Biology and Philosophy (2006) 21:41–70
Springer 2006

DOI 10.1007/s10539-005-0957-4

The notion of progress in evolutionary biology – the

unresolved problem and an empirical suggestion

BERND ROSSLENBROICH1,*
1
Institute of Evolutionary Biology and Morphology, University of Witten-Herdecke, D-58453 Witten,
Stockumer Street 10, Germany (e-mail: rosslenbroich@uni-wh.de; phone: +49-2302-669-344)

Received 2 October 2003; accepted in revised form 20 January 2005

Key words: Biological autonomy, Complexity, Directionality, Epistemology, Evolutionary pro-

gress, History, Macroevolution, Trends

Abstract. Modern biology is ambivalent about the notion of evolutionary progress. Although most
evolutionists imply in their writings that they still understand large-scale macroevolution as a
somewhat progressive process, the use of the term ‘‘progress’’ is increasingly criticized and avoided.
The paper shows that this ambivalence has a long history and results mainly from three problems:
(1) The term ‘‘progress’’ carries historical, theoretical and social implications which are not con-
gruent with modern knowledge of the course of evolution; (2) An incongruence exists between the
notion of progress and Darwin’s theory of selection; (3) It is still not possible to give more than a
rudimentary definition of the general patterns that were generated during the macroevolution of
organisms. The paper consists of two parts: the first is a historical overview of the roots of the term
‘‘progress’’ in evolutionary biology, the second discusses epistemological, ontological and empirical
problems. It is stated that the term has so far served as a metaphor for general patterns generated
amongst organisms during evolution. It is proposed that a reformulation is needed to eliminate
historically imported implications and that it is necessary to develop a concept for an appropriate
empirical description of macroevolutionary patterns. This is the third way between, on the one
hand, using the term indiscriminately and, on the other hand, ignoring the general patterns that
evolution has produced.

Introduction

Our general view of the large-scale evolutionary process is that the earliest
forms of life were prokaryotes followed by the first eukaryotic cells which
formed multicellular organisms. The Cambrian ‘‘explosion’’ added new forms
of life with hard skeletons, completely changing the fauna of the world. Within
the then existing phyla a great variety of changes led to the forms of today
which include bees, squids, frogs, crocodiles and horses. Every evolutionary
biologist feels that there were profound changes and innovations during this
process that need to be described. Traditionally, these changes have been
termed ‘‘evolutionary progress’’. But in recent years this term has been
increasingly criticized, and some authors claim that it has now been success-
fully eliminated from evolutionary biology. However, on closer examination
this appears not to be the case. There are hardly any textbooks that can avoid
using the terms ‘‘lower’’ and ‘‘higher’’ when referring to organisms. Further-
more, many phylogenetic reconstructions, especially on the level of phyla,
42

include sequences that lead to ‘‘advanced’’ forms in the traditional sense.

Usually this is also the sequence of the major groups in which organisms are
arranged in zoology or paleontology textbooks. The criticism of this notion in
recent years has had the effect that people try to avoid using the terms that
refer to it or they explicitly distance themselves from it, although nearly
everyone still thinks of evolution in terms of overall progression.
The following paper shows that this striking inconsistency is deeply
embedded in the history of evolutionary thought and its paradigms. It is
proposed that epistemological reflection is needed in order to eliminate his-
torically acquired baggage and to provide an adequate scientific description of
the underlying phenomena. The question will not be ‘‘Why do evolutionists use
such unscientific terms?’’ (Ruse 1996), but rather, if evolutionists cannot avoid
the term, what do they see in their daily work and how can they express their
observations in a scientific manner?

Three historical roots of the term ‘‘progress’’

The term ‘‘progress’’ has predominantly three historical roots. One is the
concept of the scala naturae, which until the 19th century was the most
widely prevalent view of the general order of the world. It saw the world
arranged in a linear hierarchy and was originally a static concept, but
during the late 18th and early 19th centuries it was ‘‘temporalized’’ (Lovejoy
1974), so that its elements would appear in succession. The second root lies
in the notion of social and cultural progress that developed during the
Enlightenment and gradually replaced the notion of the invariability of
human affairs. During the late 18th century it expressed the emerging
consciousness of the capacity of mankind to improve its circumstances and
abilities (Kosellek 1975; Rapp 1992). In these early considerations, progress
included the aim of an achievable perfection, what introduced a strong
teleological element. Critical reflection on the questions of change, devel-
opment and progress in human history, including its problems of linearity
and teleology, took place in Germany during the 18th and early 19th cen-
turies. Goethe and Kielmeyer were among the first to apply the new idea to
nature (Schad 1998). During the 19th century the general progress of
society, science, technology and industry was taken for granted, especially
for example in England (Bowler 1989a, b). The third root is the theory of
recapitulation, the analogy that was drawn between embryogenesis and
phylogeny. Knowledge of the embryo, developing from a simple to a
complex structure, was an intellectual help for initial ideas about the
changeability of organisms (Richards 1992).
All three roots were important in preparing the evolutionary thinking that
was increasing during the 18th century. They led to viewing evolution as a
predominantly linear progression, aiming at some form of perfection, and, like
the development of the embryo, driven by a certain force.
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Different components

For a further discussion of the function of the notion of progress in evolu-

tionary biology it is necessary to reflect on the different components and
connotations that may be involved in varying combinations and that derive
from this historical background. At least five components must be distin-
guished:
(1) Modifications in the living world generate increasingly higher organisms
(however they are characterized).
(2) These higher organisms are in certain ways better than lower ones
(=‘‘improvement’’).
(3) This progression is more or less linear.
(4) Evolution has an inbuilt force that drives this progress.
(5) Progressive evolution leads eventually to some sort of perfection (end
stage, culmination point, goal).
In the critical literature of recent decades, these components are often mixed
together, contributing to the confusion. For example, it is often assumed that
the notion of progress is always looking for driving forces in evolution (com-
ponent (4)). Or it is supposed that the view that evolution is progressive implies
a goal towards which the process is moving, thus making evolution a teleo-
logical concept. This supposition would be a combination of components (4)
and (5). But these components are not necessarily involved. It is true that most
biological thinking before Darwin’s theory of evolution was introduced into
science included these components, but afterwards the picture became more
varied.

Charles Darwin

How did Darwin deal with the notion of a progressive, goal-directed and linear
understanding of organic change? Much has been written about this topic
(including for example Mandelbaum 1971; Bowler 1983, 1989a; Richards 1992;
Rieppel 1992; Ghiselin 1995; Ruse 1996; Gould 2002). Applying the above
components to his central and epoch-making work, the The Origin of Species
published in 1859 (Darwin 1872, in the following text cited from the 6th edi-
tion), Darwin unequivocally disapproved of any idea of an inbuilt force that
was supposed to drive evolution. That the process should have a goal was also
incompatible with his theory and was explicitly refuted, as was the idea of
linearity in evolution. He argued repeatedly throughout the book against
contemporary advocates of such views, and their refutation was one of his
main concerns, an important achievement in his time.
But the problem whether the evolutionary process might generate higher
organisms is a complex one in Darwin’s thinking and hides a dichotomy. His
theory is mainly an explanation of how populations adapt to their changing
environments and to their biotic factors. The theory of natural selection holds
44

that in the struggle for existence, those individuals best adapted to new con-
ditions will survive and reproduce, while others less well adapted will be
eliminated. Over many generations, the adaptive character is enhanced until
eventually the population becomes a new species, incapable of interbreeding
with the parent form. Neither the variations in characters from which an
adaptive character is selected, nor the environmental changes, include direc-
tionality. Therefore this process can deliver only a set of meandering responses
in the adaptive adjustments of organisms to local environments (Bowler 1989a;
Gould 2002). This process would lead to ever increasing divergence of forms
independently of one another, depending upon the respective selection factors,
and, result in a network of adaptations to the respective vicissitudes of the
struggle for life. But according to the traditional understanding of progress, the
process does not include any directionality. In addition, it seems not to make
sense to compare different ‘‘levels of organization’’ when the main reality is
branching evolution.
However, through competition between individuals and victory over other
creatures in the struggle for limited resources, direction may be involved
(Gould 2002). Darwin expected an accumulation of ‘‘improvements’’ from the
struggle which would make organisms fitter and thus generate progress. ‘‘Now
species triumph because, in some sense admittedly difficult to define, winners
are ‘better’ than the forms they vanquish. And the more uniformitarian the
larger picture – the more that macroevolutionary pattern arises as a simple
summation of immediate struggles – so do we gain increasing confidence that
replacement and extinction must record the differential success of globally
improved species.’’ (Gould 2002, p. 475).
In a paragraph ‘‘On the Degree to which Organisation tends to advance’’,
Darwin (1872, pp. 127, 228) wrote: ‘‘Natural Selection acts exclusively by the
preservation and accumulation of variations, which are beneficial under the
organic and inorganic conditions to which each creature is exposed at all
periods of life. The ultimate result is that each creature tends to become more
and more improved in relation to its conditions. This improvement inevitable
leads to the gradual advancement of the organisation of the greater number of
living beings throughout the world…’’ … ‘‘Although we have no good evidence
of the existence in organic beings of an innate tendency towards progressive
development, yet this necessarily follows … through the continued action of
natural selection. For the best definition which has ever been given of a high
standard of organisation, is the degree to which the parts have been specialized
or differentiated; and natural selection tends towards this end, inasmuch as the
parts are thus enabled to perform their functions more efficiently.’’
Thus increased specialization and differentiation of parts make their bearers
superior to other ones in the struggle for life.
‘‘If we take as the standard of high organisation, the amount of differenti-
ation and specialization of the several organs in each being when adult (and
this will include the advancement of the brain for intellectual purposes), nat-
ural selection clearly leads towards this standard: for all physiologists admit
 45

that the specialization of organs, inasmuch as in this state they perform their
functions better, is an advantage to each being; and hence the accumulation of
variations tending towards specialization is within the scope of natural selec-
tion.’’ Darwin (1872, p. 128).
Thus Darwin explained not only adaptation to the immediate environment
but also the gradual progress through the selection process.
‘‘Thus, from the war of nature, from famine and death, the most exalted
object which we are capable of conceiving, namely, the production of the
higher animals, directly follows.’’ (Darwin 1872, p. 560). How Darwin envis-
aged large-scale ‘‘improvements’’ of organs and higher animals going together
with increasing fitness for survival is shown, for example, in his paragraph on
eyes (Darwin 1872, p. 188).
But a gap in this extrapolation from microevolutionary adaptive processes to
large-scale macroevolutionary progress is that the species, which are suppos-
edly more advanced, would not necessarily have an enhanced capacity for
survival. If continuous ‘‘improvements’’ accumulate towards progressive
forms, for example through the generation of complex organs as Darwin ex-
pected, the bearers of those improvements must finally be the fittest organisms,
which is not the case in nature. Bacteria as well as protists had enough fittness
to survive for a longer time than vertebrates yet. The accumulation of complex
organs and functions from single cells to vertebrates (which is usually referred
to as progress, also by Darwin) delivers anything but enhanced survival
capacity. Thus there is an incongruence between on the one hand the micro-
evolutionary adaptation process that leads to fitness, and, on the other hand,
what is traditionally called progress in the large-scale, macroevolutionary
outcome of evolution. However, this does not question the validity of either of
the two principles but only states that their relationship is not clear within the
original Darwinian scenario. It also does not mean that there might not be a
resolution of the incongruence, but one has not been definitively established as
differing views prevail, which basically still holds true.
It is hard to judge how clearly Darwin saw the incongruence, but his
ambivalence concerning the term ‘‘progress’’ may hint at his struggle with it.
Regularly quoted is his warning to himself never to speak of ‘‘higher’’ or
‘‘lower’’ which even he repeatedly disregarded. In a few passages of the
‘‘Origin’’, Darwin is guarded in his comments about large-scale progress: ‘‘The
problem whether organisation on the whole has advanced is in many ways
excessively intricate…’’ (p. 403). Some discussion follows showing that the
topic is a complex one for Darwin too. The passage from p. 127 on
‘‘advancement of organisation’’ quoted above continues with the sentence:
‘‘But here we enter on a very intricate subject, for naturalists have not defined
to each other’s satisfaction what is meant by an advance in organisation.’’
Darwin might have seen the incongruence, but could not solve it within his
theory.
The underlying incongruence has given rise to different reactions, as shown
by the following examples. Williams (1966, p. 34) wrote that there is nothing in
46

the basic structure of the theory of natural selection that would suggest any
kind of cumulative progress. Maynard Smith (1988, p. 220) stated similarly
that ‘‘… our theory of evolution does not predict an increase in anything’’, and
Wake (1986, p. 48) stated in a discussion report that ‘‘we have no a priori
expectation from current theory that evolution should be directional or pro-
gressive’’. This is clearly not what Darwin expected from his theory. He knew
that the obvious differences between organisms had to be explained. For his
argument it was important that between lower and higher organisms it was
possible to discern a difference within which an evolutionary process could take
place. The Berlin naturalist Ehrenberg, for example, had conducted studies of
microorganisms and these fascinated Darwin. Ehrenberg promulgated the view
that there are no differences in degree of perfection between the simplest and
the more complex organisms, because both extremes possess the structures
necessary to fulfill their life functions, a view with a strong anti-evolution
tendency. An application of this argument to evolutionary theory would render
any improvement of organisms in the struggle for life superfluous. Lyell also
tended to object to the idea of their being a progressive change in the pale-
ontological record (Mayr 1982).
Darwin’s ambivalence towards the notion of progress (Greene 1991; Gould
2002) has been discussed by several authors, albeit with varying presupposi-
tions and often without distinguishing the five components listed above. Some
authors describe him as a ‘‘progressionist’’ whose considerations largely con-
curred with the current views of his time (Greene 1981; Ospovat 1981; Richards
1992; Ruse 1996). Many 20th century scientists regarded the variational aspect,
with its focus on natural selection, as the essence of the theory and claimed that
Darwin included some progressionism only because he either was subject to
typically Victorian attitudes and was therefore unable to appreciate the more
radical implications of his theory, or possibly knew about the more radical
consequences, but introduced some progressionist aspects because he felt that
otherwise his theory would be difficult to accept (Nisbet 1969; Bowler 1989a;
Gould 2002). Another interpretation is that his continued belief in progress was
a function not of his theory but of his belief in a divine purpose (Ospovat 1981;
Brown 1986). This would, however, lead into the complex topic of Darwin’s
religious views.
Instead, I propose a quite different alternative. Clearly Darwin was a thinker
of his time and he used contemporary social thoughts and standards for his
biological interpretations (Desmond and Moore 1991). But at the same time he
was an excellent empiricist and very concerned with the details of his theory, as
(Gould 2002, p. 499) pointed out: ‘‘Darwin … was dogged and relentless,
fiercely honest and logical in his thinking. He wrestled with every major dif-
ficulty, working and reworking, fretting and fretting again, until he achieved
closure or at least understood why a solution eluded him…’’. Against this
backdrop I cannot see any reason to allege that Darwin was merely unable to
escape from views of his time or that he included theoretical elements just for
tactical reasons. Neither can I see how it should be possible to extract from
 47

Darwin’s work a ‘‘true essence of Darwinism’’ (Bowler 1989a, p. 7) without the

notion of progress. When Darwin’s theory is read as it was presented in his
‘‘ambiguous book’’ (Ruse 1996, p. 172), it includes a dualistic tension between
populational thinking and progressive thinking, two not necessarily irrecon-
cilable but instead complementary perspectives of an extremely complicated
matter. Ruse (1996) provided a good insight into how Darwin wrestled with on
the one hand the notions of directionality and progress, and, on the other
hand, his principle of an ever branching and diverging evolution. Darwin’s
theory abandons elements of linearity, an inbuilt force or a goal of evolution,
but still makes an attempt at explaining ‘‘organisational advance’’, as he for-
mulated it. I maintain that Darwin saw this tension but attempted to deal with
the observable differences between organisms rather than focus on a more
radical and reductionistic theory.
Since Darwin, there have been different reactions to this incongruence, which
is usually not made explicit. Is there any trace of progress in the evolutionary
process or should we stop using the term (Haldane 1932; Goudge 1961; Wil-
liams 1966; Gould 1988, 1996; Ruse 1988, 1996)? Is progress only an irregular
byproduct of the adaptive process while the main reality of evolution is the
generation of ecological networks (Wieser 1994)? Does the process of selection
lead naturally towards progressive forms (Huxley 1948, 1953; Thoday 1953,
1958; Rensch 1959; Mayr 1982, 1988; Bonner 1988; Wilson 1992)? Must pro-
gress be seen as a value judgement and therefore as unscientific, but when it is
explicitly used as a non-scientific term, it may have a place in evolutionary
biology (Herrick 1956; Goudge 1961; Dobzhansky 1970; Simpson 1971; Ayala
1974, 1988)?
However, the view that progress might be a simple accumulation of fitness is
surely not necessarily an element of modern Darwinian thinking, but the
relationship between fitness and progress has remained unresolved (Wicken
1979; Saunders and Ho 1981). Gould (1996 p. 199) pointed to this relationship:
‘‘I have long been entirely underwhelmed by the standard arguments for
general advantages of increasing complexity in the Darwinian game – adaptive
benefit of more elaborate bodily form in competition for limited resources, for
example. Why should more complex conformations generally prevail? … I can
envisage just as many situations where more elaborate forms might be a hin-
drance – more parts to fail, less flexibility because all parts must interact with
precision.’’ Remember that Darwin expected functional improvements through
the building of more complex organs, and today complexity is often equated
with progressiveness (see below).
This is also a part of Darwin’s problematical extrapolation from the level of
populations and species, predominantly studied within modern biota, to large-
scale macroevolutionary patterns (Peters 1972; Stanley 1979; Eldredge 1985;
Levinton 1988; Carroll 1997). Carroll (1997, p. 2) summarized it thus: Darwin
‘‘argued that both the patterns and processes of evolution were essentially
identical over … vastly different time scales. Although many biologists and
popular textbooks since the time of Darwin have perpetuated this concept of
48

the history of life, other scientists have argued that neither the patterns nor the
processes of evolution that can be studied in living populations are adequate to
explain the conspicuous differences in morphology, physiology, and way of life
of the major groups of microorganisms, plants, and animals, or the major
patterns and different rates of evolution observed in the fossil record.’’
Darwin’s ambivalence and the incongruences in his theory led to diverging
attitudes amongst many scientists, thus establishing at least two fundamentally
different views of evolution which at any given time have remained more or less
divergent. This pervades all of evolutionary biology with a spectrum of opinion
ranging from the presupposition that evolution generated progress, to a
complete denial of any form of progress in it. The history of ideas after Darwin
shows how these different perspectives and their dualistic tension have always
been at work.

The post ‘‘Origin’’ era

The publication of the ‘‘Origin’’ led to a quick and broad acceptance of evo-
lution within the scientific community but the elements of Darwin’s views have
quite different histories of acceptance (Mayr 1982; Bowler 1983, 1989a; Jahn
2000). Nearly all theories during the second half of the 19th century and the
early decades of the 20th century took progressive evolution for granted, but
differed according to which components of progress they included (Bowler
1983, 1989a). Most authors preserved the view that there is some form of
central element running through the process towards a single goal and they
attributed it to a variety of causes, while only a few authors were interested in
the selection principle. Haeckel in Germany included in some of his illustra-
tions a strong linear element towards progress along a trunk of evolution.
By the end of the 19th century, some alternatives to the Darwinian theory
had gained ground. Amongst the most influential were the orthogenetic theo-
ries, which tried to explain the directional course of evolution through internal
factors that were supposed to drive organisms towards progress. Theodor
Eimer popularized such a version and the ‘‘inner perfecting principle’’ theory
of the Swiss botanist Carl Nägeli was one of the most influential. Ontogenetic
theories retained the analogy of the growth of embryos. Earlier, Agassiz had
postulated a parallelism of both processes as a divine plan of development,
leading to man as the goal. This was picked up later in the century in the form
of the recapitulation theory by, for example, Cope and Hyatt. They combined
this view with a branching process leading not to a single goal but to the
different major groups. The neo-Lamarckian theories – relying much more on
the inheritance of acquired characters than did Lamarck himself – also in-
volved a strong teleological element, as life itself had to be seen as purposeful
and creative. Organisms can choose their response to each environmental
challenge and thus direct evolution through their own efforts (Bowler 1983). It
was especially Weismann who opposed the neo-Lamarckian view when he
 49

insisted that acquired characters could not be inherited and thus founded neo-
Darwinism. These examples show that the components of a progressive, tele-
ological view of evolution were still alive after Darwin.

The Synthetic Theory and thereafter

For the Synthetic Theory, which during the 1940s and 1950s moved the
principle of selection more into the center of the discussion, it was clearly no
longer acceptable for evolution to have a goal. Theories that assumed internal
forces and views that evolution is linear were also rejected. But it was still
assumed that evolutionary modifications drive organisms towards progress.
However, the ambivalence traceable back to Darwin was still fully alive, as can
be demonstrated with a few quotations.
Huxley (1948, p. 556) stated: ‘‘The question of evolutionary or biological
progress remains. There still exists a very great deal of confusion among
biologists on the subject. Indeed the confusion appears to be greater among
professional biologists than among laymen. This is probably due to the com-
mon human failing of not seeing the wood for the trees; there are so many more
trees for the professional!’’. In the text following this introduction Huxley
described evolution as inherently progressive, gradually leading to ever higher
forms of life. He defined progress as the capability of increasing control over
and independence from the environment. Greene (1986, p. 218) commented
that ‘‘Huxley’s writings display in bold relief the tensions, contradictions, and
ambivalences of twentieth-century Darwinism…’’.
Dobzhansky (1970) wrote that progress in evolution is intuitively evident but
that attempts to define what constitutes progress have met with only mediocre
success. A biologist may reasonably speak of evolutionary progress provided
that he makes clear what kind of progress is meant. Simpson (1971, 1973) was
more skeptical about interpreting evolution as a whole as progressive, but saw
some features of progress in certain lines: ‘‘Our inquiry has not produced an
unequivocal definition of evolutionary progress… We may reasonably feel with
Haldane that defining progress takes us into ‘the shifting morass of human
values’ … there is no innate tendency toward evolutionary progress and no
one, overall sort of such progress. We cannot sit back and assume that natural
selection will lead to progress for us, or for anything else.’’ (Simpson 1973,
p. 50).
For Mayr (1982, p. 532) it was obvious that there was an evolutionary
progress that can fully be explained by the principles of mutation and selection:
‘‘… any improvement, any new structural, physiological, or behavioral
invention made in any gene pool can lead to evolutionary success and thus to
progress as it is traditionally defined.’’ Mayr (1988, p. 251) wrote: ‘‘… who can
deny that overall there is an advance from the prokaryotes that dominated the
living world more than three billion years ago to the eukaryotes with their well-
organized nucleus and chromosomes as well as cytoplasmic organelles; from
50

the single-celled eukaryotes to metaphytes and metazoans with a strict division

of labor among their highly specialized organ systems; within the metazoans
from ectotherms that are at the mercy of climate to the warm-blooded endo-
therms, and within the endotherms from types with a small brain and low
social organization to those with a very large central nervous system, highly
developed parental care, and the capacity to transmit information from gen-
eration to generation?’’. Greene (1986, p. 221) again supplied the best summing
up: ‘‘Mayr distinguished between ‘mere evolutionary change’ and progressive
changes involving ‘qualitative improvement of genetic and biochemical sys-
tems’. The ‘final product’ of evolution, Mayr asserted, was ‘perfection in
adaptation’, but were the latest products of evolution any better adapted to
their environments than the earlier ones? Was the modern horse better adapted
to its environment than eohippus was to its environment? Or modern man to
his than Aurignacian man to his?’’
These authors regarded progress as the result of natural selection. Thus
Huxley wrote (1948, p. 568): ‘‘It should be clear that if natural selection can
account for adaptation and for long-range trends of specialization, it can ac-
count for progress too.’’ Stebbins developed this opinion in books such as The
Basis of Progressive Evolution (Stebbins 1969) and Darwin to DNA: Molecules
to Humanity (Stebbins 1982). Rensch (1959, p. 287) coined the term ‘‘ana-
genesis’’ to introduce an element of scientific neutrality into the concept of
progress. He described in some detail the related phenomena he inferred and
attributed them to natural selection.
Until the 1960s and the 1970s there was almost a consensus that natural
selection would lead to ever higher organisms. Although there were critical
statements earlier, this consensus began to collapse with the criticisms of
Williams (1966), Lewontin (1968) and Simpson (1973). Williams (1966, p. 34)
wrote: ‘‘I would maintain, however that there is nothing in the basic structure
of the theory of natural selection that would suggest the idea of any kind of
cumulative progress … I suspect that no one would ever have deduced progress
from the theory itself. The concept of progress must have arisen from an
anthropocentric consideration of the data bearing on the history of life.’’
However, it is characteristic of the opinions about possible tendencies of
progress in evolution during the second half of the 20th century that they were
extremely divergent, even among the more or less Darwinian scientists (Cre-
ationist theories are not considered here, as they cannot be treated scientifi-
cally).
Bonner, for example, made liberal use of the term ‘‘progress’’ in its tradi-
tional sense. In a detailed inquiry on the question of how such progress could
have appeared during the history of life, he discussed complexity and increase
in size as the main characteristics of organismic progress. He referred to
excluding the topic of progress from scientific inquiry as an ‘‘interesting blind
spot among biologists’’ (Bonner 1988, p. 3).
Wilson wrote (1992, p. 187): ‘‘Many reversals have occurred along the way,
but the overall average across the history of life has moved from the simple and
 51

few to the more complex and numerous. During the past billion years, animals
as a whole evolved upward in body size, feeding and defensive techniques,
brain and behavioral complexity, social organization, and precision of envi-
ronmental control … Progress, then, is a property of the evolution of life as a
whole by almost any conceivable intuitive standard, including the acquisition
of goals and intentions in the behavior of animals. It makes little sense to judge
it irrelevant. … let us not pretend to deny in our philosophy what we know in
our hearts to be true.’’. Furthermore, Ruse (1996) published an extensive
survey of modern biologists who still use the notion of progress, whether or not
explicitly.
In recent years, Gould popularized an extreme position on the notion of
progress: ‘‘Progress is a noxious, culturally embedded, untestable, nonopera-
tional, intractable idea that must be replaced if we wish to understand the
patterns of history.’’ (Gould 1988, p. 319). But he also sees that ‘‘… claims and
metaphors about evolution as progress continue to dominate all our litera-
tures…’’ (Gould 1996, p. 21). McKinney (1987) held the opposite view, stating
that ‘‘the history of life embodies improvement, mechanical enhancement,
advancement, or any other synonym or circumlocution’’ all of which he would
have preferred to call progress.
An interesting picture of the completely diverging views appears in the
proceedings of a conference on this topic with the title Evolutionary Progress?
(Nitecki 1988, see review by Greene 1991). This underlines the above statement
that the ambivalence of Darwin concerning the notion of progress continues
into modern biology.
With a refreshing new look at the macroevolutionary process, Conway
Morris (2003, p. XIII) recently opened the next round of discussion by stating
that ‘‘evolution has trajectories (trends, if you prefer) and progress is not some
noxious by-product of the terminally optimistic, but simply part of our real-
ity’’. And on page 307 he stated that we need not ‘‘imagine that the appearance
of humans is the culmination of all evolutionary history. Yet, when within the
animals we see the emergence of larger and more complex brains, sophisticated
vocalization, echolocation, electrical perception, advanced social systems
including eusociality, viviparity, warm-bloodedness, and agriculture – all of
which are convergent – then to me that sounds like progress.’’

Epistemological problems

The following discussion intends to show that much of the historical baggage
carried by the term ‘‘progress’’, leads to it becoming ambiguous. But at the
same time there is an aspect of organic evolution to which the term has been
applied and for which it is necessary to develop an epistemologically satisfying
approach. The persistence in the usage of related terms proves this.
Firstly, the components in the numbered list above require further emphasis.
It should be clear that accepting that large-scale trends reveal increasingly
52

higher organisms (1), does not necessarily include an acceptance of the other
components. Thus, the process does not need to be linear (3). The field of
heterochrony, especially, points to complicated time structures and complex
shifts of differential patterns within evolutionary processes (McNamara 1990;
Schad 1992, 1993, 1997). Equally it presupposes neither some sort of inbuilt
force (4) nor an end stage, or goal perhaps, that supposedly attracts the whole
process (5). But neither does it necessarily include the idea that new forms are
in some sense ‘‘improved’’ or ‘‘better’’ (2); they may just have a different
lifestyle, a different adaptation strategy or a different type of general mor-
phological and physiological organization. This much can be definitively sta-
ted, leaving open for the moment the question of the evolutionary forces that
generated such differences. Fisher (1986) discussed progress as an expected
improvement in organismal design and argued that on neither the microevo-
lutionary level nor on higher levels does the Darwinian process necessarily lead
to improved forms. However, the connection of the two elements ‘‘progress’’
and ‘‘improvement’’ may be deducible from the original Darwinian principles,
although on the large-scale macroevolutionary level, at least, this relationship
is not clear. Forms that are labeled as higher are not necessarily improved in
the sense that the label confers a more general success upon its bearers.
Organisms can be successful in terms of survival on all levels, from single-celled
prokaryotes or eukaryotes to mammals. However, this does not exclude the
possibility that Darwinian principles might stand behind this, but only that the
terms ‘‘progress’’, ‘‘improvement’’ and ‘‘fitness’’ are not congruent. For
methodological reasons it would be more appropriate to differentiate these
elements in order first to identify what is intuitively meant by progress.
In today’s use of the term ‘‘progress’’, when we are simply looking back into
the history of organisms, reconstructing and describing the sequence of basic
changes, components (2) to (5) are not necessarily included. In the history of
theory building, these components were abandoned, although component (1) is
still alive. Thus a modernized interpretation of the term ‘‘progress’’ would
accept only that macroevolution generated forms which increasingly differ
from earlier forms in such a basic way that it is necessary to provide a
description and analysis of general patterns that are involved. For this, the
terms ‘‘higher’’ or ‘‘lower’’, and the term ‘‘progress’’ have until now served as
metaphors.
Another basis for my further discussion comes from a proposal of Rapp
(1992) who distinguished in a study on the term ‘‘progress’’ (including social
progress) a genetic (in the philosophical sense) from a normative form of
progress. Genetic progress is a sequence of steps in time: the succession of
changes and the valueless generation of the new. Over and above this, nor-
mative progress makes value judgements in the sense that every progressive
step includes an improvement with respect to a higher goal towards which it is
worth moral striving. The genetic term is the prerequisite of the normative
term, but the positive value judgement of the normative term may be trans-
ferred to the genetic one, either tacitly or explicitly. However, it is not certain
 53

that genetic progress always leads to improvement. This confusion stems from
the historical link between the terms when they were first used.
The historical overview shows how closely the use of the term in biology has
been connected with the development of social thought and theories. The
notion of normative progress helped people to start thinking in terms of
developments, including those in the organic world. But the normative aspect,
in the sense of moral valuation, cannot be introduced into a scientific context.
However, even the genetic aspect may contain several different elements that at
the same time are possible elements of the term ‘‘evolution’’. This is summa-
rized in Table 1, which is compiled from considerations by Rapp (1992), Le-
wontin (1968) and Simpson (1973).
Change is the basic feature of the evolutionary process: the current state of a
system is the result of one or more changes from a former one. Pure change can
be the raw material of evolution, but for most evolutionists it does not describe
it sufficiently, as change can also be, for example, reshuffling playing cards.
Thus a different order can be generated by a rearrangement of elements. Using
the same basic elements, it leads to the appearance in a system of a new state,
that was not present in the system in its former state. The generation of a new
species from an earlier one, may be the generation of a new state, a new order.
The generation of new order and properties can shift irregularly, or it can be
changed in a certain direction, perhaps over a long time by one or more se-
quences of changes. However, these changes need not be linear. Directionality,
revealed by the fossil record for example, is usually described as ‘‘evolutionary
trend’’. Thus the evolution of early mammals from mammal-like reptiles is
described in paleontology as a sequence of trends that lead to mammalian
characters.
These trends can be described and followed through evolution, but often,
especially in large-scale macroevolutionary trends, a more basic question can
be addressed: did a general difference evolve? Are organisms in later periods
different in some consistent way, in some aspect of their individual charac-
teristics, from those in earlier times (McShea 1998)? The evolution from rep-
tiles to mammals for example involved the generation of a largely different
physiology, allowing for a completely changed lifestyle. Reptiles, being ecto-
thermic, have a relatively low energy budget with restricted movement

Table 1. Possible elements of ‘‘progress’’.

54

capacities, while mammals rely on a high energetic level with a correspondingly

greater capacity for movement. The two forms reveal different sets of char-
acteristics with respect to morphology, physiology, behavior and relation to the
environment. These basic differences of systems are described here as general
patterns, expressing integrative features of large-scale macroevolutionary
trends. Later in the paper attempts will be discussed that have been made in the
literature to describe these patterns (see also Table 2).
Evolutionary theories can be distinguished by how many of the elements
(see Table 1) are included (Lewontin 1968). Some evolutionists only include
change and order, others add directionality. Although large-scale patterns
are rarely addressed explicitly (for example Bonner 1988; McShea 1996,
2001, 2002; Vermeij 1999), they are often embedded in general discussions
and in textbooks. This clearly shows that the understanding of ‘‘evolution’’
to a large extent depends on the perspective on evolution and the para-
digmatic background of the respective researcher. Much of the controversy
around progress has its origin in these different views. In today’s evolu-
tionary biology, large-scale changes of general patterns are the heart of what
is usually called progress. As a term for this is needed, it will not be pos-
sible to eliminate it in the future, just as it has not been possible to elim-
inate it in the past.
Simpson (1973) differentiated progress, which might include a normative
undertone, from progression, which avoids assumptions about any kind of
changes for the better (Table 1). I argue here that today the term is used in the
sense of progression, as no modern scientist would want it to include a nor-
mative judgement.
A common criticism of the term progress is, that it could be anthropocentric.
However, a scientific coverage of large-scale patterns in the evolutionary
process does not need to focus a priori on the characters of man. A large
proportion of the organic world went through an evolution which does not
contain elements of the line towards human beings and so cannot be judged
with criteria that are generated from this line. Here the perspective determines
the traits being observed as well as the systematic level that is chosen. With this
prerequisite, the term ‘‘progress’’ is not necessarily anthropocentric, it just tries
to describe large-scale patterns.

Ontological problems

The course of evolution is not characterized by a process that is directed to-

wards the generation of vertebrates and mammals. Instead, early forms of
organisms were joined by forms with different general patterns. This is the case
amongst not only vertebrates but also invertebrates and plants. But what are
the characteristics of these lineages? What is the essential difference between a
bacterium and a mammal or a squid? This is the question that remains at the
center of this topic. In any case, the obstinacy with which ‘‘progress’’ has
Table 2. Patterns as measures of progress proposed by different authors since 1900.
Patterns
Increasing morphological and physiological complexity. Increasing
structural complication
Increasing differentiation. Increase in the number of cell types.
Increasing specialization of cells
Increasing centralization/differentiation/effectiveness of the nervous
systems
Increasing centralization of systems and organs. Concentration of dif-
ferent parts into new organs
Increasing efficiency of tissues, organs or the whole organism. Biome-
chanical improvement (either in general design or in relation to a
particular ecology)
Increased ability of organisms to obtain and process information about
the environment. Increase in perception of signals from the environ-
ment and ability to react to environmental stimuli. Increased con-
sciousness/knowledge of the environment
Increase in the occupancy of ecospace. Expansion of life as tendency to
fill in all available spaces in the livable environments
Improvement in adaptation. Increase in the possibilities of producing
and maintaining a larger variety of more complex adaptations.
Adaptive success
Discussed by
Bonner (1988), Grant (1985), Günther (1950), Hendrichs (1988), Hennig (1932),
Herrick (1946, 1956), Huxley (1948, 1953, 1959), Kämpfe (1985), Maynard Smith
(1988), Overhage (1957), Plate (1928), Rensch (1959, 1974), Rieppel (1991),
Stebbins (1969) and Stebbins and Ayala (1985); Tatarinov see Urbaneck (1988)
and Wicken (1987)
Bertalanffy (1949), Emerson (1954), Franz (1935, 1967), Gläser (1987), Gruner
(1993), Günther (1950), Kämpfe (1985), Overhage (1957, 1963), Plate (1928),
Remane (1967) and Rensch (1959); Severtsov see Urbaneck (1988), Storch and
Welsch (1989), Vogel (1991), Wicken (1987), Wuketits (1982) and Maynard Smith
(1988)
Bertalanffy (1949), Günther (1950), Grant (1985), Herrick (1946), Kämpfe (1985),
Overhage (1963), Plate (1928), Rensch (1959), Schuh (1937), Waddington (1961a)
and Wuketits (1982)
Franz (1920, 1935, 1967), Günther (1950), Kämpfe (1985), Plate (1928), Remane
(1967), Rensch (1959) and Vogel (1991)
Gould (1988), Gruner (1993), Huxley (1953), Plate (1928) and Remane (1967);
Severtsoff see Urbaneck (1988)
Arnold and Fristrup (1982), Ayala (1974, 1988), Dobzhansky et al. (1977), Grant
(1985), Herrick (1946, 1956), Huxley (1954, 1959), Kämpfe (1985), Schuh (1937)
and Simpson (1971)
Ayala (1974), Dobzhansky et al. (1977), Franz (1920, 1935, 1967) and Grant
(1985; Schmalhausen see Urbaneck (1988), Simpson (1971) and Wuketits (1982)
Goudge (1961), Gruner (1993), Herrick (1946, 1956), Plate (1928), Vermeij (1987)
and Wimsatt and Schank (1988)
55
Table 2. Continued.
Patterns
Improved resistance to extinction. Increasing fitness
Increase in the amount of genetic information. Quantity of coding
DNA.
Succession of dominant types with displacement of others. Domination
of the ecosystem in abundance and variation
Rationalization and economization of structures and functions.
Biological/physiological efficiency
Increasing efficiency in the usage of natural resources
Increasing plasticity of structures and functions. Increased ability to
react to
environmental stimuli. Increasing flexibility of organization
Increasing body-size (Cope’s rule)
Reductions in the number of similar organs (Williston’s rule). Structural
simplification (parts in an organism tend toward reduction in number,
with the fewer parts greatly specialized in function)
Increase in the level of energy processing. Increase in metabolic rate.
Energy intensiveness.
Increasing taxonomic diversity
Increasing mental capacity. Increasing consciousness
Increasing degree of order. Decreasing entropy
Increasing capacity for homeostasis
56
Discussed by
Raup (1989), Raup and Sepkoski (1982) and Thoday (1953, 1958)
Dobzhansky et al. (1977), Jacob (1972), Kimura (1961), Maynard
Smith (1988), Rensch (1974), Vogel (1991) and Wimsatt and Schank
(1988)
Huxley (1948) and Simpson (1971)
Günther (1950), Huxley (1954, 1959), Kämpfe (1985), Rensch (1959,
1974) and Wuketits (1982)
Schmalhausen see Urbaneck (1988), Herrick (1956), Huxley (1948) and
Waddington (1961a)
Jacob (1972), Kämpfe (1985), Rensch (1959), Wuketits (1982) and
Huxley (1953)
Bonner (1988), Gläser (1987), Hennig (1932), Kämpfe (1985), Rensch
(1959), Wuketits (1982) and Huxley (1953)
Franz (1920, 1967), Kämpfe (1985), Rensch (1959) and Storch and
Welsch (1989)
Grant (1985), Schmalhausen see Urbaneck (1988), Severtsov see
Urbaneck (1988), Simpson (1971) and Vermeij (1987)
Rieppel (1991)
Huxley (1953, 1959), Kummer (1987), Overhage (1963), Schuh (1937),
Theilhard de Chardin (1959) and Waddington (1961a)
Brooks et al. (1989), Kämpfe (1985), Nagl (1986), Patten (1959) and
Weber et. al. (1989)
Emerson (1954), Günther (1950), Huxley (1959), Overhage (1963) and
Wicken (1987)
Table 2. Continued.
Patterns
Replacement within adaptive zones by more successful groups
Increase in efficiency of reproduction. Increase in care of eggs and/or
young
Increasing flexibility of behavior. Increasing degrees of freedom in the
choice of behavioral responses
Increasing autonomy. Emancipation from environment. Evolution of an
internal environment which is both more flexible and more independent
of outer changes
Individualization. Increasing significance of the single individual for
reproductive survival of the species. Survival capacity of the individual
organism. Decrease in the number of individuals of a group and the
number of offspring
Enhancement of ecological possibilities (by emancipation from the
environment and increasing efficiency of organisms)
Increasing control over the environment
Appearance of new morphological structures
Internalization. Relocation of body structures or functions from
superficial to internal positions
Increasing integration of parts (e.g. cells) into the system.
Subordination of parts
The table does not attempt to be complete, but gives an overview of proposed candidates. The representation within summarizing categories may in some cases
violate the original attempts of the authors, what is tolerated to get an overview.
Discussed by
Simpson (1971)
Grant (1985) and Simpson (1971)
Bonner (1988), Günther (1950) and Huxley (1953)
Bertalanffy (1949), Beurlen (1937, 1949), Grant (1985), Günther (1950),
Hendrichs (1988), Hennig (1955), Heuts (1953), Huxley (1948, 1953, 1957, 1959),
Jordan (1908, 1913), Kämpfe (1985), Kipp (1948, 1949), Kuhn-Schnyder (1954,
1967), Lange (1976), Overhage (1957, 1963), Quiring (1931), Rensch (1959) and
Schad (1992, 1997), Schmalhausen see Urbaneck (1988), Smith (1953),
Waddington (1961a, b), Wake (1986) and Wicken (1987)
Günther (1950), Hendrichs (1988), Hennig (1955), Herrick (1946), Huxley (1959),
Kämpfe (1985), Kipp (1949), Lange (1976), Overhage (1957, 1963) and Simpson
(1971)
Beurlen (1949) and Overhage (1963)
Huxley (1948, 1957, 1959), Herrick (1956), Grant (1985) and Simpson (1971)
Kämpfe (1985) and Plate (1928)
Franz (1935, 1967), Heberer (1967), Kämpfe (1985), Remane (1967),
Schad (1992) and Storch and Welsch (1989)
Gruner (1993), Huxley (1953) and Overhage (1957)
57
58

remained, shows that such a term is needed for referring to the underlying
phenomena. Eliminating it from the vocabulary of evolutionary biology is not
a the solution but rather a moratorium.
From the middle of the 19th century there have been repeated attempts to
establish standards, the first attempts stemming from Meckel (1821) and Bronn
(1853, 1858). Most authors compiled lists of the patterns that should be con-
sidered valid (Rensch 1959; Remane 1967; Kämpfe 1985), but opinions di-
verged. Several attempts to operationalize patterns scientifically have been
published, but they did not generate much interest from the scientific com-
munity (Rensch 1959; Simpson 1971, 1973; Dobzhansky et al. 1977; Kämpfe
1985; Wake 1986; Storch and Welsch 1989). On the other hand there was
always a certain general consensus regarding which organisms should be
considered as lower and which as higher. McShea (1998) published one of the
most thoughtful considerations about what might constitute ‘‘largest-scale
trends’’.
Table 2 gives an overview of the patterns that have been discussed in the
literature since 1900. The very fact that there is a large number of them shows
how varied and non-uniform the views were. The following brief discussion of
selected examples intends to show, that most of them raise problems as well as
interesting aspects.
Authors usually proposed several patterns at once, often giving a list and not
always regarding the criteria as pervasive, but rather as features within certain
lineages. Among the patterns mentioned most often belongs ‘‘increasing
complexity’’ (for references see Table 2), not always distinguished from
‘‘increasing differentiation’’. In recent decades, when the term ‘‘progress’’ has
become the subject of criticism, the term ‘‘complexity’’ has often been used as a
substitute. But McShea (1991, 1996) showed that the definition of ‘‘what
everybody knows’’ is unsatisfactory and predominantly based on general
impressions rather than scientific data.
Some authors just took it for granted that evolution generates complexity
and saw it as a product of selective processes. Bonner (1988) and Rensch
(1959), for example, argued that complexity should be favored by natural
selection because more complex organisms are mechanically more efficient,
having more parts and greater division of labor among different cell types.
Others claimed that relating complexity to fitness is problematic, and that it is
not clear whether and how complexity contributes to fitness (Wicken 1979).
Further skeptical discussions were provided by Williams (1966), Lewontin
(1968), McCoy (1977), Gould (1985) and Hinegardner and Engelberg (1983).
Other authors made attempts at defining complexity and making it measurable
(Saunders and Ho 1976, 1981; Papentin 1980; McShea 1991, 1996; Thomas and
Reif 1993). McShea (1996) developed a conceptual basis for objective investi-
gations and found trends of increasing complexity in some measurements but
not in others.
Many authors see ‘‘increasing differentiation’’ as overlapping with com-
plexity and use the phrase in the sense of division of labor. Formulated as
 59

‘‘increase in the number of cell types’’ or ‘‘increasing specialization of cells’’ it

may provide a measurable variable (Bonner 1988; Valentine et al. 1993).
Obviously the number of cell types increased with the generation of multi-
cellularity. But a count of cell types seems not to describe the difference be-
tween an amphibian and a mammal. Also it is difficult to distinguish cell types
by using a standard for comparison.
Increasing ‘‘differentiation and centralization of nervous systems’’ has al-
ways been a widely recognized pattern. Rensch (1959) for example saw it as a
typical characteristic of his ‘‘anagenesis’’ and realized it within mollusks, ar-
thropods, annelids, vertebrates and others. Many biologists considered an
elaborated nervous system with centralized functions to be characteristic of
advancement. Some authors discussed it in relation to assumptions about in-
creased abilities to obtain and process ‘‘information about the environment’’,
or increased ‘‘mental capacity’’. Ayala (1988) and Simpson (1971) regarded the
ability of an organism to obtain and process information about the environ-
ment as an important feature of adaptation, because it allows the organism to
find suitable environments and resources. Some authors argued that this is part
of the more general pattern ‘‘increasing centralisations and concentrations of
organic systems’’, for which the concentration of nephridial organs within
vertebrates serves as an example (Remane 1967). Whether this is connected
with the increasing ‘‘integration of parts’’ (or ‘‘subordination’’) that is some-
times mentioned, for example the integration of cells or organs into the system
or the concentration of different parts in new organs, must be left open until
more precise definitions are available. However, recent molecular biology has a
broad interest in the evolution of integration systems (Gerhart and Kirschner
1997), so these rather old statements could experience a revival.
Some authors considered ‘‘increasing efficiency’’ of tissues, organs or the
whole organism, citing examples such as performance capacities of nervous
systems and sense organs; biomechanical changes in movement capacities;
enhancement of metabolic capacities. However, it may be argued that also in
lower organisms tissues are structured so as to fulfill effectively the functions
for which they have adapted. A similar argument may be applied to an increase
in the ‘‘effectiveness of adaptation’’. But as efficient environmental adaptation
has occurred within all different phyla and classes, and it does not describe
their general differences, many authors have rejected it as being of no value for
measuring progress (Hennig 1932; Huxley 1948; Bertalanffy 1949; Günther
1950; Overhage 1957; Williams 1966; Simpson 1971; Lange 1976).
Some authors hoped that an increase in the ‘‘amount of genetic information’’
would be a measurable trait of progress, but correlation between the supposed
level and genome size turned out to be poor. This ‘‘C-value paradox’’ has been
much discussed (Cavalier-Smith 1985; Bonner 1988; Szathmary and Maynard
Smith 1995). It is especially complicated by the current limitations in knowl-
edge about the functions of non-coding DNA and of the relationship between
genotype and phenotype (Gerhart and Kirschner 1997).
60

Body size, of undoubted importance for many physiological, ecological and

life-history traits, is one of the general features of organisms most often
discussed. Based on a very rough scale from bacteria to whales, several authors
consider that the ‘‘increase in maximal body sizes’’ during successive periods of
life on earth to be a pattern of progress (Bonner 1988). But, according to
Cope’s Rule, increases in body size have played a role on lower systematic
levels too. McKinney (1990) and LaBarbera (1986) discussed the manifold
features of this field based on the available findings. However, it is interesting
to study this pattern in relation to others reviewed here. It is worth noting that
the generation of a new group often starts with small animals (Stanley 1973).
‘‘Energy intensiveness’’ has been discussed in greater detail, not always
within the context of progress. Vermeij (1999) proposed that interactions of
individuals within the adaptation process are expected to lead to size increases;
having a higher productivity or metabolism; engaging in a larger number of
interactions; performing more functions at a higher level and, in aggregate,
showing a pervasive increase in energy flux.
In my view, one of the most promising patterns is the ‘‘increasing auton-
omy’’ of organisms in the sense of an ‘‘emancipation from the environment’’. It
belongs amongst those most often mentioned (see Table 2) and regularly
emerges in discussions as well as in textbooks, especially on physiology and
comparative animal morphology. Here again, authors usually give some
examples but there is not a single attempt to define the pattern more closely or
to describe the phenomena systematically. Consideration of this pattern began
in Darwin’s time. Claude Bernard’s theory of the ‘‘milieu intérieur’’ assumed
that the internal liquids of the animal body are defended against external
disturbances and kept constant, a concept that was extremely fruitful in sub-
sequent physiological research. The constancy would render the organism
independent from vagaries of the environment and would free the organism
from environmental restrictions (Bernard 1859, 1878). Spencer (1864) defined
life as the continuous adjustment of internal relations to external relations and
formulated a ‘‘rule of increasing independence from the environment’’.
In the first half of the 20th century, the pattern was occasionally included in
evolutionary considerations with quite different theoretical backgrounds
(Beurlen 1937, 1949; Jordan 1908; Quiring 1931) and repeatedly emerged to-
wards the middle of the century. The considerations of Huxley (1948, 1957,
1959) are the most well known. Rensch (1959) included it in a list of various
factors of anagenesis arguing that it leads to increasing plasticity of structures
and functions which allow a greater variety of reaction to the surroundings.
The pattern was either seen as being of central importance (Bertalanffy 1949;
Kipp 1949; Schmalhausen 1949; Smith 1953; Kuhn-Schnyder 1954, 1967;
Slobodkin 1964; Lange 1976) or discussed in combination with other ones
(Overhage 1957, 1963; Waddington 1961a; Jacob 1972; Grant 1985; Kämpfe
1985). In addition, it plays a role in both systems and autopoiesis theories
(Varela 1979, 1981; Wuketits 1979; Roth 1981; Luisi 2003).
 61

It reemerges regularly in recent literature (Wake 1986; Wicken 1987; Schad

1992, 1997; Gerhart and Kirschner 1997; Bonner 1998). Vermeij (1987, p. 421),
for example, implies it in the following passage: ‘‘It is possible, however, that
species have improved in their capacity to survive in the physical environment.
Many of the characteristics associated with competitive and defensive superi-
ority – large body size, high body temperature, parental care of the young, and
a tightly sealing exoskeleton, for example – also buffer individuals against
short-term fluctuations in temperature and other physical factors. Conse-
quently, individuals are able to carry on normal activity, or at least to survive,
when physical conditions are temporarily unfavorable. Without such charac-
teristics, individuals would be able to persist in a much smaller range of
physical conditions.’’ But although the principle has been noticed, it has never
been taken really seriously in evolutionary theory.

Autonomy

We focus in our work on a closer survey of this principle as a contribution to

the study of large-scale evolutionary patterns. Here a brief outline will be given,
together with some examples. General autonomy of organisms with respect to
the surrounding environment has often been part of attempts to define an
organism or life in general. These definitions point to the operational closure of
the system, within which the organism establishes and maintains its own
functional identity. The internal compartment is established within a bound-
ary, at least within a cellular membrane. The establishment of the boundary
must have been a crucial step in early evolution when the first prokaryotes were
generated. However, the example of the cell membrane shows that such a
boundary always has a double function: it allows for simultaneous separation
from and exchange with the environment. Thus autonomy is always relative.
But the hypothesis is that organisms show not only autonomy as a general
trait but also differences in the degree of their autonomization within different
taxa. The following provisional definition is offered as a basis on which to
work: increasing autonomy is defined as the evolutionary shift in the individual
system/environment relationship, so that the direct influences of the environ-
ment are gradually reduced and stability and flexibility of self-referential,
intrinsic functions within the system is generated. This is described as relative
autonomy, while, at the same time, numerous interconnections with and
dependencies upon the environment are retained.
Autonomy and the process of autonomization are considered as a complex
of several biological elements. One is spatial separation from the environment as
with cell membranes, cell walls, integuments of metazoans with cuticles, shells,
hairs or feathers. In different degrees they all serve to keep the environment
outside the organism and to regulate and direct exchange with it. Homeostatic
functions establish at the same time the internal functional stability. Another
common element, especially in large-scale phylogenetic shifts, is the displace-
62

ment of morphological structures or functions from an external position into

internal parts of the organism, here summarized as internalization. Multiple
processes of internalization are involved in building up the inner anatomy of
organisms, ontogenetically as well as phylogenetically. During ontogeny, gas-
trulation and neurulation are typical internalizations. During phylogeny, the
transition from prokaryotes to eukaryotes obviously included the internaliza-
tion of: organisms within others (endosymbiosis); functions of the outer
membrane into the internal membrane system; digestion (endocytosis) and the
enclosure of DNA into a further internalized compartment, the nucleus. The
gain in size during these transitions reduces the relative area of direct contact to
the environment by a reduction of the surface to volume ratio. These elements
are prerequisites for establishing a certain amount of physiological flexibility
within a given environment, i.e., a capacity of organisms to generate flexible
functional answers to conditions and changes in the environment. Finally, this
principle can be widened to include all forms of behavioral flexibility, eman-
cipating organisms from mere short-term reactions to environmental factors.
These elements will no doubt need further and more exact consideration, but
they serve for the time being as a useful framework.
The generation of multicellularity in metazoans is an example of how the
autonomy is involved (Rosslenbroich 2005). Metazoan cells generate an
extracellular space that is internal to the organism. This allows cells to create
their own intercellular conditions and to protect them to some degree from the
vagaries of the external milieu. Thus, the cells are internalized into a more or
less homeostatically stabilized buffer zone. The exchange with the outside
medium is increasingly restricted to specialized cells, predominantly organized
as epithelia, while other cells can be completely withdrawn from the environ-
ment and internalized into the extracellular matrix. Epithelia, establishing the
boundary of the metazoan with its surroundings, are supported by cuticulas
and other structures. The occluding junctions between the cells seal off the
intercellular space from the environment, more or less completely, so that the
passage of substances can be controlled. The extracellular matrix and the rel-
ative exclusion of the outside medium are prerequisites for the differentiation
of cells within the metazoan, as signaling processes between the cells need
stabilized and sheltered pathways. And again the transition to multicellularity
leads to a shift in the surface to volume ratio, reducing the relative area of
direct contact with the environment. Also, the building of different forms of
body cavities within metazoans represent internalizations, as interior spaces are
established away from the outside medium, allowing for specialized functions.
Internalization of a gastric cavity allows internalized digestion. The generation
of internalized organs can be traced as elements of increasing autonomization,
because the earlier exchange via exposed general body surfaces in cnidarians
for example, is increasingly replaced by specialized organs for exchange and
regulation functions.
Just to sketch how pervasive the principle seems to be, I want to switch to a
fairly different phylogenetic level, namely the generation of endothermy within
 63

vertebrates. This feature has often been mentioned as an example of homeo-

static autonomy (Bernard 1859, 1878; Huxley 1948; Rensch 1959; Waddington
1961b; Wake 1986), but a closer look at the physiological complex behind this
transition shows that the gain in autonomy is more impressive than is usually
conceded. Physiology has established a relationship in birds and mammals
between homeothermic endothermy and their capacity for movement (Bennett
1991; Ruben 1995, 1996), because the aerobic capacity, i.e., the capacity to
generate energy by oxygen consumption, is in close correlation with endo-
thermy as well as with the greatly extended capacity for sustainable movement.
The functional complex of this transition leads not only to an internalization of
heat production (while ectotherms predominantly take heat from the envi-
ronment), but also to a thermal independence with stabilized metabolic rates,
increased activity capacities, increased independence of behavior from ther-
moregulatory constraints and environmental factors, as well as a generally
more active lifestyle and widened ecological possibilities. This increases the
capacity of the individual to move and behave independently and flexibly
within the environment and to develop complex social strategies. To be more
dependent on the necessity of finding more and higher-energy food is obviously
tolerated in order to fuel the independence gained, which again shows that
independence from the environment is always relative. We expect that through
an analysis of these systems according to the proposed viewpoint of autonomy,
closer insights may be gained into what happened during the transition, as it is
still difficult to explain from existing theories why such a ‘‘wasteful strategy of
energy use’’ evolved at all.
The evolution of immune reactions parallels the evolution of autonomy, thus
establishing an immense increase in efficiency of functions of defense and
individual integrity (Sima and Vetvicka 1993; Wieser 1998). Another example
is the generation of a complex central nervous system which is a basis for a
greatly increased capacity for modulating external signals by internal pro-
cesses, resulting in plasticity of behavior, learning capacity and memory and
thereby increasing emancipation and some degrees of freedom. In evolution on
the large scale, there is an increase in the autonomy of behavior from reacting
directly to physical and chemical factors in the environment to being more
controlled and partly self-determined.
However, all these features did not emerge in just one evolutionary lineage
and certainly not in a linear manner. Increased capacities of nervous systems,
for example, developed independently in different groups such as cephalopods,
insects and vertebrates. Conway Morris (2003) presented a fascinating collec-
tion of such convergences.
The term autonomy as proposed here, does not attempt to describe organ-
isms as entities that are isolated units within their environment. The inference is
rather that each organism is deeply embedded in the systems of its environ-
ment. However, this can be through either more passive, direct physical and
chemical influences, or more autonomized processes, establishing organs for
more active and selected interactions with the environment. The autonomy
64

hypothesis is thought not to contradict the principle of adaptation but rather to

contribute to a further understanding of it and to further considerations of the
relationship between outer and inner, between organism and environment
(Lewontin 2000).
In the sense of Wake (1986), our research program intends first to define,
recognize and describe autonomy and then to attempt to operationalize it
scientifically. However, after identifying many examples it is currently an open
question whether it will be possible to make the pattern testable. It may be
necessary to translate qualitative characters into quantitative data. But as the
pattern can be detected in transitions on very different evolutionary levels, it is
obviously worth working towards such a scientific analysis. Another problem
will be its relation to other patterns that have been discussed (Table 2). Thus
autonomy is not necessarily the only solution to the problem of progress, but
together with others it may be a promising candidate. Presumably they are
interrelated. Initial discussion of this pattern has so far developed on a fairly
high systematic level. It will be necessary to extend the discussion to lower
levels. This may generate more insights into the relationship between auton-
omy and adaptation. Another open question is the relationship of autonomy to
fitness of organisms. It can be assumed that autonomy is able to increase the
fitness of the individual.

Conclusion

Greene (1986) stated that evolutionary biologists can live neither with nor
without the idea of progress, a situation he referred to as a ‘‘biological di-
lemma’’ (Greene 1991). The foregoing discussion shows that this dilemma has a
long history and results mainly from three central problems: (1) The term
‘‘progress’’ carries historical, theoretical and social implications which are not
congruent with modern knowledge of the course of evolution; (2) An incon-
gruence exists between the notion of progress and Darwin’s theory of selection;
(3) It is still not possible to give more than a rudimentary definition of the
general patterns of organisms on different evolutionary levels. However, as
there is a field of phenomena for which the term has until now served as a
metaphor, it is obviously necessary to develop a concept that is appropriate
epistemologically for this field . For this it will be necessary to overcome the old
connotations that the metaphor carries. This was basically what Rensch (1959)
intended when he proposed the term ‘‘anagenesis’’.
But developing a way of describing the underlying general macroevolu-
tionary changes is more important than having an alternative term. Table 2
contains the experience of evolutionists over several generations on this topic.
Some patterns have so far received more attention than others and they are not
equally promising ones. Several may need to be reconsidered in relation to each
other. The proceedings of the Dahlem workshop in 1985, which also include a
table of possible patterns, came to the same conclusion: ‘‘We now reject simple
 65

notions of order such as special creation, foreordination, or that life is just the
manifestation of the postulated law-like structure of the universe. But we are
still left with the facts that there are perceived patterns and directional trends in
the history of life. We must first recognize and define patterns in order to help
us detect and identify underlying structure, cause, or both.’’ (Wake 1986,
p. 47). And in the sense developed in the present discussion, Gould’s (1988)
request to replace ‘‘progress’’ with ‘‘the study of directional change’’ has been
met.
The study of large-scale general patterns is a necessary element of a multi-
perspectivity on evolution. Under such a multiperspectivity the ‘‘biological
dilemma’’ might rather be a fruitful tension – as Darwin implemented it –
within the pluralism necessary for evolutionary theory, a pluralism which re-
cently seems to be emerging anyway (Wieser 1994; Conway Morris 2003). In
addition, it may also contribute to a further understanding of the relationship,
still being disputed today, between microevolution and macroevolution (Peters
1972; Carroll 1997; Gould 2002).
Within such an approach, the old metaphor of ‘‘progress’’ may be aban-
doned, modified or understood in a new empirical light. This is the third way
between, on the one hand, using the term indiscriminately and, on the other
hand, ignoring the general patterns that evolution has produced.

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