Macroevolution vs micro evolution

Eduardo Aguirre-Mazzi

In a broad terms, Iurii Filipchenko defined microevolution as the evolution of biotypes and
macroevolution is defined as the evolution of higher systematic groups (reviewed in Adams 2021).
He recognizes that microevolution could be understood in the eye of genetics (mutations and re-
combinations) but a different approach other than genetics is required to understand
macroevolution. Filipchenko argues that the questions related to the origin of species (and inferior
subdivisions) will be resolved in a separate way than the questions related to the origin of genus (and
higher-level taxa). Theodosius Dobzhansky supports this perspective, where he recognizes that
macro- and micro-evolutionary changes are distinguishable, and that evolution must imply more
than species formation addressed by geneticist of his time (Dobzhansky 1937; reviewed in Adams
2021). He also states that no mechanistical understanding macroevolution on geological time scale
was proposed at his time, other than microevolutionary scale theories focused on organisms at
human lifetime scale. Consequently, Dobzhansky emphasizes the necessity of distinguishing
mechanisms of macro- and micro-evolution and that thinking that species have arisen only through
evolution by natural selection was problematic (reviewed in Adams 2021).

Despite Dobzhansky and Filipchenko (his mentor), agreed to distinguish microevolution and
macroevolution. Dobzhansky had a fundamental difference that led him having a key role in the
establishing of population genetics. For instance, he stated that “Since evolution is a change in the
genetic composition of populations, the mechanisms of evolution constitute problems of
population genetics” (Dobzhansky 1937: 11; reviewed in Adams 2021). Therefore, in this way he
contributed to the explanatory reductionism present in the understanding of evolution defined as
microevolutionary change at the level of genetic composition of populations.

Following Dobzhansky’s claims about what evolution is, a series of scientists started to criticize and
enrich this debate. Among these, Richard Goldschmidt, considered that macroevolution was the
central problem needing explanation and that Dobzhansky’s was somehow a reductionist.
Specifically, Goldschmidt challenged the Darwinian view, by asking: How evolution as a process
merely based on accumulation and selection of small mutants could explain: hair in mammals,
compound eyes and the transformation of the gill arches in phylogeny including the aortic arches,
among others? (Reviewed in Adams 2021) In his written work, Goldschmidt also distinguishes
microevolution from macroevolution, and he argues that to understand the decisive step in
evolution, a broader evolutionary method is required than studying the mere accumulation of micro
mutations.

In a comparable way, Émile Guyénot’s “L’Origine des Espèces (1944)” acknowledges that nothing
positive was known about the genesis of groups larger than species, and that neither genetics nor
embryology would allow us to understand underlying mechanism explaining how organisms at
higher taxa (e.g., sub-kingdoms) came to be differentiated (reviewed by Adams 2021). Guyénot then
makes an interesting distinction between “superficial evolution” treated by population genetics and
the “evolution in depth” (in other words macroevolution) related to variation in basic structural forms
and body plans (reviewed by Adams 2021). Thus, again he suggested that population genetics and
natural selection (I.e., Darwinist approach) where not appropriate to address the questions
pertaining to macroevolution.

Later Huxley’s “Evolution: The Modern Synthesis” of 1964 (reviewed by Adams 2021), does not seem
to use the terms microevolution and macroevolution nor to take an explicit position within this
debate. However, in contrast to the microevolutionary reductionism of population genetics, Huxley
states the following: “The formation of many geographically isolated and most genetically isolated
species is thus without any bearing upon the main processes of evolution.” Following, he argues that
such a process implies the development of new types provided with mechanisms encompassing
biological efficiency in the realization of adaptive radiation, thus, allowing types to take advantage
of the vast variety of environments and modes of life. Essentially, this process might involve a
progressive increase of life’s control over and independence of the environment. Interestingly, he
adds that minor systematic diversity, at level of species formation (I.e., microevolution), is just a
minor aspect of the major and continuing trends of the evolutionary process (I.e., macroevolution).

Regarding the founders of population genetics Ronald Fisher, Sewall Wright, “Jack” Haldane, and
Sergei Chetverikov, and whether they were or not convinced that macroevolution had been reduced
to population genetics; They almost fully accepted Darwin’s views. However, to start with Fisher, he
does not seem to directly address this discussion, but he proposes that among the various theories
of evolution, the theory of natural selection must be accepted due to the invalidation of the concept
of blending heredity required by other theories. Nevertheless, he also admitted that we might just
don’t yet know if there is another plausible better fitted theory (Fisher - “The Genetical Theory of
Natural Selection 1930; Reviewed by Adams 2021). In a later chapter of this book: “Fission of
Species,” he simply assumes that the morphological differences between species are the result of
the accumulation of Mendelian mutations in populations. In contrast, Wright’s book, “Evolution in
Mendelian Populations (1931)” concludes that the observation of naturally occurred gene mutations
are not sufficient to explain the adaptive evolutionary process. In a later, International Congress of
Genetics at Cornell (1932), Wright also argues that: “It is only at the subfamily and family levels that
clear-cut adaptive differences become the rule”. This suggested that the mechanism that originate
species must be a nonadaptive one (Adams 2021). Therefore, the position of Wright seems to
suggest that Macroevolution goes beyond the population level at which natural selection operates.

Consistently with Wright, Haldane’s “The Causes of Evolution (1932)” acknowledges that selection
acting on populations is not itself evolution, and that species specificity was not determined by
Mendelian genes. In this same book he defines evolution as “the descent from living beings in the
past of other widely different living beings” (Haldane 1932; reviewed by Adams 2021). Apparently,
neither Fishcher, nor Haldane and nor Wright confronted explicitly the question on the relationship
between macroevolution and population genetics. Nonetheless, Sergei Chetverikov, a prominent
entomologist and butterfly taxonomist, was the only one to tackle this issue in a more direct way. In
his 1926 book, “On Certain Aspects of Evolution From the Viewpoint of Modern Genetics” he argues
that traits determined by Mendelian genes in Drosophila (e.g., venation and wing structure) are
fundamental in modern systematics for distinguishing higher systematic categories (reviewed by
Adams 2021). Chetverikov’s position was likely to be influenced by the strong tendency that Russian
geneticist and evolutionist to think that “microevolution” and “macroevolution” were the same sort
of thing. He also provides evidence of this based on his own experience in the study of butterflies
and other insects.
A key concept to understand the debate of macroevolution and microevolution is the idea of
evolvability. In general, it seems that the concept of evolvability is more related to processes of
macroevolution rather than to processes of microevolution. For instance, Newman and Muller
(2000), define evolvability as “the inherent potential of certain lineages to change during the course
of evolution” … “For us evolvability represents the continued efficacy of epigenetic processes in a
lineage—some of them quite ancient, and some of more recent origin—and as such is tied to the
primitive morphogenetic plasticity…” (Newman and Muller 2000). Interestingly, then they argue that
cooptative genetic evolution is likely to buffer the development of form limiting the evolvability. The
definition above is completely framed in the macroevolutionary realm. Nevertheless, the conceptual
spectrum of evolvability reviewed by Pigliuchi’s (2008; table 1), in a sense encompasses both micro-
and microevolution. For instance, on the microevolutionary scale, evolvability is equivalent to
Houle’s definition of heritability, and it is described as the standing pool of genetic variation that
determines the response of organism to natural selection and operates at the level of population
genetics (Pigliuichi 2008).

In contrast, on the macroevolutionary scale, the two following definitions reviewed by Pigliuchi’s
(2008; table 1), are more related to the ability of organisms to produce largest changes that are
compatible with survival. In other words, as the potential of generating new lineages. The evolvability
defined by Wagner and Altenberg is related to evolution of species and the changes that produce
long-term adaptation and non-random trajectories through exploration of phenotypic space.
However, these changes might operate in relatively shorter periods of time and are constrained by
genetic architecture and developmental constraints. In contrast, the Evolvability related to Maynard-
Smith and Szathmary’s concept of innovation, operates at the level of the evolution of higher taxa
and can give rise to major phenotypic changes (morphological, behavioral, or physiological). These
major changes could imply an overcome of genetic and developmental constraints even allowing to
explore new areas of the phenotypic space.

The evolvability concept related to the propensity of organism to give raise to new forms by any
means (e.g., genetic mutations, epigenetic or developmental changes) goes beyond selection of
standing variation. Alberch (1991), in accordance with Dawkins, states that evolvability is a property
of embryological systems, and that certain developmental systems are better at evolving (e.g.,
segmentation, sequestration of the germ line). Apparently, such developmental systems are key in
the evolutionary proliferation of lineages (Alberch 1991). Is interesting to note that, Alberch was
interested in how variability of shapes and functions is created in macroevolutionary dimensions
rather than mere local adaptations across geography. For instance, he was known for exploring the
possibilities of developmental systems to produce variations despite some of these variations were
deleterious (Alberch 1989).

In this context, evolvability can favor plasticity and be recognized as a product of advanced evolution
(Newman and Muller 2000). Evolvability is not associated with selected genetic mechanisms to
surpass genetic determination, but rather represents the historic carryover of epigenetic
determination that could have had even greater morphogenetic plasticity (Newman and Muller
2000). In this line, any pattern generating system (=developmental process) has a limited parameter
space associated with it. Therefore, the evolvability potential can be defined by the global properties
of the dynamical system as described by its own parameter space (Alberch 1991). For example,
within a species-specific transformation diagram (Alberch 1991), the closeness to the boundaries of
phenotype is related to the ability of the system to generate new variation. As a final though here, I
can add that understanding evolvability will help to understand how the organismal variability arises,
and then complement the reductionist view of population genetics rarely interested in studying the
mechanisms by which variation arises.