CRITICAL SCIENTIFIC REFLEXIONS ABOUT
THE RED QUEEN HYPOTHESIS.
5 David L. ESPESSET
------
10 72, chemin des Baumillons,
F-13015 Marseille,
France.
15
1 ABSTRACT.
In evolutionary sciences, the Red Queen Hypothesis deals with extinction
rates over geological time. In this article, I propose a critical reflexion on this
20 hypothesis, because some implications or metaphorical applications of the
hypothesis might be considered as misinterpretations of evolutionary facts. I
review various scientific interpretations of evolution and their relationship with
the Red Queen Hypothesis. Eventually, I propose to rename it, to correct past
misinterpretations and avoid new ones.
25
KEYWORDS.
Biological evolution – Darwinism – Red Queen Hypothesis – Co-evolution
– Convergence – Evolutionary ecology – Laws of nature.
30
2 EVOLUTION AND CO-EVOLUTION.
Biological evolution is the changing of living beings in their heritable
traits over successive generations and geological time. Various evolutionary
35 processes give rise to diversity at every level of biological organisation,
including the levels of taxa, species, individual organisms, cells, and molecules.
Among other scientific theories, the “Modern Synthesis” [1, 2] explains the
evolutionary process, mainly based upon a so-called “Darwinian” mechanism
[3], which involves random mutations followed by natural selection. Another
40 mechanism, the genetic drift, explains how an allele can be randomly fixed in
small sized populations [4].
During the course of evolution, many species develop close relationships
with each other to ensure their survival: as one species evolves, it will
somehow affect other species. These species are said to coevolve [5]. Such
45 'symbiotic' relationships keep species from becoming extinct. Among many
instances of coevolving species, some are famous and have been extensively
studied, such as the coevolution of pollinating Insects and flowering plants
(Angiosperms) and the many examples of coevolution between parasites and
hosts [6].
50
3 THE RED QUEEN HYPOTHESIS.
In 1973, Leigh Van Valen proposed his “Red Queen Hypothesis” [7], also
55 referred to as “Red Queen's”, “Red Queen's Race” or “The Red Queen Effect”
(hereafter: RQ), a 'new evolutionary law' which proposes several statements:
“All groups for which data exist go extinct at a rate that is constant for a
given group”;
“Extinction rates are similar within some very broad categories (…)”;
60 “(…) Extinction in any adaptive zone occurs at a stochastically constant
rate”;
“The probability of extinction of a taxon is (…) effectively independent of
its age.” [7]
The hypothesis thus mainly deals with extinction rates over geological
65 time, which can be summarized as follows: at more than one taxonomic level,
the shapes of survivorship curves (a great number of which are shown in the
original article) are consistent with the possibility that the relative probability
of extinction of a taxon is approximately constant per unit of time, as observed
in the paleontological record. A direct implication is that longer-lived taxa (e.g.
70 species) are not better adapted than younger taxa (their extinction rates are
similar); therefore, the RQ (the probability of evolutionary change is
independent of the age of the taxa). The hypothesis can be thought of in terms
of organismal change, species change or process change.
The RQ intends to explain two different processes:
75 1. Coevolution among competing species could lead to situations in which the
4 probability of extinction is relatively constant over millions of years, in relation
with the fact that “The RQ proposes that events of mutualism (…) are of little
importance in evolution in comparison to negative interactions” [7]. For
instance, the RQ appears to be especially decisive in predator-prey
80 relationships. Indeed, predator-prey relationships are arguably some of the
most significant types of relationships in regard to the survival of species. For
example, if a prey species evolves and becomes faster over a period of time,
the predator needs to adapt and evolve in order to keep using the prey as a
reliable food source. Otherwise, the now faster prey will escape and the
85 predator will lose a food source and potentially become extinct. However, if the
predator gets faster itself, or evolves in another way like being stealthier or a
better hunter, then the relationship can continue and the predator will survive.
2. The advantage and persistence of sexual reproduction (as opposed to
asexual reproduction) at the level of individuals [8]. For instance, individuals
90 living in an area with any particular parasite may prefer a mate that seems to
be immune to the parasite: the resulting offspring itself would then more
probably be immune to the parasite, making it more likely to survive,
reproduce and pass down its genes. In other words, in species for which
asexual reproduction is possible (as in many plants and 'invertebrates'),
95 coevolutionary interactions with parasites may select sexual reproduction in
hosts as a way to reduce the risk of the offspring being infected (RQ applied to
sexual reproduction).
Numerous examples have been held up in support of the supposed
validity of the RQ. Van Valen himself “see(s) here a major difference from the
5 100 usual theory of genetic selection” [7]. About molecular evolution, he also
recalls that the “Constancy of the rates of protein evolution is often regarded
as the most important evidence for (…) non-Darwinian evolution (…)” [7]. Van
Valen also states that “the RQ, curiously, does not deny progress in evolution”
[7], something Stephen Jay Gould would certainly have denied (see below).
105 Another significant implication of the RQ is that organisms must constantly
adapt, evolve, and proliferate not only to gain reproductive advantage, but also
simply to survive and avoid extinction while pitted against ever-evolving
opposing organisms in an ever-changing environment.
6 110 THE MAIN METAPHORICAL APPLICATION OF THE RED QUEEN: A
MISINTERPRETATION OF EVOLUTIONARY FACTS?
The phrase "Red Queen Hypothesis" is derived from a statement that the
Red Queen, a chaotic and despotic character maintaining a terrifying ruling
115 over Wonderland, makes to Alice in Lewis Carroll's “Through The Looking-
Glass” (chapter two: The Garden Of Live Flowers [9]). In her dream about the
looking-glass house, Alice leaves the house to look at the garden: Alice thinks
it will be easier to see the garden if she first walks up to the top of the hill, to
which a straight path seems to lead. At the top of the hill, the Red Queen
120 starts running, faster and faster. Alice runs after the Red Queen but is
perplexed to find that neither of them seems to be moving. When they stop
running, they are exactly in the same place. Alice remarks on this, and the Red
Queen responds:
"Now, here, you see, it takes all the running you can do
125 to keep in the same place".
What appears to me as the main – and most questionable – application
of the RQ is that evolutionary change might be necessary to stay in the same
place, or, to put it differently, that ever-evolving living organisms would be
seen as staying in the same place. In this view, species have to 'run' (evolve)
130 in order to 'stay in the same place' (survive to be extant). It generally seems
that such subsequent uses of the RQ have tacitly assumed that the words
('run' and 'stay in the same place') are to be interpreted metaphorically, as Van
Valen himself apparently did.
7 But, even metaphorically, do living organisms really 'stay in the same'
135 place as they evolve? To me, such a point of view is a fundamental
misinterpretation of basic evolutionary facts.
In the following sections, I will review several scientific interpretations of
evolution and their relationship with the RQ.
140 “Neo-Darwinian orthodoxy”.
In the neo-Darwinian view of evolution [10, 11], on the basis of random
mutations followed by natural selection, evolution has no particular 'goal'
except that of adapting living organisms to their fluctuating environment (this
refers to the everlasting debate about finality versus finalism; see below). But
145 what appears to be 'adapted' today may well be 'non adapted' tomorrow: if the
environment changes, then organisms must adapt to the new conditions,
otherwise they will simply disappear. In this respect I agree: when organisms
evolve, they literally 'keep in the same place'. Moreover, most evolutionary
biologists state that evolving organisms do not 'progress' [12], they merely
150 'change'. There would be no progress in nature, no race towards complexity,
no evolutionary trends, no evolutionary strategies and evolution would be
totally aimless [11], like the entire universe itself, incidentally. Some
evolutionary biologists even state that 'complexity' is a useless notion in
biology [13]. This school of thought is related to 'reductionism' in science.
155
8 From the “Modern Synthesis” to the “Extended Evolutionary
Synthesis”.
160 But this “Darwinian evolution”, sometimes referred to as “ultra-
Darwinism”, is now more and more controversial. More precisely, without ruling
out the existence of Darwinian mechanisms to explain some aspects of
biological evolution (like micro-evolution), it appears that other distinct
evolutionary mechanisms may well be working in nature. For instance niche
165 construction, developmental bias, plasticity, lateral gene transfers and extra-
genetic inheritance [14, 15, 16]. These mechanisms are part of the “Extended
Evolutionary Synthesis” [17, 18]. I doubt that this synthesis is compatible with
the idea of organisms metaphorically 'keeping in the same place' at the species
level.
170 Consider earthworms and their ability to modify the soil they inhabit to
make it a better place for their need of humidity and wet dirt. As Laland et al.
state [14], physiologically, earthworms are aquatic annelids. Through a niche
construction process, they are able to restructurate the soil environment to
enhance water retention and accessibility. This creates for them a new
175 adaptive interface, one that is better suited to earthworms' essentially aquatic
physiology. This constructed niche serves as a form of external hereditary
memory, an ecological inheritance. In such a perspective, how can earthworms
be considered to 'keep in the same place'? I even think that such a niche
construction can be seen as some kind of progress (a notion totally
180 incompatible with Darwinian orthodoxy but not incompatible with the RQ in its
original formulation): by modifying their proximate environment, earthworms
9 have improved their life conditions, they have made progress.
Now, let us turn to mound-building termites, especially the fungus-
cultivating termites [14]. Above their subterranean nest, termites build a
185 massive mound permeated with an elaborate network of tunnels that culminate
in a finely porous mound surface. Constantly attacked by erosion,
compensated by active soil deposition by the termites, the mound appears to
be a dynamic adaptive structure, an interface between the colony itself and its
broader external environment, involved in managing water balance and
190 respiratory gas concentrations. This is another example of extended
physiology, in which the homeostasis of the nest extends upward into the
dynamic construction of the mound, thus maintaining a steady nest
atmosphere. Being permanently built and extensively remodelled, and capable
of filtering the transient energy in turbulent wind, allowing it to be captured to
195 drive the colony's respiratory gas exchange, the mound eventually acts as a
regulated wind-driven 'lung' for the colony. As with earthworms, this extended
physiological process is a striking example of counteractive niche construction
which neutralizes prior environmental natural selection pressures. Once more,
in such a complex process, termites cannot be seen as 'keeping in the same
200 place', inasmuch as they extensively modify their external environment, which,
as in the case of the earthworms, can be seen as some kind of progress.
At this point we can draw a first conclusion: the RQ is definitely not
Darwinian, insofar as it sometimes involves the notion of progress during the
course of evolution. Such a statement is all the more surprising since the RQ
205 has most often been implicated in Darwinian evolution. As Stephen Jay Gould
10 states in his 'Cosmic dance of Siva': “The Red Queen has been our dominant
model for life history” [19], 'our dominant model' meaning, of course,
Darwinian. This is the reason why it seems to me that such an implication of
the RQ may be considered as a misinterpretation of evolutionary facts.
210
The “Libertine Bubble Theory”.
In 2011, Thierry Lodé proposed his “Libertine Bubble Theory” [20] to
explain the origin of sex in living organisms, which therefore leads to new
coevolution mechanisms. This theory is original because it is based on
215 ecological considerations. Life probably began in 'bubbles' because the
molecules involved in the basic biochemical reactions of life had to be
protected from the external environment by a membrane. Basically, sex is not
seen as initially correlated to reproduction but as a process of gene transfer
involving contacts between prebiotic 'bubbles' interacting for ecological
220 reasons: food or symbiotic relationships, based on primitive cellular
metabolism. This is the beginning of 'living behaviour'. These interactions have
led to natural selection processes, as the more interacting bubbles ('libertine
bubbles') were certainly more efficient in many ways (especially gene
exchanges), which is evolutionarily relevant. In short, interacting organisms
225 can be seen as coevolving on the basis of ecological processes. Such an
ecological coevolution acknowledges the structuring power of dynamic and
ever-changing interactions between living organisms.
One of the implications of Van Valen's hypothesis [8] emphasises the fact
that interactions between two living organisms can lead to symmetrical
11 230 coevolution. However, according to Lodé, the mechanism involved in such a
process appears to be somewhat external to the interaction: in such a view,
the RQ is only descriptive and questionable because it provides no functioning
lead. This would also be the case in the 'arms race' because each organism
improves its arms according to those of its 'opponents'.
235 Still according to Lodé, on the contrary, in the “Libertine Bubble” theory,
the solution lies in the relationship itself, the interaction being considered as a
structuring power in a tenuous and dynamic balance between two living
species: the first species can only survive because it establishes a relationship
with the second species, and conversely. In this perspective, everything comes
240 back to ecology. It must be stressed that relationships are fundamental in
evolution, and that no species has ever evolved on its own: each and every
species has its place in a complex network of interactions. Such relationships
are particular in sex, in which living organisms become partners whose DNA
can be specifically modified, especially in Eukaryotes. All these ecological
245 processes appear to be part of what can be called evolutionary ecology.
In this ecological view of evolution, it can be seen that living organisms
obviously never stay in the same place. Any relationship between living
organisms is rich, complex and, above all, essentially dynamic and ever-
changing: two (or more) interacting living organisms can never stay in the
250 same place, especially in the many cases of coevolution.
12 Convergence and evolutionary routes.
255 Moreover, as Simon Conway-Morris clearly develops in “Runes Of
Evolution” [21], biological evolution is mainly convergent. “(…) Everywhere you
look evolution is hedged in by convergence”, he states. And he adds, “(…)
Evolution almost goes in circles – reinventing the wheel (…)”. As such,
evolution can be considered as repeatable and, at least to some extant,
260 predictable.
It obviously shows that evolutionary routes do exist, and that these
routes lead to somewhere: the solution to a problem (which can be seen as the
most basic definition of intelligence). According to V. Orgogozo, it is even
possible to speak of general evolutionary trends [22]. In this perspective,
265 evolutionary processes lead organisms towards better adaptations, sometimes
new ones, but often rather similar. At the risk of appearing reactionary to most
evolutionists, I think it should be considered that evolution is a progress, even
if it can be seen as an arms race. When an organism develops a new organ, a
new character, it is no longer the same organism, sometimes it even defines a
270 new species, differently or even better adapted to new environmental
constraints: it has not stayed in the same place at all. However, if for whatever
reason such an adaptation happens to be impossible, then the organism will
simply disappear: “Adapt or perish, now as ever, is nature’s inexorable
imperative.” (H. G. Wells). There is no staying in the same place in evolution.
275 Ever. This statement has strong implications about the evolutionary process
itself, as will be emphasised in the following sections.
13 Structuralism and the laws of nature.
For Michael Denton, the structuralist school of thought is a “Persistent
280 Challenge To Darwinian Selectionism” ([23]; part of the title). Indeed,
structuralism states that the basic forms of the biological world are immanent
in nature and determined by a set of special natural biological laws. Therefore,
a significant fraction of the order of life is the result of basic physical
constraints rising from the fundamental properties of matter. These constraints
285 limit the way organisms are built to a few basic designs ('body plans') which
can be seen as genuine universals. These basic forms, or types, show an
extraordinary robustness over geological time and stability over space.
Structuralists thus adhere to a strictly 'non selectionist, non historicist' view of
the biological world, in which adaptation would be secondary and peripheral,
290 not the result of natural law ('order from within') but of a response to
environmental conditions (imposing an external 'order from without').
Structuralism therefore implies that organic order is a mix of two completely
different types of order, generated by two different causal mechanisms: a
primal order generated by natural law, and a secondary adaptive order
295 ('adaptive masks') imposed by environmental constraints (by natural selection,
according to Darwinists) [24].
In such a view, once more, evolving organisms cannot be seen as
'keeping in the same place'. By respecting the laws of nature leading to
'predetermined' or 'primal patterns', living organisms might be seen as
300 'keeping in the same place', as a law is unchangeable. However, the
structuralist conception of life especially focuses on an ascending hierarchy of
14 taxa of ever widening comprehensiveness: as such, one can envision evolution
as a constant movement driven by the laws of nature. Moreover, by displaying
'adaptive masks', which by definition are changeable because they are related
305 to changing environments, living organisms lead us to the same conclusion as
before.
Finalism and teleology.
In his “Science Delusion” [25], Rupert Sheldrake proposes that evolution
310 might not be aimless, contrary to what the standard evolutionary theory
blatantly states. Such an assertion is mainly rejected by orthodox scientists,
because of so-called epistemological discrepancies . Nevertheless, the question
remains open, because it is basically impossible to demonstrate whether
evolution is aimless or not. However, in the case that evolution might be
315 finalistic or even teleological [26], then it is not possible to envision organisms
keeping in the same place. Indeed, it has never been demonstrated that
evolution is not driven by final causes: this idea is merely considered as a less
parsimonious hypothesis, which in addition would 'sterilise' any scientific
investigation. In fact, the absence of final causes is simply one of the very
320 fundamental specifications of orthodox science, an axiom, a postulate, a self-
evident truth that requires no proof. Nevertheless, according to James Shapiro,
“cells are now reasonably seen to operate teleologically: their goals are
survival, growth, and reproduction” [26], which is a genuine 'game changer',
as Carl Woese stated about Shapiro's book.
325 Similarly, in “The Long Chain Of Coincidences”, Michael Denton [27]
15 emphasises the huge amount of adequations that can be described in the
Universe, and thus intends to demonstrate that the Universe is biocentric,
another assertion violently rejected by orthodox scientists. It should be
stressed that these coincidences are truly countless: the fundamental
330 constants and forces of the Universe, the nuclear structure of some atoms
(especially the carbon atom), the relative abundance of chemical elements, the
many physico-chemical properties of some atoms (again specifically including
the carbon atom), of dioxygen, of water, of carbon dioxide, of visible light, of
metals, and so on. The list seems to be endless. In the view of a biocentric
335 Universe, living organisms once again certainly cannot be seen as 'keeping in
the same place', since their very existence and evolution would be due to the
fundamental laws of the Universe. In such a perspective, the evolutionary
process itself cannot fit the standard implications of the RQ.
340
16 TOWARDS A WIDER, MORE COMPREHENSIVE UNDERSTANDING OF
BIOLOGICAL EVOLUTION.
In this wider interpretation of evolutionary facts, it can be seen that
345 evolving organisms do not 'keep in the same place', whether it be at the
organismal level, the species level or the evolutionary process level: as they
evolve, organisms progress, species progress, they can follow trends, for
example towards higher (or lower, as is frequently the case in parasitism)
complexity, and so on. Moreover, evolution can sometimes lead to speciation:
350 when a new species appears, is it really possible to assert that the organisms
have kept in the same place?
In the coevolution of pollinating Insects and flowering plants
(Angiosperms), the huge diversity of Insects and Angiosperms in nature could
not be explained if they had 'kept in the same place'. In the many examples of
355 coevolution between parasites and hosts [6], could the huge amount of
attacking and defensive strategies be explained if these parasites and their
respective hosts had 'kept in the same place'?
Evolution is also related to migrations. Let us consider flamingos [28].
They live in huge groups of several hundreds of thousands of individuals,
360 sometimes up to one million, as is the case of Lake Natron flamingos. There,
after the young birds are born (up to 500,000 of them), the whole group starts
a special kind of bird migration, not flying, but walking (because young birds
are unable to fly). This special migration can be envisioned as a sort of
selection process. Indeed, during their long trip (dozens of kilometers),
17 365 thousands of young birds die: some are attacked by predators (such as
marabou storks and mongooses), others are slowed down by the salt sticking
to their legs, forming a deposit that becomes thicker and heavier as the birds
keep walking in the heavily salted water. In the process, it can be hypothesized
that the baby flamingos that die are the weakest, the slowest, the less
370 resourceful ones (and so those that potentially do not have the best genes),
allowing the other flamingos to survive. As cruel as it may appear at first
glance, this migration leads to an improvement of the species as a whole, by
eliminating those individuals that are the least well adapted. Thus, by walking
long distances, the flamingo species evolves. And so, once more, they cannot
375 be said to stay in the same place.
As far as sexual reproduction is concerned, it has always been considered
as a huge improvement over asexual reproduction, since sexual reproduction
produces novelty among living organisms through meiosis and fecundation, the
two major steps in such a mode of reproduction, even if sex might have been
380 originally 'invented' for quite a different 'purpose' (Lodé).
18 CONCLUSION – PROPOSING A NEW NAME FOR THE HYPOTHESIS.
385
For all the previous reasons, I consider the name of the RQ hypothesis to
be wrong. Evolution has nothing in common with Lewis Carroll's character. It
should be emphasised that for some authors like Conway-Morris, evolution
appears to be mostly convergent and, in a way, inevitable (evolutionary
390 novelties as inevitabilities; [21]). If it is the case, then the RQ hypothesis
would simply have to be renamed. Originally, the RQ emphasised the
constancy of extinction rates. For that reason, I propose that it should be
called the 'Constant Gardener' hypothesis.
This proposition refers to a novel by John le Carré [29], whose title
395 denotes the main character, Justin, who is determined to grow things, literally
and figuratively: Justin gardens as a hobby but also constantly digs for the
truth after his wife's murder, attempting to bring information to light in an
endless investigation in the pursuit of the truth; hence, The Constant Gardener.
In gardening, sometimes you have to get rid of weeds or animals harmful to
400 the growth of your crops.
The human elimination of these living organisms can be paralleled with
species going extinct in nature over geological time. As the Red
Queen/Constant Gardener hypothesis postulates, this appears to be a constant
process in nature. But this has nothing in common with living organisms
405 'staying in the same place', whether it be at the organismal level, the species
level or the process level.
19 This new name seems more appropriate. The Red Queen character is
some kind of completely unrealistic, totally imaginary fantastic figure. On the
contrary, Justin, the Gardener, is a human being, fictional but realistic.
410 Moreover, the proposed new name appears less metaphorical, more concrete,
less likely to lead to misinterpretations. In such a perspective, evolution on the
whole can be seen as a law-like, uniform process in time and space, in relation
with uniformitarianism.
However, new objections might arise, because this proposal could lead to
415 other misinterpretations: more specifically, the survival or elimination of
species might be attributed to a unique external active entity, some kind of
'divine gardener' that would make decisions according to its own interests,
independently of the adaptive capacities of each living species – a return to
Creationism I absolutely do not intend to support.
420 I will finish with a more general remark about metaphors in science,
which, although quite useful, are pervasing throughout the scientific literature
[30]. Metaphors are contested by some scientists but seem to be vital to the
advancement of knowledge. On the one hand, metaphors can be suspected to
introduce a subjective aspect into scientific theories. On the other hand,
425 metaphors can be seen as a creative element in the search for knowledge,
allowing the rise of whole new ideas and the predictive dimension of scientific
theories [30]. As far as the RQ is concerned, it appears to have often been too
literally, too metaphorically interpreted: eventually, the issue does not appear
to be about 'keeping in the same place', but about survival. What the Red
430 Queen answers to Alice may in fact mean something like “Here, we constantly
20 have to do our best to survive”, some sort of “lesson in life” just like the many
initiatory tests Alice has to face along her allegoric adventures in her transition
from chilhood to adulthood. In a way, the RQ metaphor can appear as not so
misleading, because any species unable to evolve will inevitably go extinct.
435
* * *
* *
*
21 440 ACKNOWLEDGEMENTS.
I thank Dr. Joël Espesset for his careful reading of the manuscript (for
English accuracy), and Dr. Thierry Lodé for his critical reading of a previous
draft of the manuscript and his numerous advice about the publication process.
445 I also thank Dr. Grégoire Espesset for his close reading of the manuscript, and
his numerous, precise and constructive criticisms, and Dr. Virginie Orgogozo for
helpful exchanges.
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