genetics effectively proves the existence of Conditions
2 and 3 for biological sexual organisms. Both Charles Darwin and
Alfred Russell Wallace, the co-discoverer of evolution by natural
selection, had seen that these conditions were required for evolu-
tion to work, but they had no firm evidence that they existed, as
they did, in contrast, for Conditions 1 and 4 [3]. As all four
conditions are now known to hold with virtual certainty in the
case of all biological lifeforms, it has become appropriate to de-
scribe evolution as a scientific law, defined as the logical relation
between a set of conditions with a particular phenomenon, in this
case natural selection and adaptation, which they are jointly
sufficient to produce:
Law, n.

2) The entity and its descendants have a system of inheritance “In the sciences of observation, a theoretical principle de-
that is at least to some extent stable, so that the properties duced from particular facts, applicable to a defined group or
of the entity can be passed on to its descendants. class of phenomena, and expressible by the statement that a

1Wellcome Centre for Human Genetics, Nuffield Department of Medicine, University of Oxford, Oxford, OX3 7BN, United Kingdom.
2Department of Oncology, University of Oxford, Oxford, OX3 7DQ, United Kingdom.
*email: daniel.crouch@well.ox.ac.uk

ACADEMIA BIOLOGY 2024, 2 1 of 7

https://www.academia.edu/journals/academia-biology/about
particular phenomenon always occurs if certain conditions
be present.”
(Oxford English Dictionary)
Darwin mentions four sufficient conditions (which he describes
as “laws”) in the final paragraph of his most expansive work on
evolution, On the Origin of the Species:
a) “Growth with Reproduction”,
b) “Inheritance which is almost implied by reproduction”,
c) “Variability from the indirect and direct action of the
external conditions of life, and from use and disuse”, and
d) “a Ratio of Increase so high as to lead to a Struggle for Life”,
which together he states lead “as a consequence to Natural
Selection, entailing Divergence of Character and the Extinction
of less-improved forms” [4]. Each of these has a close analogue
in Conditions 1–4 above, in the same ordering. However, (b) and
(c), corresponding to 2 and 3, are incomplete as Darwin had no
knowledge of genetics. He is clear that a mechanism for inher-
itance is not known but only “‘implied” by reproduction, i.e., the
observable tendency for offspring of macroscopic organisms to
take after their parents, without being identical clones. The con-
cept of “inheritance” was usually contrasted with “reversion” in
Darwin’s writing, which he defined as the opposing effect of indi-
viduals taking after the general population rather than their
parent, or where an apparently inherited peculiar trait disap-
pears in subsequent generations. His “inheritance” condition
therefore refers more specifically to stable inheritance.
Darwin demonstrates clearly Lamarckian views on the source of
variation in (c), but we now know that the variation that is neces-
sary for natural selection to work is not acquired via the activity of
the organism over its life, or from effects of the environment
directing its development, other than the environmental causes of
germline DNA or RNA mutagenesis. We now know that all
biological organisms use either DNA or, in the case of some viruses,
RNA as their genetic material, which is replicated more or less
faithfully from cell to cell and organism to organism. It is only
changes in the DNA or RNA sequences due, for example, to
replication errors or environmental damage or, occasionally, stable
modifications of the sequence by methylations found only in
eukaryotes that are inherited through the germ line (epigenetic),
which can be the source of the inherited variations that determine
the differences in fitness, defined in the broadest sense, which are
the basis for natural selection. “Use and disuse” cannot possibly
provide suitable variation, as there is no way for individual
organisms to respond adaptively to their environment in a general
way except in a few cases, for example shedding of hair or muscle
growth. Though a digestive enzyme can be more highly expressed
when the food source for which it is required is being regularly
utilized, there is no conceivable way for an organism to develop a
novel enzyme simply by consuming an entirely new food source. It
is hard to see how any sort of nonbiological inheritance could
account for the evolution by natural selection of single-cell
eukaryotes or indeed even to mammals up to the level of primates.
The “Ratio of Increase” in (d) is an antiquated term referring to
the population growth rate, and Darwin, influenced by Thomas
Malthus, believed that populations would typically grow rapidly
until they reached the carrying capacity of their environment,
entering a stage in which there is a “struggle for life”, in which
less fit individuals are able to leave fewer descendants. Although
ACADEMIA BIOLOGY 2024, 2
https://doi.org/10.20935/AcadBiol6158
it is now recognized that variation in survival and reproductive
success can exist in populations that are not strained for re-
sources, (d) nevertheless refers to a requirement that there is
variation in survival, which is one component of Condition 4.
Darwin later would write extensively on the importance of the
other component: variation in reproductive success [5].
Wallace provided a “demonstration” of natural selection by tab-
ulating a set of “proved facts” that would lead to “necessary con-
sequences” such as “changes in organic forms” and “survival of
the fittest” [6], in other words a series of conditions that are
jointly sufficient for the consequences, as in a law. The facts he
specified were (i) a rapidly increasing and then stable number of
organisms, (ii) heredity with variation, and (iii) changes in
external conditions, where (i) is similar to Darwin’s condition (d),
as Wallace states that it always leads to a “struggle for existence”
and (ii) encompasses both (b) and (c). Wallace’s final fact (iii) is
an extra component that would produce adaptations to a changed
environment but is not generally necessary for evolution, as
however well adapted an organism is to its environment, there
are usually further adaptations available. Though Wallace does
not specify an analogue to (a), this is implied by (i), and he was
clearly aware that reproduction was an essential ingredient.
Clearly Wallace had, by this stage, arrived at the view that evolution
was a demonstrable law rather than a scientific theory, with
Darwin remaining more cautious. It is less well known that this
essential perspective had already been put forward 30 years before
the joint publication of their seminal works, but distinguished by a
belief that natural selection could be confirmed by observing a few
simple truths about living organisms. The author of this concept
was Patrick Matthew, who published his law of natural selection in
an appendix to a book on forestry [7], but it was not widely read
and neither of the famous co-originators of evolution by natural
selection appears to have been aware of it until after their ideas
were in print. Matthew argued that natural selection, as we now
refer to it, required just a few ingredients: a tendency for offspring
to resemble parents, variation in traits between members of a
population, and overpopulation leading to competition. He argued
that these ingredients, when combined together, created a “law
universal in nature, tending to render every reproductive being
the best possibly suited to its condition”. Remarkably, he described
his idea as the “natural process of selection”. See Weale [8] for a
thorough account of Matthew’s law and the implications for
understanding natural selection. Though Wallace, as discussed,
made a case for a law of natural selection, Matthew’s was overt
from the outset, and his conditions, which are readily observable,
are a close summary of the requirements set out in 1–4. Rather
than agonizing over the details of heredity, for instance what
causes a trait to revert to an ancestral type, Matthew claimed that
parent–offspring correlations in “particular quantities”, which
were clear to anyone, were sufficient.
The purpose of this historical overview is not to determine which
evolutionary theorist came closest to correctly establishing a law
of evolution, but rather to contextualize how evolution by natural
selection is perceived today. That its first three co-discovers all
came within reach of identifying it as a law may help to explain
why there is so much discussion, and confusion, over whether
natural selection is the dominant force in evolution [9]. If genetic
inheritance had been discovered before any major progress in
evolutionary ideas, it would have been quite easy to make the
realization that natural selection has to occur, provided that there
is some genetic variation in reproductive success. Only after this
2 of 7
https://www.academia.edu/journals/academia-biology/about
fundamental breakthrough would it have been adopted as the
major candidate to explain biological diversity. As it happened,
although all three authors’ combined works came close to finding
a law and amassed compelling empirical evidence for both mac-
roevolution and natural selection, there remained doubts over
how the latter could really work, especially due to the favored
blending model of inheritance being unstable, causing traits to
revert to the population average after a few generations. Natural
selection in diploid sexual genetic systems was barely understood
until the 1910s, by which point Darwinism prevailed mainly as a
highly evidenced theory that species descend from one another
and originate via evolutionary change, but natural selection spe-
cifically was perceived to be flawed. By developing natural
selection to the stage of it being almost a law, it is possible that
Darwin and Wallace provided a sufficiently plausible mechanism
for evolution to gain major traction as the explanation for biologi-
cal diversity, but perversely, this overshadowed the idea of
natural selection itself. Instead, natural selection was thought of
as just one of many ways that macroevolutionary change could
occur. Some have proposed that thinking from genetics did not
permeate throughout evolutionary biology, re-elevating natural
selection to center stage, until the 1940s [10], but disagreements
over its importance still remain [11].
It is unfortunate then, in retrospect, that 19th-century naturalists
had little awareness of prokaryotic microbiology, which provides a
window into genetic inheritance that might have expedited a
general acceptance of natural selection and its establishment as a
scientific law. As microbes reproduce mostly clonally, in contrast
with the diploid and higher ploidy sexual macroorganisms that
were known in Darwin’s time, it might have been possible to
observe variants giving rise to clones possessing the same traits,
without reversion to a general population type caused by segrega-
tion and recombination. A variation of a prokaryotic microbe that
tends rise to higher frequency in the population because it leaves
more copies of itself per time period than its competitors, for
example in the case of antibiotic resistance, is thus evidence of
natural selection independent of any understanding of Mendelian
genetics and might have opened a path to the discovery of such a
system in macroorganisms. Natural selection is epitomized by the
way it acts on prokaryotic microbes, and diploids unfortunately
complicate the picture, though they are more conspicuous by far to
both naturalists and general observers. One of the major insights
provided by Fisher’s fundamental theorem was to illustrate how
natural selection affects fitness in complex diploid sexual organ-
isms, where this is not so obvious as for asexual haploids. On
asexuality, Fisher wrote, “It is tempting to believe that asexual
reproduction was the primitive condition of living matter, and
that the sexual reproduction of the predominant types of organ-
isms is a development of some special value to the organisms
which deploy it”. The value referred to is the bringing together of
beneficial mutations from separate chromosomal backgrounds,
which is likely to be particularly advantageous to large organisms,
which have smaller population sizes, making it less likely for two
beneficial mutations to occur independently on the same chromo-
some [12], but more likely for disadvantageous combinations of
alleles to become common due to genetic drift [13, 14].
2. The law of natural selection
Whatever the entity in question, when Conditions 1–4 are found
to be true, the following outcomes logically follow:
ACADEMIA BIOLOGY 2024, 2
https://doi.org/10.20935/AcadBiol6158
A) Within a population of reproducing entities, new variations
will inevitably arise, if only occasionally, with advantages
over the existing variations in terms of lifespan or increased
rates of reproduction. The selection coefficient S quantifies
the extent of this advantage as the average proportional
increase in the number of offspring per generation:
w2 − w1
S= ,
w1
where w2 is the average number of times that the new
variation reproduces itself in a given generation before
expiring and w1 is the equivalent quantity averaged over
existing variations.
B) Variations for which S > 0 have advantages in lifespan or
reproduction rates due to associating with different charac-
teristics, for example biochemical features in the case of
living organisms, than alternate variations. They will then
increase at an average rate (1 + S ) per generations relative
to their unchanged ancestors.
C) The probability that a new variation, initially present as a
single copy, never goes extinct is approximately 2S [15].
When the variation is selectively neutral conferring no
average advantage, i.e. S = 0, the probability that random
reproductive success leads to it eventually replacing all
existing variants is 1/N, where N is the number of replicating
entities [16]. In general, if the probability S outweighs the
probability 1/2N, then the strength of natural selection will
outweigh random changes in frequency and the variation has
a reasonable chance of increasing in number to more than a
few copies (in haploids, the equivalent condition is S > 1/N).
In this case, the probability that the variation replaces all
existing variations is, approximately, at least 2S, which can
be found using Kimura’s well-known approximation formula
for the replacement probability [17].
D) Outcome (C) is repeated each time a new variation emerges,
so that the population undergoes continual improvement of
average lifespan and/or reproduction rates, due to changes
in the distribution of characteristics of its members. This
variant may or may not occur in a previously changed
individual. However, in the absence of a sexual system, the
most advantageous individuals are mostly likely to accumu-
late a series of advantageous changes one after the other
since there is no other way for accumulating combinations
of advantageous changes.
Outcomes A–D allow scientists to make valid inferences about
what they expect to observe when a set of organisms, or more
generally replicating entities, meet Conditions 1–4, as the
consequences are jointly implied, in other words entailed, by the
conditions. Scientific theories, in contrast, work in reverse by
inferring the conditions from their real-world outcomes. Data
that do not support their associated hypotheses constitute
evidence against the theory by suggesting that at least one of the
conditions is false. Conversely, data that support their associated
hypotheses suggest that all the conditions, and therefore the
theory, could be true [8]. By this definition, a theory can only be
investigated indirectly, via the hypotheses that it implies, as its
essential axioms are assumed to be unobservable. Crucially this
means that even when a theory is very well supported by data,
there may be other, perhaps unknown, theories which support
the data better or equally well. Furthermore, confirmation of a
3 of 7
https://www.academia.edu/journals/academia-biology/about
hypothesis is only necessary, and not sufficient, for its associated
theory to be true, as there may be false hypotheses implied by the
theory that remain untested. None of this is to suggest that a
scientific law cannot ever be found to be false. Rather, it is true
whenever its conditions hold, whereas theories that agree with all
the available evidence may still be false. To find that a law is false,
it would be necessary to disprove one or more of its conditions.
What kind of characteristics might be associated with new
selectively advantageous variations? These will differ from situa-
tion to situation, and there will be a strong element of unpre-
dictability about how new variations differ, but R.A. Fisher
famously showed how they can include characteristics that vary
continuously at the level of the whole organism, such as height or
weight, via the accumulated action of many variations with
effects on that characteristic [18]. Fisher went on to formalize
natural selection mathematically for diploid sexual organisms in
his “fundamental theorem” [1], accommodating for this action of
many genetic factors on the reproductive and survival potential
of an organism, i.e., fitness. In a model in which all individuals in
a generation are born at the same time, before reproducing and
dying simultaneously before the next generation is born, Fisher
defined fitness as the number of offspring an individual organism
leaves at the end of its life. The fundamental theorem of natural
selection then shows that the component of phenotypic change in
fitness that is due to the action of natural selection on changing
allele frequencies is proportional to the variation in fitness
attributable to the additive genetic effects [19]. Organisms with
diploid inheritance systems, the only type of such system known
to science in the early 20th century, may have either 0, 1, or 2
fitness-increasing alleles at each genetic position on a chromo-
some. Of course, there may be genetic positions harboring only
alleles which have essentially no effect on fitness, and there are
typically many positions at which individuals do not much vary
genetically; there are only very occasionally observed mutant
alleles that provide a fitness advantage that is more powerful for
effecting changes in the frequency of genetic variants carrying
them than would drift in the absence of a fitness difference.
Fisher’s formulation of a simple mathematical relationship be-
tween the inheritable variation in Darwinian fitness and the rate of
increase of its average value signifies the full maturation of natural
selection into a law of nature applying to all biological organisms,
whether clonally or sexually reproducing, with confirmed under-
pinnings and predictable outcomes. Fisher never made explicit
reference to a law of natural selection, though he did regard the
fundamental theorem as a biological analogue to the second law of
thermodynamics [1]. Without diminishing the insight provided by
the fundamental theorem into how average fitness changes at the
level of the entire organism, natural selection in diploids is most
easily conceptualized as acting directly on individual genetic
alleles, which are the replicating variations in Conditions 1–4.
Fitness of a given allele is defined most clearly as the number of
times each allele copy in the population makes a copy of itself
within a given unit of time, averaged over all genetic backgrounds,
i.e., genomes, on which the allele occurs [1].
3. Implications of natural selection
being a law
There is an overwhelming scientific consensus that both natural
selection and evolution are real phenomena, yet there is disagree-
ment about the extent to which natural selection dominates other
ACADEMIA BIOLOGY 2024, 2
https://doi.org/10.20935/AcadBiol6158
evolutionary factors in producing inherited adaptations to either
the physical or the biological environment, or both. These other
factors include the availability of the right genetic mutations for
selection to act upon, and how the range of an organism’s
potential adaptations is contingent on evolutionary history, for
example, the basic structure of its body plan. Genetic drift in
small populations, e.g., where individuals are geographically
subdivided into groups, may allow genetic variants that would be
individually disadvantageous to randomly climb to high frequen-
cies, at which point they become jointly advantageous when
appearing together in the same individuals. The role of niches,
i.e., the confinement of individuals into local ecological condi-
tions to which they can then adapt via selection, is also sometimes
regarded as an important non-selective factor [20]. However,
natural selection, as a fundamental scientific law, differs from
these other evolutionary forces as Conditions 1–4 are true of all
living organisms and are an intrinsic part of any adaptive change,
regardless of which of the other forces may be having some
partial influence.
It is ultimately variation in the genetically determined pheno-
types that are associated with fitness differences, which drives
the evolutionary process. This leaves the possibility that there is
genetic variation among individuals with a given phenotype, all
of which have effectively the same fitness, or that there are
genetically different phenotypes that all have effectively the same
fitness. These latter two possibilities are non-adaptive evolution-
ary changes. In this context, genetic drift may sometimes play a
strong non-adaptive evolutionary role in very small populations,
as randomly varying reproductive success among individuals in
such populations could obscure the average advantage that
carrying a particular allele may confer. Variants with S > 1/2N
have a reasonable probability, 2S, of spreading through the
population, leaving scope for mutant alleles with, for example, a
little over 1% fitness advantage to have some chance (roughly 2%
per allele) of increasing in a population of even only 50 individu-
als, just as they would in a large population. New mutations upon
which selection can act are also in shorter supply when the
population size is low, as the rate at which they appear scales
linearly with the number of individuals. Thus, there are fewer
chances for mutations with S > 1/2N to appear in a small popula-
tion. This effect may, however, be less serious in organisms with
large genomes. Nevertheless, if left for long enough, a small
population’s average fitness would normally be expected to
increase due to the eventual appearance of such alleles, and the
population might then grow in size if there were sufficient re-
sources available, producing more mutations and allowing
further scope for selection of alleles with lower S. Natural
selection is considered to be weak, on average, in small popula-
tions because of their reduced ability to respond rapidly to
changes in the environment requiring new adaptations, due to a
lack of mutations with S > 1/2N, and their tendency to accumu-
late disadvantageous alleles. This tendency may sometimes give
rise to high frequencies of disadvantageous alleles by chance
when their disadvantage, measured on the same scale as S, is
close to or below 1/2N. A sufficient number of disadvantageous
alleles accumulating at high frequency may have the effect of
shrinking the already small population size, leading to a greater
tendency toward their further accumulation, resulting in a
feedback loop referred to as “mutational meltdown”, that could
lead to population extinction [21]. In either case, natural
selection persists, but may not result in adapted individuals
quickly enough to keep up with countervailing forces of rapid
4 of 7
https://www.academia.edu/journals/academia-biology/about
environmental change and accumulation of disadvantageous
alleles. The inability of natural selection to achieve rapid adapta-
tion to sudden environmental changes can, of course, also occur
in large populations, for example, during a mass-extinction event
as is presumed to explain the disappearance of dinosaurs at least
to some extent.
Due to uncertainty over which beneficial mutations will appear and
spread beyond a few copies in the population, it is not possible to
predict which new phenotypic characteristics will, in general,
evolve. Continuously distributed phenotypes, however, for which
there is existing variation in a population, such as weight and
height, are under the control of large numbers of variants, each
having a small phenotypic effect. If the optimal height were at some
point to change due to some change in environmental conditions,
it would only take small changes in the frequencies of tall versus
short causing alleles for the new optimum to be achieved. It is thus
certain that natural selection would shift the average continuous
trait value of the population to the new optimum. While the
average phenotype of a population lies close to the optimum for the
given environment, natural selection acts in a stabilizing fashion
[22], selecting against variants (and combinations of variants) with
large phenotypic effects, as these shift carriers far from the opti-
mum. In most cases, organisms are well adapted to their current
environment, in which case one would expect no evolutionary
change in the average characteristic (or the distribution of charac-
teristics). However, that is not to say that mutations for entirely
new advantageous characteristics may not emerge and spread
through the population.
Given that natural selection will occur given Conditions 1–4, can
we speculate what extra-terrestrial life might look like? It is very
difficult to say what sort of specific phenotypic adaptations might
exist on other planets, but we know that adaptation will occur if
a simple replicating molecule satisfying the conditions were to
emerge, possibly due to a relatively rare chance event. Matthew
provided this early exobiological perspective in 1860, 29 years
after he published his law: “The great law of nature in organic
life is competition, with a variation-power in accommodation to
circumstances: a law not fitted to earth alone, but I have no
doubt extended to the whole of the orbs of space that are in a
condition to support material life” [8, 23]. Earthly observations,
in as much as they can be generalized, suggest that most of the
adaptations of life initially will have occurred on the prokaryotic
microbial level, as prokaryotic microbes have filled almost every
ecological niche imaginable, including those with extreme
temperatures, salinities, and pH. Prokaryotic microbes emerged
only about 800 million years after the formation of Earth, but it
took a further three billion years for the evolution of complex
multicellular organisms, suggesting that specific additional con-
ditions or very rare chance events are required for this later state
of evolution to have occurred.
Natural selection may often lead to an extraordinary amount of
complexity, but this does not mean that it will necessarily tend
toward evolution of higher complexity. In some cases, it can also
contribute to the failure of complex organisms, as is likely in the
case of selection for somatically mutated cells leading to cancers,
and also in age-related disease. Adult tissue stem cells undergo
regular mitotic division in order to replenish the differentiated
cells responsible for tissue function, which become fatigued and
die over time. Due to the potential for mutations, these cells meet
the conditions of natural selection, resulting in overgrowth in the
populations of stem cells and their descendant differentiated
ACADEMIA BIOLOGY 2024, 2
https://doi.org/10.20935/AcadBiol6158
cells, at a detrimental effect to the organism as a whole. In young
organisms, homeostatic mechanisms must be in place to prevent
these cell populations from overgrowing, but these are likely to
become less efficient as the organism passes reproductive age,
due to having decreasing effects on fitness [24]. This is also likely
to account, in part, for why the risk of nearly all types of cancers
increases with age [25]. These processes illustrate the potential
for selection to favor “selfish” components within a complex
organism at the expense of the organism’s overall healthy func-
tioning, while selection acting on the organism as a whole favors
homeostatic suppression of the individual selfish elements. Ten-
sion between selection at lower and higher levels of biological
organization is the focus of multilevel selection (or group selec-
tion) theory [26].
Biological organisms all possess capacities for semi-stable inher-
itance and mutation leading to variation in reproductive fitness
(Conditions 2–4), provided that they are situated in an environ-
ment in which they can reproduce (Condition 1). It is for this
reason that natural selection is such a pervasive feature of the
biological world and yet is atypical of nonbiological objects.
Viruses are sometimes considered to be non-living due to per-
forming none of their own metabolic activity, but this does not
take into account the fact that they are naturally selected to hijack
and control their hosts’ metabolic machinery, with the resulting
metabolic actions then being for the benefit of the virus. Consider
how animals rely ultimately on plants for sources of energy, yet
we would consider them to have actively obtained those re-
sources for themselves. Under this view, it is appropriate to clas-
sify viruses as living organisms as they satisfy the requirements
for evolving by natural selection, and the same might be true of
even simpler biological parasites like selfish DNA elements
(transposons) and infectious proteins (prions). The critical
difference between a prion or a transposon and a gene or a DNA
variant, in this respect, might lie how the replicating entity in
Condition 1 is defined. Genes and DNA variants replicate to-
gether with the cell, or multicellular organism, to which they
belong, suggesting that it is the entire replicating entity that
should be considered alive. Transposons and prions replicate
separately from their host organisms and cells, which constitute
their environment.
Should then a computer virus written so as to replicate and
occasionally mutate its own code, and thus undergo selection,
also be considered “alive”? The answer should be no, as it would
have to be the computer that self-replicates not only the program.
This is for this same reason that genes themselves should not be
considered as alive, but that cells and organisms should be, as the
former are not able to self-replicate in isolation. It is interesting
to consider what life-like properties may be reflected by nonbi-
ological evolving entities. What security risks would be associated
with, for example, evolving computer viruses developing mimicry
to evade detection, forming symbiotic relationships with each
other, or undergoing forms of recombination in which computer
code is exchanged horizontally between different viral lineages?
Computer programs that develop in complexity to the point of
becoming highly intelligent and self-interested, eventually over-
throwing their human creators, are a familiar theme in science
fiction. This idea does not seem too far-fetched if one considers
how much complexity has arisen in the biological world from
natural selection acting initially on very simple materials, prob-
ably individual RNA molecules. However, an essential feature of
all four conditions for natural selection is that they must hold
5 of 7
https://www.academia.edu/journals/academia-biology/about
within a given environment, which for a self-replicating program
is computer hardware that they are totally reliant on humans to
provide and maintain.
The law of natural selection should engender caution whenever
there is a potential link between the replicative ability of a virus
or an organism and its propensity toward causing damage,
including to health or the environment, the former being a cause
for great concern in the case of emerging infectious diseases such
as COVID-19. Biological weapons represent such a risk, as the
release of replicating pathogenic organisms will almost inevitably
lead to selection for greater survival abilities within the host and
greater transmissibility to new hosts, allowing them to spread far
beyond their original target area and to persist for longer than
intended [27]. While chemical weapons share a tendency to
spread unpredictably beyond the target range, they do not have
the inbuilt capacity to undergo selection and adaptation as does
a biological weapon. It is likely that the general trend for
pathogens is to become less virulent to their hosts over time,
thereby providing more opportunities to transmit to new hosts,
though there may still be substantial health consequences to the
host of becoming infected. On at least some occasions, selection
for increased transmissibility may, however, be accompanied by
increased virulence [28, 29].
Mistaking natural selection to be a scientific theory rather than a
law has led to disagreements over its fundamental validity and
explanatory power, with a common argument being that it cannot
be successful as a theory due to merely describing a tautology
whereby the organisms that reproduce the most are those that
are most reproductively successful [30]. Critics claim that due to
being a tautology, natural selection a) contains no information
and b) is always logically true, so cannot confirm or deny
anything about the real world [31, 32]. Both of these lines of
thinking are wrong. First, (a) is only true of extremely simplified
definitions of natural selection e.g., as “the survival of the fittest”,
which incorporates Conditions 1 and 4, but says nothing about
how variations with different fitnesses are generated (Condition
3), and little about inheritance (Condition 2). Although the full
law incorporating all four conditions does exhibit one feature of
a tautology, that its outcomes are logically implied by its
conditions, this is true of any logical statement including elusive
mathematical theorems (or, indeed, scientific theories). Like a
mathematical theorem, natural selection does contain infor-
mation, as its logical relations had to be discovered, and it does
not just consist of the simple equating of synonyms as in a
genuine tautology like “survival of the fittest”. Second, although
natural selection relates statements logically as in a tautology, (b)
does not allow for the fact that its conditions are not a priori true
and can thus be verified or found to be false (at least to a degree
of certainty) in the real world. It seems quite likely that beliefs (a)
and (b) are driven partly by mischaracterizing natural selection
as a theory. Questions under empirical scrutiny such as “is there
variation in reproductive success?” are then possibly mistaken
for hypotheses generated by the underlying conditions, rather
than means of confirming the conditions themselves. If so, the
scientist might be left with the impression that they are working
to verify hypotheses that are so closely connected to the individ-
ual conditions that the overall theory—the logical relations be-
tween the conditions and the outcomes—appears to provide no
information. All becomes clear when natural selection is taken to
be a law, in which it is transparent that what is under empirical
interrogation is the set of basic conditions. Outcomes of the law,
ACADEMIA BIOLOGY 2024, 2
https://doi.org/10.20935/AcadBiol6158
such as the prediction that adaptation will tend to occur, are
understood to be separate from the conditions and unnecessary
to test as if they were hypotheses.
4. Conclusion
Mendelian genetics shows that biological characteristics of sexual
macroorganisms are inherited in such a way that underlying
heritable information is passed on, unchanged, to later genera-
tions, but with occasional instability in the form of mutations. It
has thus established two essential conditions which, together
with a capacity for reproduction and variation in longevity or
reproductive success, guarantee the action of natural selection
leading to adaptation. This establishes adaptive evolution by
natural selection as a scientific law by any reasonable definition.
Given that natural selection is then certain to occur for biological
entities and so is a law in that context, it can only continue to be
regarded as a theory in the sense that there might be other ways
for adaptive evolution to occur without it, but such mechanisms
are not so far known to exist.
Funding
The authors declare no financial support for the research, author-
ship, or publication of this article.
Author contributions
Conceptualization, W.F.B.; investigation, D.J.M.C. and W.F.B.;
writing—original draft preparation, D.J.M.C.; writing—review
and editing, W.F.B.; supervision, W.F.B. All authors have read
and agreed to the published version of the manuscript.
Conflict of interest
The authors declare no conflict of interest.
Data availability statement
Data supporting these findings are available within the article, at
https://doi.org/10.20935/AcadBiol6158, or upon request.
Institutional review board statement
Not applicable.
Informed consent statement
Not applicable.
Sample availability
The authors declare no physical samples were used in the study.
Additional information
Received: 2023-10-26
Accepted: 2024-01-23
Published: 2024-02-08
6 of 7
https://www.academia.edu/journals/academia-biology/about
Academia Biology papers should be cited as Academia Biology
2024, ISSN 2837-4010, https://doi.org/10.20935/AcadBiol6158.
The journal’s official abbreviation is Acad. Biol.
Publisher’s note
Academia.edu stays neutral with regard to jurisdictional claims
in published maps and institutional affiliations. All claims
expressed in this article are solely those of the authors and do not
necessarily represent those of their affiliated organizations, or
those of the publisher, the editors, and the reviewers. Any
product that may be evaluated in this article, or claim that may
be made by its manufacturer, is not guaranteed or endorsed by
the publisher.
Copyright
© 2024 copyright by the authors. This article is an open access
article distributed under the terms and conditions of the Crea-
tive Commons Attribution (CC BY) license (https://creative
commons.org/licenses/by/4.0/).
References
1. Fisher RA. The genetical theory of natural selection. Oxford:
Clarendon Press; 1930. p. xiv, 272 p., ii col. pl. (incl. front.).
2. Mayr E. Darwin’s influence on modern thought. Sci Am.
2000;283:78–83.
3. Darwin C, Wallace A. On the tendency of species to form
varieties; and on the perpetuation of varieties and species by
natural means of selection. J Proc Linn Soc London Zool.
1858;3:45–62.
4. Darwin C. On the origin of species by means of natural
selection, or preservation of favoured races in the struggle
for life. London: John Murray; 1859.
5. Darwin C. The descent of man and selection in relation to
sex. 2nd ed. London: John Murray; 1874.
6. Wallace AR. Contributions to the theory of natural selection.
A series of essays. MS. notes [by F. T. Palgrave]. London:
Macmillan & Co; 1870.
7. Matthew P. On naval timber and arboriculture. London: A.
Black; 1831.
8. Weale ME. Patrick Matthew’s law of natural selection. Biol J
Linn Soc. 2015;115:785–91.
9. Charlesworth D, Barton NH, Charlesworth B. The sources of
adaptive variation. Proc R Soc B. 2017;284:20162864.
10. Mayr E, Provine WB. The evolutionary synthesis: perspec-
tives on the unification of biology. Cambridge (MA):
Harvard University Press; 1980. p. xi, 487 p.
11. Segerstråle UCO. Defenders of the truth: the battle for
science in the sociology debate and beyond. Oxford: Oxford
University Press; 2000. p. ix, 493 p.
12. Bodmer W. The evolutionary significance of recombination in
prokaryotes. Symp Soc General Microbiol. 1970;20:279–94.
ACADEMIA BIOLOGY 2024, 2
https://doi.org/10.20935/AcadBiol6158
13. Crouch DJM. Statistical aspects of evolution under natural
selection, with implications for the advantage of sexual
reproduction. J Theor Biol. 2017;431:79–86.
14. Hill WG, Robertson A. The effect of linkage on limits to
artificial selection. Genet Res. 1966;8:269–94.
15. Haldane JBS. A mathematical theory of natural and artificial
selection. Math Proc Cambridge Philos Soc. 1927;23:607–15.
16. Wright S. Evolution in Mendelian populations. Genetics.
1931;16:97–159.
17. Kimura M. On the probability of fixation of mutant genes in
a population. Genetics. 1962;47:713–9.
18. Fisher RA. XV.—The correlation between relatives on the
supposition of Mendelian inheritance. Trans R Soc Edinb.
1918;52:399–433.
19. Edwards AW. R.A. Fisher’s gene-centred view of evolution
and the Fundamental Theorem of Natural Selection. Biol
Rev Camb Philos Soc. 2014;89:135–47.
20. Laland K, Matthews B, Feldman MW. An introduction to
niche construction theory. Evol Ecol. 2016;30:191–202.
21. Lansch-Justen L, Cusseddu D, Schmitz MA, Bank C. The
extinction time under mutational meltdown driven by high
mutation rates. Ecol Evol. 2022;12:e9046.
22. Kutschera U. Charles Darwin’s Origin of Species, directional
selection, and the evolutionary sciences today. Naturwis-
senschaften. 2009;96:1247–63.
23. Matthew P. Origin of species. Farmer’s Magazine Ser. 1860;
3:30–2.
24. Bodmer WF, Crouch DJM. Somatic selection of poorly
differentiating variant stem cell clones could be a key to
human ageing. J Theor Biol. 2020;489:110153.
25. Bodmer WF, Crouch DJM. Chapter 4 - Evolutionary per-
spectives on cancer and aging. In: Wasser SP, Frenkel-
Morgenstern M, editors. New horizons in evolution. London:
Academic Press; 2021. p. 97–115. doi: 10.1016/B978-0-323-
90752-1.00008-0
26. Frank SA. Foundations of social evolution. vol. 2. New
Jersey: Princeton University Press; 1998.
27. Lederberg J. Biological weapons: limiting the threat. BCSIA
studies in international security. Cambridge (MA): MIT
Press; 1999. p. xvi, 351 p.
28. Wymant C, et al. A highly virulent variant of HIV-1 circulat-
ing in the Netherlands. Science. 2022;375:540–5.
29. Challen R, et al. Risk of mortality in patients infected with
SARS-CoV-2 variant of concern 202012/1: matched cohort
study. BMJ. 2021;372:n579.
30. Elgin M, Sober E. Popper’s shifting appraisal of evolutionary
theory. J Int Soc Hist Philos Sci. 2017;7:31–55.
31. Hunt T. Reconsidering the logical structure of the theory of
natural selection. Commun Integr Biol. 2014;7:e972848.
32. Popper KR. Objective knowledge. vol. 360. Oxford: Oxford
University Press; 1972.
7 of 7